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A TEXT-BOOK
OF
PHYSIOLOGY
BY
M. FIOSTER, M.A., M.D., LL.D., F.R.S.,
rtt'-FnritjK ov i-hysiotouy in toe i'mvkfwtv ok CAiiEHinGE, and rEUX>w «f trinity
FOURTH AMERICAN,
FK>>M THE ni-TH ESIil.ISH EI>IT1I>S, THORIIUGHLV REVISED, WITU NOTES, ADDITIONS,
AND TWO HINDKED ANIl EKiHTV-TWIi ILU'STKATIONP,
I'HILADKLPHIA:
LEA KROTIIERS & CO.,
18iH.
•1
PREFACE TO PART I.
Ih the preaent ediUoo I have made oonridenble chuiges and addition! ;
bat in the changes I have tried to nudntain the cbaiw^er of the book as
{weaented in in«yioiu edition! ; and the additions, with the ezoefAion of
tbe histological paragraphs, are oaosed not by any attempt to add new
matter or to enlarge the goural soc^ of the work, but by an efibrt to
•zplain more fdWj and at greater length what seem to me to be the most
fbndamental and most important topics. I hare beoi led to introdnce
some bistolopcal stiUements, not with uie view of in any way relieving
tbe stndent from tbe neoeaity nf sbidying distinct histolc^ical treatises,
bat in Older to bring him to the phyuologjcal problem with the histological
data fresh in bis mind. I have therefim dealt very briefly with the
several histological points and confined myself to matters having a physio- -
logical bearing. Hy friends. Dr. Gaskell, Mr. Langley, and Dr. Lea, have
given me great assistance throughout, and their names might fitly appear
on the title-page, were it not that the present arrangement makes me alone
responsible for all shortcomings. I have also to thank my senior demon-
strator, Mr. L. E. Shore, M.B., and my junior demonstrator, Mr. Wing-
field, M.A., for much valuable aid. The second and third parts will follow
this first part as soon as possible. .
PREFACE TO PART III.
I AM of course aware of the disad vantages of issuing this edition of my
Text-boot in instalments, and very much regret that this part does not
complete the work. The failure to get the whole of the remainder ready
has been due to lack, not of will, but of ability and opportunity.
I take this opportunity of thanking my friend. Dr. Gowers, for the loan
of two woodcuts, as well as for much valuable advice. Throughout the
whole of this part I have been largely assisted by my colleague, Mr.
Langley, and by my friend aud former pupil. Dr. Sherrington. The
latter, besides helping me with criticisms, has prepared for me most of the
figures after original drawings by himself What little merit there may
be in this part is largely due to these two gentlemen.
M. FOSTER.
CiMBKiDcm, September, ISflO.
AMERICAN PUBLISHERS' NOTICE.
Thb Mtthor's pn&oeB to the eeveral parte, in which this edition was
imied in fiogland, will show how thorough has l>eeQ the revision to which
H hw beeo Bafajected. Tlie task of the Amerioan editor has, therefore,
bean mostly coofined to the adaptation of the work to the wonts of the
Asmioaa stad^t, Among the principal additions will be found refer-
eooee to the phyriolo^cal action of some of the more important drags,
and the test has been elucidated bj an increased number of illustra-
tioais. All matter thns introdaoed has been distinguished hy inolosure in
bnckets [— ].
Pb 1 1.1 DELPHI A, October. ISSI.
CONTENTS.
PAGE
ISTFOprCTlOX .33
BOOK I.
BL'Hip. THE TISSUES OF MOVEMKNT. THE VABCILAR MECIIAKISM.
CHATTER I.
BLOOD.
The Clotting of Blood -ii
The ( "onmscles of the Blood : The Red Corpuscles -H
The W'liile or (itolorless CorjmscleB Ill
Hl.xxi PlateletH 68
Tin- ' 'heniicul Compoeiiion of Blo<id KO
Tlie ijiiiinlit^ of Blood, and itn Distribution in the Body . . . . Tl
CHAFTKK II.
THE a>STRACTILE TIPf-UES.
The Phenomena of Miiwcle and Nerve : Muscular and Nervous Irritability 73
The I'henometia of u Simple Miiacuiur Ponlrflftion f^'i
Tetanii; l.'ontrHctiona 'J1
* In the Chanpw which Take I'hicv in a Muscle Dnringii ('mitriiction: The
t 'han^e in Form ........... 94
The i 'heraiatrv of Muscle 1U4
Thermal Changes Ill
Klt-ctrical Chaiijres 113
The ("hanger in ii Nerve during the I'lumiige of a Nerviin^ Jjnjiulsc .UN
The Nature of (he Changes through which an Electric i urrciil is Able to
(ienerate a Nervoun Imiiulw: Action of the Conwlant Current .!:;■<
The Muaele-nerve Preparation aa a Miichiiie I3,i
X CONTENTS.
rAi;y.
The Circu instances which DeCermiDe the Degree of Irritability of Muscles
and Nerves 140
The Energy of Muscle and Nerve, and the Nature of Muscular and Nervous
Action I4<)
On Some Other FormH oC Contractile TiMue: Plain, Smooth or Unstriated
Muscular Tissue 149
Ciliary Movement ..... 154
Amceboid Moveinenta 157
CHAPTER III,
GENERAL FE.\TUEEH OF NERVOUS TISSUES. . IW
CHAPTER IV.
THE VASCULAR .MICCHANISM.
The Structure and Main Featuresof the Vascular Apparatus , 17.'!
The Structure of Arteries, Capillaries, and Veins 173
The Main Features of the Apparatus 18!i
The Main Facts of the Circulation 184
Hydraulic Principles of the Circulation 193
Circumstances Determining the Rate of the Flow 199
The Heart a06
The Phenomena of the Normal Beat 20<i
Endocardiac Prmsure 213
Summar; :]24
The Work Done 22.">
The Pulse 22G
The Regulation and .Adaptation of the Vascular Mechanism : The Regula-
tion of the Beat of the Heart 240
The Histology of the Heart 241
The Development of the Normal Beat 24-"i
The Government of Heart-beat by the Nervous System .... 2.';J
Other luliuenceg Regulating or Modifying the Beat of the Heart . . 2<>:!
Changes in the Calibre of the Minute Arteriex. Vasomotor Actions . . 26-~>
The tkiursc of Vaao- constrictor and Vasodilator Fibres .... 27:!
The Effects of Vanomolor Actions 27.')
Vasomotor Functions of the Central Nervous System ..... 27i!
The Capillary Circiiluti'in 2.S(>
Changes in the t^uiintiiy of Blood 291
A Ueview of Some of the Features of the Circulation .... 292
CONTEXTS. xi
BOOK II.
THE TISSDE3 OF CHEMICAL ACTION WITH THEIR RE3PECTIVE
MECHANISMS. KUTBITIOX.
CHAPTER I.
THE TLSSUES AND MECHANISMS OF DIGESTION.
FAQE
The Oharactera and Properties of Salira aad Gastric Juice: Saliva . . 302
< raatric Juice 307
The tilriicture of tlie Salivary Gland«, ihe Gastric JIueoua Membrane, the
I'kDcreas, and the CEsophagus ''IT
The Siructure of the Stomach 319
The .'■alivary Glands ^^^
The I'aDcTeaa 328
The Structure of Ihe (£«o]>h&gus 330
The Act of Secretion of Saliva aod Gastric Juice and the Nervous Meelian-
i^mt which Hegulate it 332
The Changes in a Gland Constituting the Act of Secretion , , . 339
The rroperties anil OharacteM of Bile, Pancreatic Juice and Succus
KntericUH 351
Itile :«!
I'auiTe.itii; Juice 3.54
."■iiccus Kntericus ,...,.-...■■ 358
Tlie Secreiiim of Pancreatic Juice and of Bile ...... 3o9
The Structure of the Intestines: The Small Inleatine .... 365
Tiie Lar^ce Intestine 3i4
The Muwular Mcchanisni!> of Di^jestioii 375
Tb.' Changes which the Food I'ndergoet in the .Vlinientary Canal . 3W
The (.'hnnges in the Stomavh ■*'*■'
In the Small InteHtine ■'«!
Ill the Large Intestine .......-■■ 39;)
Till- Fei-ew 3!''>
The Lacteal)! and the Lymphatic Syntt-ni 31Nj
The Lyriipliatic Veiwels '■'•*'
Lymph ciipil la rie.* '■'■'•''
TI»'.'=trnctureof Lympliaiictilunds -1"-
The Nature and MovenieiitM of Lymph (Irirhidirig Chyle) . - -1'"*
The Chnratters of Lymph ^"'■^
Tlip Movemt-nta of Lympli ^ll
.\li-iirptiiin from the .Uimeiil:Lry Canal 419
Tin- CoumeTalien l>y the Several I'roiluct.-* of Hige-tLiiu .... IIH
The Slechani^ni of Ahsiirptiori -I-'*
Xn CONTENTS.
CHAPTER II.
RESPIRATION.
The Structure of the Lungs and Broucbiul PMssagen 4:11
The Mechanics of PulmoQiiry Res gii ration 43K
The Respiratory Movements -i*-*
Changes of the Air in Respiration 44!>
The Respiratory Changes in the BiooU *)I
The Relations of 0:cygen in the Blood 4-35
Products of the Decomposition of Haemoglobin 4I>3
The Relations of the Carbonic Acid in the Blood 4)iii
The Relations ot the Nitrogen in the Bloi>d 4ti7
The Resjjiratory Changes in the Lungs : The Entrance of Oiygen . . 4Gr
ThcEititof Carbonic Acid 470
The Respiratory Changes in the Tissues 471
The Nervous Mecbaniam of Respiration 475
The Effects of Changes in the Composition and Pressure of the Air
Breathed 402
The Relations of the Respiratory System to the Vascular and Olher
Systems . .' 496
Modified Respiratory Jfovements ■"•"O
CHAPTER III.
THE ELIMINATION OF WASTE PRODUCT^*,
The Structure of the Kidney . ■'ill
The Composition and Characters of Urine '>24
AmountH of the Several Urinary Constituents Passed in Twenty-four
hours. (After Parkes.) -J2S
The .Secretion of Urine '>29
Secretion of the Renal Epithelium 't^i'
The Discharge of Urine ■'>46
Micturition '>48
The Structure of the Skin V)]
The Nature and Amount of Perspiration ....... 'iW
Cutaneous Respiration .IliO
The Mechanism of the Secretion of Sweat .'•112
CHAPTER IV.
THE METABOLIC PROCESSK-; OF THE HODY.
The Structure of the Liver .'•(i-'i
The History of tilyc'ipeu 'iT'.i
Diabetes ■'•^'-
CONTENTS. Xlli
PAUI
The S{ile«n TiSS
The FDrmation of the Conatitueats of Bile 595
On Urea and on Nitrogenous Metabolisin in Ueneral 599
On i*onie Structures and FroceeneH of Obscure Nature .... 607
The History of Fat. Adipose Tissue 614
The Mkmmary Uland 620
Avt-rage Com poei lion of Milli in Diflerent AnimnU 624
CHAPTER V.
XUTRITION.
The fftatlstics of Nutrition 628
(.'o[n]iiiriBon of Income and Output of Material 630
The Enerpy uf tlie llody: Tbe Income of Energy 637
The Expenditure 638
Animal Heat 641
On Nutrition iu General 662
«>n Diet 660
BOOK III.
TilK TESTRAL SERVOf.S SYSTK-M AND ITS INSTRrMKNTS.
CHAPTER I.
THE piIMXAL CORD.
( *u Slime Features of the Spinal Neries 671
The ."Structure iif the ."Spinal ( Vird 676
The Reflei Actions of the I^piiial Cord 711
The -Autumatic .Vctions of the Spinal C'ird 724
CIlAI'TKH 11.
THE ItKAIX.
t >ri S(.me General Featurex of tliv Slnutiirr of the Brain .... 731
The itulh 7;!(i
The lJis|>osition and Oonneciiipii- <>!' tin' Oriiv and White .Mutter <>( the
Ilrain: Theliray llatler T4'<
XIT CONTENTS.
The Central Gray Matter and the Nuclei of the Cranial Nerves . 748
The Superficial Grny Matter 762
The Intermediate Gray Mailer of the Crural System 762
Other Collections of Gray Matter 772
The Arrangement of the Fibres of the Brain 774
Longitudinal Fibres of the Pedal Sytitem 775
Longitudinal Fibres of the Tegmental System 778
Tranarerse or so-called Commissural Fibres 781
Summary 782
On the Phenomena Exhibited by an Animal Deprived of its Cerebral
Hemispheres T>^4
The Machinery of Coordinated Movements 790
On Some Histological Features of the Brain 79^
The Superficial Gray Matter of the Cerebellum 800
The Cerebral Cortex 803
On Voluntary Movements 808
On the Development within the Central Nervous System of Visual and of
some Other Sensations : Visual Sensations 83ti
Sensations of Smell 847
Sensations of Taste 8o()
Sensations of Hearing S-'iO
On the Development of Cutaneous and some Other Sensations . S'>I
Some Other Aspects of the Functions of the Brain K6''i
On the Time taken up by Cerebral Operations 873
The Lymphatic Arrangements of the Brain and Spinal Cord . 676
The Vascular Arrangeinenls of the Brain and Spinal Cord .... 881
CHAPTER in.
SIGHT.
Pbysiolc^ical Anatomy of the Eye 887
Dioptric Mechanisms: The Formation of the Image 894
Accommodation S'J^'i
Imperfections in the Dioptric Apparatua 90.'>
Visual Sensations 908
The Origin of Visual Impulses 909
Simple SenaatioDs 'Jl.">
Color Sensations 918
Visual Perceptionii 9L'-%
Modified Perceptions 927
Binofuiar Vision : Corresponding or Identical Points 9;it)
Movements of the Eyeballs 9:W
The Horopter 9:i4
Visual Judgments 934
The Protected Mechiinisma of the Eve 9:!7
C0XTENT3, XV
CHAPTER IV.
HEARiNO, SMELL, AND TASTE.
lleurin);; I'hjsiott^icHl Anatomy of the Enr 939
The AcnuHtti' ApparalU" ... .... 946
Au<]itory ^nxAtiniis .... , ... 94S
Auditury ,Iuclginetit« !iri2
SiufU: rb.vsiulogicsl Anatomy of (be Nasal Fusiia! 9.'>3
Tafte : I'hjBiologicai Anatomy of the Gustatory Mucous Membran*^ . !>M
C'lIAPTEIt \.
FEELING AND TOL'fH,
■ ieneriil Sensibility and Tactile Perceptions 960
Tnctilf Sensations : SeasHtions of Prew'Ure Dtil
.-^f niuitii)nB of Temperature 9(>2
Tactile PerceptionH and Judgmcnr^ 9t!4
The Muscular Sense 966
CHATTKR A' I.
SPECIAL MUSCULAR MECHANiyMS.
TheVoiie; The Pliysiologlca) Anatamy of llii- Larynx . . fHiS
Sjieech ; Vowelii ..... 974
l.Vmmtnaiils .... 97^1
I.oconiiitur Mf('hu^il^mB . 977
BOOK IV.
THE Tlss^■^:.<5 ,\si> mechasisms or HEritcun'mox.
(.11 APTKi; I.
UlUiANS (H- KEl'imiin TKIN.
Tlie I'hT!'ii)logiciil .\ii»tiifny of ilie "rpirm 'irdeniTutioii . 9S2
CllA I'TKi; 1 I.
MENsrillATHiN . . us?
XVI CONTENTS.
CHAPTER III
PACE
IMPREGXATION 991)
UHAPTKR lY.
THE NITRITION OF THE EMBRYO . . WT
CHAPTER V.
PARTIRITIOS nm
rHAPTEU VI.
THE PHAGES OF I.IFE . J'"*'.
CHAPTER VII.
DEATH . 1013
A P P K X D I X .
THE CHEMICAL BA:?IS OF THE ANIMAL lODY . lOL".
A TEXT-BOOK or rilVSIOUHiY.
I
I
INTRODUCTION.
|L Dnurriox, aM«l l)f tuIcroMoptcal examination, IcitobM us ihat
Iba bodr of uuui is vamt\« un >i( c*^riain kiiiiltt uf uauHftl, «» ilifli^ring Anm
Mch niMf In uptical iio'l oilier |ili)'*ir'itl oliuractora ami *o built up logCtlMtr
a» lo gift ibt body cvrbiin olructural fmturM. Clii-iiiiunI I'xaininntioD
flutbtr UMba IM UuU iIkw IciiKlo i>f nuit'-rinl nn- r4iiii)Hi«n) of various
dMBlical aubOAnOM,* largr niiriitwr tif iihicli liitvi? lliiM vliitnuumti'' thnt
Ibftjr |M«MB • CDtuidArable luiiutiDl ol' pvlvutial vficrgy <:iii>iiblo of Imidk mI
frw. fmiUTtd actual, by oxi'Iittion or Kitno other dM-micnl cluingc. Thus
ibr bndj, u a whuk-. may. fmm a cbemtciil point of viow, Ira ct)usidorod as
■ u*m of various cht^micml substADflM, rpprawnling altogether a oonsidora-
bU capital of {Mtential energy.
iS. Tbb Inxly rnny exisl fith<-r tis a living body or (for n certain time
■1 tSBSl) as a dead body, and the living l>ody may at any time booome a
dMd body. At what is geoerallv calle<l the moment of death (Itut arlifi-
eUly to, for as we aliafl see t^e processes of death are numeroiia and
gnHMnl) tbe dead body so fkr as structure and chemicnl romiMiaition an
CBdOCrasd >• esceedbgly like the living biMly ; indevl the dittVri-im;* b«-
tWMn iba two are Hicb us can Iw deuruiined unly by vi-Ty carofnl i'i«niiaa-
tiao. six) are ritll tit a largi' I'xlcnt estimated by drawing infrivncoi rnlhor
actually <>Wrveil. At any rate the dead l»Hly at llii> luonieiit of death
ibla thi- living body id hu fur an it rcpnveoUa GUfNtal of potential
Pront tliat mofncot ouwsnl. hnwovrr, the cnptMl is expended ; by
vbich nn- Inrp'ly i!u.w of oxidiiiirin, ihc ttoi.-rKy i" gnidoslly dis-
aipBlad, leaviDg the Iwdy chii-tly i» tlie form of heal. While thc»e ohomical
[wwcBMM atw going on Uio structural features disappsar, and the btxty, witb
tbm ln« nf nearly all ittfenergr, if «i hut rasolv«d into "dust and aahoe."
Tfao tiharacterislic of the dead bixly then is that. Iwing n maw of sub-
■laaciv of coDiidcrable polcnlial eaergr, it 0 always more or tern elowly
{■lag seergT, never gaining energy ; ibe capital of energy preecnt nt the
■ooMit of oMtb is more or less slowly diminishod, m never increased or
|3. When on tli« ntfaer band we study a living body wo ar« struck witb
ibc fullowinj^ Ntlicnl fads :
1. Tbe living body moves of iUelf. cither moving one part of the body
on a04itber or raoviug tbe whole body frtm [iloce lo place. These utov*-
arc active; llie biidy in not (iiniply pulled or pushed by external
lann, but i1m> motive powfr id in the body itself, tbe energy of each more-
nsttl i* supplieil by tbe budr it»elf.
2. TbflK niovtrnuiits uo OetnrmitKil and inSnenocd, indeed often sewn to
slan4<l, by duugM in the surronndintpi of the body. iSuddeu contact
the sur&oo of tb* b"dv and wme foreign oliject will often call
8
IKTBODWOTIO!*.
fortli n nifivcciipiit. Tb« body is wosilive to chonxen iu iu surroun< _ .
arKl tlii; fcnniLivi-iK-H u iDaDifceud uoi only by niovenictila tmi by OtStt
choDgcs in thn boctr.
3. Il is contintiiillr gtroerftting heat and giving out bent losurrouiidiii);
things, the produdioii nixl Ium of ht-nl, m lh« vnbw ol' nimi uod ci^rtniit
oilier aDimslR, being to ii<iju>l<(l Omt lliv uliulti burly i« wurm, tliul t* of ■
l€iui«rtiiurc Iiiglior thiiti tliut ot KUrroiiodiiig ihitigit.
4. From lime to time It cnU, thm is to «ty tnlc<:i into itself supplies of
ccrtnio eubnlntices known m fooil, ihvm sutMlmici-s boiiig in lliv moin rirailnr
to that which compoee Iho buity, itni] bi-ing ]ik<- ihcm dicmicfll bodiiv of
OODBldeniblfi potential energy, cnpnblc tbrougli oxiiljilion or other chemiciil
cbangee of setting free n considcntble (luunlily of energy.
5. h is continually brenllilng, that is, Inking in from the surrounding air
supplies of osygcu.
6. It is continually, or from time to lime, discharging from itself into ile
turroundings eo-called waste matters, which waete matters mny be broadly
described as products of oxidation of the substances taken in as food, or of
ihv tiubstances c'omjKiBing the body.
Heuce the liviuj; body maybeaaid to hedistin^ished from the dead body
by liircb main features.
The living hoily like the dead is continually losing energy (and lonng it
more rapidly than ibe dead body, the si>ecial breathing arrangements jier-
nitting a m«re rapid oxidation of its sulistanee), but unlike the dead Ixxly
is by loeana of fond continually restoring its substance and replenishing its
store of energy.
The energy set free in the dead body by the oxidation and other chemical
changes of its tiibslance leaves tlie body almost exclusively in tbe form of
beat, nberens ci great deal of energy leave* (ho living body as mechanical
work, the result of various movements of the body, and as ne shall see a
gre«t deal of the energy which ultimately leaves tlie t>o<ly ns heiit, exists for
a while iriibin tbe living body in other ibrms than heal, though eventually
trsnnformed into heat.
The cbaogea in the surrounding aflect the body at a slow rale and in a
general way only, simply letKomg or increasing the amount or rate of
cheivical change and the quantity of beat thereby fct free, but never
diverting the energy into fuaie olher form such ns that of movement;
vhereai changes in tbe surroundings may in the case of the living body
npidlv, profoundly, and in special wava affect not only the amount but also
tbe kind of ener^ set free. The dead body leH to itself slowly fiUls lo
Eiccra. slowly dissipates its store of energy, and slowlv give« out beat; a
igber or lower leniperuturc, more or len moisture, a free or scanty supply
of oxygen, the advenl of many or few putrefkctive organitunH, tb't«e may
auicken or xlack^n the rate ut ublcb energy is being tli.-elpaied but do not
diveH that energy from heat into motion ; whereas in the living l>udy so
slight a change of surroundingn oa the mere touch by n bair of some par-
turn lu* turfacVi may so aflect the setting fixe of energy as to lead to such a
disdMlf^ of mergv in tbe furm of movemi^nt that the previously appar-
ently quiescent bcHly may be suddenly thrown into tbe mo»t violent oouvul-
sion».
The difference, therdbre. between living snljslantT and dead sulislnnce
though recondite arc very great, and the ultimate object of pbysiulogy
b to ascertain how it is that living substiincc c«n do what dnid sub-
stance cannot, can renew its substanoe, and leiilcnish the energy which it is
cvnliuually loeing, and can, according to the nature of its surroundings,
vary not only tbe amount but also the kind of energy which it sets free.
J
INTlIOItCCTIOX.
35
Thw ihtrr oiT two gmt (livtuon* of ph^Klalggv : uoc having to ilo nitli the
nBM««l of MbMaoM wu) tho rojtU-tnuhinvnt or oicrgy. tbo Uher having to
de with th« Mttiiig Art of «Dorg}-.
14, Now thtr body "f nmu (or oih- oI' iIh- hi):tirr iLnininU) in n very d^ni-
piMstvil ■trucluiv. cuii«i#tiiij; ol' ilitli-mil kiixU oC i)iali'ri«l, which nv call
litamm, Mirh le innsculur, i»rvifii«, (iMiin^ocivo, nnd cho likv, variuuMly
amspd in organs snrh lu hwirl. 1iii)g«. mutclpe, «kin. vtc, all built up i<i
fern UM bud^ awoMin); l<i o-naiii nn'riiliol'igimi Iswh. But all this vum
pUoatioo, though lulvautngt'oug and indeed nccvesary Tor ihu fulkr lifts of
ia oeL taKUtial lo the existcnvo of life. Th« amwha (Fig. 1] is a
(Fw I.
I
a«»rtnmiMa»ii»lnilllfcWial>iniiH]u».c)— nuil br ■■MwmiiuilmU.I
bctoj; il ran^WM its Mibdnnce, rvpVni»h«9i il« fivro of encrf-y, and
eaergjr now in one form, now in unolli^r; and yet tho »m(Bba may
he <•■>] ti) hare no (ueunt and no orgnni! ; al nil cvx'nt« Ihis is true of cliiaely
alltnl Uui not ft well known simpU' b«ings. U*int; iho more fHniiltar
maaaln aa a lyp<^. and. lbcr«fori.>, leaving on one Fide ihc nucleua. and aor
ifclllii linn between enduurc nud eclcKin-, wv may say that iu Imly W
boataceaMUB in thf aeoac (hat if we divided it inio small piecea, each pleoe
vmtlalM! lik« all the nthera. In another sense it is not bomoffeaeoua. For
wa knn** (bat the amioba T««eivee into its sulwiaut.'e inateriu m* fuod, and
tk*i thu frxid or part of it reruaius lodged in the body, unul it is made um
of and liuilt up iul^t ibe living substance of the Itodj, and each niece of the
Bvios Mibatancr «f the body mum have in or near it some of the material
«Ud tt •• al>»ut to build up into itaelf. Further, we know (hat ihe aniwha
^vw PUl watte mattvra mcfa a« carbttnic acid and other subttaiicea, and each
ecc of Uio aiarrtw lauu contain Kiiite nf theae wnate maUers abotit to be.
( not yat, iliachatgad frofo the |>iece. Each piece of ihe amoeba will,
tbatvlhrw, mntain ibMO three Ihinga, the actual living subalanoe, the food
at- I" living (tibatiiDCo, and the waate matter* which have eesaed
U..rftcr, wv have renauno to think (.hnl the living »iib*tanoe doea not
hnak down int» (hv waMo mattcm which K«ve ih*! Imdy ui a dngle bound.
iMt ihara arc Mui.'v* in (he d'lnnwiirti pntgnw liciwe«n the one aud the
•char, fjimiliirly. ibongh uiir knnwlivlge mi rhio (H)int in !<■** inrv, we have
naaiiri U> Ihink that ih*? lnttA is not iDOi>rj>oral<-il into (he livingnbatanco at
atio^ tie]!, but that there arc Xagea in llw nimard prngreai f^om the
' food to the llvinjc aubatancc. Each piece of the body of the am<»bii
IlfTBODUCTION.
nil], Uierofure, contain Bubstancoa npreeeiiting various etages of becomioK
liviag, Bnd of oea^iii]; to be living, as well as the living substAiice ilself.
And we may safely uinke ilib slateiiieiit, though we are <iuite uoable to
draw ihc line, where the dead foixl mi iu way up becomes living, or tbe
liring tiibnlaticc on iu way down bec<iiuee dead.
Sft. Nor id it nece*iBary for our present [iur[i<we In be able to jioint out
under thv inicnwrofM;, or li> itotcribe frurn ii bial(>l(i){ii.-«il )ii)inl uf view, tlie
part^ nhirh are living' and llie purlii uhicb an^ dfad fiiod nr drad wanu.
jIiv biHly of till- ani<i;ba in rri-<|Ui.''iiliy sjinki-n uf a* ounMtiling of " jiroto-
plasm." Tlie name wita orifiinaH > given in the niiiUcr funning thv iirimor-
dial utricle of ihu v^Mab)*: fell n-' ili»tiu|iii9ln'd fnim tlic d-ll walfoo the
on« hand, aud frum thn fluid c»nt<'nl> ot Iha- oi-ll or l-cII sap on die othrr,
and hIho wo may add t'ntm the niicli'ue. It ha» «inco been applied very
generally to Ruch parlx of animal boiltcs as n^cimble. in their general fea-
lure«, tlie primonlinl utricle. Thus the l><>dy of a while blood -corpuscle, or
of a gland eell, or of n nerve cell, i* said to consist of protoplasm. 8uch
pans of animal bodies &;< do not in their general filatures re^mlile the
matter of tbe primordial utricle are not called protoplasm, or, if they at
•ome earlier stage did benr such resamblance, but no lunger do so, are some-'
tiroes, as in the case of the substance of a muscular fibre, called "diireren-
lialed proto[ilflsm." I'roloplnsm in this sense Bometimca appeals, as in tbe
outer part of most amceba:. as a luaaa of glaesy-lonking material, either con-
tinuous or interrupted by mure or less spberica] suaces or vaouolen 6lled
with Buid, sometimes as in a |;land <%ll as a more retractive, cloudy -I looking,
or finely granular material arrau^ed in a more or lc«a irn'gnlar network, or
spnngowork, the inier*ticeH of which are occupied cither by fluid nr by *ame
material iliflerent IVoni itself. We »halt return, however, to the fvuturcM of
this " prulopliwm " when we conic to treat of white blood-corpuscbis and
olber "protoplonmic" <truclurKL Mentiwkile it is sufficient for our present
iturpose to note tliat lodged in the protoplasm, discontinuous with it, and
brmiog no part of it. are in the firvt place collections of fluid, of watery
solutions of various substances, oci'Upyiog the more regular vacuoles or
the more irregular epaoes of the network, and in tJie second place discrete
granules of one kind or another, also forming no part of tbe protoploiiu
iLsvlf, but lodged citbei- in the bars or substance of the protoplasm or in the
vacuoles or meshes.
Now, there can be little doubt that the fluids and the discrete gnmulu ara
d*Md foixl or dead waele. but the present state of our knowleiTgir will not
permit us to make any ver)- definite statement nboul the prot«i]ilnsui it*elf.
We may prubably cuucludc. indeed we may be almii>l Hunr, that pnitnplnsm
in the above sense is uot all livinjc Hubstauce, that it i* made up partly of
the real living iiulbtnni'e, and partly of material nbieh n bovuming living or
luis cpiUM.-d Ui lie living; aud in the caa^ wlien^ prtrtuplasm is dnoribco as
fenniog a network, ii is iiiK'>ible that some of tbu mnicrial occuprin^' the
mcshm of the notwnrk may W, like ^rt of the network itm^ll'. really alive.
" Pnrtopbism" in fact, »-■• in the nwiio in which wc arc now using it, and shiill
continue to use it, i» a nwrfihuliMjiml lerni ; but it tnust be iMime in mind that
llio same woni "protoplasm" L* alwi Irpiiuently used to denote what we have
just now called "the rtial living sulxrtnnre." The word llien embodies a fihy-
niologiealidfii; so UMil it may Iw applied to the living eubittauceof all liviOK
Btnicturre, whatever the microecopicnl features of tboec Htructures; in this
sense it cannot at prrcont, and piwsibly never will be recognized by the micro-
scope, and onr knowledge of its nature must be bused on iufereooea.
Keeping then to (he phrase "living subslmiee" we may say tliat each
piece of the body of tbe araai?ba consists of living substance, in which are
I
I
tXTROnVeTtOK.
87
lodnd, or wiih wim-li nrv l>uill uj) in mih^ vay or utbcr, food sod wnste in
TmiVHu lUgte.
Now. M) ftRioba nay divide itself into Iwo, acb half exhibiting nil the
phrBoHMOa of lh« whole; and we caD easily imnKine the proccM lu be
rHHU«d, until the amatba was divided idIo b niuUitudc of exceedingly
Mtaota aintrbii-, cat-h having aJI the properties of the orij;inu). Hut it ia
slnrioa*, *■ iu ihv like divisioii of a maea of a clieniical Hubttuiiee. that the
ditrUoa owild ui>t he repeated indctiiiilely. Just an tu divi«ioi> uf the cbein-
kal mtm we r<»iuc to (he cWinicul molecule, further divUion of which
ihangw iIm pruiwrtiee of the HuhDtnnt^, *» ia Ihe ooutitiued divbiun of the
amrrha we should runte lo a aiiae iu which further diviaioit interr«re<) with
tl "gicnl si-li<>»d>. we iJimitd ui>iii«toAphy«ii)lc>;;ifal unit, curnwjioiid-
iii.i ;rv«lly niun* couij>lex ihoo the chemical inoleailc.' Thid tinit lO
nmain « iibyvioliijtical unit and to conlinne to live muitt viiniaiu um only a
pwnluci of tltv livinj; KuliMaDou but alto the food for that liviuj^ itulMtance,
ID MTtral at li<«)4 of tlw Mak^. frum th*3 iniliid raw I'ihhI up lt> ihc liiml
** living" siiigvB, and niuvt itinnlarlr contain variuui ttaifui of wa»tc.
{ 6. rJow, ihf- grwil cliaraclmMic of the lypicu) aoHchn (Icarin^ out ihc
BO-'—— '- that, ■■ far ■• we mi ucertnio, all the [>hy»iii]Dgical unit* arc
ml bU do tlH^ «nif thing*. Kach and every purt of tin- body
r*!-. iif i—,l nun nr lew raw and builds it up into iUown livln); Kulwtancc ;
nach aiul ttn-ry part of tlw biKlr may bf nt oiie timv ijuiwcoDt and at another
in nntian; oadi and every part if sensilivo and rngmnds by muvvmeHt or
albiniiaB lo varioaa ehangca in tls aitrroundini^.
Tlte Wly of nun, in it> firet eta^c, while it in yet an ovum, if wc li^iv«
•aiiie the nucleua Mtd Mglet't ditTerencee caused by ihc unequal distribution
«if fotal material or yolk, may alao be raid to bo <mmpoiied uf like parts or
like |4iT*ii>lof:ical unita.
Br tlw art nf MffoneDtatloii, hiiwevcr, the ovum lit divided into PArLS or
e«li* which early aDow dillurencca from each other; and these diOerenoea
ra|tidly iiwreM* ■• ctoTelopmeut proeeeda. Some rell* put on certain ohar-
metrr' and ntlHn Other dtanolen— thai it to May. the eell« undergo hidologi'
»i 'iallon. AikI thit lakt* place in *iieh a way that a number ol
eci„ :. ...,: Kigvlhrr in a group become eventually convened into a Hmha,
and Iho whuk IhmIv boeofdM a ooltociion of such tiMn<.« arnntftiil together
acmrdio;: to morphologtcnl lawg, rnich tiMue having a definite •tructure. its
aallular nature Ining H>mc4iRi<« proaervcd. aomatimee o)wcun-d or ev<in loot.
Tfaifl hisiologicikl <It8e rent iat inn h accompanied by a phyawtnt/ienl divitioH
tflalor. Each tiaane may bo supjmeed to Ih? composed of physiologicid unilM,
UM units of the tame tieeue being alike but difleriiig (mm the unit* of other
tiaww ; and orrwjmniling to ihi^ditrerenceof «tnictiire, the units of ditfeniit
li»»iia behave m act dilferently. Instead of all the units, as in tlie amo'tigi,
daiay iIm< Kam^ things e<iually well, the units of one tinuo are told off, u it
m^n, >' ' hiu): eH{<e«ially well, or especially fully, and thus tho whola
labPT' i> ii divided amonK tlie several tissue*.
rnl lissties may thu» be clnx^itied according to the work «hi«li
ll.' . ' ' . and the Gnt great dUlinciioii is into (1) the tinues wbj«li
■!• omovnifd in the wttio); IVm- of ener[.'y in sjx-cinl ways, and CI i the tiam«e
whirh air ci>nccnii<d in n:pl<-ni>>hin); tl>e^ub?tan«'alldt)')rvlIening the energy
*f
1 :. , ivsioloftient tinSt nf tlio nmivbu while it b engaged in setting Ave
*aar|cy an a* to move itfflf, and hy rcaum of ilv wnBitivenen so directing that
flsatiQr as Iu jirtHluce a movamtnt vuiuble to ilio condillous of ita uurround*
■ Sorli • ]ilijik>luK>«l «nil miflkl la e»\Ui ■ ttmmntt.
INTBODUCTIOW.
ingit, liANat th*> nnme time to t»-jtr ibc bibur nf uikin^ in mw foni], Qfiiclrctitig
tliat pari «f the mw fmiil wliioh Li ii^criil mii) nycfling thut which if iisclcn,
and of worlciiiK up ihc nccitpti^) piirl ihrnugh n vnriHv of vla^e« into It* onn
living sub«litnce — that U In my, it hat nt tlii; Minic lim<? thnt it is lecling and
iDoring to currj on llio wurk of <ligwting niid v«iniilating. It has, more-
over, at the same time to throw out tbu waelo tnatterv niifiin); from tb«
changes inking ))lum in its own substiuice, having lirsl bmuglit l)ief« waste
nsttere into u roniiitinn »iiilnbtc< for bdng thrown out.
§ S. Id the body of tiiiin. niovemcnts, as we shall see, are, broadlj speak-
li)t(. parried nut by iiictiii» of niiiacnlar tii>Bue, and the cbaDfee in niuscular
tidaue which lead to the setting free of ener)ry id ihe form of nioveiuenc are
directed), governed, and adapted lu the surrDiiudiuus of man. by means of
nervous lissuet. Kays of light fall on ilie uorviitis liiibataDce of the eye culled
Ihe retina, and set up iu the retina changes which induce in the optic nerve
other rhaiigeH, wliieli in turn are propagaled t» the bruiu as iifrvnuA iinpulst*,
both the exdtjiliun and the propa^lioD involving an ex|iem)il.iire of enei^.
Tbcac nervous iiupiilaes reaching the brain may iudiici- otlitr nervous im-
pulMi w'hiob, travelling down ciTtuin nerv<« lo cerinin nninclvK, may lead to
change* in thoae rouscW by which they suddenly gmw ^liort and pull upon
tht MDO or other structure* to which they are attached, in which unae we
SDT the man starla ; or the uen'oii* impulsi-» rciu'hing the brain may produce
soniG ntlicr elTbct*. Similarly iMiuud falling on the ciir, or contact uelwecn
tiie Hkin and snme t()reign body, or some change in the nir or other surround*
tngs of the hodv, or Minie change wilhin the body it^rlf may w) nflcct thft
nervous tiswie of the iKxIy that nervomt impiilM-s an Marlci and Inivcl to
tbb point or that, to thv- brnin or cl»cwh<'rc, and evcnlunlly mny cither rcJwh
•ome niii-H-uhir tinuc and ko gira riM to movements, or may reach Oth
ti»ues and )>r<>iJuce some other eflf^
The muscular ti«8U0 then may be conndered as given up to the production
of muveiuenl, and the nervous tifeiio nt given up lo the generation, tranil'or-
mnttim, and propagation of nervous impulses. In each case there is an
expenditure of ener>;y. which in the ease of the muscle, as we ^ball see, leave*
the bodv partly as heat, and partly as work done, but in the case of nervous
tlaaiie is wholly or almost wholly transformed into heal before it leaves the
bodj; and thtH expenditure necessitates a replenishment of tatergy and a
lenewid of .lulistanoe.
§ 9. In onler thnt these master tissues, the nervous and miisoutar tissues,,
may carry oii their important works to the best wlvauiage. they are relieved
of much of the labor that falls upon each physiological unit of the amicba.
Tfaej are not pre^iited with mw food, they are not required to carry out tha
neonniry tranifunuutioDS of their immediate waste matter*. TIk whole uf
the nst of the body is engaged (I) in so preparing the mw fooil. and u
bringing it lo tin* nervous and muscular tissuw that lhe.ie may build it up
into their own solmtamv with the least trouble, and (2) in receiving the
vraste matters whiih arm- tn muscular and ner\-ous tismea, and preparing
them for rapid and easy eji-cliun from the bmly.
Thus to ocrliiin liwrno, nlncb ne may siH-iik of bnindly as "tiwues
digestion," is allotted lh<' duty (>f acting un Ine fiiod and pn-puring it for tha
use of the muscular and m-rvouj tiiuues; and to other tiiuu<'-'>, which wn maf
apeak of as "tinaes of excretion." i" alloiied the duty of clcnriug the bwly
from the waste mattera geiicrnlrtl hy the muscular and ucn'nn:^ liisucs.
^ 10. Thew tisauea are for the mcnt part arrange)] in machine or rarcban>
isms called organs, and the working of ihiawor^ns tnvii|vi:i> movement. The
maveroeniB of then orgnn« arc carried out, lilce the other movements nf the
body, chieflj by means of muscular tiaiuc governed by nervous tissue.
\
I
ag
J
ISTBODCCTTOIT.
B<f><v or may nixkc ■ dwliDclion betireco tlie niuscla wluck ar« coDoerued
in pfTialdciu); ■» elfcci gii the world ouuitlo man's bodj, the muBclce by wbkh
MM il<na nt« imrk io the norld, anil ilw nituotei which nre oonoerDod iu
snrinK <Mt th« raovemeota or the ioteraa) orgaiia. Aiid we ma^ atmilarly
■ako ■ •litUucLinn belweeo tbi- nervous tissue concemeil in citrrying oul the
alerokl work of tbe body aud Ibat coawmed iu re^u'ttinK the moreineuta
■ad, M we sbftll see. the general conduct of the interiiul ur^is. But rheM
t«o clawei uf muscular and nervou* tiMue Lh<>U|;h dUtinct ui work, ami, lu
M shall see, ofleti diflereot in structure, are nut nejinnited or isolated. On
llw CDDtrary, while it is the main duly uf tk« nervous tissue as a whole, the
wrmw system, an we may call il, tu carry oul, by means of nervous impulsus
raiag bitber and thitlier, what may be spoken of as llie work of miin. iirul
this aanw is tlie master linue, it abo serves as n bond of union liiitwiTO
itwlf and th«' muscka doing external work on the one hiind, and the organs
if digaatioD or excretion on tlw olher. mi that (be iictivily anil comlocl nf
iW laurr may b>' iwloquntcly udiipttd In tbi- iieedii of the former.
( U. I.^utly. tbe fiHid jirepirpd bikI i- In bo rated by the digestive urgiuin is
arriad nod preMnlcd to the musouhir imd nervous titsuea m the form of a
tfMnfdax fluid known as blood, whit-h. driviMi by incana of a compIical«d
ikrrhantsn) known a* tlic vascular apleni, circulatea all over ifae body, viitit-
iuf in turn all the tiwuct of the body, nod br a Kpccial nrraiiceioont known
m Iba rwpiralory Riix-hanism. carrying in itM^lf to the sovvnil tusum a supply
•f aiygcn as woll as of food mnro properly so callod.
Tbn motive power of this vascular system is supplied, as in the case of the
figMtiw system, by means of muscular tiwue. the activity of which i* sinii-
bnjr governed by the nervous syslem, and hence the lloa- of blood to this
jtn or that pari is rO|i;iilaleil sccordinK to the n«eds of the pari.
f U. The above alii;bt sketch will perhaps suflice to show not only how
saaieroas but how varied are the problems with wliich physiology bos to
JeaL
la ibr fini pln<« there are what mity be eallol jceiienil problems, suob aa
Uoa tbr f H«l nltrr its preiianiti'iii and idulKirntiou into bloiul is built up
isfi Itic living outwtnuce of the several ItMuc? How the living sulMtiinoe
bnaki down into lh« deail niiotc? Huw the building up and breaking
4nm liithr in tbc diH^ront tissues in such a way that energy i« set free in
dftrvat mode*, the rauscubir tissue contracting, tlie nervous timuc thrilling
•kb a oervotis im|HiUc, tbc secreting ti»iie doing cbeniicnl work, and the
Eke? Til tbaso eeucnl 4U(»tions the an»wcn which we can nt present give
can banlly Im called answors at all.
la the serond place there nre what may be called special problems, sueb
as What are the various steps by which the blood is kept rvplcnishcl with
bod aud oxygen, and kept free from an occumulalioa of ffaat«, and how is
tbr anirity of the digwtive. respiratory, and excretory oigans, whicli eJTeot
Ibta, rv-ifulsled and adapted to the stress of drcuntstances? What aro the
detail* "f <be working of tlw vascnlnr meclitini^ni by which each and every
IIMM b fc««ver baihed with fresh blDo<l. and how i» that working delicutely
adaftad lo all the varied ebanKes of the bodyT And, compared with which
■U oOwr special probleou are iu^ik'nitioaut and preparatory only. How do
mnaut impubes so flit to and (r^ within the nervous svstem aa to issue to
tbe movements which muke uii what we aoraelimes call tVe life of man? It
is la ihtve special jtrobleni.t llmt ne must obieily confine our attenttou, aud
«• may fitly begin with a study of tbc blood.
BOOK r.
BU»n. THK TISSl'ES OF MOVDJKNT. THE VASCUUR MtCIIAMSif.
CHAPTEU I.
BLOOD.
I
k
its. Ttie wvrnl linur* nm Iriivcnni bv inhiuio tubes, iIk c*)iil)arv
l>loedv«a•cl^ u> whicb blood i* brought by 'th« Mlvrirs, bik) frcnn nhicn
blood y Muriwl Kiiiiy by ilw veiiw. Tlww cntiillnrin' f«ni> ncivorka ilw
■wfcr* of itbich. rlirU'rins in rortii nnd riw in ibc diltvn-Hl ltnuo«. arc oceu-
pM by ibr clrmpiiK oi Inc tJMUO irhkh coiweijiJCDlly liv mihidf ih« Mfnl*
Tbr hliHi't flowing through Ihii cniiillnrin coiisut*. under tiorirml com)!-
lioiM, of an uliDoat «o1orle» fluid, tbe filaiwa, in which are carried n tium-
btr «f bodin, th« r*tl, «nd the u-hiu (x/rfnurtf. OuiBtde the vapillary wnlls,
fiKngnpHich ■[incnoa exist beiwceii the capillary valla and Ihecelln or (ibiee
sf Iba tiKiic, or belweeii tbe eleuitnis uf tbe tiMue lh«iiii»elrea, is fomiil ft
«okir)Ma fluid, reAcniblinj; in niaoy m>iN.ft» ilic pliuina of blood and called
tfmpli. TliuM nil (Kb elements uf th« tu*uc and the outfidn of nil iht^ nijiil-
MTW« arc li«tlM><) nilb lyinjib. uhirb, a* wcslwll «ce bcrraftvr. i* continimlly
Aowioir annr from ihc tlvtic alonif i>|Nxial chaiineU lo (MM into lymphatic
VOMela ami ihmi-v iiilo ibe blixid.
As thr bliMKl tliiwR ihmu^b ihr cnpillaripf certain ciinfiitunils of tbe
phunin Mogvlher wilb, al tiium, nhttc ixiriniK-le*. and under t'seeiilinDal
cirruni*lannii red c(ir]>itft(rK-« i pBM through tbp c«|>illnrT nail into the
lynjih, and crrlniu i:on«liiuciilt> of ibr lymph pnM tbruugb the mpillwry
«all into ihr blmwl Miiliiu tbe enpillnry. Thero is ihiie an iitlcrehango uf
Matirrial b<'l«i-cii the Mood wilbiu the mpiltary and the Irniph oiil#idc. A
•imilar inli-rchBii^ of material if at the Ftiine (iine B'^nj; oti betweeo tbe
Unipb nuil tbo tiMUe ilwlf Ueuoe. by nicuno of the Tympli nrting at mid-
atrman, ■ diwihle inierehsMKe uf maleriiil takes place beineen lh« blood
■itbia tl»F nipillary atid llie It^xite Kiitsi'Ie the capillary. In every li»ue,
•1 bioi; as life lasts and the lili«<il flows lhruii|;h the bl«<>dveHeli). a double
(■T*«tn. anw mpid. n«u filowr, is |iat»iii); Irum the blood to tbe Ibeue and
fmu ibe tissue 1» the bliHul. The sireaiii IW>ui the blood to tbe tusue car-
tir* to ibe liMin- ibr nial'rinl nhich the tinue needs for buildini; itf>elf up
amt f'f iIjiUii; it.- work, iiirtiidiii^' the all-imporlani oxygeu. The stream
fr : atue to the btoml rarrin< ii>t<i the hhiod t^riain of the products of
it. ... ......il cbaiigva which havr b(-('ii taking' plaeti in iho luiue, pnxlurta
■bkb mar he simple vtaMe. to be oM otit of the budv as soon as powlble,
•r which may bv budin ca{>nble uf bfitig mn<le txK at by aome oUier tisauc
1/ IH.'i'ili.
A iliii'l Kifiitiii, ilini IV'tiu lli'T lyififili lyiii;; in tli>r rhinks and craoQiee of
III' li> 1 iil'iHtr ll'i' Iv'l'l' <'liFinrii']4 t<> tin: lur(;t;r l_vm[>b vessels, carried
Mwiiy li'iiii llii' lii'iii'T >^<ii')i |iiirlK 'if ill'! iiiiiUiriiil coining from the blood as
till iikJ iiiti'ii iiji liy till' lir.'ii'' iiri^ir iiri'l Midi |>iirti^ of the material coming
tiitiii ill' lirn'ii iiR il'i ji'il liiid llii-ir U'li} itil" tin: hloiMlveflsel.
Ill tiiii'-i I iniiii'", iin ill iiiii-i'ji- fur inntiuiftf-, the capillarv' network is bo
I liiwi ml iiii'l ilii' iiiiini'iiliir liltrii li'-M HO nimr to the bloodvessel that the
lyiii|ili liiliv'i'ii til'' lwoc\i>ilN onlv iiM II viTy thin nheet; but in some tis-
BiiKM, (in III iiiiiilii)t<', ill'' lilci'irlv'iv'-lN lir nil ihu oiitttide of a large mass of
tlMii". Ihi' IiiIi'H'Ijiiiik" Im'Iwi'ii the ri>iilritl jiiirlH of which and tne nearest
i<ti|'lll'ii V I'l Ivcnni'l in I'lii-rii'il 'in throU|;h a liin^ stretch of lymph paBsages.
Mill III I'li'li I'lini' lliii iiriiii'i)il>' in tlii' niiik-; the tissue, by the help of lymph,
^iii 1 "11 Ihi l'lii\«l : iiii'l nlii'ii ill Hiiirifiliiiir pu^eii wc speak of changes oe-
liiriiii llii> liliiiiil mill till' liHNiii'H, il will be iiiidi'rstoud, whether expressly
al'ili'd ml 111 ii>i[, tliiii llii< i>hiiii;;ii< iiri' cUt'cli-'l by moans iif the lymph. The
bli'iiil iiiii\ tliiii Ih' ri'fviiiti'il iiH 1(11 iiilrni-if iii-'iliiini, benrtng the same rela-
Mmin 111 lliii r.<iiiiili)i'iit lix.iiK'* llmt the cxlcrniil miHtium. the world, does to
(lii> »liii|i> nutitiiliiiil. .liiM 11^ l1u> whole nrj^tiiiiein lives on the things
iiixiiiiil ii, 111 lilt mill \i^ I'lHul. HO ihi' si'ViTiil iiiu>iii>it livo on the complex fluid
lit uliwli ilii't iiiv nil liiiihi'il tiiiil whii'h is Co ihcm their imme«liate air and
( I
\n ili<< ii»uiv> t ikt' M)> i-wp'ii iVi'iii ibi' bWd and fi\t' up carbonic acid
III ili>' l>li-.',l, biU iu<< ii1».t>it III ibi- s.-uiii' r.iic or At the $ame liiue. More-
I'v.t t!i.' v'M'i^ii iiwii.s l.iki' i!|' iV-'iii ilio li!.hi.i and j;ive up to the bloid
vi/i.i .1 ilvivir, tilings .'v liu- 'ii:!!!- tbitip^ a: diilVwiit nttw or at difierent
ii.'.'i-
■.\w ^•■■•c ::A\:<i. :h:i: ti'.o ivuuv«ii:i->ii and ohsrac-
'"••.\Atr i.irx;;-; iv, .lir^Vrv:;; -.•arts ol tiie l»iy and
:■.• .-I'u- ■:'.:<." i. :.-V. :\:c :.::.:ii ai,i! a >f all the
-'■, s", •; ■■.;*■.•..,■(:■■. a:: .iv:-i_ir ■-■^::'.r'u i.''ai>.'eitii>o
V ■-■ ».vv il s''.:*:'.^,> «!-. ■..">. b'. ■•>i s tn.iwn t>i
■ .; ■ '-.■ «,-\trr*'. ;;**:;>»■-".'. xj; "x- waI: with
1 .(■■■. \;-.i:- ,■■■.;■ -v.Ut : :.t .-;i*i:i«li:cL At
. ^ ■. .■,■ ■:.:\-.-. ■'iJi'.:'>.-i <: ;!i a-^ rn»eat«il
\.iv ■■..■ *>„',;;■ .i i-:j. ,.-r-:i : - ;i-; i.amya
-.^ • 1.1-. . "■; ".■ K K-.t 7^ -.:' Z--C rj)i ai-.-K im-
V
■,\**" ; ■ "H ',
*'••* .'X**. **:*
u* ■•
.'^i " ». *v
*.M •.'
■. ■ V", i""
\^ ,A"'l *■■^
■ V«
, ■» ? ' , » ^ , , ,'
•-{ ;/ o> i>
■■
■ -" ** ■ .* ,"
' tv* ,' ■ \-
» ■ ■
._■■ i»
' 'ilV*l,'^
i ',* ■ !• . 1 I. ■■ ■^■■- :.■..■" ;>!»;■» i' t 'i^ I'd" » wrwrtLT
' ■ 1 . ^' , ^ -s ■.■ • x". ■.■!*. -Mi;;." -T-iia --.-iigm w
"•1 T^ - :-»-■ ■■: . . ••: \ ;■ . ■ 'l.a* l' ll'^i-I lOtUr
'-* ■ • -■ '":' .!«* : ■ i::i,'j r i-j» it—a sded
--'_■■. « ■ :■ ,'■ : ^Hlic *J.'iiL I^bi
< .' ■ -•..:■. 1,; : ; -ii-*ri;l" suUtKi
■ ■«. . - . ..^ - .^^j. _ \ ^ i_' ^
_ ,1 ,-.:>? «s*-. 1 -fci* iriD*
- — - - ~.-- - ; ;:v -i.:-^*: -t rhe
- , ^ _~ ■■"...- "? i>TfcJ- i T-uin
- . -- . .:, -sa liT a-fr^
-->:.■- -.-Tj. 1 ■•■■iC
TUB CLOTTIXO or BLOOD.
43
«r ArttMam<n/M«i, flonting in n |i«r(ectU fluli] xrruui. [Pig. 3.] Th« thriiikiug
•ad owMkoMtiou of tbe clot, luid lh« (virrtvpoiidiiii; iRoraaM of thr wTiim,
PMtipM fiir aune tiia& TIm- iiivikt Mirfiui- of ihi- clot b g«D«rnll}- Hli^htly
[TM. 1
;»!■. a,
>t dT (viwnllir tomoUM Uaid,
Unwl nt ■BnipiliiU'l tiluil. arm laotrv
liii<rn.tbawtii(tlwclolccinln«(ilui<ldoM>
liiK In tlH) ADid (MVui. Al)*t I'lLiulcl
Me<«va. A porUoD of lh« clot vxamincd under t)i« laionwcopo ia MW (o
oMMiK of « n-Uwork of 6n« gniiiulnr fibrils, iu lti« niMhw of whicb an
mtaafUd tlM' ml aii<l wliiie ovrpiui'lM of the bloo<l. la the mnim iiothipg
(Ml be ae«n but a few stntf curpui-
tim. chiefly white. Tbe MbriU «ro l>^" *
cnrapoaed of a hi balance nllecl
/kritk. (Fm. ■!.] Ileiicc we m»j
•{M«k of ihe ulot u coDakling of
Abrin and corpusclei; an) ibe act
of duUine Is obvioaily a Buwtitu-
tm Ibr tno planna of AbriD ami
arav. ftillonnj by a acpaimtioo of
tbi fibrin aiul «>T|iuaclei ffura the
OnfUtiMd aiola. *lM<rti>t Ik nbrltteul mq-
dMIOB. AAm Dauok-J
^> Ib nan, blow] when *b«d bocninat
Tiarid in about tww or ihn-o iiiinutut,
«mI «Dlrn tbp jvlly tin^} in nbiitit
irvur ten minutes. AOi^r tlie lapM
■ of anaiher few tninuton the lint
ditipi itf BcniBi areeeeD.nnd dolling
ii gUimally cotunlelc in fmin o4ie to
ivmal boun. The (ime, however.
■ win Iw found lo vary acoarding lo
(ireuBMHoceo. AmoD^ anitnali tbe
npidity uf duttiug vanm excee^ii);;ly in iliHerenl speeixM. The blood of tbe
bom clut4 with reiDsrkable alowoeMt; at) bIdwIv. indwd. that many of the
rod and alw loDie of the white corpuscles (botli tbeae being Bpecifi<'ally
bearirr than the plasma) bare litue to Mnk before viscidity aets iu. In con*
sa^MHim ihara ainirmii nn tlwsur&oe of the blond an up|>er layer of oolorieas
fhfift. oMOt^nlng In its dwper panions many c»ii>rl«s9 corpiuclat (which
an lighter ihnn the r*il>. 'lliis Uycr rlntu liki- t)ic ntlwr |«na iif the hlood,
fcrolnff th« »i>'Ciill<vl " butTv I'nai." A ainiiliir butTy cunt i* Mimfitiiucs Mvn
ta thr bl'--l of Minn, !n o-riuin atmurmni rnuilitiom ot tli« body.
If a [K-ttioii of txirwV hl'Mx) lie mimmndcd by a cooling miilur* of Ec*
aad mil, and thiu krpt nt ntmut il" C, clotting may benlmoot indefiniloly
puMponoJ. Under ih'we cin-urasluoeo* n iu»rt' oompltM (lesccut of the cor-
[WIN 111 takue pliu-c. imd a oontfalraabK- iiuiinlity of oolorlus traniparvat
froe from blood oorjnwpka may be ohlulnetL A |mftion uf this
BI.OOP.
piiumn rctDOved fmrn the ftvninn uiUtiire dnU tii tbc *IUM manDrr u (liit«
the vnlirv Ulo'nl. It firal bcomiat vim-id nnil i1i«d forma n jelly, which sub-
■•qiiOtiUy M-fiaintfs into n oolorlcM akruukeit clot oui) acriini. Tliicahow*
thnt iIm) niriiijuclet nrc not an caeniial pitn nr the c1a|.
It a few cubic rpntimrircf of chi« oolorltw plHtiina, or ofa niniiUr plMma
which tnnr Im' ohuiiiivd fnim nlmwt nay blixxl hy menm n'bich we will
pmcntly 'JM^TiliO. I>c ililuU-il with m«iij- liin« il« bulk of n <t.W per wni.
tnluli'Mi of Hixliuin chloride' clotliiig i» much rcUinliyl, iiiiil iho vnHoiig stages
Rutv Im) infirv Muily wMcbwl. A» the Itiiiil i* bivomin^ ricci'l, lino fibrils
of Alirin will b« Men to bo dovel»|>t.-il in il, cs|*cctully nt the hJc« of cbe con-
lniniii(,' vM*al. As tiioM fibril* multiply in iiutnltiT. the tloiil becomes more
nn<l tii'ir>'i>f (he i^nsistcncfof u jelly nnd st the Mime time somewhat opaque.
Klirntl or pulh^ nbuut uitli n needle, the libril§ shrink up into a arnall
o|iiii|iie Htriogjr iDlin; and n very eoDHideruble bulk of the jelly may by
n};itikliuii b« reeolved into n mimite frnj^menl of ■lirunkeii fibrin ilotilin); in h
(((lantiiy of what t§ really diluted serum. If a Bf>ecimen of such diluted
plaamn ho stirred I'runi time to time, as soon ua clotting begins, with a needle
or gliUM rod, the nhriii may be removed piecemeal as it fornix, and the jelly
•ta^- tiiay he ult'i^uther done away with. When fresh blood which has not
yi-t hnd time |i> clot U xlirrwl ur whi|i|>eil with a bundle of rods (or anything
proMUlinj; n InrKu umotini ot ruut;li aurfaeet, no jellydike clotting take*
plaoot Itiit till' mdit liceomo covered with a mnsa of tikrunketi fibrin. Blood
thiia whip|>eii until lihrin ccniHW to be deposited, is found to have entirely
\iM l\M (lowcr >)f cl'itling.
I'utling tlie»e fuels logollier. it i* very clear tlini th« phciiomnia of
the clotlini; of blood arc cnutcd by the apjieurance in the pliwmn of fine
flbrila of fibrin. S<i hm^ a" thaw ar« «ciuity, the bUicKl i* ximply vixoid.
Wh«U they ticcomc siiHicieiilly numerous, they eiv<- the blood the firmness
of a j'<Uy. Soon nller ll)eir rurmnli<ii] they Dcgin to shrink, and while
shrinkiti); endow in llwir um»Imw the corpu«cle«. but squeeze out the fluid
parts of ih» bItHK). llenc« the appearance of the shrunken colored clot and
tlw cidorli<eB scrum.
5 13. Kilirin, whether «I>tuit>ed hy whipping freshly sbed blood, or by
wiuihiiig cilhcr a uormni clot, or a clot obtained from oulorleM piaBtna.
exhibits the Mme ti^-ncral cltaractera. It beloajn to that olasi nf complex
utMiable niirDgvDous btHltrv railed protrid*. wbien form a tarjce portioD of all
liviiit; iNMlins bimI an uwntiid jiaH of all living stnidurea.
Our kmiwlnlge of pn>teida v- at pnwent too inii>erfecf, and pmbahly nam
of thvni have yet Ix'cn |>n-]Mml iu mUijuate purity to justify in in nltemptiiig
to Mvun '■> lliem any drtinilc fnrmuU ; but it is imptirtani to renieinbet ibeir
gtatnu ci>ii>l>>isiti'>n. UK) |iart> of a pn>tetd enmiaJu miher roon- than &0
pans 1^ cnrlxia, rather ntun than IS of ntuugco, about 7 of hydragvo, and
rather uorv than M of oxygen : that is to ny. tber eootain about £Uf their
wdsrht of carboii, and uoir about i iMr w^'glit t^ nitmgai : aul vet, m wc
■ImIi m«, tbav are emioenuy lb* iutn«nious HibMAucnt of ibe b»dy. They
wwllycaM^ aHHtllqitaalirf (I orz per cvnL) of sulphur, and manvalso
h«** mmt pkuiplMraa attariwn lo Ibaaa m sobm way or other. When norot
Umv Wra • wiaUe «|aaaUiy of aah, mtMStine of inorganic mite of whicb
iWrh— MS cMeir aoJiwn and poOHmtn. and tne ncMe chiefly hydrochloric.
ilwy all i^iwcMtain f«ni.liWM. by whicb their prMence may be neo^ited ;
of tbtw iW mtm tbniafirrirt ai« tfae folloains : Bnikil with aitnc and
ikay gin a yalhw eUar, «fckb iiw|gMa iaiw ««<uige upon the ndditiaa of
I
I
I
TUB CI.OTTIXG OF BLOOl).
4A
■awooim. Tbil k cklted ihr xanthitproiiiie t«st ; the color h due lo a product
of iMunipMJUOn. Bt^leil witli llto mixture of lOPKuric anil iiiercurous
Dimtra kiiowB M Millim't r'-aynt, Ihcv |pvc n pink wlor. Mixed nilli n
alroaf; •olmii-n of ^ndii-bviinUi- tln-vgivp. cm ihr nddiliun of a dnn>»r two (\f
m wvy ««ik s-dutiuii of cuprii' Hiil|itial(% n viokt or pink color uliiuii def|M'DK
oaMUiag. Tbcw nn- aniliml rt-acliunK, nut ttir'>uiti): tuui:)i if aor liubt
anlbtCUOMilMlioDuf proU'itU; bitl llit-y nrr iieclul as praclii-al lr«u ctinblt'ng
oa U> tiMcct llii-ir pmejica.
TIn Mvcnl mrmbon tif the pruidd group iiro at prmeiit diRtingtibbod
Ai^ tadi tUfaoT chit'tl^ \ty thdr rn>{wctive aolubilities, ee[»eciajly in viiriuiti
■tUaa atlutinna. Fibnn i» one n{ the l«uM enliible ; it is iotoluble in wKlvr,
■JiwMt insidiiblr in dilute itciilnt) Hti1in« solittione, aud rerv npuringlir wuIiiUb
IB nmns coon-nlrutw) neiitnil HiliDc tototioiis niid in dilule acide and tlkAlwi.
In (tniDK acids aud alkaliM it diasolrea, but in ibe procNa beooiDCi mn-
plaUljr cnaogcd into Bomelhio;; which is nu luiiKer fibrin. In dilute iicMf it
•■■Ik up and l)«<<(>ni«B trains pArcnt, but nben tbe acid ia neuLratized rMunia
Iji Its pivTioufl cunditiuii. Wlxrn suBpendc<l in water and liciited lo tOO* C,
or crm to To" C, it beconwa ch«i)|j[Ml, and Hiill leut Mduble ihtui tx-foro ; it it
M*d JD tliit tmm> la be eoagnlatnl l>y tUe beat, aiid, ha we «ball mc, nearly nil
|MH«Ua bav« the prujierty '>f beiiiji chaujii-d iu iiatuiv, of uodorsoiDg
OMjCtiUtion and ao Itpcimiiig leM aoluble than bcforn, hy being cxpoaea lo a
Mrtain hi|;b tt^>ii|H'r»tiire.
f'itirin, tbfii, ia a protvid dUtingublietl from oilier proteidii by itK cmallcr
wlubility; it i* fiirlber dialiufcunbed by itc iM>ctiliar iilainiMitod* stnict-
oiv, thv c4bL-r jiroleida when oliliiinefl in a •olid foroi appearing either in
aotorvbou* granulr* or, at miMl, in rUcid masKa.
I If. W« may now n-lum m the tmim.
Tbia b perfectly fldid, nnd rcmniRi duid until it deconinoeee. It ia of a
ftint ttmnculor, dueto the pirsencoof a special pigment subatUDoe, diflering
frun lh«' red niiilter wbicb Rives red»e» to the red corpusclea.
Tmed by iIh- xanthoproteic am] other lots it obvioUBlr ountitinii n Urge
mtantiij of proteid uatler, ami upon exnniiiuition ire flDa llial at Imat two
iliaiioei prulind Mibatanoea arc preaeot in it.
If rry*tnb of magnesium sulphate !>« added tn Aenim and gv^ntly atirrt^
uaiil ihny divanlvc, Jl will be iteen that thi.- »eruni att it appn«cht» uitumiion
tritb tbv Mlt lieconea turbid itutead uf renniiiiiu),' clear, n»d «veiiltialiy a
vhtir amor^dtiiiM frnuiular nr flooculeni pn-4-i|iiIit|i-- uiakM Jt.i ap|M-iiriincr.
Thi> pn.-c>pttatt' tntiy Ix- MiKiraifxl by dvcautaliun Dr BUratiou, waabud witli
iaiamtnt MiliitiouH nl' mtiv'i !•''■■ ■"■i lulpbate, in nhicli it iit inaolubic, until it
ia frwl fniui idt 'ilbcr I'lnntituciitit of lh« leruiD, and ibufl obtaimil Ibirly
INirr. It ia llwn fmjtid ui Iw a proteid IhhIy, dittiuguiabiil by ihv liiljowiiig
riiaractvni iimoiig oibi-n:
1, It in (kIii-h I'rtM-d fnim iiny iidherciit nuigoij^luni Milphalc) iutnlubh) in
■lialill**! waiir; it i* inxiliiliU^ in onft-niriiHnj xilutlona of iwutraj Balinc
bodin. Mit'h a* mngnniiitn ■ulphnt'^, Mxlium i'hluride,Me.,bul rMdily aolublo
to dilute ('. y,, 1 |>er M'Ot.i M>lution« of ihc naine n4>uiral Mdiuc boiliee.
Iliftrr fniiD it* »oluli(Hi» iu the latter il mav l>e prrcipit«tv<l vitbfr by adding
tnnre aeiitral Mline sulwtnum ur by removing bv dialyaia tlM muall ijuantity
„f .,i;.,.. i^jljatBtir-e prexent. Who'll i)t»laiiMd in a prccipitaWd Jorrii, and
* : in (liidilltNl water, it readily dixwilve* into a clear solution upon
ibrBo.iiMtiiirasiDall •i^tinntity of soiiK'iK'Ulral saline bixiy. Hy t bene various
^utiond and jirr4.'ipiljilKi»s it is not n-aliy ehaiigod in nature.
'i. Il rrndily ilim-ilves in very dilute acide {r.i/ .'m h^ilrwlilorie aetd neo
wbefi (liltneil to far Ivm than I jier cent.), and il ■« •innliirly Milubic in diluio
aikaliQ*, but ill Iwing ibas dissolved it is irbolly cbangv<) iii nature, and the
46
B 1.000.
B<>liilii>ns of h in diltile acitl nnd dilute sHulics eiv* rcai-ttons quite dtdoi ___
frDiti i.l)u§G •>{ tho suliitioii of llie siibetance in dilute npuiral snliue wdiitiooi.--
Ity tlic acid it is ooiivcrled inLo nliat is c&llod (ici(/-u//i((riiin, liy ll>e alkali
iuto alkiili-'tlbumin. butik of wliicb bodiei ve shall hiive to Hludy later on.
'i. Wben it b suspended in wnter and healed ii becomca altered in cliar-
acter, mai/ulatrJ. aud nil its reuctions are chaujred. It ia no loDger aolulile
in dilute iiviitiid aaliue solutioiis, nut even in dilute acida and idkulie*; it
haa bceoQit; coatittfaltd protxid, and in now even less eoluble than fn-fii fibrin.
When a loluiiou of it in dilute neutral aalioc eoluiiou id Nimilarly ItcBtcd. a
•iinilar cbonge lakea pliieo, n [ireeipiiale falla down which mi cxaminiiUon
if fi)un<l to hv cungulnlL-d pmteiO. Tile iciiijionitiirc at which l)ii« changD
tnkts |>lac« ia sumewberu nboiit T-t~ C, though nhilling itlighcl)' acoMrding to
the (]uantity i)f nnliiie Kubttnncv prment in Uic Koltitinn.
Tlicntirive |.bn.-i.' ruactiunii arn given by n nuinlipruf protctd b<:)dies foninn^
m griiup cullfd alohulint, tind the [lurliciiliir gtohulii) pra«cnt in blood-eenim
is vuUvd jiaraijlohnliu.
One i>r Ihc pniii'id* prenent in blood m runt in then pariiglnbulin, charac-
lerixeil by its »i)liibility in dilute neutral Mlinv x>Iution«, ii» in»oIubiliiy in
dietilird wnter nnd co u <.■<.■ n I ratol wiline colutiouF, its n-iulv Mtlubility. and at
ibe Himc time convcreiun into other bodies, in dilute acida and alkalictt. and
ID iia becutuing convertml into coagidatcd proteid, and «o beut]; precipitated
fVom its GoKitiun* at 7<>° C-
The BDifliint of it present iti blood-serum varies in ritrious animalB, am)
apparently in the same animal at diflerent timts. In 1U[> ports by weight of
serum there are penerally present about 8 or 9 parts of proields altogether,
and of these some 3 or 4, more or less, may be taken as para|;)obulin.
§ 17. If the serum from which the parajilobutin has been pivcipitated t>^
the addition of neutral .lult, and removed by tiltrstiun, be subjected to dia-
lysi*. the valt added may b<! remiivi-d, and a clear, Mmewhal dilute*) mtnim
free from iiaraglobtiliii may be obtivinetl.
This still gives abundant pniteid rvuicliouH, ko that the xerum Mill coataioa
a proteid. nr wuiie proteidv *lill more mdiiblc than the globulins, since they
will remain in sidultou, nnd nrc n-'l pn-cipitatrd, even when dialyia is con-
tiinicd until thi- nriim is pructitnlly frx-rd from both the nuutnd mIi added
to it lutd the dilTuaiblc hiIis previously present in ibe natural fenini.
When this scrum is hcnleil to T')" L'. a precipitate mokes iU appearanco;
the prolcids still present arc caa(;iitalcd at this temperature.
\\ c have some reasons for thinking that more thnn one prnteid is present,
but ihey are all clowly allied to each other, and wc may for the prveciil
speak of them as if they ueie one, and call the proteid loft in scrum, afler
removal of the paraglobuliii. by the name of alhunn'n, or, to diatingiiish it
from other albumins found elsewhere, rmnn-aibuinin. Serum -a I bum in is
diiJtlnguifihed by being more soluble than the globulins, since it is soluble tn
distilled water, even in the absence of all neutral salts. Like the globulins,
though with much lets eoue, it ia converted by dilute acids and dilute alkalies
iuluacid-or into atkati-albumin. The percentage amount of senim-albumin
ill xtrum may be put donn as 4 or o, more or less, but it varien and some-
limcd io l<Mi abundant ihaiipaiaglobulin. luBomeanimala (suak<»)it is said
to dirapjit-iir during Marvation.
TIh' more imp'irinnt chiiraclcnt of the throe pmteids wbioh we have jiial
ttitdicd may l>c atati^d a^ lolluu'ii:
BoIiiMb til <li>tlllf4 wklor bdiI In mIidc t-iluli<inf of nil ^IreuElli* - nenin-aUiimi*
InKiliiMr in ilidiUt^t walxr, nodll/ riliiUto inililiil* (alin^ KtlnUcin*,
Intvtiitl* lu omcaiiiraidl •iillno mliiUoDa paragbtMim.
tnioliiMa ID illitlllnl wjilcr. Ii-ir^llr iviuMa at sll id ililiiU aUfno
KiluUon*. an4 vvr; tliUatuIuUsIn nraiv (OdivntraiMl mIIuohiIu'
tko* .iftria.
I
I
I
I
t
1
I
I
rnS L-LOTTtA'O OP BLOOU.
47
I
Bndiln (inniglobiilln slid ecru lu-al bum iit, wruin couiniiM ii'verjr luge
n ■!■!>■ f nf subMuDCM, f^enernlly iii mduII <iiiiiut!(r, wliicli. miico tki^ )iiit«
to b* ntnMlt«) hy b|«cibI n>vth(><ls arc i-ailetl txtracliK": of lli«#c tti'mc U«
Mtr<i1t*onu>, anioe noD-aiirogeooti*. .Serum vi>ntaiii> boridca i»)|Mttaiit iiior-
(snir HtitK* mlwtaDcn: but lo th<9« ire duitl n-lunt.
t IS. With the knuwlttl^L' wbifh we liavegaiwil ofllH- iiroirtdn ofcIiKltiJ
bloi^ wr mav e*) t>a<.'k to Uw 'iix-MlioH : Clotting Wing (liii* In (he ai>iirnr<
aiK« in bliMxl |iln--iiiii «t' a proteiil MibslADCC, liliriii, vthU-li iitvvuiii^Iv din not
tairt in it aa w-h. w hm an- iliv miisM whldi Inl f > l)iv ni>|>Mirniirc of Hbrin t
W« Irani MiniHtiiiif; bv ndidying llii> iui«t ini|iiirrniil irxUToal ciiviim-
Maneat which aircci ibi.- rapidity niili which Uw blutxl ul' lli« sain« indirtiluat
dstt mhto thcd. Thm! arc m Ibllowa:
A uiDpt>ntmvnr'10° C xhidi b aboul or fllgbtly «bo\-f> tho tempemlure
of lb* Uoorf »f KHrni-bliMdcd anrnwl^. la |>i.Tlin[« the moM favorable lo clot-
tiag. A Airllwr n*t> of a few dterMs i* ii^i|<arcntl}r alio bcii«6cial, or at
toM sol Injiirimii ; btlt upon a eliirriirthcr tw lh« rHc«t chancn, and when
bleed ii npidly heated to M^ C do cluUing nl all mav Ukc gtlBce. At this
iMBpoMurt evrtain pmMidsof tbv IiIwhI an oungufntcd and prMJpiiaied
h^on elcUioc i-aii laao iJacr-, and With this cbaage tbc power of the blo«xl
to clot is whMly hwt. ir, hoWL'Tcr, the hcaliDg be not very rapid, tho blood
naijr clot bcforv this changD hns time to cone on. When ihe (emperatore
ioalcad uf being miM^I is lowered below i.0' C tliir cloltinj' l)e<x>Ttiee d«layeil
aail pmlongeil ; and at the lenipenilure of 0° or 1" C. the Mood will r«maiji
flaid, and yvt capable of clollin^^ whi^o willidrawn from llic adverec drcuin-
■cuwca, for • rtrt lonf*. it niit;ht alaraat be aaid, for an indefinite lime.
A (cnall quantity of hhMid ihcd into a small vewel clote sooner than a large
■ onsMity •htil into a larger <>»e ; and in general the greater tbc amount of
Ml ail n furrncc with which the blood conxw in I'vniait tlw more rapid ihc
dnUtBg. y>'htu vhrd blood i* Mlrrcd or "u'hip[N.-d" the fibrin niakca its
■ppMrance mh-imt than when the blood n Icll l» clot in tbo ordinary way ;
■t> that hi-iT, tiKi, ihtt acccK'niling iiitluvncv of contact with furei^ bodiea
Bakes ilwlf ft'lt. Similarly, niuvi-nu-iit of fhcil blo<Ml hnaloDii clodmg, ritice
il iscrraHK the amount of contact with foreign bodioL So aleo iho addition
■ wf MoagT (ilatinuni «r of powdered charcoal, or of other inert powderv, to
lardilf riulline liloml, will by inflncnoe of ftiirfaco. basleu dotting. Con>
vmtfV, bloiwl brought iut'> d^nlad with pure oil docs not clot to rapidly ns
la coiiiact Mill) gliiw or metal; n»d blood will continue to tluw for a
_ time without clotting throiigb n lube »ineared innide with oil than
ihlOTgb a tube not to nmenrcd. The inlliienoc of the oil in Buch cnaee is a
|itiy«irml not a chemical one ; any pure neutral inert oil will do. Ae far as
wa ktiuw thp(« inlluencea afleet "oly the rapidity uith which the clutling
take* place— that if, the rapidity with which the fibrin niakee its appear-
ancv, iMit the amount of clot, not tlie (junnlily nf fibrin formed, though
when clotting it very much retarded by cold changes may ensue whereby
tbe aoMNint of dotting which eventually takes phicc is indirectly aflected.
Iftn etpoaure l<> air eserls apparently little influence on Ihe process of
deuisf. Blood mllecled direct from n hloudveairl over mercury so as
wboUy to exclude the air, clots, in a general way, as rewlily a« blood freely
fX|NMd to thv air. It b only when blood ii> much laden with carbonic acio.
lb* prewDce of which it antagonistic to clotting, that excluHon of air, by
UlMlefio|t the escape of ll»e exetw of carbonic ncid, delays clotting.
TImm ncta icaui us that 6brin ihiee not, an was once thixight, make its
■PpMrinn» In alitd bloixl because ihe blood when shed oeosea tu share i» the
iiimiineHl of tbc circulation, or because the bl(K>l is cooled on leaving the
warn bodr.or because the blood is then nioro freely cs]Msrd to the air; they
Li
48
BLOOD.
further ftigEHl |fa« view tliiil ihv Hbrinbtlie result of some cbemical _
the cxHiviMvioa ioto libriii of mimL'thiiitc tvliiob U not tibriii, the cbuige like
Otli«r clicmica] cbangei Iwiiig nicwt uclivt; at iiii ofilimum tempenitlire, and
like fo ninnv otber chcfnicnl clinii^ro, Iwing nsHniinl by ihi' iiiHu€Deea cocflrtcd
by the [imwiiM of iiwrt bwiiw.
And nc have direct I'xpcrinivntul evidence thnl pliutitm does contain aa
SQle4.-eddiit nt' tibrin whidi, by rlit'niicnl change, it convvrlMl itito fibriu. fl
§ 19. If bloud bo rvcoivcd ditvL't rrom ibe blood vrsM-ls into oue-ttiird il^
bulk of a aaturaced soltilion of toniv tit'iiCrnl iHilt Fiich m iiiae>"'iuni :iu]|>bnti.>.
and the two gently but thoniiighly niised. clotting, <ii|iuriiUlv iil u mixk-r-
iiU^ly low tcni]>er«tiire, will be deferred for ii very long limo. tf tbs tnixtuxd
lie nllowed to stand, the corpuHi-lea will bink, iind ii uoWlov nlanni will 1^1
obtained siinitar t«the {ilaaina ^tiined froni horse':' blood by mid, cxcvpl thnt
it contains an excess of ibc neutral salt. The presence of thv neutral «alt
luu actod in the same direction as cold ; it bus prevented the occurrence of
clotting. It liaa noidestroyed the fibrin: for if some of the plasmu Iw diluted
with from liv« to ten timea its bulk of water, it will clot speedily in quite a
normal fashiou. with the production of ijuite oonua) librin. h
Tlic separation n( the fluiJ pluma from the corpuicle* and from other bodE^I
heavier lann th<> pluimn U much fiurilitiilpd by thr u»c of the cpnlrifagnl machine.
Thia cuiisintfl nLionUnlly afu lirelcm wheel with suvernl ipvkes. placed in a hori-
ziiQlal podUou and tiiude to rdvulvewltli great velocity ilWx* reToIuttons per
minute for iottaDca) around it« axis. Tubc« of metal or very Strang gloM arc
Hut|>eiided at the ends of tlie >|iokni by carefully at^ustod joint*. Aji the wheel
roinies with increaung relciiiity, eoi-li tube gradually Maumoi a horitoolal poal-
licm. bothim outward, without iipilllng any of Its conienla. Ai the rairfd TOtatioD
cofitiuuH the corpiixulcti and heavier particles are driven to the bo4tofnof th«
lube, and il n very rapid movooiont be continued for a long time will form a com-
pact cuke nl the bottom of tlic tube. When the rointion in stopped the lub«a
gradually return li> llieir upright poaltioo njrnin wilhnut anything beins npilt.
an^
the cl«ar pluania in cocb tube can then be decanted oil'.
If some of the colorleaa tranaparcni ptasmn, obtained either by ibe action
of noulrul sall« from any blood, or by llie help of cold flrooi horee'e blood, he
treattnl niib some solid ntiiirul sail, siicb an Imodium chloride, to saturation, a
white ilaky, soruewhai eiiL'ky pii?cipiiate will make itn ap|>earaiice:. If thia
precipitate be removed, the fluid no longer pcaaewct the p(iwi:!r of clotting
(or very oliKbtly so), even tbougb the neutral salt pr«wciit bi^ removed b^
ilinlyaU, nr iu influence leeaeueil by dilution. Wilh the removal of tlte suH|
niaiice prcnpiialed, the plaima ban lo!<t itsi power nf clotting. ^^
If the ]rreci])itate itwif, afl<T beiiij; washed with a mturati'd Molution of
the neutnd salt i,in whlc-li it i.i insoluble) to a* tn get rid of at! wrum and
other eonslituenld of the pbiyma. be Ireut^tl with n smnll ijuiintity of waMr,
it readily dtMolvea,' and thi; foliitinn riipidly filtered givc« a cicnr, colorleas
liltraiv, whidi i« at ftr*t jM-rfrclly tinid. i^on, however, the flinditv givea
way to vi«'idily, and lhi» in turn In n jelly condition, and linally thcjclly
Ghriukv into u dot fltMttn^ in ii dcnr Huid ; in other wonls, the filtrate clot*
like plasma. ThiiN tbon- is iirtvcnl in cooled plasma, and in pliismn kept
JVoia clotting by the pn^-noe of neutral salu. it somethiiiif pn^ci pi table by
mtnntion with uculnd ^nlUi — a Foni<'ihing which, aiuee il is soluble in very
dilute Mlini^soliuinns. cannot he tibrin itself, but which in solution speediljr
K'ves Hm to thti appearance of tibrin. To this subnauc« iia discovered
enb, gave the name of plaimiue. ^
> Tbt iiibManM luctr ti Dot tolnUe in eliUIked wtivi, i>ut ■ umiitliy ol thv nnilml mII* alinui
(Man to tbv ivMirliaw, •ii'l <Hi» Ilia a-bllUon ot w>M rlnuatfy gtm rtt lo dilute lalltM aduilLin,
fa oalcli Uw •utaiauw ■• rniOll]' wlubli:.
THK CLOTTISO OF Dl.OOt).
-49
I
TIm MihaUincc Ihii* nrvciptlmlcil U not how«rer n Kn^h; )>lh1v. bul n mix-
tan of M \auA tito botliea. Ifsudium clilondo b« i-xn'tulU- luliW lu pljuma
h> mtt •XMOl of kImuI n |i«r cvni. n wliiie fliiky vi*ci<l pmciplntv ix ilirunra
Aoma Xfrx much like iiliuniini:. If alU-r ihc rcmo\-»I ol'thv Rm |(r«(-i(iiutl«
•tijiuai chloridr, Miil <'«(ir>:inlly if iiiAgtmium sulphsto bo mlilrt). a
' precipitate u tlirowii duwii. Iu<« viitckl nnd inure ffruiiil&r tliAii the
int.
Tha aMond pnvipiUiii; whcii <'xam!n<?l i« found ut bd identical vilh Uto
fmrmgiahmiut. cMigiilnliiig at 7'}^ C. which we have nircadj aeen to Im »
«fMMUl4Mllt uf WrilRI.
Tba Ani pret-i|>iuil« ia also a proteid balongioi; to lli? (clobullQ ([roup, but
ftn lV<>iii |>;vrit;;lubuliR, not otilr in beinj: rnure r«n'lil>' prccipttutud by
waUrn chloride, and in b<-in^ wh«n procipilaied inure vineid, but ubto in
(4iMr rspww. and eepeoiallr in being coagulated at a far lowor tvinpornturv
tkao |MtnicI»bia, vix., at S(i° C, Now, nhile i^ulatvd pamglubia canuol by
may maaM known to ua be converted into 6brin. and aa ti« procnoc in the
MMxUted plawnine doea not aeem to be OMeaiiu) to the formation of 6brio
oat of itlaamiDe, tbe preaenoe iu plaamine of ih« IhhI y oiiapiUting at 56' C,
4o« aaaoi oiMutial to the otKireraiMi of pla^tmine into fibrin, and we have
fMin S>r thinkinir that it 'n tUeU ctjnverlc<), in pitrt nt least, into fibrin.
Hmc* it ha* rvceiveil the name of filiriHw/en.
d so. The n-atritLH for thU v'mv! nri; an folloWM:
Bfaidai blood wbioh clot* naturalljr when ahad. thert- are Mrtnin fluids in
iba bsdjr which do not elot nniunilly, citber io the b.idr or whuti abed, but
which t>r cvriiiin artifir'iitl mean* may b« mado to clot, and in cloltinj^ tu
jriaJd ^nttr normal fibrin.
Thoe tb« »0'i-all(Nl ecroin fluid lakrii Mroc hours after death' f^om the
perifanlial, pleural, or peritoneal t-avities. the ituid found in the volarved
tareaaakc uf the leatis, known ad bylro^-ele lluid. and iitb<-r »imiliir fluids,
«91 is the majority of oues, wlteii ubiaineil free I'roui blood or othi-r admix-
lana, rwmain fluiil almost iudeRoitoly, shuwinjf no diipiwiiiuu whnlerer to
doc' Yet, in muK t-adce at uU events. tli«M: Ituidd, when a little blood, or a
St of blood olot, or a little terura h addul lo iliem. will clot rapidly and
Ir.' g:ivin); riae to nu uoroiMakable clul of nomal fibrin. ditTiiring only
fritn tk<> clot of blowl in that, wlieu ■crum ia usad, it i« colorless, being fnt
from rtti corriuidot.
Nnw blood (or blood clot, or iwrum> contains manv thinga, to any one of
vUch the clouiog power ihuvuvcn mixhl beatlribiiUM). But it h fuiind that
ia BMuy ouea dotting may Ix.- induced in the Buidt of which we are speak-
iac hj the mttf addition, and llial i-ven in <:xciixlingly small iguantity, of a
PiMlUK!* which (»n bo cxlracl^-d from l>I'x>d, or from serum, or from blu:)d
doCurerMi fnim waithed librin, or iitiL-cl fryjm olhor aourcea, a subataooe
whoM rxMct nalurr i; unn-rlniii, it Ik-Ir); doubtful whecber It is a protaid at
alt. aoil whoM' acii«ii i* jMrouliar.
If eenim, or wh!pj>nl blood or a brokcn'Up olot be mixed with a large
riBlity of alcohol and altowHt to ataad some days, the prvileids preseai are
tine M changed by the alct^Mt) as to become insoluble in water. Hence
if tbe orpiooa precijpitote caused by the nlrobol, after long aiandinjc. be sepa*
rated hr filtraiion from tbe alonbol, dried at a low temperature, ugt excead*
la( 40 C, and exlmcte<l with dutilled water, the aiiueous extras contAiitt
tittle protetd matter, indeed very little organic matter at all. NeTSi^*
1 ir H b« tnoavwl InawluMi tlUi iIhiIi u gaiisrally cM> rMdltr aad dralv. ftilai ■ oolarlcM
CM OMiMtBe of sum •iM >(dta owiouIm.
• la BM* (faiDiHai. iM«*<t>. • ■■•MUaaint oMfoWiiB, sft.'»llr iltsiii, tat iii «xotf€iaiwl
•■■• maMf*. mtj Im ulaanwL
• tM«S»w*aaM^eMsid*»aaaruiib«tbrtiid*<v4.
«
50
The
tbelcae, eveii 8 aninl! tuinntity of lliis afjiiPfnis extract milled aliDe to oertitto
sneciiQctiBiirby<)r>H-elc tluiJ or other of ihe HnkU H[>i>ke(i of above, will brio;;
Kuotit a Hi^edy dottiug. The eaiiie a>iueoi» extract has alau a remark a b£
l^l^l;cl in biuteDiun; the dolung at' Hiiids which. I.hougb they nill eveotually
cloL, do so ver^ slowly. ThiM ptaNiiia may, l>y the careful addiiJoD »f n cer-
tain quBiittty uf neutral «alt hdiI water, be reduce*) ti> mich a condition tliat
it i:l»i* Terv'iilowly indeeil, taking perhapM day* in cDnipleli; the proccM. Tlte
addition 111 a Anall ijuaiitity of thi: »i|iie<iiiK estrart wc are iJeacribing
huwi.-vi-r. bring abiiut a i-Inlliii); which w at i>nc« lapiil and complete
The active rubotancc, uhaltrvrr it he, in thii< a<)ucons «xlmct exists
rmatl quantity only, anci iiR cintiing virtues are at odcc and forever
when the wintion i« boiled. Further, there is no reason to think thai
active 8ul)s(iince actually enters into the formation of the tibrin to whieh it
glTM rise. It appears to belong to a clasi of bodies playing ad imiw.rlnnt
pitrt in pb^'siologicn I proc^iMS, and called fermenU.ot which wo shall have
more to say hcrcaficr. We may, therefore, ^peak of it as xhv ^rin ferment,
tbe DBiue ^iven l» it by its discoverer, Alexander Schmidt.
This Kbriii fcniit-nt is prec«Dt id and tuny beeilraclcd from dolled or
whipped blood, itnd from both tbe dot' and the serum of dolled blood ; and
bince in moel, if not all. cases where blond or blood clot or serum produces
clottiuK in hydrocele or peiicardial Huid, an exactly similar dolhinv may be
induced by the mere additiou of fibrin ferment, we seem justifieu in eon-
duding that the doitiiiK virtues uf the former are due to tbe ferment whic
ibey contain.
>i«w, when fibrinogen b precipitated from planma, as above devcribttl, L ^
sodium chloride, redissolvetl, aud renrrcipitated, more than onoc, it may be
obtained in wlution, by h«lp uf a iJilute neulnil mlinv solulion, in an *E^
proximnielv ptirv condition, nt all evciiM free from oilier protCKU. Such d
solution will not dot ■poiilHiieoiisly; it may remain fluid indclinitely ; an^^
yet on the addition of a little Hbrin fcnueni it wilt clot readily and firmly,
yiddinc quite normal fibrin.
This body libriiiogcn is also present and maybe separated out ftom ti
spccimcDB of hydrocele, pcrieaniinl, and otlier fluids which clot on tli
addition of fibrin ferment, and when the fibrinogen has been wholly retuovi
from these fluids they lefuse to clot on ibe addition of Abrin fttmeut.
raist;h'bulin,on ibe other hand, whether pieparcd from ploamine by scpa-
ration of ibe fibrinogen, or from aeruni. or from other fluids in whidi it ij
found, cannot be converted by flbrio ferment, or indeed by any other mrat^l
into tibiin. .\nd fibrinogen i.wlaled. as dMcrihed above, or serous tlutda
which I'oiitaiu fibrinogen, nin he made, by mcanc uf fibrin ferment, to yi«l<l
quite uiirnml ^hrin in ihe <um|i1eie abwnce of paraglobulin. A solution of
paraglobulin obtained from »erum or blood dot will, it in Inie, clot pcricar-
car<luU or hydnicdc tltildii containing fibrinogen, or indeed a KiluliDa of
fibrinogen, but thi« U apiiarenlly due to tbe fact ihal the imirnglobiilin ban
in ihm cucn fonie fibrin fcrmrnt mixed with it ; it is aUo puiixible tbut,
under rcrlain C4>ndiiioiis, the pr«»enc« of paniglobulin mny he tiivorable I^h
the action of the lermenU H
When the so caDtd pln^mine is preelpilatcd. a» directed in S 19, fibrin fer-
ment is carried down wiib ihe fibrinogen and paraglobulin, and when tbe
pliismine is iC'diHolvcd ihe ficiment i» prrsent in the solniinn and ready
act on the fibrinogen. Hence ihe redissolvcd plasmine dole spoDtaneou»lj
When flbriuogen ts isolated from plasma by repealed precipitation and aolu
Kin-
»Iv.
i
the
i
■ A wtKrlUI wlaUon «f nbtln fcRirtiil mty l* rMdllr |>ni*n<l !■)' tUa\it
bliH'd I'lol vtll ■ lU(*ruDI. »tiiil<in or •■xlluni chloriilc.
xmrUna • wulicd
TUK CLOTTING OF DLOOU.
51
P
tfaa, ibfl CtmuMl H WMhed nway from it, vnil [he nura fcnnvnC-rrvi' (ibriii-
mtm, ultinuiti'lr obtaiiHy), i\i>ts» not dot «ponlaD<i»ii«ly.
iki fmr it mvhio cl«iir thai thciv doos oxMt ■ proteld hndy, Rhnnoga}, which
mmj by the m-tJon of fibrin ferment hv <)!rrclly, u-ithuiit tht intcrvi-iition i>f
NtMT proleidA. <:i>avi.'ttr(l iolo ibc ]*>« m>lubic fibrio. Our kiiowlnlge <>j' lliv
nnnHllilifin of proteid Ixxlics is too iin|>crfoct to eiiablo us lu iiiako unv v«ry
ddbilta M*leiu«nt ■» to the cxiict nnUira of Iho chnn^ thus dTcctcd ; but we
mj MT ibie mucli. Kibrinof;cn anil fibrin hnve iibinit iho trntav (.'li'iucntary
■OBpcaitloD, fibriu ouuUiniiiK fl triHe inoro nilro);eii. When fibrinogen ib
eMmrtrd into 6brin by nwans of Kbrin fermcDl, the weight of the Hhrin
ptvdocBd k aJwayn l«i>s> than that of ibe Rbrinogen which ia consumixl. and
tbvre y bIwos jiroductii at the aanie time n t'ertaiu quHutitr of nnothrr pro-
Uid. iMloogiug tu the );lubu1iu family. There are reaauna, liuwever, why we
OkOM* ipaak of the furmcul. un tjttitliH^ up 6brinu^D into librin and a glo-
bolin : it avcmi moro fmihHbl« that tike ferment cmverU the librinogen lirst
into m liodv whi<^ti «i- luivht call toluhU ^rin, aiul then turua this body iutn
rentable lihriii ; l>iit furihvr iH(|uine« un lite subject are needed.
It nay b« uihU-fl ihiil among thi; cnnditiom ncM«iary fur the due action
of Sbrin rr-rawDt on fibrimigCD, the pruwooe of a certain quantity uf sonn
Hairal talt Kvns to b« one. In the total abaence of all neutral aalin the
ftmmX oinnot convert th« fibriDogvn into fibrin. There arc itume reuaouB
ako Jbr thinking that the pn-aciice of a lime .talt, «uch a« calcium aulphale,
tlMOgli it nay be in minijie i^uMntity oidr, U <-]wiiiiitl.
ffi. We may convlude. Ihcu, that the plnHnin uf hhxMl when Hhed, or, at
•U «f«Bla,a(KiD after it h»» l>e<.-» *liL'd, contnin* librini<)|:en ; and it al«o Mcenia
pn>lwbl« that the clotting cmiiiii about Ifcauio ihf Hlirinoj^ is mnvurietl
Btio tMa by the action r<f librin fvrniritt. ; but wo nrcilill far fmni a <Ivfiuit<>
aanrar to the (pMsliMi, why blood remains fluid in ibe body and yet clut«
wbra »hod7
We have already uid that blood, or blood ploama, brought up to a tem-
pefBlttK of 56'^ C » «oon as |H«iiibIe sAer tla removal from the liviDB blood-
•MMk, givM a proicid pr«;ipiiate and 1u«m» it« imwer i>f clotttnu. This may
W taken to *how that blood, m it circnlaifo in the Uvinfi bloudvewela, con-
talaa flbriaoiNn a» nich, ami that nhm the hlui»l ■• heated tu 56° C. which
ia tbe cimguTating point of libriHcrgcu, the fibrinogen proteul ia cuaKulateil
aul preripitnltil, and conwK|ueully no librin can lie formed.
FurtlMT, kIiiIi- riotlod blood uiuluul.U-dly i.i>niBini> un abundance of fibrin
fcnerot, ii» fi*rn>ciil. or a minimal c|ii]tniiLy only, i» preeent in blood lu jl
Iravr* llu> blood v(tPi*l>. If ih*- blond be rcceivt'd directly from the blood-
weak iutii eloihol. tlir niiuentin extract pr\-|<an-d, ai> directed above, cou-
taioe Bii frriiicnt, or mrr\'ty a truce. Apparently ihv ferment maktv itM
appearani-r io the blood ae llie result of eliangm taking place in the blood
aflcT it hat licen ahed,
We iiii):hL from ihia 1>e inclini-d to (-oneUide that blood cloU wlM-n lilied.
bat iMit betori!, heeaoHe, fibrinogen being nlwaya prreeut, the chethlinii bringn
abuut ctumgea ithieh prMluce fibrin fernieiit, not previonely csi«ling, and
thii acting on the (ibriiKigen gives riae to librin. Itut we meet with lliv fol
kiwinf difBirulty : A very couHidt-rablo i)uaniily of very active ferment may
be injected into the b I oiid- current of a living animal without necrvvarily pro-
Jad^ any cl'>tiing at all. Obvi<Hifrly either hlood within the blixidvemcls
da« twM contain liljrinot;en ati such, and the librioogeo detected by healing
tko bloud tu >'><{' ('. i* the ivsult of change* which have already eonted Iwfore
thai lerap^mtun- i« nacheil : or in the living i-irnilaiiou there are agcocica
■Cwwk «hioh provrni any ferment which m.iy Im tuLroduccil into the cireu-
Inlioa from pruduciog iu usual efl^la mi tibrmo^n ; or there are agvnctva
62
BIiOOD.
at work which dntro^, or do away willi the fibrin, little hy little, aa Jt b
formed.
$SS. Antl inilecti when wo rvfl«ct bnw citi)])I«x btoixl ■», and th« toauj
and great chniigra U is simocptiblc. wr shnll not wonder tlutl ihe quettion
nre mittiiiK cnnnot be answorcd off hand.
I
The coriiusclftt with which liWd ic crowded uro living stnieturei^
coniMKiiictitly nro continually ticttng upon nnd being aeUd upon by the
pliuiDB, The rod corpuscles il is tniB are, m n-e AmU h«. pcctilinr bodici,
with « rc«tnct«d life and a very speeialized work, and pomblv their influ-
cuc« on the plasma ii> not very gn>at ; but we have reason to think thai ibe
relations boLwcen ihe while corpuscles and the plasma are close and im-
portant.
Then af>ain the blood is not only acting upon and being acted upon by
iho several ttsFues oa it flows tbrotigh tlie various oamllaries, but along tba
whole of its course, tbruugli the heart. ar(«riefl, cnirillaries, and reins, is acl-
io^; upon and beiog acted upou by the vascular walls, which like the rest of
(he body are alJve, and being nlive are continually undfirguin;i: ami promot-
ing change.
That relution« of Aorae kind, haviug a direct iufluonoeon ihecloUingof
blood, do exi»t between the bl»od and ilin vaMCiilur walla is rthown by the
following facts:
AiitT d«aih, when all motion of the bloot) haji ocaM*!, tlie blood r«(uaiat
for a long time fluid. Il is not until some time nfUrwnrd, at an epoch
when post-Diortom ehmigej* in tbc IjIoikI aiiil in the bloodveMvl* bas-e hod
time to develop thcniscrvi's, ihat clotting Iwgins. Thus Rome houiv after
death tbe blood i» the great veins may he found i>lill pcrfecllv fluid. Y(t
such bloorl has not lost its power of dotting; it slill cIoU wlirn removed
from the body, and clots too when received over niereiiry without exposure
to air. showiug that, though the blood, being highly venous, b rich in car-
bonic acid and contains little ur uo oxygen, iio fluidity is not due bo any
execm of carbnuic acid or abni^nee of oxygen, Kventuatly ii does dot even
within tbe ve-tii^lH. but |ierha|<!i never so firmly and completely at wl»eu
ahed. Il clots tSriil in Uie larger ve>Mel-t, but reraainsflui<l in the amaller ves-
sels for o very loug time, for ninny houn lu fact, «noc in thvse theaauie bulk
of blood ia expiKted to the iulluence nf, and reciprocally exerta an influenty
on, a larger surface of the viucutar walU ilmn in the larger venela. And
If il be urged that the re.iiiU i.i here dui? In iiillui^nceo exerted br tbc bodi
at large, by the tiasuM us well ai> by the viweular wallit, thin objection ~*
not bold good agaiDHl the fiillon'iag exiicriinont.
If tbe jugular vein of a lurg<! aDiinal, such a« an ox or horse, Iw carefil
ligalured wneii full of hlooil, and thu ligatun-d portion cxdwil, the blj
iu many caiHSt remaimt perfectly fluid, along thr greater part of the len^
of the piece, for twenty-four or oven forly-dght liours. The piece so liga-
tured mav be eu«pcndcd in a framework and opened at the top so as lo imi-
tate a living t(»t-tube, and yet the blood will ol\cn remain lone fluid, though
a portion amoved at any time into a glass or other ve&#el will clot in a few
mmutes. If tiro such living teet-uibcs be prepared, the blood may be poured
from one to the other without clotting takin;; place.
A simitar relation of the fluid to its coniainin;; living wall is seeil io tbe
case of those serou* fluids which clot spontaneously. If, oa soon after death
as tbe body i* coid and the fat is solidified, the t>ericardium be carefully re-
moved fVotn asbeep by an incision round the base of the heart, (be pericardial
fluid (which, as we ha>-e already seen, during life, and some little time after
death, posacBsea the power of dotting^ may be kept in the pericardial bag a*
THE CtOTTIN'O OF BI.OOD.
68
te > livioft Clip for Riauy boiin wiiiioul ilottiiif;, utA yet a iihaJI porti»u re-
Bomil mtli n |>i]>ette rlols st once.
Tkti rvlKiioti brinu-ii tli« IiIimmI ami the vasculur wnll may be dialiirheil
or ovcniiMf-n : cluilitii; nmy tiike place or may be induced niiliiii itie livluj;
Uoodtv**!. WIm'ii iliu liDiriK uiviiitiiuD« » injured, n« irkeit nil arury or
win h ihsrply lii;iil<in-il, >jr wlivu it w diceaKd. tu fur iDHtaiicc in Biieurwro,
• dot i* *\'l lo bv li<riiiiH] at ibo injured ur diM-tiH-tl sixil ; kikI in ccruin
■oririd eoodilioiM of the body eUitf arr r»rRi<s) in rani>uii vaiwular UitcU.
AlnmeB of iBntion, «hioh in vhtil I>Iimh1 ii» w Imvc »«en, i« unfuvorahic to
dattiiag, 1* apt "ilhtn tlH< bo>ly tu knd h> plotting. Thiu, uhen iiti nrlcry
b HpUnrrd, tlic blwid in lli« tnct of llie arlvry on the curdini: nidi; of the
S|Uan>, bolwcf'n tbt> iigaluir aul (hr hninoli Wt givvn citT by lh« tirlcry,
amtkof' to iban in tbv circulation, rvmaiuv ingticinlcm or iKarly ao, aiid
•Img thb tract a clot (nrta%, linni.-*t nvxt ti> lli« ligiiiure nod eoding atmr
■ban iIm branch i» given off: thii> twrhap* may hv cxplitincd by the fact
that tba walls of the irsrt Kiiflcr in rlicir nutrition by Ihr rtagnaiion of tJi«
blood. Ktwl itial cuQMqiHiilly the Dornial relutioD between thcoi and the cod-
Uiaad t>['-»l is dialurmd.
Tbat th« blood within tlie liring hloodrcaacb, though not actually dolling
nadar DoraMtl circuniKtanix.-^. uiuy cwily be made to o(ot. Dint the blood is,
:. av t« malt, always on the point of cloitiDK. u shown by llie fact
fonign body, nieli as a oeedle thrust into the interior ufa bioodrnael
fvad drawn lbrou;!h amt li-A iu a Idoodvcaial. is apt to becoma cot-
«ilb fibrin. .Sotoo influence exer(e>I by tli« needle or thread, whatever
ba the cfaiinu:tor of lliat iofluence, i» sulHcienl to deteriuine a cloiliu);,
Mbcfwite. would licit have taken plaee.
«a»c iiutakiliir of the hliHxl, as regarda clotliu^', is strikinjciy shown,
caM of Iho ra1il)it nt least, br the rtsiilt of injttcline into tlic blooil-
a snuill ijuantity of a solution of a peculiar proteid, prOMired from
•truotun* such us the tbymiis body. Masiive clotting or th« blood
iMt all the blmxlveAsels. snmll and large. takc« place with great
y, iMdinjc to the »iddcn death of the animal. In cotilnut to this
may br nwnti'Mied (he mull of iiijectin); inio the bloodvessels of a dotf
qitaulitr "f u tohitioD of a body called albtimote, of which we shall
h^wUrr linve lo trnit ns a product of ihe dif^Uoo of proteid sufaMancca.
aatent nf (>M jcTammc |ier kilo of body weight. So far fmio priKluc-
tiog, Ihe injcrlfd nititinione has such an e^t on the blood thai for
houn KJVr ihi- iiiictioD shed blood will refuse to cl»l of itself and
D quilfi fluid, Uiou)(h it can bo made to clot by sjieciul Ireatnicnl.
SS. All thi- forevoing fiwta tend to ohon' that the blood as it is flowing
the htwlihy^loodvcanla (■, a* far aa •■lotting is oonccroed, in a state
wguililiriutu, which mar nt any inonK'nt bn uirvt ; even within
lv«ael», iiu'l which is upnet directly the blood i* nWil. with clotiing
ramiL Our pmteiit knuwl<><Ige does not itenuit ii* to make an authori-
alatemeol n» lo the vxttvl naluni of this etjuilibriuni. There are rea-
however, for thinking ihnl the white oorpuwU'w play lui ini{ioritinl
ki the tnattrr. Wlimtver cJuiiiDg occurs natumlly, whitv corpuscles
I : and ihiu is true not only of blood but also of such spocimena
dial or other serous lluids as clot niilurally. Wlten horse's blood
kept fluid by luHnf; relaineHl within the jugular rein, lU menliomd a llttla
V bock, and the vein is hung upright, the oonMiscka, both r«d and while,
., leaving an u|-|K-r laver nf plik^nia almost rrae from oorpuacles. Thia
ar layer will l>e foun<l to hare lost largely its power of clotting spon-
■ly, though iho |iotier Ii at once refined if the white c>r|itiscla from
layrr* beneath be relumed to lU And many other argunKnts, which
■bike I
MM If If. I
UW Uyrn 1
A
the
all pcimrin^ lo the oaine con-
iinpuriai)t juirl iti lUe promss of
u'e cannot enter upon here, mny he ndduced all
rluHiim, thai the wiiiie enrpuBcles play an _ ^
clotting. Jliit it uoiili] lead ua tix> tar into L-i>ntravertaaI matten lo atleiiipt
to define what that jmrt ia, or to explain the exact nature of the equilibrium
of wbidt vc have anakeu, or to diacuBS such <|uciitiiMi8 aa : Whether ib« ordi-
nary whit« ei^r[iu*u[ea, or corpusolw of a s|»et:ial kind are cunccnicd in iW
malterf Whether th« oorpiiscles, when cloltine taka place, e.^., fibrin-
ogon or fi^rmeiit or both oraomeihiiiK else, or whether the onq>ilwIcii »im-
ply in womw way or other Hfsiitl in the tmnnformnlii'in of .iOmo prwiou*tj
«jci»tiD]{ w>ii»litn«DU of ihc plaiMDa? Whethi^r the influenoo exerted by tbc
condition of (be va»cii]nr wall it exerted dirwctly on the plunma or indi-
rectly on the corpuscle*? Wlicihitr. u •ome htive ihoughr. iliv pcailiar
bodie*. of which w(! shall presently upcsk under tJic namo nf blood plaleJete
or pia^uet, hare any share in the matter, and if tra what? Th<«« que^iionB
ar« too involved and the discuwinu of thcni too long to ho entered upon hero.
What we do know thnt in blood soon aftor it has been elied. the body
which we havo called fibrinogen \t present a» also the body which we have
called fibrin ferment, that the lutter actlnj; on the forni>!r will produce
fibrin, and that the appearance of Hhrin is undoubtedly the csu<e of what
is called clotting. Wfl Beem ju^litied in coticlitding that the clotting of abed
blood ifl due to the ConrerHion by ferment of tibrinogeii into fibrin. The
furl her inlereiice that dotting within the bmly is the same thing as clotting
outside the body, and stmiUmy due to the trans for ion tion of fihrinoji^u by
ferment into librin, though protMble, la not proveil. We do not yet know thiB
exact nature and cundition of the blood within the livio); bloodve«itla. hd^B
until we kudw that no cimnot nit ii»ftiut only explain wliy htuod in the living
bloodveMelgi is iifiiallv lliiid hut can at linuw clot.
The Coi{i'i,'H(;i.iw or thi: B1.0011.
Tlie Rfil CorpMdf*.
§ 34. Tlie redneM of blood is due exclusively to the red corpuaclea.
plasma a* K«en in thin layers within the living bloodvnaeU appenn rolor)<4«,
Bsdoe* a)M)a thin layerof serum : but a thick layer of serum (and probsbty
of plattnm) Iibh a faint yellowish tinge due, as vre have said, to the prcMnoo
of a small ipiantity of a sfteml pioiuMit.
The OorpusclM appear under the niicroM!0[H- a* fairly horaogVQKHi*. im*
perfectly traiwluccnt biconcave dLir* with a diameter of 7 to A yi and a thick-
neaa of I to 2 />. Bdng dine* they arc circulnr in outline when seen on the
flat, but rodxhajmc! when »neii in prolilf tus thny arc liirntng over. [Fig. 'p. J
Bdng biconcAvc, with n thicker roiindM rim surrounding ■ thinner centre.
the rays of li^ht in piu«iing thn)ugli ilicm, when they are examioexl by
transmitted It^ht, arr^ more n^fracled at the nm than in the centre. The
etiect of thii> i>> that, when viewed at what may ho considered ihe proper
focUH, the centre of a coqiusclc np|>cnra clear, while a eliijht opacity mark*
out indinlinclly the inner mnrgiti uf the thicker rim, whereas, when ilie foc-us
is shiAe<I either up or down, the centre becomes dark and the rest of the
corpuscle clear. Any body of the same shape, aud composed of substanoe
of the same refractive power, would produce the same optical eAeolS. Other-
wtw the corpuscle appears homogeneous, without distinction of part* and
without a nucleus. A single corpuj«cIe seen by iwelf has a very faint cohir,^
looking yellow rather than red, but when sevcml corpuscles lie one upon tk
top of t^e other die miuu is disiinelJy red.
TIIK COHPUacLKS vr THK Dl.OOb.
&s
Tb« f*d corpuscle i» i-lftstic, in the kdik ibat it tstny be deforitied lij
ptamni or tnctloo, bul wlieo Uie pnasun; ur tni(-ii<>n in removed rogaiai iu
pnviiMn fbnn. lt» shape b also touch inHiiviii-t'd bv tbi; phyncal condilioM
I
irw.».
Fio.a.
fh). 9l— NcsikS IhJinli ^ ma u> Hit VTini) «tiiiK. lUniiiiIDO'l atoHit IMDdlUBtUnkI
I I Itmfh Urt IMI I IjlUKlUi, r.r; rvnliviiinwlcion (belt odc« UMl rttwol tli t*alU*l
'•, mA iMiBwtlti ■n«niid in roulawii; r.r.crmUc iwl cvrinMiln: p, ■ HMty B'<»i>l'>r I*!" *•"•
tm^t . 9, • nanvlf (M»al*r p*te <«ir|iaKlP. U-db liarn txu uf Ihitw dliUnol ncinln. nod *cfw
wAnaoUit cbaii^M •>( (hatB at lira iDM»iri of •it'wfTiilkia ^ ill p ■ tiuclcoa tim turn rlilbUt.
r>> d-lliat.i Kiti cuHi^-nuu. I.iin-. !<t!i»LY u<»'»i.unn> Drm Iumia (At wmsnderM
Witi ^w*r at Iha nucrnBM)!*. i ]
If
irxi I.
8
«f tlMpla«Bm,Mrum.wrllui(l in which firthi- limo being it i*. If tbe {ilnsma
«r Ptmtn be diliihs] with water, the ilisc. ulMi>rliin^ wiiler, fniKlln u|i inl» ii
*pb«n> [V\f. *}]. becoming a disc n|;tiiii on tbe rciDOTsi ol' the dilution. "
iMMTUm be coocentraied, the disc, |>ivint; out
nl«r. shrinks imgubrly and aBsunrcs vaH'xi*
(^•rcM; <)D(! of thcM fornu is Ibsl of a nuMilx-r
of Uodttd |>MlubeTauu»i pruimiin;; ail uvcr
iW Mirfiue uf tbe t;ijrf>UH:lv, wliifh is then said
%m b* crvnKte; in n drop of blood (.-xamined
HMler tlw niurosopt', vrenale mrpiiM^'les are
«Aaa aeeo at ihi- nl^ie of the cover nlip where
«<npmii(iD i» leadins li> ronocn I ration Df the
ptoawK, or, U it stuKifd tWn |H-r)in|M rather be
called, (Wiiro. In hlooU jiut ahrd tlw ml cor-
pnadM arv a|it In adhere to each iitli«r hr iheir
^, .^^,,.^_ much more than tt> the giiMi or
Hti ■' with which the blood ia in rontaci, and henoo arrange tbemMlTei
ht roll* riiif tendency, howevar, to fiino n>lla very etxin diminishe* nfUr the
blood bihiMl.
8 8
4 t
oo
o-f, >iici«ari*« eilbtt* "X water
ii|>iu ■ ivl (orpiuota: / <tlR«i '4
•uliitlna uf ibI(. cniiutcd : f, vllbft
of Waiiie Kid.)
Tbouli n •iii^lp ri>rj>iiwl4- '\* Hinn-nrhal trniuliiceDti a coniparalirely thin
kjvr M klotid \» <>|iiu)iip ; tvjH-. for iiiKiance, cwtnoi be mtd tKrough eraD ft
IMCl
thin Uf ar of blood.
WlwR a i)uantitr of whipiiud hlood (or blooil otborwiM depnved uf fihrio)
ia frvMO ud ihawi-d Mvend limes ii ehangea eoJor, bocuming of a darker
66
ii3-
inuP
hiio, iiml if tbrn rotind lo l>c niiicb nioic tninKparrnt. *o i)iot ivfx- win roir
b« irn»ily rvii't tlinxigh n niu<l«rnU-lv Ihin luycr. Il u then ^fmlcrn of u
iahfhiood. The winiv rhniigo niuy lie cflrctoil by »hiiking ihp blood with
ether, or by ad(lin<' n smnll <|iiRntily r>f bill- snll^ and iii other wnyi!. Vikki
cxaminaliou of laky blood il is fouiii) tbnt the rr<t oorpuroles nre " brolcni
np " or al li^asl altered, and ihtit the rcdnns nhk'h nrvviiituly was oonlinrd
to tbem ia now diffused [broiii^h the Minini. Normal blood is opaque bveauat
each corpiiEcle, while pcrniiUiiie tome mya of light (etiielly r«d) to oaa
ihrougb, refteclB iiiaoy otbers. and the brigbliieM of the hue of Donutil blood
» due lo ihis reHeclioii of liiihl Iroin the «iirfiiceB of the several corpusoltt.
I^ky blood ifl traiisnareut because there are no longer inlact corpufo)«*to
preseut Burfacn fur tlie rellec-iioo oT light, aod the darker hue of laky blood
u (iinilarlr due to the uttsfuce of reflecliun from the several ci-irpusclee.
Wbeo laky blooil is allowed to atand a sedinieut is tbrmed (and may be
Mparated bv ibe centrifu);ul machiue) ivhich on examinaliou is found to
conpist of dUcK, or frKftnieiiLt of iWws, of n colorlew auhstiince exhibiting
under hi^h power* an olweun-ly Kpoiijiy or reticular structure^ These cotor-
lof^, thin ili»('K MTii HuI-ivim; ofirii appear a» mere rings. The aubstancfl
oonii><i»ing them »tiiiii:< wit)i vurioun rt*ag('iiti' and may tlius be made tDure
evident. jfl
1'hu rrd corpuscle, then, con»i»ta obviously of a ooUiriut frainontirk, wIhB
which in nortnsl conditioiii; n red coloring matter is tURiciaU-d; but bjr
various means the coloring matter may bv driven from the framework
dissolved in the serum.
The framework is spoken of se ^rtmo : it is n modified or dilTereutinl
protoplasm, and upon chemical analyeia yields proteid suhftunceis some of
them at least belonging to llic t/lohvlin group, and olher matters, amoog
vbichis a peculiar complex fat called /nrMm, of which we shall have tospeatc
In treating of nervoua tissue. In Ihe nuclt-ated led corpuscles of the lower
vrrichraln this difl^utiated ulroma, though forniiufi; the chief part of the
celMiriily aniuud the nucleus, is accompanied by a variable amount of undif-
fenniinied protoj>laMii. but the latter in ihe miimmalian iv<l corixisele b
either aWnl nllo^lher or n-diiced to a minimum. Whethi'r any part of
this stroma is living, in the scnsi- of being ctijrmbleof carryineon a continual
double chemical chang*.-, of i-onliuunlly buili^ng itself up as it breaks dowiH
il a qiieflion too liitliaill to he <lii'ciin>ed hcnr. ■
The red coloring matter which in normal conditions i» associali-il with thi»
Stroma may by nppTopriotc means be isolutcti. and, in iho case of ihv Mood
of many auirosls, obtained in a crystalline form. It is called hirmaghbiti,
and may by proper methods be split up into a pruleid belonging to the
globulin croup, and into a colored pigment, coniaiuing iron, ctdle<l nirmatin,
nsenioglobin la, therefore, a very complejc body. Il is found to have remarlt-
mbl« relations to oxygen, and indeed, as we shall see, the red corpuscles by
virtue of their hiemoglobin have a sjiecial work in respiration ; they cany
oxygen from the lunga to the several tittuei*. We »faall therefore defer the
furtliir study of birmoglobin until ne have lo deal mith respiration.
Tbo red corpuscle, ihrn, con»i»t» of u ilinc of colorkrs stroma with which
is acaociated in u prculinr way the complex colored btidy haimoglobin.
Thongh the hamocti-bin. an in >ecn in laky l)lood, ts readily Mduble in serum
(and tl is nUo soluble in pliisnm), in the intact normal blood it rcmaJus eou-
fined to the corpuscle; obvitiuiily there is Home special connection between
theatroroa and the btenioglobin; tl is not until the Tlromn is altered, we may
perhaps say kille<l (ns by repeuted frci-zing nnd thawing), that il kwfl its liold
on the hx'm<>>;l<>hiu, which thus set free passes int" solution in theacrtim.
The dbc of stroma when separated from [be bicmoglobin ha«.aEweharcJi
THE CORPUSCLES OP THK DLOOP.
sr
«U. an ot»eurclir «iH>ne%- Uixturp ; but w« ')» ool know aocuralcty th« cxscl
eowliUnn of lbi> Nlffuit in the iiitacl oonxiKlo nr how it holde thi> hiemo-
sloUa. Tlwn b nrluinly poilvliniu- mi'inhniiK; nr envelope to lltcGornugcIc,
nr b]r expoung bliM>il toabigli lemin-nittin!. tU>° ('., the oorpuBole will break
apialoil>on4>rlcMs|>bmciil pientt.eiich still i.xitii>itlin);ijf Mroinaatld b»RiO-
Tbtt qouitily of nromn ncccsMrv to hold a qiiuoiiiir of lunooglobiti w
«ac«T<iit)p1y small. Of ili« total Boliit inaticr of a corpu»d« tuon than 90
Cr cent '» luFtnoglolHii. A red comiMrle in fad is a ijiiantity of hieniouloliin
U (ogvther id the form of a dbc by a rainiinal ainntiiit of itroma. lience
wb»l«T«r fflK;t ihtt stroow per te may have u^ii the plasma, this, io the
ease of manuuala at all ereuti, muu be iiui(,'mticaiit ; Ihe red corpuscle is
pcactirallr Rtniply a carrier of hieinoirlubiti.
i SS. "file avemgie number of rvd ci>r|ftiicl«a in hiinian blood may be
probsUjr put down at about 5 millioitu in a cubic niillim(4re (tli« mn^ in
diflcnal ntauimals i« mid to be from '-i tu 18 milltoii*), hut the relation of
eariNHol* to nlaama Tarin a good deal evi-n in hralih. nnd very much in
dinw^ ObriuutlT tba relation may be allwicd ( 1 ) by mi iiitn-nim nr decnaae
of tba pliRnn, 1 2> by iin net mil decrcaieoriacmucof red oorpiisrlr*. Now, iliv
fcrSMT nuM rmiucnljy take nlacc The blood, as ire have nln-mly iir){i.'d, is
always bring acted upnn by changm in the timie* nn<l. indccl, h nn itidi^s df
thomt ehangn : bCDCv the puiMmi tniiet l>e «>iiliniially chnnging. though atwAVN
sUiviiW III n-luro to the nurmni L-oiiditioti. Thus irhcn a Inr^-r >jiiiiiilily of
wmtar b •lischareeil by the kidney, the skin or the tK>wols, ttint water citncs
rvmflv from the filtMo), iim) tho drain of water must tend to diminish ihn bulk
iif tDO plosniii, anil ri to increase the '•eialiit number of red corj>iiscU')i,
tbmigh tlir etftfct is pmbnbly mod remedied by the |iaf«a^ of natcr from the
tiiauf* ititii the blixxl. Ho ai^ain when ii Inr)^ i^unntity of water is drunk,
tbia piwrs iniu the blood nuil li-nil.-> lempDmrily lo dilute ibe ulasraa (and so
to dimiaish the ntniivc nuinWr nf red oorpunclev), iboufch tbii ooiKlition ia
ia tnni snuu rf'a>edir<l byihe|<a»>iigeof tbeauperfluAiu> fluid to tbu ItMuea aiMl
axcrrtury organs. The greater or liw number of red curpuide*. tiKn, in a
fhren bulk of blood may be >iiuply duo to Iimi or mnn^ pla«ma, Init we have
rraaoo tu think tluil tbe iictiml nuiuhor of ih*- eorpiiM'hv in iIh- blomi does
Tan^ fmra lin>c tu liuH-. Thi> in rr-|>i.-cially m-cu to t-ortain formii of disease
abieh may bo spoken of under tho geneml k-nn of anicmiii AWri! being
Mveral kiuils of aoatmiai. in which the number of red corpuscla is dislincUy
(Uslatilicd.
Tba rtdorsa uf blood mny. hone^'er, be Influenced not only by tW uuml>er
af red corpuscles in each cubic millimetre of blocMl, but also by tlic amount
af haaM)|[lobin in eneh corpurele. and to a le» degree by tlie siio of llie
rorpBSCWS. If we compare, nilb a common standard, the rednMs I'f two
wpmtbamat of blood unequally red, and then determine tlw relative numU'r
irt oorpuaclta in each, we may And that tbe len red specimen has u many
rorposclr* as the nilder one. or at least the deflcieocy in redness is grtMcr
than can Ik- accounted for by llw [inudty of n-d corpuscles. Obviously, in
socb a cw, the ml corpuscles have too little ha-movl«bin. In Mme coaea of
Boamia ibe dcKcicncv nf binnnglubin in each corpuscle a more Mriking
tbaa tba acantincM of red cuq>u8c)ca.
Tb* aDiulirr at (orpowln in ■ specimen of t>lo<Ml U dctetminH by mlalng a
Maall but carriulty meaiiurtd <iu»iitiiv of the blond with a Inrjce <|iiantity of some
iadiAravnt Auid. f.y,, a A per ceiil. soliiiion of tiiillum »ii1]i|iiit^. nnil then uclunlly
cowntiag Ihr. corpii*olr^ In a kn'>Hii iiiiindiiiil Imlk of ttie mixture,
' , pcfbaii", may b« mmt ci-nvcuii'otly done by Ihi- itii'ilmil gcoerally known
^^U^P
DLOOD.
■* iluit of iiuwen {HKtBKicjUMattet) [Fig. $}. ltnproT«tl b; Maluwa. A glan
bII<1«, 1b • m«uJ rnun«, ii rwtd into minute rM:Ui»t«>, •■. y.. \ mni. Ii^ | mm., m
H toglvf &conT«nl«Dt«mof Aof o*n""^ ■"■"- ThrrcuiuUI x^vm in thernm*
paraitt a novcralip to ht bratiftbl to ■ fixnd diatnnce, <- 9. i inni.. from the Nuifaoe
oflbraliilc. Tb« blood bavinr b«4i diloinl.f.^. tu lUO iiin«* itn ralonie, • amall
iiuikntiljr of die dtloieil ^«Dtl tburuusbl}' mlivai blood, iofflctent to occap.v fully
UM ■pftM between tbe eurenhp aiM the glaaa alidfi whed lh« formrr i* brouEht
lolia properpOililMi.ia placed on tti«al>ile,«tid the oovenlip broaght dowa. Tb«
trw. a.
U»
d
A liHWllbt ■Wurtot Uw iHlWlin mliiitan; fl. oiJlltJT kiIr foi <.> ..- irL.v ),1:m>1; c. mil
allb OlTtiluni on *Me, mv««ittuB uid upAiup: D, rami (ob^i iiilucliiiit. ^ inlxtri P, siMrdnl
■■•Mr'podnti-l iiordls fOrMlcilnxBniKV.)
rolumoordiliitod blood now lying over «acli ortfafirfctaneliw Kill be 1)9 I ^ X' J)
of m cubic mm. ; and if, wb*ii thr< i">r|HiM;lM have ftulMiik-d, ll» number uf cur<
piuelM lyinn within ■ recungte be L-oLinto'l, the roHuli irill cive the number of
cnriKuclM pievioualy dlslrltiuted through tJo of a cubic mm. of the diluted blood.
'riiiH mullipliwi by lOU will give the nnmber of rorpa»clM in I cubic miu. of l)i«
dilutetl blood, and wain iDultipHnl by lOO tho number in 1 cubic mm. of ihr <>ntiro
blood. Il io advinibtn lit count lli« Dumber of cofpuMle* in iier«nil of lb» rr«-
tKDglM, ami to lake ibe average. Fur the c(tDveui«nce of counting, each mtansle
la aubdivided Into n number of verv amall aquaroi, r, y. into 30. t»eb wtUi a aide
of Jk mm . Hiid DO an nrea of ,Jlg or a aqnarv mm.
Siner (Iw actual numlier of tM cnr{Hi»cle« in 1 specimon of bloiMl (irlilcli
may be tjiken lu u Ntm|)U- of ihi; whol« blood) tssomelime* more, aomctinxM
len, it is i^bviouH that either n-d (.'orpuwtea may be teRi|><>ninlv' withdrawn
from and relunivd to the grncnil blood current, or that certain re<l corjmHctea
are, aA«r a while, matlo uwajr wilh.iuid that iiea' oiiim ttiko thi-ir place, We
have no aatieficlory cvitk-noe of the former heint; the ca»o in normal condi-
tions, wbereait we bavo evidence that old cor|>u<clM do ilio and thnt new ones
nre bnrn.
^86. Th« red oorpmclee, we have already siiid, arc cyiiitinuully en-fa;;ed
in carrying o»yj[e«, by moans of their batmoglobiii. fnno the long* to the
liaauea; tlie^ load l)i«mBelvea with oxyK«n at lh« luiiii* and nnloiid at th«
tlMUCS. ll u extremely unlikely that tbia act should be rvpoated indeliHltely
rnt coRpracLcs or rnic blood.
fi»
I
villtfKit leoilins t» chkOfrefl irhicti may be fiiniiliarly described lu w«ar aud
taar. nntl wtiii:!) woub) ulLintalely leni) to the denili ut t\\^ turpiisclw.
Wfr ftbull hrtve lu stale Inter oti thnl ibe liver didcliaripa into the nlinien-
tmtj ivnal, lu a conMiiucut of biU'. a (Nminiderttble ((uaiility ul' a pit;n)vnt
knnwn n» fiUirttbiH. and tliat thin iiuhi>iauoe liiu tvuiarksble rolatioiu with,
»'■ I. majr he rcKiinieti as a derivative of Acmwi/ui, wUicb, as w« bav«
•r< . • , ii a prciiliK-t of th« deoompotitioii of hn»uoglobui. It appcan
)«T>faabl«. ID flirt, that ttic hilinibfn of bile I aud thin ii» vtir nUall woe i* Iba
ctilrf biltarr |iif(UH'iil, hihI ih^ source of the ntlicr liilinry pifjnicnls) is not
fonnnl wbollr nin-w in tix^ budv, but id maiitifucturtHl ia w>inc ituy ur oilier
ODt nf lueuiatiii derived from liiuii(>gli>biu. Tbt» mtuil cfiUiil n diiiljr onn-
•anptioB of a consiik'ntblo minntilr <>f hienioglobiii, and, »inec ivc kiiov,- no
Other sourt* of bHtuogtobin besides Uio red corpuocW, and have no uvidutiuu
of rrd eorpusclflt coDtinutne to exist after having lost their hemoglobin,
(Diut, Iherefiirc, entail a daily deatniction of mnny r«d vorpusclec.
Eveo in health, then, n number of red oorputcJes miut be continually
duppearinj; ; and in diseaM the rapid and f;rent dimtmitinn which may
take place in the ntiniber of red corpusctea »Iiowb that large destruction
aWTOcnir.
We cannot at present accurately trace out the steps of this disappearance
of red eorpusclca. In lli« spleen pulp, re<l curpuscles have been seen id
varioua wiMgn of disorganiiation. m>iuu of them lying within the suhetaiwe
of Urge coRirleflB mrpuscles, aud as ii iT«re beinj; eau-n by them. There is
alsoerMence that deatruction takes place in lb« liver itMlf, and. indeed,
alwwbvre. But the subject lias Dot yet been adec[untely norknl out.
997. Thia dralniction of red corpuscles ncovMitatisi the birlh of new
(nrjiuivlM, to kei'p itp the uominl mpply of bnnoglohin ; imd. indeul, tlie
ca«n> in which after even grcnl loss i>l blood by lieioorrhage n h4«lthy
ruddiiMB* return', ami that often raindly.ehoning that the lost corpiisclta
bavr htva replaced, as well as the cows of recovery fVnin the disease ana-min,
tifnre that red corpuscles are, even in adult life, bom soinewhero in th«
lo the deTClnpinf; embryo of the mammal tlie red corpusclee of lite blood
af* not hKniof;|(ibin'huIdinK non-uuclealeil disra of stntma. but colored
DucUaled cells which have artsea in the followinj; way.
In o-naio re^oiM of the etubrru there are formed neata of nuclei imbedded
in that kind of material of whii-h we have already (S h) spoken, and of
wbMi we shall have B){»in Ut ii|N:ak, aa unililPeretiliated protoplasm. The
Tf^f*"' faalnrrv »t' ihi* uiidilli-rvntiiUed )irotu[>hu>m are due to the maimer iu
which iu livio;: biuin 1.^ -'> . in i.-arryin;.' iin lUi omlinui-d building up and
bmikinii diiwn, di-|iii«a* of itiw:lf. iin linid. and ilti priMbn^ls. These arv fur a
■ hil«i to arranged M to lafm a cnlorlivs iniu« with niinuln colorltna soli*)
(lartjclra or c»l^csa vacuole* Initieddn] in ii, tli<- whole having a granular
appaftnUMv- Afler a while this granular-lnokine protoplaum I* in laqt"
OMMim gnuliially rvplaced iiy tnalcrial of dillvrcnt optical uml chemical
cbaract^w. being, fur inrtnnce, more homoenMous and lcw"^nanutar" in
appearatKe : this new material i* stroma, and as it is formed, there it funuod
with it, and in SOUa way or another held by it, a ooloriiif' matter, bamio-
KliAin. We cannot at piwent say aoylbinc defioite as to the way in which
and |1m> itefie by which the oriffinal pnilopbfm ii thus lo a large ctlcnt
diArantialM] into stroma and hmnoclobin. All we know is, that the exist-
of what we have calM livinj; •ttbstanoe b nectasory to the formation
atrena and hiemo|{lobin. We, therefore, seem juntiliiil in speaking of
Bvinjt Mibstant-e as manufaotnrin;: thoe aubalanoM. but we du not know
ln« living sul»tanc« tnras itself, so to apeak, iiilu stroma, or hoetno-
60
BLOOD.
globin.or both, orwbotbkrbyHmcsKnCT, tlii' iinlnn! of which i* at prcufciit^l
unknown to iix. it cnovtrU some otihr nuiti^rinl wliicli i« prcwtit in the |
C>lo|ilium, and which wc may rvgiint lu fwM\ for itself, into oue or olhcr or
h uf tliCTc b"idit'-«.
When this ilitlorcniion hiu uk«n pbuv, or while it » stjll gotnc od, Ute
matcriHl in which the nucld an imbodded diridc* into te]>itratv cetl-bo<l!M
for ihe eevernl nuclei ; and thus the n«st of nuclei is traiigfornied into a
group of niicIcal(Kl red corpUBL-les. each corpuscle coDHieting of a uucletw
imbedded in a ha<nia};lobin-holdiDf; atroiun to which is »till attached man or
lesB of the oriKinal uodifltrenlialed protoplasm. ^
8tiU later on in the life of the «mhrvo the nucleated ml oorpuavles *reH
r^laced hj- nrduiary red corpuscles, Uv n on -nucleated diaca coinpoaeil aloioat
exclutivelv of htcmoglobiuholdiDi; slromn. Ilovr the traotifuriiiution tnkus
place, nud^ e^jieoiully how the the nucleus corun to be abwnt, u at pnveiit n
matter of couniilerable dispute; Uien- is much, however, to be (aid fur tlie
view that the iiurmnl red ciirpiiscle i» n portion only of a ell, thut it in n
rrwment of cell suhniance which hav been buddcil oil' and sn haii Icll tin;
nuiileuii behind. ^
In the- ndult, as in the cmbri-o, the red corpuKcles appear to be formed oat |
of nn7(H-diiig colored nuclenlea crllt>.
In the interior of bone* is n pci-iilinr ti«tiic called marrow, which, in most
pari* being very full of blo(Hlvc*<<!|-«. is called rnrf umrrou-. In this rwi
marrow the capillaries and minute veins funn an inlncate Inbyrinih of rolu-
tirely wide paoages with very thin walls, and through this Inhyrinlh tho
How of blood is ci>mparatirely slow. In the passages of tliis labyrioth are
found colored nucleated cells, thai is to say, cells the cell substance of which
has undergone mom or Ices diSereniiation into hwmoglobin and stroma.
And there sceiDs to be going on in red marmw a multiplication of such
oolorad nucleated cells, some of which tnu)i>foriiie<l, in sume way or other.
Into red uou-nucleat<><l ilisua, that b, into ordinary rc<l curpusclea, pan* awny
into the general hhwul ciirr«-nt. In other wonln, a formation of red i'»r(>t>s-
cIM( not whoUv unlike thai which tiiknt plaoc in the- cmhryo, is in thi^ iidull
continually going on in the red niarmw nf (he liuniv.
According to flome ulnorvem tbi- ciilor^'d nucleated wlls arise by division
in the marrow from colurlc^s cells, not unlike hut probably diilincl in kind
from ordinary while corpuscles, tlic formation of biomoglobin taking pl«e«
subsequont to cell-division. Other ohsorx-crs, apparently with reamin, urge
that, whatever their primal origin, these c»1ore<t nucleated cells arise itiirinc
post-em bryooic life by the division of previous similar colored cells, which
thus form in ihe marrow a dislincl class of cells continually undergoing
division and thus giving rise to celU, some of which Iwonme i«d corpuscles
and pass into the blood stream, while others remaiu ia the marrow to undergo
further division and so to keep up Ihe supply. Such repeatedly dividing
oells may 6tlr be called htematobltuU.
A «imilar formation of red corpuwlea has also been described, though with
leas c%'ideiic<>, as taking place in the spleen, ngtecially under particular
clrcumstancm, surh iu< Hfler i;i^ut loss of blo(Hl.
The formation of red eorpu»lw i*, therefore, a t)>ecial procen. taking
place in !>|>ei-ial regiims ; wc have no Mtlsflictory evidence that tbc ordinary
white eorpuseln of the blood are, Ms they travel in tbo current of
circulation, tmii^formcd into red corpuscle*.
Tho red corpuscles then, to sum up, are iix-fiil to the body 04i acominl of
the htemoglobin, which ronslilutes to nearly the whoU.> of their solid matter.
What funciiiins the Bironia may hnve besidn the mere, so to speak, mechani-
cal utie of holding the luemojjlobin in the form of a corpuscle we do not
I
irith
iilara
TUB COltPUSOLKS OF TUR BLOOD.
61
K
kawr. Tbo prinurjr uw of the bn-moglobio is to curry oxygen rrom tl>c
UiBp Id the toauM, aod il would appear tliftl it ie mivtmitiKeout to ibc ocoif
«aay ihat th« hsMnoclobiii ebouid b« ss il ir«re boltled iiji in corpii«dc8 rittWr
ilaa timpir difflufid tlirougb (be ploAtn*. How Umn it cor|Mi8ole nii>y livv
tmnylnc osyjreii we do nol exactly know : the red cor|iuscl«a of one aniraiil,
r. y.. A hir^l, iiiJMtol into ibe reoMk of nDotli«r, e. g.. a mflininal. disappear
within a few days; but this aflbrds no roesaure of tn« life of a corpitMle in
hi ova bome. Kveutually, however, the red corpuscle dies, iu pinec bting
MpplUd by a new one. Tlie hieinoglobin svt fVe« from Uie dead i'nrj>uBdea
a|Hiwii to have a Kcondary use in forming the pigment of the bile and
{Mfldblr other pigment*.
The n'hde or CtJarteu Gnjnuttff.
t 88. Tbu white corpusol«s arc far l«aB numerous than the red ; & spect-
mra of ordinary healrhy blood will contain several hundred red curpnsclea
In rmeh white curpuscle. ihotifch the proporlioo. even in health, varies coaud-
«rably onder different circuutslaoces. rHiiKinjt from I in 300 to 1 in 700.
But iboujrii )<!» numeruus, the while i»r|tii<<.'lefl are {>robably of greater
iaipavtaorr to the blood ilM^lf thim are the red corpn^Ies; the latter an
(ftM^ Umiteil to liie special work of carrying osypti froni the ItiUKS to the
while ibe ilirmer prolmhly enerl a oin^iili-roEle inlluence on the blood
Uaelf, and help tn muiaDiiii it in a prti[ier c<in<litiiin.
Wb«D Men in a normid condition, and " at r»t " the white corpuscle is a
«ll, spberical, colorluw mow, raryinK in siie, hul with an average diameter
of abMI 10>. and prrwmio^ jtencraTly a lint-ly but ;«imetimea ■ ooartelr
^tmnular appcanuici-. [Fig. 9.J The surlincc. even when tbo corpoiolc ti
fWta.9.
O
I
^4
«. wbUtcnnmirtnor kiinwnMond; il. red roryiBolio (hl(h |n««r}.)
pHftctly at rR<t, t» not absolutely smooth and cvrn, but lomewhat irregular,
thvaby coniribuiing to the granular appcsmmr: and at times thr»c irrcgu-
lariUaaa ar« exacgcnitixl into protulwrenccs nr "pwiulo|KHlia" of varring
rise »t form, iKe corpuscle in this way asmmin^ various forms without
^•Ogiug its bulk, and br the nssumption of a senos of fomu «hining its
pWc Of iheae " amwbold moveinenis," as they are called, we iball have
ta spekk later on.
In oarryiug on theoe Brmeboid morctaents the mrpttscle may transform
ilsalf AtKO a spherical mass into a thin. Hat. irregular plate : and when this
cecan there mnr be seen at tintes in the midst of the extended finely granu-
lar RiMa r>r rW/ Oddy, a smaller bwly of different aspect and refhictive (Mwer,
the Nwe/nuL Tlie normal prcaencie of a nucle<i!t in the white corpuscle mar
■lao be shown by ircntiug the ciirpusele with dilute accctc acid, which swells
op aad renders more transparent the cell body but makes the imdotu more
63
BLOOD.
refniclivi.' an<l mora Kliarpl;r drliiiiH). imd »> iuok iitiupii-uouK, nr by llie ti*»
uf Hininiiig KMgenU, the niinririly i>t' nhicb iiiain ihc tuioltii* niort- ri-mlilr
ttud luuri- deeply lliiiu ihr nrU IkhIv. In whnt jxrhapw may lie CQtii>i«l<^ri->l a.
tvpivul while curjiiiK'Iv, Ihn iiuclvu* w a i>pboricii] ninw about 2-3 f in
diameter, (Hit it variiii in aite in diflervnt ixirpiMicU^. and not iin(W-i(\ieatly i»
irregular in fomi, at leiW after lh*t nction of rciigt-nt*. It oci-iwionnlly
■I>pcan> ai if obuut to diride into fmginrnto, imd soDiclimcg a (7nr|>u»de may
cunliiiii two or rrvn more (then gi-ncmllv >niull) nuclei. Though tlainiog
rMdily with Rlaining niigcnlv. the nuclcui of nii onlinar^' irbit« cormisclc
iloi-B not divw the nuclear network which u w cluniclcrialic, m ne »hall mv,
of tbo Duclej of many eelis, and iihieh in tbew i« ih^.- ]>arl of the uuc]«us
which especially stains; in the clwely allic<I lymph corpuftclc*, to which vre
shall have inimcdiaiely to refer, a nuelcAr network is {ireacnt.
The cell bo<ly of the white ciir|ni«cle may be taken as a good example of
what we have called undi Re re minted protoplMm. Optically, it con^ista of a
unifonnly transparent but Eomewhat refractive material or basis, in which
are imb^ded minute particlee. generally spherical in form, and in which
aumetimes occur minute vacuoles filled with fluid : it is rarely, if ever, that
any distinct network, like that which is itomelimes obwrved in other cetla,
can be seen in the cell body of a while curnnsele whether Gtainedor no. The
Imbedded particlee are generally very email, and f>>r the must part distributed
uniformly over the cell body, ^viu)> it the finely granular atipecl spokea of
above; Mtmeliuiee. however, the particles are relatively large, makin); the
oorpuaclea coarsely ^^raunlnr. the ooanejmnuleii \mi\^ l're>(iitntly CMifiiied
to one or another {Htri uf the cell body. Titcse partiileF or granule:}, whether
coarw or fine, var>' iu nature ; some of tlieui. as shown by iheir greater
refractive power, ihetr >iaiuiDg with osmic acid, and Ibeir wilutiwi by miIv
«Dl( of fat, are fattr in nature; others luar similarly be vhowii by thcirS
reaction* tn he pmteld in nature. ™
The material in which thcw granuln arc inibetlded, and which fbrntf iho
greater jiarlof the <-ell bualy. ha» notpeeini optical features; xofar aacan b«
ascertnimd, it apix-am under the micrM>ro|>e \u be ho<nog«-nc<His ; no definite
structure can be detected in it. It muKt be bnmr in mind that the whole
corpuscle connsis largely • f water, the total mlid matter amounting to not
much more than l<i per ctaL Tli« transparent material of the cell body
must, therefore, be in a condition which we may call fwmilluid, or semisoliit.
wilbout beinc called upon to defiao what we eiactly mean by these lemtt.
Tbis approach to fluidity appearv to be oxinertcd with the gml mobility nl*
the c«ll bmly. as shown in ils anxrhoid movements.
$ 26. Wlttn we submit to chemical examtnalion a sutKcieni mass uf white
corpUKlee. «paraied out from the bl<<wl by special means and obtained toler-
ably free ftvaa nd coqiuackB and plasma tor apply to the while blood-cur-
piudM tbe chemical r«sulls obtaiDM ftvra the more easily procure*! Iym]>h-
corposelea, which, aa we shall aee, are very similar to, and, indetd. in many
wajra niated to the white eorpiuelea of tbe btood i. we fin>l Ihul this amaJl
mImI matter of tbe corpuscle coomMs largely of certain pnxeidt.
One uf theee |)rDtd<U is a body dtber ideniioal with, or closely allied (n,
the pruieid called mjnMN. wbicb we shall have lu etudy more fully in oon*
i»ertiua wtih muscular tiMut At present we may umplr aay thai mrcttn !«
a body intermediate between fibrin and globulin, bein^ Icwt iviuble iluin the
latter and more soluble iban the former ; thus while it tr hardly at all wiluhlv
is a 1 per cent, wlution uf sodium chloride or other neutral Nilt. it is, unlike
fibrin, spctdily ami wbuUv dimolred br a tO {wr nal. mlutivD. My(«in is
fbrtJMf iniervtiae beoitiM, as we shall sew, ju>t a> fibrin is formed in the
douieg of bluod tnu fibrinngan, so mjixin it formed uut of a prvenliDi;
TBS CORPUSCLKS OF TDK BLOOD.
I
mjMiiio|[«ii, iltirinK m kuul of cIoIUuk wbidi lakes plnco Lu iiiu»cular fibro
wd vkirh U itiukeii of a» rigor inorlia. Aud w« bavv rmituii* fur ilihikioK
tKst in tW living white biMiil-coriiuMic Iberc dott exM a body idtnitciU
Willi or kJli^'d tn iiijrudiDiigeD, wbicn nc mtty *pc»k of uh Iwing in li fluid ena-
ditioB : and whicb nti thr dcatb of lliv ttir|>*'*c'*= 'i cnnvi-rlrcl, by ■ kind of
eloUlafc iaio myoain, or into nii nlliail hmly, wbii-h boiug wlici, give* tbs
bodj « the corpiucle a »lifliii-w mul rigidity wbtrh it did not ponoM during
BmUm thh mrmia or Riyoein-liko protrid, tbc white corpunclce nUo con-
tmia nlhtr parngl'ditilin iiwif or some ofh^^r member of ibe globulin group,
■* well u n tKMly or b'xliv* like or tdentictd nith Mruin-nlbuniin.
lo Midiliun, tbt-rr is prciwiit, in somoirhat c»nsideniblc quiuititr, h Biib-
fltaiK* of m [peculiar naturv, which, sinoe it it confined lo the niiefe! at lite
CM|»iwelw. Mid further ^eems lo be prewint in all Duclei. Iihh been called
mmtUim. Thit uuclein. Khicb lhou);h a eoiDp]«X Dttrogenmia body h very
diflhnal in rompoeitiou and nnlure Irom proteids, i> renitirknble un the on«
bani for being a very stable iuert body, and on the other for eonlaining a
brg* qoaniily (aceording lo BQue obeervers nearly 10 per cent.) of phos-
pboma. which app«an to enter more cIokIj into ibe ctmclureof tbo nM>l«cule
Una it does in ta« caee of prulei'Is.
K«xt ID importance lo ine proldd», os constant constituents of the whitfl
corpoacloi. c.x>Rie oerrain fala. Among theae the nioflt couBpieuous is the com-
plex falty body kcHAw.
Id tbe uueof niany corpuscles at all eveiiU wc have evidence of tbe pres-
^KC of a nieniber of the large rrmip of car)>uhydraie». comurising slarcliei
■ad njpir, viz., the tturuh-likc body gli/aigeM, which ne shall have In study
man fully hemner. This glycogen may ext^t in the living corpuscle a*
gljrtngvn, hut it m very apt, aAor Uto dcatli of ihe corpuitclc, to become
cfc— grd bj h) dnuiou into roma torn of sugar, nicli as nialtoM or dextrnee.
I itli the tub of the white corpuacles is chiiracteriu<l by coataioing a
frlativvly large quantity of |>otnseiuni and of phosphates and by biHng reln-
tiralT pinr in chWidea and in sodium. But in this respect thu corpuscle is
mmij an example of what weuiH to be a general role (to which, however.
llMnrw tnay be exoeplMMia) that while tbe elements of the tissues themwlvcs
are rich in potassium and ph(*)phat«fl, the blood plnsma or lymph on which
ibey live abounds in chlorides and sodium salts.
i 30. Id the broad features abuvc lueniioned, the white blood -corpuscle
may be lakeu as » picture and example of all living tissues. If we examine
llta bisbilogica] elemenls of any tissue, whether wv take an epithelium cell,
or a nerve cell, or a cartilage cell, or a muwrular libre, we meet with very
■ittiibtr feaiurea Studying tbe element morpbologteally, we find a nucleus'
and a irll lioily, llw nuHcui having the general cbaraclera described above
with fmjuetilly other charactvrw iiitniduood, and the cell body cunsiHting of
■t IcMI more than one kiml nf niiiteriiil, the mulerialH being aoroeiimea ao
dinieard ■> to produce the npiiciil i-tTtTi nimply i>f ii trauHparent moss in
■rbidi granule* are iriil><^lili.-d, in which caM- wi- niieak of liie cell UkIv us
pretop&niic. but at «[hi-r limiv f iirrnngnl that tlie oil Wly |iodfiesaes dif-
wmllated structure. Studyiuf; the clement fmm a chemical iwint of view.
W» tad protcid* alwny* prcMnt. and nnioog lh««e bodies idenlicAl with or
atm or tess closelv iilliiil to niy<«in, we generally have eridence of Uw prce-
«Dre alsn of fat of'winie kind anil of fiioe memltnr or members of tbe earbo-
liydrate gn'uji. niwl the wh always cunlains potaiwium nitd |ili<M|>)mtfii, with
•Blpbatae. chloridu, sodium, and cnlcium, to which may If adtled magnesium
au iron.
■ Ttw mlmnw rf ■ultfniiclnu ttrusmwi do» wnl tWnA tlw |i»i wit wyiim»nt.
64
BLOOD.
We sialwliu the lutroduction Iliat liring matter is alwsya un{ieT;gtNii
clieniiciil cbaDjte; (liia continued chemical dianiice itc tntty ileiiote by the
tonn nietabolirm. 'iVo fiirtlier urged tluit as looe » lirins muter ii aliv«,
llti^ (-beiukal clituit(e or luelabulttin u of ft double kiud. On tlie one band,
till- liviii); NubntniR'f Jit CMiliiiually breaking ilimn iiiti^ rimplcr bodi««, with
a nutting froc of t.-uergr ; this jmrl uf tht- miTiiihoU^m w4t may apeak of lu
roftde up of katabolio chaTigi-N. On the other baiiil, the living •iib^taDoa ia
OontinuBlly building ilvcilf np. I'mhiHlving ciivrgv into iim-ll' and ho rcplcB'
Hhing it* Htorv of riicrgy ; thi» part nt'' the metabolism wo may rpcak of at
iDiide up of nnaltolln chang<3>. Wo alw) urged that in every picpp of living
tiaiUfl there might In' < 1 ) tha actual living Kubtslance itaclf, (2) matoniil which
is nn«cnt for the puriiose of becoming, and is on the way to boeome living
Butistancfl — that is to snv, food unilergoing or about to undergo anabolw
vhang««, and (A) nintcrial irhich ha» rcaullod from, or is reeuliing from, tba
breaking down of the living substance — that is to say, material which l;aa
uudergone or is undergoing kalaholic change;, and which ne speak of as
waste. Id using the wor<I "living subetance," however, we must remember
tliat iu reality it ia not a Bubslance in the chemical sense of the irord, but
material undergiiing a series of changes.
If, now, we lutk the <)ueeuoD, which pan of the body of the white corpuscle
<or of a fimihir element of auother tiai^ue) U the real living suh^tance. and
wbieh jitirt U ftiod or waste, we ask the question which we cannot lU vetdefi-
ni[«lv anawer. We have at preM^ot no adeijitnte mornholozlcal criteria to
enable ua to judge, by optical characters, what iji really living mid what b
not.
On« thlDg we may )ierbajie stay ; the nmterinl which appears in Uie cell
body tn tbe form of distinct giiuiu leu, merely liHlgei) in the tnorv tnuisparcnt
ntatcriitl, eunuot be part of the txmiI tivttig substance; it must lie cither food
or waste. Many of these graiiulei' arc fat, nnd we have at times an opportunity
of observing that thcr have been introduced into the corpuscle from the sur-
rounding plasma. The white cor|)UBclc, as we have said, has the jM>wer ef
executing amcelmid movements ; it can creep around objecia, envelope them
with its own substance, and so pot them inside it^lf. The granulea of fat
thus introduced may be suluequently extruded or may disftp]»tiar within the
<XHpu8cIe: in the latter case they are obviously cliaiiged. and ap|>arently
made use of by the corpuscle. In other wunU, thW faity ^rauulei are appa-
rently food material, on their way to he worked up in the living substance
of the corpuscle.
But we have also evidence that dimilnr granuleti of fat rwv make their
appearance wholly wilhiu the oorpuacle; they are priHiocta of the activity of
the ooqmsele. We have further reason to ihiak that in some i-a-*eT<. at all
events, they arise from lliu breaking down of the living Huhntancti of the eor-
puBcle, that ihey are what we have calliH] v/wtU: prntluct*. ^t
Rut all the granulcK vJnihle in u cnrriiiHcIe are n»t neci^warily fatty i^l
nnturo ; aomo of them may undouhleilly he pniti^id gritnules, und it i* )>os>i-
blo that some of tliem may at timoi he of carbohydrate or other nature,
all eases, however, they are either fond material or waste products. An
what i* true of the easily distingui«hed granules is also true of other sul
•caiicat, in solution or in a nilid form, but so disjiosed as not to be optically
teowniied.
Hcoce a part, and it may bo no inconsidemhie part, of the white corpus-
vie may be nut living Guhstniico at all. but either food or waste. Furtl>er it
does not oeoeesarilr follow that the whole of any quantity of material, fatty
or othernbe, introduced into the corpuscle from without, should actually
l>e built up into and ■o become part »f the living subatance; the chau^
u9
Tirx c.'OBPi'act.Es of tiik blood.
65
\
■at raw fnod to livhig (ubclance are, u we bare already mii), prolmiMy
. u)<l it inaT Iw ibat after a ceHain number of cluingeit, frw or mnny.
•nljr of tbe material it accepted aa wcjrthy uf iKing niu*!!; alivv, anil
ml. beiDf; rejected, beconia at ODr« iradie matter ; or the malarial may,
a aftrr it lia* under^oue this or ifaat rhon^, nervr acltially enter into
tbr liriojt aubMutoe but all become wai^i' inaitcr. Wo miv wa*tc matter,
but tliia (Ion not mean umIcm auller. Tho mallrr m formnl may nilbotit
niarinit iutu the living mib>4aD0i! be of lonie ^uWidiarr use to tbc corpiiecle,
«r M (NTobohly morv otltn bappomt, bcios diwhaiwu from tho cnrpuwle,
Ba^ lip of UM- to Kime other part of the mdy. AN e do not know how the
Uthiie •uh*taDi<e bailds ilMlf up, but wo aoem <>ompcll«l to admit that, in
(•ttaia csfn at all cvi-nls, it is abk> in some way or other to produce
rkiK)^ iin maleriid while that matrrlid is Mill oiiUide the liviiif; eulwiance
M it were, before it enters into nnd indeed without ile ever actually entering
inlallu! cotnpofilion of the livio^ fiub«tiiHce. On the other hand, we muol
nptiOy admit that tome of the wnnte friibetances are the direct pruducia of
kaiabolic changes of the livingt subelance il«elf— were aL'tuallv oure |>iirt
'if lirtn^ mbetance. Hence nc uu^bt, |M>rbap«, to distiaguiJi the nruil-
nf the aciiriiy of living matter into waste pruducta proper, the <lirecl
of talaUilic <-hhn]:ni. and into hyc produda which nre the rcHullii of
dfircCed bv the living nmtli-r oui^tdi' it^ll'and nbidi cannot, tbore-
eODiiilercQ an Dcvrivarily i-itlivr iiimlHilic nr katatiulic.
ine llttebemical chanclcni of Iht' living matter itielf wo cAimot
_. make anv very dvlinit<> »talemcnu V,\: may av lliat the pmteid
>. or rather t^r protcid nntrcnient i>r unli^colent* of my win, enter in
■ay into iia stniclure, but we nre not jiiMiticd in Ntying thai the living
labiluce cMiaisIs unly of prnlcid matter in n peculiar condition. And, in-
tifcr-' '•'■ -'-oiitency with which aoiae reprewntalive of fatty bodies, and
w- iintive of carhuhyilnites always appear in living tueue would,
pnitij-. niiinrlrnd u« to m|i[Hi»e that these equally wiili proteid material
*r» tanniinl til itM Htructure. Again, though the behavior of the nuck-ue
m oiDlraati-d witli that of the ceW budy, lea<U u* to Mippote tlmt the living
nhMaara of ibo former in a itifrmnl kind from that of the latter, we do
BM ' "'tly in what iIk^ diU'iTt-nco coiim*!*. Thti niiclFiii, w we have
•K. nvctfiii whiih. peiha|t», «c mav regard w a largely modified
pnu-t . l-iii htfiuff a body which it> reinarkablf (or il» ftability, lor the diffi-
I fall) oith whii'h it la changed by ehemicnl rcngcnt». cannot be regard«il as
■I mnal ftut of the ecHenlially mobile living »ibslnnce of the nucIeuB.
Ik !&■ nmneetion it may be morth while to again call attention to ibe
kl tW the «or]:u«-U- contains b very large quantity of water, vix. alxHit
MftrcraL ftiii of tbi«, we do not know how much, probably exists in a
■Mcorin* definite combinaliou with tho protoplasm, siimewbal after the
■MBcr nf, lu u«o what i« a mere illustration, the trater of crTstallizaiion of
Mkib If we imagine a whole gmu^ of dillbnnt complex salts continually
<gwfinl in (uni in being cry«lalli»d aitd being decryMalliied. the water
lkM«Bf*getl by the *allf will give UM a rough image of the water which
MM! a and uul of the sobftance of the cor|iu>clc an the reault of its meta-
MBcactiriiy. We might cull this " water of metaboliiini." Another part
<^ Ifca water, carrying iu this esw suliftnni-eii in solution, iimlwbly exisia in
ifarmoT (Dlerriicee too small to be seen nith even llic hiKbcxt powers of (he
aitnacnpr. Btill another part of the water similarly hutdiag subMaoccs In
mtaiiim cxbl« at linKs in dt-iiuiie spaces visible under the mtcroaoopo, Dton
•rim rvf^nlarly spherii-ul. and called vacuole*.
■ WakBTed«rlt thus at leugth uu the nhite cirpUK-h) >d the fir«t plaoo
Baesna as we Imvo already eaid what takes place in U is in A atnao * picturo
I
6S
of wliut takes ;i1ni-e in *\] lirin;; elrucliim, nn>i iit ihc et-oiKl ploop
ll)L' fiic-U whioli wre linrc mefilioDod help uh (o uiiilcmKifl how the
ciirpiiM'U- may carrv on in ilic blond a n-ork of no im|u>riunt kiuii ; for from
wliat linn iieeii aitid it in obviotin itinl the white curjKi^cIo is OJUlinunllv itrt-
ing u(ioi) mkI Iwiti^' iiaed upon by ibe plasma.
$31. To II II lie ret and. however, the vriirk of these while ror|>uH)ea,
inii»t Ii^tni nbot U known of their historv.
In »iiPci.-Mivc droiM iif bloml taken at (iiflerent times fW>in tlie same ini
vidunl. llic number of colorlew coqiusde.i will be foiiii<l to vary ^t-ry luuc
not only nOtlivdy to th<! red coqiuvoles, l>ut bIsu ab«i.>lut«]y. Th«y iiiu
tbereforo. " come and gti."
In treating of ttit; lyinpliiitic *yiit«m vc fhall hnvc to point out tlini a
very largu iiimiility of tliiiii enllud Ivnijili, ci^tiliiinint; a vnry (,i>n«idrnible
number of bodi(»i, very similar in thvir gmrral chiimctcrg lo the white cor
]>ufic]eB uf the Mood, is being riinlinimlly poured Into llio vaxeiilar erntcm «(
Ibe point where llie ihoratic durl ioinn lliu grc«t veiu« on the left ai\v of the
ueok, and tv a k^ra exlrnl whcir llic olhcr tnrge lymiihaties join the venoos
■VHleiu on tho right eiiie ol' th<- nock. Thetw corpuscle* of lymph, which, as
we have just said. ek«ely rc«cnib)Ct and, indeed, are with difficulty distin-
guisbeil I'niiD the while corpu^cloa of the blood, but of which, when they
exist outside the vaHcultir eysleni, it will be convenient to speak of a^ Intro-
eyttM, are found along the whole length of the lymphatie aystem, but are
Diore numerous in the Ivmphfttic veesela aller theae have paaaed tlirougfa
the lymphatic glands. 'Theae lymphatic glands are partly eompoHed uf
whal '» kuown as adenoid tissue, a special kind of connective tatue arranged
an a dc-licaie network. The inemheH of this are crowded with oolorlesa nu>
clr-atcil cells, which, though varying iu slxe. are fur (he most part •mall, the
nucleui) lieing surrounded by a relatively small quantity of cell iiilwUtiicv.
Many of thv-w; cells show signs that ihcy are undergoing cell -4)1 vision, and
vre have rcuMnn to think thai ivlls so formed. a<-(jiiir{ng a liirger Hinoiint of
cell substance, become veritable Iciicjcytw. In other wur«ls, lencocyt**
multiply ill tlic lymphHtic glandi!, and leaving the glands by the lymphatic
vessels, make their way lo the Mood. I'att'hes and tracts of similar adenol^
liasue. not nrmnged, however, us distinct glands, but similarly oi'cupie^l b^
developing leucocyte* and similarly connected with lymfilialtc veawls, are
(bund in various parts of the body, especially in the nmeous inombraaes.
Hence. W'e may conclude that from varioua parts of the bmlv, the lyni-
pbatios are continually bringing to the bhiiid an abundant supply of leiico-
cyi^a, and that these in tbe blood become urdinarv white 4-orpUi>c1t«. This
is probably the chief source of the white corpuscles, for though tht* nhite
our|>uscles have been seen dividing in the bhxHl Itself, no large increase takes
place in that way.
£32. It follows that since white corpuscle:- nrr thus continually bciDe
sduod to the MoihI, white corpuscltni must as continttiilly either be ilcstroycu,
or be transfonncd, or mcsjh^ mini the interior of the blood vewels: otherwise
ihe bloorl Would soon be Mocked with white corpusi'Uw.
Some dn leave the bhimlvi^Mcl*. In Ircntiiii; of the circulation wc sball
have to ]K(int nut that while eortiuscles an- abb' to pierce the wall* of the
capillariiv and minute veins, nno thus to mnkc their way from the interior
of tbe bluudvnw'U into K{iaces filled witli lymph, tbe "lymph spaces," as
tbey an called, of the tiwiue lying outside the bloodvessels. This is spoken
of as the "migration of the while corpuscleH." In an "ia6aiueil area"
large nuiul>cr« of white corpuscles are thus drained away Uom the blood
into tie lymph spaces of the tbsue; and it is probable that a similar Icaa
takes pUoe, awire or less, under norma] couditious. lliese migratiug
se migrauug oc^
TUB CORPUSrLBS OP THE BLOUU.
67
I
f roDawiDK ibe d^-vioiii tract* of tlic lympli, find their wny
bluod: toiDC or I hem, huwovcr, may reniuiii, and uridi.-ri^i
vafioo* ebanra. Thus, in intlamcd iireni). when »u|iMinili<)ii rullowii in-
flaaattlioa. t£« whil« <»r|iiiM>1c!t which have niixrsied may Wi^itm' "ihi»
OMpOKlet," or. where thickoniiii; nud Kr<"*'l> follow ii|h>u ii)flnmmnli»ii,
OMT, BflnQrilini; to uiiiBV iiulhoritleE, h6is>iii« tmmrormed intn (em|iiirnrv ur
pvnnaiKnt tuMiu. ni|x'rin]ly cimncctivt- tlwue ; but this trantfitrmiitioii mio
dtm b dlifiuicd. Wh^ii nn ititlBmnialiiiii sulMtdett withoni U<nvirig any
dbct ■ few CT»q>uwl«i only wilt Iw found iu iho tinuo ; IhuM which had
prwiottily mignU'd Diu*t, ilicrefuro, have lii:rn dii>[iijw>il uf in fonie war or
MMtiMr.
In iMBkbg of the forniniioD nf rod corptucU-s ($27) we mw that not
oaljr it la not pruvcti thai tli<* iiui-lcatcd oarpturclcs wliich give rJM to red
VorfNncka are ordinary nhitv cornii»cle«, bnt that in all probabililv Ihe real
banwtobluiv. the pnrcnu of rwi corpuwJcs, arc spvcisl corpuscW devel-
•|icd in Hue »ituation]^ irhcro Iho niiimifiiclura uf rcl corfiuscles takea place.
iio 6u, tbcrtri'ore. from an»uminK, at is eomelirncfl done, that ihe white car-
ricl«a of llie blood ut« all of tnein on their way to become red corpiisclea,
may Iw duubtei) whetlier any of ihem are. In any cii*e, hnwever, «rei)
■•ktnt; atlowaa<« for thotw which mit^rate, a very coitiderable uumhcrof
tb« whiu oorpuaeUa matt "disappear" in some nay or oilier frum ih<' blood
Unam, Will in may, pcrhap*, siieak of tiieir di«tpj>eiinin<'e an being n
•* dwintction " or " dlwolution." We have, us ret, no exact kuo<rlei]gn to
■•iiiiti- ti* in tliv tuaitter, but ne cait readily imauiiie, tliiit uuun the death of
'j'liaclei, ibc •ulwtaneea >'iinip(Ming it, after unilergomj^ chaiigra, are
.._... i^l by and become part uf the planmiL If dc^, the ct>rpiiiicU's a« they
di* noat t«i>eale<tly influcooe tlw coni|Mk<jii<iii and nature of the plnsma.
But if (hey thus aliect the plainia in ihcir death, it is even more proba-
ble that ihev influence it during ihoir life Beinj;; alive they must he cou-
tiimally laliiuf: in and giving out. Ah we have already said they arc
knonn to iu;;e9l. after the fiLthioii of an amaba, solid panicles of varJou*
Unda, Micb a» fat or eariiiiac, pmeut in the pliuma, and prolwbly digcut
Mda of thno particlca u are uulrilious. ItuI if ihev iu(Cfi»t iheM xilid
etatUn ibry iintlably abn carry out the easier toak 0!* inf(e«ling diawlvud
BtatMca. li^. however, ibi-y thua take iu. they niiist abo give out ; and Ihua
br llw rvmiival on the oue huud of variotin niil»lanctA from the plainui,
aod bj the addition ou tite other hiind of othrni. they niuat be iMnliually
tafliMBdnK the plaama. We have aln-iuty :iuiil that thr nhite oorpuH-lr* in
lfc»d U4iad a* ttiey dio are aupiXM^d to pliiy un imji-irtnnt jMrt in thv clot-
ifnif of bkiod : Biniilorly they may during their nhiJc life be engngiHl tg
t^rr*''"' ■■•<\ rhangn in the pmleiilit of tlic plutnin which ihi not lead lo
di-' which prepare them for their vnri<m» uh'^ iu tb<* body.
r,>.u-..--h.<^«l facia alforil Mi)p|Hirl to Ihii' vivw. TbcdiMMue callmJ leiicn-
ntbcoiia (or Inkieniia) is chamcli-riccd by an incivaev of ibe white corpua-
cin, both absolute and relative to the reil cor|>ii»claa, the incrvaae, due to
■a sai;nenl«d production or poosibly (o a rrtarded dostruciioti. being ut
tinea m xnul ai to |.iv<> thu blood a iiinkieh-gmy nppeaimnce, like tlut of
Uoad aispd with pus. We accordingly liml that in lliia diauw the plasma
if to Rtaoy wavH pr<ifoundlv alfevt«d and fails to noiiri^ the tisuea. Aa
a ftirtWr illui-lratiun of tbe pomble acltou of the white eorpusclca we
mmf (talir tliat, acciinliiig lo ^>me ob^rvers iu cert^iiu diseasea in which
•imt« orgnnumi' micIi ns bacteria make iJieir api>e«>auoe in tbe bli>od,
Iba white mrpuMrlra " lake up" ihew baderiu into their subMaaoe, aod tbaa
pvMbahiT, hv irxeniog tin influence on ihem, modify the coarse of tbe dbtan
«f wb><^ ihoe organism* ara tl>e ««sealiu) cause.
BLOOD.
If ttiu while uorputcJcs are thus eugugtd during lli«lr life in carryiii); fM
iaiporUUit Inliora, vre may exiiert ihcm to diiTi-r in a[>))eHrance acoonjiiis to
their CQD(liti»Ti. Some of tliu cur[»uid€M arc iipikcn of lu " faintly or
*' finely" graniilnr. Otlior MimiM^lm are niwkwi nf tut" euanfcly " gmaulir.
Iheirooll Fiibntance bcinu l(M<Ivd with c»i)iipinioii»lv diicrfie graoults. It
may bo, uf coiinfc, thnt tncra nru tw» di»iinc't klDcIs of corpuitdee, baving
dHfcmit fttnctioiJH ami pmrnihly diflVrent origioa and histories; Ixit sioco
intcnnedialc fonnii nrp mtt with ontoinine a lew cunru gniniiln only, il it
more prohnb)« timl llio otic tVm ix n pbaso of the olbrr; ihni a faintly
granufar corpUHclp. by laklog in granuks ftom without or by producing
granules within itself ae product* i>f it." mctabolitn), may become a coarcely
granular corposde,
Wlicthtr. however, the white corpu«clc« are roally all of one kind,
whetlier they ure dilfereiit kind» jMrforming dilTerent functiona, mait-
preaent he left au open question.
Stood Plaltkt*.
§ 33. [n a drop of blood oxamiued with care immediately after renioi
may ho f>ccit a Dombor of esveedingly timall bodies ('into 3^ in diamrlpr
fircqiientlr disc shaped, but Dometiincti of a rounded <ir irregular furni, hnini _
geuooiu in apposrancr when quite fresh, but apt tn aMiinie a faintly gniiiiilar
l^irct. Tlier are culled blrivd platrlth, or itfooti
|Ra 10.
(tln'jHfu. Tni-y have been «ui>piK>i.il bv sonie |M
K'coniu ilcvi-hjpi'il into, iind.'iiidrcd, to be cnrijm
een
1
HiiigfS of, the red curpufcle:', and lienrv have
been called lucmutoblnate ; but this view Iiiib
not been confirmed ; indeed, as ne hare seen
(S 27), the re«l hse m a tob lasts, or developing
corpuscles, are of quite a difltrent nature.
They speedily uuder;;D chaDRe aOer reninri
from the hodv. apparently dissolvtug in ihu
StasTDB ; ihey hrenk Up, part of their Bubetjince
isRppeariug, uhile the r««t beconiea granular.
Their granular remains are apt to run togclhcr,
forming in the i)Ia§ma the shapeleda inawes
whirh have h-iig Wen knowu nnu descrilied as
■■ hinipx of proloplasm." Br appropriule rw-
ttgpiils, huwever, these plaieleta may he fixed
and EilaiiKst in the ctiRililion in which the^_
ajipear afttT leaving the bcnly. ^M
The aiiliNtaiire ei.ini[ia>ing them is [leculiill^
and, though we may pvrhaM ajicak of them ua
iHiOAisting iif living nialeriiu, tlieir luitnre is at
|>riiH-nt mmcure. They mayhcscfn within the
living htoodve£Bels[Fig. 1(1], and tbercfitre miiM he regarded ss real parts
the blood, and not as pr*idiict» of the change* taking place in blood after
has been ahvd.
When a needle or thread or other foreign hotly is introduced into I
interior of a hlo^jdvevsel, they are apt to collect upon, and, indeed, are the
precurMira of the clot which in most cases forms around the ne^le or thread.
They are abo found ia the tfiromf/i or pings which sometimes form iu the
bloodveiMb as tiUi mult of diseaac or ioiury. Indeed, it hu been main-
tained that what are called whiU thrombi (to dislingiiisb them from
m«i!< ilUKi-iri I'm Blriijip
A. network «( nhrlii. tkiowa anet
WMbtiiii aiaar lbs corT>uiclD» Riiui
« |>rti>*nll»iicrt blood Ibkt iMa br«n
■llcnrcrf lo cint: maniF of llin llln.
Btmi ndlHtc Ifota miitll clum|B
oftilowl lilatflcu, n iftDiu Otirrt.
Uux) cuH'UWln ukI vlciiicntury
|«l1ICta otIiIcKil plnlFJDU mltlilo
a >iiibU lelu.)
he
I
09
M, wlir4-li ftrc |>Iu^ of cnr|>iif>c)«« anil filirim are Jii rtfalhr aKi;ref;a-
tiMu of MoimI jilniolola: nixl for various r»uiiiiti> IiIlhiiI pluUileu Imve been
•Oppond tu |ilitT nn im)Hiniiiil jiiirt in tb^; clottiug of bltiod, rarrjing
Mit Um work wbifli, iii tlil> riv|>ix't, U by nlh«n nltribiiicd la tlie wbite
eorpinlw. Itut no vorr (kfiiiiii* xtaii-tiiKitt mii iit prc^iit hv- madft about
Udi: uid. [lulaed, tb« origin ati<l wb>iU- iinturc of Uhvw bloin) iiUielciU a at
pnaMil ofaMartf.
TlIK CniJIICAL COMI-OMTIOK OT Bl^OI).
1 91 We n»Bjr now paas brii-flir in review the chief cberoical clmrocK'n of
Uood, reHMinbMnitg aJway? tlint^ ns wo hitvi; nir«iu)y urgod, the '-bief I'heini-
lal iatunia uf blood ar« nUni'bed to ihv oban^^ « bicb it unilert;oo» in tbu
wranU tfaaiiw ; these will be cuoHidored in connection with each tisHie at tbe
BppT" prist r place.
T ,•(! siM'i-itic K"'vitr of human blood i» lOW, varying from 1045
lo I' < hin till- limiu of health.
Tbr rxTAction of blond M h lloirii from (be bloudreMcta U found to be db-
lioriK tli..ii;;b fifbly alkaline. If a dmphe plaoeil on a piac<! of faintly red,
ht^ I liiniuit paper, and then wiped utf, a Muc »tuin will be left
li.- It lilooil cuntaiiu a cvrinin ifuiintilv of gw^ vti., oxyt^i. car-
bonir Bciil, and nitrogrn, whii:!h iirv b4>Id in the bluoil in n [let'uliar wav,
whifh vary in itiirvri-nt kind-i uf blood, and »o itcrvu cfpuciAlIy b> dUlinj^idli
arterial fn>ni veft«u> blmid, iiiid which iiiny be given off from blooil when
esfwanl to an atnioipher*-, accord! nj,' to lh<- composition of llnit atmosphere.
Tmm gatea of blixHl wi- Fbnil ntudy in ronnvetion wilh rvapinUion.
the oamtal blood con«iHi« of c^Tpiudet and ptntma.
If llMeDr|KHelw be suppcwod to retain the ani^iiol of water proper to then).
bland iiuiy. in ^i-n«nil ti-rnt*. In- iMii>>idi'nr<i as con^istio^' bv woighl of I'roa
abmi one-third bi oomewbal lens Ihnn one-half of eorjiuiclea, the feat being
plaama. As we have alnauly kch, (he Dumber of ciirpusclea in a ajweinien
of hinod i« fr'uml u> vary ennsiderably. imt onlv in different aniuiaU and in
diAnuit inl*. but iu the aame individual at dilferenl times.
Tbe ; ' - rnudvtst by the cloltinic of ibe \>Wni into «^iim and fibriH,
I Si. The MTiiM eoniaini in HH) parts :
FrBtaM tobMiMw
hU. vMWua MtirwtivM, asil mliao oinlUn
■bnut H ort p*rU.
3 or I part.
m larta.
Tlw pnitiridi) arv pnrttgMtulin anrl >mim-tiibumin (there being probaM/
than lutp kind of M!nim*atbiiM))u I in varyiDK proportion. We miij,
p«H'*'- r-niiihly "peaking, my lliat tbey ifccur in about equal quaulitie*.
' <tis iind linking nt are the restilre of cloltinj*, niaiwive a» appear*
10 U ..'■ • -'-l whii'h \a fiirme<l, it must be remembered that by far tbe greater
(ttrt uf ibc clot consists of corfHiselea. The amount by wei;gbl of Bbrin
ranitired to bind togeiber a number of corpusclei, in order to form even
a larse. tirra clot, ta exceetliugly small. Ttiu*. the average •piantitv by
■r' " ' : 'niii in buuiiiii blood is <«id lo l»e 0.2 percent.; tbe amount, bow-
iin W obiaine'l from a given <|uautlty of pliumu variea extremely,
ti>t due not only lo circuinsCaneet affecting the blooil, but
I empluyed.
l> are Manly, excel)! attar a nical or in certain palbolngical
i I of the neutral fiiU — ileario, {lalmitin, and ob-io — with a
Y ihi-ir n -ji'Tlivo nlkaliiie woapt. The i>eculiar cotuplex
' r.' in viTv (uiill 'luantitic* only : ifae amount prcHQl of th«
(be ^:ll
■f
70
BLOOD.
peculiar iilcobol <'h<>1<«lenti , wliirli hiiK *a fiitty iin npiminiiKN!, is al»n sma
Among the cxltaclivc^ jiriwiit in .iL-rum niuy W [lut ijohd nearly nil th«
nitrogi'iiotiii nixl ollii-r i>tiii«liiiu'cii nhifli I'orm lli« i'xtriu?liv<'!> of the btxlf nod
of food, riJc'li B« itn-ii. kn ntiii, >ii^ar. Iiu'ticacid. tic. A very InrgR Diirnbrr
of tlic»c Imvt Item di#iiiv(.r<d in ilii- Moud under vitriquo cirvuiiiKtaiirn. (he
cotixidcrntioti of which niu^t lie IcfV for the prcMiil, The peculiar oddr ttt
blood or of Ecnini is prol>atdy duo In ihc prcwtire of volatile bodioB of tlic
fatty acid wrie*. The laint yellow cidor of eeruin it duo lo a K[iecial yellow
pij-ment. The tnoet charactvriittic aod imporioDt chetnical feature of tbo
saline conGlitulioii of the ecnim is the pre|ii:<Dderaiice, at lea«t in man and
most aniiiinli', of fodium mltii over those of potiuwuni. In thb mpvel the
aeruiu offers a tnoikcd conlrnsl (o the corpuscles. Lees marked, but still
Striking, is the abuodnnce of chlorides and l)i« poverty of phoepliates in the
aeiuiD, OS compared wilh the coqiufcle^ Tlie mlti inay in fact briefly b«
described ns cuDsUliug chielly of eu'liuin chloride, aitli ioiue amount of
sodiiiin carboDste, or more correctly Midium hii-arliuimte, and |ioia»iuiii
chloride, with small quaotiiien of mdiuin uilphali', Medium ohiwphale, cal-
cium idioHphale. and ninK"^>"ni phoq>hate:. And of fvtn tin- small <juan-
lity of ]ih«»phati.-!i found in the aab, part of the yhfinyhunit eiialti in (he
•erum ilnvlf, DOl as a phcitphnte, but an phiisphiiriiH in »>nu' orgfiiuic hudy.
§ 36. The rtd torpvtctt* contain le» wiili-r limn the H-ntm. ihc amount of
colid tnalter being variou^y Mtimat«'d m from .'iO tu -ll) or more pt^r ci-nt.
Tho solid* arv almcjat entirely organic matter, the iiiorvnnic salt* amounting
to lt»« than I jier cent. Of the iir^anic matter n^ain hy far the largt-r jwrt
coneiste of ha'muglohin. In I'lO )inrT!i of ihc driH <irgnnic matter of ''~'
corpuRclcs of human blood, about !H) jiarlii are hn-nioglnhin, about 8 part*
proteid euhnlanccs, am) nliout '2 purls niv other (ulwlnnccs. Of (he last.
Df the niosl important, forming about a <]uortcr of them and apparcQtljr
being always prexeiit, is tedlliiu. Lliolesierin appears also to be nonnslly
jimeDi. The priiicida which fi>rDi ihe sironia of the red eorpusclee ap]>e*r
to belong chiefly to the ulnliuliji I'limily. As regards the inorganic constitu-
ents, the corpuscles are diBlin;,-iiiabed by the relative abundance of ihe walla
of polait-iuiu and of |>h(L^|>baii'6, This at leirnt i» the ea^ in man ; the re'
tive i|Uiiutilies of «odium imd polaiuitim iu the corpuscle* and M-rum reo
tivvly upgimr, however, to vary Ju dilTerent animals; in aoino the »i>d
tnlle life in excrr* even lU llir corpuscles.
S 37. The pioteid uialrix of the whitr rorptiiolrr we have stated t« be
Oomp<we«l of my AMU (or an nllieil hmly >, |iiirnglohulin. and ponibly other
proteids. The nuclei contain nueli-in. The? while onrjiiKiili-* are found to
contain, in ndditiou to proteid material, lecithin and olher fa t», glycogen ,
eslractivcs and inorganic salts, there being in the asli, as in that of the red
corpuscles, a preponderance of polassium sidts and of phosphates.
The main faciN of interest then in tho chemical eompo^itioo of the blood
arc as loliows; The red corpuisclcs consist chic tly of hu^mnglobin, I'lie organ to
•olids of Mrum consist portly of serum-albumin, and partly of iiaraglobuliu.
Thcitcrum or plasma conlntMs.io man at leoM, with (lie corpuM:W. inasmuch
n« the former contains ehii-tly ehloridw mid sodium salts, whik* the latter are
richer in pbi^fphatcs and jioiaMiiiin suits. The extraciiveti uf the hluoil are
reninrlcahle rather for tluir nurnl>er and variability than for iheir almndaiicc^
the mint coDslaot and im[Hnlaut heiiig jierhaps urea, kreatin, sugar,
lactic ncid.
;>ari
1
waiia
Hum
rilK gCAICTITV ANlJ IiISTRtBOTION OF THE BLOOP. 71
TiiK Qt'ASTiTY or Blood. as& m jytgrRMnmov is the Body.
, 38. T1>.
1.1
whul«
ayrtem
ImImk* •tnii'k Wtni-eii (be lifsoes whirli |
frofB. lb* bloM). Tbiis tlic [inaoe uf llic alimenUr; »iial larjii^ly luld lo tliv
blood nier odcI tli« mmerial ilerivcd from food, ivliik- llie extri-iory Dfuiiuti
larnly Ukc anar water itn'l the other subet«iivea ooDDlituiiu;; lli« fxc-ntioiis.
OtMT Uaaea botli i;iTe and lake ; aw) the consUlenble dmiii fnim ihi' UI'Mid
tv tbo l^mph sfincea wliicli takw [iliuv iu the cit[HllarM« b met by thu flow of
lynpli intu ibe jrreat veina.
Frun (lie mult of a tew nliierriiliuuji oti esociilcil rnmiiialH it ba.i \kkii
candutlal tlint thu Ifital ({uaiitily of blimd in the huiuuu biHly in aWiit ii^lli
I at iba bxily wi-ijibt. But in vnrinii.i aiiiiiinla, the prupurtinn of the weight
I at ibe b4<mi| li> that nf the biolv iiiin lM>t-ti f»iiiid to vary vfry c<>n«idvrnbly
io i)ifl«rriit individiiaU; and prolinbly thin holds giiod lur mim iilsti, nt all
rmu aithin ccrtuin liniil.t.
Id ibr uiiiip individiinl tlte <iiiiintitv prububly doca doI vary largely. A
MMldrti dniio iijmn ihi- water of (!»■ hl<>ml by Kftnt aclivily of the eserolorv
MCkix.a* )>j {ifKfiiwdwriUin^.Qr iinuddeii ad'Hlioii to (he niilerof thu bl<x>ii,
M by dfinkini: large qnnntilice of wnter nr by inJcc-liDg Snid iii(» the blmxl-
rnarla, i> ni|>idly coinpcDMtled by the pMNiige of ntiter from the liwiint to
the hluoal, nr froin the blood to iho tiMiics. Aa we have already said, (he
IMMn mtr ojutiiiually •trifiii^ to ki'cp up an average coni|H»)tioii of the
MomI, anil in to Aoin^ ke«p np an average qiinntity. In starvation the <|uaD-
lity land i|ualUy j of the bloutl b nutintained for a 1uD(; linie at the ex{>enBe
ft Um tinuts. DO that uAer eume days' deprivation of fwid and driuk. while
lfc« fiU. llw muaclc*. nod nlher tiiMiua have beeu largely diiuiuiihed, the ijiian-
tity of Uood rcmaiitt nearly the tame.
TWima] i|Hiintily or blow) preaeDi In nn animal body la eailmnted in the fol-
InwlBg w*j : S* n\\t\i litood •» jioHible i* alluircil to mvaiic from (lie veatela ;
tkU U Riv^urcl flir(H.-t1y. Tlie rrurU are th«n washed out with water or normal
miixtn kJuiIixi. hii'I ibi; wailiinjt* cnrcrully c»IIe(il«iL mtxed. iind mcamred. A
kwnra ijaaMiiy '•( blmxl U diluted nrilh wiiler ur normal Miline solution until It
paMMMM ihc saine tint as a measured si^eciini-ii of (he washliigit. This gires the
fg^auot ■■■ 1' ■ ■! 'nr rather of hfetnog lolling in ihe nn-axir*'! itiiecimen. Trom which
iho tMa in llir vh'ile wuhliiKu u cnlniUtn]. l.iutlf , the whole tiixly la
OnfWIj t- I . :iiitl nB>b«d free from bluml. The wuhiiif,'* are eollected and
ikavad. ami the aiiii'Uiil of blood in iheni ia estliDaled. at before, by eomparlaod
«teb a Meciaieii ni' diiiiinl bl'ioil. The ijuantity or blood, a« caleulatml from tho
Xmn wMting*, together with the awapad aud directly ineaurod blood, giro* tlio
I ifoafitiiy >>( biwid in ihe body.
The »ribAd U not free frotn M>jte(loB», but other metlioda are even more Im-
P#rCKt.
Tbr blood b in round iiuinber* dolribiited as follow* :
Abuai niMyfourth in the heart, lungs, birgo nrtericH and vciiu.
About one-fourth in iIk- liver.
Aboal (mthfuurib in ihu skdeial muMk*.
Abosi ona-fiwrlh in the other orguifl.
^vrm in (lie heart and |;reai bloodvwoela ihfl b)owl is simply in transit,
^•mt\ nndcrff'itnK niiy );reat (-hiuijcea (and in llio lungs, as far as we know,
■ ! (o tt«pirali>ry rhajiiceH). il fulIowH that the allerulions whirh
II ibe IiIoihI [iiirainK throojth the liver uimI iikelelal uiu«el(« far
CMwed thorn arhicfa occur in tha ml ul^lh« body.
CHAPTER II.
THE CONTKACTILB TISSUBS.
^ 39. In otflvr tbBt (be bltiiHl may nouriiib lli« aereral ttMua* il » ■
lo and from thum )>v tli'^ voni'iilar iii^i'luiiiuin ; ami (hii nuriagc i*!;
&ctiT« niovoiDciitit. 1(1 onk'r that iIk! I>U»)iI mitr (ulvqiintfly nCKircfli ihc
sues, it tnuRt lie rcpIcniKWl by futHi from the nbniciittiry canal, and nurif
fmru Wtc by the- rxcrclory or);uii* ', nii>) )i<itli thi-wr |mtcnM» ciilAit moi
ni«Dt8. IIcnc« bolbrv n« jiiyicol'^I furllicr wa niuHl HUidy some of tbe geoer
chankcilere of tho inovcRicni« of iho botly.
M<Ml of tlie movomcule of ilie body nro cnrried out Ity mMm of tbe mi
olra of l)i« Inink nnd lituba, whiub beiii;; connccled nilli th« flkeliHftn are
fnH|U«otlir called skoletnl muaclee. A fikeleta) muscle wfaen subjected to
wrlain infliienoes suddenly aborifiis. briiiffiii); its two etub nearer together ;
and it !» tli<' Hborteuing, ai^liu;; ujiun varioris bony levers or by liotp of odier
tueohunR-nl iirrnnKeiueuu. wliicli itntduccs the inoven»nt. i>ticli ■ triniHtnu-y
febortciiiRg. cidb^l liirtb by iH-rtniti mfluenoe*, and due as westinll iM* to clt!uigM|
taking |>Itic>' in the iniiNciiliir liiuiie forming the chief part of the ii)ii.'>f|i:, ^|
technically cnllcil a foHtraidwn of tbo muxdv: and llit- niUMUiliir Imiip isfpoken
of lu a coDtrnclilc ti»uv. Tbc hnnrt in cbittfly cMinii'isnl iif muKctibir ti«*iie.
ditfcring in cortnin minor foittunt* fmm tin' mii»ciilur tiMuo of the Kkplciiil
miiDcl«^, and tbe bent of the beurt is eiueniiiilly a conlmction of tbe iiiUMular
tisiuc conirMKinK it, a .ihortenin^ of the peculiar muwular Rbm of which the
benrt Is obiclly made up. The luuvumeiils of tlie atiinentary cannl and of
many olbvr organs are similarly tbo results of Ihe contnu-tinn of tbe njuf-
eular tissue entering intu lb« coraptMltion of thrise orgnno, of the iihorloning
of certatD muscular fibre* built up into thiMC orgaiu. In fact, almml all tho
movements of the body aro the result of the eontraetioD of muacalor fibrMt
of various nature and variously dUp'Morl.
Some few movements, howevor. am carried out by »tnieMir«>« which mnnoC
Ih- called muscular. TbiiH, in the piiluionarr posfunt nixl elsewhere, more-
lupnt b eflbcted by means of cilin atlaeliC'I to cpitheliuni cells: and else-
where, W in the case of tlie nii;;r»ling white corpuscles i>f the bloud, Iraas-
ftrcnec from place to place in the body is brou^^ht about by amabolfl
mo\-ementM. But as we shall see the changes in the epithelium cell dH
while corpuscle which are at Ihe bottom of ciliary or amiebuid moveneata
are, in all probability, fundamentally tbe same as thow which take pinoo '
n mu«citlar libre wb(*n it contraoU : lliey arc of the nature of a cnnirartin
and hence we may Npeak of all tliuie Ojt diflervnt forujti of canlrnctilr tissu^
Of all these variouH finon of cuntraclile liMue, the skeletal muscles,
account of the more complete devL-lopmeiit of their functions, will l>e holt
studied tirsl ; the others, on account of Lbeir very ninipticily, are in many
respects les nlisfiictorily unden>t(iml,
All the ordinary skeli>tal muxeUv nrv onnected witli nerrn. We have i
reoaoa for thinking that they are thrown into contraetioo, under nur
conditions, oiherwiH* ihau by the agency of nerves. ^
Mu»cles and iterves being thus so clowly allied, and having bcsidea M
many nro|>ertictt in common, it will conduce to clearness and brevity if w<
treat them logother.
eat*
o ^_
isufl
:S
...any
veifl
TiTK piiffxoiiKirA or iidsclb avd kbrte.
73
I
I
^^^V Tub- ruE:(oMe.vA of Musclb and Nemvk.
^F Miuciilor and Jt^moHi IiTtlMiily.
i 40. Tb« •ItolcWl uu»clet of u Trog, th« tmiii aiid spinal cord of which
Wt* beta docnmd, do uot sxbibil tny »pi>iitanei>ii3 Diovemenu or contnu>
llniM. CTCB thoagh lh« nervm he oihorniK iiuitv inluut. Jjifl uDdietiirbml,
tba vboU IkmIjt mny dwomium; wiihuui auy ooalruocion of May of Um
•kfttfjil na»cl«)i hiiving bccii witmwfil. Neither the akeletal iuuscl«e nor
lb* nvrmt di»lrihulrd lo thpin ihhw^ss mu' powttr ul auloainLic action.
If. hiifitiTer, ft tiiuvcic 1>« hiiil Iwiv ami he more or Ivm violently dis-
turbed—ilT, for iiuUiiM-. il Ih- pinched, or tuuchud with ii hot wire, ur brought
isln niDtmct nilh ivrtuin chvmicnl milietanevM, or ttuhjeclAd to the a<;tiun of
(aK-snlr nirn>nta— il will move, thnt is. cuntrnol, whenever it b thus dis-
torbnl. Th<Hi};h nol exhibilin^' miy epontaiieoiiA iiotivity, iho muvcle id (imd
a«itiatw> for some time afl«r the j^oiwriil dejih of lh>: nuimiil to be) irritubU.
Tbouffa il remaiui <)uilo lukiti-cnt whoii lell untouched, its iiDironi are then
domBal (inly— uitt ahwiil. Thf«« require lo he routed nr "tftimulntMl" bj
•oOM ebattge or disltirbunoe in urdt^r thai ihcy ttiny nuin!l!vt thrmselve*.
Tb« MibtUDoe* or «|[enta which are thus, able to evoke ihc aclivily of an
irritable miurle are ^j>i>keii of as Himuli.
But to prtMluc* a cvnlroctiun tn a muscle the Btlmuhia n«ed not hv applied
dinBcUr to the muiich-; il may he applied indir«ctly hjr mcnana of tbi- ner%-«,
TblM. if lh« trunk i>f a nt^rve Ite pinched, or Mibjeclml lu xudden heat, or
dipp^ in crrlnin cheniioal niilMtaiMes, or acted U]>->n hv various calvnniu
canvnu. ruiitrartiuui are wen in ibe munlea lo which brancbee of tJue uervo
aiv dtrtributcd.
_ The iit-rvi.-, like the muscle, is irritable : it U throwp into a state of actlTity
■ bv a ^iniulua; bul, unltkc (be muscJc, It dom not iltcif ooiilncL The
^Kg£ni ' ' 'vs not (riv« riio in the ncrrc to any visible change of fiirm: but
^Hb: - of M>me kind or other are Ht up and pmpagmted along th«
^^^^re U'.'«<n to the miuclo is sliowa by the fact that the muRclc conlracU
mUmt ■ pArt of the aent at some distance from itself is stimulated. B<Mb
nemr and nuitole are irritablv. hul only the muBi-Ie is contractile— i.r,, raani-
tmta ita irrir utility hr a coulractiou. The nerve maniftnls its irritability by
itj[ [taclf, without any ri&ible alleraiiun of form, ocrtaia
< :.'« vet up hy the stimulus. We shall cull tbeee cbaogca
■ (has prwtwgat«il along a nerve, *' nervous impulses."
f 41. Wt Iwe slalM ahovr thai the mii*cle ruay lie Uirown into conlrac-
linaa by Mimnli appliitl din-ctly tu itself. Kui It miKhl fairly he nrf<tA that
lb« nntnictioMi so pr<><lni-4Hl are in rcsility due lo the fact that tlw stiroulos,
aliboagfa n{fparenlly applied direnlly to this muncle, in alVer all brought to
b<ar on sunte or i>lh«ir of the many fim: nurvn-bninchtii, which as we shall
«*« arr abiinduni in ihe mu»cle, piuvini; amung anil bc*irevn the niUHcular
Ohw, (n which th»y fimilly end. The fdlowing facl«. however, go fsr to
ptwc that Ihe nuscular fibres iheniM-lvcs arc capable of being directly
«tiBOl•^ ' '*' I'i the intervention of any ncrv«s: Wlicn a frog (or other
aninal ed with urari. the uctves may b« subjectcal to the slrongc«t
■linali >< i::l 'Ui i.-iuihiu^ any contractions in the muscles lo which they arc
dJtfHhutnl ; yet cvi'ii .irdinary stimdii, applieil dir*;clly to the innsclc, readily
eonlractioni. It'. Ix-fore intr>idiicin){ the urari into the t>ystcm, a lies*
ht- p«M«<l iin<tcn»-nth the »>-intic norve in one le^— for instance, Uie
! Iriiwn lightly muml the wh'ile leg lo the eichision of the nerve.
Hi- '<: thai the urari whi'n inji-clisl into the luok of the animal nill
faia Kvsa to the Hgbl sciatic iien'e above the ligature, bul uot below, while
74
THB CONTRACTILE TISSUKS.
it nil) huve frco accHs tu the r^st uf tlie body, includtiig llic ivlio)«
sciatic. If, as good as the iiniri lias taken cllect, the two sciatic nervei<
Btiraillated, uo niovemeat of llie left le;; nill be jirodiiced by ^tintulnling if
left Bciatic, vfliereaa stront; foniractious of the luuseles nf the riybl le|f below
the licsliire will follow eiimulHtion of the right aciattc, wlKther the uerve be
MiiuuTiilod almve or b«luw Ihe ligature. Now, BiDce tite iip|>er parla of both
9dati<» lire i-cjually expoeed to tlic action of the poituttj, it u dear that the
(iiiliire of llic left nerve to cause coutrai'lion i.-< tint Httribnliilik to aiiy cbajige
liiiTiti/ liikt^ii iiliicv t:j the iijijier purliuM of the iiltvc, eliae why Hhould uqfl
tbr ri}{hl, whifli lins in ltd ii]ii>er [xiriion liri.*n c({UHlly rxpoicd lu the adioifl
of thr poiiMm, aU'i fail? Kvideiilty li'e poi»iin ncl# on Mumi- |>arta of ll>e
ncrvo loKvr dunn. If a single niiL^clv Wi removed from the t'irculatioii (by
li^turing ilK ljliKidv*wi'lj'J, previous lo the poisoning with iirari. that wutcte
nru) coiitJttct whc-n nny pnrt of the nerve going to it it stJmiilnti-d, ihotigh iio
other muscle ill the Wly will contract when its nene if KtiniMliil>.il. Hi-re
the whole nerx-e right dovni tu the muscle baa been exposed lo the action of
the potwn. and yet it has lost none of its poner over the muscle. On the
other hand, if tbe muscle be allowed to retuain iu the body, and su be
exposed to tbe action of the poison, but tbe Dcrve be divided bigh up and tbe
Jnrtcoiinccied with tbe miiscte gently lifted uji before Ihe urari is intto-
uoed into tlie ayttem. so that no blood flown to it ami so that it is protedeit
from the Inltnonco of the poison, stimulation of the iiLTve will be found lo
pniduce no ciuilraetioua in Ihe muscle, though ntimuli applied ilireclly to
the niiibclv III once cauw* it to i-oniraet. From tlicse facia it b dear that
uniri {HiiHiofl the erdii nt tlx- nervt^ tvllhin the muiicle long before it alTects
the trunk ; and it in cxceedinj'ly probubli; that it U the- very extreme ends of
tbe nervcn (noRiihly the enil-|>liitra. or {Kieuliar xtriii'tnnit in which the nerve
Hhm end in the muscular tibrcs. for urari poisoning, at least when pro-
found, causes a slight but yet dislinclly recojijnizahlo cHect in the inicn>
Mopic appearance of these siructures) which are aflecled. The phcuomeDii
of uniri poboning, theretbre, go far to prove that muselea are capable of
being made to contract by slimuli applied ilireclly Iu the muscular fibres
lliemselvcH; and tbcrv ure other faels which support ibis view.
$ 48. When, io a rec«nlly killeil Ihijr, we i<timulale by various means ai
in various ways tbe uiuHcleit and nerves, it will ht: obw-rved that the mov
mentd thus produced, though very varioun. may bo distinguUhed lo be of
two kind*. On ihc one hand, the ri,i<uU may be n mere twitch, lu it were,
of thii or thai mur>clc; on the »lher bitnd, one or more muscles may remain
sliortoned or coolractci! for a ['■>ll*id^^lbU■ lime — n limb, for inslanee. being
miscd upor stretched out. nnd kept raised up or stnMched oul fnr many
seconds. And we lind. upon cxiiminiitiou. ihiil a stimulus may 1h- apptitu
either in such a way as lo produce n mere Iwilch, a passing rajiid cotilro^
tion which is over iind gone in a fraction of a second, or in such a way lu (O
kcci> ihe muscle shortened or conlriicted for as long a lime as, up to oertaia
liiuils.'we may choose. The mere twitch is called a t'niyle or sim}>{K miwru|^|
(lonlraftion : the sustained contraction, which, as we shall s«e, is really t^l
result of rapidly re|iealed tiimple contractions, is calkvl a Manie Mntriuiitm.
§43. [n onler to siudy ibefie contractions adeipiately, wc must hav
recourse to the "graphic meiboil," OS it is called, aud obliiin a tracing
other record of llic change i>f form of tlie muscle. To do thU coDvenieullj
it is best to o|>erale with a muscle isolated fmni the n-*l nf the body of A
recently killed animal, and ciirefully jircpurcd in >uch n way a# lo remain
irriinbti' for some time. The niusclci' "f pold-bloodcil iininiala ronmin irri-
table after r^innval from the Imdy far UmgtT tbnn those of warm-hl<Mi<lcd
aniniiiU, uml hiiice those of ihe frcg iire generally made use of Wo shall
1
♦BK rnE.VOMBS'A or MUSCLB AXn SERVE.
tlitHT piwtiiilv ifir ci>n'litii>D* whicli ili'inrmim- thU mninirnamv of the iiri-
tability uf muH-lc* ami imti'o! »(\vt n-movnl fmni thv bi.xiy,
A ntMelf thus iMiUlt-il. with ils dctvp kft Hllsclird to it, is railed a
m^utU'm'ryr frrpamlioH. Tlie iiK«t Cf>nvnii(iil mufcle for lliia [lUrpoee in
Um tng t*, pn"!)*!*. iW gatlrocncmiiif^ which shoiili] be diswded uut eo ts
lo IrsTi' rarvfiilK pnccrvnl the ntlmch merit to th« feniiir ubuve, kiiuc porlioa
of %bt t<-nJifn I lrnil()-j\chillUl Iioliiw, and u oonudcrublt; Icufrlh ot'tbe »dalic
mrr* kiUi iu btsiK-h«a p'iti]; lu tli« luiiad^. <Fif;. 1].)
1^,. II.
•ff
.4a.
Sp>V
A MuruMitsvK l*BKr«iunuii.
IkB ■»■>*. «w«nKa«iilUi of (K* ; ■. Ibe w4MK- nm*. «l: lli« Iinwkn Mna rui imB; exixpt
mfftjtnf ilin n><««l< , /, Inoiir: it. riaiiip: L«-. •nnlo-AcliUlli . it-f-.t'Oiol •|ii»I niisL
^L^SM. We may nfipiv In tut'h n niusclt^nerrv pr«psratioii the rnrioua kinds
^■^^tali ajKik^ii III' iiliiii'r ; iiirrhiinind, mrh m iitiikii)]; or piurhing ; iher-
^^I^Kck a* lud'li-it hcnlifi^ : i-lM-niical, rucb lU' aciil^ or ntlior wtive dwini-
■ «ail fnWlnii(v»>. or clrctriciil ; iiii<i iIk-j* wo nmr apptv either l« the intu<cli>
IdinTlW <ir In ihn wrvr. ihtu iitr<>>-tin^ ihv inii>cl« inOirectljr. Of nil lliew
•Itntuli Viv fitr lb<- nimt cunvi-iiiriit l'r>r ^^eni-nit iiurpOMS are cleclrirat »uniuli
of T«ri'iu» kind- ; mid lh«-»*. cswpl fiT (jH-cisi piiqioK*, nrc hvA appli<^ lo
IIm Denr«, aitd n'>t •lirM-tly to tW iiiiim-1i-.
Of «l«ctri<.'nl atiiituli, ngaiii, ihc rurrcDtn, «» they ere nlled, gCDvnitnl by
m voltftW ovll, itrv aw^t cv>nv('niciit. lhi>U[,di ihc elorlririty gcui-mtcil bv a
rvtBling mA);ii'-l. or Uuil iinxliieol by rrii-tiiMi mny tw tniploynl. MalciDg
iwe of K wH <>r ImiTy of rclla— DunittirK. Gruvc'fi. l.«4'liitich£, or any olbor
— «« mufi di*iin|;iii»li bclWMii ibe current prvduwd by lh« veil ilsolf, tbo
eonManl rurmtl, il* «« «lmll call it, ami the iiuliu^-d nirrvjit ohlaioed fmni
lb* conMAtit rurrrnt by m«unB of u» induction cd), u It i« colled ; fur the
{Ajifetogit'Bl i-ilfcts of [bu two kinds of L-urrent are in many ways diDcreut.
It naj, |>rrlia|Mi, bi> worth irLilp li> rtmind the rcxlrr uf the rollowini; facia:
In a galtaDic battery, ihir luluiante (jilatc of lioc, lur icilaof e) iiliich U trlnl
npno and iiwd up by the liquid ia raiird Ibe potitirt clrmeDt. anil Ibe Mibitauce
76
THE OONTBACTILK TISSUES.
which Ih not 10 acted ii)>oii aoiluDBd up(|)]Ht«,cic.. orco))per. pUUiiiim.or outiol
elv.) is called tlie ntffattie olvmdnt. A gmlvnnic action in Ret Ufi wlirn tlio jKnitin
(nine) nnil thr n«galivo (copper) eleiaonta nro conceiTitHl oubido the iMUery by
•omfl coniluclirig material, •ucb lu a wire, and (b« curredt U aaid U flow in a cir-
eult or circle from Ibe xIdc or poBitlvc element ta itie oojiper or negative element
iiutdi iKe hatttnj, and then froin tlie oopmr or nwatlve element tiack to the linc
or poailive elemool Ihroitgh Ihi^ wire ouMV^r the balirry. If the coaductiog wire
bv cut llirnuKli. the ciirtcnt ccuscs la flow ; but If the cut enOa he brought into
coiiLact, the current in rcf-fttablitbed aod coutiiiues to Uuw ao lui^ aa the coniai-i
U good. The end* of ibe wirea are called " polm," or wben uaed for pliyaiological
purpose*, Id wliich case ihoy any be fashioned in varioui wajra, are spoken of ai
tietlroda. Wbcn the pnlm nro brouffbt inln cnntact or are connccird by fitne
conducting lunteriul, ^ilvuoic n<:ti(in is set uii. mid the current Buwi Ibroujch (be
bailer;^ aod wires; ihii ia spoken of at "making; ibe c(irr<!nt"or '' ciHupIetinr or
cloaing the circuit." U'hen the poles nic (Iniwn ajmrt. from each other, or wncn
tODie non-condurting material is ititcrposeil between thum, the Knlvaoie action u
arrMted: thisi* npoken of a* "breakinir the current" or" opening lliecircaiL" The
current panMa froui tJie wire cuanectea with the negative (copper) element In the
baiterrlo the wire coitiiecledwitbthei>oritIve(Kiiiu)elGinenl in ibebaKety : hence.
^« polocoonected with lhocopper(n<ffAtive) element is called th'-z'tMi'iV pole, and
that coDDOCtcd with the linc (poaitive) cli'tnenl i* called the it'^iiitv pule. When
uaed for phyaiologienl purpoiea the piuiitivc ixile betMnK-a the posiiiire rlcclrudc.
aud the negative poltr the iieKHllre electrode. The pii^llve electrode is often
epokeu of aa the imo<U (ann, up), and the ncgntive electrode aa the kaihtdi (kata,
down).
A piece of riorre of ordinary length, though not a good coodactor, ia atill ■
coaduclor, and when placed on the eleciroilea iMinpluioe the oircnll. permiiting
ihc current to paai ibroUKh it; in order lo remove the nerve froiu the iullitcnce
of the current it must beliftod oCT from the vloctrodca. Tbii is obviously inoon-
vcnient ; and hence it is uxual to nrranfce a meana of opening nr rlooiinii; the cir*
cult at (time puint along one of tbe two wire*. This may be done in vnrioua war*
—by faateiiing one part of the wire into a cup of mercury, nnd >o by dippini; the
other »art of the iriro int>> the cup to close tbe circuit anil make the' mrreot. and
by lining it out nf tlii? mercury to open Ibe circuit anil break the current; or by
armngiiig bctwcAu the tw'i p.irts of the wirw a movable bridge of good con-
ducting material, aach us braitt, which can be put down to clone the circnit or
raised up to open the circuit ; or in other ways. Such a meniu of el'Ming and
opening a circuit, and ao of making or breaking a current, L« called a kry.
A key whidi is frequently used by physiologiata goes by tbe name of Du B'lls-
Reymond's key. Thouitb undtvirnbli' iu many reapects. it hiu Iho adTaniage that il
can be used in twodiifervnt nrsT*: when arraoguaas in A. t'ig. 12, the brass hridj^e
of K. the key, being down, and forming n nieans of good cuuduciiou bctKi-en t
braas platea to whirn the wires are •creive<l. the circnll it closed and the cnrrc
pasaea from thn positive pole (end of the negative (copper) element) to tbe poi
tire elcctmde. cir anode. An., ttirougb the nerve, to Ine ntmlira eloctrode,
kathode. K^ti.. uml thence bavk to tbe negative pole (end or tbe posi^ve (aind.
element) in the bsttery; nn raising tbe braM bridge, the circuit i* njiciied. the
correut broken. uikI no current passes tbrongb the electrode*. Wben arranged as in
B. if the bnus bridge bo " down." the rosiatance oQered bv it ia so (mall, eomp.-i red
with the reiustance ulTered by the nerve between the efectrodes, that the whole
current from the battery pWes through the bridge back lo the battery, and none,
or onljr an inflnitetinial p'lriinn. pawes into the nerve. When, on the irther hand,
tbe bridge is raised, and so the conduction beiweeu tbe two sidett <<ut|fended, the
current is not ubli> t'> pu.ii directly from one side to the other, but r^n and doee
pM« niong tbe wire tliruuuh the nerve back lo the bntter>-. Ilvncc, in srnuige-
meot A, potting down the key," a* il it called, nialcra ii current io the nerv«,
and "raising" or "opening thi> key " break* the currenl. In arrangenent B,
hou'pver, putting d':iTrn the key divorta the current from the iierv* by aendlng It
tbrou;{h lAc bridge, aud so back tu the battery ; the current, instead of making
the longer cin-ult througb the eleetrodw, make* tbe sh'irier circuit tbroush the
key; hence, this is called "short'clrculting," When thi> bridge ia miscdihe cur-
rent pMse* through the nerve on the eicctrodea. Tbns, " putting dowu " and
"raising" or"opening" thckoy havcooatrary edect« in A and D. In B, il wlU_
I
b« afcNiTcd, ili« bnllvrr U nlirkra M work, tli« ciirtent u nlwa^H IIoitIuk tUhtt
ikfwoith Ut« el»rtn>il«(l(er up) or through the kcylkpy donu); m A,ihel>ii«*ry
ia tMt kt wutk ntiLil tlie otn-uii i* nude 67 [lulUnK down the ivy. And in mnny
«M* ll U drtirablc tu take. «o in aptak. a Mmplv of ibe current while the bdtlcry
k in (tall ■wJnif. rather tbnn Juh •» it l)«fi»« to work, Mori-ottr, in 1) tlie elec-
llvilra Mr. libra Ibr key ii dnirn. wholly ibiit off from tlio clirrDnt; wherciw. Ill
A, «b*n U)» k^j in up, one electrode i> •till in direct connrriion wiib the bitllery,
•ad thh connection Icailine to what h known a* unipulnr acliim, may jtire rimi to
■tinulaiiiiii (if iht iirrve. ilencc tbc ui«^ of Ihe key iu ibe rorm B.
iJiliri rutin* of kvy may be uaod. Thin, in the Moivc key(r, Fi^. 13) oonlMt ia
k Bwl* by iiraaiDE down a lever bamlle ( An) : when the prmxiro ■• remoTtHl. the
taMfldlo, dri*eit ap by ■ (pTing, hrcnki contact. In ihe Nrrangrmrnt >liown in Ibe
Inirv, ooa win frmn itie bsiier>' being brought to the binding *«rev 6, while tbe
bbirdJiiit ecrew a iacoiincefol with the other wire, [laitiog down Ihe handle, tuakcn
ecnuiecliiin belwrrn a itnd A, nnd ihuH makn a current. By nirsnuiog the wirta
U lb* urvrnxl biixlini; *itt<vn in a ditl'rrent way, the making contact by deprtiwing
lb* hudle may be UMd to ibort-clrcuiL
fm. n.
DitntAm w* tic BiA-ltrntoiitt Kty Va
A.AaBBkla«*niltteBtU«; B, to •koiiHdKelUar
' iniluctiun i-ull," figs. 13 and H, the wire conticctinK the two eleinctita
Brrr is toivted at »oiuc n*n of ita course into a cloac •pira), called lllv
primary tvu. Tbtia, ia Pig. 13. iJiv wire t'". connected with the copprr or nrga-
Itaptelc/'.ol the hatieiT. £. Joina tbe (iriniary coil, /T.r.. aad then paiaec on
M jT''. ihrouicb iht"\ty'" F. to the poatlive Innci plttf i./-. of the battery.
Uref tbu primary coil. Irat tjiiite unconnected with it, •lide* aaotber coil, the
■laa f II J oh/, 'r. : the raA* of the wire foriDiiig tbi« coil, y" and r", are con*
tamni «a in tlie arrunKninent illuMrated in tbe li^ure aa y' and y, ukI aa x' and
T.asfl irnninato in elect r<'itea. If thme clectr"deH are in contact or connected
^h cosdacling matf rial, the citcuii of iho MTondary coil i* md to be clowd ;
etltvntlea It ia open.
In an^ an arrangetnctit it U found that at tbe loomenl when ibe primary dr-
' nil ;• Mt.i .1 1. 1 , wbeo the jirimnry currrnt la " made," a aecoudary " induced "
cat : t an exc«tilingly bTief |-crlod of liine, net up in tbe aecoiidary coll.
Thi'- ,18, when, by raovine the '"key" A' y"' aoa »'"( prcriouaJy not In
ronDf^tKiM with e^cfa other, are put iiiin cflniH-clioii, and the primary cnrremt tbaa
■adc. at that iiMtjiiii u currrnt appear* in the nircH y". j^".etc., liut almoat imm*-
dlMely dUappeHr*. A kiniilar nliiiosi liwiantiinroua current ia alao dereloped when
Iba firliuary current ia " brulcen," but not till then. So long Be tbe prisiarv cur-
mt flows Willi uniform liitenuty. no eurrcBl ia induced in tuo eeeoDdary coil. ''
!<■ tte mrdiii rli(iiibp[«(aiMliiluglb« BiiiWlt-iH.' , ,-.:..iliMi. "ni* iiiniwlp ■, K)|i|«ttail brtlw
clamixil.. ohlcli tinnif f'l* Ibrcnilof tt>i: flfmur/. Uoannwtad hf nmnidf ikc ^hook ■ uut a
iluaul mlUiiiis lever I. plocol btloii ih* idqI<i cluDbtr. !>« niims. wlili ■ ynnloa ef ilw*pla*l
oolnms K'tull uurlml bi li, ■■ iiluxl on ths •(■rtm-lMil'lat ''> i>i omUiM l•ll^ ihv win* t, v. !%•
ntholear thelotM'.rtror tbe elm com (t. It Wpt slunM wHIi innlMKM, and ikc clRlrado-bolikT
kiacMiKnu!i*llluu*i>lernor laaMriMil UMIIna'[«tcr mtjhi pUrfil ou ti irliiunii «Anilii( iBIa
wnatt wiui tha Dort*.
S. nenralTbic QtliHler bnrKti Iheiaokied p*p«"» "Olob Um lisvot wrttn.
Clhillali-RaiiiMad'ikcTMimactdlOTibDtt-clniilllac- TIicBtRB>>i>^l>arth«tliKtn9>l»-hoU*r
TOK MIK^'oMK^A or urAcLit and nervu.
79
curirni apfican lo UM ncoirniAry vihi. f u eacu com IBe ear-
f diirntiiin, gone io an inxtont almnat, aiij maf therefon be
li<ji'k," Mil liitlucUon sbuvk : beine cnlled n " nuking «hook,"
<y ttir inRtinn;, and ii " brtf^kking slioiik.'' wbva it m cjUM<d b/
^
1* oalj «hfn tti4> pficDHtT current i« «ilb«- nude or brakea. or suddoolj^ rariu
1« ialMttltj-. tliai ■ currpQi Apf««n in tint nvninAntj cM. fu each com tbe ctir-
nal b of r»Tj britff durntin
•p«k«n of M "■ s)igi'
wfcia It ta c>ui«(l by
tb bi««kin|-, of the prjinnn circuit. Th« diivclion of the ctirrcat in the making
•bo«k b "priiiBed la tbai of ibe primary current ; tliiH. in ihe ttgute, while th«
ptimMj rurrrnl flom from x"-' to y'", the jtidui.'i!ii miilcin); ihuck lloin from y lo
'■ Tfc« (tirrvBt of th« bn-akio; aknek, <io the uthi!r hand. Saw* in the ume
liirvdlMi a« tliD iiriiiiAry current frntn ^ to y. and l« therefure in direction ttaa
re<«rw <if tb* making iJMck. Compare Pig. 14. yrbere orrangtweot i< ofaown in
a iltefrmauBatlc manner.
Vu.. I*.
IRlnlia or IK iMIiirniiM Ollt.
• fuMtn pato, «Bd of n^tflf V cteuHail ; - dwiUv* pita. Ml of imKUre e)r<iHnt of IMis.-y : K,
Pa am Ki)wuwr»k«yi>r. t.^t»>Ooc<l.eiirtwmM>wnbTlhH>wp*«rro«;«r t. MonuUtywU.
nwwBi itntn bf mkiihutd am*.
Tb* cnrrmt from
aloof tne
— __ o»n to a DM
aad tbrrefim lending
om the toUerj. U|>an lu fint i>ntrHnv« int/i the pHmnry i-oi|, m II
h twiti -if thot coil, t'ivm rv* in tbe neb^Uboring tiriHU uf the
10 o«l1 to m nKnuvaUrv induc^l ciirri^at hnvinj; m liifMtmi opposite lo iu own,
to'veftken iUcir. Il UnotuuUI lltU "MlMntiucUou" Una
r
■awnvi Ikmoak hIaAlnfl temn In Hw Boar nf lb* mnlM tttuntivr wltb lli* wlrai i', y', and
aMMcand IBtMkcT.MKiHi i4ltirrM*. ItiiWuinc kfT»raklu<lxN) (lie oltMV.)'*. catnip
— WU^MTOBflu r. iif llM lixliii-|hKi<rnlt 1> Tl>i> •cunniltrT dill eau la nuOn la •lid* ai>
ia*«r lb* tntmufoM f* r.. aitb vtiloli an r^nn(rii!<t ilw laawlra i" >nd ir**; ^"b
dMcOf wlOionfolc— (cvliuuaer. lliBappn-pntcFp ctf llw ballerrX,' r*'l*e>rilt9l lo
■■■a. «.of lhi> Mfmii tvy /*, and U roiiUuuad u yi* fium wunliv UniUna Nnw, t| of
tha t<V M u* its* |i>la I p. lit iha taitnT'
•4«TilMiia h- lir Bfnntnl. aa^ the hallCTr I'hariwl : em ilapmiliia Iha haadle a*, of
tar /', a ninmi alll Iv maJg In ItH |i(lau7 will jir ( , paidiut ftum «. p Uironfft r" to
^, «. ■adtaMoaUuiHuby" !■>«, (i-ii>ia lu ti. and lu Ihivacli »■• lu ) v iinnmoTlns the fiiiaar
taM ** kaiHlla uf r. a Minnv tliraita ■!> Um baalk. and Hip iinuiBrv dnnii li In OMMsfQawa
IfkKiikn
ih»l llv rAMM«y enrKoi l* cHkw hmiI* ■■hmaMi.aalnducail BUnMil la iDrike
■wiiuyil in 111" Mnwutwy wM «. f. II iin onafclf A hi Mb Pa BitoBiTawJ^ hwt*
^
rn la iWlliirk Hiu laUwlnimi, llin «!•■ c*. r*. >. iba iun« batwavn lli«al«DUDd«i
a*(tka ali^*, ■'. ir*, IWai Uit«i>u|ilrtf<*om<lary dicult. ami (h« nnrra ooanqooiitlr *iprncii«ia
. M Imtlnv tudatt)<Mi«boek whcm-ni ilis ivlmarr enmal I* maile or bNttm If Ika
td ilv Im IM»-lti7»aMl ka; >• (aiit il-iA. u In Ilia dMIcd tlD« Via ik« t^m. tlw
Hf Ibr rnivkarU n>U(l>l cionraml nllh ikaiuf IM ncnv anrtu' I ho ali«* sohig n»in
ta* h>r *» Uw iMnc. Mat Uh) abulc accondnir (IDdocaij camni \mma iniB ^ lii v' (M Ouui y* lo
4^ aiMB tka rus'l*'. and |«ac*l«allr b«d* |«iBi tnioUi« n«rr*. Tba nom, Wns lk«a '■•bort-
.' u Ml adbctad lif aiij cbaafM to iW ciinaiil.
UUilmlol lacnl/ tnUhHUaleUw (uupral inotk«ila( KiuljtOS NiiwUlarooiilnioUun :
NN •>< 1.1 !■ MipfaadUiBiih* 'Mali* bonvtmiar* ualir«nalIradapie<l,«.iuiIaBl.ili«lisM toall
80
THE COSTBACTILE TISSUES.
8 used off that the current id the primary coil is eatHbliBbed in its Aiil atrengih.
wing to tbiB delay in the full eBtublishment of the current in the primarj coil,
the induced current in the aecondary coil ib developed more bIowIv than it would
be were no such " self-induction " present. On the other hand, when the current
from ihe batter; is "broken" or "abut off'' from the primary coi), no such delay
1b ofiered to its disappearance, and consequently the induced current in the second-
ary coil is developed with unimpeded rapidity. We shall see later on that a
rapidly developed current is more effective as a stimulus than is a more slowly
developed current. Hence the making shock, where rapidity of production is
interfered with by the self-induction of the primary coil, is less effective aa a
stimulus than the breaking shock whose development is not thus interfered with.
The strength of the induced current depends, on the one hand, on the strength
of the current passing through the primary coil — that is, on the strength of the
battery. It also depends on the relative position of the two coils. Thus a second-
ary coil ia brought nearer and nearer to the primary coil and made to overlap it
more and more ; the induced current becomes stronger and stronger, though the
current from tbe battery remains the same. With an ordinary battery, toe sec-
ondary coil may be pushed to some distance away from the primary coil, and yet
shocks sufficient to stimulate a muscle wilt be obtained. For this puipoee, how-
ever, the two coils should be in the same line ; when the secondary coil is placed
crosswise, at right angles to the primary, no induced current is developed, and at
intermediate angles the induced current has intermediate strengths.
Fill. IS,
TlTE MAliNETIC iNTEKBI ITOB.
When the primary current is repeatedly and rapidly made and broken, the
secondary current being developed with each make and with each break, a rapidly
recurring series of alternating currents is developed in the secondary coil aod
passed through its electrodes. We shall frequently speak of this aa the in/errupted
induction current, or more briefly the interrupted current ; it is eometimes spoken
of as the faratiic current, and the application of it to any tissue is spoken of aa
faradiznUon.
Such a repeated breaking and mafiing of tbe primary current may be effected
in many various ways. In the instruments commonly used for the purjiose, the
primary current is made and broken by means of a vibrating steel slip working
against a ningnet; hence the instrument is called a magnetic interruptor. See
%■ '■'"'■
The two wires x and y from the battery are connected with the two brass pillows
a and d by means of screws. Directly contact is thus made the current, indicated
in the figure by the CAi'i;* interrupted line, passes in the direction of the arrows.
THB PUKNOMENA OP ML'8CI.B AN'D KKRVK.
81
mp tito pillar a. ftlang the tt(«1 •priiig h. u Ut m t]i« iwtvw e, tbe pgiot of whicb,
um^ with plUlnuRi. I* in conUct witli ■ ooiall pluiuuaa plxte on 4. Tlie cnr-
n«l pMMM frum 6 tbrouxh e uid « connectiaf wire into tbe prlmuy coil p. I'pou
te MUrinf ioui ibv iirimorT coil, ui inilucM (laaUnfr) current U for tlic inaUxii
Jwleped in tbe wcondarr coil (not ■bown iii tb* fignn). ('rom the iiriiusrT
Mil p UW cuncni pMsee, bv a connecting wire, throueb the double tpiral m, iui<l,
dU Kotblng happen. wmiM ciiiiinni; lo p«» frnm m ttf m coontctlng wire to the
plUsT J, and wa by the wire y to the battery. Tie whole of Ihb eourw b imll-
anted bj tbe thick inlrrruplcil line with its arrows.
Pd^ W.
'V
:c.
■4
/?5
Tki MaowRW bTiaatmia with IbuniMR AuuimuttyT nm KaciuoM
twi UiKi um Unuk inn'u.
As tbfl rurreot. h«wevrr. psnm ibrouKfa the xpirsU m, the iron cores of tli««e
rw Mttde nugnetic TImjt In comMjoenoe draw do«m Uh iron iMr e, fixed at the
cad of Ihv ijiring b, the Oeilbilltj of the aptiog allowiu ibi*. But wbca i i*
dnn down, the platinum plate on the iipp«r lurTace ot fr b abo drawn awar
ft«B tlMactew <-, and thu« the current b "uoken" at 6. (ScoBttJlMalhe screw/'
b «> amuigwl that wlien c a drawn down a plnliiiitin tilnte on (he tmiirr lurfnoo of
i ia beaa(bt inta oooiacc with the platiiium-ariniii [loiiii of the ikt^w/. Tbe cur*
real ibeti riM-nt frntn A not to e bat to/, itiiil wt down tbe pillar i^, iii tbe directton
kndicaln) by the Min iiit^rrnpted Hue, aixi out to the batlerir by the win- <r, and
tt thot rni r.if from thi- primary coil. But ihia srrwifemetit i" unnccewiify) At
tk' I iDt llic current in tbim broken and •» cat otT rmn ihe priiuiirr coil,
ae ' iirenkini;) current is fur the mument developed In the ««coiidaxy coil.
Ital Uui burteut b cut olT noi only from Ihe primary oo|]. but abo ftom the aplnb
■■ ; In eoiHoinetice ibeir core* cease lo Im macneiized, the bu t CMiMi la be
•nncUd bf iheoB. aod tbe aprinii/i, br Tirtueuf it* etaiticily. rwunMS Ita former
Milllo* 111 oonbict wiib the anvn r. 'i'htt rrlurn of [he ■prini;. howerer, rcAitah-
KAm the carrenl In the primary «>il and in tlie npimU. unit the H>rliit b drawn
ihiii'B, to be reltmid onoe more in ilie Mine mnnner u Wr'-re. Tbos M long m
llw carrtnt b fwMng along ^ ibe contact of t> wiib r is sltrrnmely being made
■■d bnkM, MM the current b comlantly paasiuK into and beinx ■hui '>ir frum />,
llw pednds of alternation being determined by Ibe Mriodi of vibration of the
if*{a( A. With each pa«^e oiUie curreai iuU>, or withdrawal tr«ta the primary
<mI. an lixlueexl (msking aad. respectively, breaking) carrent is developed In a
•eeowlar; coil.
As thtH need, each " making iJiock." a* esplmned abora, b less powerftil than
iW OKrvapUBdlag " brealcing »hock ;" and, indeed. It sometimes happenn that
«
TKB COXTRACTItE TISBCBS.
iiutMiI of «ac]i mtke, u well u eftcb kieiik, >eliii( u ■ •timnliu, eiTiag ri*e to ■
conlractloti, the "break*" onlf are eiFi^ctivc, the Hreral " luakea' gimg riw to
BO ronlmclions.
But what in knoirn an HelinholD:'ii armngumeDt. Fig. IC, how«ver, the makiiig
•od breaking thocka amy be equalixed. Kor Lliis purpoM the ktcw c u niiscil out
of reach of Uie «xcuntoiia of Uie S|iria{; t, &111] n iiiuilcraielv tblck wire v. ulfering
A certain amounl only of rMUtance, is interposed between lue Dj)per biadine acrvir
a' on tb« pilUr <i, and the binding kkk q' lending to the primary coil. L'ndef
time arrariKcmooIii ibe current froin tbo battery pussM thniugfaa'.aloii^lhi? iiitvr-
pofted wire to <•', through the primary coil uud tljus as before to m. As before by
tlie ma^uellzation ofm.r in drawn donn and 6 brought iii ccictaci with/. Aa the
rwult «■ lhi« cnntjiet, the currpnl from the bnilery can now im*« by m,/. mid 4
Sbown by Ibc thin interruptt^d line] buck to ilie buttery; but not ihr whulp nf
e current, rtunii^ uf it can «iill piuH nlong the wire w to the primury coil, the rela-
tive amount being deiennined by the relative resistAnce ulfered by the two cuutM*.
Hence at cnch ■ncceuivi' mnsnotiiution ofrn. the curr^'nl in ibe jirimary coil doea
not entirely diHiippenr when t i* bruuirht in cnnlnctwith/; it is only »o far dimin-
tfhei] Ihul 'II ceuses lo ullrftt-l c, and hence by the rcletme of 4 from / tile whole
vurrrni uiice mure jiasHM alone to. Since, at what eorreHponds to Oie " break "
the cuneot in the primnry coil i« diminitihod only, not abaolutely done away with,
aelf-induclion makr« itn appearance at the " brejik " oc well a» at the "loake;"
thoa tbe "breaking" and "nukiDg'' inducvd currentu or shocks in theaocondary
coil are ec|uulix«i). They are both reduced lu the lower efficiency of the " mak-
ing" aliock lu the old armiigeraent ; hence to produce the same streaglfa of «timu-
liu wilfa thia arrHngcmenl a stronger curreni mniit be nrplicd or the *«coudaTy
«oil poshed over the primary coil to a greater extent than with ibe olJicf arraDg«>
men I.
The ilienomcna oj a Simple Afutcular Oontractum.
4
S 46. If the far end ot the nerre of a muscle-nerve urepnnitioii. Ftp. 1 1
and IS. be Inid on olectrodeB cootiect^d with ilic K-cnntlnry coil of nit indue-
tion-mAcbine, the paxsage of a nngle Induction (hock, which may he taken
at a cnnvenient form of an almotit momentary stimnliii, will produce 110
visible change in tlio nerve, but tbe iuu«clo will give a twitch, a «hort. sharp
contraction, 1'. e., will for an iDilanl ■horten itaolf, boooming thicker tm
while, and then return to ite previous condition. If one end nf llie muscle
be nitaohed to a Ic^Tr, while the other \t lixcd, the lever will by its move-
ments indicate the extent and duration of (he shortening. If the point of
ilie tever be brought to bear on some rapidly travelliiig surfnoo. on which it
leaves a tuark 1^ being for this purpose nrmod witli a ucn and ink if the sur-
(kce lie plain paper, or with u bristle or finely pointed piece of platinum foil
if tbe surlaoe be smoked gla^ or paiwr^ so long as tbe muscle remains ai
rest tbe lever will describe an even line, which we may call the base lioe.
I^ liowcver, the mascle ihorleus the lever will riae above the base line and
thus d«tcrlbe some sort of curve above the base Hue. Now it is found tliat
when B single inductiou-Hhoclc is sent tliruugh tlie nerve tbe twitch whidi the
mtucle gives causea tbe lever to desoribe some such cur%'e oa that shown in
Fig. 17; the levrr(anrra brief inltrval immediately auooeed in ^ Ihe oi>cu!ng
or diutiiug the key, of which wi; shall apeak presently) rises at lirvt rapidly
but nflerwnnl mnn.- niowly, stiowinu; tlmt llit- um»ele is correspond ingly
rbortcninjf ; then ceaum to rise, showing llinl tin- muncile i* ceasing tii grow
shorter: Uien dtvccndK, ehnwing that the nuiticlc in liMiglhening again, and
finally, sooner or Inter, renches iiiid join* tin- baw; line, vhowing that the
muscie utter the shortening biu regained! its prt-vioit* nuturat growlJi. 8uch
a curve deecribcil by a muscle during a hvilch or simple iDiiScular contrao-
tiuu, caused by a single iiiduclion -shock or by any other stimulus productng
rne phknohe.va or ml'sclr axo nebvb.
88
•a* «Atf,i>ctIlMl I curvAof H iqni|il« muiwdliu- oontraoioii, or, mora
'■hvtl*. • "miuoltt-ciifvc." U b nlimiu thnt th« rsact form of (be ctirro
dnvi^ml Uy UirM'uMl roolrncliaiu of il miiiicle will <lpp''iiil i>ii tlif rft]>ic)itv
«kb which Oir rpcvmling surr'uw i* Irnvcllioji. Thu»if ihi- ^iirfliN be (r«v«J.
Bm cJtiwIj ibe ii(>-ir<>ki' i^'im-^jHitiiliiig to ilic tJiitrU'tiiii^ will Iw vert' abrupt
mad tbv ((u«i D-«ln>kv also vvrv KUtji. l<^ in Fig. 1)1, winch u n curve froia ft
•.IT.
Fus-li
VTw
..»:..
II I I <>H iirr ri«iTiiooriHii> i>r mi Pkm. Tlili cum. likv «ll mnndlnc oiis,
iW -ti'4, i( lo iB mtd frn« kftio lUIit— ilMiltisnjr. whilpihe lentimd i«iiln(-
• •taitiii.>r) iiic MrartiBf (OfTHv oM tmiolUns ttiHn ilEbltD InlL
mtm Uir BoiiMti at ohttli ibr brdwiion-tluH'lt i* xuii Idu> ilu> nnr*. b Uh niiiiiiiiiiiimiiwii.
lf»i>au uKl it Ik* rloar nf iIh OMUnffUin.
lh*«aKl»«iirvrl>iliiic>inr']n<iub]fiituDlBc.n)[knuikliwloa<IoUtikVibniUaniaM«oad.
vplrtir fune nrfcmiiuiic (benKm ouv biiiulmlili of ■ bomuL
fencl>ralill9 iii(iarl« of n friw, tnkru vrilh a Hlowlr inuvirifr drum. tli«
fl^lbrk b«iii^ tbe tumio ft« ibn) iind in Fig. 17 ; inilfcil, njih ii vrrv' t\ow
wvimcnt, ihf twti mtty be hanlly •cpiimble Ham each oUht. Oh tbr olhvr
IhumI. if the •urtiLcc irnrH verv rairidly th« curve mfty Iw iinnii»diit«lir long
tlrmwn uat, *n in Fig. I!', whicli is » curvo from a giutrocncmiuH muscle of
a frtifi. taketi with a very rapitlly moving pendulum monvgrnph, the tuiiiiiff-
(oA mftrking Blwtit t'lOO vibrations a feeond. On exnniiiiHtioD, however, it
will ho IbuQU that both these extreme curves are liindaiiicii tally the same aft
lb* nadium one, wbea tooouat it taken of th« diSnreut rapidities of the
ii»T«lluH surface in ibe sereral caoM.
Ib oftfer to make ihc " mu*cle curve *' complete, il is nc>oe»arr to mark
Ml ilw rccordii)][ surface the exiiet time at itbieb the inductian-sbock ia seut
laut ibt aerve, and also to note the apeetl at whieh the recording etifface h
umntOat.
Ib th* pendulum mvograph the rate of movenieut can be calcuUiMl from
iK* ttogtfa of the poiiilulum: but even io this il it eouveniem, and >a the
ea» uf the aprinfc MTu^'^ph and rerolring cylinder is neoesaarr. to roeaturc
tba nklf of mm-enx-ni direvilv by means of a vibrating tuniug-fork, or of
wiB« body rihiHling re^iilarfy. IndetNl it ia beet to make such a dirvct
BManrMBMit with eai'h curve thai l- btkini.
A taniav-fork, as i* kuown, vibrate* ao many umea a Mcond according to
ila f>Hdb. if a inning fork, arrmil with a light murker on one uf iu prungs '
■M ribrating mv l"" a tccood — r. r., <is«>'uliiig a duuble vibnilion, muring
ferwmnl and liai-kwan). 1**0 timet a second —h« brought while vibrating Iu
■ak' a tn^-iRL' "B the recunling nirftee immcdiaiely Iwlow the lever l>c)ong-
iap ' '. w<* (3in um: the cnrvL- or ralhiir t-urv<v di3'i'ril>o<I by the
toD'- ' Mim thv dunition of any pari nr of tlie wholr of the miu-
clt>e«irv«. It is i-eMnnlial thai nl *(nrliug ihr [Hiint of llie miirkcr of tba
lastaf-fork th»u)d l>c trxnelly umli-nii-iilh ibi' marker of ihr Ivvur, ut rather,
■nrr the fiini <>f (he lever w ii move* uii and ilnwn dtwcrilxa not a Mraieht
but an arr 'if a circle uf which iVt fitlcruiii i» the cvOlre and ilself i,fn>m
TUB COJfTRACTItK TtSSnS.
lli« fiilcTimi to iWtipof llie iiiai'kcri the raHiu», that llie jmiDt of the luarker
oftho tuntng-rurk Hhoiitd be exactly mi the arc described hy the marker nflbv
lever, either above or below it. ai may jirove mnu
I"'"* con veil ieut. If, iheD. at MartioK the lAiDiiiR-fork
marker be thus uii the arc of the lever marker, «iti)
we tiuie tiu the curve of the tuning-fork the place
ithere llie arc of the lever cula it at the Iw^iiuinv
and at the mil of the miiscic- curve, as at Pij;. 1',
we call cdiiiil the luimlaTof I'ihralionn of the luiiiDg-
fi)rk which have taken place belnviMi the two niarkik
aivl »i> niKHMiaiii the whole timu nf the inuAcle ctirvc;
if, for inotanct-, there have I>c«-d 10 ilouhle vibration*.
<!ac'h occupying , J^ »eci>iiil, the whi>lc curve hn» takes
,'n so'oiti) to i»nl(e. In the Miuie nny wo Can uivKMire
the dtirntion of ilie ri«; of tlie curw or of ibr jall, or
of any part of it.
Though the tuning-fork may, br »niply rtriking
it, be set going long enough for the purpoam of an
otwervntion, it is convenient to ke«p it going by weans
of an electric current and a magnet, ve^- much at
the spring in the magnetic uiterruptor (Fig- 15) is
kept going.
It is not ueceteary lo uee an actual tuning-fork;
any rod. armed with a murker, which can W made to
vibrate regularly, and whoi^e time of vibration ii
known, may lie used for the purjxwe; lliua a reetl,
made to vibrate by a blast of air, is tometiioee eui-
ploveil.
Thv exact moment at wliich the induetioti'i>hock ii
(brown into the nerve may be rvcoriled ou the niuicle-
curve by menns of a " NgnnI," which may be Mjiptiiil
in varioMi' ways,
A light steel lever anned with n marker U arrancod
ovM ■ Kiuiill coil by mean* of a lifibt ipriutt in Mich a
way that when the euil by the iiiuwngc of a current
\ through it becoum n iniignet it palls ibe lever ilown lo
ilBcll- on Ibet-iinenl being lirolcnn.nnil the mnunetizaliuo
of the coil ceiHinK, ibc lever by help of the *i>rioic Hie*
lip. The marker of ouch a lever in pliiced iiniuediiti-ty
under— i.r., at some point on the arc de«cril>ed by— ibe
uinrker of the intisele (or other) lever. Hence by making
a current In ilie coil and pulling tbn li^al lever down.
nr by bicinking iin iilrcndr exi:itii>|r current, and lettinj;
the iiidniil lever By up, we can uiake at pleaHure a mniK
eorreiponding to any part we please of the miuclclor
other) eari'e.
If, in order to ningnetir« the roil of the •ignal, we tM«,
an we may dn, the priinury current which Reuerale* the
indurtioii-nbock. the breaking or making or the orimaij
cnrienl. whidiever we u»ii lo proiluce the inouctino-
■bu<^k, will make the Mgnal lever Hy up or come down.
Bvace we ibnll huTr mi ibe recording nirface, under tli«
niuarlr, a mark indimiiug the exact mometii at wliidi
tlip pritiinry current was broken or made. Now the <iai«
taken up by the generation of the induced current and ita
M0Mge into Ibe nerve betneen the clei'triidti' i* wi inliniie*imally >maU. thai w«
nay, without appreciable error, take the mouient of tlie breaking or making of
THE COSTKArriLK TlgSVEi*.
tbe primary curr«DC a« the monieiit of ibe eDtr)in(«nrili« InOitclioii-ehoci: (iitutke
nerve. Thiu wc csn mnrk Wlow ilic> niaKl«-curT«, or hy dcKribin); ih« «rc of
tlko niMcIo lovpf, 'in ilir mumOc rurrp it«iHf. clir exact niMiieat at nliicfa the
inducUon-sliock lalls into Ibc nervv between tbe elrclrodtn, na U dofic at a in
RgB. 17. 18. 19.
In the peoduluni mvogmph a «e|>arnte iiii^nal is not needed. If. harinK placed
the mufcle tever in the pn*itinn in iThi<!h w«i intvnd to make it rm«ril. we allow
tbpglaM pinto lo descend until Ilie tontli n' jwit toi>chei the nxl r(ta ihai the rod
ia juiit iiltuut lo be knocked down, uud *o Ureuic tbe primary circuii) and oiake on
tbe baite line, wblch is meannliile being dettcribed hv the lever marker, a mark to
iitdlcnto irheie the point of tbe marker u under iliv*c drcunutancn, and then
brinf; hnck the plate to ilK projirr pwilion, ibr mark wbidi we have made will
mark th« muuieut nf tbe breaking of tbe primari' circuit, aud no of tbe entraiKe
of Um ill duo I ion -9 book into the nerve. For il is juM wben. ati tbe iilaM' plaie
airings down, tbe murker of the l«Ter comen to the mark nhicb we bare made
that the rod f i> knocked hark and ihe primary current b broken.
A "aixual " like ibc above, in an impr»ved form knuwn oi D«iipn^t):'s, may be
lacd aUo Ui record liine, and ibua ibc awkwatdneM of bringing a larve tuning-
fofk up to the recording aurracc obTiaied. For this pnrnote the signal i« intro-
duced into fl circuit the current of nblcb ia coulinunlly ticing made and broken
by a tuning-folk I t'ig. 21 ]. Tbe tuninu-fbrk once net ribmlini! continues to moke
and break ibe cutreiil at e«cb of its vibrations, and ai Hinted above ia kept vibrai-
ing by the current. Bui each make or breiik cHuwd by the ttininic-fork alTecia
■lao tbe email roll of ibe nlgDal. causing the Icvrr of ihaaignnl u> fall down or fly
up. Tliuii the Bignnl dcMfibea vibration curves nyncbrorioua with lho>c of the
tuiiiD{>fu(k driving it. TbeaiKnal may similarly be worked by ineaiiiof vibrating
^^Bta other than a tuniog-fDrk.
Varioutt recording aurfacea may h« u»ed. The form moat generally iisefbl ia a
cylinder covered irith imoked paper and made to revolve by clockwork or otbcr-
wife: auch a cylinder driven by clockwork id aliown in t'ig. IS, B. ^ naiiR >
eyiiiiderof tar^-erailiua wlthade<|uaiegeaT. a high apeetl forliiaumce, in aaeoood.
eaa be obiaincd. In the ^Ttn;; mi/r/i/raph a amolced glaaa plate la thrust npidly
fbrward aloog a groove by means of a spring suddenly ttirown into action. In the
t\endiilum m-zoffrufih. Fig. 20. a amoked ^ln«> plate atlHched lo tb« lower end of a
ong fmine avinitioi: likr a pendulum, ia auddrniy let go at a certain height, and
so swiuga rapidtv through an am of a circle. The diuidrautxge tit tbe last two
methods ia that the surface travels nt a contiDually ^hangiug rate, whereas, in the
nvolving cylinder, cnreful con^iriiction and ndjuslment niill secure a very uniform
rate.
§ 46. Htivine thus obtained a time record, and an indication ol'tlie exact
tuoincnt at which Ihe induction -shock talis into the nerv-e, we may for preeent
pur[KMe)t coiiiudcr tbe muscle-curve complete. The study of such a cur\-«,
nafor inslauce that ahowu in Pig. 17, taken from ihegnUrocnctDiusof a fW)g.
teaches u» thu follivwinK facts:
1. Thrit nllhgii^h the piiaau^e of the indueeil current from oleciro<lc U)
cloclrode » |inictx«illy iuKtantuncotia. iUcflcct, mwuiiirwl from I he oil trance
of the shock into thi- nerve to the reluro of the muscle lo it« nntnrnl length
after the sharteniiig, takn no appreciable time. In the figure, the whole
llbeldta that pvTIlun lir Ihv lonlhu'OBlotiliutan ihr micIi v. In Ibetonnaof llaralna UielnMh
a* ooinlfic tn coniBin wlihihct^oJeetlnaitMlrtiilf, kii<>ckiliui>unualdolmoUie|>odtlanln4loawl
bytb« i1«ll*<l lliiv r'. Tbe nid rii In «j«trti] fuiiUnulljr villi llii> wlrv i of tbe |frtuiaiy coll of Ml
luductlon-iiiachliiiF Tlii' (rnrw rf la ■linllarly til vli<ctrld MntlniilCT "^^ 11" o^rv yof tlMMBC |>rl>
fuary ooU, Tlia fora^v tf aiiJ Ibc itrt i' ' rv armi-d wjili flaUnum ai ihv ]vilu<a la trhlcti tlw7 an In
eeutuct. and bmli ai« limiUlnl liy iiil9Ui> ii{ lliv cbniilii- lilwk t X> 1iiii( a> ' anil •! am In rontacl
UiDClrinill oT llw iitliimtrCDtl U> Hhlch i aii<l u buloiiK I* cl-awl. W lien In lu naUu (be iDuihtr*
kuockiraua; trom it. ai tbat liuTanI tb« rtmill in broken, aiid a "lumkiue" ibuek li agiil lti|ea|li
tin tliclniili'i uiiiiivL'ii>a Willi Uio iiBCOiu1ar7 Mil nf (tin nuirlilni>, aiul iti tlimiigb lb* atrn. Ibe
lover r. lb* Fill) only ^•i urlilrli i<thown tn tbe dfnn-, !• brouKhi lo btarou (bcalaa plaie. and wbni
Bimtileachbaaimliihi tlni.-, or inoTseUiOllr an on.' of a Plrelc or larito ndllK Tbo luallia-Airk/.
Iheend«nUl]":>Illiclvn>tliDla<>ri>blcbat«abnwnlnlb*ngun|dat«dltHM0lB(«lyMOlr Dntlncr.
Sttvea to niark tbe Ilia*,
TUE I'llKNOMKNA of UUSCI.K AKD XKBVK.
87
mrr* tnn a lad takca up alxiut tl>e tumw time u elcron iluuliti* vibrntiou
of ibe iDotoK-fork. Siiicv Mcb doiilile viliraliou Iktc rvpRacnU lUOlh ot a
wcwmI, U)9 diiniii<>n ol' the wbnlc i.iirvo L« ralhvr auirv iUbd I'n m.-cou().
2. In tlw tint pufium of ihiit period, ftnm u to b, l)i«ny b no viaible
rfwuy. nu niiiine nf tlw levcir, »<■ ihoncnltig of th? n)u»cli'.
T ItfenaiuDiil/'. thut i»lowy.»IWlli« lijiMof sUitit 1<lOlhM'c«n<l. tlint
i1m> •bortftttog tx^iii- Tb» thMteninr ■» *h»wn by ihi- ourvr i* ni lirst slow,
bat ■•-■m hccmuM mon- r«]>i<l, ami liirn I'tnckuii* sgvi'i until it fvnc)i«» a
tsiRium atr; thsntiok- i>liiritouitig ucctipyiug rather titurv (ban f|„»eoiiul.
ria. ».
TW nmrM tbmi altof Ik* wlcr/mniMFIrd wllh Ihr rnliliP I - > t*>'v ''r <■)') or tW wfillt*
. .VinritokkHay, Uinach IlKlnnlnii fl:>rt.>toini Uie|>lii ooUDMIal slih ltW(B>l->rihalair«r
, hi lU iiMKaij la Uw «|i hf, anil to by* itlia I'liawu In ncu(v> 1ft lbs Uitdliif ten*':
I IMa MadlBC actvir [art Of Uw viinviil Bow (limuirli ili« cnll 4 bKwcKi the pnnip ot lh»
I tbtUM ti) ike Kin < lo ilic lilniliiiR >fit« a, whll« ■nothar pul >ovi llimiMik lli«
I iIm colt of Iha t*et|inu ilunul burli by ibc nliv b, loUwUiuBiicicmr il Kraiu lh«
r « Ika eutnal IMM* buk tuiii*iir|ailv*i — > |ol#orc«<lnrth«ps«lllv«al(**iui^irf
AtUwcwmii Sinn Uifnnith ihv n(41<y ih* I>iii[iniii •icnal rrampWI^ Ike ran) (■(cull
[^■COrtlavd ilnin40Bti Uw luiKmiribaaignal Adhcrnimni llainlbinuah IbSCcild,
rofllMi iiill, alan IcoHiiliif mitcnellHMl. •Itainap Ibc lover immi u( lli« lirrt Dul Uw |da
ln»M)— I fl Ihatitwdnialiiir ■i-i'ibo iimic UfU ibt fudatof Ux >4ii niiiorilic rattTiiry. Iiiroii-
I til* oimal Mill Ibiia b(nk*a at llf, Dckiii iiritbir IbRinjfb >l tuw thfuifh ihc Imiovii
ta ww>|i»MWf. IhefvMT of Ihr liisi«i!U thoa CHUlrii to Iv iimcnvllHil. Iti* miukti flli-*
Mai OBMlly aa^ilvl liy*(prtD| nw* (bamiln tbe llfum But In iun**qiii>iip«i.itiliaeiirr«Dl
[ la Id** Ikroatk 4 liio oui* oT 4 waw lo lift op itu pronit. aai) ibv iilii, in Ui* dwmni of
t. intka OMIafl noc* stw alUi (Iw oian^rr. Tbt n^aubluhmunl nf ibc cumni. hrni-
iaui«acUa(«a Uw two mIKaaala pull* Oma itio niukvror ibcitKnal. ainl a«iklii Xir
■■■t>aMn«<4|«lka[>Uiv i«nna aii4 oma mon bnnkt uia aiiRwa Thiu ih« vurmt
%t>f HNUloand UiAt*. iw niilitltrnruir imcmipUoaaMwJatcniiined bit tberlbntiiia
> M Uw luklar'ntk. Bod ibo Ivvci •■€ iw U|[iu[ tWaf and fclUns ■nicbninuiuly uua ika
■ «f tkaunilf«-AHk
4. Arri^'«d at llir tnaximiim of sliortcning, llic tnuHcle at nuLv Wgin* in
ralas. the lowr ilniiyiiiliiig al lint hIiiwIt. then molt' rapiilly, anti at liwl
OWt* tlowly agaiD. until at J tlir niiiflr hut n^^intnl itn natural length ;
Uw wholt rMuni from th« miLsimtiin of contrnctinn tg tho natural kiiglh
nemiiyin); ratbrr more than jig Mcnml.
Tbiu a Mni|>lc muMnlar conlnietion, a simple spasm or twitch, praducMl
by ■ nonenlarT aliniuliu. such as a nnijle m<lui;lioD-»hock, cottiiMs of three
iHMii pliaaM '
1. A pboMt aiit>?oed«iit to any vuihle atteraiioD in the muscle. This
phiwi. dariD); which ini-igibl« preparutory changes are takiug place in lb«
atrr* taii muscle, is called the " l-iltitl periwt."
% A [iliaso of aborteniug or, iu tbe nwre Mrtct meaning of tbe word,
CBOlrartion.
a. A phase of relasatioti or return to the orixiual I«:ii|{tb.
In the cose we are coitftderiiij;. iht clectroijeii «n> ?u[i|>ai«<) t" lie applied
lo tfas oerrc at soDie dbuuice th>in the laiucle. I'oiuequeDtlr the latent
86
THE COXTBACTILK TISSl'BS.
pt-rioil of ihp ciirv^' contprwcs not only the pre|iiinUorif actioiisi ip-in^ <iti in
ihc titUDclc ilsrlf. liiil iil«o tbo cbaneM nevMHirv it' conduct lln: iitiinnliate
eflwl of the induct! on shock from the pnrl nf ln« nerve IwIwwd the I'leo-
trodcs itlong n cunndcrnblc length of ni.-n,'e <1«wm to tht* niii«cJr. tl in iihr(.
ou« thiit llicef Ittltcr ehiingcs might be climinoteil hv placing the clcclrudtt
on the muscli? iteetf or oii the nerve close to the nuisclc. If this were ilonc,
the muscle niid lever being exactly as tiofoiv, nnd cnra were taken ihnt the
in<luclit>n-^iock eutered into the nerve at the new epot, *l the momrnt wltcn
the jKiiut of the lever had reached exactly the t&ioe point of the trakvelling
fturfuce as before, two curves would be gained haviug the relation* »honn in
Fig. 22. Tlie two curves reaenible eaon other in aTmoet all point*, except
tliat in the curve taken with the ahorter piece of nerve, the Itilent period,
ibii <liatance a to A a« compareil with the dtstnaoe atoh' U sliortened ; the
coDiraetion begins rattier earlier. A Miidy nf the two curves teaches ae the
following two fact* :
no. -a
0('ii\'™ ii.untinTiwi Till: MiMi-suKXT Of TiiK VtiMTrr nr * Xmrow iMrviA;
Till! Mim tnOKle-iicrTi |icv|«mllnn IifMmuUlai ill uAirtu pcslblc trova ibo miMplt. Oiaa naai
M lUMOile laitui muwlc: tvitbeaauaMantam K|tl*t*(vil iii*i«:tl; llicMauitn)'
In |1> UidiliuuluiiinWnilia iienv kt Iho llatt IndlwUid by iIm Un«<i, IhcountnulloD bnrliuBib':
Hid « IidU Uunt [ie(1iK3. UwnfUrc, la tndlcklcd 17 ihadliQuin Itoiua v> V.
IB III ttiv •iliniiltiK niitpn Ilia nam U ulselly llin Hiino tliiiu n : tb( OnnlnwUon heflna U ft ; IhC
lalDQI [aloil. ilivRftiir. I> liullautd bf tha illgUUM bvlimn u anil A.
Thv Itma lakoii iiii bj' Uio iiorvmu JmtiDlH In luainR »linie ilu.- Icnttth of aem lietwwD I aiul I
li. Ihcrvfiirc. lii'tlmtc-l tiy thv i|<«laiicr balwurs ft aiu] ft', wtili'li iiu) In mmtund h; lb* limliie-lbrb
EiiTTo below: cmctadnablBtllinUouof ibFiunliv-mk mrmipuniliiul'lJilof ilUW8i«0MuJ.
1. Shifting the electrod«» ftum n point nf the nerve at some distance
IVoiH the muscle to & point of the nerve close to the muscle hiif only short-
ened the Intent period a very little. Rron when m very long piece of nerve
i« lakeu the iliflerence in the two curves ie very email, and, indeed, in order
that il niny be clearly recognized or measured, the Iravellins surface raust
be made to travel very rapidly. It is obvious, therefore, tnat by fiir the
greater part of the latent period in taken up by changes in the miiicle iiadf,
ehanns preiiumtory tu the actual visible ahortenink'. Of cuiirae, even when
tbe ^otroda are placed close to tlie luusole, the latent iteriud iuchideo the
dianeea going nii in the iihi<rt piece of nerre still K-inii biMween tbu elect nodes
and the mu«eular fihrex. To eliuiinHte thi) with a view of determining the
Intent period in the musiile itself, the electrodes might be placed iljrr«Uy on
llie muscle poiMonvd niih nniri. If thix were done, it would be found tluit
lite latent period reninined about the Nime, that is to M>y, that in all case* the
Intent period is chieHy taken up by clinnges in the muscular w ilislinguishod
from llie nervous element*.
2. Hiich iliHerence as docs exist bclwc«n the two curves id the figure,
indicates the time taken up by the propagation, along the piece of nerve, of
the chnnge^ set up at the far end of the nerve by the induction -shock. Thcae
chatigei we have already ^^poken of as constituting a nervous imi>ul«e : and
ihe wTiovc ex|)ei-imeDt shows that it takes a small but yet distinctly appreci-
THE PHK:C0]IEXA of UL'SCLB AST) SKRVE.
Mm Un» for ■ Dervoua iinpiilBe in mvtl atoag m aurve. In tlie figure ihe
dlflWwKW betw««n tbe two laleni iwri'Mln, the <Ii4Uinc« between b ainl h',
MMDi «]in«t too small to ueaMiro Mcciiraivly ; btit it' a Umk piece of iierve
tw iww) for the experiment, am) the r(>corOiiig surlacc bo made lo travel tery
&•!. the iltfl'creDm Itclweeu the diimli'ni nl' tlw tatcnl periud viheu iW iniiuc-
tioo^bock i* MOit ill ut a jMiiui diwv to th<^ ninecle, and that uliea it is seal
ia At a pnni a» far aviijr a* powhlfi frotu itic muitcle. msy be wtidfavtoriljr
■*—w<a in fnictiou of a aocoiid. If the leagili of nerve between the two
|>iijaBi ba accnnUtflf mcaaurtrl, thv nil« at whicb a nervous impuhe LraveU
akmc Hk ixirva to a niii«cl<- ran tliii« Ih! easily cnlciilated. Tbii liru lie«n
fiiuM to bv in the frug about '2'^. and in man about 33 ni«trt» |Kir micoihI,
bat raiia* ci)D8i<k-ra)ily. csiK-ciallr in warin- blooded auiaiulA.
Tboa when a inonteDlary stimulus, suoli ns a ftincle iiiduvti(>n-«liock, i« wnt
iDto a Bcrve oODMit.-tetl with a muHcle, tbe folUininK <!vent:i Inkr pbu%: a
aerroiuitnptiUeiaitsrled in the iten,'e and thin trai-ellingdowii to the muscle
ptvdacBt in ilie hiuk'Ic:. lint the invisible cb&n^ which coiwtitulc tbu latent
pwiod, aerondiv Ibf chanK*" which bring about Uw »horteniiig or cotilntction
proMr.uid thirdly the changes which brin^ about the r*.-lnKatiun and rclum
to tW original Iciii^tlt. The change* taking place in ench of these ihno
pkaaa an change* of living mattvr ; th<;y van with the eondition of tbfi
liTiBg anbatance of ibo ntiuclo, ami only tnke [ilncr ra long as tbe muscle is
•liva. ThoBf[b the ralaxntion which bring* back the muwle to ita original
length ia aamt«d by tiie muach) being loaded with a weight or oiherwiM
Mmcbed, tbia ia not Msratial to tbe actual relaxntinn. and with the same
load the retain will vnry according to the condition of tlie niuecle; tbe
rcljuatian intiit be c.xi^idered as itn essential part of the whole cuotraction
DO IcM than ihr nhorictiiui; ii^lf.
$47. No* only, an we sliall ace later ou, dna the whole eontruction vary
ia extant and diararter according to the rondilimi cif the mnm'Ic, thi-
•CrtBgth tif tbe indiiction-alKick, (hi> loail whidi the muHo i» benring, and
variottt attendant circumntanrM, but the three phaw* may varr inilepeod-
■otly. Tlie latent period may Ih- longer or thorter, the shortening may take
ft ItUftr i<r ahoTter time to rcaoh tbe Mine height, and espccinlly the relaxa-
tioe aiaT be aluw nr rapid, complete or impiTfect. Etch when the same
•Cnngth'of indaction-altock is used the oontrartton may be short and sharp
iK Terr Xaog drawn out. ao that tbo curves described on a n^irding surface
traTeiliDg at the same rate in tbe two c«isca appear very different; and
tUMbr eertain <-irammanc«. as wlien a muscle is fatigued, the relaxation,
mon peiticularly the laM part of it. may be »o slow, that it may be several
lecqaai before tl>o tnuselc really regaina its original length.
Hcntn ifweeay that the duration of a i-imple muHciilar contraction nf tbe
gasUvcaemiut of a frog under onlinary cireunuitafltiea id about ^^ second,
«f wbirh jij is taken up by (hv laivni [>eriod, f^g by the coiiirarliim, and
yfi br the lelasalioD, those must U- takt^n as "round numli^r*," Mated so
ae tn fce vaaily reiBembered. Thr duratinn of eacji pha.->e ti.i well ns of the
vfcola oiiotraciion varies in diffcrrnt niiimnia, iu ditltn^nt iniiiti'Iw of the tante
a^flMl, and in tbr same muscle und<'r ililfi-runt cnuililious.
Thr miiK' If -curve which we have been diiw-UKiiiiit in a curve of changes
IB Ihr Uugth onlv lit the rousclo; Uit if the mu»ote. Instead of lieing sus-
peaded. werv laid flat I'n a glass ptatt.' aiwi a luvcr laid over it« bcllv, we
•hould Hait, u|H>a M'ndini; an induct ion -shook into the nerve, that tlie lever
«a* raarti, ebiiwitig that tbe muscle during the <;ontraciJon became thicker.
Aad, if wr liMkk a graphic r«>eord of the movemente of the lever, we should
in a cur^-o vori' aimilar tu ilte one just diacttssed ; n^r a latent |ieri<Ml
iJm lervr wuiild rise, showing tliat the mitaek was getting thicker, and
90
THK CONTRACTILE TISSUKS.
nftern-Hn) wciul>) fnll. vhouing that the mitsclp vrxs beoiming thiti m^silL
In iillivr woriI>, in coiilracrioii th<' Icswiiinz ot' lh« iiitiM-lc loiigthMisc is
ECcxrtiipniiied hv nn iriomiw croetwiw : iiideeJ. m» ire shnll eec Inlcr on. the
muwie in ranCriicling U not ttiriiii)ieh»I in l>iilk nt all (or only lo au exceed-
iDglv »mnll pxiciit. nbout htUif of ''« 'o^jil bulk), bnl iiiiik«e up for iu
dimmiitinti in length by iiicreasing in its other diaiiKten.
§4B. A single inOiictioii shock if, ne we have »aii], the roost couvenieot
form vl' atimuTiis for proilucinK a simple luusciiUr coDiroution. but thb may
also be obtained br other aiiniiili proriileil that these afe suffidenllv sudileii
and short io their action, as. fur iiisunx'c. hy a priuk of, or a slurp blow oil,
the nerve or iniuole. For the production of a aingic Moiple rauMular con*
tmeiion the changea in the ncrvi' Itndini; Id the mu«ol« miut be of nidi ■
kind a» to coDstitute wlial iimr l>e cullod a kiiiglc iicrvoua impube, and any
slituuluB nhii'b will ovoke a single ni!rv»ti:> iinpulac only may Iw uwd tu
]in)duce a tiiinple mUK'uliir contraction.
Aa a rule, howevur, numt stimuli, olhrr thnn lingh- indiitrtlon-thockfl, toiil
to pnxlnce in a lurrvt; neveral ni>rvoii.« impiiUc*. and, »n we -ihall sec, the
ucrvous impiilxo wlii'rh tmw from tin- central nerv'itiF nyeleni iin<l m> mm
along nerrcsi l"> mu»>I(7<. arc. n' a riik-. iml Bini,'lc and simple, but. i-omplcx.
Hence, am a mmirr of fact, n *i:iiplv niusciilnr contraction t> within the
livEog bo<ly n conipiirntirely rnrv e\-ent (at toiut m far as the *keletiil mus-
clw are concenicd). and cunnol easily be produced onlside the b'>dy ollver-
wine than bv a single induction shock. The ordinsry lorm of muHciilar
coiilraelinn is not a finiple muscular contraction, but the more complex
form known as a teianic contmction, to tbe study of vhiob we must now
turn.
Tehmie Contndioiu.
-^49. If n single inductionnhock l>e fidluwed at a certain interval by a
Swond i>hcick of the t»tnt: strength, the fir»t simple contraction will be fol-
luued by a wecond simple coolraelioii. both contractions being separate oikI
dutincl ; and if the «liocks be repeated n series of rhythniicallv recurrioi
neparatv Nimple conlrai-tioiis may be obtained. If, however, the inienM
betni-on two»hocki< be made short, if, for instatioe. it he made only just long
emmgb to allow the tirst contraction to have passed its maximum before tbe
Intent jwriud of the second is over, the curves of the two eontructiona «UI
iK-dir some fwh relation to each other aa thai shown iu Vig. 23. It will Ik
Fnxa.
TsuiXQ or A Doi DLK Mtxrt.K-' tjiive,
n'liUcibe miuctc (sutrocnomlut of fmKi hm ciisucnl in iIh nm saamoiloo iubwi comiM*
<iMnr. Iiiul iiutliliit InMmnal. It liiilinUHl to Ilio diilUil Uni-i, * Hti>i»l liiilopUuii •lji>-k <rmt
thtDOO In.al metis tlin« Umi (Iw «K:ciud ronlractloii Iietan Jiirt w (he nm HM beRliiiilua loile-
<lto>. T1iaTCU»idcur*cI>M*ulaaUn ftoiiillicafM.M<toa«UiefllnfWiuibeh»»»-UlM.
observed that the second curve is almost in all reepocla like the first except
Uial it starts, so to speak, from the linn curve iuHtejitl of from the baae-liue.
TDK PSBNOUKXA OF Ur9CLK AM) NKHVE.
01
Tbc (ceoail nerruui inipulw has acted on the already eontraclcd muacle,
uiil Rutdo it conlrnii itKoin JuM u it would liav« dune if there had be«n no
lint itnpulM and (lie muM-1e bad been M rcaL Tbe tnouunlraclkius «»
•ddad tofvtber and tlie Icwr u rntHx) iiearly double tbe heigbt ii would
bsva bmo by i-iihcr aloiic. If in the Mmv way a [bird shock follows the
Mooad at ■ auftii-irtitlv •liort iiilirvnl, a iblnl curve U pileil on (oji uf tbe
Mormd: t)i« laruc wilti n fntirlli, bikI to on. A more or Im* similar nwilt
wnaU occur if tbe M^iind cunt rad inn (K'gnn nl another phaw of the tint.
Tb* eombiiiMl t^ltct i*. of eiiiinM.'. grvat<vi whi>fl iho Mcond contmution
WwJQi at ihi* niiixiniiitn <>f the lint, being U-it^ Ixilh befuni and anerwKrd.
llMin-, the mull uf a r<-|H'(iti(>ii of thockx will depend lar^ly OD the
mu of repetition. U*, as id Pig. !24, the shocks follow each oilier »o slowly
MtKiJxTvrx. NBa< iKvecTiMi iaota lurBiTCD turnvt.
llutonv runtrarlion i> oTor, or almost over, before lh« next liegtni, wch
tuttractton will Iw dittincl. or ncwrly dialiiict, and tli«rv will be littlo or no
cDBibiaed rifoct.
If, b'lwever, th* ahucka be raiMatMl mare rapidly, lu^ in Fig. Ift, each mic*
vediaif t^utilndiMl wi)] Mart mini aome {wrt of tlte prt'ctsliiig one, »ud llie
Wer will l>e raised to a greater hei^fal at each contrsc^ioD.
Miana-erati:. SaaiM, Inwcnon BMoot Rn-Miiv Ytott nirfiM.v,
If the fre()uency of the aliocks be eiill further iacreaseil. as in Fipf. 2$,
lit riae iluc to tbe coRibiaati'Mi of iMnlrction will be still more rapid, and a
Waller part oi each conlnntiou will be visible on the curve.
la cacb nf tbwe three curve» it will be itoticed thai the character of the
nrrt dwngw •omowhat durinu; iu derelopmeoi. The change ts the nwult
of cocatuneing fatipie, cauMid bjr the repetition of ihe contractiofin, the
lat^aa manlfntinf; iiiielf bv aiwt inereittiii^ prulotiKation of rach ontrac-
•bowti especinily in a delay of relaxation, and by an increaning dioii-
in tbe height i>f tlie conlrnclion. Thun, In Fig. 24, the coninctiona
distinct at hr^. bcc«me fu*ed hiier; the fiflh contraction, for iniilance,
a pcoloBged to that the ilstb bvgina beforr' tJiO lever has mcbrd the ba«
Im* : tM the *ummit of the sixth t» hardly higher than the BUininit of (he
fifth, nace the sixth, tliuttgh itartiiig at a higher level, !■ a somewhat
weskkrr contraction. 8ee also, in Fig, *j'i, iba lover rises mpidly at first but
92
TIIK COXTHACTILE TISSUES.
more stotvlv nllenvard, owiiij^ to iin iQcroii^ing diminution in the liviglit nf
tli« ein>;le oontr>iclioit«. In i-'ig. '2G itie iiiL'reii]«nt ol' Hm of ihv ?iirv<' due
to each cuutraclioo diiiiitiiahes verj* rapidly. And though the lovor does cna-
tiDtie to ri«e during the wiinle eeries, tne oeoent ttft«r uliout ilic sixtli cm-
traction In very t;'''i''"al itidee^l, mid the iodicntiona of tbe indiviiiuB) con-
tractiuiia are luucb lueii marked tlian at lirst.
Mi>''i.t<vitv& SmoLR bantctuai iuocx Rtnuicii Snix Monx RATHni-r
Hriice, when shocks tin: rp|j*:iit(.'>l nitli sufficient raiudity, it mutta that
alter a certain number of Hbocki, the aittcciMHliutj; iiiij)ul»e8 do not caiue hny
lurthur shortening of the mnicli;. any further raisiof; of the leTer. but
merely keep un tbe i-ont ruction alrcnily exixtiog. The curve thus reachea a
maxiiiiuin, which it maintalnE, subject tf> the depressing eHects of exhaus-
ti<Hi. «(> Icing as tJie shocks are n^pcitlcd. Wlieii these cease to be given, the
muH'lv mtums to its natural length.
Whun the .shocks succeed each other elill more rnpidlv tluui in Fig. 36,
■ be individual contractions, visible at first, may become fused togttbor and
wholly lost (o view in the latter pait of the curve, fflieo the snocks suc-
ceed each other still more rapidly i the fecond contraction banning inltw
ascendinfc (Kirlion nf the fintt) it becumet difficult or imnonible to ttMB
nut any of the nngle contractions.' Tbe curve then descriWl by the tw^r
nruc* ratamD wru Tire ohiihiibt Hwano tmaRi-ma or tn ixin'mosMucMa'a.
iR(<oiii>liriii»irilwau»vsUlasitiiiily.)
TIic lnl«mi[it«l ciUTeiit U lliraw> In «l 0.
Uof the kind ibowo in Pig. '27. where the primary current of an induction-
machine vm rapidly made and broken by the mit^etic interruptor. Ki|{. \^.
• Tbe <•>• nlih nhub Oir iciilivl.liul «(niinuUQM «m Ii* aMilc otii ilu|it»4t la iMn. II uwri
Innlly boBtld.iB Ibenpwlnr '■■(■>"'■"''' "'*'""'*"*' •"•«>■» mnl.
TUB rilKNOUENA or MI'SCLR axd nkrvk.
98
I
I
TIm l«T«r. it trill bo obfm-Ml, rian nl a (ihe roeording Hirface is tntrolling
too aluwljr to nllow th« Intent p!^ril>>) tt> ho dialin^nicbi'd), at firat very
lapidlr, in fact iti nu unbroken bikI iilmnrt s vertical line, sinl mi v«ry
ipffrdiW rrMtMfl tbe maxiiniim. nhich is iDttiDlJiin«d m Iod;* m the sItocM
aoattouc Id be gircn : whva tbew- rvaso to bi' given, tbe curve <le6i-etid5 at
fltiC nrj rmpiilTy and tben more nod more graduBlljr toward th« baK-lioc,
«Uel) ]t mchea just at the end or ihe li^ure.
Tb^ onxtrtion of mtiH-le, broiisbt about bj rapidly repeated aliockit, thia
ftiaioB uf a number ul' simple tnicliea into an apparenllv unoolh coutinuout
•Act. li knuvB aa lefamu or utaiiit eontraetutH. 'i h« afwve fteu ar« mott
daatli' (liowQ when induct ioii-obticka, or at leaM galvanic currents id aonM
(bfTD or <i<ber, arc enploved. Tbey are tevn, however, nbut^virr bo ihf form
of rtiniulti* enploTMl. Tliut, in the cow of nirchaiiical Miniuli, nhilc a
Bingtv i|aick bloit tuny eaiuie a ijngb; iwilfb, a pruiMunL-ed telanux may bo
obuiovd by r«i>idty ntriking iiuccCMvely frtwh {mrtionii nf a uervi;. With
rbcmieoJ Minintatinn, n.* nbcn a nrrve in dip[>e(l in noid, it a imponiibki to
M«ra a iiionMjiitarv applicalton ; bciic« t«taDu», generally invgular in rhar-
■cc«r. i» tbo normal n.*iilt uf thin mod* of slimufation. tn tho living body.
tbm diHirartions of thu fkclctnl nniFcle*, bnnight aboat cither by Ihe will or
aCbanriae, an g«D«rsUy (etanie tii vhararler. Kvcn very short sUarp raove-
Wl at I. micb as a sud4l«n jerk of a limb or a wink of tlie eyelid, are in mlity
ejUUBttlti of telaiiu!' of snort ditrntion.
Ifiht fever, instead of being fastened to the tendon of a muscle hung rer-
tleallT, be laid aenea tbe belly of » tuasole placed in a hortxonlal position
and ihf wuscte be thrown iiilo tetanus by a rt-jwiilion of indiK-tion-shocka, it
will be seen that each »lii)nenint; of tbe muiu'le is aocompiuiied b^ a corni-
■pawdtpg thickening, and thai tlit? tulal shurieniiii.' ^^f 'bt- tetanus is accom*
pntl^*^ by a r<>rrr"])"nditi^' total lliie kitting. Ami, imler^l, in lelanit* we can
obwrrs more easily than in u Hltiglo oontraetiou that Ihe muscle in coniraet-
htg cJuiagfa in form onlv'-not in tmlk. If a living muscte or group of
wpaclfti be pla»<l in ■ glaaa jar or chamber, (he cIomvI lop of whivn is pro-
IniMial into a narrow glasB lube, aud tlie chamber be filled with water (or
plWMbly with a solution of sodium chloride, 0.6 per ocot. in strength,
noallT eallol " normal *aliDe solution," which is kas injurious to the tiuue
than nm|)le water) until the water rises into iIk- narrow tube, it is obvious
tliai any change in the bulk of tbe muscle will be easily shown by a rising
or UIIbc of tlie column of fluid in the oarraw tube. It is found that wbieu
lb* mmole i* made to contract, even tn the most forcible manner, ihe obange
of l*f<ri in tbe beighl of the column which con be observed is practically
inrffiiiii am : there apfiears to be a fall indicating a diiDinnliou of bulk to
lbo«xl«iilofal>oul»ne ten-thousandth uftlw total bulkufibemuM-le. ^hi that
«« Btay fiiirly wiv that in a tetanus, iu>d henoe in a ximnle ountraclioii, tbe
l^Broing iif tlii> length uf ibe muscle causes a c<>rrt?|iunding inerva>e tn tliC
otWr rlin-<-iJoM : tbe aubstaace of ihi- muwobr in diH|ilaccd, o« diminisbad.
f M. .Si far we liaresfMiken simply of an indiietmn-aliock orof inductfen-
■bnaks withool any referencv t» their Mnmgth, and of a living or irritablu
moscU viihoui any reference to the degree or extent of it* irritjtbility. But
indvcti'ia-shaeks may vary in i4rcngt£, and tlm irritability of the niuxcle
nsr Tarr.
if «T slide tbe secondary cnil a long way from iIh^ primary* coil, and thus
anke use of extremely feoule induct ion-shiicks. we shall probably Hnd that
iteBabocka, applieil even to a <)Liilc fn?>h miuch- nerve preparation, produce
SO oMtlraottoo. If ne then gradually *lide the K-conoary coil twarer and
SSWar the primary coil, ami keep un trying the etfecis of tbe shocks, we fhall
i»d tbit kftar a'while, in a certain position of the coils, a very feeble ooo-
»4
THE CONTBACTILK TISSUES.
IrHctioii nmkiw iu sppcaranoe. As the tocondary coil comn still nnrar to
iitv [irimary coil, the coutniCtiotiH gmw grenlcr anil greater. AD«r n wliilo,
however — nml that, indeed, in onlinnr}' circiiiniitiiiiri^, vcrv ipccdilv — incroms-
injf rha^ntreiigthof the*Iii)ck no loiiu^r ini.TniuN.w thu height >rf Lht: roninidion ;
the msxiniuni contnu-linii of which the iniiwle u cnpahlo with «ucb shocks,
Imwcvct strong, hiw Ih-wi rwiohcl.
If wc 080 a tetiuiixing or interrupted cnrn'nl, we ehall ohtjiin ihe same
gcnernl results: wo Ria)-, nccnn)ii>g la the stnmglh of the current, get no
eoNtrmcli'on at nil. or cuntrncttoRfi >.'( vnrioue extent up to a maximum, which
eiiunot be exceeded. Under fnvomble eonditiaii» the maximum ooniractiun
niny be very considerable: the shortening in tetanus may amouiit to three-
fifths of the total length of the muscle.
The amount of coDtmctiou, then, depends on the strength of the Mimului,
whatever be Ihe stimulus: but this holds good wiili in certain liinilaooly;
lo this poiDl, however, we shall return later on.
S 91. If, having ascerlaioed in a wrtectly freeh luueele-nerve pre|«rnLiun
the amount of c;nlrai:tiiiu produced by this aud thnt eitreuglh of »iimulu«,
ire leave tlie preparation by itself for some time — auy for a few hount — and
then repeat the observitliuus. we shall Sud that stronger stimuli — atrongor
^flcks, for intttanee — an> reijuired l» produce the «iue iinmunt of cnnlractiiu
lift before; that in to tiay, tlie irritability of tlie preiiuretiou, the power to
respond to stimuli, hti^i in the meanwhile d i nil iii shed. Af^r a lurthor
iDlorvaJ we sliould find the iTritabilily still fnrlbrr diminishv<l : even very
strong shocks would he unable tu uvgke ci>T>travliiini> lU' large n* thoet- pre-
Tinush- caused by w«ik shocks. At laat we ■■■hoiild lind that no shocks, no
stimuli, however strong, were able to produce any visible oonlrnction what-
ever. The amount of •■ontriiriiuii. in fuel, evoked by a stimulus depends not
only on the Atren^th of the stimulus, but also on the degree of irritability of
Uie musclu-Dorve preparation.
Imniediiitely upon removal from the body, Ihe preparation poawnoca •
certain amount of irrilabilily, not ditTerin^ very materially from that which
the muscle nml nerve puuewM while witbm mid formioK an int(!(cral [Jitrt of
the body; but after removal fn)m the body the preparation loaeairritabilily,
the rale' of lorn bein^ dep)-iidcnt ud ii variety of cin-umatanoea; and thixgotw
on until, since no stimulus which wi- can apply will give liae to a oontraC'
tion. we say the irriuibilily bus wholly dinappctAred.
We might lake this diiappcaranoe of irrittibility bh marking the death of
the preparation, but it is followed aocmer or later by it curimi:! change iu the
luuBcle, nhich is called rigor mortis, and which wr 4iall Mudy pnveiilly : and
it is convenient to regard this rigor morti." iw niarkinu; thu death of the muscle.
Tbe irntable muscle, then, when slimuUlcd eitlicr directly, the stimulus
Mail aiipliixl to itself, or indirectly, the slirauhi:> being applied lo iis nerve,
reapond» to the elimulus bv n change of form which in i,«w,-Jilinlly a shorten-
ing and tliiekening. By the »horU;ntng (mid ihickiiiiiig) the muscle in oon-
trndiiig i* able to do work, to move the |uiri« to which it is attached ; it thus
tol* free energy. We have now lo study more in detail how this energy
i^ set fn-e, and the laws which regulate it« oxpvnditure.
Ox TIIK C'IIaKQIA which TaKK PlAPB in a Ml'SC'LE DCKIXO A
■ OoNTKAt-TION. ^M
B The Cbanffe in Form. ^H
^fiS- Tfie yrat* tirveUirf of mimrtc. An nrdinart' skeletal iniiscle onn-
nsta of e/onenturjr muKUjibret, bouod together in variously arranged bttodles
CIlANOES IN A UL'SCLE DrRING CONTRACTION.
95
.acctive tMue wliicli carria bli>odT««el>, avrves, ■itd 1]rmphatie& [Fig.
The Min« cMiDcdlve timoe, IwvhIm Aipjilyin^ u mora ur tcM ilistinot
tiffiaf fctr Uw wboU mutclc. Toraut thr two ■■tiili ol' the iiiiieclc. Iteing Iters
(mn KUltv. lu «l>ef« tliv miitK-lc iipix-im lo be iJirectly nltached to R
Imoc^ aai % •null amuutit oiilr ot wnncctivo titwic joins th« niusoulitr fibrae
to tbr iicri'ii4<-uni, M>tnetiiii(4 nliuiidimt, us when tli« cunnoctive tissue in
whkh tor muvnilnr tilinv imiiHilintrlr end is prolonged iuUi a tendiMi.
EmIi flkaiCDlarjr fibre, nhich viim« ev«it in the lunnimiit in li-nL-tb and
brMkdtb (ta ibt tng iW dimrn; ions vnry rcry iri^tely <, but may he «aid, on an
Avcngt, lo be 30 or 40 mm, in length and 'iOu to 30" in breailtb, coosieU of
•a ifiiitic botnogtDMiu^ or fnintl}- librilhiied «Ii«aili of peculiar nature, the
mrt»hmma. which embrmcn »nd lurms on enrelufX! fur thi: ttriaUd miu-
'jr m^antf tiiihin. [Fig. 29.] Each fibre, cj'lindrical in form, giving a
rin. Ifc
. MmoM rwiii Till nnutn-HMTOtD im Han (mapiUWil M ilmMi u. tiWi-
•■I tMtatfitan: hlMalcDhn; r. iiiwtnal pmmr^ami d.Mn.
nk. w — Fuajoan or fttRmir KitMartmy Pibmw. tiitdrDM i Clurivt n orMiran DnacnoM
I «■ aiMBHcrx.
Itlamcr. Ttao Uwcltodlntl aiiAitiianvnennMaKhiitfaMvii. Aotnc lansitndtnal
r anil *klrt ihkii tin nM, u>4 an out aHMImunD IMin end to *n>L Tliln naiilu Itom
■ of tlwMtlllr. caWllaMfonMfrcanncaBMlisrbr TlotonMiltlMtookSiinUl
.■Mlnwkad tgtimmwwtlla«»crmt||inrlilt>iotlnncoBUieatrp:t'<*miiijwiHwo
■ nnmwntf inminwl brtha MfaimioddiifleaMllv mwnMsbly nucniiMj: at r* iba
I inBmam Iiiimm* all pntVdlx ncnitanf. •'•'I ll>r mrlUilFd tfaicn ivrMilr nMaiifo-
bar M ■* llw konlcn m ^aUiifsd am] the tfiaaii Inail-Iitr. Wbcii mon (Ilnlatl and deDiillp.
am HaWfc tWBM llw fc«»it ut Ihao app^mneai.
•lai«W& Tka Imgllndlaal llnoi an tnmilj TWbl* s. inconiplrtr ftutor* bHOB-
niDuatf • 4Ih) wlilch •(>«•<*<• aCTCB Ibe lMer«a> and (ttaini the tnvfniniHmJl
TW>d|*«>4*utibc»iif tUa diM an •*•« tu t* ■iuuMl)' jmnulu. tte inaaiih*
laflnitir tiitli»iiilckM«<>rtk«dlMaAd tn ibr dMaaov Mratvn Uw Mhi laatHuilliial
■tM bomIt •StiiibDl : y. ilttaclMil AIM. mut* bifklj tnacnlacd. alioaliw lh«
r
KwiiMiIti
or laM circular outliiK in iransverw aection, ^nerally taper* oS at each
^d in a cunJeal fonn.
Ai oub «n<l of ihtt fibre the mroolemma. to which in life the muscular
■abMuKV i* lullimnl, bvoomea continuouH with fibrilUe of connective tissue.
Whao Uw «tul of lliv fibre li«« ni the end of the muacle, these conntwlive-
^mam fihrilln |Maa directly intii the tendon (or into the ueiiaateuin, etc.) and
ia (nat* CBMB of uniill muvctui which are no longer tnun their conatitueut
Ana, CMcIt fibre may ihui join at <«cfa end uf iLMlf. by means of its aareo-
Jmmos. the t«Dd>in, ur other ending of the muMle. In a verr large number
«f ■iitlw. b»wovcr, the niuM-Ie >• liir lunger than any of iiH fibres, and there
m^ be «Ten whole twndica of tibrca io lue middle uf tW muscle which do
X
96
TrtE COVTRAi-TlLK TISSfES.
DOt renc^ to either cud. In such com the ominective tinuc in wtiich tbt
t»n!o\emma ends is L''.>ntii)UouR with ihc couiicctive Iimuo which, niiiRin^
l>etweci) the fibres mid between the I>uiid]e8. hinds ihe fibm intu email
biiudles. aiid the !»Jitaller Iniixllcs intu ]ar)^r buiidlca.
Tlie conlraetioo of a riiusele is the c-uutracliciii of all or eome of il»
cknieiilury librea. the timneelive liiaue Wiuu paaiive; heoce while thoee
fihrta of the iiiuttcle which end direetly iti the tetidoii. in coutraeiiu); pull
directly on the teudon, ihuoe which do Dot ho ciid |iuU iiitllreetlT on the teudoo
bv nMuini> of tbe cunnei-live liwue between the bundles, wbicb connective
tfariie ia <^AUtiiiuoui« with the teiidou.
Th« bloodv«aseli run in the coiin«ctivtt tiwue hotwet-n the buodlee and
bctw«cii the Gbres.aiid Ihe capillariM form mori; or K«> rectangular networks
i mined lately outside ilie narcoleitinin. [Fig. 30.] Lympbaiic vc«els al>» run
iu the coDne(.'ii\'(.- ti»iui\ id llio lymjih tipncv? of which they bcE'i- Kacli
iiiufcularlihre isthu»Kiirrounde'l by lymjih fpnt-c* and capilliirj' bti>odvca«eU.
but tlw active uuiBriilar MutNitancc of the Irbre is »(<paralG>d from the*C by ibe
uroolemnia; hi-in-*- ibe interchange between the blood and tlie CDUKiilar
8ubaUui€« a i-arrieil on backward and forward througti the cajiillary wall,
tbrou|rii aomo >>t' tho Irninh gpacea. and through the earcoletnina.
Each luusclv i» «iiiiphcd by one or more branches of ner\'e# ronipowd of
luedullalod fibres, with n certain proportion of n(m-iiiedullate<l Hbm. Time
branehos running in tbe ci^nneoiive ttEsue divide into Bnmller branehea and
IwigB between tbcbuniUesaiid fibres. S.i[iie of the nerve fibres are dislributMl
Ki tlte bloodvcKels. and otherv end iu a manner of which we shall s|)eak later
t>n in lrealin<; of muscular ^nBatimid; but by fur the greater pan of tbe
iiiedullaled fibres end in the muHeuhir tihrRt, llicnrraiigement being such that
every muscular fibre is ;uppli(h] with ut least one niedullat«d nerve fibre,
which joins the niuHculnr fibre somewltere abotit
the middle between Its ttro ends or winietimei
nearer one end, in a special norve ending, of
which we shall pi'eseiitly havo to e]>cak, called
an fit'l-p/ate. [Hr. 31. J Tbe norvc lihn"» thua
ii<-»tinei:l lo end iu the niUHColar fibre? divide
as they enter the luujcle, so that what, as U
enters tbe luusole, i« a single nerve fibre, may,
in.., «.
r!
I MiM-LE, ^juicn. mini moaiuK or Ooul
CIIA>*fllI8 IK A XL'SCLK l>l'lll>'0 roNTRAl'TION.
97
I
by (IWiilingi. end iu> »cv«nil ucrve fibrcB In MrenI musctilnr llttrt^. SntiietiiDM
twoDarwlibttt j<^iaoii« tnu»citlnr ftbre. line in thin caw (lio i>iiiI-i>1bi« of c«rh
tiem ^br« i* •Ull lU totao duiunri' rniiu lht> end of lh« tiiunciiliir fibre. It
Mknn tb»i wtiCD ■ mtiMulMr fibre i> MimulAtiHl by nioiiiu nC n nnrvc tibrc,
Um DfTVow imputw tnNfllin^ ilon-n lh« ii«n-« fibiv fulU inln ihc iiiii«c>ilftr
Sbra Oflt It MM rD<) but Hi nlioiit ibi mitbllp; ii Ik tin: ^tilltlK^ of llii' fibri>
which it mitbeui firet by the ncrvout iinpiiW, anil l)ir i-lninKiv in tl>u muwTuUr
mImum* lUrUd in the nii<t<)l(? of tlu' miuciilnr Hbri;. Irnvt') ih«Dc« to th« tvro
«iKlt of tfao fibfo. In an onlinary ekckitil nitieck. h»w«rcr, tm wt have Mi<t,
th« flbrw ami Imiidlm of libm Wgiu and end al dilTiTcnit di»Iiinc» from l)i«
tads of tb*> mu«clt>, and the nerve or ncfvr» gnin]; to ibc mi»ol<> dividv itnd
»pwd oat in th« mii6clt> in sucb a way ibat rlie end-(>lat«a, in which tb«
ntdividoal fibrea of the nen-e end, are distributed wideir over the rium-I)? nt
van* tHflbrail diataacw (hini the endp of the miifcle. Hpnoe, if we iiippuao
• '(m}e nervooB impulse. Mich ae thnt eeserated by a tinjjle induction -shock,
or KMrkaofsucb impulsei* to lie started at the ^nie time at sonic nnrt of the
■rank of the iMrve in «ich of i)i« fibres of the nerve ^'oinj* to tne muscle,
the« impulMa irill mu-b rery diflerait ports ofthe muscle ai about the same
line uia tbe eontrai-ttons u'bi<^h Ihey set >r<iin|; will bcf^n.so to speak, nearly
all oTar the whole uniMk at ihv oiime time, iiud will not all stan in any par-
liealar tone or area of ti»e muscle.
f SS. 7%r •I'liw of eonlraetion. We have leeD, however, that under lh«
iaBlMPca of uniri th« nervi? fibre i^i iin:ilile to excite cnntmctiona in a mu*-
rsUr flbre, altlmugb iIk irrilahilily iif ibv muicular fibre itmlf i« retained.
Hmmm in tt miucla poiwned by timri the contmotion bl^^^« nt that |uirl of
tbe BiMDnUr Mbaunce which io finit ntlvctcd by the Mimulns, and vrt inay
•urt a tininnion in what |>nrt of the mutvlc we (ilouc by properly placinjr
lb* plirirodea.
Attar niinclea,aovh for inetaooe as the •nrforiiM of ihefras, thouf^h of some
ijctfa, an compneed of fibres which run parallel lo each other tVom one end
ibr miucle to the other. If such a muscle be poisoned with uraii so as to
If the action of the nervce and itimulaied iii one end (an iuducliou-
Atk vui ibroufib a pair of electrotlM placed at some little dblaoce a|Mn
tnm each uiber at the end of tbeuiUM^le may lieempluyed.biit betior remlia
■rwobuIneH if a mo<le of mimulaiion. of which we shall have to Mx-ak jirm-
cntlr, vtL thr application of the " cunstani current," Iw Bdo]>t«il), lh«- con*
inctinn which emueaHartit from ilwondnlimulaled, ami Inivcis ih«nco along
lb* BiMcle^ If two lever* tn; m»d<.' t<i rc«ton,or be su^^jirnded fnim.lwi [Mirta
nf Mch ft tORDcl* plac«d horijiiinlally, the pnrla being at n known dialancc
fwvn* iind iVoni the part (timnlatcd, iheprwgrewof the conimctioa
BMlr U in
The noreiDcnl* of ihr li^%'cr* indimtu in thui cn,-«i> lh<- thickcnio)! of tho
flfana which U takini; place ut the juiri* '•» which Ihi- lever* nvi or to which
ibey an attached; aiid if we take n graphic record oJ' ihme movements,
briegfaif the Iwo h>ver! to mark, one immi-diately below the other, we Khali
felt that the kver near«r the part «(imulatcd hegiiw to mo^-e earlier, rnachea
lu amimura rarlier. and return* lo n^t earlier than dric« the further lever,
TIn nAlraction. started by the ctimnlus. in travelling nhing the oiuacle from
iba pari iiimnlated raaohea the nearer lever some little lime before it reaches
the (briber lever, aud has passed by the nearer lever some littto lime Iwfore
t( baa fauaed bv the further lever; ami the fnrther apart the Iwo levers are
I In Miaali I will Iw th^-diNVrence in time between their movements. In oiher
wiinU, thv ' ti iraveU aloiij; ihe muscle in the form of n wave, each
jmn uf tbi II. succmnon from the end Blitnuluteil swelling; out and
ibartaniog aa the contraelioo reacbea it, and then reiuniioK to ils oripnnl
98
THB CONTBACTILK TISS0]
atnlo. And nhat in true of the oJlectiini of iiHratli>l fil>ron nhich vn- cnll tl
RHiwlc U alio Irii6 uf ciu'h fibrv. tar tin* s»Flliiij; ut iiiiy [ictrt uf tin- inusole if
ooly the ftiim af thi^ •wolUii); ol' lli<; iinltviiluiil &\in» ; aiKl if irv wera aUv to
uk« n Hiiigle long fibrv nml MiniiiliUi' it itt one vml. we »hoiilil be nble,
iindrr tlif mierwcopc, lo ««MniwrIling or bulging ncciimpnniitl bv» eniTfr-
■pnn<)iiig»liorlciiiiig, I. r, toMu n con traction, «vrccpuluiig (hetibre fronj end
to end.
If, !n (he graphic rccnnl of the two icvtn jiiM mentioned, wo count tbe
Dumber of ribmlione of llie tuning-fork which iDlcrreiie Wtwecn the mark
on tho record which iudicatce the ttcginning of the rise of the near lever
FthBtis.lh«nrrivHl of the contraction wni'cnt lhi« lever) and the lunrk which
indicaim the Wftinniiig of ihc ru« of the tar lever, thia will give ub the tiuie
which it has tnken the coutraclian w.tve lo trtivel from the near to the fttr
lever. Let uh suppoM? lhi»io he 0.006 wcoiid. Let iia euppose the dttitanve
between llie two levers to he lo mm. The coiiiraclion w»ve. then, hn* taken
0.006 wcdnd to travel lo mm., that is to sia.v it has travelled at the rate of 3
oieirM per tti-ruud. And indeed we liiid l\r thia. or hy otlier loethodn, ttiat in the
frog'* muHeteD the cuutractiou wave dues travel at a rale which may Iv put
down Hf from '-i tn 4 nietrai u second, thmixh ii varie« umlrr 'litfcn'iu mndi*
tioHH. In ihr wiirni-hliiiHhxl nmmiiml the rate b aonewhat greiiter, nnd majr
pnilmhl y l>e put down tit 5 roetrcii a iccntid iu the exdaed miiH'lc, rising poe-
*ibly to 10 niein:'? in x nuivcle within the living body.
If, ognin, in thu graphic recoril of the two leven we couni, in th* esse of
cither l«ver, the number of vibrittiona of the tuning-fork trhieb intervene
betwe«n the mark where the lever liegin« to rise an<l the murk where it ha*
finished ilH fnll nml returnni to the ba»e-line, we onii menxitre the time inter-
vening between the contraction wave rr-aching the la\'er and Icnving tbo lever
on its way onward, Ihikt is to say wc on mi-asurv the time which it hB« taken
the contraction wave to para over the part of tbe muiicle on which the Ivrer
is rwting. Let ua aupjHHe this time to be, say, 0.1 Eccond. But a wave which
iit travelling at the rate of 3 metres a eecond and takes 0.1 docuiiiI to paw
over anv |<oint muhl lie 300 mm. long. And. indet^^l, we find that in the
trva the length uf the couti-action wave may be ]>ui down as varying froni
200 li> -too mm., ami in the mnmnial it h not very dillWent.
Now, a* wf have naid, thL> very longt«l luiisciilar tlbre U Jttatei) to br at
Bi«iet •inly about 40 mm. in li^ngth ; hence, in an ordinary cmniraction. during
the gmiter part of the durutiou of the contraction the wbolo IflUgih »f the
Gbra will be occupied liy the eoti tract ion wave. Just at the bo^nlug of tlie
contraction then- will be n time when the front of the contrwitiOD wave haa
reached for instance only halfway down the libra leuppoting tbe stlmuhiG to
be ap(died, as in the caae we have licen diaeuBring. at one oad only J, and jiiat
at toe end of (be contraction there fill l>e a lime, fjr instance, when the »>ii-
traelJou hna lell the half of the Hhro nest to theatimuUia, but haa nnl ret
cleared away from tho other half. But neartv all the rest of the time every
(lart of the fibre will l)o in some phaae or other of contraotion, though the
fioriit nearer the stimulus will be in more advanced phaae* Ihan the part*
iirlher from tlie stimulus.
This b true wheu n muacle of parallel fibras is ntiniulaled artificially at
one end of the muscles, and when, therefore, each libri; i^ Htimulaiisl at one
eaul. It ia, of course, all the mure true wheu a muacle of urdluaiy coo*
■tnictiou b «timu1ate<l by tueaun of its nerve. Tbe Hiululu^ of tli« nervouf
inipulMr impingM. in Ihia caae, on the niii:>ch' lihn; at the cnd-plat« which, a*
we have aaid, in placed toward the miildli.- of lh<- tibrc, and the contraction
wave traveli from the end-plate tu opposite direction)' lowunt encli end, and
bw accordingly only ahuui half the Ivngih of the librr to run iu. Ail the
\
CtlAXaRS IN A MUttCLB UL'RINO CONTUACTION. 99
mure thnntot*. must the whale fibre \» io a state of coulractioii at the wnae
tian*.
It will be ohecn-e<l tliat in whni has juet beeti said tlie oonlraclion wave
tarn b(m lakeD lo include not <ml>' the onlrnetioo proper, ttie ihiokeDing
MkI •borteniii)t, but also the reUxnlioD und return tu the DatuntI form; ibe
Ar>l [Hirl of tlie nave up tu the suiuiuit of the creel comnKHuU to tb«
■ li-'rtruitii; nnd itiickinio);. the decline from llie siiiuiuit ouirard CorKe[Kin<k
bi Ihe rvlaxaiiiin. But we have nlrendy iDBiHUHl that tbe relaxolion is an
— aatial part of iho nhulc^act; inde«(l, in a certain ^eoae, aseaeentia) ait the
•hurttnlDk: il*cir.
I M. JUiiiuir ttnitinrt of mutevtar fibrt. 60 far we have beeo dealiug
m\ilk (b« miiwie a> a wbol« and ai obM.T\'eil with tiie naked eye, though we
ban itKtdeoialljr 'IKikeD of fibre*. W« have noo', confining our Htteniiou
csdaiiTely to tikcli-tal niuw:]i.'ii. lu coibud«r nhat tiiicr«ACo|>ic changut lukt^
placv doriDK a contraction, what arv tbc rulatioua uf Ihe hirtulogicut feature*
■if tba muK'iv fibre Ui tbe act of eouumction.
Tba liHiK cylindrical fbcnth of snrcolomma t* nccupicd by niiiMit sub<tancc.
AfUr ilMkUi lIm: iiiiitlc nultFinncc may M^pnr.ite from tbc Knrcolcmnia, U-4iving
lb« Wucr as a distinct 'lifAlh, liul during lite iho muscloaulwUnce is adhervnt
tM 1^ iknvdrninia, h< that no line of iicpnnUion belwcMi tlit- two can l>e
■uub oat ; thu niovenients of tin.- one follow exactly all Ui« movement* of Ibe
other. ,
.■<^lt-m) in the rniiM.-le snbslHDce. but, in the mamma), lyiii^ for the moi>t
pan rfnf<^ imiU'r iIh- ■^nnsilt'oiDin. are n nunilicr ofniirtei, oral in sha|ie. witb
iImi' ites parallel to the length of the fibre. Around each uuclein is
a ii< 1 i>f KrMnular-Iooking suDManee very similar in appesran<'e t<i that
faraBMrUw iKxIy of a white biood-corpn^cle, nnd like that oftCD spoken of
as tnuAnnilaiod prolophiain. A shmII i|imiitity of cite hbdm stmnular imb-
Ksam u pml'iuji^il fur iMime ilixtniicc lU a narrow coui<'jil Mreak from oach
ta\ of ih>? noclcii*. alon^f the lcn;j;th of the fibrv.
With the csixplion of tbtM nuclei with tW-ir grauulnr looking bol and
4* end -plate ••r i;nd-nlalc8. toW precvntlv dvccrilx^, all thereat ortbe»|iaG«
tpfliwail by iIk- snrculetnma fiMU on« en<f of (he (ibrc to lliv other apficara to
b eompinl bv u {Hvuliar rnnlerial, lirialtd muWr /mtulaiiec.
It m cnlluJ Hrialett hccaiife it is marknl out, ami that along tlio whole
laqph of Ihe Sbrv, by transvene liandc [Pig. 3:!], stretchiiig right acrom the
[Plu XL
s
Di«i«t.icNjiTR RxraaoT^raw or jt Hcaoia^uR.
■Mil— 1« f rlwii, BB«<Wli<of>l>«»»*li"<*'M«l»roJi! tfc Anld «Wuw«.I
ftiV, nf aubMance which is very Irait^parenl, hriyhl »abiUiHef, altcmaling
•itb Mrailar bands of substance which has a dim cloudy appearnncv, dim
aiMcnw; llial b to any the 6bre ia marked out alonp^ iu Wliole length by
ttUnMn brifhl baml* and 'lim h'lndt. Tlie briglil bands are on an average
about \ por !■•> » and the dini bands about 2.6 « or ^i •> thick. Bv care-
hi toattiiam, both bright bandi ami dim b«niis may be traoed tlirough
tha wkJa laickiMM of tin: fibre, .io (hat the wrbolo fibre appears to be com-
pMd of bdskt disc* muI dim Aae* placed alternately one upon ihe other
ala^ i1m whole length of tlie fibrv, tbc amingGiuent being brviken by ilie
I i -ON rRACTII.E TISSUES.
• 'k «kw>iti '(l)iv » trvaled with dilute mineral acids it a very apt
. ,. ..|- iMu.ic>n(;I> iucvdiiM-s [Fig. 3'2], the sarcolemma being diaaolvM,
■ ■iv.Mi ■& juruiv c\> divide into fragments corresponding to the dines ;
. :r4 ud,_<> .iiiiR. b< obtxinet] ao thin as to comprise oaly a single dim or
■ „.:. >H.iM, 'i-tJiru twuu) with a thin layer of bright substance above aod
\ ■.. I, ,\m •■iiMtHipf having taicen place along the middle of the bright
«t 'K-> '«uKt\.>i «iih tvrtain reagents, alcohol, chromic acid, etc., the fibre
■ >,.< if'. I" ^'U\. M)* i.aiid the splitting up may be assisted by " teasing")
-. .„.!•.. iiiiikl.k luivi ivliiiuits of variable thickness, some of which however
..-v v vv4\luij;ly thin, and are then sometimes spoken of as " fibrillK."
n.lt .iv'H' ^(ini.'ti Hiid tibritlte are artificial products, the results of a trans-
...^ >i k-u^itkidinal cleavage of the dead, hardened, or otherwise prepared
u.i-Aiv lutvitMUtv. 'I'hev may moreover be obtained in almost any tbicknea
1 i-iiiiiuvw. Aitd thvw tliscs and fibrillfe do not by themselves prove much
>, >>gi.t iht- iWt thai t\w libre tends to cleave in the two directions.
Ihv lis Ills lilirt> hiiwfver, though at times quite glassy looking, the bright
\.>i.U ft|>t«-Mi'i»K likti trans^tarent glass and the dim bands like ground glass,
'• ti othvi iiim*> marked with longitudinal lines giving rise to a longitudinal
..•i.tti^'ii. H<u»>l iiiim iHiDspicuous and occasionally' obscuring the transverse
A'MLUti III lht< iiuim'Im of some insects each dim band has a distinct p&li>
..t.'v' •t|>)>t>HiMii<H> an if made up of a number ot " fibrille " or " rods " placed
.i>lv t>t ■»!>' ami iiiibtilded in some material of a different nature ; moreover
>k.H' iDiiillii' or nidd may, with greater difficulty, be traced through the
ixi^Ki bit I >d 4, and lliitt at times along the whole length of the fibre. And
ili.'U' ii H ijivat dt-al of evidence, into which we cannot enter here, which
i^oix li> I'li'Vt' lliitl in alt striated muscle, mammalian muscle included, the
iiiMv U< iiitwtHiiiv \» nmlly composed of longitudinally placed natural ^ArtVAs
.'I n iviiaiii iiHlum, imtVddeil in an interfibrillar suDstance of a ditferait
ii.tititv III iiiiiiuniHlian nuisoloand vertebrate muscle generally these fibril lie
iiu iMiviiingly tiiiii and in most cases are not sharply defined by optical
ilniitu'tKia iVotii ihi'ir intertibrillar bed; in insect muscles and some other
iiiiiti \vm ihcv am rt-lalivt'ly large, well defined, and conspicuous. The artifi-
k'Kil iiliitll(*< uliliiini'il bv iciwiiig may perhaps in some cases where they are
I \i>'i'iliii>tly iliin I'lun'HiMind to these natural fibrillse, but in the majority of
uiii'H ihov t'ci'tidnly di> not.
Ill it'i'liiiii iiiaiH-l iiinMcUi< each bright band has in it two (or sometimes
iiiuiio iliiik liiiM which are granular in appearance and may be resolved by
.idi'>|iiiiiii iiiiignitViiig (HiwiT into rows of granules. Since they may bv focQs-
ui;' lii> liiii'i'd lliviuigh tl)e whole thickness of the fibre the lines are the
I t)iii'si)iiiii i>r liiiH'H. l''rfi|U('i)tly the tinea in the bright bands are bo conepic-
ilkU' 111 li' riinii'iliuti< a givaler share to the transverse striation of the fiore
lliiiii iln lilt' dim IiiiiiiIh. Similar granular lines (rows or rather discs of
Kiiuiuli>.->i uiiiy uiiio bo Hccii, though less distinctly, in vertebrate, including
iiiiiiiiiiiidiiiii, iiiiiHi'li'.
Il<niili<ii iliiviK griiniitnr liiici> whose position in the bright band is near to
I ill I Inn liaiiili, nHcn tipjicnriiig to form, as it were, the upper edge of the dim
l-iiiiil liiliiH iiiid tlm lowiT cilgc of the dim band above, there may be also
n..iii,ununlrHfrd aunt her Iransverae thin line in the very middle of the bright
luiiid. 'I'hix lint', like Ihit other lines (or bands), is the expression of a disc
and liiiB liiTii ludd by wmw observers to represent a membrane stretched
iiii.i-w lim nliiili> ihickncHii of the fibre and adherent at the circumference
Hilli llm BHiriilniiiiiii ; ill this sense it is spoken of as Krame'i nwinbraiie.
Till' iriiMuiiB I'm Inlioving that the line really represents a definite membrane
I
I
tM>t however ap(i«ar to he adeiiiiate.
metliiitc line."
Wli«n ■ itiiii trniuvftrM secUon of frnxcii muM'lc n cxaminetl quile IVo>U
Hod'T u tii^h pi>w(!r, tlin iini.-u-lf Kutuiniici- within the Huvoltoima ia wcu tp
ba niurlcil oul into n uutulxr '>f umiill motv or lew polygon nl Area*. AD d it
aitDilar nmtiif(<:nH-nt iuto nntu miiy al«i> bo Mwn in tmitsvunM.- xn^tion* of
prefwrad inuNcIc, though lh<- iVtiturrs of the nrcif orp somcnhat liillvrout
from thorn mod in thoi fmli living libre. Thcw areas are B|x>ki>a of M
" Cohnheim'fl nrfm* ;" they ano vcn- much larg«i: than the dinawtcr of a
RKrilla as iodicatMl bv t}i« ioniiituiUDal etrialion, and indoed corrupond to
a whole hundle of suoh tibrilla>. Their exwlence teeaa to iodicnte tnat the
ftbriUje are arrao^'ed in lort);itiiilinal prisiiia separated from each oliter hr a
larser nniount of iiiterhbrillar subitaiice than tbtU uniting to^'ether the tndi-
ridual libriliw formin}; each priam.
LiMtiy i( way be mentiun«(l that not only are the various Kranular linet
»1 times visible with diliiculiv or iiuite invbible. but that even the diAtino-
tinn belReeu dim and bright band» in uccAsii>iinily very faint or olwcure, the
whoh- muiii:i<^ »u balance. n|>art from the iiudei. apftearing almoet huiuoKetieouk,
Withuui attemi>tinx to diwuw the many and varioui inlerpretatioiia of ihu
■Imvv flixl ithtir lUtaila conoerninfc the minute vtmctiire of striated musctilar
(ibr(>. n<> may here oiHilvfit outsvlvM with the following general conclusion*:
(1,1 That ihv am»ch ituliKtiuicc i* t^ompiMed of longitudinally di«po«ed
Jibriltir I priilNttily (■ylindricat in general form an<l itrolMbly arrangcil in
h>OK>ludinn) pri'inii) rm)>r(td<)d in an inlrrfi/irlllar notiaaee, whi<'h ap|H-iir«
111 Fh- Inw ililWentiatfil thiin th^t lihriltic ihemaclvci and which i« probably
cfintinuoita with the undiircreii[>at«<t protoplmm round tin- nuclei. Thv
iiitcrtibriUnr fubrtaooe etains morv raidily with n^ld rhloridv than do the
fibriliir, and hrnce in gotd chloride epodmcM appear a* a rort of mcsliaork,
wilb hingitudinal i-paoet^ correHpondlng to th« fihrillffi.
(2i That the inierlibrillar Bubolancc w, relatively to tlie fihrilljB, more
abuntlaoi in the musclei of some animale than iu thow of others, heiug for
instaiioe very roiispicuoua in lli« nitiHcles of insects, in whicii animals we
■bnuld naturally expect the leas ditferentiaied material to be more plentiful
than til the muscles of the more highly developed mammal.
('■H That ihe Gbrillie and interSbrillar substance baviuK dilferent refcac*
live jHiwerv, some of the optical featurea of muscle may t>e due. on the one
haira til the relative proportion of tibHII» to incerfihrillar «ubsiance, and on
tlie other IuuhI to tine librillie not beint: cylindrical throujthout the length
nf tin' tilire hut contitHcted at intcnaU, and thus l>ecoming headed or
OKMiditortu ; for iiwtancifl the roiva of granules xi><>kuii ufabovi- arc by lunnc
rtpu-<lrd 111 carrM|H>»ilini[ to aggregation* of mtertibrillar maliTial lilliiig
lip the r|iar>-« where ihr fihrilhe arc nu«l coitttrinted. But it dow not Mem
pLMihln at the prTMenl time to make any ftalenivnt which nill mtbfoctorily
explain all lb« varioUN appoaraiicew met with.
t AB. We may now rettim to the •lutvlion. What happen* when a conlruc-
lioD wnv Nwiyip* over Ihe fibre?
MuMTular fibre* may be examined oven un<ler high power* of the raicro-
«CQp« while they are yet living and contractile: the oonlraclion tiself may
U (cen, biit the rate at which the wave travels u too rapid to [>crmit Katis-
faclury observations being made a§ to the minute changix' which accompany
cotilraclton. It fre<jueiitly happens however that when living muBCle
l>cen treated with certain reagents, as for instance with otimic aeid vupor,
•ubpe<]i>ently prepareil for examination, fibres are found in which a
ig, a tbickeniitit and shortetiing, over a greater or leee part of the length
i
THE COKTBACTILE TI890E3.
of the Abre, has been Bxed by ihe oemtc acM or olber iVAgeiit. Such b
bulginK ubvioiuly difl^ra ttom a normal ountrectioti in being conffoeil to a
pnrt ov Ihe lotifili of ilie fibre, v.')i«rea«, aa we ha\-(- nuiil, a normal wavo of
contrutioD, btiDg v«ry inii<^)i longer ibaii any fibre, •>ccur>iM t)i« uIkiIc
loi^t4i of tbe fibrfl at once. W« mar Iiuwrvcr n-)^nl lhi« bulgtiii; w a vi-n-
slwit. a very abbr«viiit<<(l wnv<- »f riiotriK-rion, unil nwuine lb»i th«' t'lian^
visiblo in sucb a short bulging iiU» tiiki.- jilitcv in n normal contmciioii.
Admitting ibis avuinpUon. wc learn fr»m siicb prepanttions tbal in the
cunlrnciing region of the fibre, while lioth dim nnd bright banda become
braaderncroea lh« fibr«, nnd correspondingly thinner nlong tlw Icnglh of tbe
fibre, B remarkable change takee place hetwe«n the dim bands, brignt haDda,
and granular lin««. We have seen ibat in Ihe fibre at rest the inter mediate
line in ibe bright baud t< in moM casea iucuDspicuoui ; in tlie contracting
fibre, on (h« contrary, a dark lin« in the middle of the bright band in tbe
ptieiiioD of the intermediate line becomee very distinct. As we ]mum along
tbe libre Irom (be beginning of tbe coiitrartion wave (o the summit of the
wave, where the thickening ia great4»t, thi>t line beooinftt more ami more
•tri king, until at the height of the contrai'tion it beoomes a rerv marked
dark line or thin itnrk band. Pari fintm with thid changr, th<r diMinctioo
between t)iv dim nn'l bri;;lil liiuiiln become leHs and Irm Jiiarked ; thne
aitjuMir to bcoi>mc ciinfii#c<l Ingiaher, until nt th<; hi^ight of the oontraclioR,
the whole space between esich tivo now cnnxpiciioti'' dark linw is occupied by
a •iib<4nnc« which can be called neither dim nor bright, but which m con-
triMt to the dark line apjHtars more or hw bright and transparent. So thai
in lll« contracting part there is, at the height of the contraction, n rev^ntal
of the state of things proper to the part at rest. The place occupietl bv Tlif
bright baud, in the state of rest, is now largely tilled by a conspicuoiiH dark
line which previously was represented by llw inconspicuous mlermediale
line, and the place occtipiiii by the eon^picuous dim band of the llbre at roM
now Mcm* bv ooinpnriMon with the dark line tbe brighter pan of iIk- fibre.
Th« contracting fibre ts, like tbe fibre at reit, striated, but its striaiion is dif>
Ibrent in icii natur« fr^xn the natural striaiion of tlte railing fibre ; and it is
hdd bv wHue that in the earlier phii»i=« of the contraction, white the old natu-
ral itruition is being replaced by the new uriation, there m a alagv in wbldi
all striaiion is lest.
We may add that the outline of the sarcolcm ma, which in the fibre at rest
is quite even, becomes during the contraction indented opposite the interme-
diate line, and bulges out in the interval l^tween flach two ioteroiedbita
lines, the bulging and indentation becoming more narked the gmUr tb*
con tract ion.
§ 96. We can learn something further about this remarkable cbangn by
examining the fibre under polariiHMl light.
Wli«ii urdinary light U aetil tliruu^h h XIcoI prism (which is a rhoinb i>r [ne-
laud «psr divided Inlo two in a cerlflin directloa, the faalrei being Mib»i>i)ueriily
oemeated U^thor Jn a special way) it undergoes a ebange la jMsaing through tbe
prism and is said to bo polarhrH. One e(^t of this |>oUriallon ia that a ray
of light which bus pn»ed through one Nicol prism will or will not pMS tfaroucb a
semoil Nicol Hcconllng to the ri-lallv«' jiusltion of thu (»<• iinKinn. Than, if the
second Nicol be so placed that what i« cnl1c<! ii« '' optic ai'is ' b« in a line with or
psrallel to the optic nxii' of the Hnt Nicol the light passing through the II rot Nlcul
will also (MUD through tbe Kccond. Etui if ihc second Nicol W roMl«d until its
<^e axb ia at right angles with tbe optic axis or the fint Nicol wiat of tii« light
]>aMlRglhruugbthefom>erwillpaMthrough the latter; iheprisuis in this posliioo
ar« ssld to be " crossed," In intermediate posilioDs more or Ic-m light paaiiea
ihriugb the sccoixt Nicol according to tbe nngle bsfweon the two optic nxm.
ileiHM) when one Nicol is placed beiwath the stage of a microMOpe mi tlmt tbe
k
U Irom th^ mirnif in t^iit tliroui'li it. hiiJ aiintlier Ntcol is plitoed in tli« «y«-
, r«. tlir tiriil of Ihi- inii;m(ico(i«i i*ill ap)i««r ilnrb «b«n th« ey^-pieee Niool ia
nitktfld »•> tliHi iu optic iixia i« kt Hghl Anglm to the i>|tlic ftxii of thr lower Xicol,
Mi'l coiM«<iuent1y tht? light pawing throusii (ho l<iw«r Nicol in •Ii>|i|)e<l by it. ir,
buwrvrr. tli« "plic axi" <if the t'_v<'-ii!ccc Nico] iH piirallfrl toikuturilie tower Nicol.
ilio liichl from thr Intirr will [.ftKi ihrouEh the hnaet am) the Held will be bright:
lUKlaa tbvcTv-piMie m srMluntty roiaief fVoiit oiiepmition tn theolkM tb« brlgbt<
ncM of ilie i!l«)d will tlunioioh nr increMe.
Botli ilie Niool« ar* «>RipoMHl of ihiubly rofrnctlve niuterinl. If now a ihlrd
iImiIi])- trfrHciive aial«ri«t lie pUcoil mi Ibe vtuge. aiitl tlierefore between ihe two
Nirol*. [he liicht pMai up through ihe loirer Nicol will (in a certain poaitiixi «f the
ilonbly refriLiivc mntenal on the xtaee, that is to aay. whcii itn optic *xm hava a
i-rnain r'niiion} paai ihningh it, ana alxi through the rrairej Nieol iu the eye-
(liece. ilencc tbe doubly refraclivr mntrrial on the Hlage (or -tioh [iflrtH of It aa
ar>' i" ff'i- irfnper poaition in rwjiew to iheir oplic axw) will, when the eye-piece
Ni -ed. nmiMf llliiiuinale'] ami bright on a ilnrk field. In thin way the
et I - I iluubl; ivfraeiire malorial in a prepiuatiiiii may be iletcclvd.
When tnu»cle [ireparecl nnd n>04int«<) En Catioiia bttlaum u exatained in
th« microecupt! Iwtweeii Nioo) jinanii'. oik uii tbe vUiK^ helow lh« •.ibjoul, and
tbe (itber iu the i'v(:-|>i«<x-, t])« Hbrm stand unl iw bnght objvctn on Ibi; <lHrk
DDtl of III*' li<^ld when llut axes of tbe |>riKtna nn; cnNwed. On cUmrf
linatioa it L« nvco that the parts whieh iin- )>n^lil urv chi«flj thv dim
m. Thia Indii-nlea that it is the dim iMiidit which arc dmibly n-frH«tivc,
iMtrfrD/fM-, or an- ehii-fiT made up of anisotropic subatnnce; tluire sc«rii».
eter, to be iu>nie slight nmouQt of nniaotropic subaUan in tbe bright
ada, thiiueb thot: lis a whole appear single refntelivo or itolropic. The
tibn iceonliDKly apjwars banded or rtrialod with alternate bnnds of nniao-
tnipic noi) iaotropie materia). According to most auihore such nn alterna-
tiiiD (if anisotropic and (chiefly) iaotropic bands which is obvious in a dead
■mi preiMred fibre exists also in ihe living fibre ; but some maintain thai the
iviOK filire in uniformly aubotr<ipie.
~~"ow, when a fibre raulracta,in »])iteol"the confunion preriAiMlj mentioned
reodini and bri^hi Iwodx, there i» no eouftiKiou between liie anisotnipio
nd iantrapic materia). The ani»otr»iiie. duubly refmetive bands, bright
under crm»ed Nicxtli, tMieuprin): tlie poititiou of the dim ban<ta in tbe resting
Rbrr, remain iloul))}' refract ive.liright tindiTr croiaed Nicu)s, ev«D at the very
of the eontmclioii. Th«isotr<i|>ic. Kiiigly refractive )>andi'. darlc under
' Nicolii, iKiciipyiiig thft pavition of the l>riu;hl bandi> in the fibre at rest,
iaotropic and dark uiiderenj*«i^l NicoU ni tlie very height of the
txHUradlon. A)l tliiit cnn )>vMen is that the singly refruciiveisotro)tic btinda
bevome ver^- thin indeed during Ihe cu lit mot ion, while the nni»oln)pic tiandM,
tbough of contw becoming thinner mid bronder in the coutriKriion, ilo not
mc- so thin U do the i<<>tropic bands : in other vrontf, while liolh )>ikuiU
me ihinDvr and broader, tne donbty refractive iini>^olr(ipic band si,-«nis
norcMse at tbe expense of the singly refractive isi^ronic liitnd.
$ 07. We ea)l attention to these tads because they »how him- complex ifl
the a<-t «f contraction. The mere broadening and shortening of i-aeh seetton
of ihv fibre is at bollom, a translocation of the molecules of the muscle siih<
tUQon. If we imagine a company of 100 soldiers ten nuiks deU|>, with ten
mHi b Meli rank, rapidly, but by a series of grnilutions. to ext«n(l out into
a diinble line with oO men in each line, we ^hall have a rouffh image of tho
iBDV^mcnl iif the nmleeulea during a muscular coiitrvdion. But, from what
Ilia tiM-n Milil. it is obvioun Ihni the movement, in striateil mnsele at least, is
a very nimplicntMt on<> ; in uilii^r forms of contractile tiMiie it may be, as we
■ball «<•, more "imple. Why the movement ii so complicated in striated
nu*clv, wbal purpumi it eerven, why the nkek-ul musetn* are titrialed, we do
toll:
THB OONTBACTITiB
ont *t |»mcni kimw. A|i|Mir«Dtlv wlu^rc Nwift and nii>id oontnifiiitii it
requintl iho coiitraciili- iiMtio in Mnutol inuMtk ; but how Ute Mriatioii hvlim.
•0 to )t|K9ik, th<: ciiiil tiirltiiu vri- <)<> not kniiw. We l'hiiikiL nay vrhnt Khiiro in
lh« net of coDtrnctivn i.* t» bo ttlliaCol U> lh« ocvrrul juarta. .Sincr. during
K cuuUmi^ion, the fibro hkilgaw out awrv 0|>|x)»it« to cficli dim di>c. nnd
\b indented opposite to c«ck bright disc, «inc« tliv ilini di>c i« mora largely
(vxDpoced of anisotropic mau-rial than the rwt of tbo fibre, and siooe Uk
anisotropic nutl«rtnl in the |x>sition of the dim disc iiicmiMa' <liiring a cod-
triK-iiou. nc might porhajw infer tUiit the dim disc mthor than tlic brij^bt
disc is lh« eneDtially active rmrt. Awiimini^ thiit lh« fibrillar substaDcc is
more abundaut in tlie dim discs, while the micrlihrillar substance ta more
abundant iu th« bright discs, and that the fibrillar substance is anisotropic
(and hence ilie dim discs larKcIv nuUotrupic), while th« inicilibritlar sub-
stance is isotropio, ne mifht also be inclined to iufer it is the 6brillar and
nol (be iuterfibrillar subetauoe which reallir carries out the coutractiou ; but
even tliia much iu ni>t yet definitely jiroved.
One tliiu^ mujit !>« remembered. The muscle substance, though it p<j»-
»vsii-i the ouiujiliculed ttnicture, and t^M through the remarkabUr channi
whirh We have <lt-»cnb«l, \a while it h uviag and intact in a condition which
wc are dnvc.ii to t\teak of ux nemi-Htiid. Tlie whole of itU ewcnliiilly miJtlle.
T)u> very act of coutmi-liiui indin-d nhowsthb; but it in nmttik in the sciisic that
no part of it, except of counw the nuclei ami itarccilcmma, neither dim nor
l>rif(ht futwtanee, neither fibrillar nor iut«rfibrillur nulMiaiicv. «ui tw regarded
a* a hard nnd fimt structure. A minute ueniatoid wonn has been scon wan-
<lvnng in the nmlst of tlie substance of n living contrnctiletibrc: wi it moved
along, tlvo muscle stibelanec gave way iH-fore it, ami closed upii«ain bebiud it,
dim band)' and bright bauds all falling back into their proper pMuva. We may
suppoM that in this case the norm Itircnded its way in a lluid intertibrillar
suoshioce between and anmiig highly extensible and clastic 6bri)l(e. But
even on auch a view, and .ilill more on liie view that the tibrillar aub^tance
also U'u broken and clooecl u» agwn, (he maintenance ot such definite hii<to-
Iflgical fbatures oa (hoee whicn we have deeeribed in malarial ao mobile can
Mily be eflibcted, even in the Hbre at reel, at some conudenible exiwodttitre
ofener^: which energy it may be expected hasa cJieniical muroe. During
the contraction tliciv ia a Mill ftirtber expenditure of ejier(,'y, mm*> of which,
as we have seen, may l«avo the muacle aa *' work dune ;" thh energy-, lik^
wise, may Im* cxpeclc<l to have a chemical aouroe. We miisi, tbcivfore. now
luni to the cliumistry of miiade.
T/if Cftmintry 0/ Mwrft.
$ 98. We said in the Introduction that it was difficult to make out with
certainty the exact chemical ditferencee between dead and living nub-
ittance. Muscle, however, in dying undergoes a remarkable ebemical change,
uliii-h may be studied with comiiaratire ease. We have alrvatly niA that
all miucl», within a certain time aAer removal from th« body, or. if still
n'nniining jiarl of tlie boily, within a certain time after "geocral" death of
lh«' Wly, l«*e their irrituhiliiy, and that the lose of irritability, which, even
when rapid, t« gradual, in {•uccedlcl by an event which io Mmcwhat more
sodden, vir_, ihv entrance into tho condition known as riyor inorlU. The
oocurreiice of rigor mortis, or cadaveric rigidity, lu it is sometimes caiile<d,
which may be ci>i)«idercd im the token of tbo death of the muscle, is marked
by the following features: The living muscle pooaassas a certain Iranslucency.
the rigid muscle n distinctly more opaque. The living muscle is very exten-
sible and elastic, it siretchee readily and to a oooaiaerabto esleni when a
VBTeDsUnT The entrance into rigor nortb »,
' ft dtoneniug or coiitractiuu, which inaj, un<ter oe:
liderable. The energv of thU (vintractinu In t»n ^rent, fu> tlint any
iboricaingiseui)y|)rt!veui«(l by the |irt!Miiii.-vi>l'eTeiiui'li^ht ^pjiwing
the cWuiral fviiluKu of th« t\md r^id inuwlc art idso Mrikiiigly dif'
rotii ihm^ «>' thn living tuiiiKitv.
^n lifid miijWi-, fnim which itll flit, uihIod. fnsciH, and oonoectivo
He btruiu much lu (xi^ihio n^niovm). <>i)d which hiu bMfi Awod from
^ho iiijrctiuii of " normnl " siiline solution, be ini»M-<l nnd n-])aitcdlT
with wntcr, the wnshin^ will coiiuiD corlaiii furnis of nUiiiiiiiii nnd
«xlniclit'« IxMJies.of which ««■ sbsll apoalt directly, Wtieii ihe wiish-
■ been continue'l uniil the wesb>waier Kires no proteid reaction, a
IWtion of muscle will Mill remain nudiaeolved. If (IiiB be trenled with
roenl. Aoluiion uf a neutral saH, aminonium chlonde betiiK 'be heal,
portion of it will Itecome dissolved ; llie mduiion. liowever. la iDOre or
perfect and 5Iten with dilficulty. If the filtrate be alloired to fall
y dn>]> into a Isr^rc ijuauiitv t>f ilintillnl witter, a while flucculeiit
will be {irecipilat*^!. Tlii.i ifoivuleni prvcifiitale ia myiMiii. MyoaJn
KrH), (living the oritinnry jioiU'id ri-itctionn. un<l haviii^ the turne geU'
liunttnry <'>itN|<naili(in mi other pr<>tci<li>. Il in ixilulilio in dilute MiliDft
m, aqHioiallv IbixK of ammouiuni chloride, and niav' Im i:1bm>c-<1 in (he
I fiiinily, cltmigh i( is not so soluble as {laraglouulin, requiring a
r mtuUitii of a neutral •all to diiaolve it ; (htis. while soluble in a A
nr n>tH. soluti^m of such a sail, it is far leas soluble in a 1 per mnt.
I, which, ns we have seen, readily dissolves parsglobulin. From its
IS in ueuintl saline solution it is precipitated by saturatioD with n
snlt, jtreferably sodium chloride, and may be purified by being
with a saturated solution, dt»olved a^cain in a weaker solution, and
jitaied by saturBtion. Dinotved in saline solutions it readily cua^;-
wtteti heate<t — i.e,, U couvert^sl into eoajjulaled proteid — and it is
■if tiotiee (bat it coafculute? ai a comparatively low temperature, vi^.,
6° C. ; thiM it will Im remembered is ihe temperature at which Hbrin-
congulnletl, wliere** pwraglnbulin, senim-albumin. and many otber
I dii not (VM^iliite until a higher tcm|H-rHture, 7'>° C, i^ reached,
aiif inyiMin are |ire(!i|iilat4.-<l by alcotiol, and the tireclpitate, an In the
|ker ^rolvidi. bocooMi. by continuv«l action of the alcohol, altered
88 THE COXTBACTILE TISSUES.
state. And what U true of the collection of parallel Bbres which we call the
muscle is aW true of each fibre, for the Bwelliug at any part of the muscle a
only the sum of the swelling of the individual fibres; and if we were able to
take n single long fibre and stimulate it at one end, we should be able,
under the microscope, to see u swelling or bulging accompanied by a corre-
sponding shortening, i'. e., to see a contraction, sweep along the fibre from end
to end.
If, in the graphic record of the two levers just mentioned, we count the
number of vibnilions of the tuning-fork which intervene between the mark
on the record which indicates the l>eginnin^ of the rise of the near lever
(that is, the arrival of the contraction waveat tbU lever) and the mark which
indicates the liej^inning of the rise of the far lever, this will give us the time
which it has taken (he eontniclion wave to travel from the near to the Air
lever. Let us sup|>0Ke this to l)e 0.1)05 second. Let us suppose the distance
between the two levers to be lo nmi. The contraction wave, then, has taken
0.005 second to travel 15 mm., that is to iray it has travelled at the rate of 3
metres per second. And indeed we find by this, or by other methods, that in the
frog's mnscles the contraction wave does travel at a rate which may be put
down as front 3 to 4 metres a second, though it varies under diflerent condi-
tions. In the warm-blooded mammal the rate is somewhat greater, and may
probably Ik put down at 5 metres u second in the excised muscle, rieing po«-
eibly to 10 metres in n muscle within the Jiving body.
If, acain, in the graphic record of the two levers we count, iu the case of
either lever, the number of vibrations of the tuning-fork which interrene
between the murk where the lever itegiiis to ri:^ and the mark where it has
finisheil its fall and returned to the Itase-line, wc can measure the time inter-
vening between the contraction wave reaching the lever and leaving the lever
on its way onward, that is to say wccan meoiiure thctime which it has taken
the contraction wave to \ituv over the part of the muscle on which the lever
is resting. Let its suppoi^e this time to be, say, <>.l second. But a wave which
is travelling at the rate of 3 metres n second and takes 0.1 second to pass
over any jmint niui^t be 300 mm. long. And, indeed, we find that iu the
frog the length id' the coutraction wave may be put down as varj'ing from
200 to AW mm., and in the miiminal it is not verv different.
Now, as we have sai(i, the very longesl muscular fibre is stated to be at
moat oidy about 40 mm. in length ; hence, in an ordinary contraction, during
the greater jiart of the duration of the contraction the whole length of the
fibre will be occupied by the contniction wave. Just at the bwinmng of the
contraction there will be a time when tJic front of the contraction wave has
reached for instance only half way down the fibre (supposing the stimulus to
be applied, as iu the case we hiive been discussing, at one end only), and just
at the end of the contraction there will be a lime, for instance, when the con-
traction has left the half of the fil)re next to the stimulus, but has not yet
cleared away from the other half. Hut nearly all the rest of the time every
part of the tibre will be in some phniie or other of contraction, though the
(larti^ nearer the slinmlus will be in more ailvauced ])bases than the parts
iirtbcr from the stimulus.
This is true when a mu^le of parallel fibres is stimulated artificially at
one end of the muscles, and when, tbei'efore. each fibre is stimulated at one
end. It is, of course, all the more true when u muscle of ordinary con-
struction is stinuilated by means of its nerve. The stimulus of the nervous
impulse impinges, iu this case, on the muscle fibre at the end-plate which, as
we have said, is placed toward ijie middle of the fibre, and the contraction
wave travels from the endplate iu <>p]i<>.-iite directions toward each end, nod
has accordingly only about half the length of ibe fibre to run in. All the
CRA!?OBS I^T JL MCSC1.E DL'RIXO CONTRACTION',
99
I
tnuK. liierefore, must the wliole fibre b« in a state ot' ooutravlion at lh« MUBi
lini*.
It will be observed (liat in wUat has ju«t b««u aaitl i)i« raairacti^ii wave
hiu liOFn takeu to include not uul^ ilu- cmntriictuin propvr. ilia,' thickening
Mill K boric niiifc, but aliMi Ihe lelBxiitiou and rvturn (» llii; iiiiliinil form ; ibi-
lin>L |Mrt >•( ihe \iu\-t: up to the Kuminit of ibi- crast oorratpuids to tbe
■tinrtcniii^ nml thirk<'niiig, ih*' di-clinc! from the «iimmit onwara crimepontb
lu tba n-luxBiioti. Biii w« Have aln-wlv iiuiiitcd tliiii tliv relaxation ig ao
tial [wrt ••(' ihr whole act! imlMil, tn a lortaiii wriiM<, n» fwrntial as t\w
iMi-iiini: iiM'lt*.
_ 5 44, Miii'itr linteluri:'. n) miueulnr fibn. S«> far we have 1»ecii dcalin;;
with the iniK'C'U' a* it whole and m obvcneil with the oaked eye, though ne
have iueidrjttally s{ii>kvN of fibres. We havn now, ronliuing tiur atlcntion
cxoliir^ively to skeletal aiusctcs, 1o ocwitider what iDicrwonpic changes take
[ilatsr durins a eontraction, what are the rehUioHs of the hiatologiciU feature)
tif (be DtuBcle fibre to tlie act of coulraL'tiuii.
The luuK cylindrical abealh ofuircoleuiiua is occupied hy muscle subtitaooe.
After death the muscle substance may separate from the sarcoleinma, leaving
lh« latter ea a distinct nheath, but durioK life the muscle siilwtanoe is adherent
to tlie san-okniHia, so that uo Hoe i>f sejiaration between the two can be
maila out ; liie inovementa of tlif oue follow exaetly all the laovenveuta of tbo
oii»cr, ,
SaUtend in the mmcle sulioiaiu'e, but, in the mammal, lying for the iikmI
imK oloao under tiw rtmmleuiina. are a uuiul>er of nuclei, oval in «ha|w, nilh
their long axo* paralh-l to tbu K-iiiith of th<; lihro. Around each nucli?ti» \*
a tliin layi'r of grauular-lookitig nubolanci' very Mniilnr In ap|>earancfi to that
fonunc the body of ii whitr blood'C»r|HiMrb', and like iJiiil ol^en njiokon of
as aaduTerantiatiid pr»topliu>m. A small •juniility of (ho Mtnv granular itub-
BtatMv is |in>lo»gi>il (or i«>me dinlanoc, iik a narrow conical streitk from uach
end of thu nucliip^ along tbo k-iigtli of the fibrv.
With tha exir|iU<M) of cboM nuclei with their granular looking bed and
thi? end -plate «r eiid-|ilal<?s, tolie pmentlv dtecribM, all lite xtM of tliv s(Hi«e
eni'loaod liy the surcolomma from one eoti of tbo Bbri' to the other apjwara to
tie ouuupit^l by a peculiar material, 'Irintfl niiur/i' riiittiuire.
It i» ittlli'd rtrialed boauM it is marke<louI. and that along the whole
IsAgUi uf iti« (ihrv. br tniiuveree bandft [Fig. 'i'i]. stretching right acrwe the
ir»a K.
s
14
l>i*>ia«NH«nc tanwaanxivm or i llunaoia.
iw, iwew-MWM. wa<*U^of « bUBdloctfiuu«cle-(«dsi ft, iWd «utala»M.|
lihrt. of »ulKlaiicei which is very iruiLipnrent, bri-jhl mAttntiCf, alternating
with tiniilar Imitds of >til>staii<« which has a dim cloudy appearance, -iim
mbMunfr ; that ia ta uy the tihrt! is marked out along it^ whole leugtli by
allamato bright band* aud ''na IxihiU. The bright bands are on an average
about 1 ^ or U> n aod the dim baniU about 2.5 /< or 3 ^ ibick. Bv care-
ful focuasing, both brigtit hand* aixl dim tMUfJs may be traced tlirough
t}i« whole tcickiMMt of the fibre, w that the whole fibre ap])ean to be com-
piaod uf bright disca lutd dim diacs j>lnocd alleni«t«ly one u|>ou the other
ili>ng tin* nholc length of tXw f)l>re. th<' arrmngeBMDt being broken hy the
cnd-jilBM awl here aw) there by the uueWL
TUB COSTBACTltE TlgSVSS.
Whcii n mtifctilnr fil>r« is tmi(vd with dilute mioeral itcids it » vtrr ipt
to hrvnk up tmiisvftrwly iiito diwe [Fi^. S'i], ihr mrcolemmii beio); dinolred,
or ao nltvnil lut ciuily to dindo into fragmrnta oormpoDding to the diMs;
•imI ft disc ttMy thus be obuincd so ihiD as to comprise only a Bingle dim or
bright band, or a dim baod iritfa a thin lnyvr of bright subsiaDce above and
bolow it, the clearnce having taken place alon^ the middle of the bright
bands.
Wbei) Imted wicii oertain reagents, alcohol, chromic acid, elo., the fihre
is rerv apt to aplii up (and the splitting up may be aasisted by " teiialiic")
loiigitudiiially into ouIucuqs of variable thickneas, some of which iioivcver
may be escee<liii^Iy thin, and are then sumetitnea spoken of m " tibrillir."
Uolh ihe«i.' di.-H-s luid fitiriltiu arc artilicia) prrtducts, the resultn of a trntiit-
verse w luDj^'itiidinBl ck-ava^ of the dead, hanlcnr*!. or olhenriiie pn'mr^l
Uiirt'lir :-ut»lunc<'. Tli<-t may Ri<irv<)V«r W obtained in nlmuU any ihit'KnciM
thinnciM, and thivt' ai*>» and tibrillie do not by lhriniwlv«« provi' iiiacb
l>eyond th« fact that the fibre tendit to cleave in tbi- two directJoDS.
The living fibre himcvcr, Uioiigfa at limei i)uilc glantr looking, thv bright
bands appearing like transparent giais and iIh; dim band* tikr gnxtnd gtaw.
!• at other titne» marked with loiigititdina! Iin«s giving rise to a longitudinal
siriation, sometimes conspicuous and oc<-asioRftlIy obscuring the lmn*v«rai
StriatioD. In the musclm of some insects each dim band hns a dislinc-l |»li*
siulc appearance as if made up of a number uf "tibrillie" or" rods" plaoed
side bv side and imbedded in some material of a diflerenl nature ; moreover
tbeMubrilln or roda may, with greater dilhculty. W iraoed tbrongh the
britcht bands, and tJiat at tiroee luoug the whole length of the tihre. And
there ia a great ileal of evidence, into which we cannot enter here, which
goes to prove that ill all striated muscle, mammalian muscle incttidird, ilie
muscli' nuliMtancc is rally eomnoaed of longitudinally tilaoed natural ^'^riVte
of a c«innin nature, inifieddi-d in nn iitlerfil/riKtir suMiance of a ditTereat
nature. In maiuiualmn muHcb- and verU'bnile uiusi:le generally theao Bbrillm
are exceedingly thin and in most case* are not sliarply deBi'ted by nptical
cbaraclera from their intcrfihritlar bed; in insect muKlm and aoiiie nibfr
musclm they arc reliitiwiy large, well ilefined. and coiw])ieiiiiu*. The arlirt-
oial librilla) obtjiincd by tciu^ing may [icrhaiM in some canv* when^ thi.^' an
exoredingly thin correspond to tbcso natural fibrillie, but in llw roiyontr uT
cwM they certainly do not.
In ceiiain insect muscles each bright band has in it two (onoioetimes
more) dark lines which are granular in appearance and may Iw motrcd by
iu)e<|uale inxguifying power into rows of ffranuln. 8iuoe Ihev may hv focus-
sing be traced throusb the whole thickness of the Hbro tde lines arc the
expression of discs. Freijuently the lines iu the bright bande are so ooiisuio-
uous as li> contribute a greater share to the iranerene strialion of the libre
than do thtt dim bands. Similar gniniilnr lines (rows or rather disosof
granules < mar also be seen, though less distinctly, in vertebrate, including
mammalian, muscle.
Besides Ihcac granular lines whose position in ibe bri^dit Ivand is nt-ar to
tlic dim bands, ollcn aiipenrin^ to form, es it were, the upper ed^re of tbt- dim
hand Mavi and the lowir eiigi' ••>' the dim band aliovi-, then- mav if al»»
sometimes traced nmilhiT iraii>ivtir«i^ tbin lin« in tbt? very middle iif lli<' bright
Iwnd, This liiiv. like the ulhtir lines /or luuidii), is the expn-iuion of ii ilint*
and has hvea bclil by Mwne obsvrvcm tii rcj>rei*nt a meiuhrane stn-lflied
arroas the whole thickiieM of the Kbrc and a<lhcrent at the cin'timftu'viice
with the snrcolemran : in this sente it [» spoken of n« Kt'ixuui* Hinnbroiif.
The reasons for believing that the lino milly re|»re«ents a <lelin{te mcmbnim*
»
I
ntit liov«v«r appear lo h*! Mk-quaW. Il may Ik >poke» of as tliv " inUr-
Bi«diat« Vhk"
Wlicii a thin traiwveno *cctii>n of froEcii niu»clu w I'xnniineil quiu- fradi
nnilrf a U'tgh |H>wcr, the niii»cli! KiiWtunott within tboMroolcmnin Im M-L*n Co
lie iiiiirkrtl out intii n niimbiT ot'«nuil] more or lem poljgODal an--iM, nod n
flioiilnr nrrniii^cnK'nt into antiu RtHy also be seen in transverae »vctii>n* of
prr]>art.><l niu«cle. though lh« foutiim of the arvw are somcnhat <liK'erenl
from thiMe Men in t\w fresh living tibre. Tbeec areas are tjiDken of lu
" Gnhaheim'e areas ;" thciy are very much larger than the dinmcti;r of a
tihrilla as iuilicateil hv the lon^'ituilinal rtrialioo, and iuile«(i corresuuiK) to
■ ii'htile bumlle of sucn tihrilln^. TiK-ir existence seeius to indicate itiBt llie
tilirilhu are nrranKed in hmKitiidinal prisms sep«raled from each otlier by a
lander nmouoi »f inleTtlbrillar fmbdtauce tiiaii thai uuttiajf to|{ether the iudi-
vtdual Hbrillie furmiu^ea<th prism.
I.ji»i)y il may bi- meiitiouni thai not nnl^ are the various k'^i'iIa' liaet
M tiiii<>« VMibto with ditRculty or quite invistlile, but that eveu tlie di«tii)c-
Uou t>et«i4^i dim and brij^lit bands is M-caiiioiially very faint i>r obscure, tlie
wIkiIi' niuwOr stihstaDi*. ajxtrt from the nuclei, a)i[M-arin^ almnnt homogen<Niiis,
Wiiboot iitutiiiniiiiif to disciHN the manvand varimi* iuterpnetations of the
al>ov« niul other it<-liiiu concerning the minute structure iif stHat«d muMular
fibre, «(« may ht're oHiti-nt nunxdvtii with the fDltowing j^ueral coiicluaioDs:
(1; That thi: miwle ftiilwtaiici: is iiun|>o«H>d of bngituiliiially i)itipos«d
JihrHlr iprifliahly rylli>.|ricHl in ptncml form and iiM)liably arranged in
aeiliidimil pri>uiii im)ic^dcd in an iitirrfiiiriUny fiiMannt, which appears
^M Ian dintrMitialol tlmn the fihrillig thaiast^vfa and which is jirobnMy
' loous with the unditfcreiitiatci protopleaia round Ihv nuclei. The
ilertilirillar sub*tnnce stains more readily with gold chloride than il<> the
fibrillK, and hence in gold chloride tpcdmcua appear as ii sort of Ricshwork,
vilb InnKittidinal epace^i corresponding to the fibrillin.
<'i) That the iniertibrillar subsianoe is, relaliTely to the 6brill», more
abuiMJant in the muj<il«E of some animab than in those of others, being tur
instance very conspicuous in the muscles of iiuects, in which animaU we
shiiulil iiniurally expect the less dillerentiated material to be more plentiful
tlian in the muwlesof the more hixhly developed mammal.
(.1; That Ihe fibrillH.- and iuterAbrillar subsiance having Oift^rent refrac-
the fxivrrre. some of the oiitical featura of niunclc muy he due, on (he one
hand t'l thv relative pr«>)nrtion of fibriltw lo iiiierrilirilliir Mibslance, and on
th<- uthiT haiiil li> thtt fibrillin not \ieinic cylindricitl tliroiighout the length
tlip iibri- but cuualrictvl at intervals, uiiil thus becoming lieaded or
lilif'irm ; fur liMiUice ihfi TDwa of granulni npoki^u of abitve arc by somv
_ inliil lui comspondine to a^rtgations of inlnrtilirilUr matiirial Htlitig
ip ihi- *|inrre where the 6l>rilU' arc moat coiMtrictnd. But it dow not seem
imsiblf Hi ihf) prciteRt time to make any «iatcnK-Qt which will salisJaclorily
•Xplaiii all the various appciamiior;' mot wich-
t fiS. Wa mar now r^'turn to the ijuotion. What hapjwns when a contrac-
tlno wave "wi-cps over ihe fibre?
MiiwTular fibres may he examined eT«n under high powers of th« micro-
Kope while they arc yet living and contractile : the contraction ilself may
l<e *vrt>, hut lh« rate at which the wave travels is too rapid to permit satis-
bctory observations beiii^ made as to the minute changm which accompany
crintrDdion. It freipienlly happens however that when living muscle
Itenn trealMl with t?erlain reagents, as for instance with osmic acid vanor,
*ub*«^iKtitty prtpiired for examiiuUion, fibres are found in which a
tog, a thickening: and sbortening, over a greeter or leas part of the length
THE eOXTRACTlLB TtSSCES,
of tho librc. hns )>c«n lixcd by the o«niu< ncjil or other ri;agent. Such a
bul|,nDc obviously i)itl«re f^om n normal contraclion in beiof; coofitie^l to a
part of the l«apth of the tibiv, whercne, at we have eaid. a Doroial wave of
contraction, bein^ very much longer than any fibre, occupi« llie whole
leofflii of the fibre at onoe. We may however repird this bul^ns bs b rery
■hon. a ven' abbreviate<t wave of contraction, and aaaume tlial the dion^
rUible in micb a ithort bulging also take place in a nnrmal oontraclitX).
Admitting thin ostium ptii^n, we learn fVom such preimnitions Umt in the
ooDtraeting region of the fibre, while Iioili dim and oright bamU Wome
brooder acroM the fibre, and corrcNimndingly thiuncr oloiig (Itc l^n^th of (he
fibre, n remarkable change lakce pinou between thr diiii buiuls, brigln bnnd«,
and granular Ithi'H. We have wen that in (hf; fibre at rest Ihe intiTinciiiatc
line in the briglil band is in moat caM« iiiconiipiciioii* ; in the ■■i>n[racling
fibre, on the contrnry, a dark lino in the middle of thr bright band in the
porition of the intermediate lino becomes very distinct. \t m pass along
the fibre from the beginning of the cmilniction nave to the summit of the
nave, where the thickening ie grcuto^t, this line becomes more and ntore
tiriking, until at the lieignt of the conlraction it becomes a very marked
dark Ime or thin dark band, ftiri f/'ittii with this change, the distiuctinn
between the dim and brijfhl bauds become leaa and leasjnarked; these
appear to become confused together, until at the height of the contradkni,
the whole apace between eacli tn'o now conspicuuiiA dark line) is occupied by
a subiCance which can be called neither dim nor briglil, but which in oon-
trnsl to the dark line apiMwr^ more or leM bright luid irausparcnt, So that
in the contracting jutrt there in. ut the heighl of the contraction, a rorereal
of the atate of thing* proper to the pari at re«t. The place occupied by the
bright band, in the state "f rwt. im now largely filled by a roiigpicuoua dark
line which previously was represented by the inconspicuous intermediate
line, and the place occupied by the conspicuous dim band of the fibre at leat
DOW sectnB by oompariaon with the dark line the brighter part of the fibre.
The contracting fibre is, like the fiiire at rest, striated, but iia striatitin i> dif-
ferent in its nature fmrn the niilunil ^trialion of the resting fibr«; and tl la
bdd by sofoe that in the earlier phiues of the coutraction, while the old uiilu-
ral utriatiea is liciiig replaced by ibe new slriatiun, there U a Mage in which
all Ml rial ion Is lost.
We may add thnl the outline of the sarcolemma, which in the fibre at rest
is (piitc even, bcconicj' during the contraction indented oppocito the inlerme-
diate line, and bulges out in the interval hctweon each two iDtcrme«Iiail«
linca, Iho bulging and indentation becoming mora marked the greater the
con traction,
588. We can learn something further about this remarkable change by
examining the fibre under polarized light,
When ordinary light la sent through a Nicol prism (nbieh Is a rhomb of toe-
land spardiviiied into two in a cortam dircciinii. the halve* being Mbsequetilly
cemcntrd tngctbrr in a special wav) it uiirl<incucH a change in juMJiig through the
prism and is mid to be polarited. One rflecl of this iiolarimtiea ia that a ray
of liglii whii-li lias piMcd through one Xii-ul prism will or will nut pass lhrotu[h a
•econd Nicol according to the relHtive positcui of tlie two prisms. Thus, If the
second Nicnl be so plntrd thnt whnl is cnllcrl it* '' nptlc axis be in a line wllh or
(Mirallcl to the optir axis of the lir«t Nicol the lixlit pns*ing through the first Nicol
aill also poM through the necoiid. But if the second Nicol be rotntod until its
optic axis iaat right aagtes with the optic axis of the Anit Nicol uoueuf the lioht
passing through the former will poM through the latter ; the prisms lu this position
are said In be '' crossed.'' In iiilermeHiaie positions more or lest light passes
tlir»u)[h the M-cond Nicol according ti the nngli; betn-ccn the two optic axiv.
Hence when one Nicol is placed beneath the stage of a mtcnMcope so thai the
CIIANOES IN A HVSCtE DCRiyo COKTRACTION.
r
■ Ucbt tan ika mirrvr i* wat Uirough it. anil anoilier NIro) it placed in llie ne-
' ptfc*. lb* fifid nf rtio aiictoMopc will appear ilHtk •rhrn ibo cyo-ptccr Ntcol b
n>tAi«d an that luoplir axii in al right aiifira to ihi- ofilic axborUic lower Ninil.
awl totMMUMiilr ibc light pmmiag ihrouih ih« tower Niool ia aupped br it. If,
bfvrrrr. l&c optii- nxU of ibe eye- picc« Niool i*]>aralld loiliatuf tlielowt-r NIcol,
ihr tiybl from the latlrr will pMa ibroneh lb« former aixl the Mt\ will lie bright;
iMi aa Uw pr«-t>icc« i« emduall^ roiaieif from on* pMilion tn tbe other ibe brinkt-
nmm of lb* MM will dlminidi «r iiit'r«M4>.
Baib tbi Nicola are ci>iu|mm*() of doubljr refractive oiaiciial. If now n third
dacblj r*tr»etiv* mnl«rlnl ba |>laced on the au^, and tlierefurv betwocn iho twn
INkak. I' |-a«un|c thrrxigh lb«lower Nicol will (inactftain piwilioti of lh«
dotibh - tuatcrial on the stave, tbat i* to aay. when iu optic dim bare a
f^ftaJn ^--.I'fn I I'aax thr<>ii|;h il, and alao throogb the trot*td Nicol in tb« eye-
fiwnt. llrnirt ilii- d'Mildy refractive material on the alage (or nucb paria of 1( aa
»n Id tbp proprr poaition In rtapeci to their optic aita) will, when the efe-iriae«
yteal U riMMd, Bp|war illuminated and bright on a dark Sc\<l. In thin way the
Rafa(as<« of donblj rtfraciiva niatarial Ui n preparation may be ili-tect«d.
Wbra tDuacle prvpurod «ihI nouottd in OiiiwIk balsam b examined iii
tb« micnwcope bM«eeo Nicnl prism*, on« ou the tufe b«loM' tb« ohjw!!, Aud
Ibr iTtbvr In tbo «y«-pJKW, the &bm atand out na bright ohJeciA on the tliu-lc
(rauntl of thr ti«M wh«D ibe *xe» of ihv jinmiM are crwaed. Ou cl»er
■tion il ia Mm that iIk- |iAria which iirc britcht nr« chJi-fly th« dim
Tbia indieaiM Uint il is tW dim bnii<)« which iir«- doiibty rrrrnctivc,
vtuptnpif, or an cbjcfly tn&de up of itniMHrvpic subaUuKC; thore sceuw,
•r, to be toam alight nniouni of aniaotropic mibtlsnn In the bt^ht
. tbnuffh these aa ■ wltole n{>|)«ar fingk refmclive or itotropU. Th*
Sbn tcnritoglr appcan banded or <tnatc<l with allrmale bunds of anno-
Iropi^ aod woiropir mnteriiil. Aoconliog to tnoat aulhorB auch an altcma-
in uf aniaoirMpic lutd ^chi^tlv) i^utnijiic banda which ia obrioua in a dead
nd pfvnrcd fibre exitta bIh) ia tba living fibre ; but aotue mikinUia thai tlie
MB|fllire b unifomilr ani»i>tra|>ic.
SM, when n fibrv iiintracto. la «[Mle of the coiifliAioti previouulT mentioned
bllMHldun and briicht WniU, ihrrv ia no cunfuniuti bclnceii tin; auiai>tropic
«d iaotlBpio maMnal. The aniaotnipic, doubly refrnctive baud*, bright
wiar cnwed Nicola, oocupring tb<' poailion of the dim band* in Ibe reatiog
Sbf«. rendo doubly refractrvc. bright under enixard NI«olB,eTen at the verjr
Uflil at Uw coDtraitino. The iiwtm|HC, einely refradirc bnnd.->. dark under
iMMDd NicfiKoccdpytn); the poeitii>& of the bright liiiml>t in the fibre iit r«M,
IIMUO Hiiiruiiic and dark uiidcr croeacd Nicoh> nl the vrr}' height uf the
flsUvctioD. All Uiat can becMO ia that iherRinglTrPl'mcltveiMtrujiic tMuida
t«naw very thin indeed during the cnnimctiou, while the nnivitnipic ImuiJn,
rtmigli of courw becomioK thinner and broader ia the contriittion. dn not
io tbiu M do the iiotropiir band*; in other word^. while both biitidn
thinner aud bruader. ine doubly refntctive ani!>»trqpic bimd mviii*
MiaenMe at ibe expense of tito »in(;ly re&activc iiiolnjriic Imnd.
{ff. We call altrniion (u these Mcta because they "now how complex la
fe hC "f f-iinimctiDn. The mer<' broadeuinx and «hortt'iiinK of each MClion
If tba 0bn? i* at ItoUom, a L ran* I- •en (ion of llie ntolecules of tli« iiiusi'lo hiIk
Canoa. If we imagiin' n cjiniMny of 100 »oldier« ten ranks deep, with l«n
aM« in rach rank, mpidly, but by u Mriea of );ntdiiiians. to extend out into
a doable line with 'lO ura in cnc^ line, wc ihall have a roii;;b image of the
■MVtfmcnt of the molevnlta during a muKCiilar cmtraclion. But, from what
Ua b«an mM. it ia obvious that tlio moveiiii-nl, in Ktriatei) muwle at l«aM, is
a Twry eomplicate'l otie ; in other forms of cuntmctile tiwiie it uuiv be, as we
«kUI aot. more simple. Why the muvr-mcul Is so complicatml in strialeil
mud*, what purpuaes it serves, why ibu skelcMl intisclcs an; (triat«d, we do
104
Tni COXTRACTir.K TlSSfEB.
ii'>( ai i>r»eiit kiuivt. Afitiareutlj' where xtvlfl nod ntjud contraction h
roguirLHl llit- coiitraclik- t'lMtie iti.ilriauil iiiunck': but how the ttrUtioti Mft,
an Ui upeak, tli« cuntrii<;ti»ii wo ilo ti»( kimw. Wu latirn't !i»y wlint fbnrc in
thr net of coDtrnctioii U to be alli)iu.-ij tn iIr* avvfi-ii] jiaiia. Sinci*, durine
a conlntctioii, the fibrv biilgi-n out more <i])ji'»iti.- In mcli dim dM', nnil
IN in(Iir:itri) itppotiiu- Id witli brljihl disc, since tliL* •liin ili>c i> luorr largely
('(irii|)D«cd i>f HMiKolriijiIc iiint<-riitl lliiiu l.hc Jval ul' thr fil>n-, nnd »iticc the
aiiiHDtropic nintcrhil in ihi! |»<H>itI'>ii of llie dim disc incrvtiMv ilnring a con-
l nidi on. we might iwrlmjw iiifvr ihiil thr dim diec nilht'r ihnii the brizhl
diai: It tho csk'iitially active niirt. Ae^iiniiiij^ that itic Dhrillar Euhstance is
more ahundaiit in the dim discn, while rliu interlibrillnr substance U more
abutidajil in the bright disc*, and ihiit the fibrillar subUaoce is anisotropic
(and hence the dim discs largely an is n tropic), while the intviKbrillar sub-
stance is ieotropic, we niigbl also be inclinet] to infer it is the fibrillar and
not the intertibrillar substance which really carries out the contraction ; but
even this much is not yet definitely proved.
One lIiiD]{ must be remembered. The muscle sulalance, thouj;!) it po*-
»et«cs llie ooniplioated structure, and ^oes tbiriugh the remarkable chan|^
which we have deacribed, is while it is living and iuutct iu a condition whicli
we are driven to »])Mik of ai *cmi-IUiid. The whole ut ii is e^ttciiiinlly nn/itile,
Th« very act ot'coiilrartioti indeed ahows thk ; btit it it> uiohile in the Mnse that
ii'i part uf it, vxivpl of <!ciui'<R the iiuciri and sarcnlemma, nvither dint nor
lirifcht giulKtlnnct-', iii-ilher lihrJUar nor intvrfilirillar aulMancc, cnii be regarded
an a hard and limt structure. A minute nematiiid worm hat been xeon wan-
dering in the midst of lliesulislnnce uf a living rnntritctilc fibre ; as it moved
nlonf:, the. muH-lcsnhstHnce gave way In-fore it, nndclowd up again behind ii,
dim bandx and bright bands all falling back into their proper placo;. We may
wippotv that in this case the worm threaded its way in a Huid interfibrillar
Buhslance between and among highly extensible and elastic tibrilltt'. Bui
even on such a view, and Mill more cm the view that the fibrillar sulmtuneu
also was bruken and closed up again, the mainieuRnce of sucbdelinit* bintct-
loKical features as tho«e whidi we have dwcribed iu material so mobile can
only be etfecied, even in the fibre at rexl. at mime considerable ox{i«uidituro
ofeiii-rgy : which energy it may be ex|)ecieil haa a cJicniical sourctv During
the CuDtraction then' i.-> a still further exjieudilure of energy, some of which,
■■ we have seen, nmy leave the muscle u-i " work dune;" thi* vaergy, Uke-
wiiH', may be rxpccti-il t<i have a dicmical source. \Vc miirt, thfirelore. now
lura Iu the chemistry of muscle.
The Cliemistiy of ^ftt»r!f.
$58. We said in the Introduction that it was difficult to make out with
certainty the exact chemical dift'eremes between tlead and livios «ui>-
Mance. Muscle, however, in dyini^ uudergues a reuiarkalile chemical eliaug*,
which may l>e studied with comparative ease. We have already said that
nil nm.tcles, within a certain time after removal from the biuly, or, if still
rcRuiiiiin)c part of the budy, within a cerLaiu time atliM* " p;neral " death of
thv body, low their irritability, luid that the low nf irri lability, which, even
when rapid, is graduul, is unccwded by an event which i* somewhat more
sudden, vit.. the aiirmiicu inio the condition known ns ri</or utorlit. The
occumonce of rigor mortis, or cadaveric rigidity, ns it is sometimes called,
which may be considerwl ns the token uf the death of the muscle, is marked
by the following features : The living muscle poeseBKe n certain tranj-lucencv,
the rigid muscle is distinctly more opaijue. The living muscle is very exteu-
»iii|« and elastic, it stretches readily ami to a considerable extent whtin a
OIIANCIKS IN A MI~Sl'I.£ DCRINU COXTH ACTION. 105
nvte^i b bUDi; upon it, or vlieu uiy truction is applied Co i(. bui a)ice>lily
innd, under normal circa nuuuwce. c«iupl«telv relurna lo iu oriKiual leiif|;tn
twWn tbe w«i|;ht or moUan is mn->ved ; as w« shall see. however, the
twpfdiir and cuiupleteuen of the mtuni dejie>iiU on tlie c»udilioTi oPthu
nDKlr. a xell-auuriahed activo inuacle n->:aiiiiiiK il» normal lcu;,'Lh much
Mnr* mpdiy and cuoiplelcly thnu n timl uiid cxhausiud muacle. A diwii
ri^d MiMaJe ia muelt l«a» #x[iiisil>li>, and nt ihv Mimv lime miidi lean dantic :
lh» MHela DOW nquina oonri<Uniblv I'on-v to xiretcli it. niid vrhi>n the forci'
h iVMand, Aot» not. aa before, ruturn to it* fomivt loiijclh. To the Imtch
tbt nglMl maicle hiw loK miirli <>( il* formtr sofliwff. and bun become lirmvr
■ad Bar» raaiitaot. Tbe cntmnoc iiilo rigur morti* u, moreover, itccurapn-
itini bf s ilioneniiig or contraclion, irhich may, under nrtain circunMaiict*.
W (■■Mdrnhle. The energy of thi« coiitnietioo is not great, ao iliat any
MUial •borteningMenailr [irevcttted by the prcscnceofeveu a Blight oppoaing
Now tbe cbenii<^l features of the dead rigid mu»cle are also 6trikiu|;ly dif-
I trrtnl Tnttn those of ibe living muscle.
t W. If a (if-iJ tntiM/e, from which all bl, teiidon, fascia, and connective
bate Ijeaiaa much a* puaaible mmoTcd, andHbicb baa been &eed from
,„. _ !>r the injection of " nonunl" ulineaolution, be minced and re|Matedlr
•■•bed with water, the wii!ibiii>;ii will <-<>maiii ciTlitlu f irma of albumin nnil
•vitain eitntcUve iKxtiex, of »)iich we •hull ^\K-ii\t iliroi-lly. Wbt^ii tbe nii^h-
ir^ bia bevB oontinuud until ihr Kii.->h'W'iiti.'r give* no proteid rmetioii. a
hm portion of tnuKb' will iilill ri'muiii iitnliiwilvvi). If lhi« l>c treated with
■ tOpiraetlt. aolutinn of n iiuutnil itnit, ummmiinm chloride Iwin^ (be be«t,
■ luM portion of it will Uvmu- dlwilvcd ; the Foliitioii, htiut'vcr, w more or
!(■ UBparfiKt and Hlten vith didicully. If tbe filtrate be atloncd to full
it9p vf drop into a large i^uantiiy of distilled water, a while Hocculent
MU«r will M j)reeipitat«d. Tliis tlocculeut pret^ipitale ia iiiytMin. Myosin
»t(ir»ieH. ifiviniE ibe ordinary pmteid reaction*, and having the tame geu-
titi rlvnieoiary eom|N3ailiu« a* other pr»teiilit. It is soluble iu dilute sidiue
Mhliuiw, rejieciallr tboae of ammonium chloride, and may be classed in the
^oknlia family, t]»ough it it not xo eolubh' an pa rag In but in, requiring a
Nrafn-aolnUan of a neutral mlt t<t dinolve it; thim, while soluble i»a5
•rlOptr cMt. solution nf giich a #nlt. it w far len ihiIuIiIc in a 1 per cent,
•cimfaa. wtiieb, B« nr have SM11. rvudilv diiwilvt^ piimglnbulin. Fnim ila
•(■ina la iMulral saline Mlutioii it i* prccipilntol by Mturntion with n
■Mtal alt, preferably sixliuin ehloride. and may be piirilieil by Iwing
•kM with a Mturale<i »>luiio(i, diaaolved sgnin in a weaker solution, and
rr I by Mliiraiioo. Diavilved in saline solutions it readily conx-
HI' I healed — i. «.. ia conveHed into coagulated proleid — and il in
•wiliy nf notice thai it congululcd at a coinpttratively low toinpernlure, viz.,
*b«l 08' C ; this il will be remeiulvered lb the temperature al which tibriii-
in It oiagulated, whereas paragloliuliu. scnim-alliumiu, and many other
pteifis do wA coagulate until a hichiT lem|ienilure, 75° C, is rea^'hed.
Skttirf)* of myoeiu are precipiiatt<<l by alcnho), and tlie precipiiate, as in tbe
mt <■•{ otiirr pn>leid*, IwennieK, by continued action of tbe alcohol, alterol
aa*<\«gulBie<l imraluble prnt'-id.
VTe bavc seen that par^lobulin. aitd. indeed, any member of the globulin
(nap. b vciT midily changed by the action of ilihite acida into u body
aSta 'M^-<iANiniN, chamot^riited by not being soluble viilicr in water or in
Ah* Mlioe aolutions bul rciidily tolublc in oiluto acid* an<t nlknli<«. from
ili Mladom in either of which il i« precipitatdl by iieutralintion, and by
I iW &cl that the sotuliona in dilute aciils and alkalies are not ooogulated by
tML When, therefore, a globulin is diceolved in dilute acid, what lakes
10«
piloe ianota Di«re solutioa but a ch^tiiical change; the glMmliM caniint be
got batik ft«tn ihe eohiliou, it h»s been chnn^red iiilo «ci>l-nlhiimin. 8itn!-
larly vhen (>lubulin is diseolvwl ia dilute alkaliefl it i? changed into n/^1/4-
nlbumin : and bruadir B{>eakinK alkali-albitmin pr«(npiui«d by Deutraliu-
tion i-uu Ite cbsujted by Boliition with diliiie ncidM into add-albutuin, uid
add-albiiniiii by dilotc mikalies itito alkali albumin.
Now my<H>iR i° nimilurly. nixl even mure readily tlian is t;I<>l>"li<i< M"'
vertei] imn iioidnlbumiii. niid by Ireutiii); a niU!«lc, either M-BBhe<l or OM,
directly with dilute hy()rrirhli>ri<- ndd, the myiK>in may be c»uveri«d iaUt
lt«i(l-nll)uniin and duwilvcd out. Acid-nlbuinin oblniucd by dit<Milvitig
riukIo in dilute acid ux'd to be culled minlo'iin, niid it uxed to Iw Mid that
n n)u»cl(! ointained «ynloiiin ; th« ntURclc. howcrcr, contain* myovin, nut
i>yn[iiiiin. btii it may bo ueeful to retain the word syntotiin to drtiotc nrid-
klbumin ohijiined )>y iho action of dilute ncid on niyotin. Jty ibe notiun of
diluU' nikniira, myosin may «imilarly bo conv«ricd into alkali-albumin.
From whul bti« be«n above Hated it is obvious that mvoein has mnny
annlofpes with fibrin, and wg have yet to mention eome Btnkin); annlocin;
it h, hoirefer, much more soluble than librin. and speakinf; generally it
may be said to be iuterniediate in its charaeler between librin and clobulin.
On keepiuft, and especially on drving. its solubiUty ia much diminislied.
or (he suliaiances which are feft in washed muscle, from whidi alt ibe
mvosin liaa been extracted by ammoniuiu chloride solution, little b known.
if iraabed mii^le l>e treated directly with dilute hydroobloric a<rid, a targv
p«n of the material of tlie mumle pattM, as we Imve it»i<l, at once into svo-
touiii. The iiuantity of •ynctmiii thuft ob(«in«<l may he taken aa roughly
rqnwBeniing tne ((uantity of rayoi>in provioutly esi>iting in the mutclc. A
more prolonged action of the ncid may distolrc out otlior protcidf, l>««i(lei
inyiMin, left afler the washing. The portion inxolubk- in dilute hydro-
chloric ncid oUNsts in part of the ^tatin yielding and otiier siibelasces of
the wrcolemma and of the conneolive and other timuos between tb« bun-
dles, of the nuclei of thcee li^eiiee and of the librw themselves, and in part,
possibly, of some |)ortioDs of the muscle subetsncc itself. We are not, now*
ever, at preaeDt in a position to make any very deHniie sutciuent as 10 the
relation of the myosin to the Mtructural lealurai of muade. Since the dim
bands are remtereii very indiiiin't bv the action of a I" per cent, sodium
chloride ^lutinn, we may, |«rli(ni^, int'ir that mvwin enters larKely into the
oompooition nf the dim buul^ mm I Ik relbre, of the librillni; but it would
be haznrdimx to nay much itu'ii; iluxi Uu.h.
^60. Living nuwie may be fmiccu, and yet, afYor osrtnin precauiiom
will, uo being ihawml, n:)^in its irritability, or at all evcnl4 will for a time
be fiHind to be still livinc in thu wiwc that it ha* not yot paavcd biio rigor
iaorti». We mayi therefore, toko living musole which has bemi froMU as
still living.
If liriaif fttalrnetih mtuelt, freed ns much as poMiblc from blood, be
frozen, and while frozen, minc^rd, and rubbed up in a mortar with four timss
its weight of snow cunisining I per cent, of sodium chloride, a mixlur« b
obtained which at a (em)iernturo just below Q" C «■ sulKcienlly fluid to ba
filter«<l, thouuh with ditfieulty. '1 he slightly opntevcenl filtrate, or wii-"'^
plamia iw it in calleii, i& at tirsl r|Uile fliiid, but will, when exposed
ordiniiry lem|HTaliii*, become a solid icily, and afterward separate into a
e/ol and Tiiiri. It will, in fact, coaeufate like blood-plasma, with ibis dif-
femiK'C, that the clot itf not llrni and Rbrillar, but Ioom, granular and lloc-
culent. During the coagulation the fluid, which before was neutral or
slightly alkaline, l)ccome« distinctly acid.
cnAxose tn a mitscle Di'itixo contraction.
Tlkf dot b mytmn. It givn ill tbe raifltioiu of myotiit obtninvil (Vi)iu
<1mcI wnacl*.
Tb» «ran rontaitu tn altKimin -vtry timiUr to, if not Hiciitimi wilb,
vrtim-mlbumin. ■ globulin <)ifl«rin^ ■umi^whiU I'rDni, Kn<l cmi&ulitiiig at it
)<>««r iMnpvnilun' ihuD pn»f;lobiilJii. mn<\ which lo (Itningtiifh it IVoiii tbe
globalin uf bimxl hiu bwD (.-itlleH mwylohuUn, eorav ii|h«r jiroti-itli' ohicb
Dol bv (UscHbed heiv. and various "estnclJTea" of vrhifh nrp fbaM
. dlrectlr. Such muscles as are red also conlnin a buiaII quantity of
[lobtn, and of anotltet allied jitgnient allied hitlolurmatiH, to which
ptCMaia. iotlnxl. Ib«ir r«dna« is due.
Tkoa, whil« demd muwli' cunlaiw myoain. stbuniiD, aud Mber protcids,
•ztraetir**, ami ccruin inuluble nuitten. in^ther irilh gi'laiinuus aud oth«r
wib*anm Dot referable tu tb^ niUNcJe Kulmtam-c iliwlf, living mii.*cle euniaiDe
OM aa^Min, but aonc iulwtance or •nlwtancr-!' which Ixuir MMDewhnt Ihf Mime
n-Utiun Kt mTottn thai the ant««cdfDi» of tikrin d» to fibrin, and which jitve
nM> lit BijiMin u]>nn iW denth of the muscle. There arc, imltiHl, rutuMUis for
thinktDff that thv inynHn ari«c« from tlu' (»nvcnii>n of n prcvioiuly existing
bndy wfiidi may be cntlnl myMino^fH, and that tli« c»nvcn>ion lake* [ilaee,
nr nay lake placf. br ilic iiction of n special fermont, ihv L-oovcrsiaii of
Byaaaunn into inyodin bein^ viry analogous to the conversion of tihrinogon
iMfl6bnii.
W« nay, in fact, apeak of riitor mi>rtU na characieriEed by a con^'nlation
of Um muacle-plasfna, oomparanle to tbe coaijulation of blxod-platmn. hut
JUfciny fWmi it inaanucli aa tbe product ia ixrt tibrin but myosin. Tlie
rijtidliy, ibe Um of vupplenea*, iimJ tbe diininl»lied translucency appear tu
Iw at kII e\-«iita hir;(^lv, ihotijtb pn>hubty not wholly, due iti tbe ehunife frutn
tbe lltiid plMHUt lo tlw tolid niyu«io. We miebl oonijmre a living imml«
t" a nnmber nf fine tramparmt ai«nihr«»ous tnues containing plood-pln«niB.
Wb«9 ihla blood- plasma entered into the "jelly" stage of con^ulaiion, tli«
mlvin of tubffi would present many of the phennmenn of ngor ntortb.
Tbry nouM lose much of their suppleneaa and tronshicrncy, and aoquiiv a
evrteiB aoKMinl nf rigidity.
(61. Time b. however, one very marked and important diBereoce be-
imva Um riffor mortis of muici*- and tbe angulation of blood. Blood dur-
bi^ iU onaipualion undergot* n uligbi chan^i' «4ily in its reaction : hut mus-
eW 4ariag the oniet of rieor mortis liec«'nn- diDlinctly acid.
A livm muscle at n*t in in rcAotiim miilnil, or, pumibly from aome
"TTwfrr uf lymph ailhering to it. fninliy nlkiiline. [f, on the other hand,
tW naelian of a ihorunchly rigid miiKch' he ttvleil, it wilt Iw found to Im
MCM (bitlnctlr add. This development of an acid reaction is witueaied not
obI/ in tbe solid iiiitotiched fibre but alw in expresacd mttMl»-planDft ; It
•MBa tn be awH>etnt«) in mmv way with ibc nppearnnce of the mynstn.
TW exst-l caumtioti of this acid rcartion hns not iil urrscnt brcn clc«riy
Mnrknl utit. 8iuce the coloration of the litmus priwfuccd is permanent,
nrhonir acid, which, as we shall immediately state, is set free at tbe i<ume
tine, rann"! he regarded ah the active acid, for the reddening «if litmus
neoduoril by cnrlxxiic otid »i)e«iily dbappean on exposure. On tbe other
Inod il ■* [v>wi)>[i> ut rxirad fnxii n^'id raiuKle a certain ououtity of lactic
•did, or nuber of a varii-tv of lactic uctd known an urcyilactic acid : ' and
il Im* baea thought that tdc appearanoe of tbe iict<l reaction nf rigid muscle
m da* lo ft new jbmation or to an increased formulii>ii of thin Mrcolacllc
\MmSX
■MD «wMla> or lactic uM. «Mc4i ua iMaatfto. tiarlaf thir midi ooiDpotflM),
~ ta(lMltt«HibM>»4 afsddlTlaiheuMSUiyolllKlr ilMi^ta. TbomiHty
tt dWlatiil«b«d » aFMlwUcicU.
108
THB CO,VTKACT[I.K TISftlTRS.
a€td. Then is much U> he eaid io favar of tbia vieir, but i( cannot at
pnaent b« regarded a* eeiabti^hed beyoad dispute.
Coincident vvUh the appearance of tliia aoid reaclioo, tbou^b. ae we have
said, not the din>ci cause of it. a lari:e development of carbonic add laka
place wbeu luuscle becomes rifcid. Irrilable living muscular sulutauce. like
all liviii{{ aubittaiice, in coutiuually reapiriiig, that h in i>»y, U cuutirmatly
consuming oxyjcun and ttiviug out carbuiiic aciil. In the h^dy, the arterial
bkiod ffoEug In the miia<^^ givt» up some of its oxygun, and gain* ft quiintiir
of cartioiiit' acid, tliiiii bccuniiug venous iis tC puKM* through ll>e iniiwlc
Cii]>illarit«. Eviru nlU-r rviiioval IViim the bcMly, the living niujclc coiilinuea
to taku up front the vurruunding atimifphorv a certain i]uantity of Qxygta
and to give out n certain quantity ut' carbonic acid.
At the onset of rigor niorlis there is n very lari^c and nudden inctvaae in
this production of cnrbnnic acid, in tact an oiitliurct n» it were of thnt gst.
Thi« ia ft )ihcn(imeiion dcecrving suecinl atlcntioii. Knowing that the car-
bonic ncid ivhicb is the outcjiue ol the respiration of the wbole body i# the
nwult of the nxidatioD of carbon-huldiu^' aubslances. we might very natu-
rally suppose that the increascti production of carbonic acid atteodaul on the
development of ri;>or niorlia i» doe to the fact that during tliai ereut a
certain quantity of the carbon -holding cunatituenl« of the niucole are sud-
deulv oxidized, llut Huch a view is negatived by the following facl^ : In
the itrst place, the increased production of carbonic acid during rigor inortu
is not accompanied by a correagxindiug iucreaAe in the oouauiaption of
oxyeen. In the aeooiid place, a iiiuacle (of a frog for inhtance) oontaina in
itMlf DO free or loivcly attacbe<l oxygen ; wheii subjected Ut the action of a
merctirial air-pump it gives olf no oxygen to a vacuum. oHering in thi* n-
•peel a marked contrast to bhiud ; ami yet, when placed in an atiuodphero
free from oxygen, it will not only contimifi tu give off carboDic acid while U
rtDiaini alive, but will also exhibit at the onwt of rigor mniti* the sntne
incnawd production of carbonic acid that i* ehonn by n muwlc placed in
u umoaphere containiitg oxygen. It is obvioui that iu »uch a caM- tlw
oarbonic acid does not arise from the direct oxidation of the muscle sub-
Glance, for there is no oxygen present nt the lime to carry on that oxidation.
We are driven to suppose that during rigor mortis, some complex body, con-
taining in itself ready formed oarbonic acid, so to speak, ifl split up, nud
ihiii carbonic acid is «t free, the process of oxidation by which tliat c«r-
I>onic acid was formed out of the cnrbou-holdiug ooustitueuta of llie musol«
haviuK taken place at some anterior date.
Living reitin;: mo^de, then, in alkaline or neutral iu reaction, and the
subdtance uf iia tibrtu contains a coagulable plasma. Dead rigid muscle on
ibe other hand in acid in reaction, and no longer oouiains a ctiagulablc
jiliunui, but in laden with the Kolid mrosin. Furilier, the change from llie
jiving irritaldf condition ^> that of rigijr mortis ii> accomjninicd by a largv
and Kudden dcrelopnicnt of cjirbonic aoid.
It is found, mortvover, that there is a certain amount of {wrmlleliMa
between the intcnsitv of the rigor mortis, the degree of ncid rcucUon and
the (quantity »f cnr^lic acid given out. If wc suppo«e, h wo fairly
mny do, that the iutensitr of the rigidity is dcpeiidunt on the minntity of
myosin defweileil in the fibres and the acid reaction to the development if
not of lactic acid, at least of some other subslanoe. the parallelism betircvn
the three products, myooin, acid-producing substance, and carbonic acid,
would suggest the idea that all three are the I'c^'ulls of the spliiting-upof the
•ante highly complex sulwumce, Xo one has at present, however, succeeded
in isolating or in otherwi*e ilelliiiti-ly proving the existence of such a twdv,
and though the iilen teems tempting, it mny in the end prove lotally
erruiwAus.
CHA\OX» IN A Ht'BCLR l>VRI.VO CONTRACTION. 109
I
I
I
I n. A« to ttie nlh«r prnteids of tnuscle, such ts the albuiDin and the
yMmHa, we know as yet »i>i)iin);' rotmrning tl>e parts trhich ihey pUv and
Um elMBfe* wbicli they undergo in itie livii))> muscle or in ri;;<>r morti*.
Biriiu the Jot which ta found, and that nat iiofre'iiienilr in nbundnnce,
in ibe cunnectira tiaane between the fibres, there is also ptva^ut iu the tnu«-
cwkir aalMtance wiibin ihe MircoIeRtniA, Kluays n-jme, and at timea a Krout
AmX. of fat, chieflv ordioary fai. vii.. Mearin, [Mlmitin, and olein iu variable
pfopcwtioo, but also ihe lUAfe complex fat K-Htbtn. Aa Iu the AiDdivn uf
tba* arvwrKl &ta in tite life of the minwle ne Icm-w little or nothing.
OKtiwkfiimtm, ibe third of the (hre« grnit claiuu-ii in which wc may ^ruup
Um CBVfgy-boldhlK aufaMJUtcea of which thi? animal body and it.-< t<icid an-
alike cunpcMil. tic, proteidit, &(, ami mrbobydrat^-M, are rvprvMMite<I in
iBUfcle bjr a nvculinr body, ^yerHjm, which n'l- »hiill bnvi' to xtnily in detail
btarno. ^\«mu•t here nK-rely wiy Ihnt glyoogrn i» a Ijorly cIiKwIy iillicfi
bt March, haviae a formntn, which may Ix- included under the gt'iH-ral
(maola fi>r ■taroMS.Jr (C,H,,0,), and may like it be c»nverled by the af-lion
i>f acwl*. ur by the action of paniculnr fermente known as amvlolTtic fiT-
Mtnta. into Mtme form of Migar, dextruee (CiH|,0,). or some allied sugar.
Maoy. if not nil. living niMM'lee contain a ivrtain amount, and some, under
rartaia eircuniiitances, a cun^idcrable amount of glycoceu. During or after
tianr lawiiu this gtycoji;eo is very apt to be converted into dcitroee, or an
uBed angar. The mnsclea of Ibe embryo at an early trtage contain a relu-
ilnslj eoonnoas quantity of KlvcuKeu, a fact which attgi^iB thai the xlycn-
fun M moacle n carbohydrate jood nf the u)ii$cle about to be irruuglit up
nto the living muBcular »ulMta»ee.
TW bodica which we have calle<l ejrtmefiftt are numerous and varied.
TImt art wpeciaily intcrettiiig. since It lecnis probable that they uru waite
y€vaaeU of the mclabniigm <>r ihc niui>ctilar subsianoe, and the study of
then BMT be expected to throw light on the chemical change which luus-
cabr MiHtance nndcrgno during life. -Since, as we shall see. musctilar
wbaianea forms bv far the grniter part of tlie nitrogenous, that is. proteid
poetioa of the body, ihe nitrogenous exlraclix'cs of muscle demand peculiar
attratioci. Now the body urtti. which we i^hall have to study in detail later
«a. &r nceetis in importance all the oiher nitrot^nous extractives of the
body aa a whole, since it is practically the one form in which nitrof^noiis
WHCaa loves the body; if we include with urea the cineely allied uric add
(whieli for present pur]Kisee may aiuiply lie regarded! as a varietr nf
Br*w>. we mny «ay that all the nitrofren taken in as food sooner or later
kav(* tlw body aa urea; cumpari-d with ihii> all other uiiniKcnoua waste
ifarown out from (be bwly is Inugnittcant. Of the urea which thus leaves
iIm bodv, a iv)fiuderal>le portion must at sonic time or other have existeil, or
M afaak more eiactly. ilK nilrogi'ii niLj»t hiivi; i-xlHtcd nx the nitrogen of the
pnMMa of nnsailar ntulMtan (.■.■. Ni-vrrthi-liiu. no un-n nt all I-", in unrmal
modltkaa. pieaent in muncular subvlatm' citht-r livinj,' and irritabli' or dead
aed rigid: un-a iiin-» not arise in muK-uIar Milxitanix- il»ell' ns oik- of thir
tMS0adiat« waste pniduct* nf miiu-ular Mibstance.
Tbfvc ia. however, alwuyi^ prrat-nt in relatively considerable amount, on an
avvfkp ab(«ut U.'i.S |)er ocni. of wet muKclc. a remarkable body, trrttlin. Tim
■^ in one sense, a comtmund of urea : it may be split up into una aiul sar-
aain, Tbia latter iKMiy is a methyl glycin, (hat w to say, a glycin in which
lilbyl has li^n substituted for hydrof^en. and glycin iiwlf is nmido-ncetic
acid, a ctiit)j>ouud nf amidof;en, that is a repre«entntive of ammonia anil
MStie acid. Kence kreatin oinlaint urea, which has close relations with
uaasunia, toi:ethcr with another r>>prefentative of amniDnia. and a surplus
«f carbna and hydrogen ermnj[ed as a body beloDgitig to the ftilty acid
110
TUB CONTRACTILK TISSCSS.
•eriM. Wc sIiilU I>nvc to reUiru Lo th!« kn>iiiiii niid «ondd«r its nktioo '
urra und Ui muiidi.- nhcn wc oome lo deal with urine
The oilier iiiir<?gciiou» oxirnctivcH, •ii>:)i hn kcniiD, lij'puxanthiti i.or
earkin), xaiilhiti, luuriii, vtc, ix-cur ill snitill ijutiiitity. nnd a<xii not be dvelt
00 here.
Anibug Doii-tiitrog«iiouB extractives the most important is th« sarcolacUc
ttcid, of which we hiivo nircndv i^pokcii : to this may be added sui^ar in sonie
fom OT other either cumiDt; Iruni glvcogeo or from fome other eoun.'e.
The nsh of muscle, like the a^ii ol' the blood corpuscles, and, indeed, ihe
usli of the tiMueti in frenersi, ea dUtini^'uighed from the blood, or idasma, or
lyin)>b on nhich the tit«ue« live, i« charBct«riz«d by the nrepon<IeraD(« of
|Mlai«iuni aalt> mid uf phufiphates ; these form, io favt, nearly dO per ceut. of
ihe whoh) a«h.
j 63, Wc may uuw past on to the queetiou, What art the chemicsJ
cltnnges which take pbive when a liviii^ nstinK mutcle entvn into u conirac-
linn? Tbcw! clioii^ arv mou evident otW Uie inuwle hait been aiibjcc
lo a proloDgcid lotuiiu.i ; hut there CUD be no doubt that the chemical mt
of It li'Miiiiii' lire, likd the pliysimi eruiU. dimply ilie itum of the multa 31
tliv iiinstitiivnt »ingk- contraolioiiK.
In iho tint tilnci.- llii? niii«cl<- bcconicai add, nol so ncid ait in rigor luortia,
but Etill Huttiuiciilly so Hl\cr n vigorous tetanus to tiini blue Ittmtis distinctly
I'ed. The cause uf the acid roncUnn, like tbnt of rigor mortis, is doubtfiil,
but is in all prubability the same in both niiK-s.
Ill the secoDr] place, a coDsidvmblc (lusnlity of cnrbonic acid if set free;
and the prodtiftioii of carbonic ncid in muscular ooiitrsction is ullogetker
»iiuilar to the produeti'iD of carbonic acid during rigor niorti§ ; it is not
accompanied by any corresponding increiise in the consumption uf oxygen,
lliis n evident even in a muscle through nhicb the circulation of blood k
villi going on ; for though t)ie blond pawing through a coutractliii; muscle
gives up more oxygen ihaii the bliHHl piu^iug through a iwting idukcIo, the
incmsi- in the aioounl uf uxyj;en lukitu up futlii bu-luw tlie incruuc iu the
carbonic acid given out. But it is ntill more mnrkislly nbowu in a muscle
removed from the boiiy ; for in «uch u muscle both tUv coiitiuctioii and the
incr^-iisr in tbc |)roduclion of (rarbonii- sciil will go on in the absence of oxy-
gen. A frog's muscle auapeodetl in an ntnioxtdurc of uilmgcn will remain
irilable for some condderable time, and ut cttcfi viguroua letauus an incrcaee
in the pruductiou of carbonic ncid may bo readily nsccnaincd.
.Moreover, itiere seems to bo a con«spood«ncc between the eoeivy of the
cootraotiou and the amount of carbonic acid and the degree of aoia rewition
produced, so that, though we are now trending on somewhat uncertain ground.
we are naturally led Ui the view that the emcntial chemical proc«« Ijing at
tJkc bottom of a mu»cular oontraciion as of rigor mortis is Ihe spliliiug up of
some highly eomplex subtlaaoe. But here the reBemUaooe between rigor
morlio nnd cootraclion ends. We have no satisfactory evidence of the formii'
lion during a euutraction of any lio<Jy like myonin. And thb dilference in
cJiemical nwilu tatlii-n with an iniiuirtant dilfereuce between rigid muscle
nnd contracting niunclc The rigid mUHelc, ui> we have seen, heeocuca leas
extensible. IcMs ehistic, IcsN (nin»hiii^it; liic i-outraoting muscle remains no
less transhicenl. elastic, and extensible ihiui the renliug muecle, indev<l, there
arc rmsons for thinking that the muscle in couLraeting becomes actually
more extensible for the lime being,
But if during a eontmction myosin is not formed, what ehaiigm of protcid
or nitrogenous matter do take place !^ We do not know. Wc have no evi-
dence that kreaiin, or any other nitrogenous extractive. Is incmaaed by
coniriictioD of muM-le ; we have no evi<ience of any nitrogen waste at all
CIIASOX!* IN A XrSCLB UCRINO COrCTRACTtON*. Ill
alt of n coHtmcUun ; nnd, indiml, ii» w thmlt MC Ulvr oil, thv tlxidy
ih» wsiUr prodiiCU of lti« body iw it whulw Icjid u« lu believe that th«
wtgy of ifav vnirlc don* by lb« qium-K-* of tho IkkIj: oo«im from the polpii-
tU —Brgv (if ckHmo ronitwundi. mid noi of nilngcQ ootupounds »1 all.
Bat la ihi» point wv aha)! hnvv (o return.
^ t M. \V» nay ram up the elt«tnistrr of muscle somewhat la follows ;
Duriae lift tbe tnoKtilur eubetuDcc is contiDually taking u|> from the
|blo«Ml, tnat ta, from the lymph, protcid. fatly and catb'ihydraie lunterinl,
[■■IbM iMrttMv and oxyneo; theiw it huihU up into itself, bow we do itol
kutxB. aod Ml forms ih« peculiar moipiei liviBg muscular aubstaui-«. Tlic
[•srnct oalure of tiita liviu|{ Kibataoce ii unknown to us. What «f do kium
1 1» tbkl it ■• largely coiojw«ed of nrateid tuaterial, iiud Ihai kucIi budir* nx
■jwiBogni, inyogloUiliD. aud nlbumin have voitieihiug to do with tiitt
J of it up.
DBrtei nU tlito muscular »ubiilan<<e, while litkiiijc in nwl building ilMilf
ttwtNiI of ur by moiiwof tl»eabov<:-iuentioiii;il malvtiiik is cHuittiniaily giving
wC oubaair acid ami cootiounlly funniug nilmgvnouN wnrtv tuch as kmtin.
It kW prubably );iTiH oB* aofoc amount of wireolactic a«-id, and poMibly
aiimr DOD-nitngMious walite m«tlvn>.
I>unng n oontractioii therv '» » gn-ut incrawv of carlmnic acid given ofl'
<i( nlbvr Udte acid ur some other stibstniitv giving rise to an ncid reiioltun.
• pvaUr oonmiraptimi of oxygen, though the inereaau b nut equal to the
inercaM nf rarbonic nvid, but, a» far as wi> can lenrn, no inereflse of niin^
During rigor morti* there is n siniihir increiised produelion of nirhorno
lad and of some ulhfr ucid- producing sttlielaDte. uccM(ii|)iuite<l by a remark*
■Ut OMivcrwoD of myoeioogen into myoeio, by which tbe ngtdiiy of iht
iW Ibre u brought abouL
Thermal Chaugt*.
I tt. Tbe cbemical cbangce durin)* a eontractivu set free a ijuuititr of
n, but only a portion .if this eoer^ty appeart in (be " uork done." a nm*
Ir portion lake* on tbt form of heat. TbouKh we shall hav« her«*ncr
klnat ibis subject more fully, (lie leailiui; (kcu may be given here.
Whenever a muscle Diolracts iln teinjwnitiire rixa, indicating thai hwit
iip^'u iiul. When a mercury- tltenDOnieber ia plungml into a maaii of mus-
dbb lucb n« tl>Me of tbe ibijih of tlw doff. ■ rbe of the nitfrc-ury is nliKrmt
lpNilki> muscl«« Wing ibrowo into n prolonged miilnii-timi. More exact
Mlltt, b«iir«T*r, arc olxuimil by inean:i of a thcriii'miii.-. by tht h*lp of
■Uik tb« ritv of ifin|>*-mtur« caused br a ft-m repeau-<l linglc coniriictKHW,
* illiiil liy a Mngli- oinlractlon, may be obMrvnl, mid the anioiuit of beat
fina nut approximately mcainrcd.
TW tlMnnO]Mle may mniiit either of a linjili' junclion in lh« rorm of a awdlfl
fAuffd lain tbe aubstaucc of the iniuctr, or of levtral juiiciiuuB riihcr in tbt
4if« of a Hal surCkce carefully o|>pMed to llie surrai-i.- ot muscle (ibeiHle being
IMMcvd ao »• to move irilb the r<>iittnctiiig inuRcIc. and thus to keep tn« ctiniact
■net), or )a tli« «taap« of a IhiQ nr<)j(«. ih« edge of wlilcb coraprininc the aciiisl
jactHMM. U ibnwl into a maiu i>f musctM and add in poaitioo by ilirm. In stl
OKI lb« Hrllow-juBctlon or juoutions must be kept at a coostaui lemperature.
AaolJwr lielliala mMlwd of dfteraiining the cliBiiirea of tempernluTe of ■ Ihsue
■ haMd oiHKi tli4> nMunrancnl of altera tions in electric reumanci- which a (iii«
•V*. in contJM-t vitb or plunged into tbe ibauo, ondrrgoca u (he temperature of
a» timam ehan|tM.
us
TltK COXTBACTILE TtSSl'ICS.
It has b««n oJeulsUtl that lli« beat given out bv the inuwl«ii of the thigh
ofn frog in n single cantrnt-tii)!) knitHiiiU Ui 3.1 mifrt>-unibt of heat' f!ir Mich
frrnmine of tnll3Cl•^ lliv n^ull lieinK' dbuiinM] b): <livii]iiii; by five the UAu\
BRintiiit of Itviit givcu utit in tive micct'K'tve finite contracliuDit. It irill, bov-
vvrr, bo iHtri.-r U> rcKnnl llitw; figiin.-' nf illustmtivc of iht* fnct ihiit tbr hmt
Kiven out ti coiinmcrablR, riithcr thnn il* diita iSir clnbc>ml« cnlnilaiiiiii«.
Sloreover. we have no wtiidacUin,' (iiinnlitmive ilcl«trriiiiiiilifnw of ihp heal
i;iven out by the miwi-lc* of warm -blood ei I ntiiiiinls, though there can !>e do
<li>ubt that it is much grenter than thai given out by tbe mii«cU'« of the I'rog.
There cait hanll}* 1k^ imy •luiibc that the heat thus M>t free is the product uf
chemical ehanges within the rniiscle — elinngcs which, thotigh (hey cannxt for
tbe reiiBonH given above (^ Hli ) he regarded na simple and direct oxidatioiu,
vet, since they are procoaees dependent on the antecedent entrance of oxygen
into the muicle, may be spoken of in general terms aa a ooiubtiHliuD : eo UuU
the muBcle mny be likened to a Bieam-eagiue, in which tbe cotubuMinn of a
certain amuiini of material gives riae to the development of euer^y in two
fornu, as beat and aa movement, there being certain (guautitativt- relntiont
b«iw«ein tbe amount of energy aet free aa heat and ibat giving ria« to nio^H
ment. We niual. however, carelVilIy guard ourselree again»t prewin^ **^^|
nnnl'igr too cluaely. In the steam engine we can dialiugiiish clearly bi'iwe^^
tbc! fuel wliich, ibruugh ita i-onibiiMi<in, i.i the ^ilc source of energy, and the
niachiiK-.ry, which is nut ci>DHUiued to provide miergy, and only wfl^rx near
nnii ii-ur. In the muscle ne cannot with certainly at nrnent make eucb a
dittinclion. It may t)e that the (■bimiiciil ohaiigea at tlio bottom of a con-
Imi'iion do not involve ibc real living mnlcrinr of the librr. hut only tome
»iib»lnncc muniifactun^d hv tbe living miitcrial and loifgi-d in unmc way. wc
do Dot know how, in th« living mnterinl. It may be that when a lihre con-
tracia it i« this niilKitann- vrilhin the fibre which explode^ and nut tbe fibre
it»elf. If we further suppose that this substance is some complex oouipound
of carbon and hydrogen, into whivh no nitrogen enters, we ■ball have an
explanation of the difScuhy referred to above ($6''J), namely, that tiitro-
geninrs waste is not increased by a contraction. The special ooiitractiie,
carbon -hydrogen substance, mav then be compared to the charge of a gun,
the products of its explosion being carbonic and .'tfircolaciii' acitbi, « hilc the
real living niutennl of the fibri; may be conipun-d lo the gun itwif, but toa
^n which itm-lf is continually nnd*-i^iing <'hHngv farlK-yimd nH-re wear and
tvar, among the prnihiclH of which chnnge nitrogcnouH bniiirx likr krcntin
an- convpicuou*. Thi» view will cortuinly explain whv krealin ir pot
increii«'il during the cr.ntmclinn, while the carbonic and lactic acid* are.
Hut it must be rcmi'mhcn-it ihnt wicli n view is not yet proved: it ninr be
the living tnnurial <>f \hv fibre. n» n wliole, which is continually breaking
down in nn expii.uivc ihroniposition. and as continually building itself Dp
again out of ihc nmicrinl supplied by the blood.
In a steam-engine only a certain amount of the total potential enerj^' of
the fuel issues as work, the rest being lost as heat, the proportion varying,
but the work rarely, if ever, exceeding oue-tenib of the total energy, and
{{^nerally being Icm. In the case of the muscle we are not at present iu a
pcsition to draw up an exact equation between the latent energy on the o«e
hand, and tbe two forms of actual energy on the other. We have renson to
think that lh<- pi-oporlion l>etneen heat and work varies conxiilerablv under
dil1er4-iit nrcumi'lanct'S, the work .viiiit-iinm ri*iiig nn higti u* nne-fil^ti, itoni^
linu-H poi>»ihly »inking na luw an one twenty-fourth of tn<i total energ}' : and
t llicnurTO-aiiltlieliiiBiuliliKraiiiiiwiir uAtor mlud one (legTM OnlicmW.
CRAN'OKS l>; A UrSCLK DCItlXO CONTRACTION. 118
ohMwatiow kcoi to afaow that ihe );reat«r tlie reebtaooe wlikh ilie iuu*c)e
ham to overconw, llie larger Uie proportion uf the total energy exneiidnl
wbieb ifMS out as work done. The niuscle, in fact. »eeiiid to be no tor M>]r-
nyntMia^ that lii« more work it liua tu do tiie greater, within oertaiQ
U^la^ ii tbe eranomir with whidi it mirksL
I^tfdjr, il niuM be ren>eu]l<«n-<) that the giving out uf h«at bj- the niuti'le
h Bot eanfioeil to the oocOAiunii when il U nclitiilly contracting. Wbeii, at a
laMr MrioH, no trval of tho bent of tbc body gcnenUIy, Gvi(lciK.<o will Iw
bnMgDt furwonl that lb« niuaeltis, «vi-ti wbrii at. rent, are giving ri««i to Iwtit,
•o thftt tW bmt giYOO out at a contraction is not some wholly new plHiiion-
MKwi, but a t4-inporary exaggeration of what is conlinually going on at a
mon SMble nU\
Elettrieal Chan'jt$.
t M. Braidoi ohemical and tberouil cbaiigea, a remarkable electric cfaango
ukr« plaop wbsMver a mutcle oontnirtA.
MtmU-eurrmti'. — If n muscle be rcmovrd iii an onliuary manner from
tW bidy, mihI two non-polariuiblc eb-ctrndcit,' conii«ctc<l with n delicxtc
plTanomctor of many convolutions nnd high resistance, bc plncL-d on two
ri«.».
■-if
XoK-FouuusLB ELsnnoon,
AOtilM* tmbti i.ik> anulcunkifd dBc fllpa eenMctal Mil l> ihvir tQ-r«t4]TC«i1na; t.a,ib«
■M adfkM* •■'■Una ; M /^ Uk |4iisi4 fUna-otar: (', Itn torUonof (he rtiln>-cli) |iluf teqjeH-
mimmltm^maal ikviuto; ihUcuibcwMlitnl IntDtOTrwiulntfonn.
piint* of th« surface of the muscle, a dellection of the galvanometer will
plftM. iiitlicaiitig iliv exiUenwof a current giaasing through iliu gal-
from Ihn uJw |HMnt of the muMie to the other, the direction uid
(if the ilcHrction varying accurdiug to the position of iIh> pcnnta.
IVe " DiUMlr-curivntA " thuK revealed are seen to the best advimtag*- when
ijha nuBck ebown i> n cylindrical or prismatic one with parallel fibres, and
ehm lb« two tcndiiuwis cndi ar« cut off by clean iuciaionn at right luigles to
tha tottc axi« of the Riu>i-)e. The niusrle then presents a tntnsvcnH; section
I'anlfleulf at each end and n l()iii!ilu<linitl sur&ceL We may «p«ok of the
luur as tieioK divideil into two c<ium1 \wris hy an imaginary tmn^crse line
• n«i CTlt Mn MiM^il •■•enttall;oI*iJlpn(UifindWi<mKi4}nMriMilnG.dl|>|4nCtiilOB aVarafml
■iMU^araMMllteiv. wlDcb la tnni l> biautlii into (»tui«dton with the Dorm otiDUKtctnr
•^^ HS Wtm a* MOct or •MiiB4t>r nalMtaeil vllk Bunnal lalliiiQ clilurU* (dIuOui ; It II
MIMMM (Iw) 1)^ ■<<■>' •■louM bt Uii>iniiKhlr uiul^naM. TliU fiinii ••( «)«oliDd« sIt* iIm m
M MMlaUOn Uiaii ^ rfinpU |<l*Uiiim "t miirw <*t«l(odW riir <>1>T Bllbrai ■ toaaiKOiUt
tmmJM. m i>M •nil tiw ikaiu. hIiIpIi ntOttcr mu ob ibc iIhuc iMir !• lelul do br tin Ubdc
tMbn <t •nr uauM ottdi «nppt« or plaUiuna It Id IM*lf mndcM to tfrrelop ■ omont.
8
114
THK UONTBACTrLB TISSUES.
OB \tB aurfiioe called tbe " eiiiialor," coDinii)iDK all the [toints of tli« nurfiace
midws}' between Uie Iwo end^ Fig. -il U » iTiajfrauimatii- represenUUtoD of
such a muwlc the lint! of/ hvlm; the eijuiitor. lu «uch a iuuhU iHa develop-
ment nf the muidc-i-urn-tiU b> fiiuinl to be aa follovm :
Tlic grmMvt lU^HfCltciD i.>> (rlntcrreil wlicn iiiiu electrode i* [tlacoil iit tt>S
mH-poiiit or efjtinlor of th« iiiiiiicic, iinil the o()u-r ■( oithtr cut end ; nod
the dedection is of »iich » kiml iw Ui xhow llial [lotitire cumnti' are con*
tinunlly pacing from thf equator ihrnugh the ciilv«noiDe(cr to tbe ctit end ;
that is to WIT. the cut end m oegntive. rclnttvi.My, to the equator. The cur-
rents oubdde the muticle mar Im' coiisidenxl as citmpleled by ourreots in the
munefe frota the cut end tn the equator. In the diagram. Fig. 'M, the arrows
Fim.u.
It **
:....---<y
DTiunjkH iLLi-ftainini mi Ei Jx-nuc CXnun* or Knvi *m HtwLK.
ncliiK I'Unl] •lJiiK"""'<*i'c. « '■"<} wrvE fnca i4Ma (dtbator nmvcorof ipUMElVk oxc*)* IbM Ux
(mmul* ■! ih* irsiifircw MoUon nnntit l)c uliown la ■mrre. Tbt arron ihovr ih* 4l»cUini or
the cumnt thraiiRli ili« Rtlnnoiuolor.
nti Uro oiuiMf- Tlio >lNiiim<iil riirrcrili kr* IIuiM ■hown by Uiv duk ^m, U (Mm d, ■!
F'jUDilnr, (A > iir lo y «l ibv ctjl viiilg, Tbr cutn-iil rrain <■ lo c ii mmker than fram a to y, (houcli
bolU, ■* nhowo by Uip arrout, liavi? the mhuc <ILrucUaD- A ontfvTkT [■ hhc>uD f^gm «. ulilob U a^U
UwwiuMur. tu /, wlilcb la riiiilior rmiii IIih pr|tiaiar Tlwcurninl (In tniwrloi (ruin • total In Um
ctroiiuIcnnM 10 • iioint n«sn>r the cralrvur ttit inniVDnN «ecUoii li fhmvn al tK Fram a to ho
t*am I lev. Ibvn u do nunol. wIuUcbIiM b; tbo dotlcd Uo
indicBte the direction of the ciirrenU. If the one electrode placed al th*
equator ab, Ihe elfecl ix tli« j>aiui? at whichever of tbe two cut ends, x or y,
llie other ia placed. If. uito elevtr<ide renmiiiiti|i at the equator, the other be
ahiAed from the cut etid to n sjiot (c) noaror 1" the eq^iialor, ihe current c"n-
tioues to have the >ame dirn-tion, but \* of li-s.-< iotciisitv in ]ini|Hirliou to ihe
neameu of tbe elccimde.i to each other. If thi' Ino efiHclniiltn be plactil at
nnf<{Ual diatanceii 'f and/), line n:i either «idc of the eijiintor, there itill he
a ft<i-hle current frimi the one nearer llie cquutor to the one further oA*. and
ih-' i-urnml will be the fii-lilcr the more nearly ihey arc «i)ui(li«t»nt fr«m tbe
e<|uiktiir. If they are uuitv equidiaiant — as, lor instance, when one is placed on
one cut end (jr) and the olher on the other cut end (y)— tliere will be no cui^
rent at all.
If one electrode be placed at llie circumference of the transverse seclion
and the other at the centre of the transverse section, there will be a ourretil
through the gntvanomeler from the former to the latter; there irill be a cur-
rent of similar direction, but of leas intensity, when one elecUode is at tbe
circumftnence ^g) of the transverse section and the other at 8oiDe point (A)
CIIAXQKS IN .1 MISCLB OIRINO COSTRACTIO.V. 115
I
liiwwr Um Mntn oT Uw tnimr«TBe •ection. In fact, Umj i>ointa which nrc
wtali>tly DK»l potitivs uwl m<M atpi\i\i- to each other »rv pninl^ mi tho
■laaMr Bod Um two entrw of the imiiovrnH; HcLton« ; nnd t\v! xMnmty o(
iMcnrraot Iwtwfwi wijr trro pornti will depen'l on the rcfpcctivr disUiu.'c*
of tkow points front the c^untor and from thv centre of thp ininaveree
Asilar currenu may be obserred nhoo the longitudinal surfiioct a not the
Datanl but an anificinl (^ne: indc«l, tliev mar be wiuiesMd Id even a piece
of miMcle, provided it be of cylindricof Bhape and coropoaed of parallel
6 bra.
Tb«M "roiuclft-ctimalB" an »o4 mere transitorv ourrents. dbaipi>enring
•a aooo aa the circuit M cloaeil ; on the contrary, tlief last a very c»iiiudfir>
able lime. They inuM. Iherefoiv, be inaintaiued by tome ehnnjcea ft<noK mi
in ib« mDfcl»— bv continued chemical action, in fact. They diditpjicnr a»
tb« irritabilitv of tl>e muscle vnniiibi». and are ooiioecled with tluBie nutritive.
•(>-<k1I**I vilnl. rhangtt whioh mninliUD th«' irrilithility uf the uiusd<^
MMclr-cunrntJi ouch iin have iuM Ih-<-J) dcetcribfd, may, wu repeal, Im-
nbaamd in any cylindrical oiumic niitably prepared, and similar curreiitJt.
witb varialkna which need not he diiwuNMtl litre, may b« aeea in miixcUv of
imiptlar aluipo with obliquely <>r otlii;rwii>c arranj^ed fibreiL And du Iioia>
BfyBMMMl. to whom dliefly wo ak iadfibted for mir knowlcdgv of ihcw cur-
fOda, baa been lod to regard Ihoni aji naCDtinl and important pnifn-nire of
KviBr aitMl^ He has, moreover, ndvanoed the theory that nnucle may be
WMaml m comimMil of eledro-motivfl porticlca or mokcules, Mcn of
•hidi, like the niii»cle at large, hiu n positive equator and negative cods, the
*hcile mueele being made up of thew molticulm in eomcwliat the same way
(Ut ow an illuAtralioo which must not, however, bo slrnimil or ronsideriKl u
ID Hact uaej aa a raaxnet may be aupptMed to be mode up of ma^netie par-
tick* each with ita north and south pole.
Then aiT rca«noK, however, fur iDinkinn that Uwae mnscle-currenta have
MHich ftioiUraieiital origin, that they are, in fact, of surface, and, indeed, of.
tnttdal i>ri);iD. Wiilcut entering into the cuulroversy on thin qu««tlon,
tkUlowing ini{iorlunt fai;ti> miiv l>e roeniionnl.
1. Vbcn a muKlc in esamiixsl wliJle it ntitl relainn untnjurwl iiji natural
tarminatioDR, the cvirrenUi an* miii-h wtmker than when artificial
•eelioDS have Itven niiuh- ; ihe natural ti-niliiiiiiin end is toM iMin-
lit than ll>e cut guriMc-. Hut the tdnitiiioii.- end !>i'rama at oneo Motive
■km it » dippvHl in wnler or nci<l— imU-cd, whi-n il i« in any way injured.
At la* roughly, in fact, a riiim-Io in ln-»led thv Ii^m evident are t^>: nmiicle-
qratMa: aad it is maintained that if adequate caro be taken to nutinlain a
Hsde in an absolutely natural condition, no such currents as those we have
bndeKribing cxiM at all— llutt nalunj living muscle is itotUdrie, as it ia
olM,
1 The surface of the uninjured inactive' ventricleof the frog's heart, which
iipnrtically a mass of muscle, b isolectric, no airrent is ubuinod when the
Jwtrodsn are placed on any two points of the surface. If, however, any
pn of the surface be injured, or if Uie ventricle be cut acroBso as to expose
tnit Mirfai.'c, the injured spot or the cuiHirfaeebeooDiesatODoeiaott power-
bUf Mfpilve toward the uiiinjurml surface, a strong current being developed
•Istfc fWRt* thniugh the |j;HlvBnoa>elcr from the uninjured surface lo the cut
mftoa or to the ii)iure<T spot. Tlte negativity thuH developed in a cut
wfen pHMK ofi* in tite course of aumc houn, but may be restored by making
tftnb eat aiul eiposing a ttvth surface.
■ Tlia ncnaMIr at It* iKIiil InMtln will t* «(■ NtMsquwilly.
116
THB CONTRACTILE TIBSfES.
The temptnvry duratioo of the ncgstiriiy after injury, »iiH its nae
opOB freah injury, io the case of ihe ventricle, iii coutraat to the man peiv
inineDt n«4^[iviiy of injured akek-tal mUBcle, U explained by Ihe diilerenl
atnivturo of the two kioda of iiiuaole. The cardiac muscle, aa we •ball here-
nfWrtee, is oontpoeed of ahort fibre-oella; when a cut is mode a oerlaiii uuro-
her <>f thcae fibre-cells are injured, giving riee to negativity, but the injurv
done to them iao]m witli ibeio, and is doi ]>ropagated to the oells with which
ibcy nre iu otiiituct ; beuce ui)uu their death the uegutivity and the current
<lisnp|H-ur. A friwh cut, involving new cutis, pr<Hluces trtiAx nQ{«tivitv attd
a new ciirrtint. In the lou^ librcs of the itkulctal niimclc', on [he other hand,
the eAbct« of thv injury nre ulciwly ]>r(i|uigut(.il tiloug the fibrv from the fpt>l
injured.
Now, tvhcn a mtifcle in cut or injured, thu «ulvtancc of the fibm dies at
the cut or injured Kurfm^. And nmiiy iihyxinlngivt*. among whom tho oxMt
prominent is Hertiitnin, hnvi; liccii M by ihv iihorc and ulhrr tact» to the
conclusion that muflLlc currents do not cxitt nalurally in unloucbed, unin-
jured muBcIetf, that the muHcular subetanra h naturally, when livinc,
isoelectric, but that whenever a portion of the muscular substance dtea, it
becouifD, tvhilf dyiii'j, negative to the living eubetiince. iiud thus givee rice to
ourreiits. They explain the typical curronis (as I hey might be called) inaiii*
feM«d by a muscle with a ualural longitudinal surliice aud ariiticiul trans-
vtsna seotioiis, by the fact that the dying cut ends are negative relatively lo
the T«#t of ihe muscle.
Du Buis-Heyuioud and those with bim ofler special explanations of tho
above facts and of <it her objections which have been urKe<l against the thvory
of naturally existing electro- imilive moleculea. Into theae wc cannot enter
here. Wc mu»t rest conlvnt with the atatement that in nu ordinary tauacl*
currents such ns have been described may be niln^'sscd, but that atronc
arguments may l)c adduced in favor of the view that these currents are not
" natural " phenonienn, but ciwcntially of artilicial origin. It will, therefore,
be lie«[ lo G]>eak of them as currmU vf rest.
i 67, Currviitt of action. Xri/atiii! I'ariitUon of the miuolc-eurmi(.~-The
c(introvcrsy whether the " curreui^ of rest" observable in a muscle be tif
natural origin or not, dow not atiect the Inith or tho importance of ibe fact
that nn electrical change takes place and a current is devetii|>ed in a muaclc
whenever it enters into a contraction. When currents of r«t are observable
in a miwclc, tbtse are found lo undergo a diminution u|Hm the occurrcnoe of
a contraction, and this diminution is «{iokeii of as " ilie negative variation "
of the currents of rest. The negative variation may be teen when a miiKle
is thrown into a ^ugl^ con traction, but is must readily thown when the
muscle is tvianiuxl. Thus, if a pair of electrodea be placed on a mitscle,
one at tho iKjunior and the other at or near the transvene section, so that a
cousidcmhie drflectiou of the galvnuonieter ueedle, indicating a considerable
current of roit, be gained, the needle of (he galra^»nlt^ter will, when the
muscle in totaniu-d by an interrupted current mmi through its nerve (at a
point too l»r from the muscle to allow of any vscRfm of tho current into the
electrodes connected with ihe gid vuuo meter), swing buck toward zero; it
Kturua to its original dcHcction when the tctanidng curront is ^ut olf.
Not only nay this negative variation be shown by llie gBh'anoine4er, but
it) M well as the current of rest, niiiy be used as a galvanic shock, and *o
employed to stimulate n muscle, as in tho experiment knuwu aa " the rheo*
tcopic frog." l''or this pur|>osc the muscles and nerves need to be very irri-
table and in thoroughly good condition. Two muscle-nerve preparations A
and B having been made, and each p!ac»l on a gluss plate fur theaake of
insulation, the nerve of the one U is allowed to fall on the niuaole of the
CHAXOES
A MUSCLE DL'RINO L'ONTltACTlOS. !17
Mh«r A in naeh
tllBt<
of ihe
I
f
Dorve cotiiM in coniaot witli tbe
»|B»tar of (bt mtMcU, uni) miutlicr point with one end of the muBcle or with
• potat «t Utm» diflBncc from (he oiuator. At the moment the nerve is let
fiul wmI conlart mailr, > cnireni — vir, the " current of rest " of the mnacle
A — (Mar* ibroaeh (he nerve; thi» nclH na a ittimntuB to the nerve, and m>
CHBMi ■ mntnicliun in tbe miwcte eonnecte<l with a nerve. Thus, the muscle
A BCt* a* » b«tU!fy, the completion of the circuit of which by means of ifae
Dvrre of B ivrvr^ m a itimutua, eauaiiiK the luuftcle B to conlraot.
If. while the Dcrve of JIH stilt iu «<i>nlact wiih ih« musclenf J, the Oftrre
of iha tatMt is tetaniied with an inl«rruplcil current, not milir is the muscle
>'.f .1 ifarown into teunuD, but aim* that nf B, the reason heiuft lu fulluwa:
Al eadi spasoi of which the loianun of A ia made up. there is a negative
Tsrialinnoftberonicle^urreiilor^. Kavh negative variation of the raiitck-
mrrvnt of A servcB w a >limuli» Lo the nerve of B, and u hence the cause of
m i|— > io the muscle of A; and the stimuli fotlnwin); each other ra]>i(ltv,ns
Wit^ imduce'l hy the tvtanit* i>f .-1 ihry miu>t d<i, the spaatus in li lu wfii<'h
thtif giw rwe arv alMi fuiii'd into n tt'liiniia in li. H, in fact, cuntracl« in
banaocir with A. This csjxTimrnt rhn-Hr. that thi^ negative variation ncootn-
puijriitg Uw uianus i>f a mtiKic, though it caiiiei odIjt a onzlo Kwinir of the
gstvBBonMer, Is really made up of a scric» of negative rnriations, each tingle
B^pttlv* variation corresponding to tbe single mmxam of which the tetnotu
is RMJa up.
Hill an circlrical chang« may be manifeiiled even in cases nhcn no ciirr«>t«
of m\ tfsift. Wo bare stated <§ titi) that the 'iirfnci? of the uninjured inac-
tive ventricle of the frog's heart is isoelectric, no currents being observed
mhta the electrodes of a galvanometer arc placed on two points of the sur-
botk Neveribeleea, a tnuat dblinct current is develo|)e(l whanenr tbe ven-
tricle omtractsL Tht* may be (liowa eitlier hy tli« Kalvanonieter or by the
ifcwacoptc flrea. If the nerve of an irritable musc&nerve pre]>aration tw
Ud OVCT a puMUing ventricle, each beat i» responded to by a twitch of tbe
■MBit nf tbe ]>repar«tion. In the eiue <>f onlinary lausolea, too, instances nocur
In wKiefa It vetm impoKiblci to rceard the electrical change niunifniMd
doriai; iW contrmctiun as the mere diminution of a preexisting currvnt.
Acntrdingly. thuar who deny the cxb'tence of " natural " muncle currents
yak iifa uidscte as ileveloping during a contraction a " currunt of action,"
iiiasiniiul. a« they believe, by tbe muscular sululimcv nx it in entering into
ika siste of cMntraction becoming negative toward the muscular suUtAUC*
•Udb it BtttI at rest, or baa returned to a
M(* nf net. In fact, they regard tbe
o^pttivtty of muscular sulwtance as char-
KMrwiic alikeof beginning death uud of a
kfinniiiic wntraction. So that, in museu-
br onutrartioo a ware of negativitv, sturt-
iag fniin the end-plate when indirect, or
ran the {joint stimulate*! when direct
ttinolatlon it used, (Huaca along the mus-
oilat sabftance to the ends or end of tlie
On.
If. for in*t«iioe, we tupnote two el«ctr>>d«a
flsml on two ]M>ints (Fie- ^'* ' A and B of
lUt« about to he slimuTated by a single
MnctioB-alKick at one itid. &:fbre tne
idmutati'in the Hhre is tsocleclric, and tbe
Mnile of the galvanometer dands at leru. At a certain time alW (he
4fvk haa been sent thmugb the stimulating electrodes (r), as the ware of
|ria. ».
118
THK CON'TRACTILE TtfiSUKS.
OOntraeUon ie tmreUilw down the fibre, iheBecliou of the fibre beneath .i wil
become neRnlive toward tibe rest of the fibre, and so negiklive touard lh«r jior-
lion of the fibre under H—i.c, A will be negative relatively lo /i, and
this will be shown by a deflealion of the needle. A little later B will be
eiiterinK in lu contraction, and will be beeimihij; iiei;atiTG toward the rut of the
fibre, incltidin^ the \mn under A. wlitee oefriiciviLy by tins tiniv i* iMMinj; off
— that u t^i any. B will now be negative towani A, ami ihin will be *nuwn by «
d«declioti of Uie needle in a direction oppotite tti thut nf thit dtiHrciioii which
haa juNt previously taken place. Hl-iio-, Imtwcen two el«etmil«v plno(>H along
n fibre n uugle wavu of c-ontntnlion viill give rice to two cnrrenli> of diKerriit
phn«c«. b> a <]ipliiMio diange ; and thitt, indrtsl, it finind to be th« ciwc.
This being ko, it i» obvious that the clectrirni rcault of t^lanizing a mutcle
when wnvf tttier wave followx along each tibrc is a complex matter ; but it
it nmintainedtliat the npuurojit negative variutioD of t«tanuscaD be explained
V til* net mult of a aeriM of ourrentfl of action due to the individual con-
metioiiE, the second phase of the current in each contraction being l«88
Durked than the first phase. We ciinnol, hoivever, enter niore fijUf here
into a discussion of this difiicult subject.
Whichever view be taken of ihe nature of these muscle-currents, anil of
tlie electric change during conlrnction. whether we rei^rl that change lu a
" negative variation," or as a " current of action," it h iiii|>orlant to reruemtier
tliat it takes place entirely during the latent ]>eriod. It is not in any way
tlw reault of the dinnge of form, it is the furerunuer of that change of funu.
Jurt a* a nervous impultw jiaases down the nerve to the niuicle without any
rbible changes, to a nicile<iular change of Hoine kind, attended by no vtniblc
cv«nt« kcniwn to u* at prment, but only by au electrical chaug«, rundi along
ihe muscular fibre from the cnd-phite to the end* of the fibre, prefMiring the
way for ihe vieihle chungt' of form which is to tuUow. This molecular itivi»
iblfl change is the work of the latent periixl. and careful obaervftlions have
shown that it, like the visible contraction which foltom at its heels, travels
along the fibre from a spot stimulated toward iJie end of the tibrea, in the
Ibrm of a wav-e having about the same velocity as the contraction, TU., about
3 metra a eecoud.'
T^fl Changes in a Xcrv! during Ihe I\tt»a</e of a Xtrvotu Impttte.
g 68. The change in the form of a niu»clo during itc contraction Ua Ihing
which can be socl^ and felt; but thr^ chruigoi in n norvu during it» activity
are invisible and impalpable. Wo :^timulnlo one i^ml of a oorvc going to a
muscle, and uc see this followed by a cm traction uf (he muscle altat'lied to
tlie other end ; or we etiiuulnle a nerve still connected ntlh thi.' central ner-
TOUB ayatem, an<l we sec this followed by certain movenKints. or by other
tokena which show that disturbances have been set up in the central nervous
•y«teni. We know, therefore, that some changes or other, ooustituting what
we have called a nervous impulse, have been propagated along the nerve ;
but the clinnges are ouch as we cannot see. It ia paeaible, however, to learn
something about tbem.
Strveture of a nerve. An ordinary nerve going to a muscle is composed
of elementary nerve fihrca, luialogous to the elementary muiiicJe Bbree, rnu-
ning lengthwise nlong the nerve and hound up together by oounectlre tivmea
carrying blood vcucls and lymphatics. [Fi>!. lift.] Each fibre Is a long ro<l
or cylinder, varying in diumeicr froni U-ss than '2 ^ t«> 20 *>, or even more, and
> Iiillw mttMlnoT IM troit : liui. u wrhnTciwri, tinvlns pnitubl; a lilehsr vclodir In tlie Inlacl
iiuniultui miualM. wlUilu lh« living boiljr. ftiul nryiue ncKinlliig w dttFomitUKM.
CUAXOBS
A Hl'SCLE Dt'RINO COSTBACTION. 118
Um MTwml 6brN mrr uriuigvd hj llic codhocUvo tissue into buiidlw or oordti
■ng *loDg tliv length of th« ti«rv«. A largo n«rvp. micb us thi; scinlii;,
many conk of varioiu uwm; in such b case the t'^nnevtivc tinue
(Fib. as.
^-^^.
II. - iLiii NutviorlUii. IHMuMld.l
'<<1 Imnnlln' Kk. [iruefM ttom tW f^^'
- ..iinincciHiUDuoiwwIibtbouidaiitaiMini,
'IIw MUiUpaUiv-ltoiM nbrlla oT Um vmlimvu-
I
r*Bi or t 6lan»Mi Of on at mi. i^i - 1. <
f. fiMmfllmia. anwIMIni ot ■ nnmtor ol r/
•MMBB, t«MtiHlliitolteliil«ttcra(UwliinlFi.i
« liMeHa tolUHMDu Una* taweni tlx lune-lltinji.
asB M» tMU cut KiiaM ■• nno pdn« oAan kpicarlns io mibMlli Hut lunrMlbnB with a olrcU of
■taatitiiMinrtl^MUIiaf Koyuid BtUW. Kuuicluoa DUck! uT oaDOCcUw-Ilana cslliwo «Ui-
kiMnlia ik( cBdODMUl^ : r. twtlouorsIika>liMHl )
krtWMO th« fibm in eadi corcl in niurc dolirule tliun tliut trhivh biodj Uie
conlt tngetlur: eHch oonl bw n io<irr <>r Iiw» ilixtiiicl ibeuili of ooitDectlv*
liwK, UM s BiuilAT but MautCT >l>uitlh pnit<.-clii the whole nerve. In smmDer
BCTTW the oord* ure law ia atimbcj', niiu m vrrr tininll nvrvo inny <'i>nHiiit, to
to^iwk, «f one corH irnly, ihnt i» to mir, it )iii» one ■•ht^'iith for ihv whi>l«
■trr* snd fine cnuiivotive lianie binding together all the (ibret within the
ihaUi. When n large oerre dividn or seoda oS brancbea, one or n>ar«
ofdi l«va the tnink to form tlie braoob ; when nervea are joined to form a
(JbtM, one or more cords leavinj; one nerve join another nerve ; it is. as a
ntioolrwheo averir sioal) nerreiedividin;; neurits end into delicate twigs
ihlfivMon or brauobing of tlie nerve i.- clTccliiil or owisled by division of
(kofrre fibea Ifaemaelvea.
iiulf all the nerve flbrea commsiiiK nn onlinary nerre. Much oi that goiiu;
(•■naaolv, Ihougfa varying much in lhickn«Rf,hav« thcsiiim' feAturo*, whlcG
*it K follom: ^eeo under the micrrBcopv In n ]H:rfvcl1y frunb cuadicion,
■ilkat the nae of any n-iig<-nl«. ivcb Hbrr appvan iw a trniuiparcnt, but
— iwhsl reft«etiTe. and therofoni brigb^lonkin^. rod, with ii shMqilvib? fined
Mlliiw. wbii-h I* fbnrai!(<-ristteally double, Ihnt is to mv, the «harplinv which
■■is the iMilude of tW 6bro is on nicb side uf the fibre ncconiftanied by a
•nttd UlMplinUlol to iliclf aiid following such gentle curves lu it fthows,
hliMher iu*arcr thr Kxii> of the fibre. This it ispokeii of as the lioubU eon-
^^[Fi^. •i7]. Biiit is naturally more oouspicuou*' and niorv easily even in
ll> '":'■■ ritim ill the thinn-T fibreii. The sub^litnm of the fibre betirMO
r conioiir linos apnean, in the perlmly ftvsh libre. honiogenMlW.
U uw tiknt be inu-cd ulong lU eourae for some little distnno- there will be
taaat intervals an appearancw na if the fibre bad been straiit;!*^'' !>>' a Iig«*
lue linl tightly round it ; iu IransTerse diameter is mddenly narrowed,
tal the double contour lost. Uie fibre above and below being united by a
■smi* »hi.rt isthmus ouly. [Fig. 3H.) This in cnlied ii awU. a ii.xle of Kan-
nrr. and ugwu exaininatinn it will be found that each Rbrc is markitl regu-
bth abmg its Im^th l>v nodn* nl interviils of about a roillinietre. If llie
Un kp vxamtned with ^urthrr am tbero will be teen, or may l>e seen, about
■iJoay brtweeti every tu-<i dihIw, im oval nucleus lying cmliedded, as it were,
it ths iwiilin* of the librr', with its long axis itarallel. or nmrlr m>, to the axis
IlitlUH XKHic-nnaL (UapiUlulKO
TlilW of Ihvin nn Hm: niiv iif wlilnli
It rvufttHT, one of mIdillliiiE iMclmta,
Uilcll, IHo nf whlth lire 'louUecon
loured. ui<] oiic ullhcniiuoiuoiulnnu.]
KuiVE-rtMS niuiitVruTioXEavKor KiiiKii. aitib
ACTIOS or N'mttTi or Sunm.
■L Hint Conani by ihlokonvd ntamtnnv of tehwu*
(uoda Of tuttnltfj. •», whiic iidI»iw)c« or sehwim
rtndKKd iwnrraniK liy flrmrta. rp. Crliiiil«t4Sli,
mIiIcIi JuiiabciVi.-iiidlnliiH' tbolvnl nC tboaimularM*-
•tnciliiii ]<r<-ii«tiu tliu tint* n( Fiuiiibuuid ]
im..».
IS"'
."f
the primitive nhealii, ur newiletnma.^ Lving in the axiit i)f tbiii Hbeflth, nnd
wmetimai prnjeciinx fur ADnii? <]i»lnn(v innn the torn end cifu fibre, uhctbt^r
the vboalli b« diHiifuyed or no, niiiy, in JNiniv ciimoi, bu «ocii u dim, nr very
luiiilly, gmuulRr hitncl or tiiriMid, uboutoiie-thinl or bnlf the diviivicr of tb*,
fibre ; ibis is the .inVry/ifftr (Fig- 39] ; it Wcom««
](»t lo view n:> *Fc trace it bftck to wboro the tibr« a*-
Eiinies n itoublv cotilour. This nxi»-cyliuder euio!
rcAdily with ordinary stiiining rvaeents, aod beiof; ia
this and in ulhtr respects itUied in uature la the
col] -substance of a leucocyte or lo the muscle-sub-
slaiicc or a miisciilsr libre, hoa often been spoken of
ne pi'otop)iisnilc.
LyioK about Cbo torn ends of the Hbrea mar be soea
drops or miiiule irrt-gul«r niawtt rumiirkiible for ex*
bibiiinK' a double contour Hkv that of the uerve-fibni
itself; and indeed drunt of ihi» double oo»toun-d sub-
stance niiiy tie *ecii i:>«uiiif: from the torn <<nds of
ihu fihn* [we Fig. .^"]. Treated with oeniic acid
thew dn>p« and innsaea are tltiiiK-il hlui^; they net as powerful reducing
mgeuts, and the rvduced o«niiuai givct the black color. Treated with ether
1 TMi Irnrd ti fi> lorrr^frii it wit u- ili-ri<iir ilie ooTtiiocEJvc iIhiic iliHtli wnppins round 1h« vbo^
nervr. Iikitti'. '. to iiHi»:.i>iirh unnloiiiiu umuiwurmli'niinaini) nmrl-
lODinuifiit lU' 1. aiialiiay.aml l»'ii-i; iiviirlltinau Unonnxd (brthalnui of
UKnnruuMii . 1. 1 ^ . i^ lo Ibe Mnxil«iusui lu miUEic, til., Uw ibcuU of llw Stuc-
buQiuM or SrarcTnax i^r
UlM'lUTKII linviM-iiui*,
I. !<i«nlclnlDB nr ■tintli
ofMivMitt. £. Mnlullaty
■htaUl. ^ AilMTUndcrl
I
CnA?faE5 JX A MDSOLt DCBTSO COSTnACTtOX. 121
or oUmt aolreoiB of fal tbejF moreoTer tnoiv or less rHidiiT diswrlve. Olf
viontljr Ik*; «» Urgeir conipcwd of fat. and we aliall ee« t'tiat the fat com-
ptmuta tbeoi it of a very eomplex itature. Now a itei^'e-Kbiv ahowJDg a
double contour Mains black with u>n)ic acvi ; but the MainJoi.' is abeent or
T«rT tlt);)ii wlter« th« doubb couioiir eeibiM tw at u turn end nr at the Dodm
vf kanrier: llie axidojrlinder utainii wry slighily iiidei'd with uamic acid
•ad the abealh hanlly at all. Su, nUa, whi-u a trannvfrM: Mii-tion la made
ihrounfaa ttfrvt or aiwrvvoonl.each librc nppmra i» K-cliooaou dark black
riog MrroundiDg a niui-h nwre fiiinllj rtninccl ceoErnl nrctt. Furlhrr. when
a (loabl* cualoiirvd nrrvr-libn- i* tn'^tiwl with clhrr. or other tolvciiln nf t'lit.
tbrfluobUconloiit vntii«h«^«. and tht- whole tihrc become* more lritnit|)(in-»l :
Mnt\ if lurh a tibrv, either before or aft" r the trcnimctitwith ether, U'Miiincd
■ itb oarmtne or oth«r dye. the axis-<-y1iiK)er will hv seen nt n nninnl Itaiid
or ihmd lying in the axis of n tulHitar upace dolincd by the tinarilemmn
wbicb atain* only sliKbllj except at and around the oiiclci. nhich. iis we
hse* atan. an embodued in it at inicrvnl». In tlie entire lihro the lubiilnr
MBcw between tbeaxM-cylindcr and the slieaih is filled nilh a fnltr material,
lae mrtluUa, wbirh. Tram its fatty nature, bm Huch n refritctive power ns to
iliit a ilonble contour when aeeo with Iranaiuilted ltf[bt. on which account
6tirr ii«elfhn» a double oonlour. It islhi* refractive power of t lie medulla
b Kirea to a nerve-fibre and atill more ao to a biiuale of i>en,'e-fibre« or
ta a wkoJe nerve a characteriMic opaque white color when viewed bv rellecleil
At we shall we, all nerve-fibre* do not iioeoew a medulla, an<l hence euch
■ Hire aa we are describing it cadled a meaiti/aUd fibre.
X iJP***^ luedullaied fibre conaitt*, then, of the fullowinK parte:
1. tie a/w-4'yit4'/rr, a central cylindriciil oofvof eo-oallen "prolopiaaraic"
■U«ria), (Mime in nature, and readily utidctKoing diange, soroeliinm swel-
lia|««l.^>ni>'iim<««hrink!iit;. and hence in various fpecinien^ appearing now
vilkick baud, iiuw as a ihin Mreak in the axis of the tubular gheath. and
cMig fal enm McUoq aonetinwe a circular, somctiines an oval, and not un-
nqimdjr a quite irre^lar outline. Probably in a perfectly natural con-
mi it oecapiee about one-half the diameter of the nerve, but even ile
auii] mte varies in different nerve-flbreo. Witen teen <|uiie fresh it has
■iaply a dim cloudy, or, at most, a faintly Kranular apjiearance : under the
■Mtnoeof resK^nta it ia apt to become fihrillated lonffitudiually, and has
^naiippoaed to be in reality <^(>IU)MM7d of a uuniber of delicate longitudinal
ftrilbe united by an interflbrillur tulMtance. but thii is not certain. It b
knhn Mid to be prote«l«d on ii> ciiii>ide by a truuxpareui sheath, the axis-
rrliniler theath. but tliis alf) in divputnl.
Tbt axi»-cyliti>k'r jimmisi uubmken thrmiiih luiccewivc nodes of Itanvier,
lh»r>«ttrictKHiof the node nut afTectiuK it othcrwiM) than iierbajM to narrow
il Now the fibiesof a Npiiuil nerve I omitting for the pmeut the libres
■■in frutn the *yni|)«ith«-t)c nerves) may be trni-ed bock «ithvr to the spinal
pmHno on the |iu>(f rior root, or along the anterior riKit t» the anterior
WMa of the •(linal con) ; nnd.nn weaball aeo, the uxi«-cyHndcr« nf the fibm
m, ia butli <■«<«■»'. prulongaiions oi" procrasee of nerv«-cclli', in the former
MHufoelUof ih<- ganglion, in the latter ciwe of cells of the anterior ooraua.
b ikIi ease a DnxHvi of a cell W-oming the axii-crlindcr of a nerve-fibre
nm an unbroKi-n coiirM^ pasaos as n continuous band of [leculiar living
MUrr, through node uflcr node right down to tho termination of the fibre tu
Ur iMcie in which the fibre ends ; tlie only obvious change which it under-
^ il that, in many if not nil c:k<e«, it divides near its lerniiRatioii in ibo
taaw^ and in eoinc canes the divisions are numerous, and jo\a or anaMtoraoaa
ftMlf- Obviously the axia-cylinder ti the easential |>art of the nerve-fibi^
122
THE CO.VTRACTILK TISSCK3.
'2. Th« primitive ^tvath or tKurilemma. a tubular jtheaih of tnuupnreiiL
appareatljr bomogeooou* niiitoml, not unliki- that iirnmrculeniniK iu iiitltirv.
At eavh ni)dc the DvuriloiDinti i« ccitixtrictivl »o m lo ■■lubrtoc th« iixu-cylin-
<)er cIomIj-, but is nt the vune tirno thickrnivl by »nmc kind of ocnic&t luate-
riiil. StiiinitiB rca^iibi. wpeciiilly silver nitrntc, nppcnr lo enter tbc nerve
libru fnim without more n-Atlily al a nude than elsewhere, itnining lb«- libre
tiutH n( ihe Dode, and creepio); upward nnd downwnn) from the ixxli; iilimg
tbc axis-cjlind«r ; hence il hns beeu ituppirised that the uiitritive flui<l. ibe
lymph, oncora into the fibn i: ! - < (^ts access to the 8St§>eTlini[er more
rcttiiily Rt tlie nodes than >!- i-.h- AImuI midnay lielween every two
nodes is plac«<l a Ion)* oval mn {■ n-. <iti the imide of the ueurilentran, pushing
the medulla, aa il wen;, inminl. ual so lyiiiK in a shallow bay of that siih-
Slsoce. Immediately surroundiu); the nucleui> h u thin layer of t;ranular
subetanoe of the kind which we havo ii[H}ken uf aa undifTerentiated proto-
plasm : in young newly fornKil librc at all eveiiU and iKMnibly in all fibres
a very lliin lny<ir of tliis aamc 8ub«lance lit continued all over the aegment
between tbo iio(Uv, on the inner snrfacc of tlic nenrili'mma between it aud
the medulla.
3. TV nifjUtUa. This ia a hollow cylinder of fatty material of a peculiar
nature filling alt the space between the neurilemma on the outside and the
axis-cylinder within, and suddenly ccasin^^ nl each node. It thus fomn a
close htting hollow Jacket for the asis-eyliuder between every two nodes.
The fatty malerial is lluid. nt least at the temperature of Ihe body, but
appears (o be held in its place as il were by a network of a stibstano- called
neurokenMlin, allied to the siibtttanco herutia, which is the basis of the bomj
scales of the epidermis and of other horny structures ; this network i* most
marked toward the outaiile of the mediilln.
8o long aa ibe nerre in in a frenh living, perfectly normal condition, the
medulla appcnrs smooth and continuous, showing no mark* l>eyund the double
contour; but in nerves removed from the body forcxaminntton (and accord-
ing to SDuio observere, at times in nerves stifl within the bmly) clefts make
iheir appeHrance in the medulla running obli'jiioly innan) Irom lJi« neuri-
lemma to the axis-cylinder, and freanently splitting up the medulla in lueh
a way that it appears to lie composeo of a number of hollow con«s [Hirtially
slid one over the other along the axis-cylinder. These clefts are spoken gf
as indealationM, At a later stage of alteration the medulla may divide into
a number of small irrei-iilar ina^eti separated by floid ; and since ea«'h small
pieee thus Hoparatt-d haa a double contmir. like a drop of medulla exnded
from the euil nf a Ittire, the whole tihre baa au irregular " curdy" appearance.
The esaeutiai pun tbuii i>f n medullateil nerve fibre (of a spinal nerve) is
the axis-cylindiT, which is really n ^irobugaliou of a procea* fh>m a nerve
cell in a apiual eunglion or in the spinal mnl, running au uubrokeai course
through ncidi: nuer node, never in it» cour*i% a» far as we know, joining
aoother axis-cylinder nnd very rarely dividing until it upproaeties its end,
where it may divide freely, the diviBions in sunn; casej" annstomoaiui; freely.
We may conclude, and al I we know sup|iurU the concltisioo, that the changee,
making up what we have called a nervous impulse, take place, primarily aud
chiolly nt all events, in this essential part of the nerve fibre, tlH-axis-cvlinder.
The ucurilemma and medulla together form a wrapping fbr the nourishment
and protection of the axis-cylinder, tJie fatty medulla probably serving
partly as pn>]>ared ftiod for the ax 'is-cy Under, partly as a mechanical sup-
port; possibly it may also play a part aa an insulator iu tlie electric phe-
nomena.
It is eamy moreover to see tbnt while the axis-cylinder along its whole
Icugth is practically t^whatever \nt the exact manner of its funualion in the
CUASG83 i:? X 311'aCLE DURING CONTRACTION. 123
vo> m i»n of the eall of whii^ it is ma vlongUwl pfocnii mch nocment
ttrn txrre two nodes rrprcwnls a cell wrnpping rouiid the iuct*crTiii(liir
rtiw, i>r wfiicb D)-ll the nucleii* brtwcvu thv tradu is the ODcIeiu, llio n«u-
I riWmiD* ihr mivcIom or cell irslt, and (though this a mrhupe not ijuitu to
\damr) the iiifdHlU iW c«ll suMmiice Iu^It converted into tatty tnaleria), a
n*ll in tut wbiob it raiUf uiitfide the axis-cyliiidcr or nerve libiv pro|>er.
It u alaoy the ftxi»«yUod«r that the uervoiis impul^ei sneep. and each wrap-
pinjt rrll only serves to noumb and protect the seumeut of the axuKtylinder
bp|<B««v iia two nodea. And we acrordinftly 6»d that both Mt the beginniug
nf the aerre 6bK in the ganKlion '«n or ^pianl cord, and at its end in the
. h<iUi neurilemuia and medulla dlsajijienr, the axi^cylinder only heiujt
hA.
A nem soing to a muael« is chJvtly (x)tii|iuiH!d of iiiedii Haled Gbrw aa juiit
HIhmI. the uinjority at which, i-:idiug in cud-plnl«s in the rauaculnr tlbres,
atv the fibres which cuoducl thi* nefvouN iiii|MilwM to the mtiitcle, cniuin;; it
to eoauact. aiul ma^ haDn be»pokcii of a» awtor nerve fibrcH. Some of the
Cbttv however cu<l tn other partx, such n» the lendoD, or the ci>iiiiectivc Um-
•os between the bundlw. and mme in tbo bloodvcstels. There are rcnsoni*
for thinking that lonio of tbvef ccinvey impiil«s from the muscle to tho ccn>
timl aerrous system and an consequently Rpokcn of as sensory or afffrenl
ftbns. conoeming tboee connected with the bloodvoMels ne sball speak io
iMiu with the rsseuiar ayaUm.
t tm. \errr-enitinyt in ttrialfd mttmitar /ibrtt. A nerve on eoieiiDg a
BMcle divides into n number of hrancfaee which, runnins in the conneeUve
IMW of the muscle, form a plesua rotmd the bundles of muscle 6bres, tlie
tsaller branches forming; a plexus round lite muscle fibrw ihemselves.
Fma this plesua are given off a nund>er of nerve fibres, runnini; ainj^ly,
wkef which Joinings niiude fibre eiiilf in an cnd-pl&tv. In furniinj^ these
[hniH ihr irxlividiiiil nt-rvi' libra* divide rejHTiili'dly, the diviiii»n ulwny*
tikfait plofv at a niHt<! of Kanvier, so lliul wlial in it xinglct nerv*-' tihn: iii> tuc
SSTintrn the muK-lc may give rise to sovomi tu-rvo lihnv ending in wve-
hImbmIc fibm. Tlie tier\'c fibre joins the niiiscto fibre n*- about its middle
trniaewfaat nearer one end, and occasionally two nerve librM may join one
nH* fibn- and form two efld-plales. The gonvml distribution of the bun-
fc <if n«rve fibtvs and single nerve fibres is eueh that some portion of the
■Mtin is left fVee frufl) nerve 6bres; thus at the lower and at the upper end
■'lb Mrtorins of the frog there is a portion of muscle quite tree from tterve
X mi)[k nerve fibre, ninniuf; by itself, has, outside (be neurilemma an
iHiiuiwil delicate «heath of Rn« connective tissue known as UenW* Aet^h.
sUcfc nnpeara to be a continuation of the connective tisuie forming the
iksih (if ike nerve branch from which the fibre spraii||C, or uniting the fibres
tvohcr in the branch.
The actual ending of llie nerve fibre in the muscle fibre dtflbrs in different
dHBW of animals.
la nammals and some Dth<>r animals the nngle nerve fibre Joins the nius*
difbn in a iwelUng or projectiufi having a more or Icat oval btuic, and
rwtng wliea seea iid«mys as a low conical or nmndeil cminenoc. At
Minmit of this eninaoce the nerve fibre Wot both iu slivuth of Ueiile
nk il» neurilemma, ono or other or Imth (for on this point observers do Dot
•PfeJ becoming continuous with the sarrolemma of the muscle fibre. At
tb wnaiBit of the eminence, where the shcutlis fuse, ihe fibre, now coiwiattng
«ly uf nxi«-cytindcr and medulla, loees its medulla abruptly < in Ihe mus-
chs of the lon^'ue the nerve' fibre in many caeca Uxtee its medulla at sotne
qaiiderahle distance before it joins the muscle fibre to form the end-plnte).
1S4
TKE CONTRACTILE T1S3LE8.
while llic axin-eyliTiiler liranc-fac* out in all dir«rtiom, lite Monwivliiil vni
briiiiclitv, whicli wiini^liiiKW niiiivtonxBU!, forming a li>w roniuHl niiMi, tr. _
whiMi viewbi fruiu abnvn hiui nn BrhorcMci'-tit »r labyrinthinr niiiM-nrann:.
Ou ilw brnDchm of this aborwccnco mny lie unc or more somcwhai sriinulBr
ftvnl" iiiiclci. The nborcsccncc ilsclC hns. like the nxw-cylindcr of which it b
II lUvciopmeut, a vita* taiotlv granulsr or cloudy anpenninoc, hut lying hv-
twwti it niid the iictiial muscle aubstftticB is n disc or bed of »om«Kh4il
coaraely granular inBterial, called the lole of the end-plate, on which lh«
romilied arbcire«ceiit axi^cylinder rests, more or leas overlapping it at the
edge, but with which it appears not to be actually continuous. Lying in
the iiiidat of this " sole " are a uuniber of dear oval transparent nuclei.
The end-plate theo beaoaib the sarcolemma consUta of two parU, the rami-
Bed axis-cylinder, and the granular nucleated a»le, the two apgmreutly,
thmigh iu juxlapMition. imt heiDg ■.■»ntiniiini#. Accunllngtu t>onie obsen'en
the Vile In conliuuoii^ with and indeeil is a sjiecinliztHl part nf that substance
pervadins the whnl« muitnilnr fibre which we spoke of nn iutnrnbrillar sub-
ttatic& W« cnnnol eiiliT here into a diMCUwion of the jtrolinblc mfniti»){ auil
UM of tb(«(' iilnictiirce or how ihry cfTect what xccm* obviouKly their Itinc*
lion, ihe trani> format ion of the chanpw constituting u nervoun impulw into
the changes, which running along the muscle libro in the latent period a*
fur^runners of the changes cnlniliDg actual contraction, may bu »nokvn of as
cou>liluting a muscle impulse. It is of inlereet to obwrvc ttiat orrtain
analogies may bo drawn between an end-plnle and the histological elen»cnt>
of the so-called electrical organs of certain animals. The element of the
electric organ of the torpedo, for instance, may be regarded as a muscle
fibre in which the nerve ending haa become highly developed, while the
muscle substance has been arrested in ita development and lia-t not become
striated.
In amphibia (f. ;/., in fVogs) the ending nf a ner\-e fibre in a inttacle fibre
b Mntro'hnt ililTcrcnl- A nerve fihrp about to end in a muicle flhr« dirtdea
into a hruch of several nerve tibrcs, r«ch of which, losing it* xheath of
Benle aod sarcolomma. enters th« same inusclo fibre, and ihcn Iiwing il* ni^
flnlU runs longitudinally along the 6brc for vome distano^, it and its branebt*
dividing several times in a chnrncteHslicnlty tbrkcil manner, and Iwafiilg U
iniervats oval nuclei. In other animnlfi fi'rms of nt-rve ending an* met with
more or lees intermediate between thiit seen in the mammal and that Mtcn in
the frog.
S 70. llesides the medullated nerve fibres described in S 6i!, thont are In
ini^l nerves going to muscles a few and in some nerves, going to other parts,
a large number of nerve fibres which do not possess a nieilulln, and hi.^nce ore
culled •ton-meilit!tateii ttl/rai : these are ce^iecially abundant in ihe so called
syin)>iaihetic nervuH.
A nou-medullated fibre which, like a medullaled llbre, may have any
diameter from '2 ^ or Icki to 20 " or more, i» practically a naked axi^-crliiider.
not coverr<l with medullu, but liearing on it.i outside at iuten'aU oval nuclei
diapoied loiiiritudinally. Theso nuclei appmr wholly analuguus lo tlie nuclei
of tlw neiiriieninia or n raedullattHi fibrv, and proliably belong to a aheaih
enclosing eacli fibre, though it is not uiwy to denion«lmt« the independent
existence of such a sheath in the case of n»o« ooii-mwlullated fibres. In
the similar fibres constituting (he olfactory' nerve a sheath in i]uitt.' oonspicti-
ous. Unlike the medullaicd fibres these non-mo()ullatod divide and alw join
freely: like them eaob may be regarded as a proecw of a nerve ocll.
Of such non-raedullated fibivs a scanty number are found in ncrvca g<Miig
to muKlee acatlered anion^ the medullate<l fibres and bound up with then
bj eonneotire tissue. They appear to have uo connection with llw muaculw
ji»^
CtlAXnES IN A IIUSCLB Pl'RIN'Q CONTRACTION.
126
fibns, bat to be dUiribuivil clii«Hv tn tbe bloudfesMl* : mkI lh« ftincliou of
wa-SMlBlliutl (ilinv hid beller m coDiidtied in coDn«*:ti<Ki iritli n«rve<i nf
vbidl tbtjr fonn » Inn;* pun. Mich m certuin nerve* goiog to bluwhwwU
aod lo Mcntil^ QrgWM. But it miiy W iiUt^il thnt though thej piwvtM DO
awdalia ibtj sr* capable of pri>|>iigiaiag uorvuiw iinpulvu* in thv sumo way
■» Ottdalklcd nurvw: uiti (bU Tact iniij be taken u inilknting tliiit llii*
Bvdalla caMDOt serve uiy vcn- important function m an vlectriv insiiUt^ir.
t 71. TA< rJiemiMry of« »enf. We havo apoken of tfa« lovilullii ne iHtiy,
and }Tt it in in reality vcrr largely compoaed of asubatanoe which ia nui )in
tbv Mriet mwe of the won!) a lat. When wocxiimiDechernicollF a 'juaniity
otttmrrt (or wbat la practically the Bame thing, a (tuauiilv uf thai jwit of tbe
nrntnU nervmia BystetD uhicb b called uiiilf mailer, and which as we shall
an i» duaflv cmnpoeed, like a nerve, of niedullaled nerves, luid is to be pre-
fcrrad far cotntcal exaininatiuo b«(.-uuK it cootainB a relatively l^InaII (|uantity
of omMCtirv tiauie), we find that a very large proportion, acoordiiiE to some
•banvn about half, of tbe dried matter «onsista of the peculiar bmly.
flAal(«(«riM. Now, cboteateriu it not a fat but an alcohol; like glycerin, faow-
•W, which i* at*u an ulcobol, it fornw ciom(Hiiiiiii> with fativ nriil*; lutd
ihMgll we do not kn<>n ilHinitrly the cbcniicnl ivniilition in which vhulvtittrin
txbla daring life in Ihc nH-diiUa. it i* iiiorr than ]ir<>l>iil>k' that it ex'au in
MM* eonbitiation with aoiae of the really latty bodic* also prownt in thv
■(daibi, and not in a Am mlatcd rtntc. It i* >iDgtibir ihat liotiiti.'v being
in Mcb large <|uanlitica in ncrvoua tisHio, and to a small extent in
Ikmca and in blood, cholcstorio is a normal constituent of biU-, and
tthagrcMer partof gall-Monea when th«wan preocol; in galUtonea it
* ttadoabudly preaent in a free suio. B«Nd«s dioleet«rio " while " nervous
aailer cuntaim a leaa but still coneiderabte t^iiauiiiy of a complex fat. wbtise
Man ia dbpnicd. According to ftumc authoritiee rather lau than half (hi*
iMplii &l matint of the peculiar body (eriihrn, vrhioli we have already
«tt la ba praMDt abu in bloutl curpusolo and in muacle. Lecithin contains
ititKliela of elaaric acid (or of oldc, or of jMliiiide octd) aMoelatcd tiot. as
iimfiaarr 6tU, with aimnlc glycerin, but witlt the more complex glycerin-
(btphuric acid, and fiirttier combined with a nitrogenous body. »riirin, an
■aawaia cumpound of some eonsidi'mhlv cumplexity; il is tbercfora of
iHarkablr nature, sinoi!. though n fnl, it conlniiis bulh nitrogen and phns-
tiana. Acoording to the laina aullioritits the rcmaindec of thr complex
■leMiiMa of another fattv body, also apparently containing nitrogKii but
hi lAosphonia, called etrtbrin. Other anthorilics regard both these bodice,
Imtkia and cerebrin, as products of decomnosition of n slill more complex
bl. failed pfwtayoM. Obvionaly the tal of the whitr mntler of the central
Mftoas syMcffl and of spinal nerves (of which fat by far the greater part
as« asi* in the a>edulla, and form nearly the whole of tJie medulla) is a
my tnmplex body indeed, apecially so if the cholesterin exists in coiubiiia-
lica «ttb the Ici-itliio, or cerebrin (.or prolacon). Being so complex il ia
sttatally very uiiMoble, and indeed, in its instability resembW j^roteiil
■BUar. Hence, prubahlr. tbe reason nhy the medulla obaogee so raindly
ad to profocindly aller llie drnth of the nerve. It Mciua. moreover, that a
cmain ihuugh otinill i|uautity of proit-id matter forme part of the medulla,
•4 )l M puwihte that iht* rxists in aoinc kind of combinalioD with tbe ouiu-
fln bt : bnt our knunleilge nn this point is inii«rfeLl.
Tba pcvcDOv in such large •(uamily I'f thi» complex futty medulla render*
iksebsnieal examinali^m of the olhvr con-ititueiitv uf a iterve very difficult,
lai «ur knowledge of the chemical nature of, and of the chemiial ehangra
pianon in the axis-cylinder, is very limited. Kxaiiiint-il undvr thi.- micnj-
w^ the axia-eyllnder gives ibe xaothopniUio reaction ami otlier iiMlicatious
ise
rHE ro.VTBACTILK TISafBS,
thai it is proteiit in nature ; beyond thu< we nre lurgelr confinetl lo inf
W« infer that ile eheniical niilLire im in a ^n^ral way ^milur to ibai of
oell Klbetonoe of th« uerre <.-ell of irhicli it is a proceai. \\\- iufi^r llial th»
chemical nature of the cell imbalance ut' a nerve all, heingof lh<f kind ithifili
iit fr«i]ueally called " prutoptasmic." is, in a general way, mtnilnr lu that of
other "proloplouuic" eelU, fi>r tiialiiuue of a leucocyb:. Now wbcrt^ wo rao
rxnmint: convenientir such celli we find, ati we hare uid ID $ -lO, thv nrotvidt
pmeiit in them to nc M>ine form of albumin, m>me form of elobulin, aod
either niyonii itiwif, or niiit- wdiTnix of myoBiii. or maic allied boav. Id otli«r
words, tbo prolcid hiwiii of tlie kind of cell aubitlaiicc which is frmguently
spokeD of as " imd>t)Vrvntiat«id [>roloplii»m," does not. ia its broad feature*,
differ inntcrinlly from the pnitvid tia«is of thut " diftcrcnliatcd proioplasm "
whii'li we linvi- cnllcd tmimile siihstaniT. Hpdcc we infer ibnt in their broad
chonii<^'aI featiireii the axis cylinder of a nrrvo librc and the cell iKxIy of a
nerve cell resemble the substanre of n mmcic fibre; and this view b sup-
ported by the fact that both )cr«»lin and lactic acid are present as "e:x-
traotivee," certainly in the central nervous system, attd pronably in nerrea.
The reaeniblancc is, of conrvo, only a funeral one; there must be ditloreiicca
in eberoieal nature between the asi^eylitider which propagates a iiervotu
impulse without cliange of outward liirm, and the muscle 6bre which oon*
liaela; but we eanuot at prcDeiit aiate exactly what these difference* really
are.
After tlie fats of the medulla (and the much amaller quautltv of bt
[iroent in the axiit-eyliuder), the protelds of the axit-eylinder, and ifie oibei
soluble subatuDcea preiienl in one or the other, or gathered round the nuclei
oftfae neurilemma, have by variouH inctuia beendisolveduutof a ni^rvvlibre,
certain substances still remain. Oueof tb<fleinsaiaIlc|Liiuit)tyisthe nuclein
of the nuclei ; another in larger r|uaiility '» ihe substance nnmJ)emtiti which
forms, as wc have sovii, a supporting framework for the medulla, and whose
moat marke<l charact«riitic is, perhnpt, il4 mixtimee to iolution.
In the ash of nerves there is a prepcinderaiico of potaasiuiu salts and phot-
phatei», but not so marked ne in the case of muscle.
Ji 72. 7'/te nermu* impwlte. The chemical analogy between the aubstauoa
of the mtieole and that of the axis-cylinder wuulil naturally lead up to (tip-
poae that the progrew of a nervoua impulse along a nerre fibre was
acooinjianieil by chemical cbungei^ similar to those taking place in a iniiad*
fibre. Whatever changes, however, do or tuny take place are too slight to
be recuguiy«(l by the means ut our (li»p<)e>iil. We have no satfafactory
evidence thut in a nen'e oven n.-pe'jited nervous impuKii can gi\-e rise lo an
lU'id reaction, or that the dt-alh of a nerve libn; leuda to such a reaction.
The ifniy matter of the central nerv><>us -^ystem.it is true, is said to be slightly
ucid during life and to tieconie ni»n- acid oAer death ; but in this gray matter,
nerro cells are relatively abundant; the white matter, compoeed chiefly of
nerre fibres, is and remains, during action as well as rat, and even after
ikiatb. neutral or ulighily alkaline.
Nor have we iiali* factory evidence that the progras of a nervous Impuba
is accompanied by any setting free of energy in the form of heat.
In fact, l>eyond the terminal results, such as a muscular contraction in the
case of a nerve going to a muscle, or soma affection of the central uervoiit
system in the cose of a nerve still in connection with its nervous centre,
Uier« is one event and one event only which we are able lo reoogiiixe as the
objective token of a nervous impulse, and that is an electric change. For a
piece of nerve removed from the body exhibits nearly the same electric
pbsDOTOCoa as a pieoe of muscle. It baa an equator whidi is electrically
pMgliv« rslaliTcly to the two cut ends- lu fact, the dJagnm Fig. 84, and
ODASOKfl IN X UVHVUK Dl'BINO CONTRACTION. 12"
ilk* dnoiptioD which waa siveD in $ 6<1 of tbe el«ctrio change* in mtucle
mmj Ui applied bIrkmI u well to a nerve, except that the currents are in all
cBMt anirb roun- (evMv In the caie of nerrea thanofmiMcleB, and the ipecial
mrrvata IVani tlm circatattreaix to the centre i>f the iransvene sections
cBaout wll In' nhoKU iii a sletider nen'e; indceil, it is ilouhifiil if ihev exist
at a!i.
During thv iMfMgo of a uervouit impubu the "iiuiunil iierro-vurreot"
I nrinUivo varintitiii, junl a* llw " iintumi niuwle iiurri-nl " iiiidor-
■ a nigatiii variali"ii iluriii^ a cuotrnction. Thom iirc, moreover, naMnii
Um cwa of the DUtc, an in iht raMi of tin- inunclr, which li^ul uk to douht
lb« prafxIstenM uf any «uch " imturnl " currcntx. A ntrvo in nn nlwoliicvly
lutarml niaditinti ap|>mrF to be. like a muflclc, iwKlvclHc ; hoiicr ne may say
that is a anxr (hiring llic jinwaige of a nervous impulso, as in a muscle
dsrina a muK-ular contmclion, a " ciirrcDt of action " ia developed.
^'tw " current of aclioo " or " itcgnlivc vnnation " may be shoim either
bv tht icalvaiHMnetcr or br the rhewcopic frog. If the nerve of I lie " muscle
avm pmMralioD," Jl (tee § ti7> be placed in an npproprinte manner (m a
llM*«aglify irrilalileuerve, J (to shicn. of course, DO luuscte Deed t>e aiuiched),
M»eUag for instance the equator and one end of ilie nerve, tlien siueU
tadwtioo-aliocks sent into the ftr end i-f A will cause sioffle Hpiutuia in llic
tmsrlf of //, whil<r leiniiixation of A, i.e., rapidly ni|>oated thocks »ent into
A. will cause ti-uiiiin of the muscle of B.
Hiat lilt* <iirr«iii, nhdhrr it be regarded ait au iudc)i«Hdeiil " current of
Mliso ** or as a iiejjativi' vBrmli<in nf a" prMxisting" current, ts nn nwrntial
latiirr of a nrrvoiu inipulM; i!< »lion'n by the fact that the degree or intensitv
tf Ih* one vnri<-» milh iliut of the other. They l»olh truvcl, loii. at the Htine
an. In da>cribing tlifl muscle -curve, ami the niclho<l of measuring the
■SK»br lalmt [wnod. ire have incidcDlally shown (^ ■lli) bow at the aame
tas the velocity of the oervoue impulse may be measured, and staled (hat
lis rale ill tbe nerves of a fru); b about IJ^S meters per second. By means of a
•pnil snd Mrocnhat complicated apparatus it is ascertained that tlte current
■f (rtiiai travels aIoD)( au iwlatc^l piece of nerve at the same rate. It also.
likrtW nwleciilar choose in a muscle precedioK the coutntcuon, and indfed
iW conlradion itself, traveb in the form of u wave, risiiifc rupidlv to a
am at each [loiiit uf the nerve and then morc)j;rHdiiiilly declininB
Tlw length of the wave may by speciul mcjiiis be measured, and is
laul III br about IM mm.
ITIvn an irtilaied piece of itcn-e i* .^timuluied iu the middle, lb? oiirrcnl
ittiti'a i* |>n>|>af(aied (Kfually ncll in liulh ilinictioiia, and that whether the
r Ilea cliieflyaBDMnror a cliiefly motor nerve, or iiide<-d if il he a ncrve-
OQUpaised excloiivwy of motor or ofuauury (ibrcn. Tnkiii); the curroiK
«f acftinn as the lok(.'n of a iirrvous impulxc, we infer from this that when n
flbr« ia BtiniuUti'd artiticially at any juirt of il« course, the nervous
going Irnvvls in both direction)^
naeil jost DO«r the phrase " tclanizntion of a nerve," meaning the
^pUaAioD to a n^Tve of rapidly repeated shocks nieli as ironid pr<iduc«
tAtmt in tbe muscle to which the nerve was attachiil, and we shall have
hysiit nccasioD to employ the phrase. It must, bowcver, be anderstood
Am tWv ia in tbe nerve, in an ordina^ way, no summation of nervoua
eiim|)antble to the Himmation of muscular contractions. Putting
(vrtain enseo which we cannot discuss here, we may say that the series
rfifcuks *mit in at the far eiwi of the nerve start a scrieH of impulses ; these
ttntl down the nerve and reach (he muscle as a series of distinct impuWa ;
«4 iW tint change* iu (he niiifcle. (he molecular la(en(-period chan^, also
Itm A acrirs ibe memlwre of which are distinct. It is not unul these
sdktvrt
128
THE L*ONTBACTIt,E TISSt'BB.
molecular (.■liaii;:ea bec»rae irBnaforroed into visible changes of fiina that tmy
flisioD or eiimiuultoi) tnkee place.
$73. ruttiiiK [i)|:«ilier Lbe facts oontaiii^l in lIiU anrl tlie jin'CHltDg *ee-
Umu, the fiillim ii];; luuy be taken as u brief n|]|>foximnl<: liWlonF- at what
l»kc« place in a niuMte and oervc when itie Inner t* lubjected U> n fiiivle
iiMliiction-aliuc'k. At the intitaiii tbtu tbe induwd current immki' inu> tne
□ervG. cliKiigvn uociir. of wbo«e natiin; wc know nolliiiig corlam, oxa-ii* that
they cnu»c n " current of action " or '■ negative variation " <jf the " natural "
nerve current. Thwc clmngw profiHgiitc llKrinwlvra along iIh' ncrxT in hoth
(Iirv<ctions as n ncrvnti» impulou in the form of a wave, haviug a wavv-len^b
of about 18 mm., and a velocity (in frog'« nerve) of about 'la m. per MocmcL
I'lUNng down the ucrve fibres to the niM«clc, flowing along lbe branching
and narrowing tracts, iho wave at last breaks on the end-ptatcs of the fibree
of tbe muscle. Here it is lriiusntut«d into what we may call a nmsde
impnlae, with a shorter, Elcejwr wnve, and a greatly dimintfhed velocity
(aooui 3 ni. pier secoDd). Thii) muscle impulse, of which we know hardiT
tuone than that it is nmrked by a current of action, travela fraiu each ena-
plate. in both itireciions, to the eiul of the tibre. where it appears to be Iom ;
at all eveuta, we do not know wliat becomee of it. As this impulse wave,
whose develupmeni lak<« plac« eulirely within the latent period, leaves the
end-plate, it i-> fnlKiwed by an explosive deciim position «f material, )c«dtug
to a iliHchar;^' "f carbonic acid, to tbe apgwaranoe of soom subcitaur« or
iiul)«tJUic<3t with an acid reaction, and probably of other unknown thi
with a ciin.-iideriihle development of heal. This ex]>loitive deoompoaitioii g['
rise to the vLtiblc contraction wave, which travebi behind the inviiiible m'
de impube at about the tame rate, but with a vastly increatcd wave-length.
Tti« fibre a» the wai.'e paaMs over it, awells and diorteoa, and ibiu brings its
two nwit nMtn>r together.
When repented shock; are given, wave followo wave of nervous impulse,
ninscle impulse, and visible coniraclii>n : but the last do not knep dislinct ;
they are fused into the coutinued shortening which we call tetanus,
The Nature of the Chashbs Throitoh which as ELixTRir CV brext
IB Able to Uexrbats a Nervoi» iHruL^K.
.licfion of the CoixMatU OunviU.
i 74. In the preceding account, the slimuhi» applied in order to give ii»e
to a nervous inipiiUt! hib> always been siippi^'<l to he an induction-* hock,
single or rcpeate<l. Thi* choice of slimuliu hns been made on accoiinl of
the nimoft momentary duration of the induced cnrront. Had wc used a
current lasting tor some considerable time, the problems before u« would
have become more complex, in consequenoc of our having to distinguish
between the eveiiis taking place while the current was passing through the
nerve from those which occurred at the monient when the current WM
thrown into the nerve or nt the motnent when it wns shut ofl' from the nerve.
Ttiwe complications do arise when, instead of employing the induced current
aa a stimulus, we use n eoiitlaul enrveni, I.e., wheu we paas thn>ugh the nerve
(or muscle) a current direct from the Italtery without the intervention of
any iuduelion-cQil.
Ilefore making tlie actual experiment, we might, perhaps, naturally >un-
]>o)e thiit the cou.-itant current wotild act an a Mimulus ihroughoul the wltole
lime during which it wan applied ; that, ho lonv a* the current passed along
Iho iwrve, nervous imnulitvti u»uli) be gcni-nili.-*r; and tJnU thwe would throw
lh« musde into sometliing, at all event-'*, like telunu*. And, under certain
S'TIJHTl.rS BT BLECTRTC Cl'RRBNT.
129
I. thu >lix« Ink*- |iltw«; ocoukiuiillv il does liii|i]>ai) Oiiii at tlie
b« ctimiil ia itimwti into (Im nerve IIm: iiiiux'li; uf the iiiui>cli'- nerve
p*«|)*nui«i fall* iiiii) n teuinii*. which u c<jui»ii>mJ nmil the ctirn-iii is ihut
vt; but •uch > nwilt U cxivpliiiimt. In l)i« vnsl luitjoritir of cn*es wh&t
'"rr^"' ^ ** fvllovrs: At the mooieut ilmi ih<- drciiii h raadv. tko mo-
n«U tbU tlio current if thnmn into iIh' urrvn, n tiaelij twiu'b. n >^m[i)e
ooMnctina. tbi- k^-cxIImI auikiny fonlmeiion, io witniwciT; but uflcr tbU ling
fw I sasr the muM-l« remaiiiA nltmlulrly iiiiiwcoDt, in ipile of iW current
diotinuinc topsM ibr^iugh the ncne, ami lhi« fjuiSManctH miiinlnined until
Um circuit t^ broken, until ibc current ie shut utT fnna the nert'e, wii«D
■Dolhrr Hnipl« cituintctioD, tlie so-called tirmHny mntmrlion, n observed.
TIm nrrv pHwaffe of a conatajit curnnl of unironn inlciuity ihn^n^h a nerve
4o» DM, uodvr urdinarr eircumslancee. net as n Btinmlus (•eneraliuji; ii M«rv-
utM ini|Mib*: Mick ttoimpukeitfODly set up when the current either falls iaio
ur b •but olT frxiin the nerve, ft is tli« entrance or liie exit of the current,
aad Ddt the continuKDce of tli« current, which is the Htimuliia. The •|uii»-
MOW uf ib« nerve uid mu»cle during tlie pawaj^ of the curreut i», howercr,
ifarwiiitm on the current. rtminininv unifunu in intenAily, or, at letui. not
Inn^ MtddrolT increu«d or diminiiibMl. Any mflicieiitJy middeu and Inree
iMMM or diniiautioD of itir inU-iiMij uf 'i)ie curreut will act like t£a
OMfsof* or nit of a ctirront. and by j^ncratinK n nervous impuljte give rise
H a cKMiinu-tion. If tW inlc4uil^ at the cnrrwit, however, be very tduwly
■ad ^Tvdnnlly incmi»ed or dimioiiihi-d, a very wide rutgv of iuienoily may
W (awvil ihniiigh willMdl any c»ntrac(ion lidog mcji. It » the xudden
tlufD froa MM) oooditioD to aiiirther. aoil not Uie condition it(«lf. which
laaai Uw nervous impnlM.
Id manr ooMi, bolli a " making" ami n "breaking" contradion, each a
uplc twiteh, ar« oWrvol, au'I Um ift, perbugu, the commoDeet event ; bnt
*W ibe current ia very we«k, and again when the current U very strong,
dibr tbe brmkinz »r the miikiuj^ iHniiraction may be absent; t. e,, there
Mf b« a ouniraciion unly when lh« i-tirreut is thrown into the nerve, or
•■rr afaan il i> «Uut »S fnmi the nerve.
tnikr onlinarv cirounLiiauccii the omtriictixiis wituemetl wtlh tha eOD-
Nai current rilfier at the make or nl the break, are of tbe natnro of a
'4mfit " oontractioo : but. m lia« alreiuly be«-n naid, the ap|dic«lioti of the
mmi BM/ giro rUc to a very pronouucnJ letanud. Huch a t«UinuB ia seen
■WliBHa when tiM current >» tnw<h; Infttii^' during the application of llie
wm, aometiaMi when tlw nirrvnt i* broken, laMing »onic time after tbe
(Rent baa been wholly removnl fmni tbe nerves The former i» apokeoi of
■ t " Making." th« lattt^ra* a "bronking" lelaniu. liitt ihote exceptional
naki of the application of thf onnniant current nwtl »ol <tetaiii u« now.
n* graal inicreal «tta>-hctl to the action i>t' the consl^uit current lies in
At 6m that, during the pgifnijn* of the vurreiil. in «pile of the abwn<-e of
ll avTOOB inipulc«>. ami llicr^Tore of all muscular c<inlraclionB. ihe nerve
kfcrlba line both bclni.m and on each "idc of liie electrodes priifitundly
mtiti la a owat peculiar manner. This mtHliticalioii, important bi>lb for
A* %fc» h throws on the genenilion of n«n,-i>u» impuisM and for its pravli-
•I aBpSortlona, ia kmiwu under the name of <Jr<tivt<inu*.
\ilC fitarfrotontw.— The markeil rcalur« of tbe riectrotonic condition b
llai tlta oerve, though apparently 'luiewent, is chanct-d in reaped to itA
BrittUity: and tlui iu a dittereut way in the nrighborfaood of the two
ihdnaW raspeetively.
S»ffimm that on tlie iwrve of a muscle-uerve prepanilion are plai^ two
1— [ailatiiatiliij ctecln>dea (Fig. 40, u, t), connected with a battery and
>) with a key, to that a constant current can at pleasure be Uirown
-JlIE I'DNTBAC'Tll-E TI8SCBB.
into or shut off from the nerve. TliiH M>ii«(aui c-urivnt. w)iii»e efleci> u-(
■bout to stiiily. may be c»lle«l the " |i»lariiin}; ctirix^iit." L<?t n \n: the puct
live ekctroilo or anode, aad k lite nejraiive doclriKie or knlbiMk-, both jiliiccd
St H>nie dislaDCo from the ii)usi-le. iitid nlito yrhh n cerliiiii inU'-r«-Bl Ih-Mvcmi
cftch other. At lh« jMJiit x lec lher>e be npplk-d n piiir of vWlr\>d<»
Dectod witti iiD iiKluL'Uui) eoil. I^et the tiiiiw^lt' I'lirihnr hv coniicrtnl wi _
lever, M that ila i>iiiiimc(ionii tun In; ret-orckil iim) tboir mnouiit mra»iin-d.
Before the |i»lnriziii|c current in ihroMii inln the i)or\-t^, let k niil'Ic in<bicti<>D-
(iinck of kiionn iiitrimitr (n «mk nan bi-ing chomi, or, at k-Jifl, not one
nbioh vroulil ciuim* id the iiiii»t'If ii Duixtniiini <><>ii tract toit) he tltruwn in ■!
r. A votitriK-tioD nf « oerntiD nnwiint will fnllon-. That coiitmctiQU niar
rio. ul
\
B
Itnajtrtmxi. pjttt^axTiaia, -"ixli ibn arrrv ntxmA In A ivailMRwIteffiiiuIla Sioui
■uaoiIiAfcnuiinuiriirTuaL
Inmrh n Li Ihv BiiiVli.-. (' inv lnKlimlv of iliv miutaiil fiiirfiit: i repnWDtt llM (t«l dlien lk|
liuliii'lliiii.*hijc^, uteJ 10 ini ihL- IntlHliilII} oT l)i« intrmaru anit Id.
\k tnkcu as n luenaure of tb? irritJibility of th« iwrvc at th« pcunt f. Nov
let itie |H)IuriEiug ctirr«nt be (hrown in iiikI lot the kathude or iieRmtive [Hilr
1>C m-urt-Bt the luuw^le, as in Fig. -10, A. »> thiit the curr«at pawcB alntig tbr
ncrVL* ii) a direction from the central nervows syRteui lowarxl the tnUKk;
BlK'b a current is B{K>keu of a;* a ilr-rerti'liny one. The entrance of l)i« polai^
iiing current into the nerve will produce a " makinj; " cootravtion ; this m
niD)' neglect. If while the current is possiDj; the «ain« uidueliiin-*bock nc
b«rarc be sent tliroueh X, the contraction which rceulls will ht- found to Ix-
greater tliau on tbe former occuiton. If the |i<>liiri/inK ciirroni hv aaw Mhnt
offj B " hri-akiug " coutracliou « ill |>njliably he )>ri>diU'ed ; thiH wc lUwi may
neglect. If, now, the jniiut jt. idler a short interval, he U|[iiin tested with the
same imluctinn-Hhuck ait before, tlie coutraotion will be no lonmr greater,
but of the same amonnt, nr ]ierhn|iit not >a great lu at tint. .During tbe
pnriwg.ci of the polarising mrrent. therefore, the irritability of tbe nerve at
tbe point r hi>» hcfn tvniporurily iHertnt^il, v\di* thi' >amc «hork applied to
il caiiM* a grv-ater cnntnu-tioii during the prewJiiw; than :n the abMinoe of the
current. But thi» i» only trnr iu> long il- the ])iiluri»ng current in a descend-
tag one— so long lu thv jKiint x lir» on thi- hicIi- nf tho kathode. On tbe other
hiuid, if the pijlarij:iiig runrnt had l>r<-n nn nKi-ntitiiiy one. with the anode or
poaitive pole ncnrcj^l Ihc miisclc. a» in Fig. ^0, B. the irritiibilitv of tbe nerve
at T wmild have been found \o\Kiii)iiii\i»hrd,xnt\nu\ of iiicrcasea, byibe ]>(]lar-
izjug cun^iit ; tlie cunlraclion ohtaimxl during the paseage of the cuu»tMUl
»TI»CLt:B Br ELKCTRIC CIRHKST.
ISl
eamml «(niM bv Ine tlwn bi-fuiv the pntsage of the cumnl. or niieht be
•IhM altDgaUwr, ani) the viminiclion iiftvr ihc rurronc Imic) been «Aut oiT
■■■Id ba w grtmt, or jtt'rlia|» gix^nlcr. tlinn bcfoix-- Tlinl is to tuy, irheii a
nMMtsai riirrent is iim'licH lonlicn-p. ibo irrilubilitr of llie iwtvo lietween
thm piiUriiiit); vlcclrijilM mnA ibt? musrle is. during the passage of tbc curr«Dt,
I kIh-ii the kalliocle i* nearest ibr niiiscle (and the polarmii); ouirenl
llDU.i flud illiiiiiiishcil uli«D tlie auode is iienrcMt the muscle (and the
{■lubitii; tnineiit nsccitilinK '- Ttie same result, mutiilu mulaiiditt. aud witlt
HMB i|aBlifi('aLio(ifi wliich we iKeil uot disouas, would be gained if J' were
fbccd, nut between ihe inuitole and the [lolariziiiK (.■urreiit, but i>u the far
»im »f tbe latter. Hence, it way be ataied ^■^"^"^"T ''■i*' during the
|aMBg« uf a cunnlani current (brouj^h a tier^'e tlie irrilnbility of tbc nerve ia
vtenamd m lite regtnu nf the kaibotte. and <linruiijlied In tlie region of the
uufAm. Tbe dtnngn in the nerve which give riie to this increujfe of irrita-
hililT in tb« rci(ioii of tJie kathodes are «|>(>ken of a« hilelwlrult>tiu4, and ihe
MfTc b Mid to bo In a kalelecimlonic i-nndiiion. HJiuilnrly tho change* in
tl« rnfion of tb« anod? arc vjioken uf an <i>ii-/irfroto»tut, and the nerve i* nid
to be in an aiflwiniloDic t-ondition. It ix alM oAen unial to «[tcak of tb«
UivtectmiMiir incrawc, and luii-lertrotonic ilocrtMC of JrHmbility.
Tkia Uw mMiw tnie nhnlcvtir Iw the ni<ide adopted for di.-icrmininK the
ifriubPitir. The rt^till hol'U k"-"! not only with a xinglu induction -vlitick,
bil tbo with n tctaniiiiig inli'rni[ii(.'4l current, with chrmicjil and mochani*
olriiMli. li further Hpiware to bold good not only in a diisrclcd nerve-
MkIs preparalioii l>ut also in the intact n«rve« of the livini: bo<)y. Tbe
and (k-creiisp of irritabililr arc most niarkc<l in ihv inimwliale
MUbdrbuod of the otc<;trodes, but spread for a considerable dii>tuncw in
•aradiTfcUoa in the eiimpoinr ref,'ions. The same moililicalion is not con-
laalbi tbeexlra|M>hir rcfinon, but exiata al»u in ihe in tnipolar region. In
tW intrajNilar region (here luuAt be, of uourae, a neulrnl or indilfvrvnt jwint .
tbot the katdectronic incn-ave niergcn into the an elect rot »nic decreaae,
uJ wbtffi. therefore, tbe irriiabiliiy i^i unchangetl. When the )i»lariiing
nnwiiiaifeak one, thin iudilfcr^nt jxiint u nt^nrer the anode than the
krib«dt,bat U iIk nolari«iii)i! ciim-nt increuM* in inlonsity, dram nearer
u4 Huvr tbe kathode im Fig. 41 J.
fm. II.
»■*■!« tLUnrmim TH> ViHUtnoB ur iHBttian/TT ui-uiNi Ri^tcnDMNOh wnn
rouauisa rtunvim nr IvrttiaHO tnnsiTT, irroin fRogar)
n*IKat> I* viiqnnd Id U* iJaoXl at .1. ilie k41b«d* al 8 : AB li luiuHiaDndT (hn InimiJOlit
Matt. toM«kiirUnibr«*«>u-io«,Uioiianlana(ili«tiimb«to«'llwliM«Bn<t rvjifSHnwdlMln-
■If, llut> tlbim. tnvraucil lirlutitlnr. n Wftiwim lh» (Okn ot • inak cumnl : lliB
I tuaal r, U nmi IW nnule A. In |>i. ■ nroDCH I'umBl. Ihe liidlfltinil pc4ii( t. It iwanr
I Ik iM dnnlnunan ••! LrrlWUIIiy la uwitoiniioiKit ■sJ ili* luprMw i<> knialccuuionut
rUsn Imp,: IbeeflMI ■!» ipnail* Air a (ivtar dliUum Along Ibr «i(ra]vUir nt(ini»
iHt AMH^ In rttb«<aB>n>iiuai«MeuK>beMUImoi« mulwl-
TW ■OMKUil of IncrcMM and decrease iadepewient: (1) On tb6 >ti«Dglb
tf tk» «nnu. iba Nronger current up to a cerlain limit producing the
1S2
THE COXTHACTM.E TlSflirgS.
Knier eHbot, (2) On ihe irrilHbility of ibe nen-e. lUe more irritAhle,
ttr condiiioDed nen'e beiog the more aHectetl by a current of the «une
tatvnaity.
In the cx|>erim«nu jii»t de«crtbed the increase or decreuM of irritiibilitf
t8 ukcn u> mean tbat the tame stiniuluH maru in ifae oDe ease a lar^r or
more powerful, ami iu the olber case a smaller or le» ener^Uc impulse;
bill we hav« remioii to tliink that iJie mere tirojiagation or conuuction uf im-
pulsra slarlcd el«ewbcre la aldo alTected by Ine elevtrutouic CODdition. At nil
evenU Knclf^'lrotonujt appeam to uffvr an ob*Uu-lc to the punge of m nvrvoiui
imputee.
§ 76. Eteeirotoiiie currenti. During tlie paatage of a conswnt current tbmuxb
a nerrc, varlallon* In the electric currents belonging to the nerve iiMlf mav be
obMTTed; and Uieno VBTiationH have ceriAin relation* U> tlip varinlions of tlie
■rrilability of the nerre. Thus if a convtant current supplied by Ibe battetj
Fw. e.
h
<
Oijiotun iu.i«i>Aniin SUcmnognc CcannrTK
/•ihv tnlorlilDS bAiierr.-wItblB liii<r. p ib« anoAp.auJ p' lh« kaihodr. At ih« Uft •Bdt«f <
|dttp«of nciTc lUt uniiml curronc Oouv (lirounn Ihe jpiInrHimeler Q ttvtn g ioff\ ia tiiVdlracO
oT iLu nrruHt; )i>aiirer(lou. ihenthiir. !■ (tio tmiisai Uinrof the inlvUlDR cormit: waM^ocntlr ft
■IiiaanlamunJ, ulddlailHltiTlhiMlBii t . Th> ounviil at llio othor «.tiil <rf tbe |ilN* of ii«r*«^
IhHd A lo A'. ihroUBb Uie vilranoiagier M, floni In ■ coatnr)' dIrceUmi to itw poUrfdac cnnvnt ; R
oonMi|U*QU) appBB Lu br illinliilttiiiil, u liKllCBtal by tboiltcii — .
X.B.-Pradmiiltdtjr'iMkr. ttic |»lBrlidnr tumiit IdiarttUITaWl to be Ilirawii limttha nldill*
at ■ FfNnor ncrrc, and tIWKiil»nuiaL>ivr r'*™*) >' t'lu iwovmlf. of coiin(lti*1llb»iiiiil«iMoo4
Uiat ths Akiiuit nuy I« Ibfowii in anjiwhvia. atiil tbv litter onnnn-lci) wllb 1U15 two pAlnof (dnli
wbMh nlll glT^ramata.
P (Fig. 4a) be applied to a piece of nerve by means of two non-polnriuibie
eleclrodc* p, //, tlie "current* of re«t" olitainable frura the rannuH pnims
of the nm-e will be diflerent duriuK the paMaee of tbr pulariiJuit current
friMn tbone which were uianifist before ur nK«r tlie current waa applied ; and,
moreover, the obau£«a lu the uerve-curreote produced by tbc pofarisiug cur-
DTIMDLrs BY KLKfTRIC d'HRKNT.
183
I
will ihM b* lb« MUM in the nfijthborhDud of tbc ■node (;i) m tbonc in
IW Mi^borboiid of Uw kath»<)« (p'y Tbtw let O aud H b« two gii]viui«<
M»t<r» ao coBn«cUr<l niili tlio iwo piitta of th« nerv« aa to alTord i;ood aiid dmr
•v14«BCa of tlir ''ciirrrnl* of r««l." Itvfnrci th*> pnlariKJi)^ current is llirovn
iMo tk4 B«m. thr a<-r>11o nf // will onruny ii pmition inilicnlinfc ilic puiaj(« of a
WtraatofK c«rt«in iiiitiisiir frocii A tu A' ihruugb the nlvsnoniPtcr (from the
MUivvlaticlludiiinl Mrftice' to tbe ofigitiro cut «ud or the nerve), the circoic
Wlaf e<MDpM*d by « currcoE ui lhen«rre ftvin h' to A, i. «.. tbe currvnl will How
l> Um lUnctioa of tbe arrow. t^imiUrl^ thv n«odl« of 0 will, b^ iix dcAactioR,
Mkue tlw exlMcnce of ■ currenl fluwinx from ^ to jr* tbruugb tbc Kalvunome-
ter, u»d tnm 9' toy ihruugb tli« nerve, io ibc directiixi of ib« arrow.
At lbs iaitKDt that the poUrUln); current 1* tlirowii Into the norve at ;>;>'. tbe
fwiWila at jm'. AA' will undtrgo a "oqfstive varintinn." that in. the nerre at
MCh point will exhibit a "current of action" ci>rrRtpniidinf: I« thu ncrvoua im*
|«1m. which, at the mnking of the polaririoK current. pOMUo in bath direclion*
■hMK the oerTo. am) may csuM a oootraotioo in the atlached muscle. The our-
nttt at action ii, as wo hare teen, of eitrrmcly short danUion. it Is over and
|MW (a a mail fraction of u atcond. II. therefore, munt not be oonfounded with
sMnnaaani effect which, in tli« cas« we iir« deuliii^' with. !■ observed iu both
pItaBiMMUn. nia ell«ct. which 11 de|)t'iidi'rit mi the directi'jii of ihe |H>1arle-
■(CWrml, MM follows; .Supposing tlial tbc polari/.ine current is Hnwing In tb«
Antdn of the arrow in the nftnro. that i«, passn in the nerrc fmm the ixwilive
Jmiode or anode p to the negative electrode or kathode //. it i> Tounil thut the
nrtvnt tbroagh the g^VBtiooieier O ia iacnaBcd, while tluil through J7k dimin-
xbid, Tbe (KiIariRlng current has caused tbe appearance in the oerTe outside
ihs4Klfnd^ of a current, baring tbe wtme direction aj> itself, called the "elec>
tntnir " current - nnd this e1«ctrotonic current mliU to, or Ukn nirsT frirm. the
MMal oerre-cttrrenl or "current nf rent" sceording at il is flowing iii tbeaatne
JbMieu sa that or in an opposite direction.
1W iliwBitih of tbe electrotonic current is dependent on the >lr«iiKth of the
pahiiiliit airreot. and on the length of tbc inirs)>olar region which is exposed
L to the fUmtitiBK cnrrent. When a stmnit potariiintt current in used, tho eleetro-
B Hlln fiwce of tbe electrotonic current may be uiucb |{re«tcr than thai of tbe
■ aunl Mr«»-carrent.
H Thi ttrencllt of the electrotonic currcnl vsHm with the Irritability, or vital
H rwsftliM or th» n«rve, being greater with the more irritjtblc nerve , and a dead
P Btm will not nuniresC electrotonic current*. Moreover, the propagation of tbc
[^ uiJHsi b Mopped by a ligature, or by crushing the nerve.
W* najT apMk or tbe conditions which girc rise to this electrotonic current as
a f >>wH/el«€tro«oniu nnalogoaa lo th*t pliyiolngieat elvetrotonus which is made
bna by variations in irrilaoility. Tbc phyiical electrotonic current ia probably
fo U> the aacaife of tbc (Hilariung ctirrcnt along the uerrc under the peculiar
oadllleos of toe Hrinf; nerve; but we oinit not allempt to enter here tiitu ibb
Wall subject nr ini'i tbe allied quoilion as to tbe riaci connection between the
apical and tbe phyiinlogical clectrotonu*. though there can he tittle doubt that
I latter la dependent on the former.
(fl. Tbaae variations of irritahtticy at the kath'xlc ami anode reapec-
Irwif, IfaiM brni)f;hi about by the action of tbe coitstAiii current, are inter-
Ming ihenrelicnlly, boonase we may trace n connecliou between ihetu nur)
tbi ncrvow iDipulae which la the reaiilt of the making or breaking nf n
tmtrnA current.
Par we have evideitce that a nervous impulfte is gencniled when a portion
if tbe nerve puaea auddeiily from a norniul condition to n Mnle of Icatclvc-
tMsoBS or fitm a atate nf notilec-iruionu^ hack to a uormal euiuliiion, but
telka panajfc from a oorninl <<ondilion tn aiiele«lralcuus or from knti'lcc-
inMKia back to a uomml condition Is unuble to gvnernlc an impg|»e.
Hoe* wben a i^nslnnt ciirri'iit is "roado" the impuW is gi-uvrati-a! only itt
lla kathode where th^ nerve [HMtifi ninldenly inUi kntelei'lmt'imi*; when
ttamrmt on the other hand is " litoken " tbe iiuputsc in gcneratcil unlv at
■iUiand* where iJie i»erve |MtsiN.>e suddenly luick from aneUictrotonus into
THK CO:?TYIAOTIt.B TlSSCBfl.
» nomia] oandilion. Wo have au imlin^t proof uf iIim in ihu Factii lu
which ff« <lrew atteutiou ii little while buck, rix., thnt a coiitrnctioD aoine-
liincs oceuni at the " bre&king" only, MmHintcs nl the "nuikio^" only of
the coD»lniit currvnt, tometimo nt hoth. Fur it in fotiml thnl thtg ijcponds
partly on tho mrcngth of tho current in relation to ttir irritability nt' tbr
iivrTc, pnrtly on the direction of tho current, whether iiM;cn<liDg or ^csceiid'
ing; and tho results obtained with Aitron;^, medium and wonk descending
and asronding cnrrcnls have been slnlod in tho Ibrm of a " law at' cotitroc-
lioii." Wo need not enter into tho details of this "law" but will merely
say that tlie results which it formulates are bc^t oxplaine'l by the hypocbe-
eisjnsl elated. We may gidd that when the constant current is applied to
certain structure)! comiKised of plain niuitculAr Kbrcs, whose rale of cnolrac-
tion we have seen to fa« slow, the maklnf; contraction nmy be actually ae^ii
to b^in ai the kathode and travel toward the aitode, and the break'toff oon-
traction to bei^in at the anode and travel thence toward the kathode.
Since in katelectrotoDuii the irritability ia increased, and in aneteolrolonua
det-reaaed, both the entraiii-e from the normal cuiKlilion Into katelectroioDiu
aod the return from auelcclrotonua to the normal condition arc iiulanci^ of
ji uawajte from & lower Atage of irritability In a higher Htage of irritability.
Hence, the pbenomena of electro ton iih wnuld lead us to the oonoepiiuii that
a atimulutt in provoking a nervoun impulte pntduceH itn effect by, in »oinc
way or other, suddenly raising the irritability to a higher pitch. But what
wc are exactly to uiidemtand by raising the irritability, what molecular
change i« Ibc («ti«e of the rise, and hnw either electric or other stimuli cnn
produce lbi« cliangc, are mutters which we cannot discniu here.
Bendw their tbuorvtic-al im[i.irtiiine, the phenomona of electrotoou^ hare
nita B practical interest. When mi iis<-cttding current is piuwtl along a nerve
goiiig to a mtticle or group of mus^le^, the region between the electrodes and
tne muscle is tlirown into anelectrotonti^ and its irritability is dimintshetl.
If the current be of aderiiialc strength, the irritability may bo »> much
teteened that nervous impulses cannot be generated in thai [inrt of the n«rn
or cannot paea along it. Hence, by this means the irrejjular contracliun* ot
muscles known aa " cramp " may be abolbhed. Similarly, by brlngiuff into
a condition of aneleotrotonus. a portion of a sensory nerve in whieh vinleni
impulses are being generaletl, giving rise in the central nervous syaicni Ia
scnaalions of pain, the impulses are toned down or wholly abolished, and the
pain c«BM«. ^. on the other hand, we may at jtleasuru heighten lite irrita-
bility of a part by throwing it into katelectnitouuii. In l£b way tho con-
slant current, pni)ierly niiplied, becomes a powerful remedial innsua.
We said just now that probably every ttimuIuH produces ita tdaol oq a
nerve by doing wliat ibo conntnnt current docc when it a<Ms as a Stimulus^
vis., tuildrnly raising the irritability of the uvrro to a higher pitch. At any
rate, the etimuluK «o ot^n employnd in experiinents — the induction-f bock-
acts exactly in the mmc way as tho constant current. The induct ion -shock
is a current of short duration, developed very suddenly but disuppeariDK
more grnduitlly, and thii^ is true both of n making induction shuck, a shock
due to the making of the primary current, and of a bmking shook, a abock
due tn the breaking' of the primary current. The two dtHer in direction
(beoce, if the makinj; shock be ascending, tbe breaking shuck will be deaccud*
ing, awl viet ivnvj ), and in the &ot that lite breaking shock is more sudileiily
developed, and hence more (loteot than the making shock; but otherwise
tliey act in the name way. In each case, nnce the induce<l current is devel-
uned rapidly but diMpjiears more slowly, there ia a sudden development ot
electrotonus, of kalelectn>t()nus at the kathode, and of anelectrotoiiun at the
aaoiie. and a more grmlual return to the normal condition. Now, ih«rc are
rnx XUflCLR-NSRVB FKEl'ABATION AS A MACHINE. 196
I
muiy rttmrnm for thinking (hat in »)l ohm (he jnauii}; horn lli« normal uon-
4idi>n 111 k»l«lwtroioniM ■( th« knlli<x(c is a nxjro potent siiiiiiiliis than tli^
ntani (rtmt antlectrotonna to tW uurmal cuuililloii ai ihi? iiiiudi.', and lliia
•ill b» itill Biora M if lh« return to the uormal ritihlitiott lio much «loiror
tiuui tha entiancs into el«ctruionus, as ia the caae in iin iii(tiictinn-#lu)cik.
AimI ii wixtid appear that in an intluciioii -shuck, which, lu* wc hiivomid,
clHapjNvn much more slowlj than it ii lievelnped, wc hiivu to 'teal nut with
two atimuli— one at the shook |<ttAiinK into a nerve, and one at thc*hiick
IfBvinit ibf nerve— but nith aiw uulc, thnt pruduoeil at the thwk passing
tnUi tlie nerve, ilenee, irhen itn tnductiuii-sliock in mnt into a nerve, ono
lus unlj is develo|>cd. and that at the kalhoJu only, the eslnblishmeBl
lUB. This iR true whether tliu #hock be n making i>r a brewk-
__ dinek — I. «., line to the inakiuK or brcnking of llio primarr current--
thnugfa, of course, owing to the change "f ilirvction in the induced ciirreal,
•fast WBB the kathode ni the making shock booooiea llie aoodo at the break-
luahock.
l^rily, though Iff arc doling now with nerves going to ran«clee — that is
\a Mjr, wilh motor iH-rvm only— vf« may nild that nhal ne linve said about
tlectroluaua and the dvv<'lopBwnl of nervotis impulses by it appears to apply
t^uallr wall to Pttaary nrn'ea.
( n. In a general way niosciilar fibr^^f b*'have toward an electric etirrent
r* aiurli nji do nervf fibrt^ ; but (here nre certain inijiorUmt dilfervnc*^
In Ibo lirM plnM-. niupcular fibres, dermd of nerve libra, are much more
rradilj thrown into contractions by the breaking an<l making ol' a eouitanl
niTTMit than by the more transient induction ■ehook ; the muACular Mib^tance
iMia* to be more sluggish than the nervous sub6taiii.-e, and require! to be
Mlad npoa for a hitger time. This fact may be made ime of, aim). ind«ed,
ii b wdkal praelice made mm of, to del«rmine the comlitioii of the n«rve«
•■i)iplyiDE a muMrle. I ( the intni-muAciilar nervo be Mlill in gcMKi condition,
the noMle. to a whole, rojionil* nmdily tu single induct ion -shocks, beoause
ikae tsa a4;l upon tbc inlra-muM-ular nerve*. If these nervtv, on the other
ka»' I<wt their irritability, the muM'Ie diMv not r<«|iond readily in
aa^- iua-«hucka, or to thv inta-rru ptt^l i-iirrcnl, but can still easily be
ihiima ink' c«ntraeiii>nit by the ci>n<4AnL current.
Id liie eeuind |»laoe, while in a nerve no iiupulsos, as a mie. generated
Joriac tbr |M«age of a ountHnni mrrenl. between the brenk and the make,
fntun] tliai it ia not ton drong, and ihtU it remains unilunu in strength,
Ik HI urariznl nin>M:li.-. on the oilier hand, even with moderate and perfectly
aaifiMin rumrni*. a kind of tetanus or iipp«rently a series of rbytnmically
NfBlcd ctmtractioDs ts very freiiiieully wiinesKd during the padsage of the
•anal. Tbe exact nature and cause of these phenomena in muscle, we
■wt >!«, however, diecuss here.
TnK Mtr^iCLe-NERVK PkBI-ARATIOX ah a ?tlA(')IING.
tYI. The &ct* described in the foregoing tvelions ifhuw lUatn miit«le with
ikaenrc may hv juKtIy reganled as a machine which, when oiimnlated. will
^ 1 eertain unuMini of wurk. Hut (he actual amotint of work which a
aowle-nrrvr [>rfparat>oo will do ts ftuiiid bo depend on a large number of
■vaUMaDrca, and oi>nK»)Uently to vary within very wide limits. Then
wiatinns will be largely determined by the condition of the muscle and
Wn* bl rcapoct to their notriliou ; in other words, by the degrc« of irrita>
tSqr ■lanUeMei) by the niitM.-le or by the nerve, or by lM>th. I3nt quite
■fart tnta the general influences alfecting its nutrition nttd thus its irrilit-
ISA
THE 0OSTllACTIt,K TISSrBS
bililr, n muiclc'ncn'i^ prppnnilion t!> mflt-i'tt^I nn n-tfitrtlit llic ainnunt of \t»
vrniU by a variety of othrr <'ireiitimtaD<<M, ivhi<'li vc ihmv bricHy CAiintlvr
horc. rcsprviiig to it «iiPOc^cliiig mxIioii the oltiiiy nf vsrialixii!' iii irritnltilitv.
Tlif injlufriw nf (lit nnlnrr unii madr nf njiplictlit>H of thf utimnlm. Wbrn
WO Apply n wcnk etimului — u -kv^M in(fiiition->'hopk— to R nerv« wc gvt a
smnll con Unction, n «[ight »)i'>ru-niii|i: of Ihc ttiiiM-lr; irhm ire npfilv A
8in>nt!Pr etimrilii!— n Hlrongtur in<lucli<in*hiH'k — wp gel ■ lnrj»r t-imniiclloo.
a fireutcr »IiorlPiiiii^ of tlio iniisdc. Wt- Inke. olh«r lliings Ining cmiu&I, the
BiDdUDt of <.-oi)ti'aclion nf the muide iis a mmsuif of tbi' nerroiis imptilee.
an<) my that in Ihe forni«r case n ivenk nr sli};hl, in the InlU-v cn^e n slmn^r
or liirgiT. oervoui impiilBe \\\\% he«ii ;;i?neriiio(], Xou', the mu»cl« of the
inu^cle-nerve itrepiirali'm coii^iiilti of niuoy nntscular tibree. niid thft aerVA of
uatiy nerve 6brc«: and we may fairly Buppnife thai in two ex[«rinenta we
may iu the one experiuieiit bring ihe iutiiii^tion-ahock or oiher eiimuliu to
Iiear on a few flbrea outy, and in ibe other experiment on many or even all
the fibres of thf nerve. Iu the former case, only those muscular fihreit in
vbich the few nerve tIbreH*[imulaleil enil will be llirowu into contract ion.
the others remuininj; <|uict, and the shortening of the muscle, aa a vltole,
aince only n fen librMi lake pan in it. tvill neccnmnlr be 1cm than ythta all
tlic RbrcH of the nrrve arc stimulad^l and all the fifireM of the mnacle con-
tmcl. That w to Miy, thv amount of contractinn wtU d<-]icnd mi tbv number
of librcf stimulated. For timplicily'* nake. however, we will in what fol-
low*, exeepl when otlicrwicr indicatMl, mippn*' ibnt when a nerve \* otimu-
lated. all the fibres ar« Htimulated and all the miundHr tihrri! ronlract.
In »iieh a case the Mrongvr or larger nervous impulse leading to the
crmler contraclion will mean the greater disliirliance in each of the nem
hbree. What we exactly mean by th« grent«r disturbance we mu^t not dis-
cura here ; we must be content with re^nrding the greater or more jKiwer^l
or more intense nervous impulse as that in which, by some mode or other,
mora energy in *ei free.
So far i» we know at pmunil this ilillen-'nee in anintint or intensity. «f the
eacrgy »rt Iree, is ihe ebief diflerence between \-ariiWs nervous imiMilsc*.
NerroUM impulM-s mav diRer in the velocity which they travel, in tlte length
and {NMsibly in the form i)f the impnlsi! wave, but the chief ilill'erence u in
Strength, in, su to speak the height, of Ihe wave. And our prexonl knowl-
edge wilt not permit n* to point out any other ilifTereneee. any diflercnc« in
fundamental nature for instance, between nervous impulses generated by
different slimuli, between lor example the nervous impulse generated by
electric currents and those genernted by chemic-ttl or mechaoieni stimuli, or
by lh<«e changes in the central ner^'ous system which (;ive rise to what may
be called natural motor nervous impulses as distinj^imhed from tlioee pro-
duced by nrtilicial stimulaliun of motor nerves.'
This being preutiseti. we may any that, other things being equal, the ma^-
nilude of a nervous impuUe, and so the luagtiitude of the enauing conlraclton.
ix dirai'tly dependent on what we may tall the siren<;ih of the stimulus.
ThiiH taking a single induction -shock as llie moat manageable stimulus, we
find that if, before we begin, we place the secondary coil (^Fig. 14, >en. e.) a
long nay olT the primary coil nr. r., no visible effect at all follows u|<on th«
discliarge of the iuduelion-fiboek. Tlie pawage of the momentary weak cur'
rent is either unable to produn- tiny nerroiis impulse at all. or the weak
nervous impulse to nhich it giv«« rise is unable to atir the sluggish muscular
Bubslanoe to a visible contraction. As we slide the secondary coil toward
the primary, sending in an induction-shuck at each new position, we fiml
I U will hr olwrrcil thai w«ara sivakltia nnw vxrliiiirslj-nC Ihp iirrvv (>f n KiUM'lij'nRte tman-
tlAa.l.r, orvhii >ic*l>uil henmfli-ririm h inalni iierrv. WiudierKnwiylint'liliMiIIIIkrefficatiBlljr
than nuilor iia|iii]t» u lU la fiiiiUrml laim »ii
TnR MUHCLK-XKIIVE fKEI'A It ATIUN AH A UACHIXE. ISl
k
I
lfa»l at m reruin diflUoee between theEMmodarv itii<l pHniiry coils. tht> tiiui-
Hr nmftendt In Htcli itiilufliou-shiMilc' wil'i a coutniction which ninkc^ ilMlf
viijble in Ibe ■[i)[)i[«M |Kjet»ible rise uf ihe ik(iacl>i>d lever. TIiih pavilion of
lbs onib, tlw hnlU'ry rrmuiiiinf;; tii« tame and oiliur thin^ IwiBjf e(|iinl,
■Mrfc* Ihe wini'miiV niiuulus jfiviiiK riae W the ruiuinial iM:intraclJi>n. Aa the
■Mooilarr mil i.-< Im'UKlii nearer to the prinuirv, (he ci^ntracltoiM increiwe in
Iwigbt cnfTiM|»tiJiii>: ti> ibe UiervHM iu the iHU-nnily of the«tiniiilu!i. Wry
aoaa hammtr na incmiiH; in the MimuluN Cftiued by furihrr cliilinK >1k-
MCMwianr coil ovr>r the primnry failK to enuM anr iricrvuw in thv oritrniv
lioii. Tbi* iiidii'at'i that tbv Majrtm'i/Mimiitnit ^viiiji Hoc to ihu niaitimnl
«aati»cliun htu If4'«ii rvuchw) ; thougb the shoclut incmiw in inCvtmily a«
the wcoadary ooil i* pudNd furtber and riirthi-r over Ibe primary, the con-
tnrtlona mnain »t' the nine height, until fuiiguo lower* them.
With «ingle indiK-tian-sliocks llien tht- miiKruliir contraotion. aD<l by
iaftrmce tlw nervvtu iiapulM. incn;af>v» with an incroiiw in tbe intensity of
the ititDultW, bMireon the limit* of the minimal and iniisinial uimuli ; and
lUf dMMSwkKUie »f the nervous impulse, and so of ihe contractioo. on the
MTCogtA of tbe stimultis may l>c oWrve^i not only in eleetrio but in all
UiMb of Mimall
It mavhera b« remarked that in order for n stimuluH to beetl^tire, a
<*TUin atm[ADeBS in its cetion is oeoeaaarT. Thus aa ire have »een the con-
>L>nl ctinvnt when il b fMAinj; throcigh a nerve with uniform inleiiitity does
Mt pvi; riae to a nen-oua imputac. and indeed it may lie incmiiKil or
<fiMini»he<l to aluwet any extent without ftenerulin^' nenvma impiiUta. pro-
vUmI ifaai tbe ehange be made (•radually eiioujch ; il t^ only when tlterc 'i»
■ addon «bMi|{e that the current beonmen eHi.>4:live ns a miraului'. .\iid llie
ntmto why tbe breakiii): induciion-sliuck in murL- piient iu> a miniidu* than
the malcing «back is bcxaitu>! an we luive sorn i M-t > (he current whii-h ia
iMlncwl in thr MVondary coil of an imlurtion- machine at the breaking of the
(mmary circuit, is mon rapitlly (level<>)xi(l, and has n sharper rise than the
tarrrnt which ap|>rAre nhi^i tbe priniary circuit in made. Similarly a sharp
lafi r>n a nvrre irill pmdu<?c a contraction, when a gradually increouiig
Iiiwaurv will fnil to do so : ami in evnenti the efficiency of a atimulua of any
,iad will depend in pan on the MiJdcDnesa or nbrupln«ee of its action.
A •tiinuliis in ortlcr that it may be effective, muat havo an action of a
ivrtaiu itiimlion. the time ncceeeary to produce an effect varying according
In tbe strength of the •tiiiiulus and beiuf; ditlWreni. in the case of a nerve
IVnm wbnt it ia in thecMeof a muscle, [t would appear that an eUtrlnccur-
not applied to a nerve muat have a duration of at Wet about O.UOIS *etv>ii<l
In oiuap any nintraciion at all. and needi a longer time than iht* to produce
ill Atll effect. A muac-lelibre a|>aTi from its nerve fibre re>)uires a &u]l
kdgcr duration of the ttiiniulua. and henee, u we have alreiidy rotated, a
■iMde puiMUt^l by iirari. ornhieb kaa otltcrwiM loM the acliiin of iu uerves,
■ill n«i rupcxid an readily to inductiou-sfaocka as to the more slowly acting,
hlMtkiog and making of a conuant current.
Ib Um om of clirclric uimuli. tbe Bamc current irill prwluoe a stron|^
nMraetiMi wh<m it ia *ent along t)ie nerve than irbuii it ii aent aerom tbe
wnre: indeed il ia mainminiMl that n current which {Mate* tbrougli a uerve
b n ahtolufly tnuufvenc directiun in ]>on'CrlM» Ui generate impuuea^
It would aU(> nppMr, at all evcutN up to certain liniltji, that the longer tlie
(inr of nerve through which the currviit poase*, tbe greater is the eifect of
lieMimtilii*.
._ Mihtv ikt WmUiit w nuKUiaikoekBreMkeBied. DM iMSFtlmw c— aal
«• oikrr. a*. M !*• luin <MU>d, Ibv Irni U«d* uf ikMk dUftn Iu dlDcttocT. Ik* bnaklaf
(to BM* fDiani
188
THE CONTRACriLB T1S8UBS.
When twu pain u( electroden itre pincod on the nerv« of n long uirl per*
fealy l're»h mid Hucoeuful Derve-pr«]»t ration, one near to the cut end, and
the other nearer the muscle, it in round thnl the Htmc vtimuiiis producee a
gre»(cr contrnclion irhen iipplied through the former pair uf electrodes ihtia
ihiMu^h the latter. This has been interpreted as meaning tliat the impulse
Staiteit at the further eleclrodee |^the» islrengih, like an a\'nlaiiehe, in ila
progreaii to the mu«cle. It is mom probable, however. Chat the larger con*
traction produced by Himiilatioii of the pari of the nerve near the cut end
is due to the stimulus !<ettiiiK^f<^'! a larger impulse, i. e.. to tliii part of tha
npTve beinj; luun? irritabk-. The men seotion, possibly hy deTelopttiK ii«rvu
currents, luirreaitai fur a lime the irritability at the cut end. A .similar
l^reaier irritability mav honever nl«o tie observed in tho ptirt of the uem
nearer the ipinul mrd while ii ix mill in oonucctioii with the spinal cord ;
ami it ia pus^ibli: ihut the irritnbilily of a nerve nuiy vary oon«i<lerably at
dilfereDt puiniit of it^i c'iun>c.
$ 60. We have luvii that when oin^Ic xtimuli aru ntpcated with Kullicient
frei)iiency, the individual coiitrnction* arc fuied tnli> tetanus; nc the fre*
queiicyuf the ropotition is incrc4UH^, thu individual oonlraolions are leas
obvious tin the curve, until at last wo got a curve on whifh they iwcm to be
entirely lotft and which ne may qwak of n» a complete tetanus. Br such a
tetauuit a much ^Evter contrnclion. a much ^rvalcr shortening of tl>e muscle
U of course obtained than by sin^ilc contractions.
Tbo exact IreijUcncy of repetition rcijuired to produce complclc tetanus
will depend chiedy on the length of the individual contractione. and ihia
Taries in different aniuiaia. in dilferent niuiclcs of the same animal, and in
the tame muscle under ditferent condilions. In a cold'blooded animal a
dngle contraction is as a rule mure prolon^^d than in a narm-blooded ani-
mal, and tetanus is caiise<iiifntly piuduced in the former by a lea* ft«>|uent
repeUtion of die utimuluti. A lirml muscle has a longer contraciton than a
freah muscle, and hence in many Ivljuius curves the individual ontrnctiona.
eauiy reci))(ni/i,il at lirst. dieiip{H'itr later un, owing tn the individual contrac-
tions belli)! It-n^thcned out by the exhaustion ciiii»cd by the telanu* itaelf
In many niiiiunT», p. 7., the rabbit, somi^ musi-lis? i.siich as the adductor mae-
nus fciiiorin^ arc paio, while other* (such n.i the «emiicn<lino«iis) are rea.
TIte Kid muitclnt arc not only mora richlysupnlied with bloodvcwcls, but the
muscle suli»innc« of the fibres ooutains more nwraoglobin than the pale, and
there arc other etnicturnl dilI«rcneeB. Now tho single contraction of one of
thaw re<) muj>clcs is more prolonged than a single contraclion of one of the
{Mite muM-les produc-ed by the same stimulus. Hence the red muscles are
ibiuwn into complete tetanus with a repetition of much less frequency than
that rcijuircd for the pnle nmaeles. Tlius, ten stininli in a second are qnit«
sufBdcnt to throw the red muscles of the rabbit into complete tetanus,
while the pale inusclot reuuire at least twenty stimuli in a second.
Ho long sa signs of the individual contrai-'tions are visible on the onrra of
teuuius it is easy to recoguiiii! that each Htiiuiilation priidueix one of tli« cou*
Miluenl single contrmrlionH, and Unit the number su to .4{iLmk of thtt vibra-
tiofis of the muscle making up tht- tetanus correniiouiU to the number of
stimulations; Imt the <^u(»tinn whether, when we increiuH' the number of
stiiuulatiuns l>eyond that iiccmiary to produce a complete letanuK, we still
increase the nnniWr of rtm^titm-ni single contractions w one not so easy to
answer. And connected with this r|U4«tion is another diHicult one. What is
the rule of regietition of single contraciiims making up thow tetanic contrac-
tions which us wo have »aid are ihi; kind of contntctions by which thevolnn-
tan-, find indeed otlicr natiirnt, movement* of the body are carrieil out?
WW is the evidence that lIick- are really tetanic in character?
J
I
TIIC HL'SCLB-XEBl'K fHHPARATION Aft A MACUINK. 189
WbsD a mwde is lUrowu into teuiiuii, « wore ur lew iniuioM] Mnnd li
prixlucnl. Thb may be heard by upnlyiD)r a M«Uwwopc dirwtly ovtr a
floBirwrttu^ lauaele, and a dmilar aouoa but uf a more mixixl origin and ka*
iraMwoctky may be heard when tlte maixeicr iiiUNcle* are furvibly conUmcud
or «beB ■ finger ii placml iu tl»e «ar, ami iliu muscla of ibe Mine ann an
eootneUd.
Wbfa the M«ihoaoofi« b plan<l over a Riu*clv,tbvnorvoof whicli iHvlimu-
Istad by Miicii<<ii»liod(a repeated with viirviiii> rrxviui-ncy. llii< itotc heard
vQI vary wiib tlii- iroqucncy or (he ihiM'k*. Wing of hie\ivr pitch wtlh Ura
moro rr^«lu*^ut ■hwlc*. Now ic ha* bwn thought ihal Uii- Tihntti'Mtc of tho
muack giviii); riw to Uw " inutoU- Miund" arv tdniticaJ with ihv liitgk- con>
uractioDN making uit the tvtttniui of tho n>u»cK Aod eiiit.-o, iu the luimnn
body, when a luuacle i> thrown into (.-onlnictioQ in n voluntary cITori, or
incHad in any of tb« ordinary natunil movi.-menle of the body, (ho funds-
meatal tone of tb« sound oorrmponds U> about Hi or 20 vibrations a Becood.
it ha* bera eoocludnl that the contraction takinc place in eucb cnsee ■• a
'T^f Ttr cf which the inilividuat coolniclionH fottow each other about 19
or 20 llaWi a c«4.'ond. Itui invntij^nliona st-eni to show that the vibrnlionB
giviag rne to the muscle sound do not renlly correepood to the 9hon«iun|p
aad nlaxaiioiu of the individual contractions, and that the pilch of the note
maooL tbereface bs taken as so indictiiioii of the number uf ^iugle vontrao*
ligae BUtking op the tetanus ; iiidee^l, ait w« nlmll ba iu iij)eaking of the
MMiMfa of tlie heart, a single muMular contruciiou may produce a sound
■Uch tboggh dilleriotf fruui ih« »ound given out during tvtatiu* has to a
BNlain axtent mueicaf olioracterB. Neverth6l«n the ;i|>ecinl cbarHctcr* of
Ihk noMle Muod ^ven out by muwlee in the nniural movcmi^ni* uf the
body may be taken es showing at leut that the contniotiomt of the mii*clc
a Umm Biovementa arc Ictaiiic in nalnra, and tho stmilnriLy of the note in
■U Uw voltintary eflbrta of tho body and itidwd in all niovements carried
out by the central nervons system is ai \ftut consonant with the view that
tW rvptrtitioo of sinck contractions is of about tlio Mme fret|uency iu all
(bar niiiTenienis. \\ hat that frec)uency is, and nhclher it t<> cxaetiy ideiiti-
ral in all tbeM movements, b not at pnsent perhaps absolutely determined :
b(t OBTtam mrkinjp on the my<)>craphic tracing of thew uoveweutti and
olltar Iwu eeem to mdicaie that ii is about I J a second.
I BL 7%e itijiuenee o/ Ike tond. It might be iniagioed lhal a muscle which,
«bcfi loaded with a given weight and stimulated bv a current of a given
iairtuity, had coniracietl to a certain extent, woul<f only contract M half
lluU rxt«ut wbeii loaded ailh twice the weight and stimulated with the huoh!
ttfliahia. Such, howi-ver. is not neoesurity ilie case ; the lieight to which
tW weight b ni6vt\ may lie in thv second Inatanoe as great, or even greater,
ikan in thv fint. That b to say. the neeistance ofTcfeil t<> ilio coniractioD
sttuallr auiimentd tlte ooatiaction ; the tensiou of thi- muscular fibre iucreosea
ite Csnlity wttli which tho csploMvc chaiigex nstulting in a oontrnctiou take
fhra. And wc have other evidence that anything which ti-nds to stretch
the muacnhir fibres ; that luiv tennioti of the mu*cutnr ttbrvH, whether during
Mt or during contraction. increaM« the motalMlism of the muMlc. There
Kof cvtirv, a limit tii thi* favorable action of the resistance. Aj the load
cNitimits to be incrtaM'd, tho height of the contraction is diminbhu'l, aiul at
Isai a point b reached at whicli the muscle ii> unable 'oven when the stimulus
ibosra b ihasiroogeMt poaiibU<) to lift the loiwl at all.
la a musclSTMwed as a machine wr have to dcjil. mH merely with the
W^bl of the contraction— that if, with tho amount of xhortcniitg — but with
'•wnrit dtmek Aod this is measure*) by multiply inj; the tiiimber of units
LM^t to which tlw load a raised into the nuiiiber of nnitu of weight of
140
TUB C0NTBAC1
u
K
lOO
IM
IMC
ffiO
It
•
T
a
1
u
tJ
•W
Tl»
tw
«w
u
tbe load. Hence, it ia obvious fVom th« foregoing observatioaa that the wor
done moat be Inr^ly tlepciKleiii on tlie weight itwtf Thus, there b n certuin
wcijfht ijf lotid with whi<:!i, in any given luuule Btirauliit^l hv a i^veu
BtiiniiliiH, the incut work will lie done, aa may foo Men iVom ih« follnwiDg
examfile :
Ijnil. Ill eni.iiiiii«*
Haiubi or oonmcatm. In inlUIuiMna .
WwliU'JiM, III grmni'inllUniuLrai ....
i 8S. The ttijlumi^ 0/ Iht mxe and form of the mvtele. Since all liuown
mutciiliir liliTTfl arc much ehorier than the wavo-l?ngth of a contraction, it
is ohviouH ihsc (he lon^'«r the tilin? ihe grenter wit! bo ihe sborlening ciiu»ed
by the mine contnictiori wiive ; the i>realer will be the heighl of (he con-
Iraetion with the mtnv itiiitiiiluH. Hence, in n niuecle of pariillel tibree, the
hd^bt to which tlie louil in rained nii the result of a giren fttimulus applied
to Its nerve, will dejiend mi the tenj,'lh of ihe fil)r«e, while the nMisinium
weiirht of load <:ii|)ttbli< nf lining lilted will ilepend on (he number of the
fibres, ^imt the loud i.-< dintriliulcil among them, uf two uiusclee, tliereCore,
of equal length (iind of the mme nunlity) the ranet work will be dune by
that whirh hnn (bn lnrg<-r numlM^r ol librai ; that ia (o aar, the 6bree. being
of e>|iial width, which ha* thr ^reiitiT Hcctional area, iiud of two musclea
with eijtial MOlionnl arcu.". the niont work will he d<ioe by that which ia the
loufj^r. If the two muKch-K iirc unrcjiml both in length mid actional nrea,
the work done will hu th«; grcntor in the one wbith hiia the larger bulk,
whii'h contain" (he grvntcr niimher of cubic unilit. In xpi-aking. ihcrrfnre^
of the work which cnn Iw done by a muscle, we mny uki- il-> n Miindaril a
cubic unit of bulk : or, the dpccilic grnvity of the mu.idc being the Mimo, n
nnit of weight
Wo lourn, then, from (he foregoing paragraphs that the work done by a
niui'cte' nerve prepanilion will dei^end, not only on the nctivity nf the nern
and muscle as detenuined by iheir own irritability, but tiho ou the chnmcler
and mode of applteutlou of the Rlirnulus: on the kind of con traction (whether
8 single eiia^ni, or a sKiivly re)>eated or a rapitlly repe&led tetanna) o« (he
load itself; and 'ui the »mi and form of (he muscle. Taking (he iiioel Ihror-
able circuniNlanci-ti, vix., a well-nourikhed, lively preparation — a maximum
uiniuliiM caiuinga rnpid tetunux and an appropria(e load — we may deter
mint? the maxininni work done by a given weight of muscle. «ay one granirne.
Thift in thii i:nM! nf ihc niuHcU-i* of the frog has been estimated at about four
grani-nietre» fur one gramme of niuide.
Tin: CfUfUMHTASCKS WIIICEI DlH'KKMlSi; TItE DkUKKR OK IltltlTAUILITY
OF MVUCLKS ASU NeRVIM.
*i 83. A muscle- nerve pret>aration at the time that it is rcmored from the
body [KMUMaci a certain ilet;r^<e of irritability ; it rc^^ponds by a contrat-tiou
of a ct^rtain iiniouDt (o ■ atiniuluH of a certain slren^'lh applied to (he nerve
or to the muscle. After u nhilc^ the exact period, depeu<Iiag on a variety
(if cireumstaiiccD, (he xnine Ktimuluo pmdueei^ a smaller contniction ; ■'.«., the
irri(abili(y ol the prepanitiuu ha:> iliminlKlied. In other words, ibe muade
or nerve or both have become jiurtiiillv " exhausted," and the exliaustion
Buhwinently incrrn«ca, tbu luuiie stimuliin pnidiiciug smaller oun tract ious,
until lit last all irrlUibility i* l<»t, no ^timiilnn, however Ktroug, pniducing
any contraction, wlicihcr appliitl to the iKiri'e or din^itly to the muH-le ; tuid
eventually the muFcIc, tus wc have mi:.a, become* rigid. The pnign^to of lhi»
exhaustion it more rapid in the nerves than in the niusclca; for some time
iXflRIE OP IRRITABILITY OP MlfiCLES AND SKKVKS. 141
\
\
•the DtTW-tnink Km ctattd to raepoiid to even the «troD)i[fist slirnuluB.
tractioDi niftv Iw obuincd hj apptrinf; iho stimulue directly to tlie diiib-
«)& It ia much more rapid iti ihu wsnii-hlnuiled lliiin in lh« o<ild-l>liii>i]c<)
■ni— bi TIm muscica and nervi^a of tbu former low thi^ir irritability when
naDBTod from the body, tSter a period varyiog MCcorditiL' to droumntaiiocK,
ftwB a few iDiDuta to two or three boun; thoae of cold- blooded luiiniiilii
(or « Icut of an amphibian or a reptile) mav. onder f«vonblc conditiutiH,
ivautia irritable for two, three, or even more ilaya. Tbi; duralitui of irrita-
bilU; in warni-blaoded animaU m»r. however, be coiividerably pn>UMigod
bjr ivduciug the temperature of the Dody before death-
If with aocne thin \>odj a Bharfi blow be ttniclc acruM a miucle which linn
•(Htf«d lato the later *i(agw of (•shaumioa n wheal laalin^ Tor ai-veral *ecr>ii<U U
4«««loptd. Thu wheal appcani la bt- a cODtraction wave liinilcd iti tbi' pnrt
atrttck, aad diaappearing very «towly without extending to Ihc nciubbnriu^ luua-
•alar Mihetaitce. It haa been called an "ii/ta-inunfyar'' mutraclmn, because it
■af ha bmaghl out evea when ordinary Mtimoli liaTeceaaed to produce any elTcrt.
Il ■•¥. bowever. be accenpanted at it* bffpnning by an ordinary coutraction. It
b nMWr produce<l in the livinic body on the pectoral and other mtuclea of per-
ana anlMing from phthlab and other exliaujitirig diaeaaes.
Tht# natural exhauniou and diniinutioo of {rrilnbility in mii»c)e# and
■errea removed from the body may be modilied Imib in the rum of the aiue-
de and of the nerve by a variety of drcumnljinoe*. Similarly, while the
arrre aod muacle Mill remaio lu the body, the irritability of the one or of
th» olbcr may be inodified eiilier in tlie way of inerca^c or of decrcR^e by
eanain general influeucea, of which the niuat important are acveratice from
iheccDirmI nervous ayMem and variatiooa in tem|>enilun), in blood-mpply
tttd in functional activity.
n* rffft<» of tefmnct from the etntrai nerwuu tfftietn. When a nerve,
fudi, tor inauuifle, •• iJm aciatic, is divided in tUu. in the livins body, there
», flist of all. olMwrvod a alight incrcAH- of irritebiliiy, noticeable eapcciftlly
tmx llw eat end. but after a whilo the irritubiliiy dimtnbhee and gmdually
fcyHiWra. Uuth the slight initinl incrcaw and the Bub«(]uent dccreate
bpa •! the col ntd and advance cctitrifupnily toward the |Kiriphcral ter-
■ftalinmi This ccniriln^al fcuUire of the Iom of irrilidnlity ih ofti'ii spoken
<f M tbe Ritlcr-Valli law. In a mammal it may he tu<t or l\\TW ilavx : in
tfrog, aa maJiT.orevcu more week^. boliire irriiabilltr hiu duiapiicurcil friiro
the Derre-trunlt. It is mainlaine^I in tlie >imali laiid utpv-^cially in tbe intra-
nntcalar) braodHa fin* Mill loiifjer periixie.
Thb oentrifb^ Iocs of irritubility ia the forerunner in the i>eriphera]
wrtioo of ibe divided nerve of airuetural chanif^ nbich proceed in a sini-
W centril'uptl manner. The medulla Miflera ^uu))cea iumilar to thoae seen
fa Dcm llbrea aA«r removal from tbe body; Its double contour and its
ehanuitariatic indentations booDme mure marked; it breaks uti into small,
imgvlar fracmenta or drops. Mingled witJi the fat particles of the m4Nlulla
at aaan amaJl DUUi» of pnitcid material, which appear tii Ih- derived from
iha proloplaam around tlw nuclei. Meanwhile the axis-cylinder also brealti
Sinto IVai^nMtita, and the nuclei <>r the DMirili-mmu divide and multiply,
r bitty (-KUiitituenla wbaoquently dc^-reoM- in amount, tbe prolcid material
iacnaaiog or not dinnnishtne. aihI thus the contents of tbe neurilemma
htavan each two uodm is reduced to a mass of protrid malcrial, in which
tte f^agmeota of tbe aiisHjjliiider can no longer be reooniiaed. This man,
tbich sUl) rMaina sonia tat globules, is studd^ with nudei. If no regcner*
itioa tak<« place theeo nuclei, with ihfir proteid hc^l. eventually dinpjtear.
Id the cctttrul porttoo of the divided ucrvir similar cbangen may be tracetl
142
TU£ CONTRACTILE TISSUES.
M tttV only M tbe next node of Ranvier. Beyond thk the oerre unielly
remiiili* in H nunnul con il it ion.
RweueriiLiun, n'hi-ji it iircuni, iit njipiirvutlr cnrriM] ant hy lliv pcrijihvrnl
pvtwin of till! uxis-iyliuilrni of llii! iutjit't o<»itrul porlion. WIk'II tliv cul
endu of the aurw an* clmw top-thcr the iisiK-oylimlont grawingout froiD the
c^nlrnl [wrtion niii into uuA lii-twci'ii tliv *hnink(^ ncurilMQmBa of the
IwHphrnil |H>rlinii ; hut iniicli iin<'i>rliiiiily »tiU cxii>l» u to the enact nnrt«
wliicli the prolirrrnlcd n^^iiclfi iiml llic proleiil mntvrinl rcftirred to abore,
fltid tliv iiUI nxi^'L-ylinilcrB of the pcnphornl p.irlioii respeoiively play io
jfiviiic rieo tu the new stniclurc* of the rep'nprnlod tiltro.
Saon a degCiieTatiun mnr bo uWrved to ^xteiid down to the very endings
of the tierve ici l)ie mii§cle, iDeluding the end-plnte», but does uot at firsi
atfect the inuBL-iiltir eiib«tnDee il<«lf. The iiiiiscfe, ihouith it bad h«l all its
tien'oiiaelenieiiU, still reiiiaiDH irritahlo towani stimuli applied directly to
itMlf: an additional pruof of the existance of an independent inux-'uLar
irritability.
For aonie tiin« th« irrilnhility or tho mtuclo. n w«U toward utimuli applied
directly Ut ilnelf nc toward iboM- upplivtl ibrousb tbc iinpaiird ncrvu. •ceni* to bo
ditniiiiabed : but after u u'liile a pvL-uliHr i-i>iiiliiloa (to wliidi we bnve already
alluded, { 78)_ oeta in, in wliich Ihe muscle 1h found to bo oot eaally Bliinulaie<) by
single induction -shocks but to rtvpond ritndily to tlie innke or break of a eon»lant
current, [n Tact, it ia said lo bc<u)mv nvcn nxirc lenntive to tlio Utt«r mode of
utimulation than it vru when lis nerve vtHa iuUi<.-t luid KiDCtJonally acUve. .4t the
mme time it also beconiM more irritable toward direct mechanical dtiniuli. and
Tery frequently fibrillar contmciiunH. more or Iom rhythmic and apparenily of
Bpnntaneous origin, (hougli thc^ir cHUsntion is ohscure, moko their ap|>earanc«i
TliiR plinkc of hci);bleiie(l seiisilivcnvaa of u musde.MpEicially to the constant
current, nppenra to reach ii^ maximum lu man ui about tbe seveutb week alter
nervoun impulses have tensed, owing to [lyury lo tbe aervea or nerToiu centre to
mich the muscle.
If the tnusciv thus dcprivoil of ila nervous elements bo left to il«elf iia
irritability, however teetMl, woner or later diminishes; but if (be muscle be
iwri'idieally thrown into contractions by tirtilieinl stimulation nilh the oou-
Miuit current, the decline of irrilnhility and atlendnnl loss of nutritive power
inav be poel{>oned for some conniderable time. But as far as onr esperieuce
goce at present the arlitieial atinuilalion cannot fully replace the natural dae,
and 9u»uer or later Itie muscle like the nerve suHere degeneration, loeet^ all
irritability and uUimulely its place is taken by connective tissue.
§ 84. Tiir iiijiuenK of (r-mpenilurc. We have already seen ibai sudden
heat (and llie same might he said of cold when sufhciently intense), applied
1(1 a limti«<l )iartof a nerve or muscle, as when the nerve or muscle is touched
with H hut wire, will act as a stimulus. It is, however, much more difficult
to generate nerv<iuK or muscular impnUea l>y cxjKNting a whule nerve or
tDUscle tn a gradual rise <>{ lem[Jiirtiture. Thus, according to mott obflerras,
R nerve bolonifiug to a niimcle' may be either cooled to 0* ('. or below, or
heated to W C. or ercn 100° C, without discharging any n«rvoui> impulses,
u shown hy the nbeeucc of contraction iu the attaclicil tnuscle. The con-
tractions, niorvorer, may be alMint even wlicn the heating ba:< uot been very
IfTndual.
A muscle may be gradually cooled to 0° C. or below without any contrac-
tion bein;; caused ; but when it » heated lo a limit, which in tne case of
fr()|;'s musules is about 4.^^ C.. of mammalian muscles about 50" C, a sud-
den change lakes place : the muscle fulls at the limiting temperature into a
rigor mortis, which is initiated by a forcible contraction or at least sliortvning.
< T1iaBeUon<irnldui<llie*laiiM(uary utnvirLlltr«<ouiiid(n«(liaRl«Hrp(intotio(ilioir«fk.
^DKOIiri or ItRtTABtLlTY OT VTSCLSS ASV XERVKS.
I
M-xl<raic w«nntli, e. y., id the fro^- an lucmu of t«ni|Ji-m[i)ti.- u[i (u
•oOM'wbBt below -lo^ C, ftvufs bolli niuscular and nervniu irritnbiliiy. All
|(m molFCiiIar jtmcoees are ti«st«ne<l nitil fiidliuitnl : lli« oonirni-Uun it far
* ipveo ctimulut greater aiit) nvore rapid, >. r.. of i>hon<T (|uni(i»n, mm) iirr-
X"m impillw* are geiiented morv rauiily by »ltglil »tiriiuli. Owing to tlic
<|aklcrmitf; of the cbcmical vhango*. ibc viipiilj- of new niiucriiil may prnve
iiaoutlinrnt : benee muacl«4 and ttcrvM rcmovct from tliL* budy Wme thi'ir
irrilBtnlitr mun' ni|>idly iil n hi^h limn at a low t<'iii|H'rnlure.
Thr );r>(lual njijilicntuin of 4'uld tn a lu-rvi', f»|ic(-iiiUr wbcii tW tcnipera-
tarv i* ihiH bmuglii near t<i 0° C., >tnckcii« all lli« iiiulcnilur girocfMes, so
lltal (b« mm itf iH'n-iMis im^iulfo i» li-xci-ncil mid |iix>Ibn;;eiJ. llie vclocilT of
ita patimgr Inriti}; niucli >tiniiiiiiibci). e. y.. (ntta 'lH metren to 1 metre |)er MCotuI
At abiiui {)' C, ibr irritubililr of th« ni-rvc di«appear» allof^her.
Wfa<-u a mitM'le u vx\».v*a to similar fold, <: 9., lu a tern) tern lure very
lilltv alwiTe Kcro, llic eoiitrni:tiniiR are retnorkably prDlan;;ed : iliev ari;
(liniiiii'luHl in bright nt llbc f^me timei bul not iu proporlioii to tbe iiicreaae
itf tticir duration. ExjkimhI to a temperature ut'iero or b«[ow, niuM-ltn aioh
loHtllHr irritabiliiy, wilbotit. however, imderKotu^ ri);or raorlia. AlW aii
cspmirv of not inure titan a few eei-onda to a lemjienUurt.- not nm<-b below
ibry may be mtored by Kraduiil wanutb tu »u irriliilik cotidilion,
Uxiugb Ihey nuiy npitcar to bave been frown. Wlieii kc]>l froKvn,
r, for aonie few luinuio. or when exputed for n IfM linir tu tcnipcra-
tara* nf apveral degrees below iiero, ilietr irritability is ]K-riuaneiillv
itHmytd, WlM>n al\«r tbn Ihey arc thawed ihey arc at firvt »uppK-, am),
aa ve b«v« aeon, may )>e uituh to yiuld miiM-lu pliiMiii: but th<.'y ver>-
ipwdily enter into rig<ir mortis of u lunM i>i\inoiincc<] cbaracl«r.
1 8& Tlf I'nylii/Tiw 0/ biood-wupply. When a uiiaelo still within the hodv
b ityrixtii by iiity nioBD* of lt» proper blood-«upnly, m when the hloud-
vn«M gotag to it are ligntiirvd, tbe same i^adual loea of irrilabilily and
tul ■npfliiiii of rii-nr ntiirlw nre ohsen'ed as in muscles ivmoved froin
tb bodr. Thus if iKe alidoniinal iiurlu be ligntored, (hu miieelee of the
kntr linba !<«« their irrilitbilily and tinally become rigid. ^k> also in sys-
Hode dtatb, wheD the blood supply t<:i the miiacla is cut off by the oeasBtion
of fhe dmilation, loss of irritabiliiy eiieuu, luid rigor mortia evoDtuallv
r»|hiw*. In a human corpm the nuacla of tlie l>oily eiiU-r into rifpiT murii!'
A m 6xrd order ; Brsi, thoae of the jaw and neck, then ihusM: of the trunk.
MTt %hom of tbe anna, and la«tly those of the legs. Tbe rupiiliiy with
tUeh ripH- motib ctomes en ufter death varies counderahly, Wxojt deter-
■iaad tiotb by exiemal circumsluicea and by the iniemul i.»iii]i[ioa!> of the
Wt. Tbu*, exifrnal warmth hnMcua and cold retard* ibe tmm,*!. AlVr
(nat mtucular exertion, ns in hnnu^l auimal*. and when diiitb clowe^
•Mlin^ (li»*«»r», riiror morli^ in mort mm.« cumtv i>n rapidly. \t a i;encral
nit, ii ttiuv Ih- Miid that the Inter it i« in making it;! appearance the more
ffooouDcrrl il i», iind the longer it Instc: but there are many exception*.
ad wbeit the >tiUi* i» n>c-ogiiiM-d a« boiiig fundamenially duo I0 a clotting of
attain, it b eaar to undenlnml ihnt the amount of rigidity, ■'. f., the amount
i/tbe r)ot, ana the rapidity nf the onset. ■■ «., the uiiiekiiesB with nhich
>t«|palalioa tak« place, may vary independently. The rapidity of odh-I
i&cr mutcular eser^^iK and wasting disease may, tierhaps, be in iiarl dejM'ud-
won an iiH'Teaae of acid reaction, which ia produced tinder tnoee circum-
*M(a U) the muscle, for thia leema lo be favorttble to the coagulation uf
iWatutcIe plaama. When rigor mortis has once become ifaorougfaly ntab-
b)M in a muscle through de|)rivalion of blood, it cannot be removed by
uy tuh*eouetit supply of blood. Thus, where the abdomiital aorta ho*
itnuinrd llgaiurtu until tlie lower limbs have beconte completely rigid,
144
TUE CONTUACTILH: TldtJL'KS.
untying the lignliirc trill doi iv^tore tho miisolM to sn irritable coD<liti<ia:
it feiiuply hnshrns the ilecom posit iuii of the deud tiaeues by eiipplyiii>; ihetD
vilb uxygQa mid, in the ituic i>f thp mniDiDitl. wilh warmth aleu. A miucle,
however, mny nciiiirt- us ti whole a certaio nmoiiiit of rigidity on account wf
BOiueuf the fibres bpcoiiiing rigid, while the remainder, though (hey have
lott iheir irritiibilily. have not yet advancerl intii riji-ir iJiHrtii*, At sucli »
juncture ii renewiil of the bluod-siream may leaiore the irriluliiliiy of iltuM
lilires which were not yet rigid, and ihua appear to do awuy wilh rigor
mnrlia; yet, it appears that in sauh vanen Iht- tiliro which havr a<'tu(illy
become ri>rid never regain their irritability, but und^r^o diiR-neratiiui.
Mere liAi of irriiahilily, even thxiigh oumjiltrtf, if st<>[>pin)( *horl Df the
Actual coagulation of the muj^cle )<iilHitun»>. may be with care removed.
ThuD. if ■ Ktrcum of bUnvi he aent urtillcially through the vcffcU of » K-pn-
ratctl (mtniinulian) niiiM^le, the irrilnbibty niay Ih- imiintftincd for a vcrv
considomblc time. On Ht<>|)|)ing the artilicini circidntioii, the irritability
dimiiii«hp« ntid in lime entirely dinappi^rv; if, however, the eimm Ik^ at
once resumed, the irritability will Ih- recovered. By ivgulatiug the flow,
(he irril«hilily mny he lowered and (up to a certain limit) ruiicd at pkiistirt,
From iheeiioeb, however, of inlj'rfcrcnce with the normal b]o4xi-stivam thero
is a gradual dimimiliim in the rcsponaet to stimuli, and ultimately the mus-
cle KiBes all its irriuibilily iitid Mntmea rigid, however well the artitiiial
circiilalion be ki^pt up. This fnilure is probably in great part doe lu the
blood sent through the tissues not being in a perfectly normal L-undition ; but
we have at present very little information on this point. Indeed, with
re#])ect to the ^tiit/iVy of blocxl thus emeiitial to tlic maintenance or reAtore-
lion of irritability, our knowledge is definite with regard to one factor onir,
vii., the oxygen. If blood deprived of its onygen be sent through a muscle
itmoved from the body, irritability, so far from being maintained, ^eeius
mther to have it^ diiiappeanmce baHtened. In fact, il vem>u# bloud cun-
tinncs lo lie driven through u miuele, the irritubilitv of the muscle is lost
even more miiidly than in the entire abnenct! of hlooa. It would seem that
venous blootl ii* more injurious than mine at all. If exbauv^on b« not
carried loo far the muscle may, huwatver, be revived by a proper supply of
oxygenated blood.
Ine influence of blood-nipply cnnnot be so satisfactorily studied in the
oaae of neim aa in the ease ol hium^Itv ; there can, however, be little doubt
tliot tlie effects nre analogous.
§ 86. TItr iiijtwnrr- oi Jiinrtinnnl arliribj. This, too, is more easily studied
in (be case of musclcn than of nerves.
When a muscle within ihc body is unused, it wastes ; when used it (within
ccrtiiin limiU) grows. Both these facts show that the nutrition of a muscle
is favorably aflecled by its functional activity. I'art of this may be an
indirect effect of the increased blood-supply which occurs when a muitele
contnietA. When a nerve going to a mitscie ia stiiuulalcd, the blood vawelii
nf the muscle dilate. Hence at the time of the contraction mort bhiod
flows through (be muscle, and this increased How continues for some little
white ttA«r the uuntractioii of the muscle has ceased. But, apart from the
blood -supply, it is jirobable that the exhaustion caused hy a ooutraetioii is
imnteiliulely Ibtlowed by a reaction favorable to the nnlriuon of the muscle;
and this i* a roMii, [Mwihly the chief reason, why a muscle is iucre«kM>d by
use, that is to «ay, the ItMe of sulMtanoe and energy caused by the coiitnic-
lion is subHK|ueiitly more than mode for by ini-rensed metabolism during the
fallowing period of rest.
Whether (here be n third factor, whether mu»:les for instance arc govcrued
by >o-called trophic ncrx'ea which affect their nutrition directly in some other
I
I
*ay than hy [iiSu«Dcin)[ cither tli«ir blood-siippi/ or tbeir aetivity, aiuM m
wrai b* kn tiiiilf^-iitol.
A aimelc, «v>-ii wiiliin the body, after prolaoK«<l action i* fatigued, •'. it.,
A MrmMr aiiaiuliM ix ivquirerl to ptMliice th« sami.' cuiUnitrliuii: in other
woffik. Ua Irritabililv may Iw Itiwtiwsl by functional nativity. Wb(Hh«r
fuartjuDal nrtiviiy, lb<'n-liirr, i< iiijiiniHi* or biini^fu-iiil ilci|Hiiibi on iCM amoiiiil
in rvlaiifin to tb<' comlilion q( ihv muKck. It niiiv lii; here rcmurk^sl that tu
a ntiacla bM-otim more anil more (btigucd, #lininli of «hort duration, Mich as
leductioD-afaocks. looner h«c th«ir efficacy tlian do vtimuliofluiigdrdunitioD,
tucfa M ibe break and make nf Iht- c^niitaiit curn-nt.
IImi wnee of &tig»« of irhich, aRer iirolonged or unLieaal exortjon, we arc
nMMemiu in our own b<idi«a, ii prcibably of complex orieiti. and its Dalure,
hikm ihmt of the nonnnl luuBculnr ecitw of which we sliall bave to speak
hsvaAcr, ia al preaenl ool thoroiiffhly umkralood. It seems to be ia the firM
|d«ce tbe mult uf chaiigi^s in the iDUseles thoiDBclrGe. hut ia poauhly abo
ewmd bjr changes in the nerroua apparatus conwrDi?<l iu muat-ular nclioo,
aad ntMctally in thoae parta of tbe central n«r\'ou(i •j'sleni which are cod-
oncxl ia the productiou uf voluntary iiupulaea. [n any oaae it cannot be
ukrn na an ailoiuiite mciuure of tbe actual futisue of the muaclea ; for a
■an who -■•y* he i!i iibaolutely cshauMed niuy, iitiJi-r ext-iicm«[it, pcrforni a
my luge amiiuni of work uilh hit already weary uiuKtilcM. The vtill, in
hcU niralv if cvi^r calln fnitb the grcaUH conlractiiiuii of whivh the niuccl«
■n eapabfe.
Awolul* (tcntpornry) cxbauttion of the mii»clo#, to that the strongeat
■thauli product no coiilracitun, may be producnl even within the body by
uMtiaS *iitniilaii<Hi : nwxvry lakm p)ac« on rmU Out of the body abwluU)
tshttuKion lakca plan' n-udily. Hen.', alcr, itvovery may take place,
Vbalber in any given aa»e it does occur or nut. ia determiDed by the aiumtot
•d eoBinictMin atuing the exhaustion, and by the previous cnudilion of tbe
Made. In all cases recorery it basleool by renewal {natural or artificial)
«f iha bliHid-Alreain.
Tbt BKtR rapidly ihe eootractiaQS follow eat-h other, tltc lew the iiilorval
bitncn any two contractions, the more raiiid the exbauwiou. A certain
nabar of ainuli* inductioD-^ooks rcpeateJ rniiidly, say cverr second or
<Aanr, bring about •'simuMire Iom of irHlabiliiy more rapidly than the
«as anmbw of aboclu repealed 1i>m rapidly, fur liiNtanw ewry 5 or 10
wails, RcBM l«tanui ■• a ready mean.t uf pr>iihji'itig t^'x hiiuslion.
!• axfaaiiMn) mu»cl4!« the I'hutticicy in much diniinislit-d : the tired muscle
Miras Um n-ndily to Itn imiurnl U-uglh ihnn doc* tli<- I'mit one.
The pxhauslitxi du« t» cunt nu-lt<m may be ihi' result either— of tbe con-
■isMiitn of thr More nf ready ountrartilc material prcwnt <o tbe muscle;
<4Hi tbr Bccnmulatiun iu tbe tiMiicof the proclucUof ibeact of oontraciJoD;
«i «f both i)f these iwuet.
TW rfMomtive influence of rest, in the caw of a miisclo removed from tbe
limitation, may bt- explained by supposing that during the repoee. eitlier
tti Bieraal chan^ea of the tissue manufHcturc new exp!i)t>ivc matenal out of
Asopaiparalively raw material already pr«scni in the tibres, or the directly
hnfiil pnnluols of the act uf coniracliuu undergo changes hv which they
u» nMircrted into comjiAratively inert bmlies. A stream of fro>h blood
«7 rxcft its reef>rative inllueooe not only by quickening tbe above two
mat*, bui aim by oarryint: nfT the immediate waaie products while at the
«M« linM II briitn now raw material, li in not known to what extent each
•tthesa part* i* tOaynl. Thai iIh- proilitcta of contraction are exhauatiuK in
tUroKcts, U sliown by the imiu that the iiiJM?ti<in of a a<dution <>( the
aatdMXtraclivea into tbe vcwvls of a muscle produce* exhaustion, and that
10
THK CONTRACTILS TlSgtTBS.
vxhaiisied inuBcl«* are iwovfrwl br the simple injection of inert snline
foluttoue iiitn tUuir bloo<lve«ols. But tb« matter hw Doi ytl been fullr
worked out.
One iiuix)rtnnt clement brought by freab blood b oi^emi. This, u we
have seen, i» mil ncc?<«Niry for llie tarrying iml f,{ Uie acturiTcAnmction, awj
tct n et^e'iitinl to tb« maintciiaiice of irritiibility. Tito nxTgvn absorbed
tnr Che mitn'k- npjiiireDtly ent«n in tome itet-uliurwuy into tlie fornmtiou uf
tnat oomplvx irxploaivo niaterinl the de<^uiii[iiuiti(in'<>r which in the act of
oontiaotioii. th<iiiph it jjives rise to caHxinic acid iind uthur piwduou of
osidndoD, u Dot in ilwlf a pmcees nf direct oxidation.
Thi: KxEBiiY OP Mt.>ot.i: ami Kkhvk. and the Natuhe or
MlW-ULAR AND N'ERrotm AcriuK.
§ 82. We may briefly revapitulaie »ori« of tho chief results arrived at iii
thfi preoi'iling pngeii im followii :
A inimciilur iimlraotinn ibH-tl' is MMiitially a tnu»loeation of moleoulm. n
change of Ibrm, not of bulk. We cannot my. however, nnyibintc defliiiir &•
to tlie nature of thi« triin«l(>cjition or lu to tlio way in nbich it h bnmehi
about. For in»tiinec>vc cannot stiti«riii-Ioriiyexplnin the eoDiieetinii U-ttiivD
the Htnation of n miisc-ular fibre niid u mtiiteiilur contTttetiou. Xritrlr all
rapidly contracting musclee lire strinled. itnd we must aupixve tlint the
BtriaCion m of Bonie use ; but it is not eweiiliHl to the carrying out of a eou-
trnttion. for. as we shall see, the cunlraclion of a nun-Atrinled mtiacle is
fumUiineutally the Manu' ti» that of a tiiriated iiiti«;le. Hut whatever be the
exact nny in ivhioh the tiant^ioeation a ejected, it U in w^me way or other
the result of n chemical cliango, (if an cxplnMv*! deconijKNiition of cenain
part« of the t]iii>ck- KiilMlancc. Tho vni'rg>' which is expended in the
niechaDiral work done by the muscle hiu< Itx Knurce in the eoerf-y latent in
the muscle substance and set free by thai cxplonioii. Concemiiif; the tuHiire
of that explosion we only know nl preecnt that it rmulis in the producli'jn
of earbonie scid mid in an increase of ihc acid rcai'tinn, and lliai beat ii> set
fVee.afi well as the specitic muscular energy. There la s general pnnillelism
belwceu the extent of metabotiani uihiug pisce and the amouul ot eDer;gy act
free; the greater the development of carbonic acid, the lai^r b tliccan-
Iraction and the higher tlie temperature.
It i» important to remcnitier thal.iui we have already uived, relnxalioo, the
return to llie orfgiual k-nglli, i.n an ciiAeutial |>ari ol' the wnole cuntrn'tion no
Ins thnn ihe yhdripiiing itjtelf. It Ik true that llie return to thv original
Uinglb i]> H»»i9ti.-d by lh<- afri'lohiiig exerli-d bv the load, and in the cww nf
aiUBclcs within the living budy ** nci'iired 1>y the action of aiitOgoniMie
muscles or b^' vnrioux nnatomical relalidUH ; but the fact that the cotDptete*
ncMand rapidity uf the return arc ilopciident on tlie cmiditian of the muscle,
that is, on tne complex chuigt^ within the nnucle making up wbit we call
its nutrition, the tired muscle retuxiug much more slowly than the untireil
muscle, shows that the rnl&xntion is due in Ihc niaiu tii intrinsic proceMU
going on in ihe muscle itself, proccsecs which n'e might chnmcteriie aa the
reverse of lh«e of contraction. lu fact, to )>ut the matter Ibrcibly. adopting
the it lustration used in g57, and regarding rclaxnliomutt change of molecult*
(Vwm a " formatidU " uf one hundred in two lines of fifty each lo a formation
of t«n columns each ten deep, it would be possible lo sup|>ort the tbesit that
the rrally active foreo in muscle arc those striving to maintain the latter
formation in columns, and thai the falling into iloubtc lines, that ts to aav
the pontraetion, is Ihe mult of these forces ceasing to act ; in other wortf$
J
TUB BNKHOr OP III^CI.K AND XERVE.
14;
Umi iIm mMiinuMd atotc of tlw miiftcvlnr fibn> U what iokv hccnilod Um
wUural lUte, tli«t ihc rclaxcl t-otulitUiti i* ntily hrou^'hl a\x>in »I tbn cxpctiw
of ckaogM cnuHlcratrtiii); lite nnliirni [ondt'iicice of l}ic tiltro. Wilhout gofn^
■D Ihr M tliis, bowi-ver, we may still rpco||;niz« th&l iwth cuiilnwtion nnd
nlazsUMi era the rmilt uf chnn^n which, niiof ihcy sccin to be of a cheitiicnl
BHun in ibc iiiiv <.'iu«, nrr pmlMibly «o it) the other aJau. A>i<l thoUf;h in the
iafai^Ma nf exact knowk<d);e il in (laiigerotis to speculnte, ne way iiuncioe
thai ihtOT chemical events leiuliog to relaxation or elongation w of an
ofio^U ttt BOtafonUtic cluinii-ter li> ihoae whoee issue ia coDtractiou.
It luH not boen possible hitherto to draw up a complete equalioa between
iW lalcBt enericy of the nislerial and the two fbrnis of actual enei^ set
frw. bwU anil luovenM^uU I1ie pro^wrtioa of enerji^y Kiveu out aa lieat to
that taking on tJi« form of work varies under different circuniiitauc«a : and
it w<wM Bppou- tliiit iiu ihe whole a niuM^lc wuuld nut be much more i;lBoieiit
tkaa a ttcam-eugiue in nwgiecl to the ronvenUm of rhcmical uctiuu into
BM^aaiod work, were it not that in wann-hlocidetl aniiutilH tlie liMt );iven
«nt b Dot, ai in tli« 8teaiu-eui;iii«. in«rc Iom, hut by k>'i>pii)g iiji tht- Miimal
KapcmoK nrrea tnanj auhttiiliarr purpuM*. It might Ix' iiup|>tNH.Hl that ill
aoMUfVcUoD by whidi work a actually doi», as RomptrttI with the Mow
oaMtaetfcHi wh«n uo work i* doue. ibm >« a diminuiion of the increuM of
tuipmturr (-urr«w|ii>ndlng to tli« timouDl of work d<>n<'. that i» to »y. that
Ik* BWchonit-al work i* ili<ne at lb*- vxpcnM of energy which olherwis)- would
ft D«it a* boat. ProWhk- as this nuiy ttvn, it has nut yot been expvri-
SMMaJly verilWd.
Of lh« exact Datiin* of the chemical chan^-^ which underlie a mutcular
naUactkui we know viyry little, the must importaol fact bein^;. thai the ooii-
indjoa m not th« outcome of a dirccl oxidation, hut the splitting up or
tzplaitve d«ooinpontioi) of some complex suiiMaiice or Auhttaocea. The
■Mil* dott ooDHine oxyi^n, and the pmducttt of muscular awtabolism are
btW nd prodncU of oxidation, but the oxy){i>M ap|>eara to be introduced
Kt It tiie niutnent of expl<itii<iu but ai some earlier date. As to the leal
•Mm of th» exploBire material ne are as vet in the dark : we An uui know
It (ertain whellier we ought to R-fcui^l it bh a sin^cli; suWtaiice ' in the
ctmieal tenae) or as a mixture of more «ub*(Aiic» than one. We may,
tnvvtr. pvrbapi be allowed pruvitionally to speak of it at all event* a* a
ai|tk Mbataooe and to call it " cxmlraciile aiaterial," or we may ndo|>t a
Mm vUeh hat lieen suggettei) and aill it inayn.
W« iImII hare ooowon to i>i>int out taler on. that the hving stitwtance of
ante nil* it abb lo aiaoutactiire and lo lodi^ in the tubetaiice of the ocU
aWtaly eooakiarabk qiiaiitilicii at lat whcichy the cell becomn a fat cell.
ibhl ao Jbrmed and lo<)gcd being sub«c<(uently by some means or other
fcthafgnl from tliecell. Wo shall also have occasion to point out that in a
■—■hit siuilar way the living material of certain gland cells nuiuulactum
■■I 1m)(M id itacif ourtain nubetanece which, when the cell "secr«t«»,"
taimfi man or leas change and are ejected frum the cell. Theae suhataocea
Mar to be produolaof the activity of llie living subataoce of the cell, and
tobf *o telatod to that living substance that, though discontinuous with it
oi urrrly lodged in it they are still capable of beiug mi iuHuenccd by it as
h aaikiyo change more or teea aud<Wii. more or luu profound. And we
^,ttao^oo theaaalof^of thenfiucelhiaud gland cells.auppnee that the
hncnbaUuee of the muade roanufacturca and ludgtu iu itself this con-
taw* taaMrlal or inggen which ia capoblo of lK<ing »» iidluenoed by the
liiiiy aubMaDco aj to ui>dergo an «xp»(«ive decomiNMittou. But we here
•Mwltli a difficulty.
■ he tniMcular tlhre aa a whole in emineotly a niiro^cenouit proteid body :
148
THB CONTBACTILB TISSUES.
th« rniiiiculiir fibres of tfiv hoily riirni tlio greater [lurL uf the wliolc pmicid
BUUK of the body. MiinHiv«r ihi; onltnnrj ooiitiiiutMl mcUboliMa of tb«
roiiKulnr fibrv m ■ whole is «»i-nimllv ii uilrogciiotts meubolisiD : m wv
Bbflll baveto point out luter on, th« muscles uiidoubiodljsupplyagrent i>art
of that Urge oitrogenous waste which appears in the urine as iin^: th«
nitm);eiioug metaboliitm of the muscle during the twenty-four hour« must
lherffi.>ri? be con^idernble, and under ceriflin circumetaii(;ea, as for inMUDce
during fever, this nitroceiiouB metsboligm may be still fbrther larftely
iiici-ciified.
On the other band, as we have already nhf>wn, there can tte no doubt lliat
the act of L-outraciiou, the explosive decouipoaitiun of the tiio)^ii. iloe* not
in«reaBe the nitro^uous metabolism of the muscle. 8hall we oonclude ihcn
thai the inogeti ih ofiseutially a noo-nitroftenouB body lod^l in the nitn>'
nQouf miiaJle ^ubniaucey Not ouly have we no positive evidi^icc uf thb,
out the mmlugy lietween tontraelion ami ri|{or mortis is directly »p[>o>eil to
lucl) a vi«iv ; for it U aliu<>«t impimtble to mist the voncluiiiou tluil thv >tu6r
which givcjt riiw to the niyiwin I'lot, thu carbonic acid, and lactic acid or
Ptlier Bcid- producing viibntaHOM of ri^or mortis, iii the same vliilT which
giTC* rise to thi- trarbonic acid and Int-tic acid or otlicr add- producing sub-
stances of n contraction. The dittcrynco Imwwn the two sc«ms to bo that in
tli« conlractioii the nitrogenous product of the decomposition of the inojgMi
does not ap|)car as mlid myosin, hut anumes the form of some soluble
proteid. The imimrtant fact conceniing the two aclx. Hgor mortis and con-
traction, is that, whilti the gn-at non- nitrogenous proiluct of the dccon posit ion
of the inogen, m. carbonic acid, is simple waste matter containing no energy,
fit only to be cast out of the l>ody nt once fand the same is nearly true of th*
other non-nitrogenous product, lai'tic acid ). the nitrogenous produet being a
proteid is '.till a body containing much onei^iy. whioh in the cu»e of tin* living
muKcle may aAcr the contraction lie utilized by the mu^dc itmdf or, being
carried awav into the blood -stream, bv ff>me other [larta of the body.
But if ihU view be correct the ordinary melaboli.->ni gning on while ih«
muscle in al re&t must diflbr in kind an well aa. and |X'rha|i» more thiin. in
degree fn>m the nietftlwIiHin of contraction ; for thn furiner, a* wt have just
nid, is (vM-ntially a nitrogenous luetabolisju largely contributing to tlw niiro
genoiis waste of the bo(iy at large.
Whether in the muscle at n-wt this nitrogenous mctalxdism it confined to
that part of the inuncle in which the iimgen in lodged and docs not involve
tlio inogcn itself. »t whether the iiio]^n an well as the n»l of the fibre (inder
goes metahotism when the mnnclc is at rest, going olf in puflv, ao to apeak,
instead of in a largo explosion, its nitrogcnouo factore being at tliesame time
involved in the change, are •[ueetions u.*liich we cannot at prcoent settle.
$ 88. While in muscle the chemical events are so pmminent thai we
cannot help coneidoring a muscular contraction lo be ctwcntially a rbemical
nroceas, with electrical changes as attendant phenomena only, the ease in dif-
HKtit witli nerves. Here the electrical phenomena completely overabadow
the chemical. Our knowledge of the chemistry of nerves is at preeent of the
tcaiilieat, and the little we know as to the eheudcal chaii]t;es of nervous sub-
•tanoe b gitini-il bv the study of the central nervous or^ns rather lliau of
the nerve*. We iioil that the JrHlahility of the former is closely dependeni
on an atlurpiate lupply uf oxygen, and we may infer from thin that in nervoua
as in musciilamubataiiceauibtabolinm.orin the main an oicidalive character,
b tlw real cause of the development of L'nergy ; and the axi^-cylinder, which
as we have seen is matt probably the active element uf a nerve-fibn.', na*
donblcdly resembles in many of iu eheniiual featurc:> the substance of a
muscular Rbre. But we Imvc as yet no satiifactory experimental cvidctncc
OS AOUK OTHKB rOKMfl OF COXTHACTILE TISStTB. 149
the pMMf^ of • nervous inipul»e aIoiik a oerve b th« rwull, like iht
eootncckn of a maBculBrfibre.orcl>einicalcJmDgt«,aii<l like it ncooinpuikd
bjr aa crolntioD of beaL On the other hand, the electric pbenonieiui arc no
pVDiniiM-nt that Aonie have bceu ieiii|iie<l to reftHnl u iiervuiiii impiilxo lu
■aaninllv an Hwtrical chaugo. But it ihukI be reniomhertyl i)inl tbi.- uotiml
•oamr IM ft«e in a utm'uua iinpiilao if, wi to njiciik, inHigDificuDt, no tlmt
chllWiul fihaaita ttiu iiliichl to be n-oignixc)! b_v tn<? incnriM at pitaoot at our
di^jMil MtniM limply Muflire to pmvitle all (he i-iH-rgv net Im*. On thv otlivr
imiui. the rnlc of' Imiutniiwion of a nervouo impiilHi, puttine aeidp other
telBraa, M aJoot aufllcim to prorc that it is nomelhiiiK quite djflercmt from
M onliiMTT electric cum?«t.
Tbr ininiiu* ilt*po«iliun of ihu cml-platoi, an<l their rcmnrkable annlogr
with the elvctric organ* which arc foiiDd in certain animalB, haa suggc^tM
th* new that Ihr jNUMige »t' a nvr^-oui impulse from the nerve fibre iDto the
nuuvalar mbitanoo ie •>(' the nature of an electric (lisebarge. But ibeM
■Mttra are too difficult and too abetrufie to bo discuawd here.
tt nay, however. )>« worlh nhilo to remind th« render thiit in every cod-
tnrtioDof a luurcuiar libro. the actual change of lurm U preccdeil by invisible
tlaagn pruiaigat«cl all over the libro and occupyin)* the laleiil j>eriod. aod
llal tbci* fBangea reaenibie in their fcaturee the nervous iiupuUe of whiob
tbrr are. fo lo fpeak. ihe eoutinualion rntlier than the coniraetion of which
t^y nn* the forerunu«rB and lo which they give rise. So that a muacle.evei)
fMuiif aaid« the risible i«miinatious of Ihe nerve, b AiudsmeDtaliy a
■Mill sad a ncm bMidaa.
Oa SoHB OriiER Forms or Coktractilk Tib»vr.
Jlain, Smcolh or UnttriaUil Mtumtar Tunie.
t W. This, in vertebrates at nil ovcnt«, rar«Iy occurs in isolated maoseaor
aaRba, aa dam striated muscular tiwue, but is usnally found laking pari In
litMniolure of complex orifabs. such for iiietanee as the intestines ; hence
tW inveMi|[atii>n of its properties i^ beset with cuaiiy diflicukiea.
It ii mually arranged in sbeels, composed of flattened bundlca or bauda
h«ad InKBtlwr by connective liiHue earrying hloodvea^eU, lympbatiua, and
tmn. SDiDe of these buudles or baD<ls may be split up into ^inallor batid*
aailarlr untied to each other by o'ninective listiue, but in inaiiv cnM« thu
•Ua ^oet being ibiu is made ui> directly nl' small bauds, h^ach tniall
imi is oompowii of a ouniber of elenieiiiary fibre* nr tilire i-elU, nhidi in a
nnaiti aetiac ar« analofroui to the atrial^ elementary iibr(r:<, hut in many
Mieetf diOer uldcly fmrn tbein.
Eadi uiWrtaleil tlinneDturv fibre is a luiuute i>bject, from 60 „ to '200 ^ in
impk and fmru A ^ to 10 /• tn breadth : it is therrforci. iti «iz«. of a wlmlly
wnmt nnler froB) a Ktriatod fibre. [Fig- 4.S.I It is futiform or epindle-
^pr<l, •otnvwhal fiatli-iKil in the mi<ldTr ami tapering to a jxiint at the
Md*. ahich In vume <■■■««> are branclivd; but the exact form of the (ibrc
aiii (UHor according as the muscle is in a Mate of oootraction or relusation.
Hidirny between tbo two cndo and in the centre of the fusiform body lies
■ uelaiu. which in a numml randition is elliptical in outline, with its long
■SB Inni; lengthwiie. but which under the influenoe of reogeala ia vary
^^ Uttobeeowe rod-shaped ; hence in prepared specimens the pnseooe of Iheae
b Mifcaped nuclei u very cbaracteTctttic of plain mu^ular tiauie.
H 1W Buoleus baa the ordinary characters of a nucleus, and rery fre-
H fMMly two nucleoli are coospicuouo. Around the nuoleui b gather«<l a
TKB eo;fTBACTII.B TIflSOtS.
1K10.O.
/U
UuMiK Akirhw: I. fn>m ihv
pupQIMl arlorr ; <t.Bllhniit ; b.
wllti Rcollc Mid. i;frani * loanch
or thvaalvrinMIbUl ; a, iiucJoi of
(be Bbni (mwdUlwl SM llniei I, |
nmll '(onutiiy of grnnulnr jirotuplajiin, like tlmt around the niiolei of k
Uintetl fibre, and this u C(>(iliiiut.-il alori>: the uxU of the fibre for somo
(Ibtlimce from vticb jHtle of the nucleufl, gnid-
tinlly (n]>«riiig uwav. and so formint; a Blender
grniiiilnr crn-v in the median portioD of the fibre.
Th<- n»i iif tb« fibre, fi>nnin)( its chief iNirt, is
cumjiuMd uf II tnuiHiiurcnl but sonteulinl refrucrlive
RuWtitnoe, whieh i:< either homo^ciicouiiurexhibiia
n delicate loiigttiidionl libnllutinn ; tliiit i» tiit
muRclo •uljBtnntc of the fibre nml <;orni«fK>n>li> lo
ttiG ti>u«cl« eiitwtnncc of lh« 9triiili.>d librv, btit U
not striateil. Sometimct the whole fibre it tbrofrn
into n »erice of tr«ni»vprw wrinklw, wbicb give
it a eirinted appcnrnnco, but this in n very dif-
ferent Btrinlion from thiit producml bj an alterna-
tion of dim anil bright bandi. No »m'h allerna-
doD of ban<l» if lo be feen in tho plain miiBciilnr
fibre; the whole of the eubelance of the fibre
around the nucleus and cure is homntfeneoua. or
at leaat exbibiis no diflereutialion beyond that
into fibrillie and intertlbrillar xubstance, and even
this distinction ia doubtful.
The fibre has a sharfi clear outline but is not
limited by any diiitinct sheath comeapondiug to
the sarcolemma, at least according to mo«t ob-
ftervurs.
It ia obrioue that the plain muscular Bbre i» a nucleated cell, the oell oub-
stance of which has become diRcreDtialed into contractile sub«tauce. the cell
otherwise being but slightly changed : whcrNU the niiich larger slrialcl fibre
is either a nutnlier »t' cclU fu>-ed together or a ucll which has undergone
multiplication in i>o far thai its nucleus hn» given rise to several nuclei, but
in trhich no diri»ion of cell »ul»lance has taken place.
A number of such fuHifonn nucWied cells or libree or fibre cells are
united logellicr, not by connective tissue but by a |>eculiar proleid cement
aabnaooe into a tial band or bundle, the laperinj; end uf one Bbre dovetail-
ing in between the bodies of otiier Hbre^. 8u lun|{ as this cement eubsuinee
is intact it ia very ditiiculi tu i^nlate an individual fibre, but various reagents
will dttoolve or la^iwn ihi.H ceiueni. and then the fibres se|}arate.
Small flat bandA tliu« formeil of fibres oeoiented together are variously
amneed by mciiiia of cunnective tissue, sometimes into a plexus, sometimes
into thicker birgt-r bands, which in turn may be bound uji. as we have luiid.
into shccl» of varying thicknoM.
In the pIfxuB, of cnunn', the lumda run in various directions, but in tlic
Rhe«ls or meinbruiio' they follow i<>r l)ii> most part the sntiie direction, and n
thin tninsvciw section of a winwwhnt thick sheet presents a number of
smaller or larger areas, oorrt«ponding to the smaller or larger bands which
are e»it acroe«. The limits ol each arcii arc more or less clearly drlim-fl by
the connectire liseiic in which bluodvcsietB may bo seen, the nrcji itself being
mnpoeei) of a number of oval iuiltiii»i, the soctionis of l.hc flattened indi-
vidoftl fibree; in hardened specimens the outlines may from mutual pmaure
appear polygonal. In the centre of some of these sections of fibres the
nucleus may be seen, but it will, of course, he abseut from those fibres in
which the plane of section has passed eitlier above or below the nucleus.
When a tliin sheet of plain muscle is spread out or leased oui under the
mierovrope, the baiKls may also be recoguixed. and at the torn ends of some
ox SOUK OTHER FORU3 OF CONTKAOTILK TIASUU. 1A1
I
of Ifcc Iwncb tlie imlivUlual fibrec nur Iw >ceu prujcctiag nder the fashion of
BloodnMeh mud lymphuioi are nrriwi by lh« conDoctive tinuc, uid form
owitUry networks and Ivmtihatic pWxu»n> round (h« ainiillvr bands.
1 M. Thtt anaiig«aM-(if <■/ llw norvm it) unmriatcd muscle difRtra from ihat
in (tfiaivd muncJe. WhcnMic in Mrinlci) miudc mcdiillated tibree ooiuin);
dina tnta tltv anterior mnUi nf #{)4nai nerves iiredominale. in plain muitcle
OW ■xdllUaUd fibrv« urc iD<wt abuodnul ; in fact the nervw gaiDg to plain
OismIm Mv not only smnll but are nlinoet exclufiively (.■omtmaed of uvii*
nHdollMad fibfee aud conic to iIr- miisclefl frum tbe Bo-cul]e<i lyntpaLlieiic
mum. Pawng ■ut') the coanectivc tiwuc betweftn the bundlea th« nerve*)
•iiiidt>aDd,ji>iiiin)^ ngmin, fonn b plexus around tbe buudie«; thai is tAiuiy, a
t«ig conxiftiHK of a few, or perhaps only one axis-vvliuder, coming from
bfaocb will mn alougside of or join a itiniilar «mall twig »>miiig from
Itmich ; ibe individual ax ia-cy linden, however, d» noi thcjaiwlvM
Kriim sut-b prinmry plexuiu, in which u few mrdulbiuul tibrw
H« pnaeol amoD); tbe iion-miKliillaied tibi^, arc given ofl'Mill finer, " inler-
Mfiluie" plexuses oiiwtAliag excliisivclv of nun -mcdiillaiod f;brc« : thm
fb»»ee ths smaller bundle* nf rou»ctifar fibre*. The branches of those
flkmmm majr oonaiat of a lingk axi»-cylinil<-r, or may even be (ilamenla cor-
Mpoadins to several or a few only of the librillie nf which an axis-cylinder
i> Mppneeo tn be coAiposm). From thwc intvrmwiiacc plexuses ore ;>iven ofl*
■i(ie (brills or vcrv small tiiiodlt* of Rbnllir, which running; iu tbe ceraeni
■fafance between ibv iti<lividual libru form n Kiie network around ilie indi-
titaal 6bn», which network dilfers frcini the plexuses just spitktu of, ina.4-
nneh M sotuc of tli« fil»nivnl« composing it nppear to o<ialeK«. The ultimate
MlbiK of this network has not yet been conclusively traced ; but it *eem>
■ntaible that fibrils from live network lenuiuate in imiall knobf or oirellings
tfiaf en tbe subsUuice uf the muM-ular l!breii, somewhat Jtdi-r the fn»hion of
■imile end-plfllea.
A timilar temiiDation uf nerven in a pk'xiu or network is met with in
mW tisaues, and is not cimtined to uun-ni«dullaud fibre). A mc^liillated
fln may end in a plexus, and when it does ao loses Jinit its medulU and
nAis^flentiy it* neurilemma, the picxu* becoming uUinint<tIy like (hat formed
ha Dun-mtdullntcil fibre nml con»i.'>liug of attcnusictt axis-cyiinilers with
uiokmin^ nnd Htmrliniiv with wuctci. at the nodal points.
itl. As far n« we kimw, plain nui »cii hi r tissue in its chetuical features
nmaliln strinlt^ muscular tiwuc. It cinltiins nibumin. Aoine formn of
dthslio. and anieeeduts of myosin which u|>on tlie death of the fibres
Mine mronn ; for plain muscular tissue after death heoornes ri^'id. losing
h WenribilitT and probably becoming' acid, ihouj^b the aetttity \* nut M
■vfcnl as in ftrialcd muscle. Kreatin has also been found, m well as
fl^fogva, and, indeed, it swais probable that the ftbole metabolisoi of ptab
nmtar tisaue is fundanwntally tbe same as that of the tttriated mueoka.
t n. In tlieir {general physioal feaUiru plain muscular fibres nko reeemble
^risioi fibres, aiw) like tneni they are irritable and contracltlc : when sUinu-
htol ibey riinlraet. Tbe fibres vary in natural length in ditforont •ituations,
ikss of the bloddveiMla, for iiwtjiuce, being shorter and mouivr than thoM
W the intmtiu« ; but in iW name vituation the fibres may also he found in
*• af two diflereul rvMiditions. In the one ouc the tibna are long and thin,
h lh« Dthrr onse they arr reducr<l in length, it may he to one-half or even
*f •ae-thirr), ami are c«)rr«wpoudingly tliioker. hroiuler, and Ins poinicxt nt
Asiadf, their total bulk remniuiiit; unBlCered. In the former case they are
nknd or elongated, in ihe inlter caw tbey are conlractctl.
Til* bets of the ootilractioD of plain muscular tissue may be studieal in
lo2
TBB OO.VTKACTILIC TtHSUI
tin inUMin«, tbe miiacular coat of which coiiHists of an uttter thin aheel
oompoaed of fibrea ajid bundla of tibree di9jioi«<i lonfptuiliiiiillv. ninl of on
iunerniucb iliicker sheet of flbres dispoBe^l uircularlir; m the uratera similar
«rnui]^tii<.'rii of two coatA oblaiu^
If a mechaDJcal or electrical (or, iiidted. imv uiher) ftiniuluK he braufiht
to bear <mi a [tan of a fre^h. living, still wurm iiiluiliiie (the >mall iuKwIim:
is th? Kitt to work wilh) a circular cijriU'ticiiiiii i» Mxn to take place nt the
»IKit iiliniulated: tlic iniisliiie »eeiu» nifiiHMl in ringwJM;, n» if tied miind wilh
an inviitiblf coni. mid llie [larl »ii CDiistriitcd, previously vascular and red,
becoueB pnlu and bl<KiilIra». Thr individual librcs of ibe cm-ular oooil in
the region stimnliited have ruch htnyime :<h()rtcr, nod the total effeei of ibe
BhorteniD); of the mullitude of fibres all having the same circular di8|>06iti«ia
is to coDiilrict or narrow the lumen or lube af the ioteetine. The longitudi-
nallv dispoiied fibres of the outer lon);i[uiliDat coat will at the eame time
aimilarlv contract ui i^horl«D in a loDjfitiidtnal ilirecliitn. but (his coal Iwio);
relatively much lliiutier than tbe circular coat, the lonKiludiiial coolractiou
in allojiitfiher oveti*hadowed by the circular cunUacliun. A similar mode of
»iniracti<m u also aeeu uheu the ureter is oimilarly stimulated.
Tliv contrnction thuit induced in preceded by a very Iniix latent (teriod and
lu»t> a very coniiidi^rnhle time, iu fact, several seooiitb, after which relaxa-
tiiin (lowly takni jilace. We may »ay tlieo thnt over die Hrculnrly diitpor
fibres of tbe iiitentiiie (or ureter) at the spot in queittioii iheri' lins pawcdj
conlmction-wavc remarkable, fur it« long lateJit tn-riod nnd for the tlowiii
of it* dovcl"[)mcnt. the wave being propH^tcd Jrom fibre to fibra. From
tbe »pot so directly stimulatnl the cx»ntraetion may piw also as a wave (with
a len^'lb of I cm. and a velocity of from 'iV to ■in millimetres a second in
the ureter), along tbe circular cont both iipaard* and downward*. The
loofptudinal 6bres at the sput stimulatefi ari>, ns we have nid, also thrown
into contractions of altogether ainiilnr character, and a wave of contraction
war tbiu also travel longitudinally along the longitudiual coal butb upwards
and ilowun-ania. It is evident, liuwever. tluit the nave of contraction, of
which we are now ii|)eai(iiig. u. in one rt^jieet, dillereut from tbe wave of
ciintraetiou lrcate<I of in dealing with tilriaietl miUK'le. In the latter i'AM
the ci>ntraet ion- wave i.-> a sinijile wave propa^iUid along the individual fibre,
niid starting from tbe I'mt-philo or, in the ciim> of din-ct stimulation, from
the ])art of the fibn- first ailectiil by the otimtilus ; wc have no i-vidcnc*^ that
the c-untniction of one film- can communicate conlractinn to neighboring
6bres, or, indeed, in nnv way inHucnce neighboring fibres. In iho case of
tlM inUAine or urel«r tltc wave w complex, being the sum of the cuntmcti<Hi-
WSTCS of several fibres engaged in <liliercnl phases, and is propagated from
flbre to fibre, both in the direction of the fibres, an when the whole circum-
ference of the inteetiue is engaged in tbe contraction, or wlien the wave
travels longitudinally along the longitudinal coal, and also in a direction al
right angles t<i the axes of the librea. as when the con tract ion -wave travels
lengthways along the circular coat of the intestine, or when it paaies acrots
u breadth of lite longitudinal coat ; that is to say. the changes leading to
Ooniraction aiv communicated tint only in a dire<a manner across the cement
■ubslance uniting ihir fibri-s nf a liundtr, hut also in an indirect mantKr,
probably bv nieaimof nervi- fibri^i (Vom bundle to bundle acniwt the connvctive
tissue between them. Moreover, it is obvious that even tbe oont ruction -wa\-v
whidi i}n»^ iilorig a single unslriatcd Khre dificrs from that pamiiig alone a
frtrialed Kbrv, in the very great length both of its latent jieriod and of the
duration of its contraction. lieiice, much more •.■VBit than in the ciue of ■
Striated muicle. the whole of each fibre must be occupied by the contraction-
A
ox SOMK OTHER rOKUa OF CONTRACTII.B TI&HI'E. 1&8
v«v*, ud. iodAet), be in nearly tli« mine phase o( tlie coDlractivn nt tbe
■Mline.
\^*an* of DODLraolioD tbiu puwiiiji aliinc U»e circular and loiii-iluduia)
k ■>f ibe iniesiirw! raoBtiiute irtiat b (-allt<il pemialiic at-tioo.
jk« xhv <i>iitracti<JDi of vtrimlMl uiU)M:Ii.- the coutniclioua uf plain niuecles
\t b» »t*nin] hy ttimulKtiuti of uiu-vtn ic^ng tu the part, the nerrea sup-
|i}ilif plain niiiMaUr tilMie. ruiinin); fur tlie most [tart, u ne tiave said, in
tbir*»w)e(l *yn>p«lbetic kynteni. but being, ox ni- iiball M-r. iiltiiuutelx c»u-
DRWd witb thr kpinnl cnril or brain. Hi-iv, Ii<iwev<:r. wi- runiu upuo au
Ifiiilllil dUlinrlHrn bHwm-n ibv ntrialnl nkclttal niuoi.'U'H, ami the pluiu
■wlw of tlM' vMconu Af a ^ncrnl niliv ibc AHt-liil niuiu;lt» art; thn>wn
iUm cvintnaninD only by n«rvou' impnli>e« rracbine them ubin^ thoJr nerriM;
ifMUuMMU tnuvvRicnis of tho i^kclctiil nuiBclra. ibat h. contraction* nrlalng
nuof changes in the musclca lh«inM)v«8 are rstrcnicly rare, and whcii tboy
genir atr abnormal ; au-called " cramps '' for Instancf . which are pnilvngvtl
Kinir i.-iint rati ions of skeletal inutclea lodependeDt of tlio will. tli(>U);fa tbeir
Kttmae* w larvelj due to tbe ooudiiion of the muscle itself, genernlly the
mull (if overnork, are probably ai'tually started by ncrv<iu» impultee reach-
bctbcBi truiu without. Un llw other baud, the plain muaclcs of tbe viscera,
(f ^ blflstJiw. uteni*. and ureti^r, fur iitMunc«, and uf the bloodvcMeU very
bi^wntly fill) into coulractioiia and ao carry out movements of the organs
u> whirfa ihey belong iguile {nd^JcndcDtly of the central nervous aystem.
Tltar (irgiaoi exhibit " 9j>ontimeou» " niovcmenU <]uite apart from tlie will,
ipirapttn from tbc tvntral nur^'nun *y#ten), and under favorable circum-
tiBtm continue to do tht* for »i>m« time after thev bav4- been entirely isolated
■d rmuvnl from the Univ. 80 slight, indeed, ii the connection betweem
■hanaranaitt of oi^ns and part* ^lupplinl nilb pluin miii'i-iilur fibrea and
inW, lluit thcae muKtilar tibrea have ometiince been called involuotaiy
aoitiB: but this name ■• undesirable. since some muwlM GOOneting eiitirvly
dftma moacular librea (e.g., tlie ciliary muHclea by vbioh tbe eye is bccoiu-
MataJ for riewing objects at ditl^renl diatancw) are directly under tbe
iMiimwe of the irill. and some muaclea conipoeed of striated Kbrea (c. y.,
iha* of the heart) are wholly removed fhint tlie influence of the irill.
Va ihall bt&i siuily. Imweror, the facta relating to the movements of parts
fnriiM with plain iniiacular fi bras, when we come to oonaider the parts
AmmIvml
UId* the skeletal mu»clc», vdioec iwr^'ous element* have t>een rendered
fcHtiooally iDcn|iable (i 78), plain muscles are muoh more iwnHitive to the
■king and breaking of a constant current than to induction nhurks ; a
~1|irat, kIwu very bnof, like that of an induciion-iJiock, produce* little or
.•ftvl-
Thc plain iiiuxrlfla iMm to be remnrknbly aufccnlible to the iiiHiicnt-cs uf
lm[irratiire. When exposed to low Umpi-tiitun^ tiicy rcndily low ihc (inwer
*f minrttDg : thus the movements of the intcMJnc nri' »iiid to ccim;' at n
ImpamUire below \it° ('. Vnriatii>n< In tcmperaiuro have aU'> very nmrked
•fnt nn tlwi duration and extent of the contractions. Aaaocialcl probably
tah thi> susceptibility iii ibo rapidity witb which plain muscular libra, even
b eoU-blooded vertebrates. lo»e tlieir irritability after removal from the
M* and severance fmrn their blood-supply. Thus while, an we have seen,
Ikticeleial muaclea of a frog can be e3C[)eriitiented upon for many lioun (or
nw fit two or three dava) after removal from tlie lutdy. and the skeletal
■■■tia of a manmal fi>r a much lees but alill comitlurable time, it !• ■
WUffnf rrry gnM dilficullr to im.-cuiv the c<intinuu»cv of luovemeoi* of
AtiuMtine or utherotgans supplied with plain iuu>culnr librni. even in the
■Msf Um ftog. for any long |icriu<t after removal from the body.
164
THE CONTRACTILE T198DES.
TIm contrnoiton nf nlnin muttrtilitr fibres is, u w« said, very ilnw
dvwlopmeiit iiiii) very long in it» ilurttlion, orcn wh«B sUrtmJ liy n ni'tiiirii-
tmry atimuluR, such ru n einglc inductimi-shoclc. Tho contriK-tum :illor h
MiiauUtion often ImU so long w (o rniso tho qimtion, wluttber irhRt liii* been
produced a not n single contraction but a tctttniis. TebiniiE, how^riT. thiit
IK, the luaion of n eeriei^ of contritctions. weins to be of rim occurrence,
lhou;;h probably it may be indoceil in plain muscular tiaasuo: but the eadt
of leianus are gained by a kind of contraction which, ran or ni lead not
prominent in skeletal nuiecle. beconiea of great importance in plain musculST
tisaue by a kind of coiilracliuu call«d a (onie contraotion. l^ie subject is
on« not without difficultiett, but tt would ap(>ear that a plain muscular fibre
inay remain for a very considerable time in a state of oontractJuu. the
amount of «liurt«oiny thui maiiitaiTied bein;; either sniull or i^rcau ; it t« Iheu
said to l>e in a ttLale of tunic contraction. Thin ta eapcciallr netm in the c»»e
of tbu pluin muHciiIitr tiiuiue iif lh<! nrtt^riu*, and wc shall nave to return tv
tb!« matter in di-aling with the oirenlnlion.
Till- inusnilnr liivmi^ nhich enters' into the ooniiU'uctton of the Ikcart is of
a peciilinr nature, bi-in^ on the on« hand iitriutcd and on the other in aftinB
nisneet* Mmiliir Ui plain muscular tiMUO, but thi» we Mhall considvr in dcalii
witn tho bmrt itself.
Ciliartf .VoweHiwif.
^ 93. Nearly all the iDovemeot« of the body which are not due to phyiic
causes, such as gravity, the diHiiftion of liqnida, etc., are carried out by mus-
elee, either striated or plain ; but some small and important etiecu in ibe
way of movement are produced by the action of cilia, and by those cbaugcs
of pruloplasin which are <-alled aniiebiiid.
Cilia nro generally n|>peii<lai[t!.H of epithelial cellit. An epifhrlinm cnn*ii)ti
of a number of cellit nrrangefl in a layer, ono, two, or more celU deep, the cell
bodies of the coiKtttucnt kkW* being in contiict with each other or united
mcnly bv n minimal amouiU of wmeiit onWlanee, not «panilc«l by nn
aj>preciable quantity of intercellular material. As a rule, no conoc^rtire
tosue or bloodvcasef pni«M between the cells, but t)ie layer of cell* resU on
« basis of vascular connclive tissue, from which it is usually separated by a
more or lens definite bnscrnenl membrune. and from the bloodvenolsof whjeb
its cells draw their nourishment. The cells vary in tbrm, nod the coll body
round the nucleus may be protoplasmic in appearance or mav he diHenenti-
aled in various ways, An epithelium bearing cilia is eafled a ciliaieil
epithelium. Various passages of the body, such as, in the mammal, parts
of the nasal chambers and of the respiralory and generative passages, are
lined witJi ciliated epithelium, and by the action of cilia fluid couiuiniiig
rarioin panielm und ttt^uemllv nior« or less viacid is driven outward along
the pamagen tnwnrd the exterior of the body.
A typical epithelium cell, .-luch nn may be found in the trachea, is gvne*
rally sumewhiil wedgushajied with its broad end circular or rather polygonal
in outline, forming purl of the frcR surface of the rpitbcliuni, and wilh tie
narrow end, whieli miiy be n blunt point or may be somewhat hniiiched and
irrvRiilar. plunged among small subjacent cells of tho cpilhclinm. or renching
to the ciuiticctive lii»uc. Iielow.
The cell body is. over the greater purl of its <'Xt«nt, composed of proto-
|>lasm <Tilh the usual gniniilar iip)>cnrHnce. At nbout the lowcrthird of tfa«
cell is pluced. wilh ile long aiiis vertical, an oval nnclens, having the ordi-
nary cliaraclere of n nucleus. 80 far the ciliated cell resembles an ordinary
epithelium cell ; but ilie free sur&ce of the cell is forioed by a layer
ON SOHB OTUBR FUltUS OF COKTnACTILB TISSUK. 155
kysliM trnospareut soroewluit rvl'niciiv« (uUunw whinb, whtm ih« nil is
MP. M unuftl. ill profile, mppt^in n n hyiilinc rpfrnrtivo bnEi<l or bonier.
VroiB U)i« b<iT<lt>r ihi'rc |inij««t ouiniiril n VAiiiibln niiiiihi.T, IC t<> -t'.), tklU
nttr iBpmnfc liair-liko tilftmeoM, Tnn'ine i» l<'ngih, but gvmi-rally nboiit ii
•(tiuUT or ft tliini u long U Uio eolt ibwlt'; thcw arv the i-ilia. liiinKtliatcly
Mu« llii* hy«lini:! lM>nlvr iW orll Fulwiuictt often «xhibil» more or Imr ilis-
tbetly ■ luaKitu-liiuil ftrmtiuti, lini.- linos pM§ing down from ibe hyaline
V lowHnf the lower part of the: cell euUtetici' mund the niicloiu. Tim
bordar itaelf asuallr cKhibiti^ a striatiaii as if it n-ero split up into
■rh block corresponding to one of ibe cilia, nod c«re(ul examitia-
liM loMfa to the conclusion that ( he hyaline Imrder ia really ooinpoaed of tlie
Ibtd tbiokar baeal parts of the cilia.
Hi* mail body has no distinct external luenibrane or envelope, and its sub-
KMCf is in close cuntact with that of its iiei|;hbor», beiuK nnited to them
albcr by B tbin layer of Mine ceiueiil substaooe. or bv the siuiple cohesion
«f ibHr rutieclive surfiices. At all events the oetlfl do cnhere laif^ly
ii(Hbcr, aii'l it 'm dilficull to obtain an itolntcd livinf; cell. thou)[h tlie cells
uy be evily *cparaivU from each other when dead br the help of diMooial-
iif lluMii. WbMi a cell in obuitued is»]ut«l in a living state, it u very
twiacatlT fwund to hare Imi iu ncd)pi fhii\v. and to hari; become more or
^Jb htnjtpberictl ur oven Htibcrii-al ; undi-r the Utnal cxiditiuns, and fre«i|
^|Bb tlw Mpport of its ncigiibon. the cell body change* it* f»rm.
^rksgnMralcharactenjuKt'liwrilmlnreooRinion toall ciliated cpitbeliiiui
(ilk Ml lbs cell)) in diircrciil niluntiouH vary in certain |)articular«. nivh as
Asritrl form of the cell IicmIv, the number and length of the cilia, etc.
I H. (~iliar}- actinii, in lliv ionn in which il is moM common, in mammals
ud. iodei-d. vi-rtcbrsle*, consists in tlw cilium ( ■'. r.. th<^ tsfierinp tilamvnt
ifiikto of alxtvc I being at one tnomeat straight or vertical, at the next
wt b«ia;c beni <loivn suddenly into a houk or Dii-kle form, and then more
4mh returning to the straight erect ]>oailion. WIteu the cilia are vigorous
Ail double nwremeni is repeated with very great rapidity. «(• rapidly that
4r indiridnal movements cannot be seen : it is only wlieo, by reason of
favun, the aotioa becomes slow that the moveraeat Itself can be seen ; wliat
itsMo iithtinriTC b simply tbe efltrt of tl>e movemenL The miiverucnis
ttsBslow hnvv lM-enc»unied at about eiubt (double nuyvetnenis> in aseciind;
imMkly when rigonxn they are leiNuiteci from twelve to twenty tinuv a
IV flrxiot) lake* ntim' in one dircctiim only, aixl all the cilta of each
mI and. indrvil. of all the cell* nf tbe name epilheliiini ni'ivc in the safllO
AiMfioo. Moro')vi>r, the Mtmc direction >■ maintnimil itiiring Uie whole litiB
rflheepttheliuoi; thuo iho cilia nf th<^ epiihelinm of (hit trachm and bmn-
■Ual oaaagaa more during the whole of life in »ueh > way ss to drive the
JaU Ijrlng opon them upward toward th<- mouth; a« ftir as we know, in
rsias, or at least in mamiDals, the dirvction ts not ai»(l cannot by any
be rweneil.
TleBexion it rm- rapid. Uiil the return to (he erect |>i>^ition is mneb
ikmt; hence the total etiect of the blow, supposing the ciliiim and the cell
)*)|« Used, is to drive the thin layer of lliiid in which the cilium is working,
•kl which always exists over the epithelium, and any {)articlf» whidi nuiy
Wfinstini; in that Huid in tite same direction as that in which the blow ia
fina. If (he cell be not attached but Hoatin^ frv« tbe eflect of tbe blow
aav bs tu drive the cell iLwIf backnanl; and when perfectly freali ciliated
rfiibiiuni is b»aed out and examineil in an enen fluid, such us nurmal
alia* fciliitiuu. isolated cells or amall jft^ups of cells may be ween rowing
Asodves about as it were bv the action of their cilia.
1S6
THE CONTBACTILE TISS0B8.
All th« dim of a od) mov«, a» we liave juttt Mid, to tilt mow dtTtntioil,
bill net <iuile nt lite aaoi* time. If nv cnll llio siclv of the oell townril whkb
the viliii bdiii the front of the c<:!l nud th« ornxMite n-vin th« back, tlit- dlU
Ht the bnck move n tritlt- bcf^irc thiM« at toe front. e» (lint ihv movement
runt) over the cell in tlii; dircctiun of the ixioveineDt itself. HimilnrlT, taking
nay oDe cell, the ciliu of the c«lls behiixl it move slightly before. And th*
ciltn of the L'clle in front of it slightlr iifker, ile uwn cina move. Hence, in
tht» wny, aloD); a whole elretch of epithelium, the inoveineDt or bending of
the cilia sweeps over ibe snrfiice in rip|)]cs or waves, very niueh as. when the
win<t blows, siriiilnr wnves of bending sweep over a field of com or (all grtM.
By this arraii;;emeiU the elRtacy of the moveiueiit i* secured, and a staoily
stream of 6util carryiii;; tiurllclea is ilriven over the surface iu a uniform
coiiiiniied ilireetion: if the eilta of separate cells, and atill more if the
segiarate cilia of eueh cell, moved iiule))endently of the otliers, all that would
be girudiieed would bo a leriea of niiuute " wubblM." of tu little use (or drir-
inj; the lluid dcfiiiilL-lv onward a» the ufTorla of a boat'* crew all ruwinjic out
of lime an for ]ini[H.-llinu[ the boat.
Swift bending nud nliiwer i>lniij;1ilcMiiig in the form of eiliari' movement
generally net with in the cilint<'d epUbeliinii of mammalH and. indc^], of
vertebntteii ; bill nmong the iitverU'liruii^ vcv find other kiiidii nf movonicot,
such ii> a irt ami fro moveineiil. c(|iially rapid in Imth din-ytionx, a i-urk-ecrtw
movement, a Himple uiiduliitory moTement, and many others. lo each caw
the kind of movement seems adapted to secure' a snecial t-od. Thu« even in
the mammal while the one-sided blow of the eilin of ihe cpilhclial cells
Bocurea S How of tlnid over the opithelhim. ihe tail of the i'i>erninlazoon,
vhieh is prnctieally a siuffle cilium, by movinj; to nod-fro in an iindulalory
liashioD drives ibe head of tbe speruiato/.oon onward in a straight line, like
a boat driveu by a single oar worked at tbe steru.
Why and exactly liow ihe cilium t>f tbe upitbebal cells beads swiftly ami
itmifcbtens slowly, nlway* nctinj; in the same direction, is a nroblem difficult
at proont tii aiiNweT fully. Some have thought that the body of the cell is
conirnctile, or eonlnins coutrtictile mcchnuinmii pulling ujk»i the cilia, which
arc thui' riiuplc pawivc pupp*'t» in the hands of ihe oellii. But there is no
satisfnetory evidence for such n view. On tlic whole tbe evidence U in fnvvr
of the view that the action is carried out by the cilium itaelf. ihiil the Ix-nding
is a coDtraction of tbe cilium. and that the slraightoniDg corre.<|H>tids to the
relaxation of a muscular Hbre. But even then (he exact maaner in wbich
the conlraction bends nud the relaxation strnightens tbe tilnraent is not fijlly
explained. We hnve no positive evidence that a longitudinal half, tbe iDside
we mit>ht say. of the tilnineiit is conlructile. and the other half, the outside,
eiaiilie, a suppoMtiou which has been made to explnin the bending and
Stmi};htL-ning. In fact, no adeipiate explanation of the niatler has as yet
been given, and it is really only un general grouuds we conclude that the
action is au edeci of cuntnictility.
In the vertebrate animal dliu an-, lu far a.« ive know, whollr independent
of ihe lu-'rvous system, and their movement ia probably c«nsefeea. Iu aucb
nnimnb, however, ns infusoria, hydrnxun, etc., the movements iu a ciliary
tract may ollei) be seen to s[oj> and to gu on ugaiu, to be now fast itow alow,
according (o the needs of the economy, and, ns it almost seems, acoonllng to
the wilt of the creature; indeed, in some of thew: aiiimnls the ciltnry move-
ments nre clearly under the iritliicnce of the nervous system.
Olieervations with gnlvnnic currents, constant und iuterruptcd, have uol
led lo any satisfactory results, and, as far iw wc know nt prcsenl. ciltnry
action i.i moat airecli.-<l by ehan^ea of temperature and chemical media.
Moderate heat ouickens the niovemeuts, but a rise of temperature beyond a
certain limit {about 40' C. jn ibe case of the pharyngeal membrane of the
ox iOXB OTllKR FORMS OP COyTttACTtLE TISSL'E. 157
from) hnamt* Injariout : colil raUnb. Very Hiluto alknliw «re fitvonibU!,
BCW* u* iajuriou*. An csccw of earhoiiic nciJ m mn nbofim or oxvgitn
ditsiiaUM* or arnvt* th<> movcmeiiu, nthft u-mjxinirily i>r [>crni»i>rtitly,
toooHiig 10 tb* Icogtii of the •xponiro. Chloroform or pthor in sligl'it
* ~~~ldmlldriMiorwq>ta(ls thttMUoD t«iii()oninl]r, in L-xcna kills inil dis-
Ibeoelk
AwrhtHrl J/otemenb.
tM. Tlic Khiti* blood -rorpUKl«s. m we hare aaid (_i3fi), atv sbl« of
iboMKlvM to dkuii'e iheir form uid by repeated change* of form i« move
(MB pUm to place. Kiich laovemcDtH of tM aubstance of the corpuselei on
(^M RRKeboid. siucc ihey clowlj- resemble and appear to be idcDtieal fn
wtaiTT Willi the niovemenu executed by tlie aniceba and limilar orftiinivott.
IW DinrciucDi uf ihc eiiiloplosn] of tlie vegetable cell Means al»o to be of ilic
■me kind.
Tbr aiH'pba obangn ita form (and ibifta its jtlwe) by throwing out pro*
jcrtinni of ita wbManoe, cnllei) {iM-uitopiHliu which uuiy bo blunt and abort.
MWil hiilginga lu it vtn, or may be t«o loug and tbiii a^ to be men filammta,
• may br '>f an inifrotedialc cniimcter. A« «« watch the outline ul' tJio
bslnie cvtoMrv we mny *ec n [woiidopodium bi!giniiin){ by a flight bulging
rftha oatline: the bulging incn-ntw by the tiTighlxiring pi>rhons of ihi'
td/mn moTing iniu it, iIh- movcRK'ni iiiidcr ihc niirrorcopo reniindiii); one
rf iba fl<nriiw of melted ghm. Af the jicoudopodium grows larger and
Mfia the whole thickneac of the eotccurc at the spot, ibe graotike of (he
mSmn ri«t be wen ftreamiug into il. foraiioe a core of endoaare in the
■idilk of li>e bulging of ec(oear«. The peetidoixHlium mny continue tit
pw largar and larger at tlie cxpenae of the r^t uf the body, nud eveutunlly
ik wbcilp of lhi> amn-ba including ihe nucleua may, lu » were, hiivr
fMad into the iiwudoptHliuiu: the body of the aoweba will now ncctipv
Aa bIm* of the paeudoiioitiiini iuateod of it« old place; in other words, il
«(llw cbMngina ita fonn hare aha ebang«d itd plane.
Daring aU t£s>« movementa, and during all »iniiliir nmaO>oid movem«-nt*.
iWkalk uf the oiganiam vill, a* far a* can be aNCLTtniuiil, have reinainc«l
Mckaagnl: the ihroaingoiit a fwi'iidutiadiiiin in nnedirrction U nccnm|>anie(l
bsmrmipondiiig rttraclion of the nody in other dircrtimw- If. a* N>me-
liDin ha{>]rRe, the i>r^ni«ni throwa Mit nwiid>i|KHlia in varimx directions at
ll> Hne time, tbe main boily from nhicli the perudiipotlia prnjccl is i^uced
iidileluieaB ; from being n vplu-ricnl tump, lor instance, il b<-cnn>ca a bmnched
CIk. The moreoent it hmiighc nltout not by incrcar« or decrease of nub-
Mum Iwl by nere tranalocntion of particles ; a particle which at one nM>meut
W ia one pawUon nwves into a new [xjeition. several particles thus nioriitg
ttaird the aarae point cause a bulging at thai |N>ini, and several pariidea
■uriof away from the same |>oinl cause a retraction at that ptint ; but no
t*« futielea get nearer to each olher m as to occupy together Ibm spacp and
tlai had to oandenaotton of substauee. or get furt^ tVoin each other h> a.i
teiecupT mote apace nnd thus lead to iocreme of bulk.
la tBu rMpact, in that there is no change uf bulk but only a shifting of
(■Rklca in tncir relative position to each other, the snin-butd movement
MHtbteaa muacular Mmlrairti'io; but in other reapecls the two kind* of
■amaal seem diff)-n>ui, and the (|U«Mi«« ariaea, hare we the right to apeak
xt (he aabounce which can oulv exeoutc niti<i-1md niovementa m beiitg eou-
Wa naT, if we admit that contractility ia at bottom simply the power of
4ifiaw ifie relative poailion of pnrticlte. nnd (hat niu^cnlar contraition t* n
ifaraujze<l fTw nf contraction. In n plain muscular fibre (which we may
158
THE CU>'TBACTILi: TISSDBS.
tuk« ukiinpler itmti tb<^ Atrtuied muMle) the ahiftlng of nartides is special-
ised ill the trni- ilint it tiiu iilmiiyisdeniiiM nlatioB to t&e loDg axis of liie
fibre : when t\u- RUtv. coiitraciH » curtain number of nartiolo umiiue a nev
nwition by moving nl riglil aiiglisi tn the King iixii> iif the fibre, and ihe fibre
in cdtiwi^uiMRT lin'onie* vJiorler anil brotiJer. lu n wbit« blood 'Curputcle,
UUKcbn, or other urgnuiMii fxeniting umo-boi'l miiv^mcnlii, the tliiAiiitf of tite
particles ia not limitod to taiy axis «l' itu- bmly of tbe or^nnivm ; nl the immt
moment one pnrLicl« or ono sot of piiriiclot inav be moving in one diroctiun.
mill nnolher pnrl!i.'l« or nnutlier sel of particles in lUiolncr ilireclion. A
pMudojHHliuin. fbort and brand, or long, tJiin aiid lilamcntoiis, may be ihriM
out from any part of tlio surface of the body and in any direction ; and a
Sreviously existing pseudopoiliiim may be sborLeiied. or be wholly dram
Bck into tbe siibsLance of the body.
In tlie plain nmscle fibre tbe fact that tbe shifting is speeialixed in rolation
to the long axis of tbe fibre, KMCemUtes that in a conlrsctioD tbe shortening,
due to tbe particles moving at right angles to the long axis of the fibre,
should be followed by wbal we have called relaxation due to the particles
moving back to take up a {tositiuii in tbe long axis ; and we have several
timet) insisted on relaxaiion being an eosential part of tbe total aci of con-
Iraction. If no sucb moveuient In tbu direction of relaxation took place,
tbe fibre would by rejwuted cnntracllona be tlatleiied out into a broad, thin
film al right angles to Its ongtiml long nxb, and would ihuti become u»el««.
A spherical while bloiHl-curpiiicle luay, by rv|>eaied contract ions, r. e., aimeboid
niovemeuta, tmiiHlorm Itself into such a broad thin lilm; btit in such a euo-
dition il i» not useluo. It niuy remain in that cmidition for mime lime, ami
by further eontracliuiia, /. i-., am<oboid mnvcTnenta, may awiuDie otlMir shapes
or reviu't to the »])liericid form.
So long nji we nnrrow our iileu of conlrai'tilitv to wlml we Me in a muKCular
tibre, and understand by contraction a movement of particles in relation lo
a delinite axis, necessarily followed by a reversal of the movement in tbe
form of relaxation, we shall lind a diliiciiliy in speaking of the euljslance of
the amixba or of the white blood 'Corpuscle as being coiilnictile. If. however,
we conceive of contractility as being essentially tbe power of sbiftius tJie
position of particles in any direction, without change of bulk (tbe shifting
being due to inirinsio moleculiir cbange^ abuui which we know little aare
that chemical decompowtions are concerned in the inatler), we may speak of
the substance of tbe oniceba and while blood -corpuscle as being contructile,
and of muscular contracdoti uh being a sjiecialized kind of eonlractiou.
llie protoplasm of the amoeba or of u uhile corj>usole is, as we bare Mud,
of a contialency which we for want of l>etier tenuit call iiemi-solid or mu^
tinid. Consequently when no tnteniat cbunges are prompting its ]ianicl«t Co
more to this or that direction, the influeiiiex iif the surrounding will tend tO
give the body, ii» ihey will nthi-r fluid nr M-mi-ftuld drops, a apherirnl form.
Hence the naturitl form of the nhitt* ccrpiiMcIc iti mure or leas spherii'al. If,
under the influence of K>me stimuhm inte.riml or exlenud, some oT the
particles are stirred to »bifl their place, emieboid mnvenieiits follow, and the
■pherical form is lo«t. If. however, all the oitrticlus wero stirred lo move
wtlb equal energy, they would oeutralin^ cacti other's action, no protmnoa
or rctractioD would take place at any point of the surliico, and tbe body
would remain a sphere. Honco in extreme stimulation, in what in the
OKHcle corrffiponils to complete tetanus, the form of the body is the same as
Id rvst ; and the tctaiiiied sphere would not be apnreciably smaller than tbe
sphere at rest, lor that would imply change of oiilk, but this as we have
teen does not lako place. This result shows strikingly the diflerence between
the mneral contractility of tbe a]u<eba, and the sjiecial contractility of the
macde.
HAPTER TH.
(« TIIF. MOKE (iKNERAL FEATURES OF NERVOUS TISSUKi.
tM. Ik ibc prcrexliug rhii{itfr wo have denll witli tb«! pro|>«riieei>rii«rTe«
pitf lo niHCK». ih*- ii«r\-c* wliich we citllott motor, ami linvc iiicidGUtally
HUiHl of Othif nen-f* which wr ciilli-d rrnmry. liulb tfa^ic kinds of tieri-ce
vtoowiMtttl witli (l)i- brnii) iind »|>iiinl curd, ami fonii pnrt of the general
■fmoj ajvtrDi. Wv nhall hnvp to study hcrciiftcr in d<;iail the hrtiin and
4iuJ ctiro ; IniI tlir ncrvuits »_V(U'm inlervcncs ei> ropenlcHlly in the procpt«efi
dfriid out hy other Umum thut it will he deeirnkte, before |)roce«diu|;
faltkcr, tu diM-tiM tOBM of it« mora general features.
Tht oervmu ayfrteni iv)iMiBlB(l) of the brnin and ^irnil cord [Via. 44] form*
i^loprtlier Ibe erreiiron/'imtl Oirit nr erntml iiemiu lyt'ein, (2) of the iMrivv
fwlKf f^ih lliikL axis to nearly alt jtarla of ihv bodv, thwe which are
«H«AmI with the Hfiiiial c-anl h«iD^ culled tfiiu-il and tliowt Hhich are cuu-
with the brain, tiiihlo tl»e eraniuni, being called emniW, and (3> of
ftta distributed along the nerves in varioua paria of the body,
bf tpitial cord obviouHly oonalsis of a number of veii^nieiila or nietamerca,
ving tu niorewiiiai aloog its usis, each metaraere giving off on cai^ aid*
■ [air nt Bi>iiuil uervcH ; and a Hiniilur divi.-iun into nurtaiuo-rfti may lit- trnocd
klhe hraiu, though \em di^linRtly, linci^ iIm: cranial uervv* tirv arrungtd in
amatr ■omcwhM dJfiiBrail frvm ihiil uf tho spinul ucrvce. Wv may take ■
diA apinal metAin«r«, r«(>reBeiit«d diagrunimalicalty in Fig. 45, as illus-
tnnu ihc general features of Um n«rToiU system ; and since the half on
<w aw of ilie median line reeetnblee the half on the other ude we may deal
«ilh oae lateral half only.
EMxh spinal nerve arises by two roots. The metamere of the oeotral ner-
IW tytUm i.' consists, as we slioll hereafter see, of ^ray matter Or in the
iHmur and whit« matter M' on the outside. Fmm the anterior part
tf (ny matter is given off the anterior nerve root A and from the posterior
luo posterior iK-rve root P. The Utter naaaea into a spelling or ^an-
0, " the ganglion of the |MiMerior root, ' or more shortly " Uie uniial
"; the anterior rout di>e> not pit-w into thi» nuigliou. Ueyona the
.^ _.j tfat roots join 11 form the ui^rvt' trunk .V. Wc ihall later un giro
isvidtoc* thiil Utv nerve lihrra ci)m|H>tting ihc pcaitcrior riHit /'are. a» ikt
kvc know at pri-M'nt. exdntivcly ciccupinl in currying ncrvoiui im)>uhNa
fiaa ll>v tiMuts of llw iHHiy to the central uerromi fy>t<-ni, and thai the f\hm
•Vpodng tl>c anterior rwA A arr similarly occupied in carrying impulwcs
fca thr n-nlral nerrous system to the several tissues: lluit in to say, the
iwaer ia ninde up of ssiuory fibres, or, i since the imnuhvs |wwing along
Aon m the ceolral systom may give rise to effects otlier tlian iwnMitions)
tfittmi Abree, while the Utter ia made up of tiiotor, or (.since the impulsiw
fnisf along them from the central nervous system may produoe eflocts
■ than movements), (ffemit 6br«s. Tlie nerve trunk N is oonsciitcntly
pnl nerve oorapoaed of aSerent and efferent fibre;^
I far the greater |Murt of this mixed nerve, dividing into various brnuchet,
»dNirifaated i .V) to the skin nud the skeletal luiiAcles, some of the librtt
'■nier) ending in muscular tibres (JIf ). others (iteiiaory) ending in cpithrlial
■*IU($) mnitected with the ukin, which we shall oanaMer hereailer under
[VW. M.— l^KH fKttnn OR Ham or mc CiixknHi;ii '>i> Cmu»ki.i.i,-)i, unit «r twk l^w*
Vjimui «l^ «iori.L» obiahuat*. *v«) tu« Asntuon Bir»rjict or tue t-rixii Cnu-. to Siio*
Tia UoDK iw Omii>iii or tiik Siixti, NcKris frov iiir. srtmi. i'okb. tsn the ciukiii. Xrrt
rMW mil RixKoirTili; DntiW, n, .i; nrrbnil Iit'iiiliphnn ; A, rlHht halfof cpnhrll'un: n.m^lallii
tXAtntlfm: atomhli li> trniiirrnciibllc mam. Ihc poni VanilU ; r. r*. (he uplnikt ounl. (Iiuvliigii*
cnrloal ini) IdnitxT vnlBtitiuiiiUi, knd ll*i>jliiU'd MniJtMtiuiu ; «. (lip lauditniulnii, fdnntil ligr (ha
«ltiiilist«l nvuof Ihu limbuuid mtnl oamii 1 loa. lb* Kvnrnl cniilnl ncrvn . artiln^ from tbe
hue tl Uic bnilii xnil the ulilc* of lh« niedull* oblFini^u. Bulou itioc. on cACb «l<k. arc Ibc roau
er Urifin* oT iHi; ^v^n^l ni*nrcv, vTn k'Rl. 'IrTwl LuieiTeit. wa'l Mi^t^l In «nn^ ot IbOHv lliv <lntihl«
icM «i> be iBtn, >nil (Iw nvllliiR nr eiidhIIkii on tlii> poalr-rlmr mn. n, «, ine ■1III1I17 or bnohtel
Ilium. Ibtnipd bji the ftnii luuvt ivtvicnl and Dm ilonoJ iiilaiil nerroi ; L Ihc liunlaT |ilai** : ^ itw
Mwml ptcxtH. (ImnM br (hiUut lumbar nrrv* ami llnl fmir lari^ hutyot : I, ■hiiuta idtrVoTlb*
•IwiUiar ih« cord oat ap««i,UHl Willi UBfMtloo or tht tiBftincnisin ilflnUralai-iiD wlil«U fuiiwm
rXBAL rSATURSS OP NKBVODS TISSL'KS. 161
tk^iMnwofrnworycpilfarliml cvlU. whiknihcm, .V, alter iltviding into minul«
UmiAm Attd Armiag tilexuMV ri><), in unys aot jrct detiniUily <)ct«rriiiDe<l,
to IMIM> ■■ccitled witb Ibr skin mtul >kv!piii) miiMln. &[orp)iol(ig»t« ili*-
tingviili tbo pBfts which ^ to tunn the skin, skclctiil mutclra, <-tc., n» nturi'ifu,
fno lite W&iimAniV |iim» which pi to form the vitcvra. Wr mny, nccurd-
h^j. atll Utia nuiin pan of thi- spinnl n«rve the toauUie division <if the
Sw* <lA«r ih? miscd n«rve .V leaves tiie spinnl canal it gWee nff n small
InaA r, which under ihe name of (white) ramtu mnimHuiVvjiiJ'. joins one
(f B longiludinal series of yiiny/iVi t^i cunsiiicuitus in the tboms lu the inaio
tmfilktiie chain. This Uranch is iltstiiied to supply the viscera, and raij;bt,
iliRftK be called tite »pt<tnek»ie divisiDii of the spinal nerve. We may say
M ton, witbout eotering into dvtails, that the whole of the eytnpathetJo
mliB, with its gannlia plucujwo, aiid nvrvee, ut to be rej^rded as a develop-
MM or vipAiuion of the vi«c«ral or splauchnte diviaioi» of certain spinal
Btms. By Di«uiM of thi« sv*t«ni splaachnic fibm from the central rwrvoua
tnCm «n distributed tu tbe tuaucs of tb« viw^ra. *)tne of them <>ii tli«ir
Mj pasriuc tbraagb swoudary ganglia «, and, it niny bi-. l^rtinrv gutiglia.
IWnsj*. bowerer.as wesliall see, orrlain ncrVi^ or fibnwwhirh do not run
la ikr aympnthflic syMf oi, aiwl y«-t itn> ilistribiilisl (o tbe vi.-ii-tira and an
'iplaMDQle" in naturf'. We cnnnut, [brrvfore, iikc tb« word •yni]ialh«lio
l*4nMI« all the Kbn> which nrv sphinchnic in nature. On the othiT hand,
lla~i|)4anoliiii<- iM.-r%'«i"of tli« anatomist form n part only of tfauoidanchnic
ifMn in ibv idiovo mow, ihc t^rm thus used is limiic'l to partiruhtr nvrres
« Ibtiplanchnio mteiii distrihutcil to the nlHlonteii : and (he double use of
tkiffin splanchnic might lead to confusion. The dilfiuullv may, ]iorhn(«,
bntidea by call in^ the splanchnic nerrca of the anatomist " abdominal
^IsodiBte." The majority of these splancbnic libra seem to be efferent in
nsuir. i^arrii-inf; tmpiilaeb front lbs central nerroufl system to the tissues,
aaMcitdin]; in plain muscular fibrw(mr others in other ways (x); but some
rflhaSbres are nllereni and ciinvey impul»«« from tbe viscera to the central
■VTHs iyst«U), and it it probable thai sotne ofiheae begin or end in epitbe-
U ndU of tltc visoem it).
V« shall have occnsi'Xi in tli« nest chapter to speak uf nerves which
pWB ibe bI'VHiveasels of tli4' Wly, the aCHMlled va»omi>tor iicrvm. A eor-
(i^diai iif th«4', namely, ibe Ktw-MHS(rMitor ncrvtv or fibre* are branches
<f ibesplanchniiT division* oflho cvrebm Kpinnl nerren, aiul.iia we shall see,
At Taii>.OOQStri('tor nervm nf |h<' «ikel<Hnl muscles, okin, aod Other part* nip-
"""' bgrWBUitic nervw, after nmuingfor winn- distnncw in the splnndinic
'm ( I'), turn uide (r. r and r. m) and join the eomatic division, tbe
I qf which tlu-v necompanvnn their way to the Usfuea whose bloodvoosels
iiO*«y »Mr- ■
SSMS J. ■innnvrMfKUtii'ilicnnthihrcnrd, iDihow Um fffniiorihspareKBBasrbNBK la
t»mi0rttti otiUr ■atuBiKT /r.«(iDii»airikiiicp&n>.Biul>i»iiwnwnitaaneo(ihs<asl:Bad
ftUMtr 'iiir n«uuf « [«lre< t[4iial nccrw jprlaftng Ihon utiMn [
fHC ' mi Nimvtxir * ■D-nnnornn (Mhal Vuaa, ifttmj, irwhliQiiwiMr
'qftal camt J tiiuno*. /'faialarlamnl tiRBnaniHi oa ilut poiwnar n»l. .V « bolt tiem, V
*B*l mm* fTCfM, «>4loalii W (krlttsl ui aiuuiUc iuukIv, .VuiiuUcMiiiKiry tail o lurltuie, Xln
^t^V*- r tliw d arm mhlM ami Runnuinlcknii janliis UiasmiiBlton of lh« tnotaUiaae
•IMl,*arf poaam so •* I" Id •Uffil]' Uir inoreriliUat pkOtUi'D 9. Urm ■■ F'lo Um ptrtptMnl
pai>a V wHt mill* in n itlsathnic iuumIcl f irtawtiiilc wauy tnU or nrlboc, < oUior pcwtbls
FS^aSi from 1 !• flTm nA Ihr (nrbfiu nm« r r <i[n)r luiiut iiiwiiiialiaBil wtilfib
I toctaanl inHiiKl tlMtplniil ninl.ii4>] pinlyniiHHi. ■>. in otmnetdon with tM«|4nal
\.mtaj/t*j (■aiin>ilw<otiaMnclu(>atraii>ilie midcI« (•■'ja^bUadTtaalt In oauln |iu1i^ toi
.talS* ll«li>. :b.ta««rnir*l*Mili.-(taln anlilnc ihii fUcUa a( >b« SSrlM 1 Thn linnl-
• ((IksetksrnMtOTMUas nnn> i.* c' an not sbovB,
11
(IRKERAt PKAT^RKS or ITEftrOCS TIflflPKS.
We Imvv iMwii (§ 6H) thai a uerve ),">■>■)! 1" » 'ouvcle b cotDpoacd ofneiTe-
fihrei, chietl}' m«diillated, snmc, liowever. Iieinj: non-iii6cliillaM(], bMiai)
toj^tber hf cnunei-iive tiaaiie. The same deecriplion faolds vooti tor llie
whale Boiuatic divlsioti >if eacti of tbe sjunal nerves. The «piancDi)ic <livisioii
also consists of luedulliileil and noti-medullaieil libren Unuml lojtelbcr by ma-
nectivc tiaoufr, bul in it tbe noa-Rie<Iu Hated Rbra^ pre|>onderate, Mime bmncbe*
a|i|ienriflg to oontaiii banllv auy tuedullaied fibres at all. Tbe ouu nie>lul-
laled Hbrea which are found in the soniatie divisioo appear lo bo tibn-K wbich
have joiuf d that diviHitin fmiii thct splaDcbme divUiou. Su pfimiiKiit arr
iioii-niedtiMiiled filire* in nplanchiiic nerve* ami lieiice in the Nvnipubetic
Nyalem that tbi-v nrt; »iiniiirinift> called 9yni|iutbetic tlbru.
W« hn\f. Mild tbuL ibi* axiii cylinder, whether (if a medulla led or dod-
medullaied fibre, in to )»• ci>iK-<itlvred ii» a lung drawn-odt pnM.'CNi uf ■ neire
i-oll. Nerve crIU are liiiind in thn^ Diain *ittiation». 1. In the i^-nlral dot-
volt* «yi>toni, tbn brain, anil Kpiiml curd. 'J. In the wvcral ganglin placed
along the counN- of the ncrvcti, both the spinal {pmglia, and the ganglia of
ihosnlfliichiiic oraymptitliriir "y^ti-ni. 'i. At the Icnninntionii nf nrr\'» in
certain tiffiun. .Snme of tbcMir bitter nrcr lo hv- rvgardcd a» ^ninll, more or
IcH terminal, eonglia, and similar minute ^uglia coDiti»ling uf Ino or three
cells only arc found i'rciiucntly aloo^ the ci>ur»? of x planciinic norvca and
occasionally along the coureo of spinal nvr^'es; Mich celU really, tbereforo,
bcloDg to the wcnnd grnup. Rut be»id«» tbi«, in rertuio vitnaliona. as ibr
inelance in certain »rgnn« of the skin, and in the organ* of special eeoee.
nerves, j^nerally allercnt or sensory in nature, either actually end in, or at
tbeir termination ar« connected with, cells which appear lo bo of a nerroua
nature; such cells form a distinct calei^ory by themselves.
Uence nlaun its whole course a nen-e consists esclnsively of nerve fibna
(and the ooniicctive liMue lupporling them}, except in tbe central nerrout
syiten from which it vprJngn m tbe ganglia, great ajul amall, through wlucfa
il puttt, or which an; alta<'bi-d lo il at one part or another of ita oouiw, io
boib of which sitoiUinns nerve crll» are found, and at ita terniiuatioii wbcn-
ib fibres niav cod in nerve celli>.
Jjlipniiiiiiii 1 1 niiiiii I Till MiDiii.Knr i(;ji>iiiiui(OMiii: fivni-ioR ItKiT nr 0»K or ni«
a, tum* Mot fntvniig thv a>ngtloii ^ i, libra lisilnu llie NUDitUon lojoln Ibtnoiud ((tnal Ddrrv,
<HH<i»»r<lrn llimrii ir»i f ihi- (snctloii ; •(. |inni.i1|«l i|nnip,if nrrvDCVlli, wttti 6bim fi^in ikua
ft«B *iiKiii|pi III" mIK jirobiility tu iinli« Ihv loitutiii'IIn*nr roiiiMnR (irrrc ninM br T-^hapt^
JOMtlnlu.]
Tlie features of these nerve cells diHer in these several siluations. T)>e
eharacters of the terminal cell» wbii-h, an we have »«id, are chicDy evaaory.
|Pra.4;,
GBXKftAL FKATCRES Of SKHVOVS TlSSVItS.
idI ibr Mmctiir« of (he bniin nml npiiml eon], w<> •hall sludr in detail Inter
!«. W« nwy hero ronlinv <Hir lUtemioii to ihr norvo »IU of the }rnii<jlii>
■■I ti Minitt of tho bninil fi'Uiiri.ii <>f llic ncrv<7 colU of the epinnl cx>ni.
)tT. Sftiitiil ijaHi/lia. Whcn ■ longiliiilinal skIJod of ii »|>innl gaii^'lioii
Iltuaiit»-i) uniivr n l»w iM)w«r, tl»c libra nf lh« {lostorior root ns llicy tint«r
tkpn^lion «rc ohm-rvnl ta sprond out and {inn batween r«lativvly \aTgK
nl nHMpi<^uiiti>>ty titiuknlcd c«ll>, which an- to a large extent arrnuKifl iit
piM>. «oiii«nliBl nll4T (he fnaliioa of a bunrb nf grapes. [Fijt. 4<> j These
wt tW flcrTf cisIIb; tUry h«v« frv^giiputly ■ diameter of abuiic IIM)^, I>ut
MT bt fiill lafK^r or may be much ininlkr. In a inuuvvrse ■evltiut it will
WvlMTntil that A lnri;i' compact maas of ihne lvIIs liea on the outer ride of
AfgUflion, nii>) that the raoeiDoec ^roupa on the iiiiter ride ar« smaller. A
'ly of (woiwctive tinae ratrryiiiR bloodreaseb and lympbntic* run*
Uie pntipa, UiA paauDs iiit<i each K/oup runa beinevn the oclln and
and a thick wrapping oTeooiWctiva tuaue cmtiuutMU with tlie sbcmtta
lb« u«rve fcurrvutidi iiiid fornu a aheath for the wboli? piiiBlion.
Each i>f the nerve celk — (.-nDxliouic celk, aa they arc culled — «xainin<Kl
r a hi|;h«r jHiwer, ei(J>er after having been laolatecl or in an ttde(|Uat«ly
■ad nepsrad aectioD, will presetii tbo fullovring fieatuna :
eell ooniMtSof a kH hudtf nbich is, normally, poorahniH-d (Fig. I7],
a bfMul end iu which U ]ilucci] tlic nucleu*. and a narrow «>nd whiCB
out itiKi a (udk, and is eventually cuu-
lined oii a* a nrrve libre. The xiibHtaoee of
Af cdl hotly in of Ibc kiwi uhicb we call
MygnuiaUr protoplum; Mmirtimea there
> w ap^raniw of fibrillatiun. the fibrillA
niii varioUB riirocliotu in the body of
I and beiitg gathered together in a
kapladitwl dirvction in the stalk. Some-
tioM iIm! cell body immediately aro4ind the
Mcltu* api>ears of a different i^'rain fntm that
MUtr tlHt stalk, and n<>t untrequently oenr
At gueleOB !■ an a|tK"'Ki't*^i> cf discrete pig-
Mat yranules imbedded la tJie protoplasm.
tW ntitJevi. like the nuclei of nearly all
■mealli.ia brKcand contpicuous, and when
ittoomal eumliiioa is remarkably clear and
nAirtirr', though it appears to consist like
<(Wr BDclei uf a nuclear membrane aii<l nei-
^rt and Dudcar jntervtiliul material. Kven
am nnupiciiotis, perliapa, is a very large
H>rriml. liigbly n'fnictive nveifoliu: occaaioaally more than one atieleoliu
itpraaat.
i^miODdiitg tk» ocU body is a dtttinct tUfnlh or eit/tnik oontiatin^ of a
■nKaurnt. hyaline, or faintly fibrillatcd nwnibnine, lined on (be Inude by
ia» Miferar by two layers ot flat, |Milygunnl, niiclmK^l epitMioid cella or
Mub; that is to aay, ctlla whieh rriNiniliU' e|>ith<;lium i-eli>, bnt differ not
•aljia being cilrem«ly llnlieocH. buialsoin the evil IhkIv Uiini; Iranoformed
h* untinarr graiHilar protoplasm into n anav tntmparetil dilferciitiiited
WWrial. In ataim.ll ii|>e<-imen8 the niielei of tlti-sc plnt>-e are very cmopicu-
<aa Undtr normal condition:! this •hciith i^ in chwe (.-ontaet with the wlinle
Wj of the cell, but in harth'nixl nnd prvparod specimens the ivll body is
•wrtmia iM'U shrunk awiiy from the 'heath, leaving a apac* between them.
OHHiaaally till' cell liody while remaining at(a<-hed to ttlfllbeatb at three or
tmot mora potuta is reiraeted elsewhere, and accordingly assumes a more
•rtsiimist or * PvaiTOioi oakhiJ'
(ixn: Kkkvs Ctu,
A,u«(inlln(UB«le: B.soMnUac
(DAranU.)
0E:«ERAI. riATUBBS of XKRTOCS TIS81-SS.
or lev Uellate form : but ihU aitiBcml ooudilion must not be oonfouixled
with the Datuml branched fi>riu which, as we tliali aee, other kinds of nerve
oell* iMwewL
Wnen > section is made through a hardened ganglion, tlie plane of the
MCtion psMiffi ihroiifch the stallci nf few only of the ceils, ajxl that rareljr for
nnr grcnt distance aJooj; tlie stalk, since in the oase of many of the celb the
«t«1lc is luoro or less curved and ci>nsc>|uetilty runs nut of the plane of sec-
tion ; but ill properly iwlatuil cell* we tun itee that in aianv eases, and we
liavo rensoOM tu bdiovc that in all caat^ thi- stalk of the oell b, as vre liare
nid, cnnttiiiivd on into ii ntirve fibre. As the cell body narrows ioto tlie
ttalk scvcrul nuclei niiike tlicir npiMmiraDce, lodged on it; these nre. small
emnulnr niictd, wholly unlike the nucleus of the cell body itM'lf, and more
like, ihoiii'h not (|iiilij lik<', the nuclei of the neurilemma of ■ m-rve. Ther
are probnoly of tbo «imc nature as the Inttcr; nml, indood. ii* wi- trace ihc
luirrowiiig stalk <lowiivriir(I n fine ddiwii*- shwith. which, if jin^rnt. is at
least not oIivIook over (h« cell body, makes ilii iipixTiirnncr, nnil n little further
on, between this ^hcalli. which i# now deiirly n m'uritcmma, and the Ftalk of
the cell body, which hns by this time become a cylinder of unifomi width
and is now obviously an axis-cylinder, n larcr of medulla, very line at ti»t,
but rapidly thickening, is established. Tlie stalk of the nerve cell thus
becoraeb an ordinary racdnllated nerve fibre. The sheath of the cell is con*
tintied also on to the nerve fibre, uot as was once thought ne the neurilonnna.
but as that special sheath of connective (isei'.t< »f which we have already
auoken (§(>!<) as Uenl6*e sheath, and which ultimntely becomes fused with
tiie connective tissue of the nerve.
At some variable distance from the oell l]ie nen,'e fibre bears the firat
node, and either at this or some early sucoeedinji node the fibre divides into
two; iM we have Mxn, diviaion of a medullated nerve 6bre alwavn takca
place at a node. The two diviiii<,itis thiis arising run in opposite diret-'lions,
fonning in this way a T-piccc: and whilv one division runs in one direction
toward ihc iMWtcrior root, tin- lUlior runs in an oppnsite direction toiranl the
nerve trunk. The nerve cell is ihtw. ns it werv, u «ide picc« Bllttclie<l to a
fibre passing through the ganglion on its wtiy from the posterior root to the
nerve trunk. It cannot be said that in any one f^in^lion this connection has
been traced in the case of every nerve cell of clie ganglion ; hut thv more
care w tHkeii, and the mora successful the preparation, the greater is the
number of cells which may be isolnt«l with their r(«j>ective T-picces ; so
that we may conclude that, normally, every coll of a ganglion is connoctod
on the one £and with a fibre of the posterior root, and on the other hand
with a fibre of the nerve trunk. We have reasons further to believe that
every fibre of the posterior root in passing through the ganglion on its way to
the mUed nerve trunk is thus connected with a nerve cell ; but this has heen
caUed in ((uesttoa. In certain animal" — for instance, certain fishes — the cells
of the spinal );anglia are not pearshaped. but oval or fusiform, and each
narrow end ia prolouj^d into a nerve hbre, one end thus being connected
with the puelenur root and the other with the nerve trunk. In such a traae
the nerve cell in simply a liirtct i-n large men t of the axis-cylinder, with a
nucleus placed in the eiiiarjcenient. The nerve cells above descrilieil are
simihtr eolartfemcnts, also hearing nuclei, placed not directly in tlie courae
of the axis-eylinder, but on nnc side, and connected with the axis-cylinder
by the croM-]>ie(« of the T-i)i<-ce. Hence the ordinary ganglion cell is
sjwken of u* being unipolar, tno»e of fishes being culle<l bipolar. [Pig. -48.]
In examining spinal ganglia celU are sometimes found which bear no trace
of any pructw connecting them with a nerve libro. Bucb cells are spoken
of as npolnr. It is possible that such a cell may W a young ncl) whicli has
trio.nL
VtHUCB rOUU at GUMLMmC VMBTL*
k » M«> MUsir r«IK <r1ik ilNlr pfDloovulm*. (MtB Um Mltalw honi or ih« tnv iDiilM or iba
•IMd iMl . r. tmw nil altli M* toii(i«ii«4 Wn. IhMi (In uwrionitiBl* <tf tb* ftoUl iikI auditor
MtB In Om ■—nn aiidnniliw [uutiua of the ox ; a. cell vail; t.cell ecntdui; r, iilKinmiai il,
■AMir. pnlsai^uo* lbrnila« iW tbwib af Ibt Dbr*: /. nam ilbrv: a, ntrrv nil rmm th»
iriMMa »wml>«> <if ■■■ : r. laatlar fell ftnin the iplnal ronl : macnlllal UO lUuntun.
iM. Tbegiuig)i«<irth« 'p/'inrAnir Bystcm, liko the «{>inal KftDel>>< ronaiat
*rienre«du Uid Hbm imIwHdci) in connpctlvo tiwiit-, irbicb.liowever, m
«(>kaWTUl<l len compftii tiatur« in lh<!iii tUitn in the Hpiial KtmsUa. As
kr ■ the duincten of tbeir nuclei, ilie nMureof thoir cell siil>atance. and
AtpaKSMioD of B sli«ath ur« coticenie<l. what haa been snid concemiug the
MtTC orila of apioal ^^n^lia holds, in genernl. t[ood for those of splaiiehuie
(iwlbi: »nd, iiMJeed. in certain Kiiuglia nf the B[ilaiichnie >v»t«tn connected
Mo the rranial nerves the nerve cells Atipnar to be whollv like tlwwc of
fiwU i^atiglia. In most splniu-hnic f^iin^lia, howe\'er, in t^oK which nre
piMnliT allied lympiilhetic i^aiix'Hii, two iinjior-
(■t dHfennca may be obM-rvod l)«lwe«D whnt
n may call the <-tinraotcri8tic iicrv* cell of the
^Ineknic ganj^ion and the cell of (be ipiDal
u th« lirsi plaw. while tbt- nrrvc cell of the
^inal gangliii hu one proocM only, the ncnrc
fill vt th4> KplnncUnic gnn^lin may bnre, and
^inaptly ha>, tw«, ihrre, or even fonr or tire
it ii a muitifojar rell. [Fig. 49.]
iIm viwfind pUoi'. while these proccBBtfl of
•pUnchnti- ei"'?""') cell arc continued uti ns
libraw, aa U tiu- nnglc process of the Bpinnl
mil. ihri nerve nbrea n formed nre, iu
of mi»t of tlic proceosea of a cell, and
MWtinim in the case of all the ptoeMMS, Don-
BMUillatM) librr«, nnd remain non-meditUkled as
hr af the^ can be truced. In aunie iiiHtailMi one procMi beconwt at a little
teanM miin (he cell a tnedollated Hhre, while the other proceaea become
■■hOMlu Hated 6brea : and we nre led to Iwl'un'e tlm( in this case the medul-
filtre is proceeding fo the <^ell on ii« way from llie central nen'otu
SnujTB !lEaVE CUX, tMM
TKK NLELKlk C^VIO* CXiMlrV
ll^>^^nlu•l Vimlvub Oou'umi
iir A Form or 8ii Momim,
i»«cnllM<aLI Aftei Buu.)
106
OENKRAL FEATURES OF NERVOUS TIS8t?EB.
mtcm, and lluit the Don-niedultated librw are proceedini; Jrom ih« cell '
taeir way lo m»re [leripherally [ilaced parts ; ilie nerve cell seems lo »rve u
II crntra for the diviniun of n«rve fibres, and also fur the cIibd]^ fVom ni«dul-
Intcd In DDii-UMfdu Hated fibres.
[r ci)iiiH.'i|tK'Uc-e of iU tbus poswwiug mvotaI procesMS. the splauchnic
gnnclion <-cM !:> more or lew irre^lar and oft«D alar-like in form, in otitrMl
ro tbr pi'iir Khu|ie nf tb« »i>iniil gangliun cell. But iu cN-rutin .situation* in
cerUiin auimnl* — lor iiiatiintv, in (he fVu^' — iti many of lli<- );angliu nf the
abdomm. uid in thi^ umiill nui};lia iu llie heart, puar-;*hapfM) i>pliini-hn>c
ganglion cclli^ arc nint wilb. In mich euees the nucleated shwilh w ilintinrtlr
ncar-tt]i&rM.'i) iirbnllodd-ihniHMl.aml thi' liir^ir c>niiH|>ini'>ii» nni'li'ii:! i* pla<»H, n«
ID llic Bpiiifil gntigllon ftll, mar ihe hnwil i-nd. Inn ih« cell Mib«it«nc« of tbe
cell is gathered nt thi- «tnlk, n»t into ii t>iii-_di- librc. hut into two fibres, one
of wliidi in etrnighl and tbu olfat-r twiwU-'l K|)irally round tbe utrnigbl oae.
T\w tiro libm run for somr di»tanw tojnitlier in the «ann! funnel -shaped pro-
longation of the nucleated sbealh of tin; cell, hiil eventiiaiK separate, each
fibre acf|uiring a »henth (sheath of Henle) of itn own. fienerally, if not
always, one Hbre, usuiilly ibe straight one, becnniM a mediillated fibre, while
the other, usually the twisted or spiral one, i* continued as a uon-medn Hated
libre. While within the common nucleated abeath both fibres, eapedally tbe
Hpiral one, liear nuclei of the snme cburacteras those seen iu a oorrespouding
riiluation in the spinal ganglion cell. It hiis been uiaintatnetl that the atraighl
mill npiral libros take orimn from diHi^rent part* of the nerve cell, but UtU
bnn not been definitely proved.
In the wntlla of the iniestiiie, in connection with splanchnic nerrev, are
found [K^culiar nerve cells fumiiiig what are known as the plexuses of Meias-
ncr and Autrbach, but ue shall postpone for llie present any dcacription of
thrac or of other peculiar aplanchnii.- cells.
§ 99. In the eerUral n«rvou» mjfti-nt nerve celln are fotind in the so-called
jroji matttr only, tbuy mtf alwont from the while tnatfrr. In iho gray maUer
of tlte spinal cord, in tbe pnrln .-iiiokon of ns the niilcrior comua, wc meet
with remarkable nerve oclln of the following cliarucier;. Th« oelU are large,
varying in diameter from oO^ to 140/'. nnJcach coiiai»t8 of a cell body sur-
rcHiiiding a large conspicuous rcfraclivu nucleus, in which is placed an even
still more conspicuous nucleolus. The nucleus resembles the nuclei of the
ganglion cells already deHcribnl, and tJie cell body, like the cell body of tbe
ganglioo cells, is composed of finely granular piotoplasni. oflen fibrillated,
though generally obscurely so; frequently a yellowish-brown pigmi^ut is
depiwitfJ in a jiart of tbe eell body not far fn>m the nucleua. Tbe cell bodr
is prnlonged wimetimes into two or three only, but generally into several
procewoi, which appear more dislinctly librlllat^l than the more oentral
partM of ihi- ei.*ll bmly. These proceseeH ar« of iui> kinds. One proocia, and.
apparently, "Ue only, hut in tJie case of tbe cells nf thi- anterior curuti,
always one, i» prolnugud a^ a thin nnbranched band, which n<liiiiiH a fairly
uniform dinmcle.r fur a <-i)n9iderable dlstaiioe from tht! ci^ll, and when suc-
oessfully traced is found Hooner or later to RCjuire a nKilulln and to become
the axis-cylinder of a ncrv-e fibre; tbe proccMce which ibii* pan out from
the gray matter of the anterior curnu through the while mnttor form the
anterior rools of the spinal nerve. Such a pruccM ix uccordingly called the
run»^liiuii-r proKJif. The other pmcowft* of the cril rapidly branch, and so
divide IdIo very delicate tilanicnts which are soon lost lo view iu tbe aub-
•laBee of the gray nialler. Indeed, the gray matter is parlly made up of a
plexus uf delicate tiiaments arising, on the one hand, from thedivinoD of
procCMSS of tbe nerve cells, and on the other, from the division of ifae axis-
vylioden of fibree running in ilie gray matter.
dk
OKKBRAL riATL'RBS OP XCRVOt'S TISSUES.
167
TWceltt ■ Dfit HirroiiniM like the ganglii>ii cell bj^aHislinct sheath. Aa
*r ifcall mr laur on. nhile tnatint; in detail of the ceiitnil nervous syHtem,
kU iki nvn'oup rlcnirnw of chL< spinal cord are supported by a network or
yywtfrk of delivnte peculiar liaHie called ngtiroflia, aiialofptus to and
Mm| Btuch lite nine function m Kiit diHereot in origin and nature from
cHMlttva tiaua. TIub neurofjlia forms a ahcolh to ihe nerve cell and to
|lt|>nBMa, M well ns to ihe nerve fibres runnin); both in the white and tbo
piTBdilter; li en ce within the eeiilral nervous Bjelem the fibres, whether
■MilnlUtcd ur no, poasc no Mrparate neurilemma: tubular sheaths of tlvc
pfarofUa give the axis-cylinder and nicdullA all the suppuri they need.
All Ike nerve celb Dftbe anierior cuniu probably poase«s lui axiii cylinder
mem, and other e«iU similarly prnv><led ntlh an axis cylindi-r pmcnui lire
land in oUwr parti of tbe gray matter. But in reriaiu parts, o*. for iiwtaiicc,
btlw |RMWrior comu, many of the oella appear to poiaew no axi^rylindcr
Cs; In aueh«Mtaall the procMtei appear to bmnch out rapiilly into lino
nt«L Except for thi« abaenue, apparent or real, of iia axis -cylinder
fraoaa, Mch cwlls rOMUiblu in iheir iceneml liBatumt the celU of tbo ontsrior
(■nUithoagli thay aro getwmllv somenhai smaller. Sjitqikiug gvoarslly
ikmat feature of the ner^'e ccfhi of the central nervmiK «y*teni nf distia-
fined from the ganglion cells is the remarkable nay in which (heir pro-
MMsbmneb nlT into a number of delicate lilamcnta, oorr(»|innding to th«
dsDtala tlainenla or (ibnllir in which at lU termination in (he ticeuee the
tiit-crliiuler fd' ■ werw oftvn ends.
i idO. PmcD tbe alxive dcecri|itioti« it ia obvious that in the spinal cNird
maUdl ai repnwnling (he (vnlnd ncrroiin syMem we may at proiieint cou-
iM oataslfM, iMviug the brain tor later study) aSeront tibrw < hbres of the
pauior root) an io some way by meona of the gray mailer brought into
OMictian with efi^rent 6bres (iibrca of the anterior rnut j ; in other word*.
iklMfaMi oord ia a ctoiire unitini; afl^rent and eOtrent fibrvn. The npitial
pOfM am tkot centres in thi? tteiine ; the nerve cellti oompiMiiiig tin.' ganglia
maaiply rvlnys on ih« aStnnt fdirc* uf tho pouvriur root, they have no
(MdMjad whatever with i-tTvrcnt flbroa, ibey are conn«-ml uitti lihr«» of
a* kind only. Conoeroing the ganglia of the Hplaiiehnic Kynteni we eaniiot
bill cwMa make at present a p<Htitiri; stalument, but the evidpn<-e to fur at
MrdkpoHl pointa to the ooncluntnn that iu tJiem as in the ipinal ganglia
vdl aerve call belongs to fibre* of »im' function only, that whore several pro-
lof a cell an^ prolonged into nerve fibr«, thi«« fibr«« have all the MIID«
>, tbe nerro cell being, lu in the apinnl ganglia, a mere relay. We
I ■Wiihitofy evidence thnl in a gnnglinn the fibres snringing from or
id wilb one cell join nnolhiT cell so n* to convert the ganglion into
joining tugulber cells, wbuae nerve fibres have dilfcrent liiiictions.
r shall ban* later flo to bring forward evidence that tbe nucleated cell
\»ij iif a nerve ell in a ganglion or e)»«wbcre i» in some way or other con*
td with iIm- outrilion. the growth and repair of the nerve Hbres umnsing
I it. Bcaidm thir nutritive function the multipularcellnof tbesplaaiailiio
ipprar to serve the purpose ■ if multiplying the tracts along which
B« impulses may nassL An impulse, for instouce. reaching a multip'ilnr
I line of the proxiuiul (■ympalhelic ganglia along one Dhre or pmoen
tre in very many casea being a medullated &br«) <«n pOM out of the
■D various directions akxig several proeeasea or fibr«a, which, in tbe
ity of CMMs, if not alw«ya,are non-mcdu Noted llbrta. Thux theM' nerve
lOrgansof distnlnitiou for iinpubes of the wime kind. Wlnu further
liana of the imniiLM« thtu imsalng tlirough tliem ibeae ganglia may
t about we do not kn«n-.
I only in aome few iittiaiicea that vre have any indications, and tliose of
ailTKRAI. KKATPRK9 Or WEBTOPS TlSfltTKS.
vittidratrn froni the stiinuluH, or tlir movenieDt U colcuUml Ki push ur wipe
Bff ay the auuulus. In other words, a cenain pitrpme is evident In Ike refflei
act km.
Thus in all cases, exc«pt perhara the very wmpleM, ifae rmiremenu aall«l
fivrl)) by a rv&vx action are exceeoipgly coniplt-x cutiiiiarvH nith t\>a» which
result IKiii) tli« direct iititimlation of a motor trunk. Wheit tho piTtphcnl
Htmup of a divided Miativ nerve b siiiniilnted with the iaterruptcd nurmit.
the mosclcit of the )e)i are at i)n<-e thrown into tetanus, cviiitinue in the fawt
rigid iiinilitiim diiriD)^ the )NiitM|;e uf the current, nnd nrlii\ inimii|iiit4!Jy on
the mrr(-nt l)rii)){ nhiit otf. Wlw-.n (he «amc cunejit U iipiilicd for n iieociiKl
onlj, tu the i>kin nf the llnnk ofit hraiidcw hn^, iho 1«|; io dntwn u|i und tbt
fixit nijiidly ■wept over the fpot irrilitlei), in if to vii\>r nwny ihe irritnlinsi
hilt ihi« movirmr'nt \r a complex one. rt«|iiiring tho (iinirmction of imrticulic
miiKltti in a dcRniir BniuiTiio:, with a ciirvfuIlT Adjtiitod pmportitHi iMtweca
Ihe aniuiinlH of mnlriiciinn of the individuiiC nm»rlce. And this complax
movemenl. lhii> tialanc^l niid arranged i>cric» of contractions, innr be reneal«il
more than once as the result of a sin);le filinuilation of llw «kin. W hen S
deep breath ii caiwed by a dash of cxild water, the same eoordiiiaUNl aad
carefully iirrnnged aeries of eoutradions it also seen to result, as part nf a
reHex action, from a simple stinuilue. Aud toany more examples mjglii h«
given.
In such esMs as these, the complexity may he iD part due to the fact that
thesitmuluB u applied (o terminal sensory organs oud not directly to a nerve-
trunk, As we ijJiall see in speaking; of the senses, the impulstu which sie
geaeralad bv the application of a stimulus to a sensory orsau are more ooia-
pl«X than tJiose which result from the direct stiinulntion of a seiwory uerve-
trunk. Keverliieleu. r«flex actionti of ^i«at if not of e<(uwl complexity twf
be induced by stiniuti apj>1i<^d <lirvctly u> a nerve-trunk. We art-. t)i«rcAnk
oblijtcd to conclude that in a reflex action, the proc<*»e» which are originnKd
in tite <vntn- hy the arrival of even itimplc impiilM* hIoiik afivront itvrvA
may be highly ooinplex ; and that it t* the ^vMWtitutiiui nnd condition of tbf
cmtre which determine)* the complexity and character of the movemMlU
which are cfliNted. In other uorils, a contn> concerned in ii rvllex action it
to lie regarded m oonsiitutiuK a sort of molvcular machinery, the charadtr
of tho rcaulttng movementa being dotorminei) by iIh- nature of the machinery
Ml going and its condition at the time being, the chnntder nnd antount w
tho aflerent impiilseM deterniiiiing exactly what jinrta of and how far lb*
central machinery is thrown into action,
Throughout Ihe above we have purposely us«d the word centre, avoiiliof
tlw mention of uerve cells. Hut iiuaoubiedly the pari of the spinal eora
AVting OS centm of reHex action is situated in tlte uray matter, which gray
matter is chamcterixed by the presence of uer^'e cella ; undouhiedly also [lie
etlerent fibres are coniiecied with the afTereni libres by means of cells, cer-
tainly by the eelle uf the anterior oomu described in $ !>!* aii<) pruhuhly also
by other cellf< in tho na8terii>r curnu or elsewhere. Hn that ii n-tlex nition t*
carried on undoubteoly tfirouffh cdls,. Hut it docs not follow that a cvllular
mechaiiisui is eeseiitial in the sense at nil eveiitji tiiat the unelei of the evils
have anything to do with the matter, or even that ibe most iniporinnt of the
mnleviilar processes <H)iistiinting Um change* taking places in n cenin- during
n rellex iicliim an- cjirn^'d mit onir hy the cell suliftanc*' imuuxiialclv sar-
roiiiKliiig iht! nuclei. The jiowcr of carrying out a reflex action in proluibly
contingent on tli« natun- nnd arrangement of a xis-cyi indent, a»>l of the
branching mstrrinl hy which in a nerve centre tho alfetvnt and efferent nxi»-
cylinder* arc joioeil togi-ther, the nuclei intervening only so far as they have
to do with tho growth and repair of the nervous material.
GINERAL PCATUHES OF NERVOda TI88UIS.
169
Mirmil impalws, nisy mil forth it cuiivuImvo liluf cuuf;hing,in which n verj
Ivyi auinbvr or niutcln xrc thrown into rioletil contrftcliuna; vberMB lIh-
not tiiBtirl of (he hnir wiih oilier eurliiccn of the hotly mny produce no
(ktltra cflisct at atl. Similarly, whilo in the brainlcfe but otherwise iiarmal
ftvfti ■light touch lU) the skui of the Hnnk will produce uolhin^ but a fuint
titt of the underlvin^ muitcl^ tiie Bame touch on the same part of a frog
ptMMl will) slfychiiiue will produre violent tasting teianic contntclions of
MVlt di the mutctee of the body. Motor inipulses. at ire have bc«d, travel
tbof OMlor nnrreit «ttliout any ^ttM. expenditure of eoergy. and probably
widmt IncreaMiiK lluil espeudJItiru » tbey proeii-^d ; and [W same is ai>pa-
nHlT iW CTur with allvreiil impulH» paunne alonj; aft'en-ni n«rv«). Wheti,
hwVrr, in u n:f)i-x mti'iii alli^reiil impulses reach the nerve oeuira, a cliSDjte
fallwMtDrv Btul uw)^iitiid« of the iiupulM« lak«s pinoo. It is uot that in
iWMrr« ectiin.- Ihe alfi>rvut impulse* sn simply turned aside »r reflected
bill riltcvnt impuliMX: aud hence the name " rellcx " uctinn is a bod one.
It n rather that lh« nllStreni impnitea ad afreah, at it wen.-, tu a Mimulua to
Ibanrv contiv, producing nccnrding to circnmstancea and cvHidiiions cither
■ Ira >cak vflintvnl inipulMa oraintiltituileof utrong one*. Thv nerve rcnire
Btt W rvgardeil m a callcoiion of c xpluaive chargi« ready to be dUcbnrged
■il M> lo atari oHervtit impulmy along cvitain vtlm-nt ncrvcv, and thote
ikmtarvaoarraUMd rim) w rcJatnl ^l crtntn ttfli-rcnt uerris, that alli;reiit
isjalHa loichinr ua ocntre along thom ii(Tvn:> may in one t'iuw dwchnrge a
Iff anljr of the cnargot, and so give rim to fwhle movonionU. and in nnutlMr
^ttiBlAmtffi a very large numbor, and eo give rise lo large and violent
Mnmnta. In a reflex aolioD th«i (h« number. inlen«iiy, chnracler. nnti
ttuflntlon of the eOerent inpuleea, and so the kind and amount of raove-
■nt, will depend i^htefly on what takes place in the eciilre. and this will, in
Hro, ibfwnd, on the one baud, tin ihe cunditioo uf the i^enire. and, ou ihe
mWt.oh il*e qiectal relatiooi nf the oeulre of Ur- afTen-Jil iinpultiee.
At tilt ttme lime we ore able Ut recognize in uiunt tuDi-x notions a certain
between ihe Mrenglh i<f ihe utimuluH, or the niaguitudi; of the aller-
impuhea and the extent nf the movi^mrnt or tbc magnitude of the etlVrenl
The nerro-ocn Ire remaining in the unmv ronditiuu. the sironger
iraan inimeroua atftrcnt impnUcs will give ri^ t» the more forcible or
■or* KimprahrnMiTe niciviMnentH. Thu«, if a flank of a hrainlen iVog lie
nn lightly tuuchfci, the onl^- reflex movement which is vifihle is a nligbl
ttiidiiDg of tbo niuaclm lying inimtvliately tindnrnralh the spot of skin
itiaaklMl. If Ihe stimulus Ik- int-rvnsc-d, ih« iDovoments will sptvwl to the
Ukfrkg nf (lie ■aim- ojde, which fn:(|tiently will execute a niovemenl ealcu*
hud lu puab ta wipe awny the rtimulu*. Ry forcibly pinching the santc
^>if skin, orotkerwis'- inrrcnj>ing the stimulus, the rt^iilling movemenlp
aai In- led to rnibran^ ihv forr-leg of the same side, then the opivwile side,
•M Gaally. almiwt all th«- inusclee of the body. In other word*, the disturb-
I RioiBg in Um' cvntre. confined when Ihe stimulus is slight to a small
of ifae centre, overflows, su U> siieak. nbeii the stimulus ia increased, to
pane of the eenirv. and thus throws impulses into a larger and larger
tt of e&rent nerves.
Wt laay Mid. without going more fully into the subjeet here, thai in mon
ntrxai^oua a sjiectal reialion may be observed h«tweeu the jiart stimulale<l
' tke resulting mureraeoi. In the simplBSt eases nf reflex action thb rela-
ii turrely of such a kind thai the muaoJca tbrowu into action are tbwe
I by a motor uerve which b tli« ftiDow of tbe sensory nerve, the
latiiMt of whiHi calls forth the moveittent. In the more complex reflex
iiiM of the brnJuleai fnig, and in other cawa, the relation is of aucfa a kind
tbe raaulting movemeiii heai» an adufAatioit to llic stiniulua : tbe foot i»
170
UEN£RAt. FUATlfKUS Uf N£KV01IS TISSL'CS.
withdrawn rrom the slitnultu, or the
In other wo:
llUB
urd?,
renient is ai]eulftt4!d to push or
a c«riBin purpow is evident in the rvBez
awftv the stimii
IK-'tioD.
Thus ill nil cnsee, eioept perhaps tlie very Mtnpleat, the inovenienu unlM
forih hy a reBex action are exceediii);ly complex cDiupnred «iih thoM' which
rtnilt from the direct atimulation of a motor Inink. When Ihe f)eH|>h<^raI
B|iiui[> (if a divided sciatic nerve is sliniulaied with the inlerrupted etirreni,
ihe itiuiiHes of the leg are at oiiee thri^vrii inln tetuuui>. continue iti ih« ^stne
rigid Liindiii'tu (luriii); il>e puiMky:e of the current, aiid relax iinmediabely on
the curreiil bein^ abut oS*. When the Miiuie current u applied for a aecoad
ouly, lo the skin uf the flajilc of a bminlenii {nig. the left iiamurn up uihI the
foot mpidly Nwrpt iivur the «pi>t irritated, tu if t» wipe away llie irritation :
but thi* niovemi-nt \» a complex one, nyjiiiring tlie (■oiitnictiuu of jittTticular
hiumIm in It defiiiilu *ei|ut:iiM!, with a cnrefullv udjuflcal (irojiortiou bctwcail
the Binouiitu of contraction of the individual nnMtcloi. And thtf complex
movement, thi* biilancnl and nrnin^i Hcrioi of coniraotiomt, mav ht rcpmitcd
tuurv thiu) onov la tlit- rcsidl of a *inglc MimulalioD of thv slcin. W hen A
deep bnutth i» oiu^'.d by a dni>h of coh) water, the Minie cnonlinntcd nntl
cnrciwlly nrniiiged Hcrit* of coiilrnclioni' ij« also w«n lo nviill. n> part of a
rcflax action, finom a simple stimulus. An<l many more oximiplM might b«
In such cams iis thesis, the complexity may be in pari due lo the fact that
the stimulus is applied lo terminal tvuf^rf organs mid not diric<ctly (o ii uerve-
truuk. As we shall see in speiiking of the senMs, the impulses which are
generated W the application of a stiiuiihis to n sensory organ are more com-
plex than those which result from the direct stimulation of a sensory iierve-
tnink. Nevertheless, reHex actions of great if not of equal complexity mar
be induced by stimuli applied directly to a nerve-trunk. We are, therefore,
obliged to conclude that in ii rutlex actioti. the |>rocucie« which are origiuaic<l
in the reiitrn br the arrival of even simple impulua along oflitreni ncn-a>
may Ix) highly complex ; and Ihul it i» the couMitntion nnacondition of the
centre which determine* the complexity and chnriieter of the movements
which are elTecied. In other nords, a Mtutre cuncerned in a rtdlcx aciion is
to be r^nrded as constituting a sort of niulecular raachinorr. the character
of the mulling movements liemg detr:rminc<l hr the nature of the machinery
Mt going and its ondrtiou nt the lime being, the chariicter and amnuni of
|]ie aRi-rent impntm-s determining exactly what |>arts of nnd hoir far the
ccnlntl machinery is thr-iwn into iietioD.
Throughout the above wo have purtMisely used the word centre, av')iding
the mention of ner^'e cells. Itut um)i>nblcdlr the part of the spinal cord
acting as centres of rcllex action is situated in the gray matter, which gray
mailer is chanicteri/^d by the prciience of nerve cells; undoubbodlr also the
eflerent fibre# are connected with the atleivnt libres by means of cells, eer-
taiiily by the cells of the anterior cuniu (le«cril>e(i in !i ti!> and probably also
by other cells in the jx'iAti-rior eornu or eiaeivhere. So that a retlex acliou b
carried on undoubtedly Ihrougli celk Dot it docs not follow that a cellular
m«chanism t< cwentiul in the sense at all eveuU that the nuclei of the cells
have uiythiog to do wilJi the matter, or even that Die niosl importam of the
tDolet'ular proceiKii conntituling the chanKca taking plai-e* in a centre during
a rellcx action ore carnal out only by the cell Hul(siant« luimediaiclv mir-
rounding the nuclei. The power of eiirrying out a reflex action in ]>n)liahly
routingtmt on the nature and nrranjiement of axiii-cvlindcrt, and of the
lininching material by which in ii nerve (centre the alVrrcnt nnd eHi-'ivnt axis-
cylindcri are joincil Ingethcr, the nuclei inl^-rvwiiiig only so far as Ihey have
10 do with the growth and repair of the nervous material.
OKffftllAL rKATL'tlKS OT N8RVOCS TISSCKS. 171
: 101. Aidamatif iKii&nM. Efforeoi iin|)ulM» froquently iiiiie frmu the
~ I tad vpioal cnrd mi)<l su give ri«c lo mnrpmcntfl without Mag ubvioutlj:
by ftnjr BtiinuUtiuii. Sucb louvciovnts bit *pcikfti uF na HuloitiKtiv
auft. Th« d&fvnt ioipulMs in xucb cuws tire started bj chaDgw
I Bwva ccnire irKich tirv tuA tbi.* immediute r«Hilt of lti« nrrivxl At tlic
lOMUv of «llen-Dt impulse* from wiihoiiE. Hiit irliich appcitr to ariu* in
mv* OADltx' itwlf. CliHDi^ of Ihis kind may r«i-tir rlirthmicslly : lliue,
llhall te*, we tikve rcaMn lo thinit ttml in n cerlttin part of the mt-diilU
. cluuiKfa of the o^rvotiB materia), recurring; rhythmically, lead to
kythmiv dwrltarKv alone c«riaio nervM of ei&renl imputic* wherry
I conneoted wtlli llii< cbcei nr« riiytbinically thruwn iuto soliun and it
_ . nically r«peaioil brealhinv is brought about. And otli^r Kimilnr rhyth-
Htaioiniilio inovt:iii«iitt< may m atrri^ti out br other parts of tli« spinal cord.
Pnn Ibo bmiii itself a much mure varied ami npfiar^uily irrt.-|{ulur di«-
ttgpat rtlirviil iiii])ii)»t«. net lh« obviuiia nvull of any iminvUiaivl^' for?-
fill illtTeiii impuliMtt, and t)M-rff»ni not formiiijt part of reAex actiona, it
WJ atintnon, miidtitiitinjt what we apcalc of as vulition, eflereui iinpulsM
ikatuiiiib); being called voliltunni or voluntary itopuUe*. The npinal oord
qwifRMn lite limin dow not appear aipuhl« of «zAoiiting thiM! voluntary
BMonmi* : but to Xhm au1>J4«t w« tludl retuni when •m come to ii[ioak »f
iWnnual aarvoiu nyvtem in detail.
Wtmid juRt now that th«rc i* no antisfaciwy ev!i)cnoo that t)i« ganiili* of
ibifUiichnic •nrMnm «vcr ad w (M.-ntrc8 of ra flex action. TIm eruleiio*,
iuiriv, that tnow ganglia may Mrvc as conlra of rhythmie Mitomatlc
■MOM at firf>t eight of »»mc stirngth. Several organs of ihd 1>odr
sing naecular tij«ue, the most notabln being the heart, are during Vitt
'\ IB riiytfamic autotnalic iuovenieDt«. and in niiinv cato* oonlinne thtf*
inis aA«r renMVal frf>m the body. In nearly all titeae caaea (rau){lia
vtpnacnt in oonnectlou with the mutu.'ulur tiMM«: and the preaeooe axul
nMd euulitjon of th«n puiiiiia t-am nt all evenu in many ouea in tone
«iti«ent«al to (hi- due |iprformBn(« of (he riiythmic auiontalio niovemi-nla.
bjnd, it ha* brvn ihmitihi thai the iDovementa in rjueition are DMilly due
biW rhythmic auiimialtc generation iu the cell* of thcai- ganglia of uiTerout
■ifalm which paaHngdowo to (he a|>proprial« muwular fibrt* call forth
kr ifcrtliraic movameDt. WIku wv come lo otiidy tlicao mi>vemci)t« in detail,
Wifaall find nMoiu for coming to the conclusion that this view in not sup-
|Mlal by luieqttMc «vi(l«ocfi ; and, indeed, though it i* perhaps immacurc to
•iki a dogmatic ateiWMMit, alt tlw evidence goes, aa m« have alrcailyMid,
titfew tbat i1k- great use of tlw ganglia of the snlanc!hDic ^sleu, like that
rflbtlpiltal ganglia, is connectmTwith the nutrition of the nerves, and tbat
Aaaanifllam do no* like the central nervous system act aa centm eitlicr
Mt«atic or reflex.
f 108. Inkibiliiry nenf. We have said that the fibrea of the anterior
nNiboutd be calM eflerent rather than motor, becau» thou);h they all
any inpiilaea outa'ard from (he central nervous aysteni l<> the tiwttw, the
tataUa vhich they cany do nut in all casea lead to tlie eoutractton of miu-
(w Cbrva. Some of titeae eifervtit l)br«« are distributed to j^laodular ttruo-
>Mm, ftir instaoM U> the aalivary glanda, and impulses pusaing along these
lad Id changea In opiibdlal eelu and their surroundings whereby, without
uymucnlar oonlrai-tion necvMMrily inlerveniDg, xecretion u brought about;
tkt sdioD of tboM libret of accrbiiou we shall study in coniteciion with
BaHia (his ihert are efferent fihre)> going to muMuIar tinue or at all
enats tu musmlar orgaiw, (he inipulM* piii<iiin>[ alung which, so far from
Miimif almtit innN-nrnr I'ontractiun. diminisli, hinder, or Mop moveniculB
172 QENEBAL FSATUBBS OT NERVOUS TlflSUBS.
already in proeress. Thus, if when the heart is beatin); regularly, that is
to Miy, wheii me muscular fibres which make up the greater part of the
heart are rhythmically contracting, the branches of the pneumogaatric nerre
going U) the heart he. adequately stimulated, for instance with the interrupted
current, the heart will stop beating ; and that not because the muscles of
the heart are thrown into a continued tetanus, the rhythmic alternation of
contraction and relaxation being replaced bv sustained coatraction, but be-
cause contraction disappears altogether, alt the muscular fibres of the heart
remaining for a considerable time in complete relaxation and the whole heart
being quite flaccid. If a weaker stimulus be employed the beat may not be
actually stopped but slowed or weakened. And, as we shall see, there aie
many other cases where the stimulation of efferent fibres hinders, weakens,
or altogether stops a movement already in progress. Such an effect is called
au inhibition, and the fibres stimulation of which produces the effect aic
called " inhibitory " fibres.
The phenomena of inhibition are not, however, confined to sucb cases u
the heart, where the etferent nerves are connected with muaculu- tisBue&
Thus the activity of a secreting gland may \t^ inhibited, as for instanoe whm
emotion stops the secretion of saliva, and the mouth becomes dry from fear.
In this instance, however, it is probable that inhibition is brought about not
bv inhibitory impulses passing to the gland and arresting secretion in the
gland itf^lf, but rather by an arrest, in the central nervous system, of the
nervous impulses which, normulty, passing down to the gland, excite it u
we shall see to action. And, indeed, as we shall see lat«r on, there are many
illustrations of the fact that ntferent impulses reaching a nervous oratre,
instead of stimulating it to activity, may stop or inhibit an activity preri-
ously going on. In fact it ti< probable, though not actually proved m every
case, that wherever in any tisiue, cnei^y is being set free, nervous impulsn
brought to bear on the tissue may affect the rate or amount of the energy
set free in two different Wat's ; on the one hand, they may increase or quicken
the setting free of energy, and on the other hand, they may slacken, ninder,
or inhibit the ^letting free of energy. And in at all eventa a large nnmber
of eases it is possible to produce the one effect by means of one set of nerve
fibrcH, and the other effect hy another set of nerve fibres. We shall have
occasion, however, to study the several instances of this double action in the
appropriate jtluces. It is sufHcient fur us at the present to recognize that a
nervous impulse passing along a nerve fibre need not always set free energy
when it reaches its goal, it may hinder or stop the setting free of energy and
is then calle<l an inhibitory- impulse.
The blood, ns we have nid, is tlic inlenial luedimn on which the
tiatt lire ; from it these drair their food and axvgea, Ut it tber give up ilio
ptdmU or vme inattere which the}' Ibriu. Yhe tiaauM, with toiiic fuw
aaif&out, MTt tntreraed l>r, and ihua the eJeraeolB of the tiaiuei Humuwled
h, Mtvorks of mtuute lliin-wstled lubee. the ra/iiflarii bloodvetneU. Tlie
timmiuar Mriuled tuuscle fibre, for iostaiiee. ia surnHiiideil by cnpillariM.
nmaf in the (vnmevtive (iaaiie otileitle but iluiM; lo the aarcolemma, nrrangan
biMKcrk with uKirt- or leM n?ctaugular nit'.ibeN. Thriu- cupillnrtm urv
dtNd tub<« nith <NjntiniiuuK nidti, and the hhmd, nbich, nit nc xltnll ttx, u
flMmallf Hrcamiog through tbem, a at a whole »>iili(i«d lo tJicir channela
ml iom But cacapv fn>m theiu. The elementa of the tiwtiM lie out«iile tin
HulkriM «od forai ettravimcular xMrit, nf iliHbmit forin nnd «iie in tliC
diMat ti«UM. uirroiifiiU.ll hjr I'ttpillury nHworlu. Hut the nali» of the
afQUnia an •» (liin and of nuch n nutiirr thul certain of the const itii elite
iftW blnid pim frtiiu ihe inlrrior tif the rapitlan' thrr>ii)^h the rap) I In ry
nil III the clcnicnls of tbt- Iimuo ouUid* tlio capillary, and siniilarlr certain
•f iht cuostiluentfi of the tissue, to wit, certain sutwtanoes the result of tlte
■rtiimltiiii coolinually K'^inK on in the liaeiie, paaa from the tiwue i>iit<iide
llNi;af>illarTUir<>uyh tlte <:apillary wall into the blood flowiiig throujih tbe
afilhrj. Thua u we have already nid, $ 13, there ia a coDliuual int«r-
daim of matenal between the blood in the capillary and the elciaeudt of
ikuaite outside the capillary, the Irniph acting aa middleman. By UiU
bhtthiaxv the tifuue liv« on the blood, and the blood \a aflected iiy il«
(Mife tbnHi);!) the tiMue. In the aruull aruriea which end in, aud in the
«*U niiM which Itejcin lu the capillance, a dmilar interchange take* place :
ta the amount of iiitcrchajice diminishea us, paatlng in each direction from
IW capUario, thu wall* of the arteries and veiux become thicker; and
iKkH, in all but tlu' minute veinH luid arlerin, the intrn'hange is m Miiall
ika ii may prBclically be ncgU'Clod- It in in the cnpilhinnt (and minute
antria and Tcin^j that the bui<iiieM> of the blond Lt itntin^ it i» in ihi.'w that
tW inierclianp.- Inkm |>tnoe: an<l the olijecl of thr vnu'iilar mcchnniiim is to
nan the blooil to tl>.>n- ihroiij^h tht-ve in n nianner IkwI ailuplcd for currying
« tUi bt«rchaiiuc under varying ctrcumstanoca. The ii«n of the art«nw la
ii the main aimpTy to carry the bluod in n auitablc mauucr from the heart to
thi apillnrici. t^e uae of the veina ia in the main aimply to carry tlic blood
hn the capillaries back to the heart, and the uae of lh« heart is in the
•ain viBply to drive the blood in a suitable iDaaitor through tlic arteriee
<M« the ca[Hllaric8 and from the capillnrim back along the vcitta to itself
■pin. The structure of these several p«n« is tidttpt«d to these several uaaa.
Tht Sirvtiiift of ArUriet, C'-apillarta, and I'eiiu.
}10S. On tom«Jt»t»rtM of mnfKWtfw (wti/-. The hrarl and bloodvaaeli
ut, broailly speaking, made up partly of iniwcular liaMe with its appropriate
174
THE Vi8CI
nervous elenienie, mid parlly ufcciiAiH vurielim of the imut- known n« con-
nvclive tii»ue. We nha)) have di xpcnk of Momv of ih- ftwlun^ of counective
tiuMO of i-ihynulo^iftil )iii|Hir(nii('i^ whi-ii nc cnint' to dcul wiili tho Irniphiitic
Sfsloni. for lliio synlwii w iiitimuU'lv iimormtct) with ooimi-clivi; liauv. But
an iiiWH'iatkiii rnilv Ii-m <-1>w OxUli^ lictwwn llio blixjilvnitrh mid oonncctiw
lisxip : for ojiintntivc ti«<iii- not nnly ontm Inr^ly. in ono or otiier of iu
fonnM. into iho viruciurr of th<i blood vrsvlo, but nUu torrn:* * sort of bed bulh
for Ibc Inr^r vctwln on tlicir way to and from llio several tisf ilea aud orgaiu
and for Ibc siniillor rcwde, including thtr cnpillarics, u-iibin each tiasuc and
or^nn ; ii>di.<vd. n cupillarr may bo regarded as a minute tubular [WMage
hollowed out in the vi>i]Rcctive tissue whlrb binds together the elenienu of a
ti»eue. It will bo dmrabK thorefbrc, to point out at ouce a few of the
chnractc^rs of cunncctive lissup.
Tlie coBDCClive liwuc of tbe adult body is derived from certain imviiblutic
i-elU of the embryo and cDDsietS eseeiUially of certain celU, nbitdi do not He
in close contact niib eacb other as do liie celU nf i^pittieliuni, but am
if[iafated by more or lest) inler-cellular luaterinl nhicli tuny in <x'rtnin c«Ma
be fluid or semi fluid but which is {(enerally wHd, and in oitiiinioiity ojiokca
of as matrir. [n mu»l fonuB of connective tiwuo the matrix in rclutivvU- ao
abundant and proruini'iit. that the (%1U, or mmeetivf H/trw. rorpuitciM as tWy
art' callcil, become incouiipir'nousi; and, speaking gon^trally, the value of
cnnn«ctive ligsue to the body degMtnclit innoh mora on tho ([ualitJes of tbe
Rintrix ihiiu on tbe activity of the connective lL-«ue corpneolm.
Tbe kind of coiincclivo tStnur, itomclinii'H cjilb^d " loose conncctivu tissue,"
which wrajw round and fnrmn ii WI for thi; bloodvessels, cunsista of an
irregular iDCuhwork formed by tiitorlncing bundles of various snes wbidi
leave bctwucn llicni ipacM of ver>- variable form niid aiie, eonie being n>ere
chinks or clen«, uthcfs being larger but generally Haltened paawges. all cou-
laiuiog lymph and having as we shall »ee s)>ecial connections with ibo
Iviiipfaalie vessels. The larger spaoee are someiiiuee called "areolar," and
i)ii» kind of connective tissue is aometiines spoken of aa" areolar ti»iu«."
When a itnmll portion of this tissue is leased out carefully under the niicr»-
scope, the larger butidli'a may be Hejiaraled into liner oundlM, and each
biiitdle. nbidi gi'iieriilly pumui-^ n navy ourxe, has a fibrillated apjitnmnu;
W if made up uf (?x(^t.itia^d y tim: fibrilltc; treated with linie-watcr or haryta-
waitr the bunillendfiartually njilit up into fine wavy fibrillicof lott than 1 (■ iii
dinmetcr, a aulMtancc of tt iieirultar nature which iirevioiMlvcciucutcd tbe
(ibriihe tiiftether being dii9olv<«l out from belwuen th<!iii. Wlion a mass of
micb libritln i* bnikil with water, llicy beoonie converted into ^tnlin. n sub-
Htancv containing, like proteid material, carbon, nitrogen, hydrogen, and
oxygen, with a siiinll ijunntily of "ulpbur, but diflcrinjj from ]>ri)tt^id material
both in its percentage, compwition, and in it« proi>crlie». A remarkable and
well-known feature of gvUiin i* that its solutions while fluid at a tempernturo
nfliailing waltT or Iciw, become solid or a"j(illy " at lower temperatures. The
untouched (ibrillic. In their natural condition, bohavo as we shall se« in
B|>eaking of tbe digestion of connective tissue, somewhat iliiTerenlly from
prepared gelatin : ihe natural Hbrilla, therefore, docs iiot consist of gelatin,
unt of a substance which by boiling is readily converted into gelatin. The
subetanco soluble in lime- or baryta-water, which cements a number of
Bbrillae into a bundle. up|ieat« to be allied to a body, of which ve shall
speak lat«r on, called mucin. Since the fibrillie form by far the greater jMrt
of tbe matrix of connective titwiie, a ijuautjty of this tissue wl>en boiled
seeniB almost entirely <Hinverled into gelatin.
In collective tixaue, Uien. a number of exceediuu;1y line gelatin! ferou«
fihrillui ore ceinenteil together into a fine microHcopiu bundle, end a auinbcr
riATUKKS or VASCULAR AI>PABATUS.
175
•f tbtM finer buiiilk« muy be aitnilarl}' cetDetitetl tocether or aiiuply appowl
HfitWt to fnnii l»r)i;«r buDdl«e ; Mtue of the bundlee U leut «p|)«Br more-
•nr 1.1 be dr-tinnl by ■ ile)ic«tc tnin!t|iar«nl ^alh of a Minewhat peculiar
oiUtT. A tiiinilirr of these buiKllM, tinall and In rev, are arranged aan
wihaotk, (hi- irD'gular tpacn of nbieb are oi^upieO by Iviiipb. On lh«
«ln<i ibr tiiiiiiMtv rnnani iho ipiiCMi or b«tnw:ii llto )>viKli<-s wh^^re titvw
m a iiifuBilti)!!, iil\r» lyiii)f in nitimlr tpuctd hullnwe<l out in ihi- ii-inoiit or
pMnd Nibkiitnc)- uiiitior ibv binKlIra, aro rmiiiil the i-nDUiM'tivi! (twiK- cor*
fchfc I'jich >if thcM- it a ri-ll niQtiiiitiiiu "f ii mic]i-ii>, trritcnilly oval or
ilwpf r . •urriiuitili.ll by it pnitoplai'mH' ovll body u»iinlly inv^'uhir iti lurni.
Majt Kinirliii)i.-a invrvty xpindli- fhit|icd. bill nxm Tttf^ueutly dnlindly
kwM or iii-llatc. and iH-iirly hIwmt* inuoh HatiitKi) in ■ plaiw rorre-
ifNtdin).' to tbr ilirrvtion of the fibrv*. or buadlc* of the Rinlrix. Although
■ or hnvc mid lbi> librillir ara ccnmud toscth^ into ■ buitdic, nch Kbrifla
mtia* iiilRdriitly dixtinct to hav* a roareoil rolractm- vlK-c< uii raya of
%ld lUliDg upon or traiwinitml ihrough tbr ttmiti'. «u thai iho bundW
iffor while and opaque ; ht-nco this titwutf, and specially n morv dense fi>rui
(< it. it toraelimo s]>oki-n of me irhilo tibrou* tiemio. Oirinz bi this onority
ik nofr df limle connective linue corpuBclcw arc not rctinily visible id the
MUinl nxulituiu of (h« tiHue. They may, however, U.^ brouglit (o view by
ikaclko of dilute acid «uch as acetio acid. L'mler the intliieiiee of lbi>)
mIocIi flbr^lta swelb out. and the swollen tibrilliv prcv^ng u|>i>n each other
na* Id rcfnct li^ht so mush M befbte. Hod thua become more trnnspnreni,
niyiDWPh as an opaque maes of ttripa of tringhui beooinn tmnepareni when
iWtffiia are swollen by bojliiijf ; ihia incnaaeof irannpnrcncy aJlows lli«
■wpamn, whieh are not mrullen nul rather •hrunkeii aixl made mor« opaque
fcf ibr artitm n( llie arid, to become visible. The prcwnoe of these oor-
fad* nay nltm be rovcali>d by lh<! ui«: of MUch Muiuing rea^ccnu a«. whil«
■M 4ainio^ ibo fihrillatol matrix, ftain the nuclei and the proioplanmlc
kiCa of tbtt ciirptivJoK
BmUm time hmnchrd irregtitiir-llnttrned oonnwtive timiio cor^iUHclea.
«kMi do Dot oalurnlly exhibit any nnio-boid moreinentf, Icucucvte*. exbib-
iuagmmor Inam-tivc moTcments. are fiitind in Iho spncoa of llMtianw,
kftUer with cnrpitw-leaof a third kind remariinblo frxmi the la?]^ cnartr
fmaaWa «itb which the cell Nihalance ii etudtlod, and known na " fdaama-
tufmim."
(lot. When eonnectire liwiie ia rendered tnin«pare»t by the action of
•iilnr acetic arid, there come into rieir befldes the <»rpiiiie]ee, a number of
Ua« iGfletvut I'nHn the ([^laiiiiiferous fibres not only in not beins swollen
•adiOKlered tranapareut by the action of (he acid, but also by tneir slae,
thlinly acuity number, clear bold outline, and sharply ctirvetl oourw. The
Amwjr much ia uie, aoine licin)t very fine »> as to appear mere lioea,
vkQi Mben are very )arj;e with a diutin<.-t double outline. Whether small
vbiye cwb 6bre is a linvlo llbre, mx a bundle, and cannot be split up, like
albviirHnill bundle at the ordinary matrix, into fibrillie. ^ol only id
thiir nxirvF iharply i-urrcd unlike tlx.- gently aweeplng outlinea of the gela-
liaifmius fihree, but iltcy divide and aiinntamoae fte«ly, thus furmiDg Dct*
•wi* of vnrvtnic shape : the gelatiuifemus fibrillie. a» the other band. uev«r
diriia. aiwl tlie Bundlea int«rlaee into a network rather than anastomuae.
Tie Bumbrr of tbvvo librea occurring in fonincctive iiwue varv much (b
difcent «i(UBlionii, and ia aone pliiii% a.*, for instance, in the li^mfntmm
■aricr t4 certain anim^ iMBrly ihc whole liii:<uc ih ct>mii(ii«d of lar|^ librea
(f tha kind, having in the man a yellow color, (he oniinary gelatinilcrous
ihiv biiiig redurca to a miniinuni. In nich a ttttualiou a remarkable phy>
<Wcku«ctar of tbtoe fibrei it catJly reeogniiMMl ; ihey are in a high degree
176
TllK VASCULAR MBCHAKISU.
GxtMwi1)l« Mid elaMic; hence lliey are fret|uetillj' called e^aW>>jt'i>'^>: lt»»Bl
ihvtr )'i-tlowUh nolor thev are sometimes oalled fellow elastic fibres. The
whitr grlntinifi'rouH iibrilfwi, uu tbe contrary, pMseas vary liille ex(«n>it>ility
or olairticity.
WIk-ii a {Hirtton nf HKiuoentiiiu diiuIi% \a frettl hy iirolonged boiliug fv>ra
th<! n-mniiiil of gclBtiniliiroii* fibrt« mixed iiji wilh ilio velluw claolic inaU-
rial, tbs Inll'.T t« limiul on cbemiail tn-iiiineiit to yi«l(l a mibtiiaim called
tlattiti, nhicli very clowly nveinblts [iruteid matter m clemtDtarr cotuno-
sUton, excvpt llint it contain* no fulphur, and wbicli yet probably diftera
wid«ly IVom it in nntiirv.
Conneclivo tiwuc Ibwi con»xti> of a matrix of iuexlciisiblc in«la»tio irhile
navy );GlatiiiifrrouH tibrilUv, ccmcnU-il inm bundlr* (iht- bundl» Xitat^
arranKeil. in Io<«o coiiiiecitrc tiAtic, in irregular mt^hunrlci) nilli which are
ttwociated in varyitii; abundnncc anastnnnMiiiig euHcd ypllow clashc tibrea,
flud auion^ which arc embedded limnclirii coniictivc tietuc ciirgnifcle!!. I^U-
ooejUs aod pluma cells are itl«a found in th« raMbn or aroolie of the dimIi-
work. We may oou- r«lurn to the striicttirc of the b1ood<r«Mela.
% 107. (^tpUlarie*. A capillary if, iis vc wid abttve. a tubular psfsage
liollotved out ID counective timue. Without e|)cc)ul prejMralion all inat can
be K«n under the microscope is the outline of the wall
(ni>.U. of tbe capillarv, shoirinj' under hi);h powent a double
\ \ contour. and marked with oval nuclei which arelodced
in the wall at inU-rval». and which project aonieirOBt
into the lumen or canni of the vessel. IF iff. 51] Wbea,
however, the tis^'Ue coutalninK the capillaries u treated
vflih a weak eoUiltou of silver nitrate, and after being
thoroughlv washed b exposed to light the wall of tho
oapiUnry U seen to be marke4l uut oy thin black line*
into »pitidlc-«lm|)ed areon, da\-«ituling into each other,
and mi rrlaliil lo (lie nuclei in the wall, that each nu-
cleus occupie:< about lh<- ocntrc of an area. Krom thi>
and from other liicU vrv concludv that ihe laipillary
wall i» built of Hal luxifnrm nucleated plates cemented
together at their cdsrs by some cement subalancc^
which more readily abaorbs and retains silver nitrate
than do the plales themselves, and hence aAer irtAt*
mcni uilh llic silver salt shows in tbe form of black
lines the silver which has been absorbed, and suddenly
reduced. Kach plale is u Halteoed nucleated c«ll. the
cell body of whicn, except ti)r a remnant of undiilV-rni-
tialed protoplasm round ihe nucleus, haa becuimr con-
vened into transpareiU. diOereotiated material. Sinoo
the cells, except )br the minimum of ocnK-ni i>uhi>ianoe
botwceti tlicm, are in dose coiilaet with each i>iher,
wcmiglit Rpeakof them asformiiifc ana'pitliutium ; but
on account of their cell InHly boiut; reducnl to a more
plate,and(>tiaecountof their connection both hv origin
and nature with mcNoblaHic conncctivp ti<vtiv corpua-
IHm, it is convenient to speak "f ihi-m ns rpH/irlioid a-Uf or plnto^ They are
■pmelimes R|>okcn of as rndothrliat cells or plates. In a small capillary tbe
widtli of one of these epithelioid ))lal<« al its widest part, where Iho nucleus
lies, may be of nearly the wtmc size ns the rircumfervncc of the even-distended
capillary ; the cells consequent! v arc placed not side by side, but more or len
altarnate with each other, and their nuclei priiject alternately into the liinien
of the TCiML The larger capillaric» may, however, be su wide that two or
*-
'fs
\V
Hratimiiror * uvi>r.i-
nu. triKii Tkutub't
Hfia Sonmon ft Ni-
laiTS uj' SiLvn.
d. CMt. >. Thdr nil-
rBATORKS or TASCOLAR APPARATDS.
177
>
i-ellii lie iikore or l^se abrcasL OuUide the CKiHtlnrv, U'liich is tbus
ft Ikto whJ dclicale menilinine. a m«re {latcliwork of thin epithelioid cells
OMBottal toK«tJ»or, i» alnava fuunil tk certain nniouiit of couuective tintiue,
tlw wall of Uie capillar/ furniing at places jkan of llie witllfl >if the lymph-
boUiM eannective tiwiieqncta, aiid at oltier places Imiujj uiiitt^<) by cetui^nl
aataml tn the buixllt*, bands, or alwets of tbe same coiiitcciive lusuc. Not
itaf>«|ijmtly, in youuii; iimmka, branched counective lisnne c(trpu»c:les He
•i[*ia utd rtntiraor u CAj^llBrv, fonw uf tW i>ri>cesiet of the ckII beiiij; sitai'lied
U> tho KUlaidi,' of tbc rpitlx'liiiid iilaliv of t^i> eit[tillary. Kvcii in llic rapilla-
of aocb a Uiatie as niuscle, tW ni-lwurk i>f ciipillaric:! i-nilirii<-iii^ a nius-
flbra b alwavR surmuixli-d by a crtAiti ihuii^h sonictitnrg' a miiall
•MMai aaW of nmm-clive linue ; iiidwd, nhcrcver capiltnntii nin. tliey nro
aOMatpaainl. mt wv haw wiid, by coonertivp tiiwu«, to itint overywheru all
0**r lb*? b«dy Uie blmxJ in tbc capillnry i« «c|ii>riit«d frnin the lymph in th«
tfttcr* of the c>onnectivo tissue br iiolhiii)f more thati the oiceetli ugly thin
Wiw of the cemented epithelioid plates, ll must he added, however, lliat
th* ifiaees in the cuoDeclivo tbsue are lbeim>elvca fometimefl lined by similar
wuliclioiii platea, of which we slinll have to treftt in sfteakiog of the lym-
|M(KB, ttt that in placw tbe partition between tlw blood and tlieM lymph
mar Im' a double one, aod consbt of two layen of thin plut«t.
Ife KDJ ciiae, howovcr, the [lartitJon U an exceexliii^-ly thin one. and no |ior-
nniMi that it alhm-s nii Hile<iiiately rapid iuterchau)^ of inntenul lietwi.i'n
lWbto«lai>d the lymph. As wu ithall pri-rt-'ntly M^e. not ufily tluid.t— that
ii^mtten in wliilioo — arc able to piura thr»U);h the riurtitiou into the lymph,
UtinUct eurpusetm Wh rod wid while, ea|>ecially tlie latter may, in ovrtain
cinanMUnc**. niabf Iheir way thniugh, and m> paae from the inurior of tho
■fiBatr IdIo the Irniph »pace« ixitxide. It is probable, however, that then
mkt ueir way chivtly. if not cxvlitiiirelv. through the cement lines, and
■IMJally at ihe point where the <«inenl linc» of three or more cells meat
(DMlier, and where the cement Fub»ljiDc« exists in larger amount than elafr-
TW «te of (be capillaries is variable. In some regions of tbe bodj, for
OHUiiet, in the Iudk^. the capillaries arc, on the whulo, wider than in other
nti'ias, fur iiRiaucc. the xkin ; and all the minute vessels joining arteriea to
wisi and pfwwMint; the structural featurra jiist described — that ia, bdag
mil! spill arte* — will out always have the $au>e size, even iu the rante region
<l (W Dody ; tbe artrrr may gire rise to Uiffe capillaries which branch into
•■all QBpiUnrMM, and these iniir again join into lar^ecapillariee before uniting
tifctn raiiH. Thus one i-api^larr outy be so narrow tnut a single (mamma-
Su) red oorpuaclo passe* through it with diliieulty, whereas anuliier vapiil-
hr? may he ttid* enough to afford rotim for two or three such corpuscle* to
tr- i>i. Itt-viiles (hi*, ilu' same capillary, may, in Ihe living boily,
ii i:ii frxiii time m time. At one mi>meni, as when (lie etiininoe on
tt utrrwl rii\r is blm-kcl, or when liIrnMl. fur Home mason or another, ceasea
lalow iDt>> it. (he ciiiiilhirv may Ik- i.-nipty and eollaiwed, its walU in eon-
tut and its lumen nholishol or nciirlr so; and, in liawes taken from ihc
■bbl budy and preparwl for niicmtcoitical examination, tbe capillaries are
smmlly thu« cm|>(y of blood and (-iilliiji*"), »i (hat they can be seen with
etCnilty, appeanuK as (hoy then do as ulinu»t mere tine* with swellingd at
>iu«rvalii comspoiulini; Ut tlie nuclei of iho eoostituoiK celU, At anotbar
(IBS. as when blood is flowing into it a( high prwetm-. the capillary may bo
viddy diitnidfi). In the variations in calibre, the walls of the capillary
fJsj ■ paHTv |iart : the maleriul of the eirithclioid phites is extcnsihk*, and
lis fnman of tho bl<Kid within the capillary diBlen4ls the wa)U, aotl the
~ baing also elastic, tbe walls shrink and colUpae when the prewire
13
ITS
TUB VASCULAB UBCUANI3U.
ia itnovcd. being iiMwtcf) in this by thv prMsurc oT \ht lymph in (he ipM«»
OUlnd« tlic cnptUtiry. ItnC iHifidcs thi», in i> vnung nniinnl, itt all wcntf, Um
citpillnry wnll is, to » oi^rltiiii cxtpni, cvnirnuiito ; thi: <:{utbelioid ci-Ili', which
ihtrii a[>|M-nr lo cont«iii a liti't'O imiutinc of iimliftl-ienlinUxl protoplaixii. »n-m
able, iiii()<.T the intlticnco tif Mliiiiiili. to vhati^ tlirir fiimi. piiMiii); t'mni n
loiigirr aikI narrower nhape lo a fllvirtcr and hruiidpr otic, and (nii» iiiHucndng
ih? calibre of the iiibc ot* whioh lli«y Ibnn the weiIIh. How far f\ich an aclivv
chaii^'c of form lakes place in the cnpillariot uf the adult body haa not yet
been <letiiiilclv determiited.
Tlie ^iruQlure of the eapiDary then seems adapted lo two ends- In tke
firel place, ilu walla lieiiin penueablu iire adapted for cnrryioK out that im-
portant iuterfhaiixc between the blond uud liaatie. which, aa we have more
than nuoe Mid, laltcs place uliuiwt oxduMvely in the cunillary rejijioiM. In
tbcsec-ond pluru, the extenBibiltiyundelHAtiiity nf lut wallit permit it to adapt
■U ealibru lu tho amomit ami r>iri.-e witli which the bWd ii> llnwing into iu
^ 106. Arlm^. The wall nfmmintUi^ artrrtj, i. c, nf one whiuh imHHin abuul
to breulc up inlu capiliaricv, and which i:< •onieliniM i^pokcn of iu> an (irtm'«t,
coiuiitt* uf thtt following parts :
(Ptd. u.
Pm-a.
t'
1, ■twlkMlnfUTT ', ^ Ummiaun voaicl ; 3. «sn«r raiilllnnw : 4. Hiier cB|41l«tla a. Mruciurekio
nmbniie Mill wlUi waan i>ii«lvl. rt>[im(ntoll*e o/ Ihe tantca vitliiu: b, uuold of Uie biuku^t
Mf»««IU; I', iiuckl wlihlu tlin miiaII artciy, larbjti* apiionaiiillig lo ui mdoUwlliuiii A nuclei tu
Fbl ^— aiUU. AaiEKT Tn SaoH iiiK V<iiiui-i l.trt;iT!i uuicii I'DHi-itn n* IVauo. ■, tniiy
tfetSmn ;;ft. InUmal cImUi- luuiD*;f.rJrp>ilaruiiuculnr fibrrt ariliemlil>ll«ri>at:it. ib«vonn*ctlt>*-
danaiHUHrcpM— tunica •dTnillU>iLuiM«'«i«»>Mi'iv*. «!''■■ b? iSUiliiuif,)
There ia within n lining of fnHifono epithelioid oelts. very eimil»r to thnw
of a capillary and aimilarly ccmmted together into a membrane t^'>^- S-]-
lirac (liBiticicr of tbne fueiform cells, irhich «re Mmoiimee very narrow,
!■ pIacwT luirnlli-l in tlii! axis of tlio artny.
Oauitit' tbu> ipiilii-lHiiil liniiif; uoiaea * iliin trenitpBrent aM-ueturelew or
finely lihrillali'd mrmhmiie. Aeeii tii an itpticfti or other secti'm of lh« ftttm*
_ a* a mrrc liiir, wliirli ivrviw hi> a fiu|>|x>rlii)i; m«tu)>r«n«, baiaciDcnl inciubranc,
■ W mtTfirtna finrpria, for i1m> «pi(lt«lHii(l velllt. Tin* inl-iDtiniiw \* similar in
^U^niiii->l nnliin' nncl in pmixiriin In tittt olrurtii.' fibnu foiiiiil in con inactive
^flfair. aoil licniv U »|Kik<'n of an the tloMie memhrnnc. Tliv f [lithi-ttoid colli
^^Sd tW clultc fi>«ml>nin<- t<ip?lh*r arc often cfiukvn of m» forming tbo inntr
mtt (ttmiVn i'n(iui) uf thp artrrr.
IVi'ni)>|iril inmaverwly in n antn or Ipm clutinctlj wpirsl manner round
lUi iiibrr niut, and imbedded in n Kmiill c^uanlily ol eonnective tiwue. tie a
nitnber <if [ilaiii musetilnr fibre*. nrrnni.'''<l id the i>malle«l arteries in a einijle
Uftr, in the larger but itill »iiiiil1 iiitcrie* in mora than one layer, liiia
iwtna in tkese arteriea ihe mUlille or niiisctibir eoaL ( tiini'iM mMia). Outude
Am nuMuhir coat come* the external eoat {lunifu esiinui) conaiMing of con-
nctiva liHue ilie buudlee of which are disposed for the moat pari I'>n}{ituili-
DillT And contain a nitmlior uf eunne«tive-tinue corpiudea and u relatively
brp number of elsHtic librra. This outer cuat ia continuouH with the coa-
Mtivr-ltMue \yiA in which the artery lies.
A uioule sKery then dilfeni from a eapillary. In ihi? ihickncw of lUt walls
vkmhr (be pertneabilitv tu cbaraetvrjstie uf the cupillary i^ tt> u gn-at extent
hH. ia the itintiuet development of elulic elemeuti, tbc! Gla*l<e rocmbranr of
A«iutf coat, and the clastie librvn uf the outer eoat, vhervhy eliulic >juali-
ti« wt definitely ntureil tu the walk of the veinel. and laHlv and chiefly by
iWMtmiee >4 diriincl muscular eleroento. It ia obvioiix, that while by tho
<lnwfiMPl of ela)>ltc clenx-'ntii, pimve rlinngM of calibre have a grealer
mpttkan in the capillary, active ehan|.'c« in calibte, irhich in tbe capil*
jnrm al best doubtliil. arc ujsured to the artery by llie muscular elementiL
Hmi these trnnsvenbely digpnoed muscular fibres contract, they must narrow
ih calibre of ihc artery and may do that against even rery eonsiderable
tWraal prc»ure: when tliey relax, Ihey allow the internal preaaure which
■naiM. ut diHieiid ihe veuel and tenipomrily to increase the calibre.
nbeii Ruch i> Hmall anery break* up iniu capillanftt tlie niuiicular fibrca
uA flattie nicmbrarie dtwopear, the remnant uf the niiMculnr coal being
noMiBtit coiilinueil fi-r a shorl diittunce in thi- furm of a single libri' stnig>
(liwtD a i>]iiml fiu<hi"u muud the artery toward tlie capillary, and all that
altsii the epithelioid lining of tlic inner coat with a little oonDcctive tiMiie
■•nnment the outer coat.
i 108. Tha toryer artrritt rcMinble tbe minute arteries in to far that their
■dia maj h(> cons>(lere<l n* eompoeed of Ihrre voate, but rjich of \iuxe coats
iiif • miirv itT lew ounplvx nature, and the minor detail* of their struc-ture
ilh in dillrnrnt artorie*.
biMieh an artery as the carotid or radial, the three coats have the fblEow-
■jtmeral characters [Fig. -MiJ.
iW inner eoat is composed of a lining of epitbeJioid cells resting not on n
■a^ delicate basement membrane, but on an elastic layer of some thick-
>■. I iMWistilj||, diielly of a so-called " fenealrated " elantic membrane or of
M« ihao itne ttich ntembrane, together with some amount at fine elastic
laad in some enses at all events a small uuBDtily of white conneclive
L A " fen<-«trated " membrane is a membrane conipnewd of the Mine
am a* the elimtic fibres, perforated imeKidarly with bolea. and more nr
l^aarkvd with indicatioosol fibrH; it may be reyanled aa a felinork of
•latii: fibn*. fused or beaten out, as it were in a UMrv or K-m mmplete mem-
Wua. BDina of Uw meidm of llie feltwork remaining us ** fenestne " and
and ^A
180
THE TASOVi:.AR MB0HAN18H.
tncwof thellbna being xtill hfi. Suc-li ri-ncAlrkted nciiibnats, »ntnc thick,
•OIU4 thin, occur hut!) in tliv iinK-r niid riiiil<ll(> ooaCa of tho Uw^r url^rius ;
Dti'l ill dm iiini'T cant, ti»iinliy iinmctiiitolr iindpr th« e|>ilhvii<ml lining, ibei*
is in niu«t Inr^v tirtvri«« n vooNpicuoiis mcmhrnun of tliis kiiiJ, sometimes to
thick nit tn give n wry distinct iloubic ojlline in mentions of the arlcrr even
umlor nioilernlc powcre. nrn«filh this there tnny be other eimilsr fenestniied
membrnncs, or u felt of tinv cl«tic fibres held together hf a very siuall
({iiiintily of white coDOCctive tiwue. In iho norta, and in some other arteiiee,
tlie cpitholioii) mIU rat imraeilialQly not oa an elaatio membrane but i>q h
thin layer of eo-called " iiub-epiihelioid *' tteauc, which cun^isla of oi>iiii«ctivc-
licBiie cor|>u8f!e§ imbedded in a homogeneous or very faintly 6brilla(ed
matrix or };round eub^lam.'e.
The epUbelioid <.-ella are diiposed loni;itudiiiallr. that is, with ttieir loi
diiiiii^iers parallel to the Kxis uf the artery, and a similar hmgiludia
armtigi-ment obtains to a greater or \tss cjttent iu the undprlyiiic elnitlir ele-
mentA. When utler death the arteries, emptied of bloml. bei-ome narrowed
or cotutric'ted by the r-oii tract ion of the muKCiilar element of the middio
colli, lh« inner coat is throwu into longitudinal wrinkle* or fotdi. M UiKl io
tniiiitvi-rr>u necliuiiM of an artery in thin condition the inner (wal hai • dianwy
t«riiitic ptiekereil appejirance.
The inner cout in .lomewhat delicate, and eunily torn, »t that in injiirin to
arlcriiv, an ndieu nn artery in forcibly ligatured, it in ant to be brokeJi.
The middle ctmt, which in gcnernlly ninny time* tnick«r than the inner
coat, <im»ixl» of clnNtic Inyere an<l mn»cruiar htyer* plnircd in more or lem
regular alternation. The muscular biyert L-nnsist of band* of plain muscular
libn» placed tniiisvenely and united together by a very itmiill amount of
white connective tiseuc. The ela«tic layer* eonsiot of somcwhnl (bick fenes-
trated membranes or of feltworks of elastic fibres ninning on the whole
loDfpludinally, but not uufiequenlly more or leas ublii^uely; these are aUo
bound tof^ther by a aiuall quaulity of while coniiectire tiineue.
The ou[«r coat eonaista of feltworks of eltietic fibres, or in fome iostancee
of ftDeelraied inembrauee, diftpueed chiefly lougitiidiDally, and aeparated by
buodlee of ordinary white connective tUieue, which become more and more
Itredominant in the outer portions of the coat. In many arteriea bund* of
j>)uin mn!>eulHr fihrr» ore pre!<ent in this coat also, and then run for the moat
\»n but not excliixively in a longitiiiliual ilireetiuu.
Bloodvci«iel» for the noun.ihmeiit of the tiiniie of the walls (mm wuorun)
are procrnt in the larger arterieii, beiug iu<i»t abundant in tlie outer cont, but
penotraling for »ome dint luice into the middle coat. Nervex, eonaia^ngcbioBy
of noD-iDcdullat<t<l fibre*, may he iraired through ihe outer coat into the
middle coat wberv they apptrar to end in connection wtib the muscular libree.
Lastiv, in iJie case <if mwt larger nrteritv, the bed of eonncotivc tinsue lO
which the artery run*, t; fomicil into n more or lew distinct shnith. In this
•bealh, tlic white connective tiiwic is much more abundant than are tbe
rellow elastic elements, so thnt the sh<-iith i' far lens clastic than the artery.
Hence, when an artery and it' sheath are cooi|dctely cut acroe*, (hb artery
is, bv elastic shrioking. retracted within its sheath.
Tlie nut important structural features of a largo artery may then be
Bummed up, by saying that the artery conMsle of a thin inuer cuat onosisling
of an eiiitheliind lining reeling on an elastic basia of no ooutipicuoiu thick-
nan, of a thick mi<ldji'! coat consisting partly of muscular fihrw disposed for
^e moat part tranHvcrxely, and partly of stout elaatie elemeutA, tliia coat
beEog the tliiektat and must im|>oriant of all three coats, and of an outer
coat of variable Ihieknec^ cou^intinK chiL.'fly of elastic eleueota ioiermixod
with an increasing aiuouiit uf white connective tissue.
rSATCRSS OF VASCULAR APPARATUS.
181
k
I
rteriM powow the uhore fealum.- It n>aj furthpr he «ai(i, that as n
nile lli« Minfctilnr ctemeni bean a lurger [Toiwrtion lu the clastic
io the (lualler tliao in the larger arterif«, that is to eav. (he itiiialUr
Otarin kn more cboapicuoii^lv museular. aod the lar^r arteries more coii-
il^ODCNMljr elastic It rauat W n?iiieiiihered, huwrever. that the several arteries
vf IImj boily difler coiisidenihly in minor ledlures. siich as llie relative die-
poattiun ftnd UDount of tuuscttlar aud eluiic eieiuenu in the middle coat,
tlM amount of miwcular ibsue in the outer coal, and Die proportion of white
ovavctivo Umup prcxent and the like; iu llie aorta Tor imtanee, a vonMder<
able quantUjr of while n>nti*«iivo tiMiu« U pre*eiii in (be middle and, indeed,
b ill* laner coal, a» well tu i» the outer coat. I-eavJog the*e (unaller dilfer-
neca on one aide wo may tmj. that while all thnM^ couih, but cw])edBlly Um
teporUDl middI<^ coat, ooDtribute to kIvc- nii nrtcry it» chflmclcrintic elaHtio
■Mlkiw. by virtiH- nf which it expands remlily under iiilcriiiil |>r(«.-<iire, and
■riab again wbvo the pm»tiro w rvmoved, it i* (h« niiddk coiK which 1>y
■laMef lira abtiiidnnt circularly dispoced inu^ciilur tihriv, now through tliv
(■inrtioii »{ thow tihres narrows and conslricti!. now through their rclaxa-
Am famiilfl thf widening of the vessel. The iinporlaDce ut ihe inner cunt
a fmbably cenlrMl in the epithelioid lining ; in ircatine of blood (^ 'i'i) we
«« RMon to think that the bloodvesets exerted a mariied, though obscure
iilMMe oo tfae blood BtreaniinK lhrotif;h them ; that inlluence in all proba-
tA; IB eflteied by ihe e[Hthcli<)id nelk The elastic elements of the inner
(Ml are pnbably chiefly of value in jwnnittiuK this coat to follow the
iImm of the nxkre important middle coal. The outer coat while increasing
lb* wutic power of Ihe whole vewel, in especially useful, by means of iu
nail blond vesae Is. in (-onveying nourisbmeut to the middle cuut.
illO. The m'iM. Thc)« vary in diflVrent purm of the itody to very
■iJily, that it is difficult to eive a gcneml dncription »f strurlurc suitable
bxDvnnsi It mav Ih- ui«l, huwevvr, ihal ihry dilTer from arteries in having
Wtth tfainnrr walls, and in ihusc walls containing rclnlively much more
■Ut* otiDDeclive litduc and much lew yellow elastic tissue.
A large vein [kdvcsbcs like an arlcry an inner coat consisting of an epitho-
M lining, the celts of which are eliorter and broader than iu the cornv
riding artery, resting on an elastic basis, which is leu conspicuous than in
EMTMpoodiiig artery, and conaiMB of a tine feltwork of tihrw rather than
afaulrated menibraue, and ci>niain« mure white connective lifsue.
laa medium med vein auch as the sikphena vein it is porsihle Io disljn-
|iUi outside the inner coal, a middle and an outer coaL The former
eaMUof white conueetive tiMiue. with a scanty supply of elaalio Sbrea; il
Hnuias, aometimea iu couMdeiable ijunntiiy, plain rotiscular 6hres, the
kmdka nf which form a meabwurk, with the m«»hep disposed for the iDnu
fan trannertely. The latter consists also of white coiiucvtivc ti«*ue with
■W elastic fibrta running lnOfritudinally and <)hli<|Uely, plain Ku»c»lar
flms being sonietiniM |irtM-»l and wlien prevent diajHMNHl chKfly in a longi-
laiim,\ direction. Small vasii vusonim arc pro«util in the outer ouat and
Ottnd into the middle eOBL In many large vein* lliero in no sharp diMinc-
lisB betwcvn a middle and ouir'r i-oai ; the uhnlc wrapping round the inner
Mt cennile of white connivlivc with n vniiublc (|uautily of clastic liHUc,
ud i^f muscular fihre* which luu chiedy longitudiunlly or obli>|ucly and
•kick may be very scanty, or vhicb as in the vena portie may be abuitdant.
TW Mructure of the v«'ins iu fact varies very widely; on the wlmle tl^y
Hy be said to be cliennels. the walls of which are elastic enough to adapt
iMMlrcs to considerable variations in tlie ()uaiitiiy of blood passbg ihrtiugh
Iha, wilbout poasewiDg, as do the arteries, a great store of elaMk ]H>uer to
■*■! fTtat variations in prcuure. and which are not so uniformly muscular
182
THK VAHCULAK MKCHANISM,
and ooiitractil« ax nre tlie iirteri(!!'. - And we ahull see Utnt this [:«iierul char*
aoter uf paaaive channels is adapted tu the work which iht.- vuiti« Imve (o do.
Tht* (:«Dcral chai'acl«r, however, U modified io certain situntiuii* (o m«et
purtieiiitir wanls ; tbu!> while the vein4 of the bones and of ihi: hriiiu are
devtiM of mtiituuiar fibres, uthera such m tlie v<-rin |>i)rl:i' mny be very niuv
I'ular; and in .■niriie veioH such iio llia-x; iif th« cxtri'tiiitic^ ii c'l.invtdcrahk
<(ijHiilitv tif eliL->ti<' liHiio io ]ire>wDt.
A minutK vfin jiint einergiii){ I'runi i-a^iitlikrics dill'ors very litllv from an
artery of i.-iirfV9piiiiding siw; it is of riiiJivr wider bora, hiu ikcidvdly I(m
muMciiIar and vluvtic (i#iio, and ihn c pillmlioid rails arc shorter and broader.
Many veins, eciiennlly thosv of (he limbs, are providi;!) witli valves [Fin.
&i ami 'io\. which arc pouch-like folds of the inner coni, the muulh of we
pouch looking away from the uipillnries toward the heart. The wall of each
valve cunKii^ts ofa fining of cpilhi-lioid colls uii the inside and on the outside,
and bcCweoD the two a Inycr of while connective tissue sirenf^chened with a
few elastic libres and somewhat thicker than the connective-tissue baeis of
the epithelioid lining of lUe veins geDerally. The valves may occur sin^^l/
or may lie iw» or even three abreast. The veins of the viscera, those oftba
cenlml nervous system and lU membraiiM, aad of the boDed, do not poMcaa
valves.
TimvtTii VjLTisorRK- Alter Dalton-I
lattlac lilt t9y laWnl rbaanel.
i)it>wm of Mend
AMr Dil.TOW.]
SUl. The details of the Alructni-i- <if the fiifi^uliar muscular tis«i« forming
tbe grcaU'r part of the heart we nhall n^ierve to a later seotjon : but we may
here sar that the iiilerior of thf heart ia lined with a membrane (frido-
Mftiiwwi) ciirrCBpiindinK to the innur i-oat of the bloodvessels, and consistioK
of u lavcr i)f epithelioid cells, which, however, are shorter and broader than
in (he Moodvntwls. being polygonal rather than fuHifurm, resting on a con-
nective tissue basin in wbiob are present elastic fibres and in places plain
muscular fibres.
The valves of the heart, like thow- of the vein^-. arc liilds of (his lining
roCtnbraae. sirenglhcncil by a cmatderablc development or(<onnectivc tisane.
In (he middle of tbe ihin frw border of each of the srmilunar valves of (he
aorUi and pulmonsn- iirtcry biiiidlefl of this connective liKiiie, meeting
(ogMher, are mixed with i-artili^ cells to form a small nodule of flbro-
carlilaice called the rorfmi ar'intii.
In the auriculovenlricolar valve* muiiL-ular Bbres pass in among the ooo-
neelive ttwue fir sonw liulc distance from the attached border.
In one mipect the endocardium dijfont from the inner coat of tbe Idood-
reseebt; the conneeiive (issue in it licar* bloodvea^ela and lytu{diatira. In
rtATintss or vascular APi-AKATrs. 183
tbfOM nf ihci auriculo- ventricular t»1vm littat lilondvcBeb of ibo tad<h
ariiun irAVnno the whole ralve, but in Uiq caae of Utc setnlluiiiir valvw
•up thort w-*r lh<' Atutdwd bonier, m that tlic greater purt ii)' tbc vuItv ii
MwdtMk
J/inn t'raturf* ef the Apparalut,
ills. W« IDB7 Di>« raw* briefly in r<>vi»w mime nf the mnin fMtunw of
lU Mvuttl pans of toe vn»niliir appiimlui, heart, nrtcriM. veinc, aad
ounUarin.
The heart ta n niUHiilnr {>iim|>— tbat is, a pump th« forcv of whnw Urokw
pMpplinl by the iTininu'tion of muKtilar fibres nnrltin]; inlennitleolly, the
■trans bchi): repvalcit n> many tim«s (in iiiaii about T'i timea) a mioute-
[1 ii an ron^ruclr*! ami funitshed wilh valvea in such a way that al each
itn>ki il ilrives a ceriaiu iiuanlity uf bloMi with a certain force and a certain
a|>idity frwu the left wniriclp inro the aorta and so ini<> ilie arlerio*, receiv-
b|;durio|; the stroke and the interval lielwi-cn that dtnike and the next the
anrquHUtily of hlouil t'mni ihe veius into ihe ri([tit auricle. We omit for
ani'licily'!) take the |H)lm<>tinry circ^uluiton by wliii^h the aarae mianlity of
Um k driven at the llI^tkc fnim thu ri^ilit vciUnde into the lunpt and
itciivtd into the Ittl nuHclc. Tlit.- rliyihra of the lieiit. thai U the fre^iiiency
of NpMilku) of the alr»ke). aod tin.- cliaracter9< t>T each heal or «trok«, are
drinmiaed by chanxv* lakinj: pinoc in the tuMum nf the heart itMlf, thouf^b
tlvi air aW tiitlurnof^l by cauMs uorkiiit: from witboitL
Vtf •rli-riv* tin tulws, with rctattvelv :<toul uhIIk. bninching from the
Mfta all ovfrr the body. The (tonnliiulion of tbi-ir udIIh. ax n'C have rttfo.
•pdatly of the rDiddleGnul,givi,w the ancric* two client propi^rliea. In
ibtlini plaor they an- tfry r/'utie, in thi- iK-nw thitl ihcy will mrvlcb rrwlily,
koih IfDj^hwaVi and eroaswtae, when pulled, niKt return nuulily to their
fcmvr aize and abape vhen ibe (hiU is taken ofl". If fluid he driven into one
foJ uf a piece of artery, the other end of which is lied, the arierr will swell
oai til B very |;reet extenl, hut returo immediately to il» former calibre when
ihtlaid ia let out. 'I1ib elufiiciiy is. m v« have aeen, chiefly due to ilie
flMiie eJrmeuta in the ooatit. elniuic Diembranea and leltworks, but the miu-
nilnibns, being ibenuelvee abn elastic, conlrthule to the result. By reuou
rf Iblr MMOiIiig Hich stout, elastic walls ibi? nrteriw when empty do not
MiDipw, iKit ntmnin iw o|>eo tulien. In the second place tbe nrli««. by virtue
ifllitir mutoiilar elemeata, are ranlrarii/'- : when itimulaieil either diTts:llv,
tthfmfiviag an elvctrio or ntcchtmical •timuluit lu the arterial walla, or
Wn(Uy,i)y luoans of the Mvcallml vaxo-niotor nenm, which we A»\i have
la Nady prMvnlly, the urli-rii-a shrink in calibm, Uie i-ircnlarly dispowd
"awllii' fibtta contract iuL'. iiml Mi, in proixirti'in t4> the amount of tbcir
iTClractMD, Darroaiii); ihnlumcn or bori' of the vctvel. The i'»iiLmi'tiou of
■Wr arterial louacular tibn'o, like that of ult plain, iion-nlriuUil muM'iitar
IIW. b eluw and l<mj;-<>->niinuvd. with a long Ut^'nt jM^od. ii^ com|iar«>l
«uk the conlmction of Hkelrtnl Mrialeil mu-HMilnr AbrvM. Ovrin); ta thig
■■nfair eleim-nt in the arierini wall«. the calibre of an artery may he very
tun* or very wide, or in an intermediate condition hdmwo the two,
adlbvr very nanuw nor very wide, according aa the muscular libru* are verv
wuit niniracted or not conlrncted at all. or only moderately eonlrac*4Ml.
V| have furllier aeen thai, while thi- relative projtortion ol eloMic awl
Maoplar elvmenU diHers in <lil1ereDl arteries, as a general rule the clMli«
iImhoU predominate in (he I arcer arteries and the tnuaoular elentents In
(btmaller ■itvrii-i.x tliiU ihc I ■r>,'er arteries mav be spoken of a« eniittcntly
abitic, or a* r«{><-<-ially uveful im accKWint of tbeir elastic proMrlic:>. and tba
Mnaltrr artrriw n» eminently muscular, nr lu etpcciallj useful on occiiunt of
thnr iBuw;ular pto|>t'rti<«. Tbui, in the minute arteries which are just ^uaa-
184
THE VASrCLAR MECUAXISM.
iog into C^IUrin the niiiKiilnr i-iml, though ctiiii]inff() nfton of n single
layer, and thst •oinclimM nn imiH'rii^ot imc, of miiiiciiliir Itbrat, in ii miicti
Diore (^ODHpictiftiiB anil imptirtjint (mrt of thcnrlcrial wnlllhaii tlmt furniKhcd
by th« otnxtic <■!(.■ iiK-iiu.
The arttirw* brnmhing out fn>m n nitiglc Hurln down m niullitiidinou*
capillarir* in murty i-vi-ry pnri of the body diminish in htire m ll»cy ilivi<)c.
Whera un nrlvry dividt^ into two or gives olT n branch, though the bore of
ODch division i* len thuii thiil of the am-ry Ixtfore the division or brandiiog.
the two together nr(> grcntcr: th»t is to say, the united wclioDal area of tbe
branehes is greater Uian the wctional area of the trunk. Henoe. tbe aec-
lional area of the arterial bed through uhicb tbe blood Aova goa on iocrcM-
iug frDEu tbenorla tolhecapillariee. If all the artorinl braochcv were throwa
together into one channel, this would lorm a hollow cone with iu apex at
the itoria and its base at the capillaries. The united sectional area of tlte
capillnriea may be taken as several hundred times that of the tectimial area
of tbe aorta, ho greatly duen tht.- arterial bed widen out.
Tbe capillariee arv^ channels i>f variubk' but e:(«'ediiigly vinali utA. Tlw
thin !>heet of cemented epithelioid |)hiti.i> which fonn.n the only irnll of a
cnpillani' is elaitic. permitting thi^ rhannel ofli'nil by thr Miino capillary to
dilfcr much id width at iliflt-ri-nt tiincN to widen whun hluiMl and blood*
corpu^tclcn ar«! being pri-NX-d through it. and to narrow again wben the
prr«8un? is loncned or cut olf. I'hc Minic thin sheet porniitt wntor and sub-
sta»i;ps, including canes, in solution to pics through il«elf from the blt>od Lo
the ti«ue outside the cnpillnry, nnd from tho tissue to the bliiod. and thu§
carries on the interchange of malcrial between the blood and the tisautt. In
certain cireumsiance*. nl all events, white nnd even red corpuscles may also
pass thn>ugh the wall lo the lis.sne outside.
The minute arlerics and veins with which the capillaries are coulinuous
allow of a similar interchange of lualerial. the more ou the smaller they are.
The wuIIh of tbe vein* are thinner, iveuker, aud 1cm elastic than thoac of
the arteries, and |iOH«e.-4t a very variahle aiunuiit of niuacntar tiwuc; ihcy
oollap«(.- when the vetus am i-m|>ty. Though all veiu.H are more or lent elaMic
uhI lonie veins ar«> ilistinrily niuncutar, thu veinii as a whole cannot, like tint
artericVi be charactcrixed «« eminenllv elH.'>tic and conlnictile tul>v*; ihcy
■re nlJier to Iw rcgunlcil ii;< nimplc cfiuuni'li> for couvcying lh« hkmd from
th« capillaries to tbe heart, having just sit much ehutieily an will enable
tfaem to aeconimoilatc ihenixelvi-ti to the ijnnnlity of blond piuning through
Ihem, the same vein l>cing nt one lime full and distended and nl another
time empty and shrunk, and only giflol with any i;reat amount of muscalar
oontractility in s]>eciul cases for »]>eeiat reasons. The united sectional ari»
of the veins, like thai of the arteries, diminishes from iho capillaries to the
beart ; but the united sectional area of the venncavie at their junclton with
the right auricle ia greater than, nearly twice as great as, that of tlie aorta
at its origin. The total capacity alao of the veins is much greater tlian iJiat
of the artcrim. Tbe veim aluue can hold the total masa of blood which iu
life is distributed ovtr bolh nrterim and veins. Indeed, nearly the whole
blood is cnjiahle of being received by what is merely a pirt of^ tbe venous
iystem, vix., tbe vena porUe and iijt branches.
The Mais Facts op thk Circulation.
$ 113. BcJbre we attempt lo study in detail tbe Rorkitig of iheoe aeverm)
pans of tbe mccluinism, it will bo well, even nt the ri^k of »ome Aiture repi-
tttion, to take a very bri«f rarvi^ of tone of the nlivni pointit.
TUX MAIN rACTB or TIIK i^ BCULATIOX.
165
Al tacb (mU i>f tho heart, which in miin ia ivi>piUi<<I nimut I'i liroee a
jtoatr. ibr cotiiniolion or ctbIoIc of lh« veniricteii drives a cerlaio ijiiuntUy
I'STblml. iMxilwblj' Binoiiniing to abmit 180 c.c. (4 lo (I ok.), wiib very ifreal
1m iatn the aurta (awl ihe aaioe <iuanlit_v of blond willi l«w force into the
pliniiaarjr an«ry). Tli* diKharge of blond f^om ibe ventrirfe into ihe
HTla » trry rainil. and tlie time taiteD u|i by it a, a>) we sbatl itee, niui-h lea*
ihiB ibr iiin« wliii-b iiit«rTen» between it and the next diMrhar^ of the next
hat- 8n ifaat tlio Aov from iht< hesrt into lh« arteries 'u iwm dialincUy
iMtfvkleui, •udilcrii r»{iii) discburjices alt^niating with relatively b>iig iiiler-
nlidurinK wbtch lh<< iirU-riai receive do bUxul rnim thf heiirt.
AlMch brai of tbv heart jiirt a» uiiii'h bloml fluux, un we ibnll aeo, from
ib( mill inin ilkc right aiindr an OMuipc* from lliv left ventricle into Ihv
tMta; hat, aa wc «hiU iiUo ree, thi« inHow ia mudi tlower, lukoa a longer
tee, ihan the dbcbarg* (ram th« WDtricle.
Wbro the finger U plncT<t on an nrten- in the living body a sense of rtsist*
iOc» i* felt, and ihi* rv«i!>iiincc tevm* to be incrcnsed at intcn.'als, conwpond-
ilf la the hMirt-bcata, ihv arlfr>' nt eueli hcnrt-Ix-nt being felt lo rise up or
aatnil under the fingor, coiutituling what ne shall HtU'Iy hercuilWr aa the
fit In cprtntn tirU'ri** this pol»e may be eG«n by the eye. When the
fapr ia nniilsrly plaivd oti n cur ree ponding vein rery tittio reaislanoe is felt,
mA.muU'r ordinary circunlslaDOe^ no pube can be perceived by the touch
•r bt Ihe ^e.
nlwa an artery is »evered, the How of blood from the proximal cut enil,
itaM Uw heart side, ia not ciuable, but comes in jets. correepondinK to the
iMK-beala. though the tl»w duw n<'t cenM" beiweeti ihe jela. 'llie blond U
•MMJ mth conmderable furoe, ami may in a lurj|:<^ arlery uf a Inr^e animal
m^mneA nut lo the di«lanc« of some fecL The lurjicer Ihe artery and the
■mr W tbf heart, the greater the furn: with which the blood ijttiics, aiul
ik aon narked the iiilerRiillenceof tb« flow. The flow from thp dirtnl cut
Miihat awav frtiiu the heart, may be rcry flight, or may inko place with
■Milrnihle turn and marked iiiKrmiltcnce, aeconling to the amount of
alhlwal coRiniu»i(-alion.
Wbto a CO rrcfpi Hiding vein i« aevcrod, tho flow of blood, which la chiefly
6m tlM ilifllal cut i-nd. that in connection with the capillarim. is not jerked
bn raDtinuous : the bloiMl coniea out with companilively little foree, and
'•tU* up" rather thnn " »puTts oiil." The flow frwii the proximal cut end,
ikd ta the heart Mde, may amouut to nolhtni; at all. or may t>e Hli|;ht. or
t b* eDBfltderBliIe, depending on the preeeooe or abociice of valves and the
Laf collateral couimunii-atiou-
I an orlery is liuauire)! the veswl awella on the proximal side, lowanl
Ibbovt. and the thrubbiu^- of the pulte may lie felt right up to the li^-M>
m. On the diMal tide the veiu^l is empty and Hhrtiuk, and no pulae can
hUt in it ohImb tlwre be free cullaleral communication.
Wbm a rein la liffatunil the veMvl iiwellii i>n the diilul i<i<[o, away from
tb heart, but no puuc is fell ; while ou Ihe proximal t>i<ie. toward the lieort,
liirmptv and collaptfl unloM there h« liro free collateral riimmunicatioO.
1 114. When the interior of an artery — fur inatancc, tho carotid — is
in oomrauntcaliou with a long glaitt tnbe uf not Ina groat a Imre, bold
Jly, tl>o blo«<]. immediately u|)on the oommu mention being elfeclvd,
She Mm to rt)»b into and to fill the tulie for a certain diflance, forming
a mlumn of hltxHl of a certain heighL The column risca not sleiwlily,
^ by Im|», each Inp corn* |>»n< ling to a heart-ttcnt, and each leap being
tlhoii ita prtsir«e»or: antl thta gooi* on, the incn-nso in the height of the
Da at rui-h heart-beat each time diminiahing. until nt laitt the oulumn
nhiDi)
Itftl TUli: VASCULAR MKCHAKISH.
ceases tu rise aud remains fur a while at a mean level, above and below which
it uscillatea with slight excursions at each heait-beat.
To introduce such a tube an artery— say the carotid of a rabbit-^is Iftid bare,
ligatured at a convenient spot. /' Fig. 6ti, and further temporarily closed a little
distance lower down nearer the heart by a small pair of " bull dog" forceps, bd,
or by a ligature which can be easily slipped. A longitudinal incision is now made
in the artery between the forceps. M, and the ligature /' (only the drop or two of
blood which happens to remain inclosed between the two being lost) ; the end of
the tube, represented by c in the figure, is introduced into the artery and secnred
by the ligature I. The interior of the tube is now in free communication with lie
interior of the artery, but the latter ia by means of the forceps at present shut off
from the heart. On removing the forceps a direct communication is at onceeitab-
lished between the lube and the artery below ; in consequence the blood from the
heart flows through [he artery into the tube.
This experiment shows that the blood as it is flowing into the carotid ii
exerting a cuDsideritble pres.''ure on the walls of the artery. At the moment
when the forceps are removed there is nothing but the ordinary pressure of
the atmosphere to counterba lance this pressure within the artery, and con-
sequently a quantity of blood is pressed out into the tube ; and this goes on
until the column of blood in the tube reache:^ such a height that its weight ii
equal to the pressure within the artery, whereupon no more blood escapes.
The whole column continues [o he raised a little at each heart-beat, butsinks
a.'' much during the interval between each two beats, and thus oscillates, as
we have said, above and below a mean level. In a rabbit this column nf
blood will generally have the height of about !)0 cm. (■! feet) ; that is to say.
the pressure which the blood exerts on the walls of the carotid of a rabbit is
equal to the pressure exerted by a column of rabbit's blood 90 em, high.
This is equal to the pressure of a column of water about 95 cm. high, and Kv
the pressure of a column of mercury about 70 mm. high.
If a like tuite be siniilarly introduced into a corresponding vein — say the
Jugular vein — it will be found that the column of blood, similarly formed in
the tube, will be a very low one, not more than a very few centimetres high.anA
that while the level of the rolunm may vary a good deal, owing, aa we shalL
»:c later, to the influence of the respiratory movements, there will not, as iw-
the artery, be oscillations corresponding to the heart-beats.
We learn, then, from this simple esiwriment, that in the carotid of tb^
rabbit the blood while it flows through that vessel is exerting a considerable
mean pressure on the arterial \v;al1^, equivalent to that of a column of mer^
cury about Til mm. high, but that in the jugular vein the blood exerts on
the venous wnlls a very slight mean pressure, equivalent to that of a column
of mercury 'i or 4 mm. hifrii. We speak of this mean pressure exerted by
the blood on the wiills of the bloi>dveBsels as blood-prensure, and we say that
the bliM)d- pressure in the carotid of the rabbit is very high (70 mm. Hg.),
while thai in the jugular vein is very low (only 3 or 4 mm. Hg.).
It) the nonuiil state of things the blood flows through the carotid to the
arterial branches beyond, and through the jugular vein toward the heart;
the pressure excrlcfl by the blood on the artery or on the vein is a latent
pressure on the walls of the artery and vein, respectively. In the above ex-
])eriment the pressure measured is not exactly tbis, but the pressure exerted
at the end of the artery (or of the vein) where the tube is attached. We
might directly measure the lateral pressure in the carotid by somewhat
moaifving the procedure described above. We might c<mnect the carotid
with a tube the end of which wa-s not straight, but made in the form of a
T-piece, and might introiiiice the T-piece in such a way that the blood
should flow along one liuib (the vertical limb) of the T-piece from the
ArroMM NIK iHVBRmiTUHi UiiMit'nt
M •■ ■)(•* fteM-lwDd rmvt l« wdo, ob ■■■ tBluvBd mta. Ibc afoUil uim. *l*ai|iwt bv Ite
M. arUkiteMcauMTMljIaaliir lUiMr, T»a •rivnr tiw b(i>ii t1palun<1 •! T •■■d IW |(UM
r kM ■■■■• IMta)iii>*>l Iniii ihrwurir IkImmii ilir lltnilniTraii'l ilx lVi*n|aM. uHlaHwrtil
^ ' •- t>.l1lBca aiiimlail •uluUiMi cf laUuin i«rUiBaw nr • oMUiai oT
ma^ >( t|i, sr. lira, u»1 (■■BbtoiiT Ivlna ralaw) ur luovrrd at pliBUan*.
naalalMu ikn> Itt iIib ii»tK p.' ncuUUd k; Ika iduup C" IMO Ite lub* l, A iiiliicc. ■Hh ■ mofr
188
THB VKSOUhAH MErHANISK.
proximnl to the dUlul part of the cAixiiid, and at tbe »in« time by tb« oUwr
(bonzoutal) limb nt' the T-piece iiilti th« luaiii upHj^bt part of itio gla«
lul>e. Tlic tubulin ii)' l>Iti(icl ill tbu tube would then \>f a an-asarv t>f lh»
pressure which thi* bhioil aa it iH llowing along thu carotid it ^-sorting on ■
portion of iu wuIIh correvpondiiig to the mouth oftbL- hnrizontal limb of the
T-piece. If we wt'te to inlroduw inio tlie Hurts, Kt the pinco of origin of the
carotid, a •imiUr (larger) T-pirn:, aiid to connect the glum lube with the
borizdiilal limb of the T-pivn by » piece of cImiSc tubing of the nmc length
and borv ok the carotid, llio column of blood rinng tip in the tube would be I
the memurc of the lalenil preesnre exerted by tbe blood on the walls of the
torta at the origin nf the carotid artery and ttnnBinitted to the rigid glafv
tube through n certain length of elastic tubing. And. indeed, what i^H
tnciieiireil in the experiment previously described i» not the lateral preaauJ^H
in ibe cjirolid itself at tbe spot where the glass tube in introduced, but this '
lateral pret»ure of the aorta at tbe origin of the carotid modified bv tlM
influences exerted by the length of the carotid between its origin ami Ibe
apot where the tube is introduced.
§ 115. Such an cxixrinient as the one deei-ribed b»A ihe diaadvantagea
tliat tbi.> animal is tviitkt-ne'l by the loait of the bl»oiI which goe« to form the
column in the lube, and ihut the hloiHl in ihi- tulK- noon clotv. and w> bring*
the exjieriment to an cod. Blood-pretciure may be mure conveniently aiudicd
by connecting the interior of the artery (or vein I with a mercury gauge or
DUDonietcr (Pig- 06) the proximal ileiicendiiig limb of which, m, ik tilled
above the mercury with fonic iniioeuoun fluid, ba in alto the tube connecting
tli« manonielcr with the artery. Using such nn iniilrunicni, we should
observe very much the Mime fnct* na in the more simple experiment,
Imniediniely that communicnti'in i» «*tnbli!«hed between tbe interior of
the artery and the manometer, blood ruithi')> from- the former intu the Intler.
driving some of the meri:ury from the descending limb, m, into the n^icending
limb, wi', and thus cauuiug the level of the mercury in the aacetidiug limb to
rise rapidly. This ri«! is marked by jerks corresponding with the heart-
beats. Having reaeheil a certain level, the nmrcury ceases to rise anr more.
It doea not. however, rvmiiin ubfululely at redt, but undergoee oscilfatious;
it kee|i.H risring and falling. Eiich nse, wbicli U very slight compared with
the total height t<i which the mercury has risen, ha.H the i&me rhytbin aa ibe
Ryiitole of the ventricle, similarly, each fall cornatjionclii with the diastole.
If a float, swimming on the lop of thi! mercury in the aiicen<Iing limb of
the manometrr, nnil bearing a brush or other marker, be brought to hfnx on
a travelling surface, some ■'ucb tracing as that rcpreaented in Fig. 57 will be
dcscribcil. t^vch of the smaller curvi» (p/)i corrc*|ionds to a heart-bent,
the rise corresponding to ihc systole and the fall to liie diastole of the veu-
eocli, BBT te iub«Uiutv4 Silt llio IwHIr unil ■lUi'lin] si r". Tlll^ iti<1>h>1, I> in uuuijr nvimil* ■
more tadTCDlcDi plan- Tbe mi-- ' lonnri.i'iwi uiiii ihc Iim-Ich mtit 1, iiml iw itopaDek e aUli ilw
BunoiutUir.or U'bleb « lathi' I <. :iiilI>, bqiI tifac aiipjiart Tbv nwr-
mrrlnlliiSMaiiillDB lliDb U'. >. :i,>^< ii>> 11 > < nxl ailaclinl in which 1* DUad
wUbUieiicap. wraiSEoa ihc iuuMIbk •urtaucr. Tlic «I>in|.oL at the ciul al tbt tubal bM «n
enaiifanwtn ahoua oti » lar^nr waloul the rliilii hand miliar cunier.
n«4anMvlla|luto not (ho inauciindat Hill] ihg mbalboing nmuplalal}' Hlled tiimit It* irbela
lanfUl Wllb fluid to ibe exoliulmi of all air, thn mnuta « la ail«d wlih ftuld, (llppeil lula Ibo opoa
mi ot Iha thick -vallfil Iiiitia-njIiUt lube i. unUl It mtou tbe tubs ( (wboaa poritko wnblu tlia
iDdla-nblsr (iilw U ikown hr the itiillnl llnnV and 1* Ibaii •viTimlj' Dxsd In Itila p«alllni by iIm
Tbt •idpi'Mkir t aud f* an anr npcoM. aad the proauni bniilc talHil or (luld drfTOD Is bjr U»
tritniir until Ihe mcreaiy tn lb« uunoncMir t> ralMd in lh« iVi]iilmI belglit The rlamp r" Ii then
Blnatd aiut tin loreeia M nuioml Irani the adefT- T^ pmaan: ur Ebc XiioeA in Ibe earottd m U
' la eonsqiMne* Iinuijbi lu I've lhn-<U(h ( u[idii tb* nieiriir) lii llie maoomvter,
^
THK MAIS rACTS Of THE CIBCOLATIO JT.
189
trid*. Tlitf' laripir iiodulttions (r r) in the tracing, irhioh are retipirntor;
In origia. will M dbciiastd h«n>«ller. In Fig. oH are given tiro tnciu^pR
1^ X,
nt^Hlto ovm* )• V arv U>r rolawiiTC*. Tti*i|«M IKnn r (n r vmbmi'ca n rcrfdntiay and*-
M« 1Wln<ilai9>ukiiqnvB»dcc.B<>4U>B lrn«uliulll«TWIiialiittucUKiwrn>|iM(illr M«l
nha fromltbe rarotid of a rabbit ; in ili« lower curve the reci>nliiig mirfiire
iiinndlia^ inon; rapidlv lliuii iii tlie umier rurve: utbemi«e iliv curves iire
ittaMd repeat the Jteneral f«tur«t of Ui« curve from tlie dog.
no. M.
'*.J\y\^/Kj\J\^
liiflnnrKBmiac Cnrai meat rai C^ion* or B>MfT. nix lau "*■"*
in uo HUM Uittom Bbcdsm.
fl—yltw ^ rxfirriHtrnt. In a CHrotiil. or other bloodvMM], |iroparti] at ex*
(UmJ, ■ Rdnall RIM* lube, of »utUbU- liori'. called a amata a introduced by Ibe
Brihod tleaoitied atiove. Aiid ia suliw<]UFDtly ouniiecled, by eacaos of a aliorl
rwoaf India rubber tubing { Figi. M >), aikI s Iciulfn or other lube < which U al
«n^lrlllil« and yet ool exiciuihic. willi lhi> i)nii-en<lins Itnih. m. of ihc nuuin-
«M<crot mtrcury cauiir. The cnnulit, (ubr. anil <ltiici.-ii<liii); limb of the nianu*
twan all filled with toine fluid, which tenil* lu {irrtent clotting of the hlood,
innc chnaen bpinK generally a atroiig tolutloii («[». ^r, |i>S3) of aoiliiim bicar-
•, Uii other Hums oiaj ha cbo««n. In order to avoid Iom of blooil. a quao-
' flulil i* iiijrtMd into ibe flexible tube auffieient lo nuBO tbe morcory in the
( limb of the manometer to a 1»vf1 a rery little Iwlow irhni niny be
■d gn«Mtd at iw the irnibabic meiiii [ircMure. When llie (oref|M M are
h\, the prMHire of Ibe bl'iod in tbe carotid in irannniiiied Ihrougb Ihr
luiUr tnW u> the muioBieter, the lrv«l of the mercury iii the lucciKliiie limb
«f ehkh falU a little, oraioks a lillle al finil, or uiay du iivilher, aecorJing lu
lb MkfTvM «iib nhieh itic prubable riivaii prrwaiire hnn been guooied. anil on-
llnai In eihiblt tbe chiraetetinllc oM-ilUll'init until thu experiment U brought to
u ml by iba Miioil ilotiinx or otbnririM;.
ITneli^p of the im<iv«<ii
_ . ^ iiv«<incnli of ibe oitumn of mercury in tbe manometer may
wukra riihmm a mnohed surface of a revolving cylinder (Fig. )l>. or by m«anii
«f abnuh and lok on a oiniinuoui roll of tiaper, aa in the more C"ai|ilcx kvaiO'
■id-ig.W)-
IWl
THK VASCL'LAK MKC'IIAMSM.
S 116. B^ iht help at Uio taanAtuot«r a])pl!«d to various art«r»eei ntid vt
vn iMrn Uic folluwiuic tiu-i^:
1. Till' iiii-uii 1>lii<iil-['ri'!«<iinr la liigli iu ull tlie arit-rioi, tiui i* greater in
llic lurgiT nrtcrit^ iiritrcr llii? h^-nrl tlinri in lli« miiEillur urieries I'lirthi-r tfum
the hmirt: il iliuiiiii.-'liot, ill fnci, ulong the urterinl Lnicl frutn ihe liuirt
l»vriiKl (Iu- (-ii|>illnrii-9'.
'J. Till- nicnn Wniid jircMuru id luw in lln- *'dii» but, io grrali-r in tin'
emalkr leina near<-r lh(> iiipilliiric* tliun in llic Inrgvr vi-i:i» imrtr ttir UiMtt,
diniioishing. in fiicL, Oom the i-apillariof lowiirvl (he Iiviiri. 1» tliu largo
veins near tlic heart it may bo ifj-ilix-r. thai i* to rny. the pri'S^ure ol' hlmti
ID the vein bcarini^ on the proximal dctudiiJitig limb of the ninnomclcr ma]r
be leas llinn the pr^MUrc of tb<< alnio^phtrc uii ihi' sHCfiiidinc iliital limb, eo
that when coiuniiinication in nindo Wwccd ihe inieriur of tna vein and the
wunomeler, the nterivtiry lunkt in lh« distal and rises in tbe proaiiual limb,
being nuk.'ked tip Mwanl the vein.
na.U.
Si^l
■--a ■
tjCDWM't XvaooRirii roa kaamnjiu tin a CViNTtvunis Kou. up I'atu.
Thti tnanomctor cannot well bv apniied ui the capillarioa, but wo maT
ipcamirc tbe bloml-premtirv in the cnpilhine« in an imlirbct war. It is well
knnn-n that when any portion of the skin in prerned upon, it becoin« pal«
and bloodless ; tiiio is due to the preasiire driving the bliHjd out of the ca]>il-
lariea and miautc veawls and prevenliu); niiy frtvh blimd t^ntcrtng into lb«D).
By earvrtilly inveetiicittiog the iimouni of prtmnre mxiitfiiry to prevent tb«
bli^xl eiili^riiiK ll>^ eapillarii« and miniii<< nrt^riux of |h<< nt-b of tbe frog'a
foot, or of (he nkin bi>uvulh the oail itrelMwhcrv in nmn. the iuternnl pr<«-
Hire which thtt hbHHl id exerciiiiug on tht? walla of lh« capillariea and minute
TIIE MAIS rACTS OK TIIK C1KC17LATI0K,
lOI
■llfja awl v«itu mav be Rpproxiinaiclr <l«termiaoiJ. In the froR's web
bMD fuuiid to lie e(|iiul to about 7 or 11 mm. of merciiry. lo tlie
tb* capillarT liltKxI-gtre&tui^ n naturally higher lliaii thi* aiid maj
!•( ilowa at fruin 'JO to ^U rtini. It is. tlieivforv, c»»»i(l«mble, beiiiK
ptawr llian thm iii lliv vvmn lUonKh \tf» itian lliai iu ihe Brl«ri«a^
JL Tliprp bi llius K oniiiiitiifl di-cliuL' of bliiiMi-pn-BMire Trnm tlie nxil of
tk aurla. llinxigli thv arli^ncii. t'lijiilliirirs, hdiI vi-iti.-t Iu lliv tigltX aiirirl«.
Vrfiod, bi>«rvrr, iiii I'xiiniiniiiinii tliiil tbe mint nmrkt^l Inll of [iritc^nre
lU* |>la<v bi-t<iti.-rti tin- kiiihII ntte rim un lUu utiv xiilir of the nipilliiruv am)
Ibtwiall t*riiu OD iIm> ntlier, iIm' i-iirvo iif {imviin- IkJiij; itomcwliril of tiie
tm given ill Pig. t>0, wlik'li » >iiii{>ly iiK^nilftl li> tlmw thw fact gnii>liinilly
mi bat nM bMO ranMriirtei) by exact ni«ii»iin-miiiU>.
Fill. cu.
I'MOIUI* or niJUnlVPBBMtTlta.
/*. tcH|iliemln4tealmliiiili(Knnte*.<w|il1lrulei.aD4 vdai); V.Miu.
i In tW art«Ti» thk meuo pmaure b niarked by oadllations cam-
In tbe tienrt beats, each wcillatiou coosMtiiiii; of a riwe (iDcrease of
in abote tlie nictm) eorratpondiiiK to Ibe systole of tlie veiitriol«.
by a Call (dfcreaaeof prctwire below tlie moan) eorRS{ionding tu
idiMtoM of the vvutricle.
Vum owdllatioDH. nhkb yn may .-jirak nf iia the pulse, an- lurgcvt and
■M CM^iinwua ill the l«rg« arteric* iK-iir lliu lioiirt. dimiiiiiih fnni (ho heart
Wiwd ibt capillarioi, niid iire, iiiiik-r uriliniiry (lircuiimtuiii-'uii, vrholty hIumiI
fha iba Ttina alnng tbcir nbolc exirut frvim the mpillnrim lo tlii^ hcnrt,
ObriDualy a ^rvat obangv Iaki9 pliin- in tlint piirliim nf thr i-irviiliitioti
vUdloamprion the mpillariiv, the iiiiuiiU^ arli-rii-n li-iidin)i; to am) ihc minuto
*tJH laadmg away from llio capilliirii.^, luid nhich w- miiy apvak of n» the
'pwiplmsl rf^on.*' It is hen- that n great drop of pmviiro tnkM plaec ; it
iikftalwi that thr p^ilse diMpmare.
|UT. If lite neb of n lixiga fixit be ciaminni with a tnicnwcope. the
bliul, aa jiidgt*! of hy the nwvemvnts of the corpuBclcs, is sevii to be pnssin),'
in a DuotuiuiMis rtn-iim from the small arteries through the ea[^llaries to the
im». The vidoeity is giealcr in tlie arteries iIibii in the vv'im, and greater
ii belli llian in tlie capinnries. In the nrleries fuiut puliuitiunB. ^ynehronoDS
ailk iba heart's bent, nre freipiently visible ; b4it these dirapjtenr in tliv
■nillariea, in whtvh Ihe llow is even, that it, not brokeo by piilsalioRs. and
t^ iTeDDceB of flow is continued ou alon;t the veins as far as we enn irnco
iImo. Nut infre<(ueiitlT vnriiilii>n-i in velocity and io the distribution of
lb*h|nud, due tocauM* whieh will l>e hereafter ilit«uai«d. are witiwBsed from
liawif) liine.
102
The chRraclcr of fho tlow ihmugh the smaller cspillnrira is wry Tmriabl«.
8oiiietiin«« the curpiiecW nre seen prntsint; through the i-hnnii«l in siDf^le 6k
with great ref^iilariiy; nl riiher liiues ttiey inar be lew luid far betn'e«ti.
twnie uf the capilliiric^, as we li&ve said in ^ 107, are wide eai>u]{b to permil
two or more carpiiBcle« abreaat. lu k1) caaee llie blnod aa it paasos ihrou;^
ttie cspillary Airetcbea and ex|)iiii<!s the walla. Sunieiimcv a corpuscle uuiy
remain aiaiiuiiary at Ihe eiiirHiice itilo a i-a|iillartr, ihe chauoel itself botng
for some little distance enlirt-ly live frcnn Gur|iuioles. Saraetime* many
corputteJes will appear to rvmaiu uratiuuary in one or mon vapillaric* for ■
brief period aud then in niiive oa uKaiii. Any one of ihettc coitililioiu
readily \vtMti inlo iinulher; and, cii|>i.'cially with a aamewhat feeble cireula-
tiim, itialniii'CAof all of them may )ie«eeR in the name lli^ld of tbe niicrniH'opc.
It in only «lii-ii tbe vt^itN.'li' of the w<:b arc uHumially full i>f hUiod that all
the cniiillnriw r-an he «cn vaiinlly filled with corpuscle:!. Tbo long, oval rod
ourpu»cle inovra wJlli iv long axis parallel to ihc stream, oceaeionnlly
mUttin/^Dii iu lung axis, niid ■i>inrl!me«, in the ]nr;gcr channels, on ir^ short
axis. The flcxibilily and rlnxtiiiiy of a Rorpusclo are well seen when it »
being driven into a capillary narrower thtin itself, or wben it bccomva tetn-
pornrily lodged al the angle between two diverging channels.
Thno ana ntber phenomena, on which we »tiall dwell Inter on, may lie
readily seen in the web of the fn>g's foot or in ihe stretched -out tongue or in
the niewntery of the frog; sad e»entially similar phenomena may be
observed in the mesentery or other transparent tissue of » msmmal. All
over the body, wherever eapillaries are present, the corpuscles and tbe
plauna are being driven in a eonlinuous and llinugh somewhat irregular yet
on the whide Meady flow ihron^h channels so minute that tbe poasage U
maniftBtly attended with cousiderable dilUculliea.
It is obviouH that the peculiar ebaracien of Ihe flnw through Ihe minute
arl^irie*, capillnriia, and veinn affonL* an explanation of the ffieat change
taking place in thi- |icri|iheni] rt^ginn between lh« arterial flow and the
venous fluw. The muted Mretinnal area of tin* capillaritv tt, n.* we have seen,
BOiDO faundrvils of times grvnier than the sectinnul tin-a of the aorta; but
this nnic<^il wclionnl area is nindc tip of thousandH of minute piuwigca,
varying in mmi fnim ■> U> '.'(I u. some ot them, tlien-fore. being in an undis-
leaded condition, smaller than the diameter of a red c(jr[Hiscle, Kven were
tiw blood a simple liquid free from all eorpiiccles. ihcw extremely miniile
piWUflfii would occn«i(iu an enormous amount of friction, anil thus present a
oontaderable o!>stacle or rvaiatttuee to the How of blood through ibom. StjII
greater must be tbe friction and reftisinuce oceasiontu) by tbe ncliial blood
with its Tvd and white corpuscles. The blood in fact meets with great difG-
eultiee in its pasan^'e through ihe peripheral region, and somelinies, aa we
ahall aee, the Iriciiou aii<l resistance are so great in the peripheral reoaela of
ihia or that area ttial no bJmid pa^tiea through them at all, and an amat of
tlie Jitiw lakes place iu the nrea.
Tbe nwiHtauce t<> the (l><w of bloud thus caused by the friction f^uerated
in Ml many minute pawagen i» one of the moet important ph_\-sioal fiuita in tbe
drculation. In the large art«rie« the friction is small ; it increases gradually
M tbcy dividi', but nn-rivM iu chief and most imporluut addition in the
minute art<iriv» ttn<l mpillnriv*, it ix relatively greati-r in the minute arteries
than in the ciipilliirii-* nn accoiinl of tin- IIdw being more rapiil iu the fnrmer,
for friciinn diniinii>h<« riipidty with a iliminiilion iu the rale of tlow. We
may speak "f it as tbe " i>eri[iheral Irietion," and the roixtancu which it
oflersus Ihe "|>eripheral resiniance." It need perhaps hanlly W mid that
this pheripheral resistance not only op|Hisc» the flow of blood Uirough the
c*|^Iartue and minute arteries thonisvlves where it is generated, but, working
THE Mais wxcra of tok cibcdj-atjos.
193
■ckvard Rlvng the whole irurial iyittftin, has to b« uveroome by th« h«irt
1 «acli ■jraloJv of Ui« ventricle.
JlgdraHlie Priaeiplr* of fh^ CirttutntloH.
8. In ifav drrulntinii, th«n. ihc fotlowing* three Gictx of fuDdnnwital
taiiM MTt net wiib :
Ttw >jr*iuJe n( the vcDlricIc, driving at intervals a certnin tjuaDtilr of
Uood. with K eortaiii foixw, into the tiorUi.
3. Tb»|Mripbnal rtsistance just detcribed.
S. A km; MnKch of elastic tubing (the arleriee>, reaching from the
to tlu! region of peripheral resisuiDC«.
<m tfanc fnds wo titaj explain the main phenomena of the circulatiou,
Aith «« have previously sketched, on purely physical piiadpl«s without
U<r arptal to the »{)ecial pn>perti«a of liviuK tiasuis, beyood the provisioD
Uai tKr veoirirle remains c»]>aDle of (pMvd rhythmical con tractions, that lh«
■fkrial walls retain tlieir elasticity, and that the friciiou betwoeu the blood
u4 Ih* lininj; of the peripheral vessels remains tlie same ; nc niuy thus
nyUo the htgb prasurc and pulsatile flow iu the arteries, the steatly streuin
luMffb tbe capillarM«, the low presanre and the unilnrm pulscltMS lloir in
(bniBa. aod nnally the continued flow of the blood tVom the aorta to lh<
moiIm air the vean cavir.
AH ibe atHive phenonH-na in fart are the simple nviiItM nf iin iiitrrnillteot
firw(lilte that uf the sys't'Lilc of the vrjilricle) working in a cl<.iM-d circuit
*f Ismching tulMs, wi iirningul that while the iudiviHiiiil tubes lirel
^iamili in calibre (fmni ihr heart to the cnpillnrie*) anil then incrvusc
Am (h« capillarifv td the hrurt). the area of the bed lint increoHw and
Wi dimtnishts. tlie tubes to;^iher thus forminf; two conee placed base to
it the capillaries, wilh their apic.'cs converging to the heart, aud pro-
at their conjointed bases a conspicuous peripheral reaistaiifv, tlio
cm one side, the arterial, bdtig eminently eluittic, and on the other, lh«
, affiwdioff a (tee aud easy panage for the blood. It is the peripheral
aae ({or tbe reaistaoee oftVred by the friction in the larner vmmIs may,
•hso nniDared with this, be practically ueglecl«d), reacting through tlie
•Wc walls of the arteriea upun tlie intermtlteut force of the hiuirt, which
pirn the drculatiou of the blood its t)cculiar fcaturu.
iU9. Virtumtlatuet ilftirmmiuQ Ihr. charaeirr of lAe fiow. When fluid
k dfivfO t>r an iDiermittent force, as by n piinip, through a [lerrnctly rigid
M^ soeh aa n glnas one for a fy*t«m of xuch lulim), there nu-apvs at eavb
mb of the pump from the di*tal end of tbi.' tube (nr .ivi>lem of 1u1m») just
■ ■aeh fluid AH cutirt it at the proxiiiinl end. What bnpiiciM is very like
ah* woald hapjien if, wilh a wide glsn tube completely Bllixl wilh biliinrd-
Uls lying in a row. an additional bail were puabcd in nt one end ; each
hsD would )ir p(i»li<Hl on in turn a tl»fp further and the last ball at the
hither end «i>uhl tumble ont. The eacape, morvovor, takes place at the
—«» lime «a the entrance.
Ihi* rcanlt remains the §anie when any resistance to tbe flow is introduoed
Mo the tube, as for instance when the end of the tiit)e is n.irrowed- ThA
: of the pump remaining >he same, the introduction of the resistauoe
•btedly Icawns tlie ijiiantity of fluid issuing al the distal end at each
s, Imt it at the same time let*«iis tbe quantity entering at tbe proximal
lh« iolluw and outflow remain cijual to each other, and slill occur at
saw time.
la an alattic tube, such as an India-ratiher one (or in a system of suoh
nW). whose aectional area is suflirientiv great to oJler but little reuatanoe
13 "
TIIS VA801II.AR UBcnAXltsW.
Ik iIh- |>rogr«M of ihe iluul, the flour ciiused br an iDtermiUeal force b rIw
inli'rnuttcut. The outfluw beJOK uearly as t-oAv as the ioSow, tlie elulidlf
of till- \>flllii (>r thf tuW i> H-anvlv iit «)l talletl inio (lUy. The Lube behant
SnctH'til))' Itk).' H rigi<l iuIk'. \Vbi;ii, h»w«ver, auliickiit rmMaDce u intn-
ucc<l into liny iitirl of tke voiinto, the lltiiil being unnbte to pnat by ibt
mwlancv m ntfHmr ni> it eautn tJw tube fmn) i}i« |iunii>, lenilfl to acminiiliie
on thi> nroxiniiil inilo of iIhi nxiialaiu'iv Thi* it u Able to ilu by ex[i<in(lia(
tbeeinstic wuli* of tbitliibv. Atuivh Mrukoof the fHimp n vertnin ()iuuil)l}'
of fluid ctitcn' tli« tube at l)ie pruxiuwl cud. Of iImh only n fV«<-lioa cu
pBA tJirough t)i« reMctiincc duriag the »troli«. At ttic iw>m«ni whw thi
Mrokc CM8W, th« rait »til) nrmainM on (he |ir(>xinuii aide til' the nvislUM,
tlK elastic tube having «xpaud«d to reoeivi; it. During tb« inivrAiil bvlwwa
tiiia and the next Rtroke, um dietendnl elanlic lulxi, Hnviag to rrtum in !tt
natural tuidulanded condition, prcnn on lhi» extra ({uanttty of Huid whidi
il coiitaina and tends to driro it past tlie mistnncw.
Thiia. in lh« rigid lube (and in the oln^tic tube withotil the reoitaocf)
tliere ie«iic&. f'runi the diolal end of iho tube at each stroke, just as nnifa
fluid as eniere it at the proximal end, while between the strokes then !■
perftet (juiet. In tha elaslic lube with reustiuice, on the ountrari-. the qiuu-
titr wHioh jMxto the renalaace is only a I'rauiiou of thai which euienilit
tiilte fnun the pump at anyone stroke, the remainder or a portion of tlw
reinnindcr <'(mtiuiii»K l*> [>sm durinjt thv iuler^'al between the strokes. U
Uut fonuer ruav. Llie tube u uo fuller at the end of the stroke than at tW
b^innin)! ; in the latter caae there in an necuniulation of fluid between ibe
pomp and the nsiHtuum, and a eorreiipmditi}; distension nf thai part of ibe
tube at the rtiBC of earh »lriikt! — an nci'timiilatiun and dislciiBtitu, liowever<
which 20 on diniiniohiiie during ihir iiitcrvul hrtwevn that Mtriike and tb^
next. The nniminl nf fluid ihiiii remaiuin); ufter the Htruke will depend uO
tke amount of n-»i»Ca»pe in rclnliim In ili<; force of the sLroko mm on ll>*
tliateiuiibility of thit luhr ; and cbr amount which jiaflMM tbo mistance Wnr^
the next rtroke will ilcpviiit on the drgriv of elnstJc rottction of which tl»*
tube is capable. Thus, if l)i« miRtnnco be very iiinsiderabic in relation t^
the force of ihu rtrokc. and the tul>e vtrry di«tcn«iblc, o«ly a »miiil n>nioav'
die Saul will [lOW the rc«i«(iu)ce. the gn-nier [Hirt remaining lodge*! betwve**
the pumn and the rwaistanoe. If the elastic roactioii be grent, a larife po^'
tiun of Ihis will be pmed on through the mistanoe before the next stroke
conies. In uther wunU. the greater tbe resistance (in relation to the for^
of the stroke^, and the more the elastic force ia brought into play, the lei^
iulermittent, the more nearly continuous, will be tbe Bow on Ihe far side<»
tke resistance.
If Ihe first stroke be succee<led by a second stroke before its tpiautity of
fluid has all passed by the resistance, there will be an ailditioual accumula'
tion of fluid uD the near side of the resistance, an adililJonat dbteurion of
tbe tube, an ad<Utional strain on its elastic powers, and, in con*er]iieiice, tbtr
flow hetneen this second stroke and llie tfaini will be even niont miirki-d thUL
that b«iweeu tbe first and second, though all three strnkcM rn-ri' of ilw sani*
fom, the addition being due to the extra aiuouiit of diuttic (•<nx called int»
plav. In fact, it u> PviiH-nt ihut, if then.- 1h' a sufficient store iif eliu^ic power
to lall back u|>uu. by i^oniinually repealing the strokea a state of thin|f< will
be at la*t arnvwl ut in which ifie plastic force, callc*) into play by the coa-
dnually tncn-iuiiug dinicnxiou oi' ilxi tube on tlie near side m trie reeiuanoe*
will befuRicii'iit l<> drive thnnigh llie raualauoe, between each two siroksa*
jtlst a» much fluid iw cnlcm the nirar end of the syalem al eneh stroke. Ik*
olber words, the elastic renclioii of ilu- wnlU of tlie tulx; will ha?e OODVMtcA
the iut«riQitt«iit uuo a ooutinuMia How. Tbe flow on the br aide c
THE MAIS FACTS OF THB CIBCUI-ATIOS.
IM [« in lh!s <-«M not tlie •/!>«¥< reeull of the strokes of i)iv ]iuii]|>. All
v» nf ihc pump ie epenl. 6ret iu getting up, and afterward iii keopiiij;
i^ tlio ilieleo^ion of the Uibe on the near eide of tl>G resistance; (ho immft-
Kte caiiac of the coDtiououa How lies in the disteiiMon of the tul)c nhiob
his it to eoipt}' itself into the far side i>f the resistonee at suob a rui« that
dtschar][«8 tlimuifh the rrais(aii<-e dnriii^ a stroke and in th<! viiccecding
l«ri kI Just la luucti as it recirivui frum (he pump liy thi? ilrokc itiitlf.
Hiis id exactly uhat Inkm plm-^ in the viiscuiur avfllein. The friction iu
p minuie artc-ritv and ciipillntie* pr<4eiil)i a coiini<MrruliI(! rexuilaiiw to the
f/n of hloiHl through ijieni iiito thi* Kiunll veins. Iu coni«-r|uence of tliis
UatMice the ibroe of thr ht-iirt')! heiit m «pMit in niiiintainiiig the whoU of
te arla^rial pyoiem in a ilntv of grent di>Et'n>i>in ; ihr nrlerinl wiillii are put
t^Mif iiu tli« Htretch br tbe preHura of ihi.^ hliiuil thrust into (1r-iii bv the
t|ar«trd olroko of the heart ; this U (lie prtroiiro wliich wo Bpuko of above
I bluHl-pnwiiire. The ureatly dii^leudtd iirtrriul qretcm in. br the elastic
MolKMi of lis elastic walls, eontiuually lending lo rmptv itself by uvertlow-
p% ikmuf^h the capillnrtes into the venous svMt'ni ; and it ovcrtlows »t such
tnUi ihni juBl ne much titoixl i>aai«s from the nrteries to the veii)» durlug
Mi ajstole and iia sucL-eediug diastole as enters the aorta at each sfelole.
«
ru.<t.
■:l/fi
Aatnu*t Sirnm.
— „il. ball clMI'* inlxtn n(«tMiil ih* trtmal •jitem, bnuicblni*! X nii'l V. and (ndlns
*kn^ninr |i<Tl|>lwnl mMtaim. lacli>llnii(hocai>lllkrl«^(rblchBi«iinlutol br olUav IoomIt
'*'^IMU jtvm bI tpiinc* i*» |«ru iiliaoii u dilated tn llie ngmr. Tlw ea|4UiiTlgi air galliond
*'MUi tmim (j-tiiB. ibiulol. H'hKb icnnlnaWaatU. WaMr I* drlrtn into the anntal vj»\ma
I'T** "" '■ er ail i-laMt* fe*|t<yrii>«D <» anr o(hcT ream drpimpi, C1aliiiaB» ptu«<J on Ibn iindl'
"'''Mar, (•, p". W'li(-n llivw Dlani[atanll«lili-iird, Uiruiljr accaa Ibaltir aatiir it'Oa ttiaanarlat
'"'•••■aBMitalalliR'iialinii;4IMrd parla niird wini iirODg*, whif h e An a raiuldanbt* nitat'
^*<killO*ar I<ildUm<ti|ibtbaln. Wtttn Ib( plDin|danDlilaaK<! HtcHiild pwaca. lalUi OMtcfa
"■MtNutra^ Hiimifh ilio iindllaHd litbM> Tliiu. Iiy ll|[hltaIuK or louMitii( lh« elamt* tba
'"Mmt' naliiaiun mtf In Intrt— <l or dIjnIoMml M jjwjnw.
'**.<• llMtfiariai -hie. aiKiai V. oa iho nnon* ilitc, iMiwanetcncan be altaobed. Aiatnd*
^■Mtt I «iid f I, bj nmiiii or riaini*. Itw nuir i>r dald ttt^ an ait^vy add team a vidn, ondar
"^tcMIBwk awr l« «t«tnO. M fa. fa. and t^r. t|ih]iiiia«n|ibi may be appBtd.
IIMI, ladeeil, the tniportant Incls of tbe circuliitioii which we have not iu>
T*i i>udtti) may In roughly bm Hicccwfully iroitnled on an nrlificial model.
Fig. 61, ill whioli an elastic syringe reprefieoCs ibe heart, a'loii(f pi«v
dasiic liidia-rubWr tubing the arteries, anoilier pie<.-e uf Iiibin;^ ttie veins,
and a number of Hiuallcr (.-ounecting pieces the minute arteries and onpil-
lari«e. If these connecting {liecea be uutde at Ant aomcwlint wide, so sa to
oSer no great resistance ti< ihc tluw tVuin the artitidal nrlerii^ to the artificial
vdns, but Iw eo arranged that thev maj be made narrow by the .tcrewiuK -up
of clainporotherwiiic, it i^ pouibli; tti illuHtrutc the behavior of the vascular
luechamain wben iJie i>eripliural resiAtancL- in lt«s ttinii umiul (and &• vv »)iall
•e« later on it is poMible in the living organism either to reduce or to iiicrcajc
wtiat tnay be considered as the norinnl jieriplieral nwiAiancft), and to conijwre
that behavior with the WhMvior uf the inecbaiiium wbon the pt-riphcral
n«i«tauce ia iucreaMd.
TIh! wliole apparatUK being placed Bat on n table.KO as to avoid dilTerenocs
in level in dillerc.nt parta of it, and with water, but so ns not to distviid the
tubing, the iwa iniui<>inet«ra attached, one (A) to the arterial si<lv of the
tubing and the other (V) to the vchoub side, ought to show the mercury
■landing at eipinl heights in both Mmlw of both iu^irumcnls, since nuthiae
but the pressure of the atmosphere is bearing on the fluid in the tubes, ana
tat equally
If,u<)w,thc
that equally all o\'er.
J pieces b
reaistiince being verv little, we imitate a ventricular beat by ihi? stroke of the
he conni'cting pieces being fn>oly open, that is to say, the peripheral
pump, we aliall observt: the following : Almost inimediatefy after the stroke
the ntercury in the arterial manometer will rise, but will at once fall again,
and very shortly afterward the mercury in the venous tube will in a similar
loaiuier rise and fall. If we repeal the? stnikes with a ni>i too rapid rhythm,
eaoh stroke having the same furoe, and make, h^ may by a iiimple contrivance
be eflevted. the two manometers write du the Katue i«coi-diug surface, ne shall
obtuu curves like those of Fig. 62, A and V. At each utroke of the pump
no. an
TaAOirai T*SB( PMM AR Annnnii. 9rniuni, with rax FKHiruniAi KnoBrunx Suuur.
A,ut«U: V. Tvuoui nuiunnoler- TlitiOiiure, ui wntiocc, !• m t ■nullwMSle Ikan tboror-
n«poiidina r%. <y.
the mercury in the arterial manoineten riaet, but forthwith fnllt again to or
nearly to tlto l>a»o line; no mean arterial prcMum, or very litth^ in •■tabliibcd,
The cionieoi* of the ventricle ("yringiO thrown into the arterial »v*t*tm dis-
bsnd it, but the jiMmgo through the jieriphcml region is no free that an cQiial
quantity of fluid pas*v« through to the veins inimedintelv. and hence the
iDorcurv nl once falls. But the fluid thux pnxfing casilv into the veins dis*
taoda tteao too, and the mercury in th<nr manometer rfsce too. but only to
&U Bguii, as a corresponding quantity i^ines from the ends of the veins into
tbe basio. which »ervo!i as an artilicial auricle. Now introduce " peripheral
fesMtonoo" by screwing up the clamps on the connecting tubes, and set tbe
TDK
^■i[>to «Drk again u bcfure. With Uw liret vtroko th« nwrmry in tbe
wuwil Mnowrttw (Fig. 68. A') riw ks before, but inHeadaffiUUiiff rapidly
t bSk alowlf. becBiiw it now uk«a ■ longer (imfi for a quantity of flui^
iqMl ta thai wliidi hns Wn thrun into th« arterial snum or tl)i> veutrica-
«^i«ka Id |HUa llirt>ii)(li the narroireiJ |icri|ili«ra) region. Defore ibe mirre
Pni.«.
V
twaw T<as9 raoH ui AimnatL Scmc, wim nrx PranmiL tUMmuta OmnKiuHa.
A'. HMrlal ; V>, nnoaa iia mill nil
bwUIni til tbr baw lin«. boforo the arterial i^vtem has had lime to dia-
thai]gc lhniu)[b ibi- narnivroil iH<ri[ib?ml rpgioa aa roitoh lluii) as it receinjd
I^M tha TTOtrirl)', a woiiii) Ktrukv <lnvr» more fluid into the urterieA, dis-
Mtfiac ibrm tbi* limi- inori* ihnn it ilid before, aiul raitiiiijt lite luereury to
a ffill ht);h«-r l«Vfl. A thini, a foiirtli. itnd suoeeeding Mriike- ginxiuce tbe
■airrflcrt, cxcejit tliHt tbe tulililionul Jveigfat to which tbe uifreurv h roitHHl
•t taeb (trc^v becomes at each 'trxke le«s and )i?s8, uulii a ntatv i>f Iblii)^ is
nocfctd ill whieb the mercury, briof; uu the fnll wbei) ibe ntruke lalcvn {>[ac«,
ft bf tb« Mnike raiMii juHt as bi);b as it wa« before, and tlieii )i«f(intiiuj; to
Ua|P^ is aicaiu raised just as bi^cb, and so nil. Witb i.-Ath siii'c«<diiig
Mnlw the arterial ByUem has becooK mure and Riom dixleixleil ; but the
■en ilisiMKled it is the Kreaier ia tbe elastic reutriiun brougbt iuto play : this
flMcr clastic lettctinn more aiid mora overcoaKS tbe obstacle proaoDt«() by
la* fcHpbrral mietance and driv» the fluid more and inoro rapidly tbrouKii
tht i«npberal refcion. At laMt tb« iirtcrial systam !■ so distandoo, aiid tLe
faftc of llie elastie rcmotioii Mt ffreal, tbat durinc the sirokv and tbe sucoeeil*
m( iaurval just ua mucli fluid ]>aNN,x> ibraugh tbe peripliemi r^^ion M oiitara
tb> ■ftJ'rtm Mt (he ■ln>kir. In ntlier vrimls, the rcpratt^l ttruk'* have catab-
bbtd a menu anrrinl iirawiire which, lit ibc |H>itil where the mnnometor
ii sAsad, is raised sligtilly at eneb vMitneular Hroku and falls slightly
b<t«m 1^ strokes.
Iteaiag now u> tbe vcn<Mis maiKviocter, Fig. 6S V ', w» observe ibiu each
198
v^
Ctmlcc nf the pump [>^}<iLlcca on itiin much less ofTort than it dit) Iwfon L_
intriKluclioii of the iiicrcuwd ])criphcml raeiitAnce. The iiiercurr. inntewl^
diftinoUy rising aiwl faUiDg at ench Htroke. now shows nothing more th>n
v«rv gentle uuduUtions ; it fe«lB to n very slight degree onlr the dirf^i plfe<4
of the ventricular stroke; it ia simply raised slightly nbore the base line,
and rcmitinB lairly steady at this level. Tlie sli;;!it riae marks the nieMl
prenare exerted by the fluid at the place of attaclioient of the manometer.
This tDefln " venous " protHure ia a continuation ul' the mean arterial preeuure
BO obvious in the arterial nmnonaeter, but a much leu tlian that because a
large part nf the arterial mean pressure has been exi>eiideil in driving the
fluid past the periphera! reeistant-e. What reiuatDx i*. hciw«ver, ttultideiit to
drive the fluid aloiiji; the wide venous tubing; ri^ht t<> llie upcn end.
Tliud ihii aritfifial niiidcl may be made to illuntrat*! liovr il comi:* about
that ibe bli)iii] ttiivi* in the artdrie* at a nOativrly high pn^3t«ure. which at
aach ventrioulnr )>yst<d« in mi»ed (lightly iilii)Vi\ lux) at ■■Jkch diikdolc falls
slightly t>ejiiw, a cttrtitiii nicjin lcvl^l, and llowii in thi^ vi-ini> at a much lower
prt^Mure, which iiutt not tihi)W thu immcdi.itc i'iret:l« of each heart-hwit.
If two ma nomut •.■»>, instead of une. were atlAcheil Irt the arlfriiil *y»lcm,
one near the pump and (he othor further ofl'. close to the peHpheml miM-
anct, the prrasuiv shown by the near mnnomolcr would hf fuund U> be grralcr
than that shown by the far one. The pressure at the far point l* l«M
becaHio some of the pressure exerlod at the near point has been uwd to drive
tlie Huid from the near ]ioint to the far one. .'Similarly on ihe venous ude, a
luaDomotor placed close to the peripheral region would show a higher prea-
aure ibaD that shown by one further off, beeaiise il is the pressure still remain-
ing iu the veins near the eapillariee which, assisted, as we shall see, by other
cv«nts, drives the bloud onward to the larger veins. The blood-pressure is
nt ilM highest at the root of the aorta, and al irjt hiwott at th« mouths of the
Tcnii; cnviL', luid is falling all tbe way from une point to the uthi-r, because
all the way it i.-> beting usi-d up to move the blool from one point tn the otlier.
The great drop of prewure i», a^ we have said, in tlie pcrijiherai n^on,
because more work has to be dono in driving tlic l>l<x>d through this rvgimi
than in drlviiig the blood from the heart to this region, or from this region
to the heart.
Tbe manometer on the arterial side of the model shows, as we have seea,
an (^'illation of pressure, a pulse due to each heart-l>eat, and the same pulse
nay be felt by placing a finger, or rendered visible by placing a light lever,
on the arterial tube. It may fbrther be seen that this pulse is most marked
DMTMt tlie pump, and becomes fainter us we paw to the periphery; but we
must reserve the features of the pulsa for a sjiei-ial study. On the venoiia
side of the miwlel no pulse can be detected by tbe manometer or bv tbe
Soger, provided that the [leripheral re*i»tancebeadoi{uatc If tbeperipWrol
NMStaocc be iliminished, aif tiv uiiKCrewing the ctamjvt, tbeti, as ueoeaaarily
follows from what bus gixie biifore, the pnlxe psjiseii o\-er on to tlM veixxM
side; and. ns w'l- iihall have uccneinn to putnt out later on, tn the living
organism tlw (>crij>b«'rHl nvistancu in {mrticular an>ai> may beat linKtao
much lessened that a distinct pulsation anpcJin in thv win*.
If in tlie model, when tho pump !■ in lull swing, and arlvrial pNifun.' well
eMabltiiied. ibe arlerial tube be pricked or cut, or the sninll xidn tulw a
be opi-ned, the water iiill gij*h out in jets, as dues blood from a cut artery tn
the livin/ iMidy ; whereas, if the venous tulie i>e similarly pricked or nil, or
the small lulie r t>e ojieoeit, the water will simply ooze out or well up, as
does bliKxl fnrni a vein in tbe living body. If the arterial tube be ligatured,
it will swell on tin- puniji Bide aud shrink on the jteripberal side; if the
venous tube W ligatured, jt will swell on tlie side Dearwt the capillaries and
TMK MAIN rACTS OV TltK CI KCL'LATIOS. 199
•Kriak on the atber sidv. In short, the dead iHDdol will tliuw all tli« niiila
fbrta of lh« circuliition which w« have as yet described.
{ m. Id tbc- livinc lioily. hi>wever. there ore certain bel[is l« titv oircalii-
tioa which oaniiot be iiniimed by *v<-h n roodd wilhuut intmdnciug great
•ad UiMledrRble coiupIinaiuiiB ; bul ttine diietl)' ittri-vt the How iilniig tlio
xtiBm.
TIm TviDS uv in nuuir placm provided vith valvts so t'nnatrudod as to
oAr little or no naiMance to ilie llnw from the cii|Mll«rios to tliv liparc, but
aallv to block a reitirn lowanl lh« eaplllarirs. Hencv may cxtvra*]
n Drought t'< War urHiii ii vein U-nds to help the blood to mnvo for-
lownrd the Itrnrt. In tbt.- various mo vera en tn carried out bv the
1 mitK-li-f, [>u<:h an exleninl prcmiire is brought to bear on many of
vriii*. and 1m-dc«- tht-sc niovcnients imi«t the circulation. Even juiiiaive
KioTrini-nla nf the limbn have n nmiltir ef^t. So, also, the in ov emeu is of
the alintentarr canal, carried out by means of jdain muHOUlar tissue, promote
ihe 6u* aliMig the veins coming fnini that canal, and when we eouie to deal
vtlh tbr spleen we shall see ihat ihe plain muscular fibres which are so
■htiadatit in that organ in mnie animids, serve by rhythiuieal contractiona
toPsnip Ihe blood re^ularlr away iVurn the spleeu along tbeipleuic veins.
Wlwo we t!oni« to deal n'lth respinilion, ue shall see that each enlargement
rflJM cbeu constituting lui iDi>|iiniiiou tends to draw the bl»o<l tiiwanl the
that, and each return i>r reiracliou of the cliesi walls in expiration lends to
Mrs the bhiud away frotn the chnt. The nrrangenienl i>f the valvi-» nf the
ban cauaea thin actiun of the rt^iiiradiry pump li> prriniuU' the How of blood
io Ike dlrvctioti uf (l>e normal circiilaliun ; and, inilc^il, nen? ihr hciirl per-
iuljr iDntinnl<M>, tlie working of this rir«piraiorv ptimp atone would li.'iid to
itin t)if> btood from the venit oavie tlirom^h the heart iuio the nortu, and so
In ksMt op the circiilalioii ; the force so exerted, however, would, without
iktiia of llie iK-art, b« able to overcome s verv »iuall part only of ihe resist-
lon ta Ihe capillaries and snull vessels of the lungs, and so would prove
idiislly inefTectual.
Tbefe are, then, several heljM to the flow along the veins, but it must be
imnabered that, however useful, they are helps ooIt. and not the real cause
tl thi> cimilaliou. The real came of the flow is tlte ventricular stroke, and
liii i* iofficient to drive the bliMnl from the tel\ veiilricle tu the right auricle,
when every muscle of tlie Ixidy i.i at nnl aii<l hmtlhing ii> for a while
1, when, tlterefore, all tlte hcl]i« we are speaking of urc wanting.
CCVrcHMMtoiuvi DettrmiHmg the Rale of (Ac Flow.
i 132. We aaj now nun on to consider briefly the rate at which the blood
Of through the vfmelt>. and fim the rate of flow in fA«r artfrie*.
Vhen even a small artery is severed, a ronsidcrnble qiiantily of blood
faapn tr\na the proximal cut end in a very short sjiaee uf time. Thai is to
■t, ihe blood moves in tlie arterira from the iiearl lo the capillaries with a
Tfty ninaidemble velocity. By various methods, this velocity of the blood-
annir ha* been measured at diflerent parts of the arterial systein: llie
■»oli>, owinK to iinpeifi'<-lio<u» in tlte meibods employed, cannot be regarded
m Mtldaii'irily exact, iHit may be Bi-cefneil as approximately true. They
sluiw that the vd'M-itv of the arterial >tivani u. greutnt in ihr largest aiteriea
acar the heart, and liiniini^lxv from Ihe hmn toward the cajiillari(«. TbuiL
ba UtYT artery of n Inrgi- atiiinal.sucfa a»th« carotid of a dog or hone, and
jirshab^ in the carotid of ii niiiti, thi- I>Ioih1 Howm ai t)ic rate of SOU or oOO
■u. a irroud. In ibi< very tmull arteries Ihe rale i* probably only m ftw
mtaiceaiKL
£00
yfrthaiii. The bicmndmiaiiTnelcr nf Vnifamann. [t'ig.SI.] An •Hrry— '. y.. »
carotid— in rlnmpod in two pliCM. ntid dividi-d bclwecii thrclonip*. TwocanulM,
of s b<)r<! an iiMirly e4unl ni riuMible to tbut urtbo urterv. or of ■ Icddwd Irotf. mre
Inaetti-il iii ilie two i-nds. Tbe two cauulie iiri- cotiiiecieJ by mi'iuu of two siop-
coclCD. whii'h work togtxher. wiili Ibe two endit of a long ^last lube. b«i)t in iltD
Rhnpo of It U. and filled nilh nnrmnl inlino Holutinn. r>r wilh a calorri) inn<icui>a*
fluid. The clnmpK on the uricrv lieini; rnlciwud, « turn of the stoficiwk* pennit*
tbe blood to ealvt the proximuf end of ibe lone U'lul^i along which it MMirae*.
driTing tbe fluid out Into ib« arterj' throuub tlie dUlnl «iiil. Attached to the
tube !■ a graduiil^d scale, by ii)e«ns of wbiiili Ihe vclociiv wiih which iho blood
llovra along the l<ib( mny bo rpod ofl'. Even vupjiwiiig the cnnulie xa he of tbe
•ame borp im Ihr iirlorv. it in evident that tbe oenditioiis of the flow ibrouch ihe
tube are BUch w will only aduiit of the result tliuii gained being coutlderea »« an
approximative eitimatlaa of the real veloolty lu the artery llselT.
I Flo. M.
/
7="
/
w
J
VnumASir*! HjoiiniioianiicriEu.
"nn ronlaJ [■mumi i-i <1<d ln<tnimaii( an Inwrlvl hi ilii- i-ot gn<ti of* rein or MHrr. Rjrl
dMpIc utameninclil of n dbulilo uoiixick Iho bloal-ciiiTciil can Lc mule l-i T'Uk ImniadlMcty
UuDivli tlw i"aiT*n* am, •■ 111 A, or (n )dM [littiugb ilioBailiiaMU-thapod tube, w lull.]
The rheomeler (Slrorauhr) of T.iidnig. Thii cornuo* of two glan bulb*. J nixt
B, Kir. C5,GamiiiunicatinK Hbovc with eueh other Hnri with the common tube '*,
by which ^ey can be filled. Their I'lwi-r end* arc llxed in the metal d)«c b.
Wbii-h can be mftde to ruUte, through two right anglea. round the lorrt^r dine B.
In Ihe upper dlftc are two hole?, a and b. cnnlinaon* wlib A aod B rtejiectively,
and in the lower di*c are Iwo nimiliir hole*, a' and h'. limilHrly continuous with
tbe tub«e G nnd //. Hence, in the ptwition of the dliirs tthown in the flgure. the
labe (1 i" conlinuou* throueh the two diao* with the bulb J, and the luhe //
vilb the Ifulb B. On luming tlie diK- D thtbii):b two ri^jht anplei. tbe tube Q
beoonea contlnoou* with R insieud of A. and the tube fl with A Jmlead of B.
There b a further armngemeni. omitied fioni the fiKuro for tbe ntkeof oimplieitjr,
bv which when the dii>e />iii turned Lhrou;:h one iniilend of two right anflea from
eilbcr of (he alHire poiiitioiiii. fl brevniw directly cuiilinuoiiii with H, boib being
campletely shut olT from the bulb*.
TIIK MAIS KACTS Of TIIK CI K<ULAT10K.
201
Tb* niila hT llie iub« H and O an oiute (o fit exnclly into tmi c«du1» iiwrttd
I lb* i«» cni enib of Ibe »,nttj ubnut to ti« eiptvi mealed upon, and Uavio); a
I BMrlf *(|iia1 u jiOMibl« t(> Umt of tti« artery.
FM-tt.
^<
:5te-
Lrwvti>> Svaomnu did t DiAi.aAXiMttc IlKranarrtnoK or Tiik i*xt.
Ttt sirtbod of experimcDlin^ Ih ho follows ; The dim* D, liciiig pl«c«d in thv
IHmiillal* pontion. tn ibai u and h nrp liotli cut olT from «' and b', tin- bulb .1
ii DM whb pun olive nil up l'> thi.' mnrk x. and Ibc bulb B, tlie rrst of A. aud
Itejuncilon C, viib JclilriDnli'd blood; aad V M tbcn clampnl. Tbe tube* II
ni'itntiao Bll«il with dclibtiiiiit«d b1u<Hl, ami 0 U inaWod into tbe canul*
if tt( tMlral, B into thai of the i>eritib«rBl, end of tiiv iirt«r>-. On rcmuviDic
Aadmn from tlw arWry the blood Bom lbrouj;li O to //, mid an bark into tbe
•fifty. Tbe obavtrraiioD now brgins by taniltig the due D into thr poailioa
•km ia the (iiture; ibe blood tbrii fltwi inln .1. drinng tbi- oil there I'lmtaini^il
«tWfor« It into iIm bulb B, iu tbv dircclioD of tbr nnon-. Ibe dofibrliMied blood
"' — \j pnwBt In B poaunc by H inlu tin.' iirtrrj. and au into tbe nataoi. At
HMftit tbat Ihc blood la an-n la irae to Ibe mark x, ttie disc Dw with all
lie fsptilitv tunieil Ibrougb two ri|t;bi noglea : and thiia the bulb B. now
J ftlltd viib oil, placr<l jn commii nidation nilh O. Tbe blood-rtreum DOW
lira tbe nil hack iiilo .f . aitd tliu new blood in .1 tbrouicb II into ibe arler^.
Xumm'a 1.1 ibi- nil ban wholly tvtuinvd lo ita original [Haitioii. the iltM {a agnm
Rrii ind A raiev taorr placed ia conmunlcallon with O, and the oil onc«
ma- , :r<>m A to It. j^hd tbts la repeated aeveral tinici. jndrcd eenerally
■bJ Utr i-li'iiinK if lb« blood (It ih«> ailmixttire nf tbo oil with Ibr bloud iiuta an
tu Ibr eiiicriEnrnt. Tbiia tbo (low nf bl<)o<t i* uied Iu fill uttcrtiat«ly Willi
' HT «dl tbe ainiv of the iMilb A, whuac caviiy a* far ■■ the maik x biu Wen
,' MMHured ; boni-L' if ibu imiubct of linitn in aay j|;lve» time tlie dUc D
ke lurneil nxind bi- kuowD, the uuinbcr of line* A baa beoa filled b alao
a, aad tbn> ilio quaniity of blooil wbti-h baa paatod in thai time througb
Hia connrftt-d with ihe tube O in directly meanurcd. For instance, an p-
;lluit ih<- 'piNKlily hrUI by Ihc Uiilb J when tille'I up to the mark z iaSce,,
,, tj,„{ |',„g, ti,^ moment of allowiog Ibe dxtX li c.c. of b1'>od tn b<f in
it<« to the monienl when tbeCKa{>e of Ibe laat !^ c,c. fmra the artery
. ;... . *■■ eoniplcte. M) Hconda bad elatwrd. dnrinc »bich time A ex.
'kal been recvived ten timea into tbe tube from tbe arte/y (all but tbetaatG c.c.
'Mi
THE VASCILAR MECHANISM.
W\it)( rcluriietl inti> ihe dit>(al |>ortioD of the artery), obviously 0.5 c.c. of blood
liHil Hl>^^'t^<l t'miu the pruiimkl a«ctiuD of the anerj id one Mcood. Heoce, up-
)>iniiii>; ilint tW diameter uf the caDulK'(and of the nrtery , the; being the bum]
were 2 mm., with area Ibt-refure of 3.14 wguare mm., an outflow tbroueh the tec-
liiiii iif (t.f) i-.o. <>r .MHi cm. in h lecond would give (?"'') a Telocity of about ISt
mm. ill H niHiinil.
'I'lic linnmilHi'liometer of Viemrdi [Fig. 66] is constructed on the principle of
iiii'iiHuriti); the vt'locitv of Ihe current bv observing the amount of deviation nudet-
>I<iiLit by u )H>iidiiliim. the tree end of irbich b»nga loosely ia the stream. A sqnsn
or reotangutar chamber, une side of which is of iIm
anil marked with a graduated scale in the form ot *a
an- iif a circle, is coonected by means of two slwrt
tuliea i(ith the two cut ends of an artery ; tiie blood
ciitige>|uently Howh from the proximal (centrml) px-
lion of the artery through the chamber into the disti]
^H II '' -^t I )'«>i'tion of the artery. Within the chamber and wf
^\ '^ '\^^\ I l«'inW fri>m its roof is a short pendulum, which when
I ''jiji^J^^M f ||],> b!iH>d-»tream is cut off from the chamber huigi
miitionkwH in a vertical position, but when the blood
is allowed to flow through the chamber, is driven b;
the fon'e of the current out of its position of rest. Tbi
iiriiiliiliiiii in i>o |i1m'tit lliMt a murker attached to its free end travels close to [be
joiii-r HiirfiiiT of till' glitMi nidf aUDig tlie arc of the graduated side. Hence tlie
himiiuiit iif deviulioh fMUi u vertical position may easily be read off on the ictle
(rum (111- ouliiile. The graduation of the scale having been carried out by eiperi'
iiii'iitlii^' nitli HircnuiH of known velocity, the velocity can at once be calcalited
lioiiL [lii> iiiiiiMiiil of ilevialion.
All iiiiilnimeiil liiiMi-d on the siime |>riiK'iplc has been invented byCbauveanud
ii»i>tovi-d liv l.orli't, Kif;. tiT. In Ilii^i the part which correeponds to the pend alum
ill N'iri'oiill'i' iiiKtruno'iit is prolonged onteide the chamber, and thus the portion
ullliiii till' rliHinlii'r is iiiBik' to form the fhort arm of a lever, the fulcrum of whiti
in 111 ilie point nhiTi- ilie wall of the chiimlier is traversed, and the long arm oi
n I yi VI M nitMi LI i; ^ii V i>k-
■ PIII'l. 11. I'. »11< Mil to '11-11-^ ]
Kii., 67.
1! 1 ^IMAi-lllOll.n.Jl Ml^ f'llA^'VKAU AM> LliRTET.
» Nii'li iiiiijn In on(Hi<]<', A soniewliat wide tube, the wall of which is at one poiD*
i'iiiii|io?>i'<l i>f nil I III lilt riiUKiT iiiembranc, is Introduced between the two cut ^**?
Ill 111! til ii-r> , \ hnin lij-lii K'vcr pierces the India rubber membrane. The Aot*
ixiiiiiiiliil HI 111 III lliin IfViT jirojirliiig within the tube is moved on its fuleruni »"
ilu' hidiii iiililiir riiij: liy llie cnrrciit of blood passing through the lube, ^"^
jiu'-ati'i ihv u-loi-iiy of lli'e current, the larger being the eicursion of the lever-
TOE MAIir PACTS OT THE CI KCfLATtO.V.
n> BMnnila <iT tb* ihitrt anii glT« ri»e to fxMTMpoiuliiig iiioir«nirnli in the
Iffrito dlractkm at the loag um buu)d« ilie tube, and ititac, hj ni«iiis of u
BWlW MUcbMl to tilt end of ibr long arm, nirty bv dir<>i:tlr iriiicriticil »n ti
NCMliaf MufiiCr, Thin iniitriiRivnl i* vrr^ wall iwh|ic«(l Tor nfa^trliiK chmiffra
It tW t«Ificlt]r of till' flow. In drtcrmininK kctditl vvlocilics, for whicli |>ur)>i(te
h tai to be exfiertnraullr gnuluat«d, it U not k> tueftil.
la tbt capUiarw, llifl mio ia Rtowmt of all. In lh<> w«l> nf ihv frag th«
Im u iudgeil Ur tho movement of the md corptisclfw mny \k diroctljr
■■MfM ua<ler the microeeope br me«ti8 <>r n mk-romvter, anil i» fuuml to
U iboul half ■ millimelrv io a wmad ; but this it pmlitiMy a tour t-slitnnttf,
or* h m ofilr when th« rircnlstion ia somewhnt slow, slovrvr [nrrhiiiw thin
ilal oQffht Ut he cotiaidered the normal rale, that the red coriNifclea cnti Iw
ditiliMljr seeB. In the mummnl th« mle has been (Miniated at ibout <J.7j^
lAliaaMrM * »ecoa'l. but U pmbubly ()Liiclcer than even ihia.
At nnnts tAe rein*, llie flow b verj aIuw in the small veins eiuert;ii)t; from
Aa(H|dTlann, but JncreueiM these join into larger truuk;*, <intil iu « larj^e
WD. rtirh aa ibe iuK>ilar <if the diij;, the rut* is ai>out 200 tnm a »ec()ii<l.
ilSS. It will lie neci). tliMi. that the vcliidtj of the tb>w i.-> in itiTenw
fnfOrtkM) to the width nf the bt-d, Ui the united nectional nrtOM nf the
HML It i> grMlr*t at the aiirta, it diminuht* aloD); the arlerial xyKtcm to
tb» (Kpillarki. to the imiicd baaei of theconw «|H>ktii of in § 112, wliere it
iltwi,BiKl fimm theniv inenttmi again along thv. venotu avRleni.
XaA, EimIhhI. it ix tbi" width i>f the bed, and thi> nl»n«. which tl«tcrmi»es
ibevowrW ve|i)eil_vi>ril)c flow nt variou* [)artii of the ayrtem. The »lownc«
it Ike flaw ill ibe niplliiriea u not due to there Iteiiig »o much more friction
htWir narrow cbanii«U tbnn in the wider mnaU nf the larp-r arierif*.
r« lW [•eripfaeml reeisiance atusctl by tho IriotiuD in the capilinriu> and
•■all arleriea ia «ii obuaele not only to the flow of blo<xl thrciti>;h ibe»e xmail
nawb where the rCMMaoee b actually generated, but also tn tlte escape of
thg bliNxl fVoni the Urge into tlie small arlcried. and indec<l from (be nenrt
iMtlW Iwve arteries. It eserta ita influence alont; the whole arterial iraet.
Aail !i \t i^viouH ilini if it were this peripheral rrsialanoe which cheeked
tbf flini in the cnpillnriM, then could be no recovery of velocity along the
now* tract.
lit lilood ia flowing through a cloaeil aratera of tubea, the blooilreiHeK under
tkr influence of one propelling fi>n-e, the svstole of the ventricle, fbr tliis is
llw li>rfe which dri<rc* the bliHHl from ^-vutriett? to auricle, though, an we have
■■■.iltaetlaDWmialifierl in thi- >eveml parU of Ihesyatem. In xiich a >y>tem
AtaiM <)aantity nf fluid muni gum each aecllon of the «)-*ti<in at the wim«
!(■(, nthrrwiai- ihen- would Iw a block nt one place and a deficiency at
i»<tkrr. If, for in«tanev. a fluid is ma<le to flow by some one Ibroe, premurc^
M fravlir ibrough a tulw A (Fig. W) with nn eHl«r;g«aH-n( B, it i» obrioui
iW the *ame quantity of fluid miL<t pan
<^n^b tbe taction h os \»bk* through the
wii* a in lh« mme tttne— for insinnce. »
wa*4. Othcrviec, if leas panea thrmigh b
tJaa a, tbe fliiid wuuld aceumtilate in B, or
V*M«. B would be emptied. In the same
'< juU ea much lausl y»^ tn the nntc tinte
■ifh tlie aeciioB c u paasea through a or
t". ikit if ju"( ft* many particle* of water
hm U' get through the unTnui teetiim a in the «ame time n* tbey have to
(H ihruugli the br'vi>!er 'eeiiiin v. they mutt move <)nirki-r lhfi<iii;h a than
linM^b c, or more nlowly thmiigh c ihnn thrutigh m. Fur lh» oamr r<-iuion
flawing along a river impelled hy ana fbrc« — vix.. that of gravity —
FU. S1L
904
nisltea ntpidly ihitiuiijh a "nnrrovf" nni] flows sluegUlily wlieii ilic rivor
widens oui iiiio a " bruail." The flow tbrotigb B will be iimilarly 9lack<*»cd
if B, instead of beioft simply a smgh enlarge men I of tlic tulie A, couslstK of
a number of small liibeti hraiioliiu^ out from A, wilti a uuiied wHlioiial nrra
neater Uiau the seclioiwl area of A. fr) i-aeh of auch smmll tulxw. nl tbo
nn« c, for instuiire, the Row will be ^lawer thitu at a, wlierc the *ranll tubci>
branch out from A, or at h, wliere they join RKsin to form n MR^le labo.
Heiicv it is thai iho blood rushes swiflly through Lh<- nrliu-i», tarrm slowljr
through the citgiillarius, but ipiickeiis il* |inco i^aiii in ttie vdi».
An apparent wnlradii'tion to thiH principle t-tint tliv rat^ of flow in depoo-
dont ou the widlli of the bed Ls srcri in the emu wlivrv, the fluid having
allemativu mutra, one of the route* U temporarily widcmxl. Suppose a tuM
A dividing into two brnncbes of equal length x and y which unite again to
form the tube T. Suppose, to stnrt with, that J; and y are nf equal diameter;
then tlio resistance oHi^rcd by each bein^ equal, the tli>w nill be equally
rapid through the two. being juet to rapid that as mucli fluid dbmcs ta a
eiven lime tbrotij^h r and y together a;^ pas8L« thmugh A or tnrough f.
But Don' suppose II to be widened ; the widening will diminish the re«i!taDoe
ofler«d by y. nnd in cotiseqaeooe, supposing that no material chnnge lake*
place in the pra^ui'e or furct> which is driving the fluid along, more fluid nitl
now \Mae alonj; t^ in a Kiven time than did before ; that is to say, the raptdi^
of the flow in ,v *^'>11 h^ incn-ased. It will be increased at the expense of the
flow thruu);h ^, since it will still hold good that the flow through f and y
together is equal to the flow through .4 and through I'. We shall hare
occasion later on to point out that u vmall arlery, or a set of small artertee,
Diay lie more or Inat suddenly widened without materi<llv affecting the
general bhMKl-jjrWMUr^ which is ilriving the blood through the artery or set
of ait«'rii». In such caaea the flow of blooil l.hrongh thi- widt-nvd artery or
arteries is lor the lime bein;^ increased in rapidity, not only in »{utc of, but
actually in consequence of. the artery beinK widened.
[t must be understood in fact that this dependence of lite rapidity of tlie
Sow on the width of the bed applies to the general rate of flow of the whole
circulation, and that, besides the above instance, other special and trmporary
varttilions occur due to particular circumstances. Thus cJianges of pressure
may alter the rnjiidity of flow. The cause of the How through l)i« whole
system is the prmsnrc of the ventrtt^ular systole mauitesied as what we have
Qklled blood- pre»u re. At each point alou^ the system nearer the left
veotxiole, and therefore f\irlbcr from the right auricle, the pressure is greater
than at a point further fVoni the left veutricTe and so nearer the right aurirle ;
it is this aitTcrenoc of prewure which id the real cause of the flow from the
WW p(^nt li> tlic other; and other things being equal the rapidity of the flow
will depend on the umounl of the diiPcrence of pn!*ture. Uence. temporary
or local variatiorm in rapiditv of flow way he cau*cii by the eatai)li»lunent
ftf IciDitorar}- or local diflcrcnccs of prewure. For (example, at any point
«l<mg the arterial itystem the How is incrcjiixil in rajiidity during the tern*
Sorary incri'ase of preeture due to the ventricular ayntole, i. r^ the puUe, and
iminishcd during the sulmcqueiit Icmpnniry dccrcn»i% the increaie and
deorease being the more marked the n<Mrcr the point to the heart. And we
•liaU probnbly meet Inter on with other mstnncefl.
S IS4. Time of Ifie ftitirv Mmiil. It is obvious from ihu for^>ing that a
red corpuscle in j>erforming the whole circuit, in travilling Irom the lel\
venlricle back to the left ventricle, would spend a large portion of it* limo
in the eapillarivs. uiinule arteries, and vetne. The eniirc time taken up in
the whole cin-uil has been approximately attimaled by measuring the time it
takes for an ca»ily recoguiied clieuical substance at^r itijectioD into the
TltK MAIS FACTS OF THE CIRCULATION'.
205
»
I
Jnnkr nin of one aide to «ppesr in lh« blood of ilie jugular v«io of the
olWaiift.
Wlii1«MBklI')iunllti« (if blood an* being ilrawn at fiei)U«Dtl7 r«i)eaf«d iut«rriilB
A«fn thp jufnlar vein of iin^Hiclc. or wlillf Uieblowl from the voin Uboing allovc^l
«0 bll in a uiiiiutr Klrraui on in ahaotbrnt paper cnverinK loatti traTrlUug xurfacc,
tfa Imn nit mrli aa putamiiim rrrrncyiniiJi' (or iinrfcrably loiJiutn frrrric^rnniile ■«
BMtiag nora innoruoiii) b injectPil iiii'o tlie jugular vrin of theotlit'r nide. If Uie
siwfl of tb« Injecltod be Doled, and ibo time nder Ibe Injection into one aide at
«i|iidi vniltaco of the pratVRca of (be iron aalt can bv detected in tho aaniple of
l>too<l fron llic Tpio of the fltber lidr ))« nntod, tbi* Ki*^*" tbr time it baa taken ibe
«aU tm pflrCwm Ibe cin-uit ; and on the «uppwiitiun that mere difTuiiioB doea not
anlwUly aifrct Ibc n>«uli, tbe time nblcb it takca tbc blood lo perform tlte tame
dftaH i« ihenby girvn.
In the hone Hub time baa been experinieDlallr detenoined at about 30
■MMwb moA ia the dog at about IG eecuada. In man it ia ]>robablv from 20
10 35 wteuoda.
TtkloK the mio of flow ilirou(;b tb« capillaries at about I iniu. a second it
•uM tue ■ oorpuiwlc as lou^ a time lo net ihrouifU about 20 miti. of capil-
bm na to perforin (he wbole ciriMiit. Ileo<«, if aiir corpuscle bad iu iu
cinBlt to poaB ibroui^b 10 iiiiu. of rapillariiis. balf ibe abole time of ila
jmef would be spent in the narrow obuiinelR uf the capillarieiL Iniuinticb
aiW iMir]nee0 served by ibe bluod are ebietljr carried uut i» the c-apillariv*,
it ii oWiouslf uf Btlvaiilage tbat iU atair in them should be |>ruliiii^il.
Hon, bnwerer, the avera)|:v leiiEtli of a capilWy is sbout O.S mm., nlK>ut
UfaieKind isspeui in tbe atpillaries of the lIsHuex and another half tecond
is the oapillMries of the lunjpt.
iUtSi We BIST now briefly fuiuniariM' tlio broud f«iturt» of the dreulH'
oco, vhicli Kc bavr wen may be explained on purely ph)>i«it nrindples, it
biWuKiinex) Ihat the ventricle delivers a eerlnin quanlUy of blond nilh a
(ffUui force int'> ih* aorta al regular intervals, and tliat the physical pro|>-
ettki of th« btoodvcen-ls remain the eame.
Ws bare seen that owing to tho poriphervl rvcislanee ofl'ered by the cap)''
IniaaadKnall vnseI»lhedi>Fvteflectof ibe ventricular stroke is to establisb
to tlie artAries a mean arterial premiire which is greatest at the root of tbe
aorta and diminishes lowsrd the small arieries, some of it bcinj* useil up to
4rffs ibe blood from the aorta to the small arteries, but which retains at tbe
n|>M of the small arteria euffldeiit power lo drive through tbe small
sttHin.capillanci'. and reins jusi as much blood as is beioE thrown into the
aula by the ventrirular strukt-. We have seen, further, that iu the large
ift«i« at eaeb stroke ibe proauro lUiti and fnlla a little above and belov
tisHMan, tbu-- omstiluiitig tbe puW, but ihai thti vxm dbtension with tla
•okuteat recoil dtminisbca along tlie arterial ltbci and liuallr vaimlua ; it
diaiaUies and vaulBhes beoiuise it loo, like the wliole force uf tlie ventricular
Hnlw, of a fraction of which it i* the cxprcMtioti, in i»ed up in eatabliohing
tka nam pranure: we «hall, however, coiinder ngnin later on the «jiecial
falaras of ihU pulse. We havi; sura, further, ibnt tbe laak of driving tbe
Uwd through thi* ]H'ripl>i.-nil rmslnnee of tbe sniall arlerii.ii and canilTaries
nOMiiBea much of this im-an pmuiure, which ooiuequently t* much \v» in
■Wsmtll teioH than in ibo en rrc« ponding rmall arterii.v, but ihat sullicienl
HBahv l» drive the blinx), even without tl>e help of tbe auxiliary agenU
*Ueb arv generally in action, from the small vetos right back to the auricle.
Ui^^ wtliav* seen Ibnt while tbe aliovo ia ihe cause of the Bow from
vnlnde to auricle, lh>- cban^iing rale of the How, the diuiininbin^ nwiftncsa
ia ths arterini, the sluf^fpoh crawl through the ciipillarice, Ibe incrcnung
20ii
THE VASCl'I.AR M BCIl AN'liiM ,
qoicknesB through the veins are determined by ibe cliUigtDg wliJlh ot tbe
Tuoultr " bed,"
Befiire vre proceed to cunsider any ftirllier dnnilH lu lo ih"- |iht>iii)TOi.*iw iif
the Boir tlirbUjih the veaieU, we aiuu turu iiiiulo tu fttutly tliv hi«rt.
TlIK HiLAKT.
^ 126. 'I'lie heart » a vitlvulnr piiiii|> which works on inecbsiiiciil ]trincU
)>te:d. bill ihe motive (wwcr of which is supplied by tbe contraetiuu of it*
niuaouhir 6brci. Its Bt-lioii conso<]uoiilly presents probleiuii which ate piinly
Riecbaiii<nl and partly vital. lie>;arded lut a pump, its eflects are determiii«il
by Ihe fre'iueiiry ul' the beats, by tite force of each bent, liy the vluiracUr of
oavh beat — m bether, lor ioatance. slow aiui liiiiti-rinx. <>r Minldeii and Mhar|i —
and by the quantity of fluid ejected at each beat. Ilen{«, with a given
frttqucuey, force, ana character uf lieal. and a given quantity i^bctMl at vach
h<'tii, the pi-obleiiiB which have to be deiili tviih uiv for the miitit inrt
iticchaiiictil. Tlie viinl prubleiuB are I'hictty coiinecie<l ivilh tlic c-humv which
(letcnuiiie the frci|ueucy, fiirce, nnd cliaruut^r of the bcMit. The qiinniity
eivctml III wich liuil i* govrrned iiiore by the iitnto of tins r«l of the Iwdy
loan hv ihiit of the heart Ilnelf.
The Phnwmena of the Normal Beat
The vuibk movenimts. When tbe chest of n iiianinml is opened anil arti*
ficbil reapimtiiiii kept up tbe heart may he watetied beatinff- Owinjc to tb«
ivinoval of the ctiait-wall, what ia aeeix is not abwlulety identind with what
tak«« phtcc within tbe intact chest, but tlie main eventa are tlie lamc in both
ewes. A complete beat of the whole heart or catdiac cycle, may be obwrv«d
to take place a» t'ollnw^ :
The grvot vetni>, inferior and superior renn cavic, and pidmouary reina
are seen, while full of blood, to contract in the nuighburhood of the bean ;
the contrnctioii runs in a peristaltic wave toward tbe auricles, increasing in
inieiiHity as it j^oes. Arrived st the auricles, which nre then full of blood,
the wave suddenly spreado, at a nitc totj rapid to be fairly jud>,'ed by the eye,
over tbe whole of those organs, which accordingly contract with a sudden
(harp fytitole. In the systole, the walls of tbe auricles press toward the
auriciilo-ventricular orifices, and the auricular appendn^'cs arc drawn inward,
Wcuniiug timallcr and paler. I>urinK the auHoulitr systole, tlie ventrioloi
may be iteen to become turgid. Then follows, as it were imrueUiately, the
the v<'ntncuhir «ystule, during which the ventricles become more conical.
Held Wlwi-en the fingerii tbey are felt to become tense and hard. A» On
Kitolc progn«M>i, the aorta aii<l pulmonary arteries expand and elongnio,
e apex u tiltcid tilighlly upwanl, and llie heart Iwidta Homewhal on ita long
azii, moving frnm tne \t(t and behind toward the front and right »o that
more of the left, ventricle beoonie* da^layetl. Ait the MVNtolo given way to
Ibe •uoceeding ilitwtolc. thp vpntricl™ n-sunie iJitir firevniu!t form and pOM-
tion, the norla and pulmomiry nrirry shrink aud MlK>rli^n, the heart tnmi
back toward the left, and thnu the cvcln is I'^iniplrifW.
Ill tbe normal l>cat, the two venincleM are |H'rlci-lly nvnobronout in action,
they contract at the same time and relax at ili" :<nmc lime, and the two
Uindee are similarly synchronous in action. It has Wm maintttim-d, how-
•rer, that the synchranism may at times tint be [lerlect.
Beibre we atlempt to study in detail the several [wirts of thia complicated
•orica of evenia.il will be convenient Intake a rapid survey of what is taking
plao« within the heart during such a cycle.
THK IIKART.
507
I UT. Thf oirtiioe rgtlf, IVe tany lake aa ilie end of ifa« cycle the moment
HvWb the vcolrkW liitvint: mptipil th«ir (.■onteois have relaxed and
nCnnriJ (•> tbc diaslolic or rt^lin^' [vi«jiiiiii nnd form. At ihiit inoni^nt the
kbtd k flowing frc«h- with n fair rapidity, but h« we have aeeu at a very low
pmtn, ihrodgh the venv l-svh- into (he riKlit auricle < we inuy I'uiitiiMi
Mfflnt at firat lo (he ri>,'ht side), am) ^iiir« ihtftv a now ocithing lo kmp
lt*liwiM|nd valve shut, ^iii>e uf Ihis bliiod j)r<>biil>lv KmU ita nay into itu)
nnliM* abo. Thia guea on for aome Hltle lime, aaJ iheu comt* ihr nharp,
Awt n«toI« of lh« auricle, whJeh, since U heuiioii ai> wo have mmmi iw> a wave
if cninctioD ninniiit; forwarxl ulimg ilic emb «r the row cnvio, driviw Um)
Uoij net baekwanl iu(i> iho vciii.-> Imi Kinvanl iiidi (he vfiotriclti: thifl end
b fclthar aeeurvd by thf liivt thai (h« ayttolv bti* bchiml it un (hi- veniiua
MlJw fmwurv ul' ihc blixid in (be rcin», incrtuuiiiit: an no have Eevo back-
Mnl Inward tlie cafiilliirU^, aiid before i( tbt^ ivlatively empty cavitv of lb«
muide in wbich the ]>rw»iiire ui at fintt very low. By the eomplotc oon-
tflKtMO of the auricular wall* (li<- oi<mplclo or nearly complete emptying of
thttaTft* i* imturvd. Nn valvi-fl an- prtvcnt io the mouth of the eutterior
*nacava, for (he^ arc not ntx-dtd ; and the imperfect Eustachian valve at
ifenovtli of th*- inferior vena cava cannot bo of any gmt use in the adult,
'i in its BU'Tv <lt-vclo|x<(l mate in the fo-tiis it hail an important funeti'iD
' the blood of the inferior vena cava thrmi^h (lie fommen ovale
; anriclfk Tbe valvee in tbc '-orouary vein ari-, howercr, probably
' DM in pT\-<rcn(int; n rertns in(o that vessel.
A* llie blooil \a being driven by the .-itiriciilar systole Into the rentiicle, a
irflut current ia protmblv tel up, by which the blood, passing along the
I of the ventricle, |>elB net ween iheiu and the flap« of tbc triiiiAjml valve
B iMtda to flo»t these up, IFigt. t>9, TO.j It is further probable (hat tbfi
' ntax current, continuing komewhat later than (be niin inii> (ho vvn-
', h tufficient (II hrinz the flaiiH into ap|witiilion, without any regurgitation
inu likC snricle, at the eroac <>f ihc auricuUir avfiole, before the ventricular
rnawi
(riclr,
[no-O*.
7io. 70.
PuiiatM w ViLTD -ir illl (lutil. AHk Itiunh
mute haa beyiiit. According to wme authoiv, however, the cloeure of (he
nln m vflected, at the very beg^nntne of (be vcn(ricular svHole, bv the
Matrmctiua of the papillary mn«cle«; the churrUe tcndiue» of a papillary
cU are attached tu the adjnoiTni edge* of two Hajs. »i> that (lie shortening
^ • moacle (end« tu bring lh<M edgoa iolu apposition.
208
TUK VASCCLAR M ICCEt ANiaU.
The mirit^iilnr avHtolc is n" we Imre Htii] immcdiKtvly folio'
t]ie ventricle. Whether th« contrnclinti of the vmtrii-ular vniiie (which ks
we fhitll a'« ia n «iinnl« though prulungv.! contraction antl not a tctunus)
begin? Kt one point una swiHtlr tnivels over Ibo rest of the Hbres, or )>eeiiia
alfovcr the ventricle ut unce, ie n >(neeti()n not at present definitely #eUled;
but ill any cose the wrnllB exerl on the contentM it preesure which in aoon
brought to bear on the whole conlcnte uid very rapidly rises to a maxitouio.
ITie only effeel of tliis iiicrenaiug iiilru-ventriculiir pmcure llpoD the vnlve
ia to render the valve more and ruore tense, and in consequence more tecure,
llie cbordot tendiaie (llie slackeoinj; of which through the change of form of
the ventricle is pmbably obviated by a regulative coiitraeiion >if the impillar^
ntuscles) at the eam« time preventing the valve from beinj^ inverted or cv«a
bulging largely into the auricle, and indeed, accur<ling to onuie uhoerven,
kiKiping the ralvular aheet actually convex to the ventricular cavity, br I
which tuean» ibi; complete emptying of the ventricle is inore fullr eJlecWo. <
[Fipi. tJO, 70-1 Thtt connection, to which we have juM rderreil, of the chordte |
of tlie Mimv pnpillan* inuxcle with the ndjaoent etjgee of two tia\», ntiHi iiMttU i
in kvi;piu^ the HiipK in more complete appiiiitiou. Moreover ihii extreme
bordcn^ of the vaiv<9, oul*ide ihc attach men Li of the chordiK, are oxomnvtly 1
thin, so that when the valve i:< closed, ihete thin ;iiirti<iuii an? prcn>cd flat
together bsek to buck ; bencc, while the tougher wnlral piirl» ol the vslve»
bear the force of the ventricular tynUih, the iip|>ntcd thin memhranou*
edge*. pmMul togvahcr hy the blood, niort- cuiupletcly Mccuru the cloewre of
the orilice.
At the cimi men cement of the vontricular syHlolo the semilnDar vslves of i
the nulmonarv artery arc ckifcil, and nnr kept elwcd by the high prcsnire ot '
the nlood in the artery. Ae, however, the ventricle continue* to pre» with
greater and greater force on it« contents, making the ventricle bard ai>d
teoBe to the touch, the pressure within the ventricle becomes greater than
that in the pulmonary artery and thin greater preeeure forces open the wmi-
lunar valves and allows the esca[>e of the contents into the artery. The
ventricular arstole nuiy he seen and felt in the expiiseil heart to he of wmie |
dnraliou ; it i» xtning enough and long enough to empty the ventricle com*
]>lelcly ; indeed, n* wc ehslTsee, it pniliably lusts hinger than the diseliargtti
of blood. Ml that lliere is a brief period <)uring which tlie ventricle is riaptr:
but yet contracted.
I>iiring the ventricular systole the .wmiliinar valviw are prtascd outwnni
towan) but not close to the arterial walln, reflux enrn'nl» protmbly keeping
tbeiD in an intermediate position, so tbiiC their orifice fornui an e<()iii lateral
triangle with curved sides ; they thus offer litlh- ohi>tneIe to the escape uf
blooa from the cavity of the ventricle. The ventricle n» we have seen pro-,
pels the blood with great furce and rapidity into the pulmonary artery, and!
the whole contents are i^|>eeilily t^oc1«d. How, when a force which is driving
a fluid with great rnpidiiy along ft closed channel suddenly ceases to act. the,
fluid, by its momentum, conlinuee to move onward aAer the force has ceased ;:
in coDEequence of this « negative pressure makes its sppeanncc in the rear:
of the fluid, uid. sucking the fluid back again, seU up a reflux current. Sol
when the last portions of blood leave the ventricle a negative pressure makes'
its apMArutce behind them, and leads to a reflux current fnim the ar1«ry
toward the ventricle. This alone would be suflieient to bring the valreii
Kweiber ; and, in the opinion of some, is tlie real cause of the closure of tht,
vdv4« ; others, however, as we idiall nee later on, mainlniu that su1)se<p)eut lo'
tbi* reflux due lo mere negative preaaure a somewhat later reflux, in which
the elastic reaction of the arli-rmi walU in cuiiri^rncd. mure cumplet<'ly (ills
and rvndent tense llie pocketx, cauniug their free margins to con>e into close
^
THE HRAHT.
20B
od Irai omUuit. «oi) thus eoliv«ly blocks tlte my. Tim corpora Arnotii
■■I ii iIm etntn. mkI th« tliiu ineuibniiiouii teabiona or luuulw an bn>ii)^t
■Wtim amavition. As in tbe tni^ua]>iil valv(«.aa here, while the iiittHura
4f llvbhioa ■ bont* bj ilie tAu^bcr bu'lies of i)ie iwreral vfth-M, eii«h two
duD idjMHii luniilw, prettied to;^tber br tliv bliiol aL-iing no both nidca of
lbaa.Bl* kcM in ci>i»|ili!lf cuiiloct, n'tlbotil nnr Mraiii being put ujioa thuin ;
iitUi wftf u* orifice ia cbdol iti » iiuutt eflincni itinDnt-r.
nn»l«Baai4«|iwtef6and«tion rorthcmw put hnrnt'l by BrQckv Ibitt durinc
ilttHlrioiUf (TKUile the flaps are prene<t bjuk ll*t nxainst tlir arliTii! witlt!>. uiicf
li itt ou« of tM Kuri* coinpl«teljr cover tip lfa« orifiMi* of the corutiurj- Hrirries,
■ ikti Uw flow «f blood from tbc aorta ioto ihe ooronarf arlerioa can ukc plaL-e
«lf darinc lb« vanlricnlar clia*lo)e or at tfa* "nrf beginning of th« ayatolv. aoil
Malal)<larlngihf «7«tole ila«lf.
Tbt vootriciiiar ■r«to]« now ywM olT. tbe mitMulKr wnll« reliix, ibo
taDUide nrtitnu to it8 pnvioas form Ktid positiun, aiid tiit? cyd« is onoe
anudwi
What tliui Uiko pUee in th« rivht tidti tnkr» ninety in the teft side also,
tWi it ihe Hnic Kiddon »bnrp uiriouUr ayHtfilc Wginnin^; »t the roi>l« of
the pilmuoary vcin». lb<' kkhp «y>Uile of tho venlricli?. but, n« n'e ithni] see,
"M murh Hi'irc |H>wiirfiil wild I'xvrting niiurh rioi'p fi>rce ; tho mitral Talv«
w'tik iu iwu fla^ig n4-la exactl;- like tlii! trictispid valve, and tbe aotioD of the
wilnnar valvm of ibc aorta simply rc|Mtata that of the v«jv« of the
pafanaarjr arter?.
Wc tmT now proceed to sludv some of the oardia« eventa in deuil.
fW. IV rJtaivfr of fartH. 'I'lie exftct detemiinulion of the chan^ in
Un nd poailioii of tbe heart, especially of the veiitriclcs, during n wdiao
tjek b au«ud«() with difficullios-
7W vmtrielea. f<>r ini>lanov. are c'>ntinuallv cbaoging their form ; they
(faug* wbik llwir cavitico are l>eiii)i tilleil I'rom thu auricles, lliev chan^
nVk i1m oiaifmctioD of their wiilU i-* iMiinjc up the prwaure on dicir con.
Kan, Umj ehaiw* while under ihe iiil1ii(-n<<e of that pnwure tbeJr oont4«ta
bitag dbdiargM imo tbo imeritv. lunl they cbiui)^ wlico, their cBviliM
hanig bosn enipti«d, tbi-ir mii«.-nliir tnitl» relax.
. Wa may tnk*- it f»r icninteil itiiU the intenml ejivitics are oblitcraied by
tb( qntou. for it a probable Ibat pmciir-iilly tlte nhotc oonlcou are driven
•01 tt cacb struko, and probulily ido each citvity is emptied from ii» apox
maanl iha mouth of lb« nrli^ry.
VTitb ragard to cbtiiijji-s in oxterniil form, there sMou do doubt that the
wle-ton'te iliiuiMiter is much leaseawd. It Mti-ms also clear that the fro»l-to-
kack dtanteter b grciilrr diiHttg lh« nhole time nl' the »y»tole than during
thciiaKule, the incrcow Lakin;; |>Iii(M! during tlw finl part of tbe sy»to1c. u
i%bl lever be {ttaccl on (tit- sorlnce of the heart of a mammal. tl)L> chesl
hinsg baao opened and arlilicial respiration beinj; kept un, some such curve
« thu raptCMnied in Fig. 7 1 is obtained. 'Die rise of tho lever in dcMribing
mA a curve is duo to the elevation of iho part of tbe fr-int turfac? of lli«
heart on whicli Ihe lover ia resting, ^^uch an clevnti'Xi iui)<ht be caused,
«ipMially if the lever avre placed near the apes, by Ihe heiirl being " tilted "
' ntii the ayatule. but oiily a siimll portion at most of ibe rise can
. Ill tbia cause : the rise is perhaps best aoen when tbe lever ia
. ui lUc middle (loriiiii) uf the ventricle, and inusl be chieHy due to an
I ia thr fr'ini-li>-Uick diumeler of tbe ventricle during llie heal, Wu
dwniaa ihit curve later on iu oonuevtiou with other eurvee and may
•|tly aty that the iiurt uf the curve IWtm b' to </ probably oorresmn^
lathe actoai ajntole of the ventricle, Ihal ia to the lioie during which the
II
310
THE VASCtTLAR UECHANISU.
Gbr«a of ilie ventrit^le are iiDd«rKoiu|t contriictitm, the suilden full from <£
(iiiH'iiH r«]>reKDtiug the relaxntion whii-li i'onm tlic Orel purt of lli« <)i»>tale.
If (hU iuterpretulion of ihe curve fie eurreet, it U ukviouii llinL the froui'to-
bitck diameter ia K'^»'>r duniifr the whule of the nvslitle ilmii ti 1* during ,
diuatole, siiwe tlie lever t.i raueil U]> all thin time.
nn.7L'
THACIMH nMI IIUST or I'AV, OUTjIIXKb (V rL.lt'U<i> A iJullT LtlKn UX TUB VCNTVIULI, tH*
Cnon iuvi9(o Bnn nvciiMt. Tar Ivsisn-rciiLK cuuve hjhui M VmiuTiixai r*a Sannnv
Tilis increase of (he froDC-to-bacIt dinmeter oonibinc<l uith a decrease of
the »i'k--to-eide diameter has fur u rotult a chaoge in the form of the Mclioa
of the bnso of the ventricles. I>uriiiK the diajitulc thin hiis somewhat ibe
form of ail ellipse with the lon^; axis from side to ude. but with the fr<><it
part of the ellipse much more tunvex ihaii the bock, wnoe the back siirfsc*
of tbe veDtriclee is soiueMlmt llatiencd. DuriuK ihosyMnle Ibis elli|M>e is by
the short«Diiig of the siik-lo aiile diaint-ter luji) the increiue of the fniiiMo-
back diameter cimverUil iiilo a ti).'itrc: much mure Dearly ri'aeuibliiix a cirrJe.
It ii> iir^tM), moreover, ihat the uhole uf the bii-ii: i> const rii-lol. and that lite
greater dficiency "('the iiurioiihi-vrutriridar x'ulven U ihurtrliy secured.
An Ut the bchaviur of the long diameler frum ban; Ui afiex «bgH.-rv<:ri arc
not acr(«d. Some maitilaiii thai it in Khurleiieil, n:ut tithere that it u prac-
tically iinchaDgvil. If iiiiy ■>hi'rlcnitig dm* lake plarv, it iiiuttt hv largely
conipennuted by the clongatioii of tbe gnmt vvmcN, which, lu iilalcd above,
may be MtMi in aii iQ»|>ectioii of tbe healing hcnrt. For (here i» evidfocr
that the apex, though ns we have neon it i» during the aysttdo «omcwba(
twisted round aiid at the same tiiDo brought closer to the chMt-wall, does
I "Rw Tenia*) oi tnilua ennai llnH (icxincrii of elrr In} inlrudUMO inio ilili niul many ulWr
eUTTWUVOf <u* fat tbe burpotD-f uiLiuurlDx luiuor Ihr nnt\-r. A mii>|il<*U' I'linu iliuuld eiblMI
sn "abvelMS" Una. TliU hiht Ut iln^n Ity jiilnwitiii ih^ U-vt^r^ nmuatfnl t'tt E.h<^ nivrliiivitl bgl
icmalntiWBl mt. toiurk ullli 1k |<:<1ii[ <m ihv monlliix •urfaco tti Id wison : n atnttalil line, tlip
■liiiiiwi UEic, I# ibiu 4c«ct11jh1. anil nia) betlmwn tvSipear aAcr Oit^ oiirvu tiHiir H inue. aurl nuj
b* pUcAd BU-n ur tn*a>rKtily brlrjw Uicvunv. Wluin a lunlnK-6>[lt or ulhi-r llinv-iuaikDt li> luol
tbvUni^nr (tif ij]-nk"tTijir^k.r .-.r n i;nriln«^^i in*-"!!;-!! t>»< c>nnw if iIjd liiiilriK-'irk will «4vitf m 40
StaclWB titiv. Af: liiu bn-[i iriro <rilltiE •iifftti?!.- iliinilil rt- hrdiisht t<*(\ imncli
SfOAlUun ihuT i^j. i< luvc: ix^m <in<: iTiIni ■>f ihv ptiitv ulik'b It liclHlrcd bi
aiuti if tlicluvv-.- . . ijilv mnrnl n' ■ iIl. ,..i r ..( rliu 1m tt will (l»<-rib« • wcnuail
nf ■ olrriv lUir ri^Mirr til itliijila Itr* mi t^' '<i H^tffiuiii •ifniM Tiv nudv louf
E«ioai[h to cm luili IIil- ciitv* rniil ilic >ii< ' <it: cimy nr nitici Uni^iniirklni
Ifakei Wberv UilaUiIruwii- H}' itidvIi^ iTir^ ' ' ^ l[>I iiTt<l forfaiil ^linlbit vv^mmli
oT rlrclw nujr t» liruwu lti(T>u«li iiiluT .• tin' liii» <i. 0. / lu Kie ;i won Ibn)
4nwu. Tht dittJ>n« tM.<twrrii inv tv>< '' vhu- l»i rjuiitunT) uri Uw tunlrii^-lhTk
0SI1C or DtbDr Uiiiocurrc. oroQ lli« nlivcb^i - n,.! ,r irio* ni'tv t>,- ilmwia oi> lUn inicliitf ancf
lumuuval ttoiD Ibu ntunlliit; liuimnuiut in liiu loUouii^i: *aj i TuKi' i> inirof «>ii>[iaB«. incliiu
pdntt «r wlik'h an lixnl iiui m far ifian M Dii' IitiuiIi ■.•! [)iu li>vi*r uvl In ihu e>|«r4nirni.
iMaNiPod ttnnt Ua axil tc> lla uruinf i">JLii l^i iikrj^n- •<< \Ui- ruEiiiiimMi^ eI^ipI Uic imviliinn mi tbv
mttncof ilicMnirvof tin: cln-lcnf ubn-ii r itir |>r>.'vi<iii-lF ilniMti <'i]rtnl liniH Umnt a
•KKiMiK- Thniucli it^ orati.iT Omv a Jr ' < iTk- hiivlkvA. [Iy Vorg^rte imp |kj4nt nf tbe
eoDpaM oil I 111* line iiui riii.iJnu ii niium : i -. ' ■ iiiM or CKHnnI a •ci;iiiv<t( '>( a ffrtl* ina;i b*
ontwn ■!> K* I'l r<ii piriv |."ii' 'i ' ii' -nrrv itu: ;^;4lj Imi Jnlr>«l. ikTiilnl«'i ibi' nli«4<k>«4i hfiponhM oiiir
kloe. Siicli aacciDijni oj ii> iir luwl for Uie tmmc git^ri"^'*'^ "^ 'l><^ nrt^liinl \ttiv atifl may
nnnibrror nui^li H^rnnr'i- "I'll '• 'I'lmo
TUB IIKAKT.
211
bM doD^ iti [HMition up or dowo — J. «., iu tJie lotifi iixU oi tlto body. If
iairabliii or iIi>k u needle be tbiuHt tfanmicb ibc fhtHi-utill m Ibai ita iioiiit
fliiDp* i[il)> ibe n|>ex of tlie heart, ihouxb tbo uevdio (piiviini, itM IkwI
MW neitber up our down, fts il would do if iu poiitl in tbu apvx moved
d«*n M ti|i.
Bnadly ape«kiiig, (ben, dtiriuE sj'sUtle the v<uilHcl*x> uiidvnjn n dlmiDiilioQ
tt bul vulufuD, njiijil Ut ibe voltime uf cxkiIviiIb diM'bargvd Into Uh) grvat
imit{tor tb' wmllv tlwnuvlvts, like nil tiiU!«uUir ittruclum, niUiii Uieir
nliS* dariaK contractiuu, **ve for i'linngni wliicli ninv take pliicc in the
qoutitf of blood cKintiiiDrd iii ibvir bUH>ilv»M'!i>. nr of lyni|>h in cbt.i iiiler-
■nmkr t|wot«), vrbilc tiwy iind«rgn a chunsc nf form whiih mov be
ilMrilMd a» that from a ronghly bemisphcricariigiira wilh un irrcgularly
tUlMiad irction t» n iiii>r>' rvtruliir com- with a circular biiw.
fill. I'-inUar imi>ul*r. If (Ih- haiiil Id- plitircd oti ih? cbcat, R shock or
llfalM will be fell Ki rncb iM.'at. mid on I'xnmiimiion ihis iiDpulse, " CArdiao
bfviK" <ii)l be limnd l» Ih< >yiii-lironoii8 with thi- s)-«lolf of tbo vontricle.
In mm, the cardiac inip4iUp niciy \vt must dislinclty fell in the fillfa cuetid
intfipac*, alMUt an inch lioluw und a tittle to tli« meditiii «ide of the lell
■fftfc Id an animal (he wime impulte iiiny also be felt iu ttnoUier way —
rit, by makiui; nn incision throuch (he diuiilimgin from the abduinen, aod
phnntC (be fin^vr between the cheM-wall and the apex. It ibeu can be di»-
(intllT rccuentied a» the retiull of (he hardeiiin); of the ventricle during the
■iNiH. Atid the impulse which is feU on the outside i>f the cheM it chielly
lM«Act of the same hardeuini^ uf the itatiunury |>"r(iou of (he vetitricio In
amaa with tlie cb(«t wall, tmuimitted thruuith the che«t-wull to (he Rugvi,
la Mm dacfid elate, during dia!>lole, the apex is (iu a atmidin;; )i<i»iti»n. nl
karti ■( (bin )Hiiiil iti contact with the cheit-wtti], lyin^ Wini-en it and the
MhaUy nviMaiit (ttaphrazin. During the »y>t<>lc, wliilc being briiiight
cMwr to the chnt-aairby (be lihiug of iW ventricle and by the move-
Ifl ihr frrnit iwd to the right, of which wc hnvc nlrcadv tipoken. it
ly gmwB Iciiw and hanl. The vcntriclw. in executing ificir «yttale,
l/i cuntnict agaiiiKt n-«i«tnncc. They have to pru(lu«^ within (heir
anvm pretMirrs greater than those in the noria and pulniuiiiiry arteries,
(■yoctifclr. Thin ii. iit fact, tltc object of the •yctolf; Hence, during the
•idl avtfote, (be ventricular portion of the hmrt beoomca suddenly (eme,
acamnat id Iheaame way hs a bladder full of fluid would hcconie (eose and
hwd aban fbrciblr Mguee/^x). The sudden pressure eser(ed by (he ventricle
tW twwiiata suadenly tense and hard, aided by the closer coo(ac( of (be
Ti "tlh tbe ebea^waII Iwhich, however, by {(self withou( the hardening
tonuacliuu wouUl be lOdutficieDt to produce tbe efl^t), gives au itupuM
ar ihM-lt Intb to the cbcet-nall ajid to the diaphragm, which may he felt
rmlily both uu (he cheat- wall, and also through the diaifhragiu when tliv
aUomni i> o|icued and the tini^er iuicrted. If the miidilk-ation of (bo
tfj^^uaiiti Bph (of w hieb au nhull s|)eak in dttaliug. later on, with the i>uU«),
ba cardiograph. \k placed on (be spot where tbe impulse it felt u>net
'. Um \t\KT t* N-trn to be raiwd during tlie HVMute of Ibe ventrtdt^,
All again a.-< the syttule |ia.-uie9 away, very much aa if it were placed
lh« heart dirvdly. A Inicing may thun be obtuiiM^l laee Fi^, 77), uf
ir« aball have to ^-ak more I'ully immediately (itee ^ W.\\. If tbe
of the lever be placed. ni>t un (hv exact •«>( of (he imimlse, hut ut a
iHancv from it, (he lever will be >/epreue>' during tbe iyatole. While
*pnl of ini|iulH- ilM'lf the roniact of the ventricle U incraaacd duriug
, away fri'in the «(>»( the vi'niricle rctirci from the uhe«l-wull (hy th-
ilJoD of ita rigbl-to-leli dinnicter), and hence, by the mettinMiuul
tttackainilt of (be pericardium, dmws the che*t-wall alUr it.
»n t-
S13
TnS VA3CVI.AB MKcnANISM
S 130. 'Hie *OHnd* of the hmrt. \Vli«n the ear u applied La llie vIicbI.
eiilior cHrectly or hy lueaiis of a, Htetliwcope, two •ounds are Wari), the ftnt
a cttinfiuraiively loiit;, dull, boomiug snund. ihe second a short, atiarp. *uc)(l«a
one. Between i\\v liral and eecDud ooundu the interval of l!nie i« wry «liort —
loo vhort ti> be uiensui-able— Itiit between the fUMond nud the nco«>'<lin^' tint
pound there i^; a dtstima piiiise. Tlic ^uiidH linv<- lit.-«o liknxsl |« tin* pro-
niinvifltioTi nf llie sylUlilcs lulili d(l|i, ah thtit the inrdbr cycle, lu far a* the
«i>iind« nn'. ooncenit-il, iiui^ht Ih* n-piisteDt^cl by: IQbh. iViy, piiiiov.
Tiir tffouil mund. whlc^h U cbiiri niid ithttrp, prt^cnCii iii> didipulltts. It is
oiinclilciil ill jiriini of liino wiib the cli»iiiv nf (he wmiliiQur riilve*. and it
h««nl t<i th(- iKiit lulvHiitn^t uvi^r ihu a^ciiiid righl ooEal carlilafru cl<we to il«
junction with ihe ftcrniiRi — i. r., at tba imiiit where the imrtic arch niiii«C
n(^»ri?«r to Ihr ■iirra(«, and to which snimd* ^nvrHt4.t) at th« aoriic orifice
iTDuld be bent cDndiiclvd. Itii oharuntcn ar« such u WDiitd belong to a
sound genemtod by memhratie* lik« the semilunar vakte bein>; suddenly
made tense, and »o thrown ittl» vibrations. It is obscured and altered, or
replaced by " a murmur." irhcii the semilunar ralvcs arc allecled by diaeaw;
and may ix nrlilicinlly obliterate!, n murmur taking its place, by pasaing a
wire down thv arteries and hoolcint; up the auitic valves. There cau be no
doubt, ill fart, that the second sound is due li> the semilunar valves being
thrown into vibrations at their sudden closure. The sound heard at the
■econd right costal caititage is chiefly that f^norated by the aortic valve*,
and murmurs or other alterations in the sound cauited by chaii^is in the
aortic valves are beard nioit clearly at this bjmH. But even here ihv sound
ia nut excluMvely of aoriic ort).;iii, ii>r in certain cases in which the semibinar
valvm on the two sidea of the heart are not wholly syuchronuuii in aiuion tbe
»>iund heard here i^ double i " twlu plicated second sound "), one hdiig due to
the aorta, and miv to iht^ pulmonary artery. Wlien lh« sound is listened to
DD the lel\ «i<lc of the; sternum at the 3>nme Invel, the pulmonary artcrr is sup-
|ioecd to have thccbiff sharr In produciii); what is hejird.and changes in tbe
Bound beard more clearly hem than on the right side are talten as indicati mm
of mischief in thu milmomiry valves.
The jir*l Konnri. longtT, duller, and of a more " b'wming " character than
the oecond, hcartl with grenteKt di'tinctneM nt lh« s)>ot where the cardiac
impulse is felt, nnwenix many diflicuhin' in the way of a complete explana-
tion. It is hcanl diKtinctlr when the ehi'^l-wallo are removed. The cardiac
itnpulse, therefore, can have little or nothing to do with it. In piiint of lime
It 19 coincident with the >^ystole nf the ventrioUv. and may be heard to the
greatest advantage al the spot of the cardiac impulse — thai ih to say, at tbe
plac« where the ventricles come nearest to the snrlace, and to which si>uuds
generate<l in the ventricle would be best cnuduetHl,
It ii more closely etjincidcnl with the cloxuiv and eonseipient vibrations of
the auriculo'Ventriculiir valves than with the entire systole; for, on the one
hand, it dies away beliirelbeseooud sound begins, whereas, as we shall see, the
acttinl st-stole lasts nit to, if not beyond, the etosure of the semilunar valvct;
aihl.oniheotherhaiid, the Hurioulo-ventricular valve ceases to beteiucand to
vjbrnir tm soon na the contonia of the ventricle are driven out. Thin aue^^tf
that the sound b caused by the sudden tension of the auriculo-renlncular
valve*, and thi* riew U !tup|M)rted by the facts that the sound i» obscured,
altered, or n-plBir«<l by ninruiiim when tbe trimiitpid or miira) valre» are diB>
eR(e<l, and (hat the r/iund is also altered, or, according tu some ot>M*rven,
wholly ilonc away wiih, when blood i* [wvenied I'roiu entering the vcn*
tridee by ligature of the vatiir cava). On the other baml, tlu; sound has Dol
thai sharp chamclcr which iiue would fxjiis:! in a »)und geiu'ratnl br tbe
Ttbration of membranes xiicli us the valves in <iiicMiuo, but in its booming
Tng HEART.
SIS
I
I
<nMUiua ratb«r itti|ci;rau a luuflculHr souimI. Further, acconliDK to some
(mMrvrrv. the Buiitxl, iboucb ■i>iii«wbiit motlifici). mny Blil) l>o hcanl wheu
ibe Wkc fcins mn (-btiti|KHl an tluil lui IiIchhI <riiti-r> itir v«niric)i*. iiml. imlM'il,
nor b*! rrcuciiiM^l in the fi'w bviil* aivtm by n iiiiiniiiiMlian vcnlrii'lii nipiily
rut <iut nf thf living IhmIv 1>r an loraiJoD ciirrivl W- 1 on' tin; miri<-iil<> ven-
(rirular rin^. Hcni^ tU- view liiu Itcon ndoplctl that tlii* lirvt miiiMl it n
ntnculRr »i>uii<l. Iti (ljfcii8>iii^' itif ni(MCuUrw>itnd »f aki-l'-lnl musclo (wo
illOX Wtfwto mi»i>n» l» •li'tniM tlif view ihal I his foiinH nas gwertWd by
Um rvpoUcd indivi'luitl »iiiiiilp coDtrnctioix nhidi mniif^ iii> ihfl tctaana, Uiil
Imbcb cormpondm) in lotw to llie nunilx'r tit' lltiiai.- aimjiV ra>t)tnctioii8 rt-
pMitnl io A woiind, and to ndopt the view thtii ihv s'tiind nna renlty due to a
n-fvlition of ui)i.-iti>i>l IMMiong oi'ciirniiK in n muscle iluriu^ llie cuiitnujtion.
Nu«. ibcT«ntricular:^y»(ule isuiidoubleilly a»im|)lec<)ntra<.-tiuii.a prolontp^]
nmple mntnu-liou, wA a letanua. mtd therefore uii<Ier the old view of the
Mtiiin- nf a inTwcular soiiiiil. could not pnjiluce such a sound ; but. acceptiug
tktHihrr vic-w.nnd n^lleclint; how cixuplox must be the eouneof theavHtolic
mvf iif roll trai.-l ton <iver the twiiied fibres of the ventricle, w« tball nut fiitd
pnl dirtit'ultv in impiMBiug ibat tliul wave iti capable lu iu pnignn of pro*
^ariiii; 'ui-h t(<)H.-tili<>iu>i{f uni-'|uiil tMisinns lu luighl give rw to ii"mii««ulitr
wiimI." and ri>fi»cc|urnltj in n-j^inling ihe fint toanti iii> mninly •>) cmutit.
A< -iich n vii-w nf Ibe origin of the nound, w« fhould cxptM-'t lo find
•■ if ihc niUMulnr lihnf, mid w) the luitureof tioiiml d(![ieiidi'iit on
t!*' .jjniiiuv ■■r Hirid prrwnt in the ventricular nivilivf, and h<nce nn)dilird
bf t>i,-alur« >if the iiwiil vrinj. nnd *till more hv the total removnl of tlm
auricir' itifh llip niiriciilo-vrntricul.tr vnlve*. \\V may add ihnl y,v »hi)iitd
rijrri 111 f^nil it iiii'ititini by the eecnjie of bhiod IVdid the ventricle* into the
anrriri during; the syBtole iiwlf. Hnil niit,'ht rr^^rd this m eiplaining nhy it
ifaiaway bc^n* the venini-le ha.-, eeutud to wiilfncl.
MMTorn*. "i-i'iuK thni the auriculo-venlrieiilar valves innU be thrown into
■Mm irtuion nl the oiufI of Ihe venlricuhir ^vitiole. vrliJcb. aa we have aeon,
it ArttloMiJ with eoiisiderBhle raimlily, not far removed at all event* fti>ni
At npMitr with whicii tfa« teuiilunar valreit are el<iiw<), a rapidliv. ihertr-
far, npabV of giving rive to ribraiiun> of the valves iideijunte to pmdure
■ mand. it iadirtii-iilt to <«enpe Ibe condu^imi that the eliiinirc <if thivv ralve*
■Ml abo irnenttr a tiiund. whieli in ii normally be^tiug heiirl ix mingled
(a MO* «av wilb tb« Mund of mtitcular origin, allliough tbv car eannot
dUKt Ibe muiure.
If we accept lhi> view, thai ibe Miund Lu of itoiible origin, partly " miiBcu-
Uf." jMrtly " valvular." I»>lh eauiMrn being dependent on tbe tension of the
tnirteular cavilic*. wf can pt-rhaii* more euity uiitlorstand hon- it i« that
Ikr wirninl fint round is nl limes to larsriy, indi-cd w« niay say so com-
(llttly, altered nnd oWrured in disetifes <>l the nuriculn-venlrieiilnr valvea.
^celhe lett vi-ntriele foimf (Ik- entire tell apes ofthelieiirl. the iiturmiin
tctther ehnBgesuf tbe lirst si<und heard most diftlinelly at I tie i<|>ot of cardiac
ittpulw belong to tbe tiiilral vnlve of itie lel\ ventriele. Monnunt genemteil
b the tricii^Did vatve of the riphi ventiicle tirv beard wore dbtinctJy In th«
wdiu line oelow the end of ibe Kemuui.
Kuiloenrtiiae PrtMUre.
*
f m. Sinee lh« heart oxbis for i1m> pur|iaM of vxerling presMire on the
Utni within its rat-ilie*. hv which premurc (be eirenlntion of the blood is
(it«t(d. the etudv of the e^anicters of this rndocardiae pressure poawsMa
fital intvtrsl. t'nfonunately. ihe iibHTvalion of thia pressure la stieoded
314
TIIK VASCULAR MK<'HANISM.
witli );TPat r]ifl]ciilli«i. The ordinary mercury iiianr>iueltir wliicli m on user
ill studying tlio prcmiri? in tlic iirlerica fails uit wheu applitN) ui ilie lieart.
It ti true tliut a loit^* canula, or lube open ai (he «Dfl, lillad vriili sorliurn
carbonate solutiiii. may Iw inirixiuccd into the jujtular vein anil wi Kliniiefl
dnno iuto eiiber the riglil aurit-le ur ihe ri]j|ii venirii^le, or may 1h« iiimilarly
Intrniluoo'] into Ibe (.-aroiid artm* and with <-nrf ulipjied dowu thnut){li liii'
oortn, put the soniilunar valves, icii') Iho left vt^ilnrle, and baviiig been
tbua liilroduued may, like the ordinary caoula uned in studying arU-nal
1>reururc ($ ll.O), be broiigbt inui c()iiueci)i>ii with a niert-ury mnnoinoteT.
n [biji way, nit in the ca^ of an arlury, a prapbic rtniord may uc 'ibtiiinvd o(
tbu chnnjjcs of |irc«ure lakiiijc {)lace in thither of the nb»vo thret cavitJM.
But tile diniiges in llic vcDlririilar carici«< are m grvat aint raptd, that the
inertia of iht: mrrcurv, an evil in tlw cn»o of an iirtery, mmcs w) Iar),'*ly into
piny that ihc curve (l(Hcribvd by the flout on tlio mercury is far from beiag
ail accurate n-cord of tbc chaTigcit of prcsMire in llic cavity.
The mcrt-iiry nmiionwlcr may. htiwcvcr, be wade to yicfd vnliialde reeiill*
by ad opt in (I the ingenious conirivanc* of convening the ordinary mnnomei«r
iolo u niiixiruuin or a minimum inlrument.
Tile priiKi|>te of tbe maxiniuu mnnaineler, F\g. 72, uonitiiiO in (lie inlroduetion
into ttii; tulie leading fVoni ibe heart to tli<.> mercury' eoluntn, <i( n (mudrficd cap-
aod'ball) valve, opening, like tlic aortic aeaiiliiiiar valve«, eu^ly frani tbe hcan.
rta.Ti
Till! MtituiH Uxaeanna or 0*u> mid <".i i^
At ' ■ contwoUMi U auili; sriih Ihcliil* iMdlni lo tbe hMrC Wbrn the •pn;v<UBp t Ic eioavd,
Ibt mlTB r (•mil* liibi nrlluii, *n(l llio luilrumcnl. ID llio poililon of Ui* rain ilwun la tb* l^nk
l> a iMxlmiim nutnoniitUf, lly rvivnliiii iIhi illnicilloii of t li !• niii>«n«(1 Inlii a nlnlmum nuu»-
Omrtn. Wliirn t Imptnad, llic (arlHUims uf |<r«Mun ■» miiitcJw) Blong a. nod Uie UMmpRil Uioi
Mtt llfc* an •mil iiui] iiuuiOQiDler.
bat cloMDg firmly nhen iliiid atlttmptc t» return to the heart. Tbe liighmt pres-
Hire it that which dritra Ihe lungect column of fluid Tiwtt the voire, raining the
mercury ouluma lo a currtHpoiiding height. Since thii culumn, oan? pa.it the
valve, cannot return, the mercury reinaiDs at the height lo whiuh It niu raised
hy it anil tbu> iccotd- the Biaximum preanurc By r«ver«ing the direction of the
lairo, tile manometer i* convert<<d from a maximum into a minimum instrumeiit.
TTw mnximum manometer applied to the cavity of either ventricle or of
tbe right auricle, gives a rcoorJ of the highest pressure reached within thiit
TH-E IIKART. S15
nrliT, ami thr niinitiium mauoincler aiinilnrly iliowtt the hiweu prvwure
utrifi, (liiriDb' il>c lime tltiii lbs iiuttrtiiuetil in np(ilieil.
TWni««tntuiii iTi&ii»met<>r itiusemployMJ tliow.-rtlint the maxitnum jiroi-
HTMO llic Uti vciilriclr t> (Iminctir (^reurcr thuu Itie mcsii urewiuK in the
MUfllifi ortlitiBry ntPriniry niMKHDcicr ImviD); previiiunly giveu lti« [hita-
dunl rpMili. due In tlx' ittenia or Ihf iiH>n-ury, l!int U)C iii«nn jin^Muro ta
ibt IrA vtintrkl^ iiiit,'lit \tf li-w iIirii in lli<- iionn ^ thut llir nnixiinimi jiro-
■itin Um Hfcht TOiilrii-l<- ii Ir^t ihiiii in lh<' left, iiii'l ill lliu right nurivlc b
4111 Imil la Uivdofc. lor «'xiuii|>lc, the iiri-Miirp in ihr left vciiiridi* nwcbM
luiimum nrnlxiiil 140 iiiiii. : iiii'riiiryi, in ihv right vciilricio of about 60
Mi.uul in iIh' rii^ht aiirii'h- ol'iilimit 'i^O mm.
Hal lb* chief inimMt ntlnchi^ in the miiiiiniim (irvH^iirr oWrvnl ; fur ihfl
oiiiMiiu ttwiKHnotfr nv>r>l^ ii uryliif prcmiirc? in (he cAvilieaul'the hnut—
iL t. ibmw tbst tbe prwMirt- in ihi-m mny fail bclxn thtit of ihc iitmmpherc.
Tbnio lb* IcA WDtrklp (iif thi- d<%'i n minimum |irt»<ure v.iTviii}; from —
Ut* — 20 IBID. ttMy hv rcncbiil, tin- minimum of thv ri|;hl ventricle being
lh«B — 17 to — 16 nim„ iihi) uf Ihe righl niirii-Ie from — 12 to — IT mm?
hitof this dimiDUtion of prCMiire in the ctirdiav cat-itiee luuy be due, as
•ill b* I'Xplnined in « Inter part of tliie trurk. \'i the iui>iratiou of lh« (borax
ii ite ft«pir«l<>ry movenM-nt*. Bill even wlieii Ui« tlionix i» opened, and
Biiicnl mpintitw kept up. uniter which circuinslaDcn do such mpirati'm
ubi plu«, a ncftalive pn'&Hire i^ «till uhwrved, the ptvasure in the left veii-
trirlr Jllll tinkiiiK o^ l<"^' *^ — '^^ t^'Ri- Now, what ibe instrument uctuallv
ik^iis thai at ituoie lime or other during tlie number uf bentM which took
pbf* while tbe instrument wu applied i and thes« mar lutw been very few)
Ihf fMMUR In tbe ventricle lonk »o many mm Ih-Iow thatof tbeatmoflphere
nhi* mTTBlivf |>irM>iin.- is olimrved when the heart iti Wating nuiti- regu-
,nc\t U-nt Wing exactiv like iho wlhrre, we may infer that a nt.-gativ«
Of* im-un at s»nn- p<!ri<Hl or lahcr of ench canliiu' i-vflo. Itut tho
roiiK-ni ■ibvlinii'ly givcd ii* no inl'orminion w to the exact phaec of the
hwiB iihirh the ncg"tive pnwiiri' iM-curv; to thu point Its wvll M to tho
iaaurtnnrr of (hi> iii-gntiTe jircsurr "c jhtill return pnwently,
\\M The diHiruIlir* duo to tlic iiicriia of the mercury may he obviated
kf (ilcipting the nx'lhixj of Chauveiiii iinil Marvy which cuii*i<t« in inti*i>-
Htiif in ■ largo aniittiil iucb as a horce, through a bloodvcaeel into a cavity
Wlhr hnirl, a tulw ending in nn olu#lie bag i t'ig. 7-t, A ) fashioned »i>inclhing
Kk( • wmimI, both tulw and )>ag Ixting tilled nith air, and the tube being
naotcMd wiih a recording "inmbuiir."
A taW of appropriate corvniurr. A. A, Fig. 73. la ftimioheil at ita and with on
*laiti«ba]C or ''ani^utla" -i. Whi-n it ii ili-nirrd to cxplurc nimultaneoualy Uotli
diritl* and veninele, the mund i- I'liniinhed vith two ninpulla- with tiro amall
•tMlr l>*g*> on« at the extreme end and the other at such a diatance that when
tbf fannvr u within thr cavity of the ventricle tbe latter i« in the cavity of the
•■kir, Huefa an iaatraiDOOt iaijiokenof ann ''cardiac souad." Knrh "aiupolla"
mnicam by a aeporate air-tiglu tulw with an air-tight tambour (V'\g '-i, U)
'»cb a levrr rml*. mi thai aiijr iiri-^ure on the aiiipulla b cuiuinuiiical«(l to
irity of ilH re>t|H<rtiTe tiiinbiiur, the lever of which is raiwd in jfojiortloR.
two anpullB> are iiaeit thr writin); |»'iniit of biith leven arc brought to bnr
I tmmt fvconling furfaee pxiictly niidrracstli each irther. The tobe U core-
'iMroHarei) through the right j'l^iilar veta into the rijchi side nt the heart
lb* hrviT t ventricular I ani|>iilla lo (iiitly \n the cavity ol' the litcht ventriola,
I tcawrijarntly the UJ>i>«r (nurivulnr) ani|>ul1a in the cavity oflhc riirht aurici*.
(loanraihl prvTi-ureon Mtlicr aiu|iullri Ibrn cauw iniweiniMili'nriliv eurm|H>i>diaf[
When tlir preaMre, fur instaoce, on the aiupulla ia the aurkle in iDcroaaeJ,
' n^ kanikan am In t* nail tuvt mnvlf a> Innaari i irhich kin tn^ olBcnvd. aad OM ai
Hp>4««n Itrm ■ tus« ai^ln of n**a
St4
THB VASCUJjAR M Ei.'[I ANISM.
With grwii (liffimltioB, The onlmary m«rcury muiionieter which \a m tiBemT
in ituih'ing tlu- pn^^iine iu the artvrie* failH us wbcu npjiHoil in iho heart.
It i« tnir ihsl II long cuiiiila. or lube open iit the eiiil, filkd with fto^lium
citrWiintn :i'>hili<in, may bi: iiurniliicuil iutu the ju)culnr vein hdiI ita vlifiped
(l>>vrn iiil'> i^lliT i\\v right nnricle or the rl^ht vtmtHt'lc, or may be i>iiailiirij
iuir^xbiccul tnli) the rnmtii] tirt«ry lun) vtiln I'ltn* iilipTieil Hnti'ti tlirnugh th«
sortn, piwt the scniiltiniir vnlvM, into the \ett vcntru'lo, ami linviag httn
thtXi intro<)i]CiC<) mny, like llie unlinnry miiiiln iiwrl in ■lintying nrlcruil
pmaon ($ lI-'>), be brought into (wnnt'ctinn with ■ roemirv minKimcter.
In this wity, nf in tlio caw; of nn itrtcry, ii ^riipbir n^cord niny be obtninetl of
the changes of premure taking pUiM in cither nf tlic nbor<i tbr«e cavitiM.
Dili the chtiiige» in the ventricn]:ir caviliiM nrc to gnwl nnd rapid, thnt the
inertia of the mercurr, nn evil in the ctuc of nn artory, cnniM m largn^ly into
play that the curv« describod by the float ou the mercury is far fruni being
nu accurate reoord of the changes of promire in the cavity.
The mcrciirr mnnoinel^T niay, howerer, be made to yield vabiable mulls
by sdoptiiii; the iiigonioUB contrivance of converting the oritinary inauometer
into a manioiuio or a minimiiin iDtriimeat.
The principle of the maxiniuin tnniKimvler, t'ig. 7:2. conamts in Lbe introdiietioa
into the tulip ImdinK fnim tlie heart to the imrrcury column, uf a (mwlilieil cnjr-
anil-bnll) valve, openlug. like the aortic Miniluuur vatvea. easily from the h«an,
no. Ii
TiiK HiiiKiH UutoHsm or OvLis «)n> oivix
At » • oaMiecQon l> awlu Miili tlie lulio laullnclQ Ibehoin. Wlicn the lercixlapiii * l« Clowfl.
IktnlrofoiiMliitaBrtloii.anillliD limlruiiiviil, Iii iliv innlUuu of lliu nlr* tltaan la ItW SfUl^
It • iiniin«uni muunneUr, Ur nnnliis ilii- dircciloii of i' u l> convcrici) iMu ■ rnlalmwin nuuk-
DiBiAor. Wlu-ii tlio^wsad, Ibe vulnUuui ur iircatiirc aru copiDfvd almif u. sad Uic ttuUanunt Uwn
Mb Ilk* nn ontlEiMT aananKMr.
but cloalng flriair wlien fluid atlcnpt* to return tn tlio hearL The liighott praa-
•ure is tliai whicli drive* the longeat column of Ruii! {>iuit the vaWc, raiting tha
mercury L'olumn to a corrMponiling height. Since ibu column, oncv pa«( thB
valve, cnnimt return, the mercury remalas at the heiglii to which It win raised
liy it ant) iUuh reconin llie iimviiiiuin prMxure, lly reversing the direccitin of the
valve, the munoineter ia converted from a maximum into a minimum iiittiuaienU
The iDBiciinum manamelvr applied to the cavity of cither vcDlriele or of
ibe right auricle, gives a record of iho higbi^<t pKuiire renchod within thni
T1!K HEART.
215
artr, uxl th<> tniiiiiiiuiii itiAiioincipr iiiinilnrly >li'>«r8 tb« lowMt |)r*mira
imHwI, durinj; iltc tiiii« tKiit ili« iti*tritn>cnl W opplitvi.
Thr mkiiriiuiii maiioiRftiT iIiih mi)|»I«v«<I dinwrthat ihv mitximttni ]>rM-
■I* lu the toA v4-nirii'l« » >lt»iincllj grcnlcr Ihno Iho menn prcmure in tht
■■rtiilbr onltnan' mvreury iiiniuk racier Iinvinc previously giveii ihe |iuni-
iniai malt, due to iIm- inonia uf lh« nier<-ury, that the mcnn prcesurc in
(blifl mitmte nii^'lil Ix- leiii' ilmn in lli<> uorta'. thnt ihe niniiimim proft-
Mnin Ihv ri^l)t retilrirle b tfiSB tfawi in iW l«ft. iiod in ihe H^ht auricle is
Mill Urn. In (tie dot;, fur exiiiDpl«, the pressure in ttie left \-eiilriele reiichoa
latiimiiin 'ifatratit 1-10 mm. I nicreury), in the rij^lit Teiitricie of nbout 60
M^ Aiid Id iIw Tight niirit'le of nbout 'i<) mtn.
Dm the rfaief inlervKi Htlncltfe to llie niiiiiiiiiiiii prwaiire uWrved ; for lh«
afainsn nanonieter nsH>rtl' u ne^iliit presgitre in the eavilieanftiie liean—
i(L.«bm> lliAt ihe pre*«iir« in l\wm may fall beluu ijjitt uf tli« atmiupli«re.
Tbiln Uie left ventrifle (of iIm* ci-ifii n oiiuiiiiiitn |iivsHurc varyiiijr from —
VlU — 20 IDID. (UHT be reuelivit, l)i« iiiiiiiiitiim of th<^ ri|;lit v«D(riel« beinjr
fioa — 17 lo — 16 mm.. ainl uf ilw HkIiI auHi-le from — 12 m — 1" mm.
E^nof Ibi* ilimiDUtion nf ]ir«Biir« in the ciinline i^vitin may he iluc, a*
■til W npliUDetl iD a ialer parioniiiu worlc.t^ the ttamralioti (ij'th« thomx
ii lla lOfiiratory nxivinteni*. Bin uvea when iht' thorax i* upeiieil, itiid
arliftfial n>pinil>o<i kv|K up. under whieli cin-iiini'runi«« no i>iieh anpiration
tabi place, a n«^tiv*- pivwurv i> riill •iImitv<'<I, the pri'miirc in the lefl vcn-
iriflr«lill linking n» Ion m» — 'li mm, Now, whiil the in»tritmrnt actiiiillv
abuii i* that at i>om<- linn- or other Htiriuji tlio nnnibiT nf beittx whii'h took
plat* vliila thv iiutrunirnl wan npplir<l i iind ihive ninr haw bc«n very lew)
ihifniiw in the v«nlrid<t!uiMkf>o mnnr mm In-low ihiil of tb<^ atmnfphvrv
9btn: tbi- ncftnlive tirmxtiri' in iilui<rvi-d when the heart a ln-ntinj; <|iiile re^u.
Itflt, i-arh iM-ut being exnelly like tlie i>lher«. we miiy infer thnt a ne|;Dlive
«tv iMxiirf at Kidie period or olber of eaeh eardiac cycle. Itut tlw
DSiTDt obviously f,'ivefi u* nn information as to the exact phiLse of ihfl
til whkh the oeKalive pre»ure ncenr»i (o ihia pojol as well as (n the
on if ihis ne^rative preaaure we bHaII returD prvaeotly.
I US. the dilficiilliefi due to the inertia uf ihe oKrcurr may bo obviated
by iiloptlax the mcthitd of Cliuiiveiiu lujil Marer wliit-h conttiiU in iutro-
Mrip^ in a iarye animal such as a horse. tbrou([h a bloodresMl into a cavity
«ftbe benrl, a tube ending in an ehutii- Imik ' ^ig- 78, A> fashioncit something
likt a miiixl, tioih iul>e and liaj; beiny filled with air, imd the tuln; being
mmtcu^ «ilh a recordinif "lumWur."
) labe of siifiropriHle curvature, A. ft, Pip. T^, is lbrn!«Iied at its end with an
klisj; !•! '■aBipiilln'-r, Wlifn it i« il.*ir(.I lo explore simultaneously botli
it aitil Tvfiirlrli', (lii> Hiund i* riiiiii>hi-il with two ninpu11«> with two small
*(aulc iMfi*. one at the extreme cnil Hiiil the other SI inch a ilblancr tliat when
ibc f'TTRpr ta within Ihe i-Mviiy of the ventrii-le the latter is in the cavity of the
Mr- I IB iaitriiuient iii*iinkenora«a "csr<lia<: sound.'' Each "ampulla"
(DC. by a MjianiU' iiir lijilit tulie vith an air-tiKht tamUiur (Fijc 7-E, D)
•« •Uifli a Ii'K-r r<Bts, vi ihul uny ptnaure on the ampulla ii oJtumuaii.'itli'il lo
(br (arhy nf its resiiective la'ulH>nr, the lever of irhli-'h ii raittd in profiortion.
I iwp aoipullie are usi-il ttit- writing points of botb levers sre brought to bear
Fsamr reeurdinjt >urli*i'i' rinctly underneath cacli other. The tube t* nre-
'{fliraduatol ihroueh the ri|{hl jiiRutsr vein Into the riitht siilc of tbr heart
IllMkrwer {ventiicularl an)]Millii is (nirly hi the Cavity of the richi renlrU-ts^
iMSMnawtl^ the upfier tauricuhkrl ampulla in the cavity of the riclit suriclv.
■BBtof piv«uroon either ampulla then caiiMt inovemeots of the c«rrr«pi>ii'lina
WbMi the prcwure. Tor inslsace. uniboiunpullainUioaorickiit incmumT
*tb— WMattwamobs nxMlitrnrt nmlv >• lawaiMai aUoh Imv* )■«■ utrvsJ. and bMh
<—Wihwwa a— a toiye nusUw al ■—
i
216
THE VASCLTLAR MKfHASISM.
tfa« Hiiricolar l«vtr i* rniif il and drocribpa on Ui« r«corcliii|{ nirikec u ■«ern<linK
CDivr: Klirn Ilic |iirsiiiirc ii liik«n ofl' Ihe curve tl«»cen<la; and to alio ir lib die
T* lit rifle.
The "iKiiind'' miiy In n iiimilnr munnvr he rpiiHUr iitlmilurei] ihrougti ihp
carotid nrivn- itilo tliv /^ vfntririr and the cbunKn ukiiig pinve in ihui chnml;
a1«> ciplotrd.
A'
Hr,
rf
A
MiMT** TAaaiii'K «'iTii Cakdi*c Mi'miv
A- A >Liniile isTtllAP totiurl viicli lu Enaf be lihd Ibr cxplormUoti ut llw Itft nfU'iclo. Itw iort3£m
• olltii' iiuii'illn >1 llii> villi i* t>( III! 11 liiiI!ii'itiM>cr, ttn>ti'l)vil uiut aii ii|i*ii fnniwuiirk iitlli imrlnUle
nppiflti gljuTciinit bt'luiv. ITii: long iu1« ^ wrvn lo [iitnxliici' 11 Into ihu cavllf Btilcli It liitBlnd
loci|>l'in
R Tilt utnlwiii. I'lio cnrUl cliainlii'r m In <nrigiv<l 111 mi nli-tlghl iiiBiiiicr irilh Ihr IndU-mM**
c, bcarLiijE & ihlii Qicliil pUle n'lu vlilcli h atlvh^ Hit Itvcr I iiio%inie *>" the bliict A, Tbe itlHikB
UuiUiiir iMii W iiliicol liy incmiiof Ihc i.-l*iii|ifViil*ii)' lieidOliiu Itiu iii'ilglil r. Tlin lii'Ua-nililivt
liibr rxtvn lociiiiiivot Ihr Inii.'rl'itortliounit'iiirvltiipr n1(b ihoarltyornivuminillii uf.t or mil h
Mnr olhoi CBVlir. Siipiitnliii! liini ilio tube ( vttn connci'lril * lUi b. kiiy jiremun; «■»(■«•! on ^ wuuld
CMi« Itw >u4(«f tbo Umlnurlu rite anil Uia point af lti« Ivvvr woulil b» jirnjiutllciiiiild}' mliwl.
WIk'ii lliLi inHtniRK'nl ia iipplir-'I (o tliv right ntiriclo iin<t vrntriclr' Knn«
ituch rvcori) t* uliiaiiic^l nil ihiit i^liown iii Fig, 7 1, whrrc the ui)p«r curve i» t
trnciiig Itikrn I'nim ihc right iiiiriclc Bin! the Inwrr ciirvo from the right
vciilriclc of Iho hi,ir»«.'. Iwih curve* liiJHg tJiki'ii simultaneously on the smne
n;cnriJingfiirface.
In ihno curvu lliv tisc^ uf tho lever indicatefl prcwure exerlcd upoD the
corrciwiiding aniiniUii. niid the upper curvo from the right niiriclo shon-s
tbp sudden bripf |>rcs<iir<' ib) pxcrlcd by the ■uddcn and brief auricular
syetole. The lower curve froin the righl ventricle ^howit ihnt the preanire
exerted by Ihe veutriciilar sVfilule beginii nituoat immedistely after the auricu-
lar BVfllole. increflBee very rapidly indeed, so that the lever risee in alniixt •
elraight line up lu c, is coiiliuued lor eoiue ounaidernblfi time, ami then
ftlls very rapidly lo reaoh the base line. Itul it may be duubted wlwtber
ifae iDBlruineut can be tnuted lo tell much more than tbio. The prasure
noorded by each lever is the pressure exerted on the imipuUa, and ihb
nay coniinue lo be exertc<l after all hlmiil hm \ttren i)ii><-harged from
cavity, the nulls of the emptied ca^nly eliieing round and prci«iiij[ un
amiMilla. Uut. iia Me ohall presently M-e. it j^ of gniit inien«t to i)e<ermiiM<r
Dot only the furcu and duration of the prevaure exerted by ibe ventricular
Ttric n«A«T.
Flo- -I
,but alio wbetber or do tlie Gbrea continue cmlriict^d iiti<l i-x*'rlitig
■n for RD npprvcinble lime after ibe blooil has been force<l oui o( the
miir. Tb« figure. nii>reover, it i>eeil banlly
WaiM. dwa uot by ilftclf tfive any iRfurtnu-
liga ■• lo tbe relative ainuuiils of pivwure «s-
ttbd tw the auricle and veiitrid« rMpeclively.
liihorarve ilie auricular lever nixa almiit
Ufwhifth a* lli« ventricular lever; Iml vn^
■Ml not luAr ftuni tliw ibat the aurit-iiUr
<nkc ii half aa »Iroug ut the veatrioilar
Asko; tba fomtcr in amogoH mi ux !•> n»>v>-
nnth BDre nawlily, to be much more friivicivf
IIm the lalliT. The iiwiruiiKritt. it i* [rui-.
iMf be cspcrinienlally gra<liiu(r-i], ami ninv
ikil be u(«d li'i ilctrnuiiti; lln' nclunl ■ramitil
rfuraailte; but fur ihU jMirjirHO if iiot uhnlly
■mclor^. W«> may a<l<] dial ihi.' irrt'^u-
hMn Men on thu wiitnnilnr vurvc ilun'iig
(b mtriralnr «yMoI«. ami un the nuriculnr
nitr Bt itic xiini« limv, hiivu givi-ii ri»c la
fluuh iletiate, uni] ncnl n«t be ilitcuwvil here.
0> die nbole, thv tnethod, though useful for
Jlnug a gnif>liie T>ew of ihe fcrira of eveniB nithin the canllar cavilira
ung a canlioc' cycle, the «hort auricular promure. the long* eon tinned ven*
tncaltf fn^nn, lafling nearly half the nholi- period, atid the $iibHx|UGnt
KlniiT \i ( ■ ■■ ■ ■■ r. I, r iinifui! or
tat. Boi^' uii anil
no-Ti,
B
n-a*a* or bDociuiuc |-uo*uaa; Pxia Ltrr Vixtatcu or tmo.
Xa<tnratr«mllkC.II,*liHim>Mil}liniUuN. boul. Tlw Mlvn tu tM* aa^ IM fiirMKaiaa
2a>
TIIK VASCULAR MEfll A SI3M.
d«Keiil lit' tlie k-ver i.'linii^v» In rnli'. liMimiiinj; Inw rapid, c<>riv«pou<l'
end of ilie oultltiu iVom tht- vt-iitriHc ; )>iit thii> m imt n-rUiiii. iind. in>k>c<l,
Um exacl internrelHtkin ur Ihin pnn nf ih<- nirro i« r«i>i>cislly difficult.
Ute oeape rrum the vuniricl^ iw mptd nnd fi^rciUlf ; tht^ rtnw c<'ii>cs fiid-
4001}". Hence, a» wc hnvc ntrr«ily »iati-d, S I'J". nwinp Ui the mlutan of
bloml teudiiif; U> move "n bv virtup nf im iiiprlin iillcr the pr'iiTlling force
IiMCMMhI ti> net, n ticentivi- ;>rx«»iire m»kc» its appcuninre bchinil the column
of blrxKl ilWharKfil friini tlic vcnlricte. iind us eoon its the culiiinii is Indgvil
ill till- iKirlu IchiIk |i> ji irllpx toward the ventride. Thii retliix wnuld nf
ilH-lf liiivo llic cflbct of oliwiitg ihc vnlves even vrere the tiurtn ii rigid lube.
But the iiortn i« extensible and elnstic and the efl^(-l« of ihc movement uf
the column of fluid arc combined with the effecls of the movement of lb*
trterial nails ; the clastic action of the arterial n-alU, in a manner nhioh "«
shall di«cim later on in dealing with the pulM>, also leads to a reflux. It lias
been iirgeil that the reflux due to the negative pmnin of tbe were move-
tnent of the column of blood being more rapid, occurs independently of uixl
earlier than the reflux due to the elastic recoil, the former closing the valves,
the latter securin;; iheir complete closure. Be thin a<> or no the valves are
probably clo^d almost imme<]iately after llie escape of the ventricular ma-
tents, though observers nre um afreet) upon this |*tiint. some urging that the
valves are not closed until so late a period as the puint rf, just tu. relaxaiioa
U about to beigin. In the curves we are now wmsiileriug, n notch, futluwcd
by a t4w, or at least a more ur less abrupt diunge in the course of the curve
■te'. is aointiinics observed in that narlnf the curve nhich intervenes between
tJio int lurge Tvfe and ibe Rnnt sudden fall : atitl thi.s secondary rise has btt.-»
taken to indii'ate the oloaiire of the nemilunnr viilvc:>. Soraetimcxi tw<> such
notch (V and iH-aksnreaeen, and theni-ciirn'ni!*- nf the two ha* )>c«n attributed
to a want nl yyncbmniym in cliv dunure <>f the pulmonAr}- and aortic semi-
lunar valve«. the tatter closlnj^ some little lime before the former. But it is
by no means clear that these notches and peaks are thus due to the closure
of the valves: they may pomihiy have another origin, they arc not alwaya
pmeiit, and indeed it dues not seom certain thai the chisiug of the valves
should necessarily make an impress on the ventricular curve.
^ 134. In the |)erf<irruanee of the ventricle then <aml wliat has been said
of the led ventricle applia> also ti> the right ventricle) there appear to be
four stages :
1. A mpid"gelting u)i" of pressure within the ventricle, all the valves
being as yet cli>sed: this continues until the pressure wiihiu the veutricle,
becoming j^rcaier than that in the aorta. thn>w* oiwn the aortic valves.
'2. The <vcn|ie of the contents of the ventricle into the aorta, the oontrac-
tion« of the ventricular walls still continuing.
:!. Further mninletiunce of the contraction for some little time after the
main body, at all events, of the cuntenLx bare paawd th« aortic valvos; by
this the compktc emptying of the ventricle Miaiut nasured.
4. f^ddcn and rapid rvTiiXBtion of the ventricular walls.
These four events together nmko up a large |K>rlion, and tu « quickly
be«ling heart the greater portion, of the wholo cardiac cTcle.
Meanwhile, that ts during the lime froin t/' to u. blood has been flowing
from the j^reat veins into the anricio : during the interval from h' to d Douo
of ihifl can pass into the ventricle since this is slill contracte<l. but with th«
oomniencenient of relaxation fnmi >' onward there is no longer any obstacle;
on the oonlrary, as ue shall see, an induc«menl for the blood to pass from
the auricle into the vciilritle.
For n brief time, as we have seen, there is probably an unbroken flow
from the great veins (pulmonary or vens cavic) tbroujjb the auricle into Die
ide, lt»diii){ ii> m Mvftdy l>ui aliglit in««aM of tlie frouMo-back diam*
bi > »Jt]cht |>r<i«ure of ibe a]iex on ibe cbc«t-inill, nml in a xlight
bicrtaM iir inim-vriiiriciilar ]>rcMur«, cmiemlljr Hlmnu in the curve nr the
riovljr bmtlng Itiwrt i>f tlii' bon>e(Fiy. li). lu Pig. 76 the suilili-n riw i!uG
luthv ventrkular »p(i}lo im precwk^H by a riw fc luIlow«d hfn (all. forming
Ibo*. a* It wen', a *lii>iil<k-r im tb« curve. ThU hiw l»p<in iiil('r|>rolixl u» inili-
(aitag tbr aliarj) Iriinnii-nt aiirkolnr kvhKiU-; tlic Huildvii iiijci-tioii of llt«
aaricular coDti-uia into tlic v<intridi- iiirn.-iiv:s tbo fronl'lo-back diatn«tcT of
tU natriclc, anil tlw iiiuiiiuiiUini >'!' ib^ rtipiil •tr»kv being coneidcrablo, tbo
Uvn i* in MU-ti ca«e carriod Um> far furwar^f, mt tlinl tbc riM u Ibllowcd hy a
bU. pTQituciag a notch. A fimilnr thougli eomcnhat diflcrvnt shoulder is
■Iv ttm tn ibn cnnliogntra Fig. 77. In the curve of ventrimlar |)reMUT«
tabs by moaiut of the cardiac Miind (Fig. 74) iheni it a rituilar leinpnniiiT
iHraM A^ in the vi>otHi.iilar prrssiire coincident nith the auricular iitroke o,
•ad in i1m> " pi<U)U " pressure curve of lli« rapidly beating heart (Fin-T.i.A)
that b a ninilar thotilder 6 Jti»l preceding the ti^ of the veotriculur systole.
Tk amtirnf; of ibe lasl curve is, bovrever. doubtful, for io tbc siniihtr curve
otiht iBon alowly beating h«art (Fi^. 75, B) it occun inimediuiL-ly al\er ibe
niniliiin of ihe ventricle, eume lime ii^lore the nccurrt-nee of iIil- ntiriciiiar
vnlola. aod )<■ tDany mrven takvti by ih« tuime methixi h ahttent uluigeiber.
m ftact nuiininK. tberefure. of lli« tibouldvr b in the other curvea niuni l>e
Mstiinwni iindn-ided.
t lis. \V>- liiiV4' Mlill «i coimider tbo DC)(ativo prcMure abonii br chc mini-
■an nanomi-U'r. Thi> iuHlrumeitt, an we have Mid, merely •hnws that the
fnmnn in the virntridc (nr aurtcle) beoumea oegativu at ft^mo pbaae or Other
af the canliac cycle, Ixit ilovx nni tell ui Ln which pbaie it occur*.
}i<im there are tno iriiv> in which irucli a negative prwduro might nriginata.
Id Iha flni place, a» ne have just Hxn, u negative prewurc makcM its npjwar-
aaea in the rear of Ibc colomn of blrHHl driven from the ventricle into ihe
aota with great suddenness and rapidity. But this negative pressure, ns we
hiTt ako Mxa, fulluws the column iaio the aorta ]>a«t llic semilunar valves,
aad in pan ai all event* determine* the closure of tbc semilunar tnlves-
Utwe if thifl i> tbc negative prrasurc which the minimum manometer
fteanlik it ouaht tn I>e abonn not only wlicn the end of tbc tube conueetetl
vU ibo nuuiuiueter is in the cavity of the ventricle, but uIm when the tube
M •Upfwd nut of iJie ventricle just {>asl the semiluuar valves. W'bttu ibe
tA»t Bowewr, la in the latter eitiialiun tbe manometer does not shuu' the
«Me Bisrked negative pmMire that it does when Ihe tube is in the vetilricle;
(he nmaiirr pnMure which (lecura in the aorta at each beat h iusuffideiit to
mdace Mich an elferl on tbe minimum nianoineier lu is produced when the
B«ot i* in the \-enirieIe. iieucv we infer tiiat tbe uegaiive prewute
by tlw minimum uiaiioineler u not produivd in tbi.i nny. ^^ e may,
rmr, outidude that tbi; .'«niiltinar vah-ee are cloHeil In-fore thi* negntive
\n oinkta ita ap|wanuice in tlie ventricle; otlK-rnis-, iiowever pmiluivd,
il wiMild l>e traikMnittcd from tbe interior of tbe veutricio through the upcii
talm to the root of tbc aorta lieyond.
But tliere is another event which might give ri« to a n»ativc pfwsure.
Tha rrlaxation of the vetilricular walU in, a« lite ciirvcH (Pigx. 7-^, 76, 77)
4feir, a rajnd prooeas, something (piile dinlincl frMn tbe mcro tilling gf tho
iwtrirulnr cavitica with hlooi) from tbc anrieteai and, though cumo bava
4ijii Hd III the view, it tniiy be urged tluit lhi:i return of the ventricle from
in mklnuled (and empUeil) oindilinn to itM noroiul form wmild devi-iop a
■^aiin> praauro. This return io proiMbly niniplv llw total mult of the
mrs of each Hbre or tibre cell to it# nniurul comfitiou, tlmiigb some have
ojgw) that the extra quantity of blood thrown into tbo ooro&ary artrrita at
S22
TUB VJ
LR XKCIIAS'I
the E}-slot(> help* to anfoUi the ventricles aoinoirbat in ibc wnv ihnt fliiM
(Irivpii Wlweeo tlie tvo walls of a. clou ble-wal led c»1Iiijjm>(1 balfur mip vrill
unl'tilil ii^
Ai'cepuug the return of the veutricle« to their nurmst form ii> ibi* cnuM
of ihe lu-eiiiive prawure (au<l it miiy l)<> rerunrkfil thni Ihi; rpliirn of lb«
thick-v>alleil \etl veDtricle oaturatly exerts n K<^riiit-r ni.-gutiv« [irowtin- than
the tbiii-K'uIled right ventricle), it in itbvioiis that tJie iieguiivf jinittnirD vill
aBrixt the circulation by Duckinjc the hlmn) which hiw mMiiiitbilr li«eii acca-
niiilated in the aiirli-le iVom that cavity iiifi the ventri(!1l^, the aiiriiiilo vcti-
trietilar valvun easily )(iv)iix way. At ihf Hanie timf^ thi« vcrv flow from the
auTicli- will at once put an end lii thi^ UL*gative prvwurp, which nbvioui^lv can
Ih: i)f brief (luratiun •mir. It may further bi* urgvd iu :iup|>ort of this vtow,
thai evGii when l\w thorax in i)p<-Heil, *o that thi; nt^piralorv niovemcnta caa
ii(> loii^r act lounri) producing a iiegiiliv« nrewiircr in the auricle and fireat
v«-iii» < S 1-11 ), a tnitiimuni ninnonictcr placi-i) in lh« right nuHcle sli(>w» fre-
quently oil prtwiirc at nil ilhnt ir, a prt^mrcciinal it> ihal orih«attn(»pher«l
KD<I unmet inic» a drdih<tly nc^tivc nrcmiire. .Sdoinj; that the blood under
thp>c nrcunMlanfra i» being driven along the great vvina by a )>ref«ir« which
though low is alnayx abovo that of th« atmotphere, w« may uuuclude ihal
the negative previirf produced in die vcntriclo ix the cause of this IdweriuK
of the prewurc in the auricle, ihotigli it it unable to make itself felt ahiii);
the greitt veins.
§ 136. Th« •liiration of Ihf trrernl phntfr. We may, lirat of all, dbtin)fuUh
certain main pliaiteH: [\ ) Tbesyatoleof theauriuW VSj The systole, proper,
of llie vfBtrieleB, during which their fibres are in a state nf contraction, Itut'
inn to '/ in FlgB. 75, Ttt. 77. (:1) Tbe dijiHtole of Ihe vent rides— that u to
Bay. ilie time intenp-eDing between ibeir llbres eeaaiufi to contract and cnw
Diencing to contract again. To tliew we may i>erha[s add t4) The iiaui(c »r
rcsil iif the whole heart, comprising the [leriml Irom the ejidollhc rvlaxatioa
of the veiilriclw to the Id^inning of lUe systole of the auricles ; iluring thia
lime the wnll# are undergwtig no active change*. Dcitbi-r rt>ntrnctiug nor
relnxing. their cavities Iwiug simply piusively filM by the inllux of blood.
The mere inspection of nrnuwl nay wrie? of cardiac curves, howeviT tnkea
— Ihoae. for instance, which wc hnw just i)i*CUMcd — will show, apart from
any accurate measurements, ihnt the svslole "f the ntiricles is always very
brief, that llie systiile of ths vcntricUs is alwuys very prolongwl— always
Qccupviui,' a conBidcrablc portion of the whi>!c cycle- -and that the diastole uf
ll»e whole heart, reckoued from the end either of the systole or of Ihe relaxa-
tion of the veiilricle, is very various, tx'ing in <]uickly beating bearta very
ahurl and in slowly beating hearts decidedly lunger.
When we desire to arrive at more complete mensuremenis, wc are obliged
to make me ^>f calculations based on various data; and theee give only
approximate results. Naturally, the most interest ia attached to the dura-
tion of eveuta iu tbe human heart.
T1i« datum which perha|M ha§ been most largely used is tbe interval
between the beginning of the Hrst and the occuiTenee iif the second sound.
Thitt may be iletermim-d nith approximate correctness, and ia found to vaiy
from O-Sftl to 11.327 w^imd, occupying from At) to 4G per cent. i>f the nbule
|ier[t>d, and tiring fairly ouiniant tor <li8erent rates of heiiri-beai. That i» to
aay, in a rapiilly iH-nliug lii-url it ii the juuses wbicli are nhortencd, and not
the duniliiin nf tliv aeliud hi-at*.
Tbe Dbaerver, iMtenine to (he sounds of the heart, make* a •ii(uaJ at eJich eveiiL
•<■ a rvcurdiii]( Mirlace, lue dtflrrencn in time litlwci-n tbe niaila being lOMunired
kby BmnB iir the tihralions of u lutiitii;-t'ork recurded on the tame aarface. I)r
imutice it is fuuod [Nmible to reduce tli« errorsof obaerratJoa within very smalt
limits.
THK UKART.
^2S
Xow. wKikiCTrr beihv exnd cmiMiiiun of ihv tir»t »i>i]n<I, ii in ani)oul>lcdly
11 «illi ihi- Kvxtoli- of lh« vpiitm'U**, thi>ii<!li tii<s<ili!v llw iicluiil com-
Dtni of iu Imiiiiilii^ aiiilihir nuir Ix* iili^lilly iH'tiimi ihc itctntil iicgin-
Tiin^ nf ilir tiiiiKCiiliir cotiirmcltniia. i^imiliirly ihc nrciirrvncv of the tecciDd
■juntl, whidi, iiit wr h»v<- -vu. U (H-rminly due in llir clinsiirc- nl' the ■enii-
loDftr Tilim, hiu )>ct.'fi litkvii !•> tnttrk lln- cliwc nl' (he veiitriciilar »vst>ile.
And iin tkU ■D|>|NN>i(iuD Ihv intonal hrtnci-R the hc^iiiniiiif; of tlie tint aud
llw iicrum-iiri' of tl>e i^cniKl tumnd hns l)c<n M-iiHnictl lu inilioiitin); »|>pr<>xi-
ouilrty lilt' iliinitiiiii uf thr v-cntri<:(il«r rvfIoIc — ■'. c, ihv pvritKl during wliicli
ih* trnlririiliir fil>rii> arv coiiinictiiig. \Vt> liiivv, howover, orj^d abuvc ihat
tli« vmlni-lri fttill rvniain coDirnctcol fur ii brief period n^er ibe valvn are
tk»l: if tlii» viv«T 1m> cwrrccl, llicn tbp iwcond cuuiid dms not mnrk the end
tf tkf fifiole, UMJ the duraiiiMi of tbe sjHtole is milwr longer that tbe time
■imi above.
ibf I lot ertiii nation of lb« tcnaralf diirnliou of eucb of the thrue jieriutU (d'
tbt iFiilricuIar Bvatolt' — vii., the ]>t^iliii^ up ut' lb« prtsaure, the <lUcharge of
tb( niiit«nl£. nnd tb«- reiiiainiiij- eniplieil but cuiuracled — is mibjec't tn no
BuA unnrtainty ihiit ii ueeA not be ioBistcd ou iii-re; it may, buwcvcr, be
ml tkat, ruuKbly ajieakiiiK. «^:>L-b phiue occupies probuhly iibout 0. 1 wound.
In 0 li«uri tumtini: 72 timcii a iniiiui^, uriiieh mar t>e tukeD as tbe normal
nif, Mcb i.tiiin: ciirdiiii: cycle would la^l abiiut 6>^ d^Muiid. ami taking 0.'^
mod a* ibc duratioii of ibe venlrietilur ByHiokMbc deduct Juii of tbU wiuild
Invr a^ wctiud for tbu whole diaalole of (hu vciiirii-le. iiicludiii); il.-i n^laxti-
Unn. tlir Intlrr orcupyiiiK ubotit or sumowhiil Wji iliaii 0.1 iu.t?i>iid. In thu
liatr part of tbiii period iht'ru occunt Ibe M'stole of tlir aiiricln, tlii; cxucl
duruii'i of wbicb it a ditlii-iilt to dcterniiiip, it being ban] t» aity wlu-n it
mlly U>'ina. btil vrlii«b, if tbv cviitrnction of the grau v«iiis b« iucludul,
luy [Thara be taken ns la«titt);. "U no Mvemue, 0.1 wond. The " |iuB«ire
bimal,*' iberefore, during which nHther iiiiric^ nor ventricle are undci^iiiig
BMfaUiufM, lasts about 0.4 hocoikI, and tbe alji»olut« paiii»e or rest during
viich iMritber auricle nor ventricle are contracting or relaxing, about 0.3
■cowl: if. however, a longer period be allotted to the ventricular eyslole,
lUw perioiU muat be proportionately Hhurleiied. Tlie iVBtote of tbe ventricle
Mloiraao iinmediaiely upon that of ibv auricles, that practically uo interral
aiirt» between the two evenli.
The duration of tbe iteveral phases may. for convenience luike, be arranged
is s lalnilar form m folloun; but in reading the table the foregoing; remarks
■t la tbe appruKiRiale or even uDcerlaiii character of some of the dalA must
W Imnw In raim).
^tfole of vetitr(c)« belore the opening nf die Minilunnr
Talvei, while picMure ia atill getting up {probnltly ralhrr
iMthaa) Il.l
beaiNr of blood into aorta (about) 0,1
Cb«tinu«d ooairactlon of tbe emptied Tentriele Ipoaslbljr
rather mure tliant 0,1
Total *y«l«la of the ventrit'le (probably tailivr nxxg
ihani
IHaabile uf both aarii'le Rn<l vMilricte. neillM'r oontracting,
••r '* paaaive Inlvrvul " (pn>l>ahlv rather leas than) . . 0.4 I
Ryatole nf auricle ^abtxit or leM tlmn) IM j
lKa*(i>l« or vciilricle. iiiclu'linj: rrt>xiiti"n and filliiiir. up to
lb« txf inning uf the vcnuiuulur ivsiolr ((irotiably tiiilicr
Ina Ihao)
0.8
0..>
Total cardiac cycle
0.8
224
rs VASOULAB llECnANTSK.
Summari/.
! 137- We mny iww bricfl)r nninintiilitU' l]w main fitcte coiinecloc) willi tlx
posnge of bliHMl through the liritrt. Thi- right auricle ilitring il> din»litl«,
by the rclitxnlinD of iu imisculur dhm, nntl bv lli« liict (hat nil bnckwnni
preAuro frum rbc vriitrii-Ie is rcinovixl by the closiug ofUM) tricuspid vnlrea,
uHon but Unit; ri-siftdticc to (he ingrivs »t' blood from the veins. On lbs
other bncid, the lilood in the tninks of buth the BU[>crior miil liirorior vena
cava i» under n prcMUre, which, though diminishing b>«-ard the heart, re-
luains higher thuti the pretaure obtaining iii the ititerior of the auricle ; (be
blood in cotiwjijciice dows into the empty auricle, its prourcse in the case of
lh« superior vodb mva bcinj* atdisted by gravity. At each inspiration thb
tiow (as wcchall aee in ^pealciitg of respiration ) is favored by the dimiiuitioD
of prcflBurc ill the heart aud great ves««ls caused bv the reepiralory loove-
nients. Before this How liau gone on very lon^. the diastole of the ventricle
be};iii6, 'H» cavity dilates, the IhipA of the tricuspid valve fall back, and blood
for fiouie little tirne tloiva In itu unbroken stieaiu from the vena- m\x into
ibe ventricle. In n »liori limii. howevtr. probubiy before very much i>luud
h»H hud time M enter the- vcntrii'k-, the auricie id full ; and furthnith ita
Hharp sudden sysiok- tnko place. Partly by rea^in of the buckwan) pr«B-
jiuru ill the veiun. which incnruofs rupidly from the heart towmni the cupib
Inricn, and which, at foinir diataiici- IVoiii thi- heart, in OMinted by tin- proenoo
of vulvi-s ill ihr vtnoNit triinkf, but Mill miin> from thi.- fact that the xvrioU;
begins at ihi? grrat veiuN thrmM-Ives and spreads thence over the auricle, the
Ibfxw of llie auricular oonlnielimi i> £|>ent in driving the bigod, not liack
into the reins, but into the ventricle, where ihe pri-wum in utill excceflingly
Ii>w. Whether there u any backward How at all iritu the great vcIr«. or
vhethor by the progressive character of the systole the flow of blood oon-
tiuuee, io to speak, to follow up the systole without break, so tbnl the Mreain
Irani the veins into the auricle is really cuutiniioue, is at present doubtful;
though a slight positive wave of pressure synchronous with the auricular
ayaUile, travelling backward along the great vcina, baa been observed, at least
in OUHB where the heart la beating vigorously.
The ventricle thus being tilled by the auricular svaiote, the play of th«
tricuspid valves described above cornea iniu action, tlie auricular iysiole is
iblluwed by that of the ventricle, and the jire^urc within the vetitricle, cut
<dr frQm the auricle by tht- iricii^pid valves, in brou^-ht to bear ou the pul*
monary leinibiDar valven and the I'libimu of blood on the other aide of tritno
vatrea. Aa noon aa by the rapidly inereti^dn^ shortening of the ventricular
fibnn the prc^aure nitliin the ventricle beouiuea greater than that tu the pub
uionnry artery, the iiemilunar valven open and the alill continuing nyatolv
duobareCH the contonts of the ventricle into that ve^isul.
As the ventricle thui> ra|>idly and lorcihlv einptien itni-lf. either the trnnjucut
Dcgative prauurc which innkra it« aj>pearance in the rear of the i-jected colunia
4>f blood, or the eln»tic action of the aortic wall*. IcuJh tu a rellux of blood
lowanl the ventricle, the eft'ect of which, however, is to close l)ie iwniiluiiar
volvw* and tlni# to i>hut off thu blood in the dwteiided arlericri from tlio
emptiefl vQiilricle. Kither iinme<lialely at or more probably fi}mv little time
■Aer this dooingof the valves the venirieulnr Kv»iole emlif and relaxation
bt^iDs; then once aum the cavity of the ventnclo bueomtw unfohlcd and
finally divtciKlcl by the influx of blo<Kl, a negative prexiure devolopc^l bv
the relaxation probably niiliiig ihc How from the auricle nnd greal wins.
Utiring the whole of this time ^e left eid« bag with still grotMr eaotgy
been eJteoutiug the same inanceuTn^ AtlhesaniQ time thai tha von» cavn
THK UEABT. 22S
•n Uioy (be riKht iiuricle, the pultnoniu-jr veim are fillioi; llie left ■iiriclv.
Al l^mne time thai die risKt auricle ia cMntrai-tin];, itie left auricle it coR-
ITUtiDi; Uw. The BTBtole of tlie left v^eotncle \a syDcbrnnous witli thrtt uf
Uttcunt \-rtiLri(;le, Imt executed wiili frailer foree; siul tlte flow of blood
a gnnn) •■n ilie iHt liile by the iiiitrul aod aortk valree in th« aamc war
ilni it is on tlic right by tbe lricuj»j>iil valves ani) the viUves of the pul-
murT arwy-
"Hftati in a givcu |)erio(), but pi-rluiDS the must importaut factor of nil in
ih dtUnuinatiiiQ of the nork of llie vasciilur niecliuDisni, the <|Uttntity
qstted from the renlricle iulo the noria at i?acli systole, cannot as vet be said
The Wort Oow.
)18& W« can muiMire with apiirosioinlivc exactness the intrn-rcntrii'ular
in. tbo U-iiglh of Mcb «yBlolc, and the number of tinKv the systole ia
tobtTe Iieen aecuralely determined ; we are largely ublijjed to fall back oo
nlealationa liavinf^ inanv sourcee of error. The jceueral result of soioe of
ihaamlculaiioosgivca aFiout ItSO cramuia (<j oxa.) u the quantity of blood
vlitb it driven from each ventricle at each eysu>le iu a fiill-j^rown man of
I sice and weight, but Ibiii e«tiraale k probably tui> bi;;li.
Im tb« dog tlifl iiuauiil^ liBB been vi|ietiiu»n tally <lei«nnined. by allowing (be
fcauitoddlver iia eonteobi iliroujcb one branch or the aorta, all oibera being
"p^T^ or blocked, into • reooiviT. (he contents of wliich are at iotervnl*. by an
Upaloai coBlrivjini:*, rriiirni-rl in the right auricle. Tha time takon ta fill the
nniw aitd tbc [luuilwr »f Wata eiccuted during ihiil time hviag aoted, the
annn quantitT aitict«d at a beat la tbiuciveii. It ia found to vary very widely.
VatttM Betbovi have been adopted lor cal(;ulatiag the average amonnt of
Und ijected at each rantrictilar Hystolc. Tlie almpleAt method U to meaaare the
oifadtj of the rfcmlly rpmavcd and m vet not rigid ventricle, flllod with blood
•Mf 1 rirrware e>iua1 to tbo calculated nwnfe prea*uro in the veoitricle. On
At ■ >ii that tbe whole cunienta of t£e T«alricle are iMMtcd at each
■fi: -juld give the i]naallty driren Into tbe aorta ul each Htruke. The
w«tbwl« arc ret; indirect.
It ii CTHleat that exactly the came ijuantity must inue at a beat from each
fntride: ibr if tbe right ventricle at eaih beat gave out rather leae than
ll> kft, after a certain number of l>eatA tbe whole of the bkwd would be
plLiiil in the svstemic circulation. Similarly, if tbe left ventricle gave
Ml !■■ than tbe right, all the blood would soon be crowded into tbe lunjf*.
1W tun that the pr«aaure in the right ventricle ia »o mucb leM tlian that in
(he left (prabably 30 or 40 mm. aa comimrHl with 200 mm. of mercury), is
Att. not Iu dlflbraioes in tlie tfuautilv of blixid in the cavities, but to the liitct
tbal ibn ]>rri|>licnil redttance whidi baa to be overcome in the lungs b so
smefa I^ than that in t^ iwM of the body.
Il moat be mnemWred that though it ia »>' nilvaninjei^ to nwuik «f an
tnrage rjnnntity tjerted at vach atnike, it ia more than pn>bablc that thai
^•aaiiiy may vary witliin very wide llmila. Taking, hawcvvr, 180 grammes
» Ibe r|u«ntity. in man, tjecteil at each stroke at a pnxauiv of 2-V> mm.' of
OMTury, which ia oquivaieni to 3.21 melren of bloud, this meaiia that lite
Ul Teatricle e capable at its systole of lifting 180 grammes 3.21 melna
y^ i. a., it does 57H gramme-metres of work at each benU Supporing llx-
Mft lo beat 72 timex a niinuta, this would give for tbe day's work of tbe
Ul v«Btrid« nearly (JiMXX) kilogram in«>metrcs. Calculatingtlwworkof lbs
fifU reotricle at one-fourlb iTiat of the left, the work of the whole h«an
< A Wall caOmaM I* |art«Hlr UkcQ kuv^
IS
S36
would nmouiit lo 75,000 kiiograuime-melres, which ia jii«t ubotil the aniotint
of work tlciiu ill (111' nHveiit ol' Sijowdon by a tolerubly heavy iubd.
A cnlciiliitiun of more [intcliciil value- U the lollowiDg. Taking the
riuatitity of bUiod n* i*i n( ihe hmly wei^ihl.. the bloud nf a man weighing
76 kilus would Iw about A7G0 grniiiiiivM. If 180 ajaminea leA ibe veolricle
•t each beat, a ciunntity C(|UivuIi:iit to the whole blood n-ould paoa ifarougb
the heart in 3*2 beale, *.«., in k«t than half a minute.
Tiis Pruiii.
^ 138. We have seen that (he arteries, though always dislcnded. undergo
at each systole of the ventricle a ti?mporar}' additional disteunoo. a Icmfuirarv
additioual expansion s« that uheD the tinker is placed on an artery. *iich ua
the radial, on inlerniillvnl |>reature on the linger, comio); and going with the
Fro. IS.
n«1t tmm MuioHirrn.
in* flalloued Inlic 111 tlw f^irm Of • liaop t* ami)- ttinl u uiio md, vhflo Ihc al>»t tn» •nd U
•KmIknI Ma law, Thu liiUilur of llm liilir. ttlM wltli iiilril. Ittuniufhl. hf lurvuof ■ inteooi)-
ntnltic Mdluia oniboniitv Kiliilloii, liiro ninnrrili.n wtih nn aMcrT. In mui'h tbe Hmc hajmId llie
(■■I tit Uio Dtimi; DiaiuuuelQi. Tht luvrcue vf [ircBurc lu tbe aner} Uilug tiuminliwil lo u«
ImUowIiDop, UiuUluflnlKhUn tt, aDdc>'iTM[ianillDirly nKn«> tb« Bllwb*d t«itr.
beat of the heart, h fell, and wbeu a lt):ht lever is ]>laci-d on tbc nrterr, Iho
lever i* raited al «ach boat, falliuE hetwt^en.
Thi* intermittent L-ximnsion which wc call the pulw, corrwprmding to the
jerking outflow "f hlood tVinn a severed artery, is im-wnt in the artenc» only,
DeiDiF, except under panit'ulnr circumNtnncca, aWnt from the vrius and
cafiillBricM. The es]Hui»]on i* fn-iucnlh- vinhlc to tbc eye, unit in some
cases, us where an artery has a hend, may cniifc n certain amount of loco*
motion of the Tnsel,
THE PUt.SE.
S27
iitatyonrj iucivntt of preoHire wliicb is tlte cauM of the temporary
01 oxiMiiMOD makes ilself fell, as we faavo seen, in the ourra of
rra.T>.
IiusRiH (M > BrnvnuoaiiArii iDiiiA>AN*n.
II1|H<IIIH |auM uc oflilncd n that Uie ■aUlptjliiE tctrnmiaj'bedliptoTod. d tin nuall
»(if UMknrtalimiiiMtMdpabitil. Tlw* an rominunlaiWd w uid nu«-
I If tt* ln«t « wlilcli iBona roaiid ib> Bzwl fdnt /, Tbe live 'nd or (hit lenr BoiiUi b tight
■MMr <rUtk iwHoaftMnpottanlud raptr.tr- Tt« |iai«r Si plKvd bvocaUt two mull
•( IMM <■ ■ ioSm abMi «■ t« rotolod bj ausoi of rtock-xirk ociaUJn*d la Uw bpt k.
1kt|i9«rltihai«a«i(1totniTeUt>anllbrmm*. nttrnw gnAuiua lnoiuioa(TTaT)talmiBlM
• ta* M Ih* ifAliB * tV niMiia of a MBm. and t>)r iMt Ihd pnmuf pul oq Um iirUfy (nn b* Ntfu-
■■a tkt ItfW* OMpaltr Ik* |uIm mtanaaviM llttjr ItmM.
MnU pWBW lalcciii bv ihr mrrcnry mnmimclcr ; but the ini-rtJa of the
[jmrcoto th« epCL-ial vhiu-uctcn of ctu:h incrt^uec bot'oming visible.
HiBit'i t!riiTn»mR>rli.
VXb*kOTiitaw<ii|irr'n'V*i>iii*<>M'i>'<i i»tbo«m. lt,if(tiia>klcfefaiiiipiNi tadlaluMtrt
te adfutl^ niarkla« tiii-i I. : II, rlnrkiAck ; l\ iMtfenl ymtm vpM ■faloh tradaK tt
,MMn«HllH ft ttui 1 1 iif I ml ■ftii r>l>lilt 1
Mf'<«Mn*i«t
228
In Fick'« fprine Diiini>iiiel<T (Fig. 78), in whii-Ii titc invrcMo of prcMUtv
unfolil* a cunofl iipriuK iiml mi movt» n hver, the inrTtia » much Ivm, uul
MtiafiuiUinr tmcingit may be Uikcn l>r \hif iiiiitriimettt. Oilier inetrumoaU
bsve aIko men cl«vU«(l tor roounJing ihc spocinl characters of each incr
Pia.il,
» AfkcJ-ii
I-i'i*> "nuoKii nnoi TiiE Raciji. aitkut ur sum.
Tba Tvnioil curTEd lluo, L.glTw tba na«]i>EHtiIviiilior««riltiit; luira iii«l«iib*n Uia Hm^
•nol psiwr KU mntloiila*. Th* <iitm4 InUmpLcil Uno ihuw Itnr ittirlanM (tam tne iwottia In
aiDO «< th* chlcl plum at the pulw-WAni, tIk. ;g — commenMmont uid A eiul ut vrawton of
wiair. p. piCKlicnllc uoteta. d. dlcfoUc noteb. C, ifleniilc cmi n, INMt^kKitlo nM. /. Um
iM^rMlc nolcb. ThtM BM «l|ll*liied tn Iha Wxt lal« ■>□.
of pnaeure or of \h\: cxiiimeion of the iirterv which ii> the result of lliat
tneMaaei The eiui«8t sod nioet comnmn method of regiaterioK the ex[>aiuion
of aa art«rj is that of simpi)' bringing a light lever to b«ar on the iiut«i()«
of 111* artery.
V.
f^-
BI>B rubber puiDp, .. .' :i . i: . >iiuciiini-nl. to |irtircnl* nsanrl<A"l <.-iim.iit: t,f,f. uv Icrrn
nMosiiii kciiui tuba, *t liiiermb «1 3) cm. ufiubliis: C.drom upon iitJeb uuliif Ii maA»-. II,
fliockiwivk 111 rvroUc itrum.)
A lever s^ially ailaplcd to reconi a pitlw traciiiz u called a sphygnio-
granli, the iostniRMUit gcoenlly comprising a itniiill Iruvolling tvcordioR
Mrtace ou which tha Uver wrHea. Thorv are many diflvrvnl fbmia of sphy^
THE PDL9E.
229
tMpipb, bui llie g<>tienil nlnn of nnictun^ in th« samr. Fig. 79 ri>pr«KDtfi
ID « duf runnwtic form the cakuUhI P^* of (he »phygnio^«pb, ItDown ss
pii4geaa'« Tum] Fig. S2, Marvy'ii, which is id moro corhdod use]. The
iainumt is gMOrally n[i{>lt(il to tho nxlinl iirtrrv bccniiee the ana offonJa
• (■inn[«imp))OTt {*> Ibc Aili'mtii <>f tbo k-Vf-r, nnd because the position of
pm. t&
^\/V^AAA/^iAAAAAAAAA/
ftiM ravm na^BM> st a Hun or ^JiiT'inoaurnic Lxnw-plkndMtnicrTalaol Vem
^■M*Mtarakiii(*ncOjallClubelnIoiihlchfliUi)Ufan<dbTthoMdd«nMRitwc>(«puia|>. Tbe
^■»*«>* It mMlllliK nuu krA to itclu. u InJIoUrt bf lh« umn a*«r Uip pninaor (a* Bml neond-
iQA<lf«lv««i«L TlMiMM v«mc>lllnndnwiift<an thaMinimltorilioieT«nl|iiliiiu7'«m<r«
■*<telMlB|Ajik DOnv litlDvr, ouh tmoiplQla TtbfMloa of wUcti aceui«i> 1 JO aeiaBd. ftUowUi*
^tab*Baa*nn<Hblchl«ul«upbjtb*miT«lnpMl4>lans3a(Tni.orib*tublBc Tbo««T«
^■n>MW«<tftoMf>nntbark«dilMkl*nit of Um mUm : tkli la tiullcaittl by Ue dtrMiloii of
^ ■*•» II win be obKtTDd iliai In ll» wan dtauui tera VL Itic nOeelod wan. tiNT&w tol •
■^MAaumlo tivpel, bfoOBiM nawl Willi lh> |iriinu7 Hun. iKnni MaKST.)
tk ulfrr, n«Hr to tbe lurfncc aii<t with tbe Aipimrt of the radtus b«lo«r ao
ilat Mlc^iute prewure can be brought to bt-nr by the lover od the artery, »
fcrcnble for niakine obwrvatiomt. It vuu, of cuurao, be applictl to other
amritm. When ap[iiit)il to cIm radial ortvrr lomo such tntcing a» that
4mM in Fig. 81 in obliiiiiccl. At acb bcvl bi-itt Iho lever rises rapidly and
Ami hiU mote gnulually in n Hdc vrbich U more nr le» uneven.
280
THE VASCULAB MKOQANISM.
S 140. Wc have now to study the nnUirc ivDd characUrs of ihi' pul«« in
greater detail.
W« rnny fnr at once, und indeed have alreadv incidentally eccn, that llie
pulse is eBSpniinlly due to the Hction of ptiyeical cuiieeB; it is the phyHcal
result of the sudden injection of the conleote uf the ventricle intn the elxatic
tubes called arteries; its more imporiani features may be explained on
physical principles and may be illustrated by m«atiB of an artificial model
[Fijr. 82],
If two levers be placed on the arterial tubes of an artl6cial model Fig.
61 &a., iS".(r., one near to the pump, and the other near to the peripheral
reiUtance, with a conuderable length of tubing Ixiween them, and both
leireni be made to write oa a reo<brdiug surface, one iniDiMliiitelv below the
orhvr, mi that their curves can he more easily oomporeil, the fullowiug facts
muv be ob.ierved, when the pump in Met to work regularly. They an- i>vrhn[M
still better neeii if a numher of levers be similarly arrangeo nt ilitlcnmt
di.ilunc^ from the pump ha in Fig. 8.1.
At eiieh »t.rolc<! »!' the pump, Midi lever rises until it rctncbn< a maximum
(Pig. 8.S, la, '2ti. i.-[c.") and ihi'u fall* again, thus d(«cribing a curve- The file
is due Ui ihe i'xpiitiiii<)ii iif tin- part nf the lulio under tho lever, and tho fall
IM due to that part i>f the lulw> n-liiriirng after the expansion to its prc\Hou»
calibre. The curve in, thi'refi>re, the eurve of the expausioD (and relorn) of
th« tube at the point an which the lever rests. We may call it the pulse-
curve. It is obvious Ihttt the expansion |ins«c« by the lever in the form of a
wave. At one moment tho lever is at mt ; the tube beneath it is simply
distended to the normal amount indicative of the mean prasurc which at
the tjme obtaius in the arterial tubes of the model; at tlie next moment the
pulse expansion reachw the lever, and the lever begins to rise ; it continues
to rise until the top uf the wave reaches it, after which it falU again undl
finally it comes to rest, the wave having completely pacsed by.
1^. KL
/
A sovon DiKTRxwHUic i(KrKi»xv(iiTi'H< riF t fruc-wAV* N«q]ra ovu m Aktuit.
It may perhaps be as well at once to warn the reader ihut the figure which
we call the pulse-curre is not a leprcnonl^ttion af the pulw-wai-e ilMlf; it n
^mply a repieMniation of the movenient», up and down, of the piece of ihp
wair of the tubing at the spot on whieh the lever nste ijun'n^ M4 linu that
the wave is passing over that spot. We may roughly repreM.'nt the wave in
ihiiiHiiiii Pie< M in vrbicb tho wnvu «boira l>r tho i)i>tt«d liito i* pM«ing
«MlM lab* («D(Mni in it mudilion of ruet hv the tliivk double lint') in Iho
JiNdloB fromHbi C It must, however, (>e remeitiberecl tbnt tho irnvo
itaiSpind is a mueh aborur ware ihao is the pulse-ware in reality (that
Uk m w* BbBlI aae. about (1 nwtrw loi^), i. «., oocupia a smaller length
WtM arterial airM«ni from thv heart /f toward the capillariw C
Tbe curre* below A*. 1'. Z repreaont, in a similarly <tia{;raiDm8tic fluhioB,
iktcorret deacribed, durio)* the passage of lli« wave, by leran plocttl on
ibt points 1', y. t. At X the j-roiilcr part of the vrave has already pMoed
atoUie lerer, whi«fa durinff its passage haa already described tbe greater
)Mt of its curve. sIiovd by ibe thick line, and has only now to describe the
Mall fan. Hbown by the <loiied line. eornM[M)i)diug to the remainder of the
an* m>m Jf lu //. At I'lhe lev«r i^ at the xuiniuit of tbe ware. At JT
ds lerer baa only described u Hniall pun of the tieKinning of tbe wave, vis.,
bm C to X. tbe rest of the curve, us sliown by the dotted line, liarin? yet to
hid«eribed.
Bvi to retora to tbe OMisideraticm of Via. 8^.
t liL The rise of each lever i* soiitcwliat sudden, but the fall is more
psdual, and is generally murkei! with Home irrcfrularititra whic^h we shall
niv pnacDily. The Hmc u *tiil<U-ti bcn'Aiii'C tho sharp iitrDlce of the pump
— Uwily drives a (jnantity uf tliiiil into the tubing and «•> »ttd>l(tiilv expands
itilabe; tlte fall u more t;nidiinl because the elastic rutt-tion of the walla
rfiha tab*, wbic-b htin-j* nbont the return of the tube to its foroMir calibre
lAirlW expanding jxtwcr of the pump has ceased, is more gradual in its
BrtiaD.
Tbaa Aaturai, the Rud<lenn<M nf the rise or up-etroke. and the more
pailaal slope of the fall or dnwn-Hlroke, arw seen ako in natural pulse-curves
Iskearrooi living arteries (Figs.81, 85. etc. 1.
Uied, lira didereuee between the uf>-«lri>ke
nri ibe down-strvike is even more marked
ia ibr latter than in the former, the (k-livery
(/ blood iVom the veotride being more rapid
Ikia tbe iMue of water from a pump an onli-
Birilr worked.
It may here be noted tliai ibe imtual lixe
<f fb« curve, that k the amount uf exciiniion
<f ibi lover, ilcpeuibi in part (an does aluo to
Sgrat axtent Ui« form of tlte curve) on the
laawil of prcMure cxcrtiNl by the lever on
Ik labe. ir ibe lover only juHt tonrhcs the
tub* ID lu upaoded aute, the rim will be
NigBMauit. If. iMi tho other hand, the
bnr ba praned down two firmly, the tube
haath will not bt^ able to cx'paDd as it.
Hlaiise wovld. and the rite of the lever
■fli be proportionately diniiiiUhed. There
btcertun prasure which must be exerted
hr tbe lerer on tbe tube, tbe exact amount
■bjRsdtng on tbe expansive power of the
taUifand on the pressure exerted by tlie fluid in the tube, in order that tbe
■ndig nay be best marked. This is shown in Fig. M5 in which are given
thne trwdngs taken from the same radial artery witli the Kauie inatruraeat;
in Iba lower curve the prewtire of tlie lever is too great, in the upper enrve
H» mall, to bring out the i-haniut«ni seen meat distinctly in tJie middle
<WTt with a medium presaure.
PM. Ul
JltL^^^
Pci«i TsiaiAH i-Bos inn >4>U R^
lUAt, AsTUr rxpKi IHrnman
I'UaHIKM or TBI LlTU.
iTbB Icltenmni sxiiUnsd In ■ IsMr
■■ft of Uio (Ml.)
282
THE VASCULAR )ISCBA^'IS1I.
Fia ««.
Fro. M.
li 142. It will be obttrred tliat ia Tig. 83, eurre I., wliteb is nearer
|iiim{t. tiaes biglier, and rises more rupinly than curve II., wbiob ia fiit
away from Ibe )Hmi{> ; tbal h t(> sny, al the lever Airther awnr froiD ifi?
pump, tlie expautiioii U Uta hdiI take* place miire tloirljr ibmn at tbo lever
Dearer the pump. Simil»r]}- in curve IV. tbe rise is sUll IcM, itnd takes
place still lew rapidly limn in 11., and tbc wmc cbange is teen Ktill miir«
marked in V. a* conipnn'd «ilti IV. ]ii fuel, if a number of Irvi-rw «i-re
plae«d at ciuul diKtaiii-es nioiig tli<- arterial tubing of tbc model and the
model were vrorklni^ profNTlv. with iiu iiili'ijiiiilp jHTiphcnil n.wlam'o, w«
mi^bt irar'f out step hy step bow the rxpimiiioii, m* it tnivi-lleil nhmg the
tube, gnt IcH niid Ici>e in amounl and itl lh<? «enie tira*^ Ivcciime more gradual
in Its development, Ihc curve becoming lower and more flattened out, until
in Uic nciglitiorhood of the artificial capillaries there wni> bnrdly iinr trac«S
of it left. In other words, we might trace out step by et«p tbe graJua] dia-
apmaranco of the pulse.
The same changes, the aame gradual lowering and Hattening of the curv*
may be seen iu natunil piil^e'trnciDgs, na for insunce in Pig. im, which it a
a tracing from the dorsalia pedi* arlenr,
ooiupared with the tracing from the radial
artery. Fig. 85. taken from the ean>e indi-
viduiil with the same tnalmnieut on the
same ocenuon. litis feature in, of course,
not ubvious in all puls^ourves taken from
diflerenl iudividuals with diHerent iti^lni-
ments and under varied circumtitauc-es;
but if a series of curves from diderent
Brtcric* were carefully tukt« under tlie same conditions it nouhl Iw found
that the notlic tracing In higli*^r and more sudden thiin the carotid tracing,
which, agnin, is higher and more sudden than ilic mdial tracing— the tibial
trat^'ing being in turn »till lower and more flnttcne<l. The pulse-cnrvc diw
ont hy becoming lower and lower and man? and more flntlenod out.
And a little consideration will show us that this niual bo so. Tbe systole
of the ventricle driven a quantity of blood into the already full aorta. Tbe
sudden injection of thi* ijunnliiy of blood cspauds the portion of tbe aorta
next to me heart, the part immediately ailjaccnl to the semilunar valves
beginning to expand lirst, and the expnneiou traTclling ihenoe on to tbe end
of this portion. In ihetsme war the expansion travels on from this portion
through all the sueoeeding portions of tbe arterial system. For the total
expansion required to make room fi>r the new quantity of blood is not pro-
viaed by that jiorlion atone of the aorta into which tbe blood is aelually
received: it is supplied by the whole arterial system; the old ijunnlity of
blood which is replaced by the new in this first |>ortion has to find mom for
itself in tbe reet of tbe nrterinl B|Mice. As the expansion travels onward,
however, the ijiergntc of pressure which each portion transmits in tbe suc>
ceeding portion will be less than that which it received thtm the nrecediug
porliou. For the whole increase of prt^uure due to llie systole of the ven-
tricle baa to be distributed over the whule of the arterial system ; tlie general
mean arterial preMura ia. as we have seeu, maintained by repeated sysiolof.
and any one systole baa to make il^ euutriliutifin to that mean prcMore : (lie
increase of pressure which starts l>om the ventricle nmst, therefore. Icavc
behind at each stage of its progrcM a fraction of itself; that is to Htr. the
expansion is continually growing lew, as the pul.w travels from tbc neart
to Uk capillaries. Moreovtir, wbde the expunnioi] of tbe norta next to the
BmK is. so to speak, the direct etfectof thvM-ntolcof the ventricle, the expan-
sion of tlw more diilaul artery i* thu cDivt of the systole transmitted by tbe
' the elastic reaclion of the arterial trncl betwMU Ihe bmrt am) the
auawnery: utd uaet this eladk mftimi in «Iuwer in i)i-\-eloiin)eiit tlian
lb* kUmI antole, ibe ezpanaion of tlie mmv dixtant bhit}' in tluwrr lliau
tkM «f 1^ lorta, iIm up^vlroke of the ptibe-curvc i» In* MiddcD, udJ the
«W« palK-«urT« is niure Hatti-iiisi.
n* object of tbe mtiilc b tn mpplr a fiinlribiili'>n to lliv mcBa prcaHir*,
uJlb* pubebui cwcillntioci aborc aiiil hi-lon thiit ineiiii prcfRun — im OMil-
lUiMwIiicIa (liininulm from tlic hnrt onminis hiding diimprd by the elARtlc
mill «f til* artrriiv, ■nd po, Uttlv l>y little, colt Tcrtcd itita mvaa pmaure
nXil ia tli« cKpillnms llic ibmu prasRura alone ramniiM — the oenllatioiw
ianne A'amppt^mi.
(143. If tn tikc mo<lel the {Hiints of ibo two lerere at dilli-nMil di»tniir«»
ftm ibr pump bv niaced cxiittly one under the other ud the rceordJng eiir-
fctt,il i« obviuuB tnat, the levcre being alike cxeept for their poeilion on th«
laht.ur difitreiK'e id time between tlie movenienta of the two lerera will be
^•n bv ui interval between the beginuin;^ of the curves they deiwribe, the
KMding surface bHng made to travel suliicieiilly rapidly.
If the novcmeuta of the lirn levers be thus compared, it will be tffio that
6t far lever (Fijij. sy II. i oommenc-es later than the near one (Kii;. So I.) ;
dc further a|)art ih« two levers are, the greater is the interval in time
\tdina their curves. Compare the series I. to VI. (Fi{[. K3). This nieatift
iiai ibt^ wave of espausioa, (he pulse-wave lakes some timt; to tmvcl along
l&(Ub& In ilw same way it would be found that the rise of tlie near lever
bnaaone ftaetion of a sccuml itHcr the .iirukc of the pump.
The veloci^ with which the »u]Hi-wavf tnivcis depcuil* chiofly on the
uwunt of risidily poaieMml l>y Itie tubing. Thv morv extensible (with cor-
npOodlM Mutio reactinuj tlio lube, the slower i* the nave ; tho more ridd
lkttnbebeeam«i>.tl)C ftster tbcwnve travels; in a perfectly rigid tul>r, whnl
btWflaftic lube would Itc iIk pulse, beeomes a more Ehoek travelling with
nrrgTMl rmpidity. The width of the tube is of much less influence, though
■RMttng to MiDG observere the wave travels more ilowlv in the wider tuliw.
The rate at which the normal (mke-wsve travels in tKe human body has
kwn vaHoti'ly estimated at from 10 to 5 metrra per Mcond. In all proba-
Ully the lower estimate is the more correct oue : bat it must be re«ieiiib«n(l
thit (he nilo may vary verv cooaiderably under iliflerent condiliona. Aooord-
ilgloall observers the velocity of the wave in pnariinii from the (jroin ta tlie
bat is gTftater than in na^ioK from the axilla In the wrist 1 1! metres againat
i mttnt). This U proiwbly due t.i tin- fact that the lonionil artery with ila
taBehw ia more ri};id than thu axillurA- and in bnmchi-^ So also in the
I of ohildreu, ll>e wave traveU mor>e slowly than in the more ri^d
I of the adall. 'I1i« velocity in al»o increiued by circuni;)tAiicea which
baijrtta and decretued by thinv which Icmiii the mean nncrtal preaanre,
met with incmsiiii; urcwun; the arturiiil witll.t become more, and with
dldRiilliog pTBMure, Itm rifpd. Pnitiahly. aUn, the velocity t>f lh<- ]>tihie-
•nv (lepeD<ls OQ ooiMlition»«f thearlfrinl wall*, which wecnnitot ndcciunttly
^Mribt u mere difTert'im* in rigiililr. In exiierimt-nliug with artificial
it ia found that different c|tuilit>cs of Indui-rubbar give rno to very
mulu. '
I muM bo ulccn not to oonfmind the progms of ibe pnlte-wnve^i. e.,
rf tha fXpanioD of the art^rrial wal Is. with I he actual onward morenu-iit of
lb btum) iunl^ The pitW-wave travels over llic moving blood somewhat as n
npidly moving nattinil nave travels alone a iluggi»hly flowing river. ThiM
iiila tfa« velocity of the pu)se>-wave is 6 or powibly even 10 metre* per
1, that of tho current of the bhKid is not more than luilf a metrv per
I vren iii (he lar^ arteries, and is still Ices in the smaller ooei.
234
THK VAflCt7I.AB VKCllASlt
^ 144. Referring ojritin Im the ntulinn a'ivcn above luit to r«i;nn] lUo pulse-
curve as a picture af the puliK-wavv, we miiv new sdi) that llio imlMr-wttve b
«r very conaidentble U-tijcili. It' tv'i; know how luii)[ it iakir> lor the |>ulse-
tritve tu ptaa ov«r iidv [H>iiil in thii nriorit^ luid huw fam il i« travel I i tig. w«
can euMJIy calculate the k-nglli nf the wave:. In an (inliitarr jiuUe-curve the
arlerj', uwiiig tn iho Hlnw return, i» nrrn nut to rc;(aia tliu valilirc which it luul
before the nxrianbtion, until jii»t iu> ihc wxt vxpauvion t)e^iis — tliat i» to «j,
the pubc-wavv tiikm the whole time of n uunlinc cycle, vii. : -fisiht seoond to
paw l>y ihtj kvi-r. Taking the velocity "f llie inil»-wnvc a* 6 metere j>w
■econd Uie le4>gth ofthc wave will be /slhs uf 6 moirea — or nearly A metraa.
And even if we took a »iaaller estimate, by enppo«ine that t4)« real ftzpaa-
flion nod return of the art«ry at any point Inok much lew time. t*y tV^i'
Mcnnd, the length of the piil«e-wave would still be more than 'Z nieLn». But
even in the tallest man the caitillarieH furthest from the beni't, thoKi in the
ti|w of the toes, are not '2 melrefl distant from the heart. In other woril^ the
Mlgtb of the pul»e-wavL- is much greater than the whole length of the arte-
risTeyBtem, bo that the beginning of each wave has become lust in the Hmnll
arteries and caiiiltariea some time before the end of it has finally passed away
from the beginning of the anrliu
We muBl now n^tuni to the conaiileration of certain special feaiiires in the
pulse, which from the indication)! they give or auggeat of the eundilion of llie
vascular system are ol\uu of eri^at iniereat.
049. Ditrotism. In nearlr all pulite-tracinga, tbe curreof tlw expannoD
anil recoil of the artery im broken by lito, tiiree, or several anialler eleraliona
and deprea«ion»; «ecoiidurv wavin are imposed ujun the nindnmeuta) or
primary wav& In the spWgmogruphic tracing fVntn the carotid, i-lg. 87,
no. a.
rvi««.TNACi)iw raoK C*aonii Aktieiit or IIultiiv Mas (ftom HonW.
a, DDmnwoconiMI of upuulon oTBrtcrr. J, lununlt oftlieDninte. C, dlcnU; teconOur waT«,
Aii^dluMlaMiMadaTT war*: p.Qouh prtetdlnf tliliL D.'upnodlimiMaailuy unTa. TtMimtva
■bnali llMlof ■ nndnpDHt with un daulile TlbnUon* Id B Mooiiil,
and in manv of ibo other tracings given, thcM cecimdary elevations ar«
tuarke^l, as /{, f, />. When one such seoondarv elevation only is con^icu-
ouR, so that the piiUe-eurve presents two notabto creets only, iha primary
creit and a aeoondary one, the nuUe U said to be "dicrotic"; wnen two
wcoodary creats are prominent, tne pu Ise is often called " tricrotic " ; where
Mrcral, " polycrotic' As a general rule, the secondary el evatinns appear
only on the deacending Hnib of the primary wave as in mml of the curves
given, and the curve i* then spoken of aa " katacrotic." Sameliine», how-
ever, the Srtl cl«vniion or cns<t i* not the highest but appear) on Ute aicend-
tng portion of the main curve; Hiii'h a curve in apakeu of at "aDacrotio."
or tbeae aeoondary elevations the moM freiuent, eonspicuoua, and impor-
THE PULSK.
285
iMt b ih* one which uppeara w>iii« nj down on the dewmding ltntl>, nnd !•
Mrbd C on Fix- ^~ itnd an mcMt uf the curvw hera given. It in nion: or
luiJiitiiiiHr viiuble on all sphysmogmmf, and amy be mnmi in (Imjuv of the
MTU ■* well ss of oUier itrtcrica. SomeliiDen it is m slight «< to be hnr<lly
_i[liiilJhlr U other timos it muv be so marked «a to givp rim to a roally
■Ut |iulM(Pig. 89), !.<;, ■ puW which oao )m fHt as dotible by the
f\.
n^m.
Vta
L
M- M-Aitnii>Tii^Vaii.H'Mii 41-11 Thvctiu ri-'U nil Mcwii!!!) AaltL lAniurtaa.)
fta ■ -T*I> U*4liD or MlBKIl' tlll-IKITBN I-I It •IIIIIL Puw OT Man, «TfTlloi<l (ClVf,)
hfif ; httMre it has been called the tliervtu^ ok'vutiiiii or the dicrotic ware,
t Dotrh pr(>cr>]ing the clevntioii hv'we spoken of as the "dicrotic notch."
Abfr it nor any other secondsry mvatioDi can lie raawnbed in the
Mitft of blood- presBu re taken with a niaimmeter. Thi:* mnr l"> rxpliiitied,
iVt have said $ 13!}, by the fact that the tnovcincnis of thu incrcunr
Ama are too BlufEgisb to reproduce theee liner vanutione ; but dicmtiim t*
in compicuoiM by its ahMOce in the tracin){« ^vcn by mora del iceloty
'■^fpnttve iiMtmmeiila. Moreover, when the normal piilm? i» fcU by the
V. Ridsl persons BikI theiuselvei imabk- !■> deloel any dicrollnm. But
it (lo«« n-allv exirii iu the nunnikl pulse is ahonn by the ftu-t thai it
toparp in a iaa*t unmbdukahlt^ manner in the tracing oblained bv allowing
nt UojI to afMirt dtrm-tly fr»in an ojiL-ncd tmall artery, itucb as tKe dorsalia
, friv, upon a rrconling lurlacc
Um runstwit and consplcuouti than the dicrottP wrav4>, but vet apfteering
'* I BMt •phvgR>ogr*inR, ix an devntiDn which appear* hi^lier up on this
jBoiIine hmh of the main nave ; il in markeal R in Fig 87, and on several
rda other ciini-a*. and i* frequently called the pre-dipro(ie wave ; it may
tkuBW very nromiaent. Somctiinoa other atoondary wavr*. uneii called
"fatdienitic. ' are »eeii following the dicrotic wave, a* iit D in Fig. ^7, and
■■elher rurres; but thcM aronot ofttn pmH'nl, snd imuully, nven when
pMnl, iDPomptFU04i».
When tracing* are taken from mveral nrlcriea, or Irom tlie same artery
tadir dtfltont coDititionn of the body, thcM aeoondnrT waves are found l»
nij Terr oMHiderably, giving rite to many dwracteristic ibrma of pulse-
eirrr. Were we able with certainty lo trace hnck the wveral featurcaof the
111 their rmpective uiuses ; an adequate examination of sphygno*
hie tracings would undoubtedly disclwe much valualde infornuitiflti
cnuiig the contlitiou of the body presenling ihem. Unfortunately, the
pnU«n of the origin of these teooDdiiry waves is a n>oet dilBcult and' oin-
fbienv: so mueh ao, that the detailed interpretation of a sphygmograpbic
Inaojt b still in n»<itit coses extremely uDoertain.
\ m. The chief intenst attaches tu the nature and meaning of lite dicrotic
•atn In general, the main cotidiiiuns favoring dicroiimu are (1) a highly
■ niwilili and elastic anenal null, (2) s\ cuminiraiivcly lnw mean pressure,
letfing the oxtensilde and eluAlic reactimi of the arterial wall fre^ M>opo to
act aiM (3) ■ sufficiently vigorous and sufficiently rapid Uroke of the ren-
Hiek. 'The developmait of the dicrotic wave niay prol*ably be explained as
Mloin:
266
TillC VASCtTLAit UKCltAXISM.
At eiicli heat the lime durbijc which Uic coiireuu of the left v«>Dlr
iDJei'leil iulo tbe aurta is, ai ive have neeti (^ I3ii), verr hri«r. The ex]Mii-
MOD of the aorla u very ■ik1iI«u, tunl thu oenMliou of tnat cxpannoa U aln
very wuddeii.
Now, wh«n fttiul Li b<,^iig driven with own > Rtciiilv prrK^iire through an
^Ulie tube or it .■'viitcim iif diutic tiilMW, Icvora plucoii oil thi- lubo will
detcribc ctirvc ludiniting vnriiitionsin Iho (linmc'Ierof tholtibo,)f iheinHow
Into the tube be Hudiluiily slo[i|io<l, lu by Hhnrply turning a stop-cock ; and a
cotnparisoR of IcveTS puiood at JiffOTeiil distancea from the stop-cock will
show that thcxe vanatioas of dtamel«r travel donm tJic ttibo from the stop-
cock ill the form of waves. The lever near t)i« stop-cock will first of all (all.
but Bpeodilr begin to rise again, and this subsequenl rise will be followed by
another thil, afler which there tnnj be one or more siiccceding rises and fnllft— •
that is, oscilliitions — with decreasing amplitTido), until the fluid comes to rest.
The levers further from the ston-cock will describe curves, similar to the
above in ibrm but of lesa amplitude, aud it will be found that tbeee occur
nmewhat later in time, the more ^ the furlher the lever is from tbe stop-
cock. UbviouHly these waves are generated at or near tbe stop-tywk, am)
tnvel thence along the tubing.
We may infer that nt each licat of the heart aimilur warea would be
frenerated at the rmit nf the uorta u[>on the xudilen enuuiliou of tlie How from
tbe ventricle, and would travel theuee along the elantic arierie-. Tbe fai^
liiat each beat in rapidly nucci-eded hy another, ntiil that the How which *ud-
denly ceases ia aliu, by the uutiire of the ventricular stroke, siiddvnly mt'
ented, may render tlie wavia more complicaUid, hut will not changv loeir
«wentinl nature.
The exact intenirelatinn of the generation of th<'«o waves b [wrhaiM not
without difficulty, but two factors seem of especial importance. In the firtt
place, as we have already more than once srtid, when a rapid flow is suddenlr
stopped a negative pressure makes its appearance behiud the column of fluia.
In n rigid tube this simply lcn<ts to a reflux of Huid. In an elastic tube its
elleci^ are complicated i>v the second factor, the elastic action aud inertia of
the walls of the tube. Upon the sudden ceMuiliou of the tluw, the expanMoii
of the tube, or as we may nt once say. of the aorta, ceases, the vensel beg;iiia
to shrink, and the lever placed on it» waliii. tut from .1 onward in tite pube-
curre. This abrinking i« in part due to the etastii- reaction of the walU of
U)e aorta, but la inereaaed by the " suotion " action of (be nirgative preM^I
apokea uf above. In thus shrinking, however, under theac cumbin*^ cwimH
tne aorta, through the Inertia of iltt walU, nvenihoots the mark, it in carrietl
beyond ita imtural calibre — i.e.. the diameter it wuuld ]>(MU»ti if leli to itcclf
with (ho prcMure innide am) uulcide euual ; it i<hrink;i ttiu much, and ooitae-
•e<)ueii(lr hcciiu ngnin Ut expand. This sucomlnry expansion (taking for
simplicity sake a pulac-curve lu which the so called prc-dicrotic wave. B, is
ftbaoat or inconspicuous) ciui<w« the necoiidarv riw of the lover up (o C— tbat
IB, the dicrotic nse. In thus cxgnindriig again, tlie aorta lends to draw bock
toward the heart the column of blood which by loss of momentum had come
to rest, or, indc«d, under the intlucncB of the negative prcasuie s))okeD of
above, was already undergoing a roHiix. In this secondary expansion, more-
over, tlie aorta is oy tbe inertia of its walla, aided by that of the blood, again
carried, so to speak, beyond its mark, so that no sooner has it become ex-
panded and filled with iluid to a uertain extent than it again begins to shrink
ae from Oooward. And thi» .shrinking may in a siintJiir luantier to the firvl
be followed by a furllier expansion and sbriukiug. giving rise to a poctt-
dicrotic wave, or it may be to poet-dicndic wavei. And the »ucoeMn\-o
changes thus inauKurate<l at the root of (be aiirta travel na »> mniij
TOE PCL9K.
237
riH|(tte utorUI ayatvin. iliminlshiiig H tbrr go. It irilt lie olwcrvvd Uinl
brtSFJavclotmient uf the«- wave* n certain quality in lh« walk of Um tubing
■ iwrrwary. Ttu> tnlw niiul ItPHichaspnanPwrawIiviint mtauop«n liiiDen;
tkrvaUi niMt be '>f Huch a kind that tii« tube romniiw open wben emptv —
i t, ■bea the atiiuwph«rlc prcautv i« equal inxiilo and outside — «K tlint w)i«o
il thriBkl too mtlcli it expands eigain in stiiving li> rc-;!Ain its natural calibre.
1k> *e bavt* Men to be a cbaracteristic oF the arteries. A (x>]Iapaib)e liitie
•f lUl BWQibraiie will not »hi>w the pbetionicna; Hiich a tube n'beii ilii- iiu>j>-
Mcfc ■> tarunl i-ollaieea and ciiipttea iifeif. conlinuiD^ to be collaftfieil witli-
gd >iiT diort tu ex iMud aj{ain.
Is llie above ex plaitaliun no nM-ntion baa been made of the cloaiue of tlie
•MiluDar valvro ; we almll Imve lo >.\vak of these a little later on in refer-
ni| lo the pr^v^licnitiv wave, ami biiall M'e tliui, nmler the view w<- have juat
|i(*ii, tho clgting uf lite fieDiilitnar valves is to be regunli-d rather a» the
tttCt tlMn iho cauw of ibc dii-mtie nave. Manjr auihom, however, givo no
hRnntation of th« dicrnttc wave diflerent from tJiat deliiitol nbiive. Thus,
kbliabl tlifll ibo primarv sbrinlciiig I'runi A onward, being brouglit to hear
MlWcolamn of blood already come to nat, in face of the great prcwure id
fiwl.driim the blmid Imck ngainst the *eniiliinar valrcw. iliiin cimiug them,
ni that the imiMCt of tlie nilunin of IiIihhI ajruiiist ihn valves tttmrie a new
nn of cxpanawD, which relnrorcintr ihn natural tiniduiicy of the ehutU-
nib to exgiand again after thi-ir pritmirv nbriuktng, pniduen the dicrolie
■m C On this vi*-w, it i» lli" blowl drivi-n baek from ihr valves which
•ifMik the arti-ry ; on iIm- view given nlmvc, it is iho oxpaiiding artery which
4i»s tlic hI'X"! twck tinrnrd ihr vnlnw,
Uonover. quite other virn» hnvc been or arc held cuncorning this dicrotic
nra Art^>^dinK to many nulh'irv, il is what i* ciillcil a " reflected " wave.
Tbw.*)ien the lube of the artitieial luodcl bearing two levers is blocked just
kinad ibeiar lerer, lbs primary wave beeen tobeaooompanied byasccoiMl
aant which al the &r kver b seen doee to, and often fiiaed into, the primary
ant (Fig. f*3, \'I. <■'), hut al the near lever is at some distance from it
(Pi|.83,l. i'). lieiu^ the further from il the longer tlie interval b«tnt«n the
mm aaij the bli>ck in the tuht?. The second wave is evidently the primary
••*« rtAeded at the block and imvelling backward toward the pimip. ft
tkm. of eouiae, (MWCa the far lever liefure the near one. And it has been
m^Kii that the dicrotic wave nf the tmliw is really audi a reflected wave,
Mulnl riilM-r St tlw iDiniite arierius and i-apillariea, or at the points of bifiir-
Bttiwof the krger arterie*. and liavidling backward to the aoilA. Dot if
lib mr^tba c»V. the ilistanci^ l>ctwe(-n the primary erMt and the dicnitic
<pM oaglit to be Ion in arteriei more diitunt from Ihan in those nearer to
ibtbmrt, JDSt M in tbo Hrliltcial •cbenie the ri-tlecteil wave is fused with a
|*iury wave near the block (Fig. 83. VI. •> a. a'), but bcoomes more mid
HR trparattd from it the l\irtlier back toward ihv. pump wa trace it i Fig.
I3,L 1 ir. ')'). N»w. ihi^ is not the caae with the ilicrotic wave, i'areful
iimneiiis show that the distance between the primary and dicrotic crtata
■ ■tier grCBter. or certnioty not Ims, in the xmnllcr or more distant artvrice
tbn in the larger or nearer ones. This feature indeed prnve« that the
difMie wave cannot be due to reflection at the peripherr. or. indeed, in niiy
nf ■ Teimcrade wave. Uendee, tbe multitudinous peripheral division woul^
ndtr ooe large periphically rejected wave impooible. Again, the more
^iQy the primary wave is obliterated, or at least diminished, on its way to
■b peripbery, the las conspieuoiis should be the dicrotic wave. (lenoe
bunwed exictnubilily and increased elasUc reaction of the arterial walla
■dick tend touaeup rapidly the priiuari,- wave.ahould also lessen thedtcruiie
■ \>v;i.AR MECHAX19M.
;■ 1 -..jt::- 7 : raa th«e conilitioiis, as we havD said, are favnr
..::■-. : ;hr liiiTniii' wave.
■< . rluit? sv,<i till' other conditions which favor dicrotUin
i:» \;v.".,'. :."..#<,' whidi would favor such a developmeat of
'.-.- 13 !.i» '.-t.>:: di'^'rilKil above, and their absence would be
.I !.■ ■; -t^ka; '.I'sui'h wuves. ThiiH dicrotiaiii is Icaa marked
v- -..' ' 1?:-''^' of old people) than in healthy elastic ones;
. :., , wa-j ■> so naUilv nor shrinks so readily, and hence
1,1.; ■ ,■ '. -■* '■? siu'h s*fondary waves. Again, dicrotism is
. 1 ■■ I .. -['v.i', irtt'rial jiresiure is low than wheti it is high;
, .. . ■ ,*. -1 - : uvd wheu absetit, or increased whoii sliebtly
; -.■ , i; ' rf way or another, the mean prusnure. Row,
v.. • ■> : ^'1. :'.••: iruries are kept continuallv much ex]Hinded.
.• "< w .-n:M.'i; 'f further exiianeion ; that is to say, are,
....1. ' :<iu'- !ii ::ll:^ii[i<.>naI expansion due to the systole is not
. ,1 > t <•« ^-ll■:■.^^'v tor the arterial walls to swing backward
•i. 1,,. tii.t nu^f a Itss tendency to the development of
., -. ^^ U.I ii- mail pn'ssure is low, the oppusit« slate of
- ..., . x,,i., 1 ■.■iir>f. chat the ventricular stroke is ade<juat«iy
. ,»»..- hihl; in, n.'[ to diminisherl cardiac force, but to
. , i-i:i;iiv . : !n' rvliitively empty but highly distCD!<ible
> I ,.,..:i.i, i;ni, -iiiliti,: r.ipjdlv back, eiitci^ uj>on a secoDtl-
1, ,. ... <...! -ivii .1 third.
. .....< 'I 'ill <,.ii> :ii.tv be applied to uxpluiii why sometimes
., ^,. ......^'.i 1 .1 :'arti<.'ular artery ubilG it remains little
■. -!■ 111. [u t'xiierinieuting with an artiticial tubing;
. . , ■■ ,.\'.. i'lsical I'hnrapters of which remain ihe
■ ■. •.--tiA-.y .uid the secoiidury wiivw retain the same
..ii;; :■■-■ tubing, .-avf i)nly that both gradually
,..,..1. i> .11!. 1 ill the natural circulatiou, when the
. . „ ' I I :.':iii ihroiigbuut, the jiulse curve, as a rule.
.... ., ,i..-K-;irs througbijut, save that it is gradually
,.-,-.■ »iiv to mtbsiiiute for the first section of the
..,.-.,'■ ■-■.■ -.iliiui;; this at the stroke of the pump, on
.,., i..„ ■! -Ii.'w neither primary nor secondary ex -
. i:.., .■;..■ .'[' the pump's Stroke Would be transmitted
;,,.i . Mvti.'ii. and liert the primury and ^coiidary
• .'\ '.it'Lti. This is an extreme case, but the same
,. . I 1 -.■. iii;ri.t' ill putwiiig from a more rigid, that is,
, . . .11 .i.iir. !.. K le^s rigiil, more cxtc-nsibic and elastic
i 1.. ^,.i;irv t<xpansions, in spite of the general dauip-
,.,..% ;i, i.-iiac. Similarly in the living body a pulse-
> .I'liu'Iling along arteries in which the mean
' ...1..1 !■.■ ilivivlun.' practically somewhat rigid, is not
. . '.,, [III.' wry markedly dicrotic when it conieij to a
. .. ....] .1. 'iii'iiii pressure 1:; low (and we shall soe pres-
, I.. 1 1 . iiiid the walls iif which arc therefore for
. , > ,. ,.' .i'..>[vii-^il>lt.' than the rest.
. . I. 'i i.-.i'1'viiiion made above (^ 141 ) that the curve
■ K- :.'■ nniiilii'd by llic jtressure exerted by the
. . , i;i,i ib:Lt hcni'i', ill the saniL> artery and with the
. .. ii.ii.. and cvfti tilt; sjiecial features of the curve
n. .i' |.ivs.*ure with which the lever is pressed
V ;:._,^ \w amount of dierotism apparenl in a pulise
...>..!!■ i\trted by the lever. In Fig. 6'>. for
TUB rULSK.
I
I
I
iiMlnnn!, th* dlentie wnvc 1* nxtre erident in tbo mUMl* tlian in tbe upper
1 147. T\w pro-<llcrotic wnvi' (mnrkfr) !i on Fig. t^7, and on wveral otlier
flf lhi> puU«^urva>, which prt-ccdi-* Ihc dicrotic wsva and is still nwre
vvriililr thxn thai vfsvo, Ih-idi iu)mciimc» slight or even invisible and some-
tiuMi onoRpiciiDUH, Una given rue Ui rauch conirovenir. Id th« iiitorpretati»n
of UiN ilicnitiu ware givvu io Uto prccMliQg paragraph it was staled that the
nrgntivn prrMur^ 'kvcluj>rd on the wsMtiun of the How in the rear of the
cohmtti of hlix)d, led by itMlf (o a reflux toward the veniricle: nni) it ha*
been suggeet^ that at this reHiix mcclinf; and cloeiuir the seniiluoaf ralvc*
■tKfbt n RDiall wave of expansion b«t<>re the larger dicrotic ware bu hwi
tinKf Io develop itself, On this view tbe nniiliitur vmlvea wouhl be actiinlly
cIimmI l>ef<ife the oecurreflee of tbe Mcondary dicrotic expansion of the
artehnl vinlls, thixiKb tbe larjcer, more powt^rfnl reflux of tliio laUr event
iDiut rvtider tbe cliMure more complete, and in doing ta [ximitilv givra riee
to tbd Moond sound. Accordini;. Iiowerer, to lite tecond viviv glvat in the
Mne Mimgrapb, which reKanbi (be reflux due to the vjiriuliing of the
mtlery in face uf (lie ^ratt prt-Mttrc in front art firmlir chMine Ibe soniilunar
vmlvet, uxJ an ibtu startiii); tbe scctMidury dicmtic wavu ot cx|Mnsion, itie
firm doaiDg of tbe temliuniu- valve* nust Uike place before the boginniug.
iMt (Itirini the developmeul of the dicrotic wave ; il is mill pomJhlc. however,
even on tnie view, m od the other, to iiU]>piN(C that an antix'cdeDt rvflux, due
to the negative prenure wci-oeding the ucmatioii of flow from the ventricle,
doMa lb« va]v4« and slarti tbe p^^di<'rl>tic wave. But the matter is one not
ytl bcvrind the (tage of controvervy.
g 14&. Ill ao anftrcrotte yulu- the lirat rise is not tbe highest, bnt a second
rise t II. Kig. aH) which follows aiul in Mpnnitcl from it by a notch is hif[hcr
Umn, or at lewt aa high, aa llaalf. Sucb an aiiarcrolio wave, tJiougfa it may
MMiMtina be proiluoed tem|H>rHrily in healthy persons, is senerally ama-
dhtted wltb disnued conditions, usually such in which (he arterieB are ab*
noniially riKid. In dt-Hi-rihing tbe vctilricular aysiole, wc spoke of tbe
iinwHirv within the vt-otriili' as reaching its maximum jnst before the open-
Dg uf the seniihinar valvcn ; and ibis is apparently the uuruial event ; but
then are <:urves which seem to show Ihai after Ibe linti sudden rim of
prawuru wbi«b opeu» tbe valves, followed by a brief lenening of pniKuro,
which appears on (be curve us a notch, the pr««ieure tuay again riae.and that
to a |Kiint higher than before. AihI a similar curve is xoinetimed deecribed
by tbd frunt-to-back diameter of tbe ventricle. The syslule o|ieiu tbe valve
■• it wvtt with a burst ; thia is followed by a slight relajxto. and then the
systiile, strvDgtbeuing again, dischargee tbe whole uftla' ventricular contents
inlii tbe aurta and su bringti about a tardy maximum exiiiuiaton. And what
is tbii* started in Ibe at*na travels onward over the arterial Mvstem. It is
ditficnlt U> »ce bow tbeae anacrotic events can lie priHluceil, except by a
certain irregularity in tl>e ventricular systole; aiHl, indi-cd, the iinacrotic
pulac in rn-i|iii:ully awuciatitl with some disease or defect of tlu^ ventricle.
( 14S. I'l Hom fiuhf. l-'nder ct-rtHin i'iri-umstance!< the puW may be car-
rierl on frtmi th<i arleriiii thn^iKb iIk- CHpilliirii-x into the veins. Thus, as we
•ball see Inter •>». whi-n iht- Hnliviiry ^^liuid ia aeltvcly K'crcting, the blood
nmy iw>ir from the gland through iIh; vcini> in a rapid pulsating stream.
T]>r nrrvoii* vvcnis wbieh give ru« to tbo secretion of stilivii, lead al [be
tame time, by the agcnry of vasomotor ocrvce, of whieh we will presently
•peak, to n dilatali'iii of the xmall arli-tii-a of the gland. When the gland is
at rrst tile minute arteries arv, u wc sbnll kc, sonicw but constricted and uar-
rnwnl, and thus contribute largely to the peHpberal resistance in ibe part ;
tliia iwripberal recistaaco throws into actio«i the elastic projwrtkti of tbe
240 THE VASCl'LAR MECHANISM.
email arteries leailing ti) the gland, and the remnant of the pulse reaching
these arteries is, as we Ijefore explnined, tinally deatruved. When the minute
arteries are dilated, their widened ehanuets allow the blood to flow more
«aidi!.v ihrouj^h them and with less friction ; the peripheral resistance vhich
they normally offer is thus lessened. In consequence of this the elasticitv of
the walh of the small arteries is brought into play to a less extent duu
before, nnd these small arteries cease to do their share in destroying the
pulse which comes down to them from the larger arteries. As in the cue of
the arliHeial model, where the " peripheral " tuning is kept open, not enough
elasticity is brought into play to couvert the intermittent arterial flow into a
continuous one, and the pulse which reaches the arteries of the gland paaKS
on through them and through the capillaries, and is continued on into the
veins. A similar venous pulse is also sometimes seen in other organs.
fureful Inieings of the great veins in the neighborhood of the heart show
elevations and depressions, which appear due to the variations of in tra-cardiac
(nuriciiUr) pressure, and which may, perhaps, be spoken of aa constituting
a " vcuoua pulse." though they have a quite difierent origin from the venous
pulse just descrihetl iu the salivary gland ; but at present they need further
elucidiition. In casea, however, of insufficiency of^ the tricuspid valves, the
systole of the ventricle makes itself distinctly felt in the great veins ; and 1
distention travelling backward from the heart becomes very visible in the
veins of the neck. This is simiettmes spoken of as a venous pulse.
Variations ()f pressure in the great veins, due to the respiratory move-
ments, are also sometimes spoken of as a venous pulse; the nature of these
variations will be exp1aiuc<l in treating of respiration.
The Regulation and Auaitatios ok the Vawui-ah Mechamsh.
Tile Jte/jiihitioii of the Beat of the Heart.
§ 150, So far the facts with which we have had to deal, with the exception
of the heart's heat itself, have been simply physical facts. All the essential
phenomena which we have studied may be reproduced on a dead model.
Buch an unvarying mechanical vascular system would, however, be uaeless to
a living body whose actions were at all complicated. The prominent feature
of a living mechanism is the [>ower of adapting itself to changes in its
internal and external circumstances. In such a system as we have sketched
above there would be but scanty |><>wer of adaptation. The well-constructed
machine might work with beautiful regularity ; but its regularity would be
its destruction. The same quantity of bloo<l would always flow in the same
steadv stream through each and even' tissue and organ, irrespective of local
an<l general wants. The brain ann the stomach, whether at work and
needing much, or at re.-^t and needing little, would receive their ration of
bloiwl, allotted with a pernicious monotony. Just the same amount of blood
would pus.' througli the skin on the hotten as on the coldest day. The
canon of the life of every part fur the whole period of its existence would
be furnished by the iiihoni diameter of its bloodvessels, and by the unvarying
motive power of the heart.
Such a rigid system, however, does not exist in actual living beings. The
vascular mechani.«m iu all animals in which it is present is capable of local
and general modifications, adapting it to local and ceneral changes of cir-
cumstance. Tliei* modilicalions fall into two great classes:
1. Changes in the heart's beat. These, being central, have of course a
general effect ; they influence or may influence the whole body.
241
»
1. Oiaii|;» in llie peripbentl rewlimre, due to variations in tlie celihn- of
iji« niioiite arteri«v, lirouybi about by tJie ageucjr of tbeir couirarlil« luiiv-
itiUr coaU. TIkm cbangai mny be dtber local, aflbctine a uarlictilur
vuctilar area only, ur genenJ, atfiKUiig all or nearlj all ihe Dloutlva»els of
tbe body.
Theae two da«e8 of «v«dI« are cbiefty governed by (be ncrrou* tvMeni. It
)• by ineaiia of lli« ii«rvoU)i •rctcm thiit tl;u liciirt'* Ixtul imil the ciilihrA of
tbo minute oriemti are bmiigbt into n-lntion wiili ciicli otber.eixl with almost
vvorv [«n of the IkiiIv. It ii by int.<nns uf lliv nvrvous «yatviii scliog ciihrr
Ob tM heart or ud tbe »inal) iirt«rM«. or on IkiiIi, tlint it change of drcnm-
MaocM Bfieting citiwr the whole or a port of (hi- lioiiv is mvt by c^nijiciwitinj;
or ngulativc cliaiigci> in ibe (low of blood. It i* by niiL-»nc of tbi- nervous
tjfUim ibal an orgnn Ims a more full supply of blood when at work than
when at net, that the lide of blood ihrcuch lli« «kin rises and ebbs nilh tbe
rift and (all of the (eniperaluTe of the air. that the work of the heart la
tctppered to meet the strain of overfull arleriee, aud that the arterial gates
• ipTD and Hbut as ibe force of the central puni|i waxee and W8u«8. The
study of these changes ljecoiiie«,tbei«fore,b> a iar^e extent a study of uervoua
art inns.
The drcutalioD mar abu Iw modifled by events not belunj^n^ to either of
the above two classes. I1iu», in thin or ibat peripheral area, obaiigivi in iJie
capillary walls and the waltn of the minute arteries and veins may lead to
an increase of tlie tendencv of thL- blond cirouuclea to adhere to Ih*! vu.-H'nlar
walls, and to, i|uile apart from any diauj/e in the calibre of tli<! blood v<rtte Ik,
may lead to increase of ihi^ periplx^nil moxtttnce. Thin is wen in un vxtreme
cKM ill in dam mat ion, Inti may [Hjwibly intervene to a Uw extent in the
nnlinary mndilion of the circulation, unit niuy al»o be under the iuHuence
I ihu nrrvouB system. Further, any deciilwi chunge in lb* quantity of
actually in circulation must aUo inrtiience thir working of the vascular
ninisni. But both lho« phnngcs arc iinim|H>nant compiirod with the
two kiiulfi of chiin);t'8. Ilciice, the two nn»l ini{H>rlHnt problems for
us tit Mudy nr^'. 1. how the nervous system regulates tlit^ bent of the heart,
anil 'J, how lid' iicr\'uu!- system regulates the calibra of tbe bloodv^sels. We
will finst cNiimitler the former problem.
<i 16L It will be netmaary now lo take up certain points tyiiiceniing the
minute Hnicture of tiM heart, which we had previously jxwtjMtned ; and
iinre much of mit knowlcilgt^ nf liie nervuus mechanisti) of th« b<«t of the
benrt i» drrivtsd from expcriincntH on tbe hearts of cold' blooded nriimiils,
iiiof« particularly of the fru^. it will be dtwrablc to consider tbew as well as
the tnatHtnalinn hi-«rl.
t iir<lt<tr iiMi'fulnr tU'iir. The vcntrii-le i)f the frog'n heart is composwl of
miiiolc i»j>iii<n<-KhB|>ctl tibn« or fihre-oell*. vuch i-oniAining a uiiclciis tn its
middle, mid tapering i<> n pout itt each end ; Koim^timiit, honevcr, tbi.': end is
forked or even branched. Th«Me filtr>^ «r tilinf-ocll*. in fact, resemble plain
eular librt» save that ibey are somevrhal iarvvr and that their substance
Wriatnl. The strialion is duo, like the striation of a Ktrialed muscle
t. 111 alternate dim and bright tHtuds, hut is rarely so distinct as in a
rial rtbre: it is ver}- apt to be obscured hy the prownce of dispersed
ilinlinrt granules, which, in msny cases at all uvenle, are of a fatty nature.
Like th*- ])lai» oiutculur fibre, tb« cardiac muscular tibre has no dtetinct
Mifi'i'Ieiiiiiia.
242
TIIK VASCUIiAK UBCUANISH.
A ntinilwr of tbew fi1ir«a are jniiied br oenwnt Bill»Uui«e infi tinuJI
Uiuillt«, aiid iheee bundlefl are, by cbehelpot connective lisBue wbicb cam»
DO klocMlveawls, woven iuto an iiitricale network or B[ioDge-woric, which
forn» tli« greater part of tbe wall i>l' Itie vealricl«. Imnitidiateljr undvr At
[WricKrdiM coating, oooMatiug of a layer of e]>illi«li<:iiil {ilatea raUng on a
fioanectivii-tinue aiaa, tbe mutcalar tbsue fornu a lliin coiitinuoua ■baet, boi
wilbin thw it Kpnads out inta a sponge work, tbe n>e«beM vf wliioh pniwDt a
Iiibyriiitli of i>iuwik«« voiitinuouH witb the caritjr of the vi^niriclr. Tbe baa
of this spoDge-work, varying in tbleknea* ani], ibougb iipimrvtitlv irrvgular,
amogea on a definite tvMna, oonmt of buudl«» or miiKuUr lil^rw united
br oonnwtive tJMiiv, ana are coated with tho itanic endocardi*] racmbnuc
(flat epitheliwd plitt<'« ratling »u a connective- iimuo bam*) that Htm tbe
cavity of the ventricle nitd, indeed, tlw whole interior of the henrt. Thr
cavity of the ventricle, in other wotcIm, opon» out into a Inbyrinlh of jwii'Mgiii
reaching nearly to tbe surface of the ventricle. Wh^n the veniriele ■
dilated or reluxc<d, blood flows freely into am) fills this Inbyrinib, batbing
tho ban of the sponge-work, whicli, in the a)«ence of capillBriea, de|wnd oo
thu hloud for their nourishment. When the voiiiricic contracto. the blixvl
i* driven out of this Inbyrinth a« well as out of the central cavity. Ht-iior.
the ventricle wbon dilalod and full of blood is of a deep red color, «hi-ii
oontracled and empty is extremely pale, having little more than tbe color of
thr muscular fibres themselves, which, like siriated fibrea, puMcsa in their
own aniMiiance a certain amount of hicmoglubiii or of myohiSfnatiu.
The much thinner waits of the auricle OoiuiBt of a much Ibiiinnr network
of similar librcM uuiit-d by a relatively lat^r quantitr of coniKclive ttnue
into a thin sIhwI, with the pericardial membrane on tbe outside mod tbe
cadocnnliHl metubrane on the inside. Tlie libra* bnv« in the auricle a much
groHtor tendency to be branched, and many, oaaning to be tpindlo^baped,
become nlmoxi tttctlale. Among the obwurcly nLriate<l but atill striated
fibres arc found ordinary plain mu*i!ular HbnM which incrcaie in relative
ourabor along the rmitii of tho veitis, rimw c«vmi, and pulmoiialce, until at
80HM litth' dintancv from the )it-jirt jilain muscular 6bres only arc found.
Bloodvenots are absent from the walla of tbe aurieW aW,
In Um bulbil" nrl(■^i•uu^ mixed up with mtii-h conncclive and elastic
liasue, are found fu*ir<irii) fihro' which clo'e to the vcntHole >m striated und
form a thick layer, but at A certain distance from the ventricle luee their
atrialion. or ratbor become miied with plain muscular iSbree, and form a
thinner layer.
$ 1$8. Ju tbe namnal. both the ventricles and the auricles are formed of
buiHlIeaof muscular tissue, bound together by connective tissue, and arranged
mon wpeciallv in the ventrictee in a very complex system of aheela or bands
ibpoeed as spirals, and in other ways, the details of wbicb need out detain
_^ In ibe auricular Appendices and elsewhere, tbe bundles form irregular
■networks projecting into the cavities.
The ounneciive tiaoue binding the muscular fibred t4;^ther, unlike the
atfnspoodiug ootinective timuc tu the frog's heart, is well supplied with
bloo<)vcsMils belonging to the coixmnry aiy*tcm. This ouiincdivc tivuc forms
on tbe inner surface of the cavities a couliouous sheet, tbe connective- tissue
basU of the fiat epithelioid cclU of ihe endocurlium, and on the outside of
the heart the viaocrat tayi-r of the pitricanlium.
The hintoliigical unit of tbi^e miiKt^ular bundles is neitlier a fibre nor a
fusiform fibro-ecll, but a more or let* columnar or prismatic nucleated cell
generally provided with <mc or awn short, broad prooessw. [F\g. 90.] The
nucleua, wnich is oval and in general rv^emblos one of the nuclei of a #tnaled
fibl«, b placed in about tlie middle of tbe cell with it» long axis in the line
THE VASCULAR UECUANtSM.
iKni-fti.
■ir nil lIUHi. MUlK IK i UfHal'
ntimti. »ni.titi%. On lbs ttshl, (ha
IlulUur ll>« ■g|».-«la vrlb wllh Ui^
iiucIpI on i-iMHliil Moicwlul 41a-
cnummuloally.)
of tbi' luDg (liiiDxtttT af iIk- cell. The coll IkmIv, wbicli is nut liixiiiiind by
any ilvfiiiile niirruti-mmii, \* KtrinUNl, tti»ii);h (riiocuroly <o, ncriMS tbu tuBg
■licinntrr iir tho c«ll. llio Ririaliuus m in n
skeUul musok Kbre being diH> to iho tUivrna-
Una of dim mmI bright baixla. As in lli« fn^'*
bmrl, i;rutiul0' are IWi^nenlly abundant. <th'
acriiriii^ ihe stnatimi, whicli iiHlecd tven in the
abdCfM^eot' umntileB ia never so disliticl as in
ili«flbn8<»l'skel«Ml miuclw- Sucbucell is at
«ftrh end joiiieil by reinent Biih«laiic« to HimiUr
oelU, no<l a n>«t •>( mcU celU i-imstituicfl n
Mitliac "{rmriitary fibre. Heni.-^, a cardiac
flbra b II fibn- !<triMt«<). but wtthoiii iinr<!i>lomma,
mm] divt<)r<l by tiartiuon* of reJiieiit 6iib«tBnoo
tnu> M>tiH>» bat rlou^itvd ilivi«(<>iu or culls, each
anntainiti^ a nuclMW. Many of tli« nib in a
tibro bavv a uliort bmod, Ulvral pruceai. Buob
It prac«M b uniti-<l by cMnent Mibn«Doe to m
lirailar pr»i;tw> of n wll twionging to an
adjoining tibre: and by tbo uninu of a number
of tJirM |iroocHu-«. H nniitb«r of parallel fibm
ar<* forni^Nl into i\ uxixrirbat oluw nvtwork.
Kacb buDtlle of thu carditu' miidcular tiwuu in
thus itcelf a iiatwork. THmo bundlca ara
Airlbvr wovirn into ni-tworks br oonnoctive
liaeu« ill which run car>illarioe anil liirgor blood-
veaaels ; aiid dhoeu <>r oiindira comiMMed of such
notworlu atv arraii^'^d, tw we have said, in a complex manner both in th«
auricle and ventricle. Hencu, the miucular dub^iunoe of the mammalian
hnirt is, at boitoni, on exoeediogly complex network, the elem4:nt of which
i« a wtniewfaal branched nncleatetl striated coll. U may Iw rvmarkud Ibat
tbe " tnusculi pectinali " of the auricle and tho " coliimnLV carnca; " uf tli«
vvntricle suggeil Ibe orixin of the inammaliau hunrt fn)ii] a iniiw4ibir
Ubvrinili like that of the iWi|t*s ventricle.
At ibi' cifinuienceiuvnt of tne great arterieii thin [icoulinr eardiac muscular
tMMue onucH abruj>ily. beinff refilamd by tbn ordinarv iitructurei of an anery.
but tbc •triuted miimiular Bbrea of the anricle may lir traood for aome du-
taocv al»n|[ bitfh the veiitt oavie aatl venie [lulmonalcs.
tTniler tb« endonmlitini are frmwintly prw^nt ordinary plain muscular
flbrw. and in koidc cawii )>«cul!ar ciOl* arc tuund in this situalion, the uelU
nf ['urkiiij'-, which »r>- intcreitlinf; ninrpholngicallv becaUK the budy of the
ndl anxiad the nucleus is nriliiiary clunr protoplasm, while the outside b
■iriaiM] lubatanoe. Plain muHcular fibri-a arc wiid al*u to snread frum the
oadiinintium for a ccrlnio clistMtc« into the nuricti to- ventricular valves.
ilSS. Tlui nrrTTM of tho kimrt. The diMribution of nerves in the heeut
varii-a n guttil Ai»\ in diflerent vcrli-hral>- animals, but neverlhele^ a k;eneml
)>lai) i> tniit« uT IwH evident. The vrrtclirate heart may be rcpirdetl m» a
muaculnr lube (a niiiglu IuIm>, if lor the m'>riient we disrettanl the complexity
(if a double rircnlation occurring in tlie hi;;her noimals) divided iuio ■ wrlea
(if ciiaitilipn. sinus vetioeus (or junnioii of great veins) — auriele, rejitriole,
and bulhud (or cuuus) arteriosus. The nervm (with the exception of a Hmall
or-Tvc which in aotoe animals reaches the he«rt by the aorta) enter the heart
at the vut'iuB end nf this tube, at the sinus renoaiu, and pa^ on toward ibe
arterial md, diminishing in amount as tbey proceed and •lUappeunng at tin
aorta. ( InDoected with ibe nerve tibree thus paniog to the heart are grou|n,
286
THE VAl
rBCHAKISll,
At ench beat llie time durip); which the <.'ont«nU of tlie left wutricle are
injecwil iiilo (he aurta U, tu we hnvt leeu (.^ 13t>). very brief. The expnu-
sion of the aorta b very sud^ten, and the ocaiaiian of that cxpaimon is alao
very >u<)den.
^ow, wlioti lliiiil U l)«iiijt dri\Mti tvith o^t'o n stviuly preniire tbrtHigh an
elastic tubt! or n syattm o£ ehmtic tiibvH, levcnt placed On the lube will
de*cril>o curvtw iiidiating VArintictnii in the dtnmctvrof th« (iibclf lUc iiifluw
int<.i the tube bo vuddcnly Rtop[)cd, u* by sharply turning a etop-cock ; and a
comjntriitoo of Icvcn plovud at diflcront distancn ffx»iD lb« »top-cock will
ahavr thill thoie variaUoiM of diamuter trav«l down the tube from the stop-
cock in tho form of wavoa. The lever near the stop-cock will lirstof nil Dul,
but t>pce<lily begin to rlsa iignin, and this subflcqueot rise will bn followed by
anqlticr thil.afler which there mny be one or more ancceeding rises and fnll«—
thiit is. oscillfitions — vi-ich dccreaaing amplitude% unt^l the fluid conior tn rest.
The lot-ore further from the slop-cock will describe curves, similar lo th«
aIiovc in form but of tese amplitude, and it will bo found that these MMrtir
somewlial Inter in time, the more so the further the lever is trom the stop'
cock. Obviously Ihese waves are genemted at or near the stop-eock. and
travel thence al<in^' tlie tubing.
We may infer that at each beat nf the heart similar wavea would be
geoeraitMl at the rout uf tlie aurta upon the sudden cessation of the San- IJroiu
the ventricle, and ivoulil travel thenc« along the einittie arteries. The facia
tJutt each beat is rapidly succeeded by another, and that the flow wliivh sud-
denly ceusex is also, by the nature of the ventricular atruke, suddenly geo-
eralcil. may render the wuvoi more complicated, but will not change Uieir
essential nature.
Tlic exni-i interpretation of the gencnttion of th««o wavex is perbajM not
without diflieulty, uul two factors Mxm of esneciul importance. In Ihv first
place, iia we have already more than once said, when a rapid flow isauddenlv
stopped n negative pretaurc makes its appearance behind the column of fluid.
Id a rigid tube this umply tends to a reflux of fluid. In an elastic tube iti=
eflocte are complicated oy the second factor, the ela^itic action and inertia of
the walls of the tube. Upon the sudden oeeeatioD of ihe flow, the axptttisioit
of tlie lube, or as we may ut once say, of the aorta, ceases, the veeael becilia
to shrink, and the lever placed on il^ walls, as from .1 onward in the pube-
curve. This shrinking is in part due to the elastic reactiim of the walb of
the aorta, hut is increued by the " suction " action of the negative iirtHoie
siioken of above. In thus shrinking, however, under these oumbined cutseft.
the aorta, through the im-rtia of ila walls, overshoota the mark, it is carrie<l
beyond its natural calibre — i.e., the diameter it would poasew if left to iudf
with the pretturc inxidu and outaide ^ual ; it shrinlu too rou<Ji, and ooiuse-
*e(]uently beeins again Co expand. Thii> secondary expatuion i^ taking for
simplicity take a nulMi-eurve m which t)i*^ ho called pre-dicrotic wave, B, is
ubtHMil or inconapicuouN ) cau^tc* the HM;uudarv ri»o nf the lever up to 6^— that
is, the dicnitic rim.*. In thu^i cxpunding nsiiin, thv aorta leiida to draw back
toward llio heart the (Hiliimri i>f blood wliicl) by Iom of momentum had oomo
to nvl, or, indeed, under the influvncc «f tho negative preasure ()>oken of
above, wim alrviidy undcrgi>inf^ ■ reflux. In thin sccnndary expaneion, more-
over, the aorta is by tho incrii-i of ii.-< wulU, aidud by thai of the blood, again
oarriet), w to jjK-nk. beyond ila mark, wi ihnt no sooner hiis il become ex-
panded and filled with fluid ft a certain extent than it again Itegins to shrink
u from '_' imwnrd. And thi* shrinking may in n similar manner lo the first
be followed by a further expansion and shrinkint', giving rise to a |Mist-
dicrolic wave, or it may be lo post-dicrotic waves. A.OU the Mocotsivi-
changes thus inaugurated at the root of the aurta travel aa so many waves
TOB PCLSK.
237
I
I
kliMS the vtariaJ mt«iii, dimiiiisliiiif; ns tkey go. It will be oboen'et) ihat
fitr ibw ileTetoiiiiUMit of tl>p«e itnvee a oorUuu quality iu iho iralls of Uie tubing
ta WTiCt— TV. ThetulwiniiM licaucb fts powMSeB when at rcfll an o|)ei) luuten;
lh« walls uuK be of such a kind thtX the lube- reniaiiia open wlten eiuptv —
L *,. wheo Uw atinca|iberic pmaure is equal innidc and •nilaide — A that vdeti
it ahriala too macb it exptUMlR again in Mrivinj^ to re^^n U» naiiiral <-iili1>rc.
Thi> w« have K*a lo be a cfaaraeteriBtic of the arteriea. A culliifMJbU- lube
of lb'' '-.lae will not ibtm ib^ |>li«uotiK-na ; audi a ttilw wlioii ihcxtup-
eoek ' .1 1 i-ollnpws and cuijitit* ibH^lf. vouliniiiug ii> be cDlltiiAii] witlf
out Msv «Dwft U> ci|inDil nuulD.
In tfw above ri|tlBiiHti»fi nu meutioii had been made of tli« clnainL' of the
Mialliinir valvw; tv« •ball hnve in ^lenk of lb««! ii litllo Inter nti in rvfor-
ring to ibe ppn^icrotio wave, und nball wee that, under lh« vi«w we hiive just
giwB, iht c\tMag uf the !>eiiiilLiiiar valves b to bo rci^rdiNt rather lu tliv
•flbcl tban tli« cauw of the dii:ti>tic wave. Maay nulbont. buwevitr, give no
fanaravetatlon uf Ibe dicrotic wave diflenut fnim ihiit detailetl above. Tbus,
h it beld tbat Ibe primary tlirinking Ironi .-I onwani, being brougbc to bear
«■> tW (Mlamn of blood alnady come to rest, in bee of tbo gnat preatnre in
fVtnt, drivt* ibe blood back aeainM tba wmilunar valves, thus clwiof* litem,
aod that the iaipncl of ttic eolumn uf blood against tbe valves siarU n new
war* of espaniion, nhicli ifinforcini; the uaturnt tendency of tbc oImUc
wsJIa to expand again aficr lliHr prlranrv ehritikint;. produces the dierotle
mn C On this view, it if the blood dnvcn beck from the valves whicb
•xpUMb tbe artery ; on the view given abure, it is the expanding artery which
dmtrrs ibe blo<id baek townrd ibe valveH.
MoteovtT. fjuite olber views have been or are held ooneemini; ibiai dicrotic
wa%-c. AevirdioK t" many authora, it is wlial k called a " ivtifcied " wnvc.
Tlma, «brn lite lube nf the artifteial niodcl bearing tnu lev(irt< is blocked jnat
bwyunl tbr far lever, ibe primary wnvu is wen to beaoconiiuinW byafecond
waw, wbteb at the far lever in wi-n rlotw tu, ami often fusecl into, the priniair
wan (Fig. h3, VI. a'), but at the itcar tever is at some distance from it
(Fy. H3. I, »'). living the further IW)ai it tbo longer the interval between Ibe
Urrr atxl ibe block in the lube. Tlu) seennd wave t* evidently the primarv
wave reflected at tbo block and travelling linckward toward the pump, ft
thaa, nf eour^, pospes the far lever before the near one. And it bai) beon
ATfon) that the dierolie wave uf tlie nuUc is really sueii a reHectetl wave,
•tertnl either at the minute artfries and capillaHe*. or at the points uf bifur-
«atioo of ibe larv'er arteries, and travelling hnckward to the norta. But if
tUs wervlbe eaw. ihe distanee between the iirimary cmt ami the dierulic
cXHt ougbt to be less in arteries more distnnl fmui than in iboae nearer to
tlM* Iwnrt, just as in the artificial scheme the reflected wave ia fined with a
primary wave oear the bl<»ek (Fig. i<^, VI. H o. n'), but becomes more and
man sefanued from it the further hack loward the pump ite tract.' it < Fife.
A3, L 1 M. <>'). Now. this is not the caw with the dierolie wave, f'an-ful
imawirrmrnts abow that tite dislam-e between the primtiry and dlcmtic c-rcsla
>• aillMr gmter, or ecrtainly not leas, in the Knvaller or more distant arteriea
than in the Inrger or nearer oitea. Thin feature indeed prove* tbat iba
dicrotic wave canniM l>« due lo reflection at the |ti-ripherv, or, iniltx^il. in any
way a retrogrado wave. Besidea. the niullitadJnous |ieriplieral division wuultl
mtder one large periphically rvAecied wave impoasibfe. .\fn>in, the more
rapMly ibe primary wave i« ublilerated. or at leiitt diminulwd, <>» its way to
tba ptripbery, tbi^' Iom oooapioiiou* should be the dierotio wave. Henc*
iaoaMM exienfibility and Inereased elastic rtnctioit of the arterial wxlla
vbfafc tcod to Use up rapidly i)h- |)rimary nave,*li€iuld nlso IcMen ibe dierulia
^
SS8
TllK VAHCTLAB MKL'IIAMSII.
vmw, Btil DM n nmtlcr of fact Uinw conditions, ih we have Mid. ure fnTur-
mblc U> ihc proniiiicntT of the dicrwlic wave.
Od tlie other hand, thitw nnd thv otWr condilioRE which fiivor dicrotiain
in the pulso aro cinctli' tho»c which would favor such a develofiiDeat of
gc'ondiii'T waves ns hoe dcvd dcH^rituHl nbove, and their absence would b«
unfavoriihio to the ocrurrcnco of eiich wave*. Thus dicrotism in les iiiur][«d
ID rigid arteries Tsuch ns thoM of old noopio) tban in healthy elastic ODU;
tiw rigid wall neither expands so readilj- nor Bhrinlu so readilj, aod heme
does not so rendily give riw to such secondary waves. Af^in. dicrotiaiD ia
more marked when the mean arterial prcssure is low than when il is bigh ;
indeed, dicroiism may be induced when absent, or inoreased when slisluly
marked, by diminifihing, in one way or another, [he mean pressure. Now,
when the prenaure is hJKb. the ai'teries are kept cuntinuallv niueb expnnd«d,
and are therefore the le&s capable of liirther ex]»ausion ; tliut ia to ny, ar».
so iar, more ri^id. Uence ine additional expausioD due to the aystolo ia iioC
vcrv great ; there in a lew teodeooy for the arterial wulU to awiog backward
waa forward, so to speak, and hence a lens tendency w the dcvelapm«ni of
Mcondary waves. When the mean preasuro is low. tb« opposite itata of
things exitiiH ; tiuppuainj,'. of counte, that tlie ventricular etn>ke i* actcquMcly
vigftroutt ( the luw prcwure beiti^- duo. not tu diininiiihed cardiac force, but f >
(liminixhud |>tripfaeral resiataiiOL-). the relatively empty but highly di.-xriii^iblc
irtvry in rapidly expanded, and. fallinf; rapidly back, cnlcrt ujkjh k wcond-
arv (dicrotic/ expansion, and rvuu ii third.
Moreover, the .name principlis may bi^ anpliod U) explain why anmctimc*
dicnilivni will apiwar murk<.-d in ii pHriii-iilar artery while it remains little
narked in the nwt of the Hystem. In *!X|iirrin)cD[ing with an artificial tubing
euch as the arterial nitxlel, the physical cbimictcn of which remain the
■anio throughout, bulb thu |)riRinry and the sccoodary wnv» relaio the same
obaracters as they travel along the tubing, save only that both gnulually
diminish toward the periphery ; and in the natural circulation, when tlie
vascular conditions are fairly uniform throughout, the pulse curve, as a rul«,
poaMsses the same general characters throughout, tave that it is gradually
•• damped off." Bui suppose we were to substitute for the lirat section of the
tubing a piece of perfectly riKi<l tubing ; this at the stroke of the pump, uu
account of its being rigid, would show neither primary nor secondary ex-
pansion, but the expanding force of the putup's Htroke would be traniuilled
through it lu the second elastic secljon. and here the primary and aeoondary
wavM would ut once become evident. This is au extreme case, but the aame
thing would be aeeu to a leas decree in passing from a more rigid, that ia,
lam cxtctuiible aD<l elastic wotJon, to a less ri^d, more extensible and elaxtlc
Mction; the primary and secondary expam^ion^, iu npice of the veneral dump-
ing «flVt, would suddenly increase. Similarly in the living Doily a pulito-
ciirve which, no long aa it U travelling along arteriu in which tlw nu-AH
prcsaure ia high, and which are therefore praulicully ^mewbut rigid, is not
markedly dicrotic, may l)ec(imti very markedly tlicroiic when it <umn to a
|Hkrtini1iir artery in wbidi the mean ]>reMture U low (auil nc shall kc pres-
ently that nuch a case may occur), and the walls of which arc Ihervforc for
tltc time ttcing relatively more diiiteniiible ihau Uie r«cL
littslly, wo may recall Uic observation made above (§ 141) that llio curve
of expaiieion of an elastic tube is motlified by the preseura exerted by (he
lever emplovod to nK.'or<l it, nnd that bciicc, in the same artOfy mid with the
same in^trnmcnt, the nine, form, and even the special fcjiiures nf the curve
vary according to ihe amount of pnwmrc with which the lever is pmssvd
u{".>n the artery. Accordingly tlic amount of dicrotlum apparent in a pulse
may bo modified by the prcmurc exerted hy the lever. In Fig, 80, for
I
TOE rULSK. 289
iaaiBMtt, l)ie dicrotti' wave it moro ovidont iu Hw muldlo tluiii iti tlic U|>per
$ 147. Tbv pr»-<licn>tic wmvv (nmrkctl A on Pljr. HI, nod od seveml other
of tbr pulw-^tmcs), which pit-Mdcv iho dicrotic wave mid u still mure
Tarisblv Ihmi thai wkvo, beiug soniviinii-s slight or oven invisible niid soiue-
timM ooHpKilMH, has given nw to miifh controversy- In the iot«ri>retatioD
of tlM £erntio irav« given in the nrcccdiii]; imni^riiph it wu slatM] tlmt the
OMBlIr* praMure developed on the eesaatioo of the flow iu lh« rear of the
CDMBUiar blood, kd bjr iuelf to a retlui toward ilic ventricle; and it hn*
hrra 9afgm,ted that at thif reflux meeting and elosins the aemiluaar valvi.»
•buta a BOiall wave of ezpnnsiou before the larxer dicrotic wave )uu> bail
tisM to develop ii«eir On thia view the seiniluiiarvaJvei would be actuull}'
dflMd Ikefore the ocetirreoce of the seoondary dicrotic expansion of the
arl«n»l nails, though the larfft^r, more |M>wer(ul reflux of thi* lat«r event
mnu rvuder tlw tlmure n>ore L-oinplete, uud iu duiug so prxvihlv gives rise
to ibm peoond muud. Aecordinu;. however, to tiie aeicoitd view xivui ia the
MUM Mragraph, which regard* the reflux due lo the Kbrinlciug of the
arterjr to fooe of the great pntwire iu front as Krinly closinz the semilunar
■BlTaa,«lid U thus starting tlie semniUir}- dierutio wave ofexpaDiion, the
film dMiag of the somliiinar valvCM must take place bofbra the bc^nning,
Ml dariiu the dev«)i>po>enl i>f the dicmtic wbv<> ; it is Mill poi^blc, however,
ima OB tail view, a* on the otluir, to mpjiotu; that au antccttdcal refiux, due
t* A« iMeative preanira wioccediDg the c'cssation of fluv from the ventriele,
dotes U>» valvM and Utrts the pr^icrollc vravc. But iliv matter is one not
jM bevond tht' flago of coniroverey.
)t40. In an anarcrotii- y\i\fV the lint rise is not the high4«t, but a seeund
rm 11, Fig. Ml which f<>lliivt:> and is sijpnnitrt! from it by n notch u higher
tkBB,or at least as high, as ileolf. Such an anari-rolie wave, though it may
■wdoes be produced ieni|xintrily iu healthy pentoiM, is generally MW
(bud with diseased eundiiioits, usually such in which the arteries are ab-
Mnally f^^i- Iu des«rribing the ventricular iyutule, w« spoke of the
■■m* vUbin tlie ventricle as reaching its tuaximuin just before the open*
■gof tba ■emiliinar valves; and this is appareutly the nornuil event ; Imt
iWn are ctirvea whicli seem to show that aller the fimt suihluo rise of
fnpurs which opens the valves, fullowei) by a brief Uwcning of pressun,
efcich appcara on the curve us a notch, the preeaun: may again risc,nml thai
Its putBl higher than liefure. And u similar curve is •nmeiimri) il««cribed
\if the Auot-to-hai-k diaincter of the ventricle. The tiyalole opens the valve
■ ilvSf* with a hunt : this i* followed by a slight relapse, and then the
ntdt, stimgthoning a^in, discbatget the wholt^ of llie vriitrici>lar conleolt
klalM aorta and »u brings about a tardy maximum ctjinniinn. And what
■ tliin tlarteil in the nuiiM tranll onward ovrr the Btlerial system. It is
<Unih to ter hiiw tltew anacrotic ercnts cnii Iw pnxliiM-d. excupt by a
Riiaih irrrgulantj io Lbs vcairiailar •vsiolv; uid. indeed, the anacrotic
pobt is fn-i|uenily Hwoeialed with some iliMtnw.- or defect of iliit ventricle.
iU9' l'ri*ou4 pultr. Unitor n'rtain (-ireum»tnncn ilie pulse may be car-
■W •■ from the arteries ihrriiigh tbr uipillnrics into ihc veins. Thus, as mc
liall sea later on, when th« salivary eland is actively Meriting, the blond
nnyiiaue from the gland through the veins in a rapid pulsating etn.-nui.
Tfat aerroua events which ^ivc rise to the secretion of ualiva, lead at the
•Bs tline, by the agency ol vasomotor nerves, of wfatrh we wtll prcHcntly
i|nk,t» a dilatation of the small arteries of the glai>d. When the gland ts
Ufrst \iw ininuiv arteries are. as we shall see, somew hat ooustricled and nar-
iMtd. and thus contribute largely to the peripheral resislanoe in Ihe part ;
Iki* psripberal rtsistaure throws iuto action tbc elastic pm|ienics of tlie
240 THK VASCfl.AR UKCHASISM. ^^^^^^P^
SRUlI tutcrif* loAiliii^ to lUv )clai><), nml l)ie ri^miiiint (if the |)iit>e renctiiiig
tliCM' nrU'-ric's i,*, lis wc Ix'forv ex|)l)iiiM'il. tiiinlly (K-ntriivol. Wlii-ii tht? miauie
nrtvrit^ nn; ililiiti>i), tlieir nideiinl cliunutils nllnw tlio Muml (o lti>w iitoro
c3«i1v ihrotij^h ttiviii iiiiil with leav frii;ttou ; Ibc {H-ri|)hcral n'^iiitiiiirr wliirh
thi-y normiilly itffvr in llms Jf.-wticil. In Gpn*v<^ucn«- of tbU tlu) daolicitv of
thcirall» of lliv Hmnll iirtcrii^ m brought into })Uy to » Icn «xU.-iiI tWu
beforo. tind tlien? «iiii>11 BrterieM cotMO to <)o tlifir ahnro in dcstrojri'^ ^
pulse wliirh oonics dnwti to tboin from the lnrfi:('r ancrim. As in th« case of
the artitidal niod<;l, where thu " |>i;ri|ihrriil " lulling i« kept "]ii;ii. n»l <'tniiigli
«liuticity i« tiroiight into jility to convert Ihc intermit t«ni Krieriiil How into m
continuous one. nnd the jiiilw which rnichcs the 3irt«ri«B of the giniiil peuncJ
on through them nnd through the ni^illiirics, iiad U <:oiitinucd un into the*
v«iiia. A Bimilnr vcdous pulee ii also '•onietimea seen in other orvaus.
I'arvful trneingH of the grent veins in the nenghborhood of llie liearl shnw
elevations and deprcanons, which appear due to liic varialtansof iotra-cardtsc
(nuricular) prewure. and which may, perliaps. be spoken of as cooslitultny
n " v<<nnu!i pulse." though tliey have a quite ditferent origiu from the vraoue
pillie just dcHcribe-l in the salivan- glaml ; but at present they Deed further
eluciiMtinn. Iii uiises, however, of insutiicieney or tJie tricuifKd valves, the
•v«lolv of thi^ ventricle makee itoelf disliuctly felt in the great veiits ; uiil a
^fisliMiiion travvlliiJi: backward from the heart beoomee very vUible in the
veiiiH of thv neck. This i* xotuelttncii :<poken of lu n venoUM ptibie.
Vnrintion!) of preixure in the ^reat veiii», due to the rcKpirnlorr mor^
nienls, arr nlno mnietttuea iipoken of an a reuoti* palw; tJiv nnlare (if ihew
variation* will bn uxplaiucd in treuliug of rcspintioit.
The Redulatiox akd AoArrATioy or the Vascular MecnANtoM.
b The Itfyufatlon o/ the Bmt of the Heart
§ ISO. !v> Gir tlic fiu-ts with which w« have hnd to d«nl, with the exevptioa
cf tlw heart's beat itaclf, have l>eei) simply physical facts. All the ^vMntiftl
pbenomcna which we have studied iriay he rrjiroducFd on a dead modfll.
Such an unvarying mechnnicid vascular system would, bonevor, be itaelenio
a living body whose aclionn wen? at all complicated. The promiodnt CwtoK
of a living mechanism is the power of adapting itself to cbangea to iu
internal and external circumstances. In such a svstem as we have sketched
alMive there would be but scanty poircr of adsptittiun. The well-construcMd
machine mi^hl work with beautiful regularily : hut its regularitr would be
Ha tleBtructioD. Tht? Dame quaniitj of blow] would always How in the ttme
•teody BlroaiD through each and everv tissue nnd ori;un, irrespective of local
auil general nanta. The brain and the stuTuacli, whether at work awl
mtvling much, or at rest and needing little, would receive thetr ration of
IdiHM). allotted with a pernicious monotony. Just the same amount of blood
would imns, through the skin i>u the hottest lui on the coldest day. The
oaoon of the life of «very part fur the whole period of its exittence would
he funii.-*hed by the inborn diameter uf iia blood veateli), and by the uuvaryiog
motive power uf the heart.
8uch a rigid svslem. however, does not exitit lu actual li\'inf; beings. The
vascular mf'hniii.'U] in all imimabi in which ic is present is capable of local
and general modifKntiouM. adapting it to local and nocral changa of cir-
eiimetaiics. Tbv»' Ri»diticntir>n!< fall into Iwn great duns :
1. Changes in the bearl's beat. These, being central, hareof eoursom
general edbct ; they influence or may influence the whole body,
Tttt TASCCLAB MECnANISsr,
S'tl
^
'2. Cluuig«a in ihv [icripboral n«ii>iiiuc«, <lut' lo varinttuii-a in the culihre i>l'
Um miootr iiri«ri<«, brouglil abuut by Hut agtacy oT tbdr contractile tnii*-
vnlar o<uit«. TIk-m ctuuigM n»y m either luca), HfliM-tiuv a uarticiilar
vawulnr «m uuly, or g«Dcnl, iflectiiig all or nearly all ihe olooilvotitelt of
the body.
TIkm twodaaMs of events arechieHy ^vernrd by the nervoiu «yMo(ii. It
it bjr BMUs of th« iKrvouB «\vtcn) thnt ibi: h(-nrl'» iHrnt nii'i llie <.'nlibr« uf
iba uiaiile artcrice iirc brougiil into rolalioii with onvti nlher.aDd with almoM
•vcnr ]art of llie Wly. It in by mtviu of the ntni-oiiF »yntm acting cither
on lb« heart or on the bdwII arteries, or un both, Ihnt a change of circiini-
wumrtB aJStctinz either th« whole or a part of the l>ody is met by compeiiwiling
or rt^litive cliaOf^CB in ibe flow of bloo<i. It is by nicaiie of thv n^'rviius
mytUta lliat an orfcan bu a more full supply of blooil when at work thitu
wkeo At r«M, that the tide nf blouil (hroucli the sltiD riaea nod ebb« with ihe
riw and fall of the tttiiiitralure of the air. thai the work of the heart ia
leiapered to nieel the strain uf overfull arteries, and that the arterial ^•atee
open and ihut u> the furue of titc central pump waxes and wanee. The
■tody '-f theae ebaDgea becotges. therefore, to a large extent a study of oervoua
■otiwtuL
The cimilaliou otay aim hr. iDmlified bv eventa not belonguig to either of
the abow two Flamn. Thug, in tbii or ifiai peripheral area, Hianges in the
otpillary walla and the walU of the minute artenea and veina may lead lo
■a inciuw of tl>r (vndencv of the blo<Kl corpuiK'Iea bi adhere lo the vascular
valll. and to, quite n|>art from any cbnnj^ m the culihrt) iif ibti bloudviawdf,
nay Wd tn increiue of iho [icriplicrnl n-nt^tanoe. Tlii* in wen in an exlrvni<r
catc in influRimalton. hui may puwility mt«rvvne to a Inic extent in the
unliuary oiindilioD of Ihe circulation, anil may aVi hd un<h-r the inHum^v
of the uervous system. Further, any decided chan);r in tlie (juantily of
blood aclnallv in cirrulutiuu 0111.M also iolluencc the working- of llie vasmlur
mechanism. Due Ixnb lhe»-- chaii),-<h^ are unimportant compared with tlte
other two kindH of i-hiinun). Hence, the (ho nioil important problems for
u» to study Hre, I, how the nervous syiitem re^ihttai the beat of tbe heart,
•ad 2, buw the nervoui' »yi>ti-in regulateA tlu! cnlibre of ilw bloodveasela. Wo
«t1l Rrvl conai)ier the fomter problem.
Tkf llUtobtyi/ of thf Hrart.
% Ul. It will b« neccMary dow to take up certain {>ntnt* coDcerning the
■ninute ■tro«-((ire of ibe heart, which we hnil previously {loriponed : ami
■inct much of our knowle^))^ of the nervous m«-hani"m of the bent of the
hisrt is derived from eK|>criiiH'Dt« on the heartJi of I'old-bloodeil animals,
niorv particularly of the frog, it will be dciirabic to coniiiler these as well as
the mamnialiaii heart.
t ittdi/t^ rnvteaiar tisme. The ventricle of the frog's heart H composed of
miDMla «}>il>d]e-«haped Hbres or fibre-cells, each coninining a nucleus in its
middle, and tapering to a |>iint at each end ; foinelimes, however, tbe end la
ferfccd or even hraneWd. Tbeae fibres or fibre-cells, id lact, resemble plain
ataamlar flbres m\-« that they are somewhat larger and that their nibtitance
Is striated. Tbe Mriaiion ts due, like the strialion of a striated museic
Ibre, 111 allvmato dim nnd bright ImmU, but is rarely so dintinct tis in a
•kclrial lihrr; it ts very apt to lie obscurei! by (he presence of dispersed
dWtinrl grttTmlfH. which, in niiuiy ciuat ut all events, are of a fiuty nature
I.ikr ihf ptoiu muscular fibre, the ear<llae miiHculnr fibre has oo dUtinet
laroilenima.
tn
2 12
THE VaSOOLAB MECQAKISH.
A number of ibcM librim nre jninecl bv coment MulkWiuiM Eoto rruII
buniilcfl, nnd tl»i>i:biiiiclli.-a are. by lh<- belp uf CMiiucctivv tittniv which cttrri»
no blood v<-«M!t>, wuvon into an intricnU^ network or sponge-woric, wtiicll
forms the ercnicr piiri uf iho null of thi; veiilriclri. Iraiii«Nlial«)y uiKler tb«
pericArtlmfcnnting, cuiMxliiig of n Inyt-r of upiihclioid pbilcs restinj; on a
couocclive'tisiur- bnvix, Llic niuHciilitr liscui; fornit! a tbin continuous »b«ec. but
within this it spread* out iQto n (jKiiigD work, the iik'»)i<« of which nreecDt »
labrrintli uf paMSg«s continuoiii> with tho cnvlly of the Tentriclc. Tlie bars
of thiit apon^fvwork. varring in ihickncva nnil, llioiigh ■p|>ni'cntly irrejcutar,
urriiu);ed on a dutinile erslcm, congist. uf bundle* of niuHcubtr fliir«A united
by conuective tissue, and nre coated wilb the same (.'iidoeardial lui-mltfane
(Ant epithelioid plates rvsiing on a connective- tissue basis) that liuu the,
cavity of tbe ventricle and, indeed, ilie whole interior of the bean. ThM
cnviiy of tbe ventricle, in other words, opena out into a labyrinth of iias-^iges'
rciiching nearly to the eurface of the ventricle. When the ventricle is
dilated or relaxed, blood llowa ft¥«]v into and fllte ihia labyrinth, bathing
ibc ban of the apiin^ work, which, lu tbe atiHeuc^ of eapillarirai, deiH-rnl mi
this blond for their nouriahment. When the ventricle contrant*, the Moodj
ifl driven out of this liibyriutb an well at »ii( of the central cavity. H«aeeJ
tbe vcDtricle when dilated and full of blood i» of a deep ml colrjr, wbe4i
oontmclod and empty it extretnely pale, having littlki more than the color of
the muKular fibre* ibemMlveH, which, likr MrintiMl fibm, posKMi in their
ovn Rubetanco a aerttuii amouut of hKnioglobin or uf myohntmatin.
The much ihiiini-r wall* of the nuricio coii»i*l "f ii mwcb thinner nctworU
of similar libr<« tmitti] by a relatively larger i)uantily of connective tiasilS*
intOsUliD Rhcvt, with the jH^ricardinl racnibranft on the outside »»d tbe
endooaidiaJ membrane on the insiilc. The fibres bave lo tbe aurieJe a much
snater tendency to bo branched, ntkI many, ecasing to be spindle-ebaped^
Moome almost stellate. Among the obscurely stjiated but still Mriated*
fibm are found ordinary plain inuBculnr fibres wbiob iDcrease in relative
number along the roots of tlie veins, veme cav%, and pnlraoDales, until at
•one little diat^mce from the be^rt plain muscular mtn» only are found.
BloodviMiels are absent from the walls of the auricles also.
In the bulbus arteriosus, mixed np with much eoonective and elastic
tinue, are found fusiform 6bres which close In the ventricle are tiriat^td and
form n thick layer, but at a certain diiilunce fmm tbe ventricle Iohi; their
Htriation, or rather become mixed with plain miMculor librw. and form a
thinner l«j-er.
S 163. In the mammal, both the veiilricltH mid the auriclm nra (brmed of «
bundles of musculnniMiie,l)[)und together by coiincclivc tiMuc.aad arranged]
mora etpeoially in the veiitricl<« in a very comjilex Jiys[«ni of aheeti or bundai
diapOMa us spiraU, and in othi-r ways, the details of which need not delaio
ua. In the auricular appendices and elsovherc. the bundles form irregular
networks projecting into the cavities.
The connective tiMiic binding the muscular lihru' togolher, unlike tbe
oonespunding connective tissue in the frog's hciirt, is tvell supplied with
bloodvessels uolongiDg to the eoronary system. This cunneetive tissue fornn i
uu the inner surface of the cavities a continuous sheet, the connective -tbdue
baMH of the flat epithelioid cells of the endocardium, and on the outride of
the heart the visceral layer of the pericardium.
The bUiologiea] unit of these muscular bundles U neither a llbrc nor a
fuiifunn libre-oell. but a more or lew columnar or priamatic nucleated cell
generally provided with one or more short, bnmd procebses. [Fik. ^0.] The
nucleus, wlii eh is oval and in general rewmhlesoueofthenucleiuf a^tnatcd
fibre, is placed In about tbe mi<MIe of the cell with its long axis in the line
TIIK VASCDLaR UECIIAKISU.
248
irvi ai
or iiiK Hurt, a*» ix t Uiwii-
ti.'MMI. MiTI»*, On ihr rliU. tfec
limliior ihc KfaiBte oslli wiib ibdr
nutttl an alilMlal tonivwhal <1la-
KmmnMiimllT.f
tt Ibt hmg iliaiwur of ihe «e)l. The c«ll bixlv, whicli ie iiot boiiwled \if
WIT ilefinito Mrculcmmii. ia uriBl^l, thauKh uWureljr ao, 8<iroaa the loDg
diaiuKrf of t\tt cell, tlie niHiiUuiu an in ii
•kakul Btncltf Itbn btiiit; ilm- to itie sltenis-
tioD of ditt Uti brigbl ImikU. A.i in iIk- frott'it
iMUt* gnuiihw an tn^ioi-filly nlximlniK. olt-
■ciiriiif Ui« •thxtion, Mhicli idiIm-iI uvrn id th«
ahMDOtof gntRiitM is ntvor to ilimiiu-l an iii
thm 6br?t ul' akrlvuil idumIm. Such ■ ortl in ui
«aeh Mid joined hy cemrat wibstAtict tn »iinilnr
fidli, ■0(1 ■ n>n of i>ucb cell* mnfilitiilrK ii
cardtkc <'li-ii><.'riliiry fi)>rc. Ilniiie, s cardiac
6bf«MafibrvalriBtr<l, hill n'i(h<Hit wircoloinina.
■ed diirided by partiiioni of ninviit »iilNitaiii-i.-
bibi MHWK'lial elon gated diviMom or <-pll>.4-H<:h
aMtaJniniE a nacleua. Many of tlic m*!!* in a
flbra faavo a ahon broad, lateral proocos. Su<!h
m [noma la unile«l by ceiii«Dt mbetaiwa to •
■iiallar fuvii^eitt of a coll WIoDg;ing to an
•djotniui; libre; and by the union uf a number
4ir t)M«« iTiK'fw. a number of parallel librca
■n> loni»i'<l into a ntnewhai oIu«e neiwork.
flarb iMimllr of ibe cardia« muscular tinue is
titu* itH-lf a Mtwork. TiieM bundlet are
tofthrr woven into nelworictt by connective
tianii ID which nin capillariai and lar^r blood-
vaaah; and ibiv-i* <ir b II ml lt« composed of auch
Ol»Bfb arc nrriiii^o), k> w« hare said, in n oomplox manner both iu llie
auricle sod vi-nlriHo. Heno.v thv tiiu»ciilnr xtibilnniv of the mammalian
hnrt is, at holl'im, an exceedingly complex notwork, the element of which
iiBBMMwhat branched nucleated »Inatoil cell. It may be remarkMl that
■kf>**iilusculi peclinati " of the auricle and the "columaie canK«"of the
*«a(ric)e aoKK^t t)ie origin of the mammalian hearl from a niuacuiar
labrrtoth like thai of the iroe'a ventricle.
At the eummencenient of tlie F;reiit arteries this peculiar cardiac muscular
tiwiip cmu>e# abruptly, being replaced by the ordinary structurei of an arlery,
faai tfae stniiied inuwular nbrea of the auricle may be traeed f>ir mime dis-
uaea along both the venw carte and reoie putmoDalea.
Uadar the endooardiuin are freaiieotlj preaeot onltnarv plaiii muscular
flbrea. and in some ea«es peculiar cells are fuimd iu this silualjon. the evils
flf Puricinj/-, which ure inteiestiug nmrpliologit'ullr hecaose the body of tlu
call an>utid the nueleus is aniioary elear protO|>lasni, while the outside is
•Iriatfd (iilmtaiice. Plain miucular fibrM are said also tn iiiirrEul from the
andof rdium for a oeriatu distance tni'i the aiiriculo-venlriruliir ralves.
I US. TKf Hfrif nf the hearl. The dintribiition of wrvca in the heart
Tarios a good daal in diirereni vertebrate animali, but tirvcnhalMa a general
fiho is HKww or Icm eridunt. The vert«brnU- limrl may be r^ardad aa a
■MPCfllar tubo (a *ingle tube, if for the monirntwo diaregnnl the oonplexily
«f ■ double circulation occurring in the highor animals) divided into a series
«f obanibtrs, sinus venosus (or junction of great vaitu)— auricle, ventricle.
■wl bulbus <or mnus) arterioHiSL The nerves (with thu eioeption i^a small
asm whirii in some animiiU ranches the hoart by the aorta) outer Ihe heart
al ibe voiMMa end of (his tube, at the sinus vonosus. and pass on toward Die
aitcina) end, dimini«htng in amount as tbuy proceed and disappearing al the
ODtuiect«i with tJie iMin-e 6bra» thus passing to the heart are group*.
24-1
TUB VASCOLAB UKCIIAIflBll.
■mailer nr grvAier, of D«rv« wHi^ Tbwe, like (lie mm fibres, are moat
abunilaiil at tl>c vctious end (uppMrlng on tin; iK-rv« branches before tlicw
actually mnvh the heart), a« a rule, become fewer toirard ibe nrlerial end.
Mid finally disn{)pear, m tlint (aceordiog to mwit (>b«erven; at tbu bulbu*
(oODUi) artrriiieue they are entirely abMtt.
Th«M) colleclii>ii» of iK-rve imiIIn or ennglia may ht arranged io grou]
aceonlin^; to tbeir [toMlinn. In nuinv lower vericl>ra(e0 tber« b a i£at[D<
riug or collar of gitnglin nl the jiin<<tiiiri of the 8inu« veiiosua witli tl»e aiirielt^'
where the prinuliTe circular di»p<uiti»n of muscular fibres is mainiuiiiMl ;
and there if a Hinilar ganglionic nillar at the juiictjou of the auricle with
the veniriclc. where nlxo there \» HiRiilarly retained a circular diaptMition of
the muBciilnr tibree forming ihe so-ealk-d mnaiit nurieulari*. And. imleMl,
iu all vertebrale* two similar eotlcclions of ganglia are more or letn iliatinctly
proeiit. There are ganglia at the junclioii of the sinue with the aiiriek- nnit
along tbe entering nerve branches : theee maybe called ihu oiiiuo gaiifilia.
Tliere are i>lher ganglia at the jiinclioii of the auricle ajid ventrick ;
may hv nailed the auriculo vciiiricular ganglia. Ue^dc* tll(«<^ Iwo grou,
tbcri! an- aim) ganglia overthe auricle in cunneotion with nervtw passing frvim
the Mnu.-> l<> the veniriclc.
IvHiitlv.aii a general rule. Ihe main nerve branehea and the ganglia are not
plunged dee]> into the suItstaTice of the bean, but are placed auperftdalljr
immediately u»d<'r the jieriiardial layer. From the eell.-< and nervw so eitu-
nted finer brancheit ami lihrm |iaH» lo the ^bstanoe of ihe lieart.
til the frog land other aniphihia) llie arrangvnicnt ditlere somewhat from
the above jilun, and therefore needii a special deacrijitiim.
The only nervea going lo tlie heart of the fmg are Ihe two vagi, riKht and
left, which may be leen niniiitig along the two tuporior venoi cavie, and
becoming Ium 1u view at the siniie, where tliey ihm from the surface tu
deeper part*. Kach vagus is not, however, simply a vagua neire, but, im
we aliull sec, contains tibroe derived from tlie splanchnic or Hyui[>atltetic sj»
tem. Ae tlie nerves appnwch the sinus, gruups of nerve cell* becooM
nbtindanl ii> ooniiectiun with the fibres, and as the fibres spread out nt tb*
dnus many ganglia are ocaltered among them, forming what ia emlled •■»
whole the o'liu^ rjitnijUon , or the 'i<iHplio» of llemai:
From the HinuH the two vagi, leavini; iheir po-<ilion under llic [lericnrdium,
plunge ititu the heart and run alon;; the bepcum between the auricles, on ibe
left ndc of the seuluni — one. the auleriur nerve, ptiniiing near«r th« fnini ef
tlie heart than the other, llie posterior. Several gruu|Di of cells or small
ganglia are cunncded with the two " septal " nerves thus pa»>ing along tlie
acptuiii.
The nerves reaching the auHculo-Tenlriciilur ring on the anterior tide of
the heart end in iwii ganglia lying at the \uun! of the two large surKiulo-
ventrieiilar valvea.
Front ih<«c two ganglia AiWr/crVf^oji^io, or the i>iineiiya-t<eii(n'cM&iryaHj7/t>i.
nerve fibres pass into ibe suhstanco of the vcniricle. Nerve cells mar t>e
traced on tbc (ibrm going to Ibe ventricle for oome little distance, Imt tot a
ItUle difflnnce only ; over the emler giart of the ventricle, the lower twu-
ihirds, for instance, the nerve 6brfs are free from nerve cells.
Thus, in the fro^' ihero are two main ganglia— sinus or liemak'a gangUoo,
auriculo-venlrieulitr or Itidder's ganglia. From lUe former there pass, on
the one hand, ^'altered librcs, in connection wilh which are amall groupt* of
oclb. to Ihe auriculnr vralls, and to the sinus walls ; and, on the other hiiod.
llie two main nerves running along [lie eepiitni, tn ciuinection wilh which
are omall gaii];lia which may lie enlled "seplul " gauuliu. From the latter,
Bidder* ganglia, fibres uDaccompanietl, except fora uTiort distance, by nerve
TIIS VASCL'UAB UKCflANISM
24d
I
to lb» Mihttitnor of tbo vrnlricle, Hn<l jxiwibly to tliv bulliu*
In Ui« mKmnial tlw nrmnf^Ricnt nppeara to conform nioro cliwuly tn lh«
gwni«l plnti dncribt-il uliovv. Tito «i-vcnil vtinlinc ncn'n. from thp HjinjiK-
tbttio cnun, lo^etlicr irilti ttniiii-liM frum the viigiis, includtug Hbrm from ilia
rwomnt Inrytifsenl, rnrm the supcrlicial and deep card iiic plcxuMs below
•ad beoeHth the urch of Ihc aortx. From these plexuora fibres nr« dia-
tribaied to tin supenorveDB cuva nod tu the pulniuiiary veins, and thence lo
ttw mrii>u« jtmrta of the lienrt. Gaofflm are abundant on the euperior rena
oiTM and are also found on lli« pulmonary veins, in llie vralls of tne aurielea.
in lh« auric-ulo-veiilrieular Kr<WTe, and in the bnaal portion of the vonlriclet;
further. aLxnrdiui; to wme obserTers, in L-ontrast m the fru;;'^ hearl, u num-
bor of *mall ^itn^lia may be observed over a \^rge part of the veiiindc fur
4oim luKurd ih(- ajH-x. Tbo auricular ^ptum, nl lea^i in iu cunlrul partt.
id fnv fruni jjaiiulia. The nerv«« and gauglia lt« for lh« most part superli-
ciailj imnuiliatiny undvr the pi-ricardimn.
!■ i1h> frog th* libmi furmiii^ the vagUK nvrves an lb«y nin alone ihe
•Mfrior TriUi cSTiv nrr (-ompDwd of mtslullaU.il uiid n(in-m«dullntcd finm,
tht lut«T tmng chifflr. if nm. uhollv, di-rivcd iV'im tho iiilanchnic or lym-
CktiMic ■yatrni. MMulUtwl Hbn-*. with ■ larger pnipirtion of non-iiiRduU
I»tI ffbmt, an' f"unil in tlir g>'pliil ni^r^'w riinnlni; In BiddcrV gnniflia. but
i1m> fin« fibtt-^ nhicli (mw from Itiddcr's ^iinKlia to llir «ub«lunoo of ihe tc«-
l/ict« an- ficluMi-ely nuniiK-d'illeilod Son*. Thi- iicrvo cells Jn the siDtu
(;■&«)*> and alou}; ih« c«ds of (lit- vagus ii«rv(«, us wHI ns some of the mIU
cif iJh> ^ii;;ltu M:Hlter«H over the Kcpliim. nr« of the kind prariituiljr (§ 96}
dorriheal as iipirul e«Itf. The coll$ compneing Bidder's Kunt^lia, at well M
maoT nf the celU in the seplum. are said to be bipolar and ru^iiform.
In the mnmmnl tli« fibret puasinK to the heart are al«o meilullated und
afNi-Rir>lulIatr<l. Some uf tlic raedoUnted flbrM are nf Hne calibre, muy Ite
tracnl back to thi^ vniciu. ami up|H.-ar to be Rhn^ of wliich vrc *liull (penk
nrcMinilr a* inhibitory. (Ilher* of the uiC'diil]nle<l fibres urv of larger call*
on. aiM sum* of tlxvK at ull events upiieur to be aeiuory, or at ti-aal allcnut
in ftiBCtloti. Of till- uon-mealullulvd libres, itome may be lnui->l luck aloiig
the ccnlUc ai'rvr* |o ihr infmnr rvrvicul gnnzlion, und nri' of thv kind we
»li«ll speak of us augnM-nling. In cmtnut to Uie frig, many uf thr tibr<« in
th* voairicta (wlicro thev lie rtufi' itndt-r the pericardium) are mmlullatcd.
■ad it b probable thai tbrsi- art- uflVrenl Hbrvf^
Tb« eMf (nrmiuji the rnrioii* ganglia XTUltCTed over tbo miimmnlian heart
■nr. pertu4«. h^ elasMd af unipoliir und multipolar, the fi>rmvr being ape-
ctally mnn«oie<l with medulUted libres, the one cImm being praniincnl in one
•bawon. the other in nuolli«r.
TAf liewlnfitneitt of the Ntrrmat Rfit.
Tba heart of u miinirnal or of a warm bliwitlvil anininl gvnemlly
U> h«M within a fen minutin uftor bviii/ rvmured from tb« bwly in
tbr iinlinnry way, the liisirt* of nnwiy-born animilii uoniinuing, liowcvvr, to
tvBi for a tongvr tittiv ihaii thiMp of adult«. Hunee, though by spcoial pro-
caationa and by mcam of an arltlicial circulation of blood, an Nolatvd
nuimmaliun lionrt wny lit* prnfcrvi'd in a puUatin^ coudittoo for a niui*h
Inoyrf linw; our Itnunli-il^i- of lb" exarl naturv and of the cniMU of tbo
«anl(ac l>«at i* ai yel vry largoly bavod oti the study of tliv hearts of ixild-
htwtA^ animals, wliich will conltQiio tu bral tot hours, or undc^r fnvorablo
<iiTt*iiaM(anoM enit for d»y«. aAer they havr been remiive<l fMm the body
w
THK TA8CDJ.A11 MKCBaKISM.
witti only ordinary care. We have r*iisou to think that Uic meclianixru by
which the bent i« c-arHcd ou varies in some of its Becuodary featurvs ia clifTer-
em kinds of animala ; that ihe hearts, for insuin^e, of llie eel, the snake. lh«
tortoiwi, and the frog, difTer iti some minor details of hehavior, boili finin
Mch othrr mid I'roni the hiril and the luaintiial ; but we may, at Gr«t at all
evenlH, tiikc lh<> hmrt uf the fmg iis illutttnitiuK the nmiii and iinportsni
truths coiiocruiiig ihi; cuiiim and incchanisin of the beat.
In HiucIyiiigclMely the phoiiomeiia of tliel>i-iit vt iLe Iienrt il becoiives nect«nry
to oliiaiti a grajihic recurd u( vurtoiia movrmnils.
I, [ii tlic (tQg or Dllirr coldlilomlcil nnimnl ■ lii()it IrTOf may be plMvd dirirctif
on the vcntrtdc {<>r on an iiuricli'. rlo.), iind ohurisn of totm. due eiUit-r to dislMI-
lioo by th« influx of blood or to ihe ivmole. will tuu»e luovi-mcnts of Ihe levw,
wbicli may bo recorded on a travelling tiirfncc. The «ame mtlhods, an we bavt
•een, may be applied lu tlie nininmnlinn hciirt.
3. Or. nil in C.iiukcll'« mrthod. the bntit may be fixeil by n clamp rarerulty ad-
jiiatfd araund the •uriciild-rcutricular Krimve. while ihe apex of tlie Tcotricle and
aome portion of one aurii-le are attached liy ibrtiidH to liori/Antal levers placed
respectively nbOTc and below the lieatt. The anricte nnd the ventricle each in iia
sVKlolc piitlH nt the Irver attached lo it, and the limea and eitrnt of Ibe oonlrac-
ttoni niny I bun be recorded.
3. A record of «ndo-cardiac preaaure may be taken lu the frog or lorioise. as in
the tuauimal, by nieanH of an appropriate manometer. And in tliose animals, at
all CTcnls, it is cnsy to kec^p up sn iiTliReial circulation. A csnuU i* inimduvcd
inin the oinna reooKUs luid nnothrr into the ventricle through llie nurtn. Seiuin
or dilute blood (or noy other lluid which it may be denired ta employ) is driven
by moderate pressure ibrough ihe former ; to the latter is attached a tub« con-
nected by meaos of a side piece with a small mercury mnnometer. So long a*
tlM exit-tabs is open at the end fluid flow* freely throu);h the hriut and appaimlua.
Upon cloiiog Ihe exit-lube at its far end the force of the vcniricular systole is
brtrashi to bear on tha muiomel4)r. ibe Index of which reaisien in tlie usual way
the movements of the mercarv cotunin, Newell Msrttu hu* succeeded In apply-
ing a moditlcation of this metliod to the mammalian lieiiri.
4. The movcmenis of ihe ventricle may bo regipiicred by inirodueing into it
IbrDUgh the auriculo-veniriculnr orifice n ■o-called "perfuiion'' ranuln, Figs. 91
and 'Jn ]., with a double lube, one inside the other, and tying
F^-91- ibe rentrlcle on to the esnula at the aurfculo-voBtriciilar
ETOOve, or at any level bclon that which may b« desired.
The blood or other fluid is driven nt nn udequate prennurc
ihtuLieU the tube a, outers ibe vi-oiricle. und returns by ihc
lube b. lib he cuniieeied with a niauometer ss lu method 3,
the mnvemcnis of the ventricle niny be registered.
&. In ihe appnmtnsof Itoy, Fig. 92, II., the exil-tabe is
free, but the ventricle (the luime method mny be ndonlcd for
Uie whole heart) is placed m an airtight chamber filled with
oil, or partly uilh norniid saline si)1uii>>u and partly with oil
By means of the Inhe A the interior of the chamber a is ooii-
linuoiu with that of a tniiill cylinder e in which a pUloti */.
■erured by a thin tlexibli' aniniul membninc works up ana
down. The pislon sgain liesr^ una lever ebr mean* uf which
its movement* may be rcgiatcrcd. When Ihe veulrlcte con-
A PurnatDi C*K\i.^ Iraels, nnd by conlriicling dimlni*he* in rolunie, there is n
letBCuing of presHure in the interior of ihe chamber; this is
limitsmilied to the cylinder, nnd ihe pisioti correspondingly rises, carrying with it
Ihe lever. Aa ihe veniricle suliMqiicntly becomes distended the pWMwre In ibe
ekambrr is incicnsed, nnd the iii*i»n und lever (.ink. In this way varialiona In
the volumu of the vrntrirlo mny lie reiorded, wilhuut noy great interference with
Om flow of blood or tluid Uirou't'h it.
The heart of the fro^. as we have just said, will rontiuue to beat for hourv
afler removal from ihe body even alter the cavities have been cleared of
blood, and, indeed, when they are almoet etupty of all fluid. The l>eais tlius
M
TIIK VAMCt'LAlt JIECllANtSH.
247
cmirird out ■!« in all iiu|>»nant rM|iocts idcntiml with th« l»rA1»«x«:uir(] by
th* bout in iu tiunn»l coattittoii wlthtu the living ludv. Hoooc vn- mity
loftr that Ibr litul of tlic haul » ui Mitmnutic ■cti'iii ; ihc aiuHtiliir mti-
tiaciiaw which coiigiitiil« lh« beat tre due to ctiiiM-s which arifc Rpunla-
OMHMly in lh« Itmrt iueir.
II rw, ta. I
t MMiM mute tint IMn ItaCt rmtili'lc n, «ntrmn<p; b.«i](-lul«. < . wall oT notriclt: fi.
KloTikpiMmlai mulUM br lluklll. a, (kuDlnr llUtd wtlll wIIik •iiliillun iiid nil. «cmlalii-
llfc imiiWto * iMcntoforfUMoucaiiula/. ti,iBbe1iadIiittooTllui]i'rc.liiHUtiIiiiiuTaplMan
tn th« frog'* honrt, u in Uiitt of tlic mnmnml. $ I'JO. ihrri' it> a dirtioel
iwftiico of tnrniU which ia ihr miik- whi-ther tlw heart be removed from, or
WniU io ill normal coodition withiti, the body. Fim cone* tlw beat of
lit MDus Tcnoeiis, precr<lr<l by n titonr or !<■» jhcriBisliic «>ntnictton of the
iirg» wins leading into it, next lbllon» tlw oharp beat of the two auricles
VfHittt, then otrmea the longer benl of th<! ventricle, and lastly, lh« cycle ia
■plated by tlie beat of the biilbus arlerioeus, which does not, lilte the
Jiau norta, simply recoil by elualic refleli»n after diatentJOD by tb6
_ 'Jcular Mrtike but canioa out n diHlinci muscular contrMtJoD pMeing io
iwtve (htm the venlrii-tb outward.
Wbon lb« ii«art in dyiti^ oeosee to beat, the oeveral niovenienta ceuK, aa a
mle, in an order the inverw of ilic alMve. Omiltio]; ibe bulbus urteriosus,
■ hkh aometimtit exhibits i^real rhyttiinioal power, we may say that first the
Ttatriole faib. then the auricle* fuil, and htnly (he aiituti vtmasua ikila.
The heart aAer it had c«aaed tu beat >ii>uniune«Mi»ly n-tuainit for aoow time
iiritable — that i», rapubk-uf executinj; a beat, or a ihorl wTica of beata, vbeu
■tinulaied cither mcchiiDi rally, a* by touching it wilh a blunt iiccdlv or dec-
Itiially by an iixluciino xhuclc or in other wav«. Thu artilii'iid l>ont to called
fcrib maT b« in Us muin feuturcn kli-iitical with the natural Ih.'AI, all the divi*
■ioaii>r tlie ht-urt taking jiarl in ihp tii-jil.aitd ihom-iiucncxtol'evfni* bring tho
an* a* in th« nntural beat. Thun when the (inu» in |>riokvd ihv beat of the
MBM may be fdlluwnl by a Wal of [he auricle* and of tho vcntriclo; and
trtn mhvo ifaa r«nlHcl« u Ntimu luted , lh« dircrlly following boat of tlie ven-
liicl* may Iw MeOM<l«d by n i-omplrte Iwat of tti<> whole heart.
UacWr nirtain circumstADOoa, how»v«r, tiM< divtAon directlv stimulaicd ia
246
aUy the
pownrn
tbe onljr III10 tu Iwitt: n-hrn the v<Milride is pncked tbr iustaiic« it kIodb
beatii, or uhcn thti tinug is pricked it nirtiio hcalK T)ie reeiilu ol' ittimiilBtioo
moreover may iliR'cr iiceordin;; to lli« conditioD of ihe bean and accurding
to tbe niirliutilur i|mt to which tbe etimulus is apfiliod.
Wilt) nu incretising Iusb of irrilabilily, tbe response to atiaiiilatioD cmms
ID the Kveml divisioDs in tlie sanie order as that of the failure of iIm natunil
beat — tbe ventricle cea«« to respoiitl drat, tlieii tbe auriclai, and lasllv tbe
uiiuB veoosufi, which frequently reepundtt to slimulatiou long allcr ibe <
divuious have ceased to make any »ij;ii.
It woiihl ap}>ear us if the s'mu.^ veiia«u«. euriclcii, and veiitriol« fbrmedS
d(MC«ndin){ ^ri«« in rtf»i)>ecl Ui ihiiir irriliibrlily utid In rhr [inwor thrr P«m4
of carrriii); oti MpiinianeiiUE rhythmic hcjil«, the »inii* being the matt pat«llf
Thill i» aW ^len in tbe foll'uvrrig exjMrimoiitK :
lo order that the frog's heart may hi--iil iiA-cr reinnval from the body with
Uie nwircKt nppn>ach in rapidity, regultiritr, and cndurancfi to the normal
c'ltidiiinii, the removal must l>e carrictl out so that the excited heart dill
rotaiiiK the itntif vcnwiix inl.nrt.
When the inciiion is corricd through the aiirides so as lo leavv lliv sinus
venoeus behind in tbe body, the rvaiilt ie dilferoiit The iiuut vcnoeus beats
forcibly and n?giilarly. having Fiiflered hanily any interruption fmm the
operation. The excised heart, however, remains, in the majority of cases,
for some time motionless. Stimulated by a prick «r an induction -sliock, it
will give perhape one, two, or several beats, and then cornea lo r«et. lu tiie
majority of cases, however, the animal having previously been in a rigorous
condition, it will after a while recommence its »|K>ntaneoas beating, tlte systole
of the ventricle following that of the auricles; but the rhylhni of Iwal will
not bo the same aa that nf the sluiin veuosaa lefl in the body, but will be
slower, and thi^ lieniv will not coiitiuiKi to go on for so long a tiRx- a» will
those of a heart mill retaining the nnus venosiis.
When the incision is carried through the aiiriculo-vtaitriculnr groove, so
as to leave the auricles and sinus venosu? within the body, and to isolato ihs
TCiulrlcle only, the renult^ are i^itnilnr but more umrkeil. Tbo sinus and
auricles beal regularly and vigorously, with tlioir proper Mouence, but the
ventricle, after a few rapid contraclionsdue to the incision acting as a stlinu-
las, generally remains f<>r a lung time (|iiie^oent, W'h^u Mliniulaled. hoiv-
vrer, the ventricle will give one. two, or several beatn, niiil after a while, in
many ciM^^ at lea^t, will eventually set up a apootanenui puUation with an
Indeiwndent rhythm : and this may Inst for some couMidcrable time, but tbe
beats are not ito regular and will not go on for so long a time m will those
of a ventricle to which the aurioiea are still attached.
If a transverse incinion be curried through the ventricle at ahrmt Ita upper
third, leaving t)io hone of the ventricle still attached to tbe auricle*, the [lor-
tlon of (he heart Irft in tin- body will go on piil.-uttiug rojiularlv, with the
ordinary setjuftncn of sinus, nurides, vuntricV, but tbo iHvlatw] lower two*
tliirds of the ventricle will not heal spontanoously at all hnwover long it be
U-tL Moreover, in rt:>)>on«: ^J a single stimuhts such as an induction shock
or a Ktntle prick it gives, not as in the <-ni<c of the itntiru ventricle, when
sUmulaled at the base or of the ventricle to which the auricles arc attached,
a serica of beats, bat a simple l>eat.
Lastlv. in complete the olory we may add that when the heart t> bisecteil
longttuiiinally, each half continues to l>eat aponluneiKisIy, witli an indepen-
dent rliythni. mi that the beats of the two halves are not iKCeaaarily -vM'
chronouH, and this iMUiinuani'e of s|xinlaneo(tp pulsations after longituiHnal
bieet'tlon may be SL-en in (lie conjoined auricles and ventricle, or in the iso-
Intcd auricles, or in tbe isolated but entire venlriole. MureoN-er, the auricle
TUX VASCULAK UKCIIANISM.
210
imf he dividrd in many wan and yet luiny of lh<> »0|pii«iits will r<iiilinutf
bsiing: unall i>\w^ er«n mmy W neen iindcr ibe niit.-ruBCO)>c |Milmiii);.
fnUy. k B* tmv, but ilMinclly and rhvthmii-nlly.
b thcM expenmenu, tl)«ti, ili« variouB pariB of ihi> fro^'n Iwart alw form.
H rvnb tmt power of 8|>oiiianeouB iml^'atiou, a dncvnHins serin: siniM
nSGN*, •uncles entire ventricle. lo«»r jwrlioDs of Ttrn(ncl«. Ilie )ut exhibit
lif asdtr (irdinMr)- circumBtuicM no B|)i-iniaoeauB pulmlion^ at all.
i IM. Mow we have seen <§ i53) tlint these parU form to ■ certain extent
■ MDikr (kaoatMlinf; aeria m nyarJii ihe preaence ofjianKiia : ai leaHt bo for
tki the nojtlii are verr numeroiiB it) ilw itinuH veoosus. that the)- occur in
tW Mrieles, and that wliilc Itiddur'a f[0»)clia are preeuit at the junction of
tW mMride with the aurM:l«^ ganglia are wholly absent from Ihe rest of
iW natriele. Hence, on tho aMumplion (which we have already, g 100,
iwg namm to doubt) thai (h<: nerve cflU of ganglia are aimilar id general
AactiMU I0 Ihe nerve rclln of the ci-nlnil ncrroun «yMem, ihe view very nalu-
aHj pnaata Etaelf that thr rhrthmic niHintani-ouH heat of the heiirl of the
#•{ !• doe iQ iImi ■[HintaiMyHiii pMicrali^^n in the gaoglioDic nerve cells of
rlnllmie imrfur impulMii which puvitii: down to the niiMCular Bbn« nf the
Nnnt pana auMS rhythmic contnicti'<nB of (hrw (!bn-«. ihn M-i^urncc and
aMioBtian »f the beating of the ■cvcriil diviiinn» of the hcnri lifin^ the
mAtoft coiinlination Wtwcrn the scvcnil ganglia in n&gard K" ihe p-nera-
l^if iBi|i<il«M. Under this view ihecsnliac muKciilar Itbre Dimply rrajiondt
l<tbe aolor irii{>iil«ea nsacfiing it along ite motor nerve fibr« in the ■nmi- war
» Um •kel'-inl maseiilnr fibre rmiMindi to the tnnlor iinpulsee rcnching it
ilmg ila HNXor avrvt' tihrt- : in bolh caaee the museiilar nbrc is, as it wcrv,
1 pMtve inntniRM-nl in the handf iif tbe molor nerve, or rather of the nervous
nain- i ganglion or spinal cord ) from which ibe motor nerve proceeds. And
ii» Tww. thus baaed on the fact >if ilic frog'ri heart, ba* benn extended to the
kaitinf (vertebrate) aaiiuaL< generally.
Tkrra ai* rvtwona. however, which shmr tlint this view ia not tenable.
For laaiaae*, the lower two-thinl*. or tower ihini, or i-ven lh« mere tip of
Or frps'a vcfltHolc — tluit ijt to mij. {Mrt* which are ni)mitl<-d not to onniain
wr*e etll*. luity, hy R]iei'iiil nieiinn, l>e indiicol to carry on for a coiiiidorable
ttBsa rhjcibmic beat, which in it* main fcHitir<» ■• iib-ntiral with ibe 'pM)-
taaMoa baiat of the ventri<:lo of the intact hesirt. If mi-h a ]iiirl of iho fnig'a
nntride be tied on to the end of a ]>erfii>ion canuU (Pig, 91 ). the portion
•^ the ventrinilnr cavily Monging to the pari mnv he sdei)ualely disteiidei)
Mil at ibo *»itii- time l>e " fed " with a suitable fluid. «iieh as blo'^i'), niude In
'ibmugh the eaniila; il will then be found that the portion of ventriclo
atcd will, nfler a prelim in iirr pi-rii>d of qiiieseeiice, C!>mmence to boat,
Rally «[Htntaneniis]y. and will continue a-> beating for a long period of
It mar be sui<l thai in this ease ibe di^lentinn of the cavitv anil the
ily of hliKnl or other tlutd acts a* a aiiniuluB ; but if ao the stimulus is a
'nuous one, or at least not a rhythmic one, and yet the beat u mnst rega-
'rhrihmio.
Then, apiin, lite reliictaiice of ibe ventricle to rxenile apontaneoii* rhytbmic
1 1> til n certain rxteni |<oculiar to the fro;. Ttie ventricle of the tortoiiM),
oManoe. the grenD-r pin of the MitMlancc of which in n* tree from nerva
I a> U that »f the frit^. will heal iiiK)ninneogiily ithen iaolnttMl from iho
I nilli grimt turn- ami for n limv' titni', Kiirther. n mere Htrip of tbi*
ibr niUK'ti' ti»'.i'-. ifki-pi gi'iillv e^ttcnded and Cixitinunlly riioisteiiod
I hhmd i>r other -iiiiabic llnid. Hill e inliniie In bmt njnnlanoutly with
'irrral rv^ulnrity I'nr bount.nr cvi'n 'tiiv»,i-«|v>eiiillv if tli^-MriCM bo started
b pivliiiiinarr application of ind>i<iion-flio<-lif Hiythmtcally repMtwI.
fn runnei-tiiKi with ihi* >pir*iifiii we mar call atlenliofl to the fact that the
S60
TUK VASCri.AR MECHANISM.
cntvlinc fUliiciilar fibru i* not wholly like the Hkclvtal lutttcular i
inanj n^ped* iIkt niulriictimi iir lii-iil of the fornivr is in i1« very luilutc <til'-
fcrcnt fivin itic coiilnictioii of thii lutlcr; ihc Hinncr canoot be coiisiderwi.
like the laitcr, u mere in«(runu-nt in the hamis of lh« nolor nrrvc (ihtv.
TIm^ IV-jiturc nf the bcitt or contrsclivn -f t^rdinc mitfcW mny be rtudieU o«
th« isolated mill (luieccent vcnlriclfl. or part of llie ventricle of ihe frue.
When HiK'h n ventrielo is stiniiilsteil by u eingli.- r<timulu«, such rs » HUgTc
in dilution -shock or a single louoli willi a blunl ni'edle. a bent moy or may
not result. If it follows it ivseiubles, in all its (■^iieral feature* »( leaM. a
spontaneous beat. Ueiweeu the application of ilie stimulus atid ili»' firei
appearance of any coptraction iu u very long latent period, varying aecordiDg
to ctrouttistaDces, but in a viKiirous iVesh frog's ventricle bein^; abouC O.-i
Mcond. The beat itself lastH a variable but vensideralrle time, rising «loKly
to a ntaxiniuiu and declininx slonly aKain. Of course, when the bent is
reconled by means of a light lever placed uti the ventricle, what the iradng
altowfl is really the im-reai« io ihe froni-io-bKck diameter of the ventricle
durJDfc Ihe beat — that in to say, uoo of the rciiilfn of thci eonlnic-tion of the
inrdiae llhri'!' — and givM, la an indirect manner only, the cxt^t of the cuD-
traction nf the filin-x thcmM-Ivt-n ; ami the Mime u the cniie with the oilier
methods of rocorditig ihn mnveiD<:nli( of the whole venlnclc. Wr may,liow-
cver, study in a more direct way ibe contrnclioii of a few fihrm by taking a
slip of the ventricle (and for this purjioM the tortoise i» prefembh- ti> tlic
frog) and siisiH^^ndiiig it to n lever after the fashion of n nnm-le- nerve |)re|<*-
mtion. We then get a curve of eontraclion, chamcterixed by a long latent
period, a slow long-conlioueil rise, and a slow long-continued fall— a con-
tmclinn, in fact, mora like that of n plain muscular fibre than of n skcJetali
muscular tibre. In the tortoise Ihe contraction is particularly long, tlte oon-'
traction of even the skeletal muscles being long in thai animal ; it is InM
long, but still lonu. in the f'tog ; sliorter still, but yet loug as compared niih
ihe skeletal muscles, in the mammal.
The heal of the ventricle, then, is a tingle but relatively slow, uiolon^ed'
eonlraction wave Hwe«ping over the iieculiar cardiae muHcIe-c«li, paesiug
through the cement suiutance from cell to cell idoug the Rbn, from Hbre to
fibre along thi; bundle, and from bumlle t^i bundle over the laliyrinth of the
ventricular walls.
Like the e«se of the skeletal muscle, this Mngle contraction is accompanied
by an electric chung<r. a current of action. The intact ventricle Si res* Is, a*,
wo have nlrcjidy Mid ($ 65), iwK^leclric, but t«ch pari just as it is about la
enter inUtastalo of eontriiclion iH-oomes negative toward the mt. Hean,
when the electrode!- of a galvnniiMn-ler arc phiccd on two points, A. fi, of lb*
•iirfaoo of tli<- ventrirle, a ilijiliiihiv variation of the galvanomcler needle i*
seen just a>a U-ai, natural or exc-iied.is about to occur, i^upposing llint the
wave of eontraetion reache" A first, this will become negative toward the rest
of the ventricle, including Jt, but when the wave some time afterward rmcfaei
B, h will become negative toward iho rest of the ventricle, inclading A.
Compare § fJT.
The beat of the auricles, thai of the sinus venosus, and that of the bulbui
Uieriosus are similar in ibeir main features to ihul of the ventriule, so that
the whole beat may be consi<lered In be a wave of contraction sweeping
Uirough the heart from sinus to bulbus; but Ihe arrangement of fibre» til
nich that this beat u cut up into sections in such a way thai the sinus, Iha
auricles, the ventricle, and the bulbus have each a beat, so to speak, to them-
•elves. In a normal state of things these n'verul uart* of the whole beat
follow each other in the secjuciice we have deM-ribtsl, hui under abnormal
oonditiona the seijueuce nuiy hi: revcnted, or oue rcclion may beat while tb<
TnS VASCCtAR KECRAXIAU.
2ftl
or Um scverml wclioiM nwv bcuL out of lime itilb vach
fe fiir, tko dmcHpiino nf
tat iliirm fmtn llial of a «k«
.Ibr.
th« c^nlrncliou which ia lli« fouoHfttioD of tlw
k«Ietal luiitcle in de)>ree only; but now oMnw an
b^rtuil iliflereDM. WIteii we tttiiuiitate a skeletal muscle with » strung
Miuhia we fiei it \trg^ cnnimction ; trbcii we apply h o'eitk itliniulus wc gti
■ *>ll coDtraniiiti ; uitliiii certain limits (we ^ 7'J) the contniclion is pro-
ptiiunal to i)ie •limuhiB. Tliis isi doi thi^ chso wiili the quiescent ventricle
It bran. Wheu we npply a »tnuig imluction-ahiick wo get a beul of a our-
ma Mjwagih ; if we now upply a wcitk ubitck. no ^i either no lieni at all or
q«W U •tnMtjt > beat na with the glroDgiT riiiiiuluv. That la lu say. tho
■uniltidv of tbe In'nt <Ie[ieDd« on lli« conitiiinn nf thi:! ri-iiiricl« 'or liearl),
aitot na the magniluile of U)c itliniuliis. If Ihf miinuliis t:uii viir the veit-
tridf i>p to l>eiil at all, llut brat i* ihr Ihi>I which the ventricle lU the time
aa utitniplith; the ttimiilu* either proclucc* it« maximum cfTcct or iinoe at
tIL U would weni m if the stimulus dow iiot pmOuco a oonimctiou in ihe
«nir war that It doea when it is hmugbt to bciir od a skdutal niuoule, but
athrt *ti» up tbc be«n in Huch a way os to enikble it to execute a spon-
tumuthnt, which, without Ihe extra slirotilus, it couM not briog aboiiL
Aid this » further illustmtcd by the fact that when a rcntrtdo is beuling
rbnimiatlly. eitlier ■pMitaneously or as the result of rhythmic «titDuluti<>n,
ihrkind of sObcl produced by a now etimuhis thrown in will dojieuil upon
ihianet phnae of the cycle of the bent at which it is thrown in. If it b
ttnnt in just m a relaxation is taking place, a l>eiit fullows pram a lii rely,
Kdar* the next btnt would naturally follow, this premature beat being obvi-
miy produced by the siimuliis. Itut if it be thrown in just as a contraction
i> biglflniiig, no premalure bt-al followH; the ventricle <toe£ not seem to feel
til itimulus at nil. Tht-rt.' L-> a jxnuil •litriii}; which the ventricle is insen-
■iUsloBtiinnIi, and that hnwrvi-r itnmg; ihio jKrioi] ii> cjillcrd tlie"reA«0-
period. (There in, it mny Im mentiuiml, a similar ratVactory period
muscle, but it is of cxceeilirigly short diimli[>n.) From this it
lliHt, when a succowion of stimuli repeated nt n certain rate are sent
llie ventricle, tbc number of beats doca not corrcs|iond to the number of
ttinuli : H>mc of the •limuli foiling in ri-fractory periods are ioeflcciive and
— Wc« no heal. Hence, also, it is iliflicult if not impotsible to produce a
tetanua of the ventricle, to fiieo n iiuinber of beats into one. And tlier«
■Olhtt faclf tending to show that the contraction of a cardiac muscular
ffven when induced hy nrtilicial stimulation, is of a peculiar nature,
hat the analogy with the contraction of a skeletal muscular libre, in-
by motor im|HilM« reaching it along its nerve, does nut bold gotMl.
and other conf^ideratioiig, taken together with the facts already nien-
', that portions of cardiac mitbcular (istue in which guiigHoiiic celh> are
tfttaialy not prcseni. can in vari'ios animals be indiicni. either eawly or
•ttli iliraralty, to execute rhythmic boats w hich have all the appearance of
bsiag Hioalaneoiis in nature, leiul uh to conclude that tbe beat uf the heart
BMI ibr mull of rhrthniic impultea pmceediu^ fruu) tbe cells of thi.- ganglia
lS|aMitc muscular tibrE«. but i^ mainly tbe result uf cluuiges taking; pUc*
fa tfca moKuIar tisane itself. Ami here we inay cnll attention to tlie |>eculiar
Uulos^fal features of cardiac uiutcular tjisue ; ih^Migb so far diflerenliatad
I* In be striated, its cellular ooostluition and its " pnuopluMuie " features,
iKluHiu)^ the obfcuritr of the striation. Dhow tliat tbc diirereiitiation ia iucom-
pbte. Sow, ooe attribute of uuilifrrreniinti'd prioionlial protoplasm is the
puerr nf •[■ontaoeouH tnovciueni.
t U6. We have, moreover, evidence that it '» the ninsctilar tissue, and not
■W amngeuent of ganglia and nerves, which is primarily concerned iu
biQl
(nxlti
fifi2
TIti: VASCltLAK MKi'IlAN'ISM.
ntniniaininj; the remarkable neqtience of sluii^ heal, aurictc beat, uti<t tea-
trick- Ileal, ThU h jwrhaps l)yiter seen in ihe bean 'iT che lortwise than in
that of the fru)>.
In tliia animal llie iiervea paniiiK IVom tbe sinua to tbe reatricle maj be
divided, or ibe several a*")?!'^ nm}' be re*(H!etiveIy removed, and ret the
normal ee<|ueiii'e i^ maitiiuiiteil. On the other linni), we Dnd that inieriereitce
with tiie museulnr xib^tiinoe of th« niirii-le, whea rnrrit-d to ii otrtaiu ext«Dl.
prevents the bf ul ol' ih« uuricK^ pn-ttiiig <ivvr lo ilif veutricic, hu that th<:*cqi)ence
IS broken adur ihc niiricle beat. If, for inxLanw, the aiiriclt' he cut ihrnusb
until oiily n narrow bridge i>f iiniticl^ be Irft (.itiincL'tin^ the part of the
■nricle adjoining ihv siniif with the [Mrt luljmiiinjc the aiiriculo-vcniriculu'
ring, or if lhi« purl Im- C'iniprc»*(^d with n clump, a Klutr of tbingit maj he
brought abiiul in ii hifb fvrry sci-ond honi <inlv, or cvnrv third beat only, of
the *1DU« nnd nnriclc is followoil by a bi'iit of tfic vinitride : and then, if th«
bridge bo still further nurrowed or the L-Iiimp screwed tiehl<ir. ihi* vcniricle
doc* not nl all follow in ils bent tbe 'eqiiorice of ainiii Hiiit luirii-tc. thmigh it
may nftpr a while eet up an indeimndrnL rhythm of il» own. This ex|H>ri>
meiit I'liggwJ*. nnd other facw flnp|>nrt, the view ibnt lh« normnl »ei]"cnoe it
mainlninrd ni^ follovvH ; The beat begins in the sinuf (incliidin)* the uuds of
thi> veins) ; thv coniraction wave, bet;iniiing at the ends of the vein». travels
over the muscular tissue of the sitina, and reaching tbe auricle starts a cun-
trnction in that iiegment of tbe heart : similarly the coniraetion wave of che
Biiriculnr beat reaching the ventricle «tarU a ventricular beat, which in luni
in tike fashion ciana tbe beat of tbe bulbu». And in hearta in a certain con-
diliou it is pos^ble hy stimuluiioii to reverse thin sequenee, or lo produce, by
alternate stimulation. lui ulieruiition nf a normal and a reversed aeqaeDoa;
thus in the heart ofihe «kate, in a ocriain conditiou, meohanioal slimiilatioD
of llie bulbus bv iiidiaiting a brni of the bulbu« wilUtarl a sequenw of tb*
bulkiiH, TvnlHcle, nnrielc, and Riuiin. nnd similar ■liniulntion '^f the ainnf wilt
produce n normal fiequ^iipa of sinus, ntiricle. vcnlricli>, tind Imlbu*.
It nould. pitrh»|)s, be prt-mntiirc to iiisjfl that the m-rvous vlenieiita do not
Intprrune in any way i" the mninlunanuc of thi" si.«|u<'ncc : but the evidence
•howv tliat they iirr not ihi? main fnctof, nnd we have at prcwuC no snliifue-
tory iDdicntioiis .)f ihe way in which they d" or iniiy int«rronc, »
Two iiuestion* iialurnlly Bugp?Bts theniseK'ee here. The lint is. Why d'»ea
the cardiac cvcle begin with the sinus beat? \Vc have pi\!vii>iislv, ^ l->4,
given llie evidence ihnt the sinus has a greater poteiiliality of beating than
the other parts ; iu nnd by itself it heals more reiidily and with a quicker
rhythm than the other parls. When wc ask the further question. Why baa it
this greater imlentialiiy ? the only answer we can at pivaeut give is that it u
inburu in the subttance of the (inns. The problem is somewhat of the same
kind OB why the heart of one animal hoaia so much quicker than that of
amitber. All we can say at pn^eiit i* thai the rale is the outcome of the
molecular constitution of liwuv, without Wing able lo detine that uiul«culAr
coMtituiiun.
Till' soonnd •(itration i*. Why doM not the cnntrncliou wave starting at tbe
ninua spreiul m a coiitinuonit wave over thfl whole heart? why i» it broken
lip into j«iuuK Ix-al, aiiricio ImuiI. venlriok be*l? We may here mil In ndnd
the fact inenliom-'l iii S IM of tb^i existence, mon- or U««i mnrki.'d in all
hearta and well m^-u in the hciirl of tbe lorlniae. of a niu.<rular rine or collar
between tbe muii* ami the auricle, ami ()f n ximilar ring hrtwecn Ine auricle
and v**nlrif|e. The muscular tiwue in lbt»e ring* swim* to he of a wmenlial
diHvn-tit nature from ihe muHodar timue fortniiig the body of the sinus, or
of the nuricle. or of the ventricle. If wp suppnec that this timue has a low
t-o lid II cling power, it may oflVr oudicienl rwialaiice to the progren of the
TUB TASCt'LAR MECHAXISM.
25S
I
The (Jiivemmait of the tfetxrt-btat bi/ the Nervou* SgiUm,
ctAiruttun lo prrtoit lite «iDU» ruri'SHmple lo cum out or to be far on ia
llw dr% rln|)i»«iit "t'ila hcul, bpfuretbe auricle be^iDs \u beat (uiil thus bitect,
•o tu tpmli. tb« Ileal wbich otJi«rwiee would bo ooinmon to the two), atid yet
Dot nflcr ao much nftiatance na to jirovpot the cmintction nave iMw&iDj; ulli-
■Mlalr (A from ihi' linua to ihc auricle. We may ia the tortoise bv carelul
dUDptiie or Kctton of the auricle in ila middle, by which an obstude to the
fantractiou wave ia introiluccd, bisect the eiiicto auricular beat into luo Ifeiitn,
OOB or the |i*n between the ftinuti and the oluttacle, aod another heluecii the
afa»Ca«:Ie and the wotricular. We may thus coniuder the breakiujc up the
priroitivv uubrukeit periualic wave of o-iUraciiou from »i>ius li> bulTiuit fi he
du« lo the iti trod uc lion uf Iwtue ul' louer (.-ouductiuf; jiuwcr at thi; Junction*
mt Hm Mveral pana.
We dn tux My liiat tbii ia the coiuplcte Mlution of the nroblem, but it Vt,
Ht oflkn an apiwnxlmate aolutioo ; aod hen^ u.h olaewlicra we haw no
Mtilfiwtary vvinlence of nervou* elenient l>eiu)c luaiu facLore in thv matti.*.r.
la the above wc have dealt chietly with th« heart of the cold-blooded
Miinal. but at far na wu know the tame ouiicJuHoiis hold good for the
aMmnaltiui lieart alto.
Tbe c|u«(iioti iiow arise*. If the jpuiglin are not the prime caote of the
bctftn'a rbrlhmie t>eat. or of the niainleDance of the normal ee<)ueuce. what
parpow cfo iher terve? Rut licforc weevvn itt(cn]{it lo nnxwer this iiucfllion
m* Mint <]c«l wilb the nervou( mcchnnitms by which the Ih-hI of the heart,
tfaoa sriaiDK aponlaneously within the tintjrs of the heart itself, \% modiSed
umI re^latetl to meet the rc(|iiircnK'iit8 of the r««t of the body.
^% 197. Il will becooTCnient lo be^in with the hcnri of the fruj;. which u«
> have teca v conotcted with lite ecnirat nervoui* sytteni through aiid
llwnrfbfa govenied by the two va|^ nervits. each of which though afipareotly
a liBSle nerve rontaint, ae we »hflll tee, fibree of <liHcrciit oriein aii<i natarc.
If while the beal# of th^i heart of a frog are being carefully rogi»tere<l an
ialcrtu|Aeil uurrenl of ntoderutc strength be sent through one of the vaui.
lb* twa/t ii teen to stop beating. It rcniiuns for a lime in diastole, perfectly
■liliiiillrii umI iJacc-id ; all the niUKrulnr libree of the MVeral cbaniberft are
fgr tba time lii'iug in a Hate of relaicution. The heart has been inhililni
Int Umi impuleea devceodiug dowu the vagus from tbe jMut of tike nrnrv
Mnulatnl.
If ihK duralioti of the stimulation be shurt and the mietigth of the current
t, \\»« •taiiddill may c^itiiinue alter the current hat lieeu abut ofl'; the
it* abni they r<-ap|Mwr arv generally at fir^t feeble and iafretiueiit, but
reach or even i,"' Iwyond their previouit vigor and frei^uency. If tbe
dnratioB of tlie current l>v vtiry h'og, the heart may recuninu-nce heating
vUklhe rtimolntion i* Mill going on, but the lHat>nre leeblcand iiitWguent
ittMigb sradually increating in Ktrength and frcijuvncy. The vircrl of tbe
Mtnolatloa ia at its maximum at or tnou aftrr the rommeuceuicnt of the
spftlicBlioii of the stimulus, grudunlty di.-('lining afterward ; but even at the
(M uf ft Very prvilongi.il stimulation the beala luay Mill be tew in fiircv or in
ftVfatBCY, or in loth, than they wen before tli« oerra whs stimulated, and
"■ the removal of the current may show etgna of recover}- by an iiK-reasw iu
toKe and fm|iteney. Tbe cfloct is not produced instantiuieoii!<Iy ; if on tbe
emve the pciiiit be exactly markeil when the current ia thrown in, as al on
Flit. S'L, It will fre<|ueDtlr be found that one beat at least occurs afler the
timvni ha* pMr!«d into l^c Dert'e ; tl»e developmeot of that beat has taken
c:
THS rA8Ct't.AB MKCRANI81I.
place before itie ia)|>ulM« (ItMeiidiii)^ iIh- vngiis hsvo b>H time to ii^ct tbe
The slitimlua ne«<) i»i>l nciMMMtrilr \>r tho interrupted curreul ; median icu I,
ctieniieal, itr (hernial mimiilutKin ot iho vagus will also produce inhil>ilinn ;
bill ill order lu ^'t a iitnrki-il ett'erl it it lUwmble tn make use ut not a singU
iiervouainiuulte bulascriM«rDervoiiB irapulseB; tlius it iadillieult toobUiin
any recognizable remit bjr employ ios ■ Nnf;le iiidu<-iii)tt-^b»ck «f niudoraie
inteitnt^ only. A* we shall eve later on "natural" uerrous impulses
docendiiig the viigun from the central nervous system, and staru-d there,
by aHerent iinpuls<v or otherwise, us jxris of a reflex net, may produce
inhibition.
«n
ImWIMTMII nr KDUin IIEIKT Sr STUtULjITKlfl Or VlGtS NUVt.
■IB nwrlulliaUiiuM wlikb lli«tlRi«TTupial curnni «•• iliniuBlBioiIianRuat qf wlwnii nit
•liut off. Tlir UtDP-nurkar tvl'>« msikii Mniinili Thi' Uwtt w«iv r^l^•*tn•l by nnpcndlBS Uw
ventricle rroBBClampUIMtivl to lticnurteinilalU(TlilD(a lIcHI leTor lo Ilu^Up e< Uw VdiONte.
Tlie stimulus may be applied )to any part of the< courm of the vagus from
high up iu the ueck right down to the sinus; indeed, very marked r«Bull9 are
oIitAiiiml by applyinc the electrodes directly to the sinus where as we have
M«n the two nerves plunge into the substance of the heart. The Biiinulim
ninr aluo he applied to either vagus, though tn the frog, and some other oiii-
iiinU, one vagua is aometJntes more powerful than the other. Thus, it not
unfre(|ently hap|>cns tJial even strong stimulation of the vagus on one side
praduoos no change of the rhythm, while even ntoderate stimulation of the
ni^rvi' on the olhi^r side of the net;k brings the heart to a stand^ill at onoe.
If duriiii: the inhibition the ventricle or otlter part of tht- hnirl Ix* ntimu-
lated ilirc^ctly, fur inittanoe mt.vhauically by the prick of a needle, a Wat
may fullon' ; that is to tny, the irapiil«ai dectoiuiding tli« vafcus, while inhibit-
inc ihc spontaneous boat*, have not wholly abollthed the actual irritability
uf the furdiac tiRHUc«.
With u current of oven moderate inlemitv, Kuch a current for ini^tAnoe as
wmild produce a marked totouus of a muai-V-iiurve preparation. I he stand-
still i; complete, that is to say, a certain numlwr of beats are entirely
droppeil; but with a weak current the inhibiciitn i« partial only, the Iteart
doe* not stand absolutely still but the beats arc slowed, the intervals between
tlieni being prolonged, or weakened only without much blowing, or hnth
slowed and weakened. Sometimes the slowing and sonietimM the weakening
is tbo more conspicuous result.
It Kinietimes happens that, when in the frog the vagus is stimulated io the
nedt. tlie eflbct is very diOerent from that junt deacriiMd; for the beats are
iaoiMied in frequency, though they may be at 6ret diminished in force.
And, oocAiHonaily, the beats are increased both in fon^ nud in frequency ;
¥
TlIK VAaCL'LAn UKCilANlSM.
S55
iW rvNiU b BuicnwatatioD, not iuhibttton. Bui this !■ due tn tho fnci thnt
a Uw ftw the va^^ alone the i^reater {wrt of iu nnirae is a tuu«d tierve
■■4 onaiMiM flbm uther tbnn ibone of the vhrva jtja\>t^r.
\ 1S& ir we eKHiiiiiii; lli« \amit uvrve cIml-Iv. Irnciiic it up tu tlte bntiii,
*m finil ihni jiul as ihc nrrvi- biu iijvn.'ti) (he crsniiim, jiidl where it paasM
IkfDVfti ibe gmngliuii ( (i. i'. Pig. Vi), oertalu fibn« puM into it Ctom iba
1^. M.
ftMMUitM' RamauuiitrKm n» ra* Ooomm o# CicutclAcianrrDB rtiaci ui ths Fm*.
rAtmutir t^» lud tiiftiMm. I?. I'. gmgUiMi itf wt. O. Uii« oT onnliU nil. 1^. ncni
VMt. 11. uBiUi. tli^tdi*t)»»iwl nwTg. A.rc wpntor mutjMnt. % ijaiikibatlc iMrm In
■■fe. « r. |«MrtUii a< iTinpalhFlIc 0tn/I1iiii with rttmt ttntfifa HDdlOf l.r. lutn cnni*) nliiw
t Mt C— !«■ gingUna Tbn mi -if ilw Dl.na fmm ftlaiiir thB(TacTi> mink O' aptitiif biile
I wlllt lb* Sn> (ptiwl iiwvv- </>ai>UnahBlrsia«ll(aior UiaMQODi] iiiiiial imrc
>K r OToalw i>r ViMaanft Xi^ ■utoli«lin>(i«iT. iT"' itiUnoliaic fuifUaanf iliu ibinl tiiiiutl
■••<• iit. IhIM •«<imI a*n«. r c. nmiu (.■miiiiiviiluiin.
Til ■■III lit lh>a'i(inMili>cnhn«l« ■tbmu l>f IIif llilak tiluk llriv. rtaer m*y bv (laovl tlvin
••«■•) «MdtTlb«aalalMi nwlof thcUilnl ipliul Mrn^thiousli iba mnu* maunu&lcaiB lii Uw
•w^imIIw ff4M*ito vwalloii 0<" aiHa Uhbm l>r tta •Mond laoallaa O" tba •snulnc trf VIm»>
■«Mi Hw Rnt mKMiHi 47 (# ift« iwilntl irmpalbiUc .%*nd n by Ibe n^ Uunlr to iln wfvilor
*TBpalbaiic D«m) uf lite Douk, S3, of llie rurlhar ioufineotioiia of wbiob w«
tlwltpcak pfOHotly.
Ttm }k'm$ tba fluc^ w« uay expect that we abuuld set infTerfut nwilta
MMnUi^{H w«atiiniilatetH')il>fl vaj^m io the craniuiD befur« it wan )oiiii>d
266
rilK VASCULAK MUCllAMSM,
by lli« ayiuputbeiic. i^2] tbe lympathetio fibres before tb&yjoln the voiiud.
>»il I .i) tho va^iifl trunk coulaioiiifc the ronl vn^uB and Ui« BjnifMitbelic fibre*
adtlud. Wbai we have previousl,r dwcribed are tlio ordinary resiili* of
aiiinutBliuK tbe mixed iruuk, and Uieae, an we bave said, are uottibolty
ooiiMiuit, utuugh UHually and in tb« maiu moHl distinct inlubitun' nsulu
fulton .
If wc Hiimidnle Ihe aym|iatbvti(r in llic neck, us at Si/, Fig. 93, cutting tbe
nervv bolun, >»> us U> bluck all inijmlses I'roiu pajiiinjj dowunanl. aiid uuly
klluw inipulieit to iitm u|i Id ibi- vuguti nod tb«DC« down tbe mixed vagu*
trunk U) tbe beurt, vr<- gi't very reninrkuble reiulla. The beat of tbe heart,
iiwtcad at being inliibiicil, \i aiigmcuted ; iIr- bcutt* are increaaed either in
frequency or iu fnrec, ur most f^eiicrally both in fr«<jUGiicy and in form.
Tbe 9tbcl it), [wrliap*. Wnt Men when the heart hvfure nlimulation i« Ifeatiug
elowly ami fiwbly ; uixm stimiilalion of ihe ttrvic*! »yDi]iathetic tbe b«at#
at onct imjirovc in vigor and frcaiiwicv ; indci-d, a fiwirt which, for on«
ntflwiu or Biiothf<r, has almnst ccaeet) to beat may, by profivr vlimuintion <tf
the symjiatbtrtic, be cnlk'd back into vigorous activity.
If, on the other hand, wvAliinulalc iTic vngus before it bae been joined by
the sympathetic tibrcH (and to insure the result not being raairad by any
escape of tbe etimiiluling curreni on U} the sympathetic fibres* it if noresmry
to atinmlate (he va^'us within the cranium), we get pure and cooslnnt inhibi-
tory resultii, the boats arc for a lime wholl}' abulisned, or are slowed, or ar«
weakened, or are both slowed and weakened.
Obviously, then, the heart of the frog is supplied through the vagus by
two seta of fibres coming from the cenlrHl nervous eyatem. the one by the
vagus proper and tlie othyr by the cervical sympaibelic nerve, and these two
seta liave opposite and ftntugoiiisiic etlecta ujiun the heart. We find ujxin
examiuatiiiu ibat w« can make the follotviug utaiemenla coucerning them :
Tile one i>et, (bow belonging to the vagun pr()|ier, are inhibitory ; they
wcjikcn Iht' iiy»t<ilc and prolong the iliaelolc. the <-IIi.'ct with a strong stimula-
liuii bring comjilcte. so that the Iicnrt is for a lime bnmght to a atandstill.
Hometiinoi the slovring, eomotimes the weakening is the more prumincriiU
When the nerve and the heart arc in good condition it needs only a slight
stimulus, n wenk ciiirvnt, lo produce a marked effect ; and it may l>e meii-
tioned tbnt the more vigorous the heart, the more rapidly it in beating, tlie
easier i* it to briii]; about inhibition. Alth^nigh. us we have said, the effect
is at its niaiiiuiim a»on after the beginning of stimulation, a very prolonged
inhibition may be j>rodueed by prolonged stimulation : indeed, by rhythmi-
cal stiniulaiion of the vagua tbe heart may be kept perfectly quiescent for a
very louj{ time and yet beat vigorously u|n>u the cca&ation of tbe sUmalus. In
otiier wurdH, the mechanism «f inhihitiou^lhat is, the fibres of the vagua
and the |iart or tiulHitauce of the heart u[ion which these act to produce infaibt-
tion, whati-ver that jiart or Aiibstanee may lie — are not readily exliauaicd.
Further, the inhibition whi-ii it cauva in, Vre'iiiently at all evenly, followed
by a period of rencli»n, (luring which the heari for a while beats more vig-
orwisly and rapidly than bcror*'. lnd<:cd, the total eftect of stimulating tbe
vagus fihrvs is not (o ex hau:<l the hnart, but rather to strengthen it ; and by
repealed inhibitions cnrvtiilly adminialervd, a fcubly )>vatiug hcajt may M
nursv<l into vigi>raus activity.
The other set, tb<Mc joining tbe vagus from tbe synipnthctic, are " aug-
mentor" or "accelerating" fibres: tbe latter name is the more common, but
Ihe fbnner is more aecumte, since the effect of stinnihiting these fibres is to
increase not only (he rapidity but the force of the beat; not only b tlie
diastole shortened, but the systole is strengtbeoed, sometimes the one reauli
and sonMtimes tlie other being tin nor« prominent. In contrast with ttie
THE VASCDLAR MECQAtflSH.
25T
I of the ragu* fibre*, ■ numcnhiitalroOKitiiTiiiliilioii J* rt<iuim3 to nrcMliice
»a effett : ih*- lini« retjuirvd fur tltc mnxTniuiD eHucL to lie pnxlucira w also
tvmarkatiU lun^. Moreover, Ht all event*, tn ilie cuM of a heart in vhkh
tb« ctrrulntttui i# not mnintaiiK^), nnd wliich i» therefore cut olf from it»
ounumi nulrilivc diipply, tlir MUciTM'nior tibrv* nr>' far !««■ etinilv ■■xhnii«ted
tiuui arr th<* inhibitory Abnt. tinnv. when in fiich H heart ooih Uti of
fibnw are Miniulatcd together, a* when ihv vngiin trunk in the Deck m ttlma-
Ut«tl, Ow 6nt rfle«ls prochiccd nrc thiwe of iRhihitJon ; bill tbiwv on cxm-
linuetl ttiroalatioti mar become miicil with ibiMe uf auffmcntiilioii. nnd
fioallv the latter alon« remain. Lottly. the contmst it oompletml by iho
fart thai the auf;menlAlion re»iiltii)j; from the slimutaMon of the ermpathetic
■ m fella«ed bv » period of reaction in vbich the beats are feebler ; in other
wwda. aiigioeotatiun n followed by exliaustion; aiu), iDileied, hr repeated
atimalation of ibcM rrnpntlielic fibres a fairly vif>DrouB bloodleaF ^ean may
bv reiluoed to a venr ieeble coiiditiou.
By waiching the eifecU uf atirotilMing the sympatbetic uerve at various
poiata uf iu coune we may trace thew nugmeDtor fibrw from their junction
wilk Ibe Tsgiu dowu the abort aympalbetic of the ueek ilimujib ilie first
■ apjanchnic or aympathctic ganglion connected with the fint «ninal nerve IV'
< rig. 94 >, through one or both the loofw of ihe iinnulua of Vieunieii*. .4m. V,
duoagb the Mcood ganglion cw)niii>cted with tlie i^cond npinnl nerve. &",
to tk* ibinl gM^lloa cooiMctvl with third »pinal itcrve. fw'", and theuoe
thraagk Ibe nuniia nMomiinican* or viM^'ml branch of that ganglion, r.c,
to lb* tblrd apjiud nerre. III, by the anterior mni i>f which itiey reach the
apioBl oora.
I IW. Both aeti of Bbrc* may then Ixi traced to the oentml nervous ays-
nm : ud we find aoconlingly ihul the hfArl may be inhibited or augmented
Wdcttoim inpuUee, which are Marled in the nervous aystem either by
mMmvot tmpulaes as part of a reflex act or otherwise, and which paaa to the
basft by the inhibitory or by the augmenting tract.
Tbo*. if the medulla oblonjjata. or a particular part of the medulla oblon-
C, wbicfa ia apecially connected with the vagus nerve, Iw «timulaicd, the
t i> iiibibiied : if, for iiiMance, a needle be tnrutt into this part, the h««rt
ataocU aiill. Tliia region in qutMion may be stirred intoaitiun in n " rt-tirs "
■MUMrbvaflcreni impulses reaching it mim various pnria of the boily. Thus,
tf tWabJoiDenof a fri>gl>e laid bare, nnd t!i« intetitine Iteslnick nharplywith
lb* hftodlv of a tcwljnl. the lituirt will Miuid Mill in dianlolc with all the phe-
BWnWMi iif vafcus iohibiliun. If the ncrvi metenUriei, or the connections of
tbM* nerves with the spinal ooni, tie stimulated with the interrupted ciirrvnt,
euttiao inhibition is similarlr prodm-cd. Tf in these two experiments both
f«gi an> (livi<led, or the tncoullR obh'ngatn i* dwlroyol. inhibition is not
{wmhohI. however much either the inlcHtine or the mesctiteric nerves be
rinBulaled. This shnws that (he phcnomeim arc caiuwl In* impiiUes asc«nd-
tiif along the mesenteric nerve* to the inealulln. and to alleclinj; n portion of
tiuil r>f|:ao as to give rise l>y roflci action to impulses which descend the
nfi as inhibitory impiil*es. The jwrtinn <if the medulla thus mediating
Wwntp the atlercnt and eSerent impulses may be spoken of as the rardio-
UkOiik ry er^iln-.
1ts0ex inhibition through one vagus may be brought about by stimulation
flf tbe c«Dtral end of the other. In general the alinienlary tract seems la
dner niooection with the cardio-inhioitory centre than other parts of the
hdy : and if the peritouewl surface of the inieslioe l>c intlnmed, very gentle
MMalalion of the inflamed surface will produce marked inhibition. But
a|ifiarratlv Ntimuli, if stifBciently powerful, will through reDex action nro-
■loee inhibition, whatever tie the ]hu-1 of the body to which they ue appned.
26S
TIIK VASCUI.AB MECIIAKISU.
Thu», LTU^tiin^' a frog'ti foot irill Atop the heart, and ailequKte MimuUiion of
niofit ulIbrv'Qi iicrvea nill prcdiice some aniounl ul' inliibiiian.
TIte detaila of tlie reflev ehiiin ati<l the portion of the i-entro oonnnm) in
itie iJerelopment of augmenting iiTi|)ulM« have not been worked iml so fullr
an in the cnse of inlilbiCdrv im|>ulM.'». hut there can b«i lit(l<- iloubl that lh«
former, like the liillt-r. are govL-riied by thi; (.'ciiLnil imtvuuh hvtU-wi,
i:I60. Si> t'lir we buve bi-en deullnft with thr heart of the fmg, but the
main fad>> wbieli nx* bavc- eljiied re^rding inhibition and nugmcnintion of
the lifurt-heiit a|i|>]y hIbo to other v^iebrntc! nnimaln, inchiding mnminak;
and, iixleed, wo meet similar phenuntciia in tlw hcorla of invertebrate
aniuinhi.
If in II mammal the ht-Art be esnnacd to vieir by openinj^ the thorax, and
the vngnN nerve be t'timtiliile'l in tnc nrek, the heart lony bo se«n (o stand
still in (linstole, with all the pari* flaccid and nt rest, if the onrreni em-
ployed Im- twa weiik, the result a« in the frog m not an actual arrest, hut a
Hlowinc or wenkeninf; of ihc beaU. If a liuht lever be placed od the heart
a graphic record of the i^tindetill or of ihc slowing of the complete or incoai-
plcte inhibition mav be obtained. The reaiilt of etimnlating the vaguti is
also veil »howii on the blood pressure eiirve, the eflect of complete i.'arHiiie
inhibition on blood-jireeanre oeing most striking, If, while a traeing of
nrlerinl picture is being taken, the beat of the heart be suddenly nrrmied.
eome fiuch curve oh that represented in Fig. 96 will be obtained. It trill Ih.-
riu. Mk
TaxrnmSaov\%n Jilt lvn.va«!ti or CtMHMi tsHontioK ■>!■ Bino)>*yu»nmx. Fnut a EUarnr.
I. Ihr maiKt oil ilip ilgnul lino w)i*n Ih« current !■ Itiruwii Intd, niiil y. utiut olT tmm Iha niftii.
nui UuiC'iuukct bBlaw mnrb Koniili, Ihe htmrt. lu it CKqucMlf thv eao In tha nbMt. Imbubc
obaenred that two builn follow the application of the current marked by the
point a, which currenponds tu the signal -r ou the line below. Then fur a
apace of time no heals at all ore aeeu, the next beat b taking place alinuit
inimediiilely al\vr ibc Hhuttiug off the current at y. Iiumediately oiler the
Wt lival fotliming a there u a sudden fall of the hlood-preeeure. At thv
pule*- due tu Lh<! Iiwl fvstiile the arterial ti\->tt-in la ill itja niaxiiuuni of diirieu-
tion ; j'firthwith the eliiaiic reiii'tioR of the arteriid walls pi'opels tlie blood for-
waid inlu the vein*, and, ihere Iwing no fresh lliiid iiijci-Ied from the heart,
ihe fall of the mercury in unbroken, being nipiil al iirwt, but alower afterwanl,
w llie elsnic force of the nrtcriid wall* u more and more uiwd up. With the
reluniing beats the preMuru corresjnndingly riac* in encccwive leMpa uuUI
the normal mean preasura ia regained. The sii» of these rvtumiug leapt of
the uiereury may seem dispronorlionalely large, but it miMt be remembered
tliat by far the greater port of the force of the (trat few strukea of the heart
TIIK VASCULjIB IIKCUAKIISU.
969
I
I
r
m mimmIwI En dbuodiog the artvrU] sjsl«in, s rrisII portion <Hily of tlw
blima whieb is «j«<4vd into iW urteries paoring on into th<i vriiu. A* lh«
aruriAl prcwnre ri»M, rioiv niid iiiun- blood pamm at «-HL-h I>«hI through the
eapiHann. and the riae of tho prtaaure at each bent bwomcs Ivts aim Icbh,
until at Ia«t the whola maUota of the ventricle poM it eneh stroki- into the
win*, ajxl the maaa arminl preMurv is «tablishcHl. To thi« it may Iw
•ddcO that, aa we have ttVD, tiw force of the individual bcnU mar he eom»-
what greater aft«r than before inhibition. Bcaider, wlwo the raerc^irrmano-
laKer ■• uacd, the inertia of the mercury lends to magnify the rffects <il' the
initial beau.
In the mamma] mhibition may be brought about by iiDnuteea paseing
•iuog flbra which, starting io tlie medulla oblungats, run doirn over the
ragiM nerve and r«ach Ibe heart by tlte cardiac nerves. Ii would appear.
bowvrcr, tfaat the iuhibilor}' fibn^ do not belou); to the villus proijer, but
lean Ibe central nervous sys[«ni by Ibe suinal avi-essary ner^'e. Thuit if the
RWO of tb« »j>iuai tuxeaaary be divided, tnoie of the va«us proper being left
tntact. the «pinal acoeaury likrea tn the vagiDt trunlt iKgenerate, and when
this laka place »tiniulaiiou of the tu^ud trunk faila to product; the ordiiinrr
inhibilitrr rlTectii. In th« manimni, a* in thu frotc, inhibition may bu brought
ab>Mt nik imly by artificial ntiinulation of thr voguK trunk, but by tiliniula-
tioo in a redcx manner or othvrwiiic of the carilio-inliibilorr oe.nlre. Thu.t
Um fainting which often follow* ujwn a I>low on th« »toin»cli ix a repetition
nf tlM rvult just nientioned u obtained on the froc by striking the stomach
or atimulaung the oerri roeMntcrid. Ho also ihr fninting, complete or
partial, ithich ncoomnanics teTare pain or mental emotion, is an illustration
nf <»nliac inhilntion by tho vagus. In fact, cardiac inhibition so far from
baang a uiera laboratory experiment entera rcpealcdiy into the erery-ilay
trorkinK of our own organism as well as that of other living bdogs.
Indee<) there is some reason fur lliinkiuK tbal the eentral nervous system
by means of the cardiac inhibitory ttbrea keejM as il were a coniiuual retn nn
tM heart, for, in the dog at least, section of both vagi causa* a quickening
of the heart's beat.
In lb« dog the atigmentor llhres (Fig. 96) leave the apiiiul con.1 by the
aotrrior roots of the second and third <lorta] nerves, ])owibly aba to some
extent by the fourth and liftb, jitua along the rami (ummunlcanles of those
Bervsa to the ganglion >tcllatuRi, fir^t tlioracic ganglion, or respectively to
saa or other of tho ganglia forming [mrl of tin- thoracic .iplandinio or
vvnpathrtic chain immediately below, and thenco upnanl through th«
aonolu* of Vicuiwvns.tnuiiug along one or other or L>oth loo[)«. to the inferior
Mat kal ganglion. Tneir further course to the heart i* along the nerve*
sprini^ing oiDwr from the inlbrior corvical ganglion or from the loop of
Viaiaaens dim-tlv. Their exact path from the ganglia in fact teem* Io vary
ia dlSrsnt individuals.
Tbe path of the augmentor fibres has not been worked out »o fully in
•Unr BMmnuila as in the dog. but it is moat probable that in all ca^es they
bare the sninal cord by ihe anterior roots of the ^cond and third donal
aarres < possibly also by the fourth and tiftJi) and. pawing up tbe sympathetic
ehaia to ilie gaoflion stellatuni and annulus of Vieussens, proceed to tho
bout by nerves branching off IVom some part or other of tbe annulus or
froai tbe lower and middle cervical ganglia.
Tbe oflbcts of stimulating these augiueutor fibres iu the mammal are, tn
MMral, the aame as tliuse witnessed in the frog. In the mammal us in the
tnf Iraplilars along iboN; augmentor fibre* may \>t ur^innliHl in thi- citilral
1 nai I inn srstcin, and that pr<>biibly in various waySL That jialpilaiion uf the
1^ bnrt akich is so conspicuouii an efled of cvrUiin emotiono is protmbly due
2ao
THE \'ASCL'LAR MEC1IAN18M.
to the aiiddeii poeilive boUod of augmmling impulH«. thuuj^li il ma; poadbljr
be rlue. lu pari at Itinst, to snddMl wtthtlrimni of iiormat, coDtiouoiM, tonic,
luul inhibitor}' impulses.
IiixiHiMiaaTii' KiTtiJannimoK or laa cui>
iirjiL iHiriiiiioBV t>o Atjoatma Pibnb
!!• TUK IXKI
II1B upper poitioii of Ui> titan npnasnU
U>g InhltilLoiy, tbfi lower l)w wisnuDior Abrai,
T.Yv., raotaoT the new; rJip.Xe.. rcnu«if Um
■l^nitl ncrauory: boUi drawn •my dhjft
iDfttlullr. (iJ.RaiiRllos jnsulan: u.Tt.Vc-.
^[ikIIuu tniiii'l ngt : ^p-Af., i|4iib1 amnwy
liunk : Kxl J1|i. Ar . cilmuil ipliuil w c«»i.ifT j
ISr><Ac.. Iniemal (plnalaucnaurr; Vr., tnink
of v*tiu> iivrvB : 11.'',, bninchBiiroliii; lolinut-.
C.lji)r««rtlMlt}mi«thi!«li<: (i.C Inner cwrvt-
Ml vuigUoD : X.tb.. futwlavlan arury ; Aa.V,
JUmiilluor Vtmmam: O.W. >Tb.<), fUfUoB
■fcllatum or am ihoiulc ssdrIIoo ; U.Th*.
n.Th.'. o.Tb.i. Hcond. tbltis. kiiil latinli Ik^
raiUccttnilla: n.II., [>.I11., I>1V.,I> V. Mcond,
Ihltil. fbnnb. uid nnii ihcirnr)cii4iAlDtrr«>:
ijt., TKiDui cummuiilnuu : n.v.. ii«na Kal>'
dlacj lawlnt 10 licatl inipeitca raoa M**)
fraoi nrvlnl (uigUbti unl Itoui tlic uinuliH
of V'leaaciui
Tlw labltdtoTT flbra*, Aowa hf lilncl Ilnr.
run In Ui« iipp« imtduUur tnuu. <a ibo
■l*nkl ■cotMOty. b; the Inlenwl bnacb of
itiu >4>iniU kMKaiory. |M>t ibe MnRMoo Iranel
vuKi' iiliiug Iho iRink <■( Uin nKiu. kud M by
limncbn la ihc lUperlar nna can aiul Uw
Till' >iii|(in>'ii|i)r nbm. alfo •hown t>r tdMk
line. iBH [n>iii (Iwiiidiuileoidli)' ibeuMstor
Kvit^uF Ettr *4<i-i*ijii Aiiii ihjnl ihara^c nfrrvn
.jitHiilily ii\f rroin itoiinii and anii u In^i-
.'ndfl ti> brukiii black lliio). paai the Mwoil
uid i)t>i tHiollHivi thoracic (■iiirlla tiy tli» u^
iiiilut III VlsuHriB l« Uia lowrr nnlokl tu^
rUuii. 0(1111 wbtncc m kIm (kuni tb* uioulai
ItwK, thrj' lan alnng llw cartlac ntrro ta
In tlio RiunmBi tben as in the hog, (lie heart !» g»vciti«<t bjr two fels uf
avntt, tbi> on« ■nlagonietic to the ol]i«r. lii the dug llic moU of tl>o «j>insl
ncixetoTj Dcrre, bf which inhibitory &bn» leave the (X-ntntl nervous sTsiem,
TBK VASCl'LAR UKCIIAM8M.
Amoog tbwe are fibres of line
be usom) linwa the tniok of the
^HHJ^^BbUrIj of medullated fibres.
raguB, ftkmji tbr hnin<-b«9 t:iiinf; to tbe bmrt, right i)i>w» to tbe beart ilself.
TImk mn Kb llUle doubr tbcit Hicse rootultiitoi) fibm of line calibre are tbe
inUbilorr flbm of th« va^nia, noil inilr<^l Un-n' \» evitlenoe whicli r«uders it
pmliable tbit tin: iiihihUorv tibra of tb« bi-nrt arr nlwByi> metlulbted fibres
«t fiM CftObrf, H liHi cuntiiiiif a« modulUtiil librr# rif;l>t ilown to the beart,
bat •waltnllf loao thtir medullx in th<< hrnn itM-IC.
Tb« nucrior root* of tbe tocond nti'l thin) ilorMl iMin,'m. nod the (ithiu)
tmal eofDraanicanla* belonging to th«Di. which. a« n*u hava jtwt swa. contain
ia th» dog augmenlor fihrw, alsci consist osclnRively of mod u I lit led lihita.
Bat tbft n«rr«s which convcT tbe augmenting inipuJK* finni tbo |nin>r
carTifJ noglion or from the anuuluB of \'ieiiMcn9 to the bmrt coniuM of
aott-KiedallaMd Sbra, Hence, the augmentor fibroe must bave lost tbdr
Mtdulla and become contiitaouii with non-medullnle<l Rhnt somewhere in
tWr cuurw alontc the ivinpathetic cbaiu. It \a probable that the chnng«
oecan in tbe^'anj-lion Bt«llaliim aiid lower cervical x''>')Rfion, and it i» further
probable that thi- ihant^? L-> cfll^-ied \>y the mediillato<l fibre pna§in|; into one
flf Ibagaoitlioti celU, ami Bvluing ita meilulla, the iuipul»cfl which it coorova
pMriDK out uf the nerve oell by one or more of the other procewn of tM
e*ll wftkh ar* oontiiiuad on as non-mrdiillHti'<l iibres. Cf. § t»*.
la lk« dog then thcM two sats of ni-rvc fibn«. antagonblic to each other
to function, difler in Mnicture, lh« atigmi-iitor Kbres early lonng their
Mtdulla and heneo being over n large part or lh«ir course mm -medu listed
flbna. wbrmu th<- inhibitory Hbrea are in«lullat«d fibres whit;h lluiu^b they
may paoa b^ or Ihruiiub g«iiglia (as the ganglion jngularc and (.-un^liou
tmoci rasi) do not lose thfir medulla in ttiwc ganglia, but remaiu aa
■srdullalea librts right down to tlie heart. And this dinereooe in Ktructnru
afpeara lo hold goo*) for all muniinals, and is possibly true for verU-bratcs
fUkermlly.
I in.' Tbe qnflKkn, what i» the exact nature of tbe change braught about
by tbe inhibitory aud augmenting impnbfti reamctirely on their arrival at
tbe beanT or, in olhor word*, by virtui^ of wliat events nroflnccd in the
beait itself do tbe tmpuUvs of •>iic kind bring alKiul iuhibition, of the
StlMr Itiad augmentation T b a very difficult one. which w« cannot attemtit
to ill can ftillv here. We may if W please «]wak of an ** inhibitory mecli-
ttimm " nlui-il in the ht-jirt itMlf, but we bavo no exact knowledge of ihe
aUan of such a mecluuiicm. dtill los do we powow any wittiitactor>- inform-
•lioa ■■ lo am augmenting mechanism. It has been »ugge«t«l that Mme of
tbe paglia in the heart serve as such an iiihibibiri- (ut augmenting)
■KUtusm ; but there ia evidence thai the inhibitory impulses prnduee their
•Am by aettng directly on the muscolar fibres, or at all ovenu do nut pro-
daca tbsir e^ct by acting exdiiaivelv on any ganglia. One evidence of
this jttftd is sapplieil by tbe action of tne dniji atropine.
If, eilher iu a frog or a mammal, or other animal, after the vwub ttbree
Wic br«n pn>veil by (rial to produce upon Kitmulation tbe usual inhibitory
efct*, a ■mall quantity of atropine be intnKlnced into lite circulation Cwlien
ih* •iperimsot b conducted on a liviii); animal, ur be applied in a ncalc
snlutirai to (he heart iiiK^lf when the ex|ierimeut b eonduoted, a» iu (he i-*»e
ofsftif.ua an oxi-iiaHl l>eart or after ilie rircu)atioD baa ceased), it wil) al^er
a 4wt tinve he fotind not only tliai the Hlimiilution, Die application of a
cnrmt for inMance, which previously when iipplii-'i to the vagus produce"!
Barktil inhihilion now producci no inhibition, but even thai the i>lr(inmi
NiSNilu*, the strongest current appli<i) to t)io villus will wholly faif to
sftvt ibe heart, provided thai there tw no 0Ma|>C of mrrent on to the cardiac
•>62
THE VASCCLJLIt ITl
tiMUM ibvmsdvts ; under the inHtioncs? of even n nnall iwv of ntro))in?, ibe
BtrongM slimulation of the vngii>s will not p^^duc« ilaiiijslill or npprcaablc
slowmg ur weakening of the k-nl.
Hon it tnighl be suppoeed thut the atropine producer this remnrkable etTwt
by acting on some gnngliooic or other mecbanism intervening belive«n ibe
va(;uB librei and ihe cardiac muscular tissue: but we have evidence that the
atropine acts either on tlie muscular tiaeue itself or on the verjr eodingB of the
nerves in the muBcnlar fibres. We have said. § 155, that a properly prepared
Htrip of tbe venlricle of the tortoise will execute for n loii^; time spoutAiteoua
rhytbiuic contracliuns. it will go on " beating " for a long time. A strip of
the auricle will exhibit the same pheaoiueini even etill more retidilv. If now
while auch a strip froui the auricle ie saliafnctonly beating a f^utle inter-
rupteil Diirrent he pan^t^l through It, it will Htop U-ating ; tbe current iuhibits
the s|)ciu[aiieoui beats; a very j^itle iiiternipKit curn-^t mu>l Iw u!>ed,
olhurwim: the eflbct 1* obacured by ibn mon^ direct stiniulatin^ m-tion of the
currvnt. If now the xtriii hi' gently bathed with n wenk nobilioii of alri>pine
no Much inhibitory clK^t is prudiiceii hy Ihe intiTnipi<'d rnm^nt ; ihi' Iteata
go on ri-gardtetiH uf thi; urtion of the current. Thi; ititi-rp relation of thU
expfrimi-jil in that in the first case the iiilernipled citrrcnt slimuUtrd tho
fiiK termination i>f the inbihittiry iibres in the muacnlar strip, and tbnt in
tbe second caw- the alrcipino pr<iiim«d smne ofiect eith^rr on lh«e line librcw,
or on their connections with the muncuUr sulislanoc or on tbe actual mns-
cular sub«t»nro iUclf by virtue of which they ceased to act. But if this be
so, if the entnc inhibitory ctlcct« tttw produced alike by etimulating tbe vaRua
trunk, and Btimulatiug the vorv cndnigi of the nerva in tbe muscles of tbe
heart, if not tbe actual muscular tiesiie itself, then there is no need to 6up-
p(«e the existence of any special inhibitory mecbaaism placed between tbe
tibrc8 in the VBg;us branches and tbe cardiac muscular tiasiie.
Till.' action < f atro|)in« on the heart is. so ti> apeak, com)d«m«Dlcd by ttie
action of muacarini;, the active principle of many imiNoiioua mnnbroora*. If
a small iiuuntitv of mtiscarino be introiluced into the circulation. orapplM
directly U> the ficart, tbe beats bwrjuie *l'iw and fetible, and if thedoM be
adequate the heart is brought to a coniptet<r iiliindiitill. The fflect it in vonie
respects like that of powerful »timulatinn of the vaguf, but tho staiiil^till is
much more complete, the ellecl is much more prul'i'urid. Now if. in a frog,
tic heart be brought t" n flandiilill hy a d<i»e of m»»cj»riiie, tbe application of
an adetjuate <|iianiily of nlropini- will bring back the beal# to quite their
ni>rmal strength. The one drug Is a* far tui tho heart is concerned (and,
indeed, in many other respects) the antidote of the other. And, as in tbe
aae of atropine, so in the case uf muscarine, there is evidence that the drug
acts not on any ganglionic mechanisms but on tbe cardiac tissue itself.
Tbe conclusion tbat inhibition is the result of changes in the cardiac llssne
itself may »en'e to explain why in inhibition sometimet) the slowing Mime-
tiroes the weakening b tbe more prominent. When the itibtbilorv impulse*,
by rcawHi of particular fibres l>eii)g atfected or otherwise, are brought to
bear dti«fly on ihooe [lart^ uf the heart, such as the ainun, which pu»»e«lng
higher rhythmic ixitejiLiality (tutt ^ 156) deiermiue the Ae^uence and set the
rale of rhvthm, it iii thti raiv which i» moat markedly airi'i^led. Wlivn, on
tbe other bBn<l, ihr^ inhihitory tnipnlnes fall chiefly on llic paria {i><w<i>e>ing
lower rhythmic potentiality, Ine nuwt markc<l i-Heet is a diminution in tlie
force of ibo coniractiona.
There is no ade(|uatc evidence then that tho citnliac ganglia act n» an
inhibitory iuecbani*m in tho sense that they prmhicG imiKirtaiit changes in
the nature of the impulse* reaching them along vagus inhibitory Iibres before
tJiose impulses puss on to the muscular tivue. We may add tlial there is
TUE VASCtTLAIt UBCBAXISV*
ass
I
H in;
ftmilftrly no MlMjuate BvidcDoi- ihat nDV of the ganglin act ns xii " nu^mt-
iBg " uodutiUni. We have piTvioiHly iwn, $$ lu'i, 1-06, rca«oiie for thioking
iIm nogUa nrv Dot ccniras for tbd ori^oulinii or reeiilntion of the gpoutaiiA-
iiiM liMli Tbe qucBtioii ifaeo ariipw, what are Uicir fimdionaT To this
i|ilHliaa «« rnuiiot at present fyn a wholly luitisfacloiT aoBwer.
TTmi inblliilory RbrM reiuuin, bb we have Mea, niMlullated libm until tbcy
i««rl) ihc bdArl, btil it would apjK^r that tb^ lose thdr medulla, somewhere,
in tbe h«»ri before tJi«v actually reai-h the tnusculkr tiwuc, and it is probable
that the luM takes place in cuDiiecliou with aowe uf the cardiac gaDglia
much in the Mioe way thai tbe auj^eutiuK librea loae their medulla in Ibe
ganglia of lh« rrBipathetio obaio ; but we do not know whiil is tbe pbyai-
olopal effect or Uw purpoae of this Ion of the medulla, aud we caunot
MppoM that tilt* i* iJk aole or eveu chief use of the ^njilitu Coinddenl
with ibr luM of tbe iiwduUii nn inrreaie of fibres fretjueutlr lake* place,
Bvorv than one non-iiirdu Hated fibre lea<niig a uervi? cell into which one
aaadullaied fibn t«ter> ; an<l we uajr auppow that ihla ntoile of branoltJag
baa ninoMa am fulfilled by tbe mere divuion of a fibre. Then awaio bearing
in awM Ibt nutritive or " (mphio " function of the apinal ganglia alluded to
in i 100, w« may luppn-e that the cAnUac ganglia arc ia wme way coDcemad
ID tike nutrition of the cardiac ii«rv« &hr<«. Ittit our knowlodge is not yM
Mttcieotly ripe to allow cxart itatciDmta to be mailv.
Oiker Injla«neei Regulating or Modifying the Beat of tite Heart.
i 169. Iniporlant as is tite regulation of tlie heart by ilie nervouji iTiileni,
it ntMt be burae in miml that other inAuencee are or may be at work. The
bwtnf the lieart niav. for iii*iiiiu>- h« modified by inflm-iit'ot bearing directly
t» iIm tiiilritioD of the heart. Thv (iiwtiCM of the heart, like all other tissues,
D«e*l an adequate sopplv of klixHl of a proper quality ; if the blood varyiu
quality or i|uantily tne Val of the heart w oorretp'mdiiigly aflbcted. The
U(C)M<1 fruc's heart, as we have aoon, cmliuiHa to Mutt fur some OOtuiderabla
titoe, lluiugh apparently emptv of bknd. After a whilv, boirever, the beata
dimlntsh and dimppear ; ami their diMppearanoo is greatlT barieoed by
waafalng out the heart with a nornial saline lotutioo, wnieh wnen allowed to
flow through the cavities of the heart readilv pemeaiea the tiiauei on account
OiT tlM peculiar construction ($ ]ol) of tlie ventricular walU. If sue b a
" wobed nut " nuicttcent heart be fed with a perfusion canula, in Ihr manner
deecribtd (§ 155), with diluted blood (of the rabbit, sheep, etc.), it niay be
iMIiiiiil to Amctional activity. A siniilar hut lew complete natotaiioD may
be wftavMcd if sonim )w umhI i&Htead uf bliHHl ; and a heart fed r^[ularly
with froth Mlppli<« of blood or eveu of ^rum may lie kept iMitlng fur a
fWT gnat kngth of tin>c. In treating of tlie akt^lelal iiitiwliii we «aw that
fal UMir caJr the exhaustion following upon withdrawal »f ibc blood^otrcam
aaicht be attrilniud either to an inade<iuate xujiply of new ntilritive material
and oxygeo, or to an accuiniilalion in the niuiu-iilar Hutkttancc of the proilucta
of muscular tnetabolUm. or M Ixilh cnuMv combined. And the aiue cod-
ndentioiM hold good for the ncn-ixiR and muwular itruc4ura of tbe beart,
though the aubject hsc nt>t yet ttciu ^udicicntly well worked out to permit any
vary definite MaleaienLs to be mn<le. It cwms probable, however, that an
impoirtant flutor id the matter is the accuiutilntion in tlte muscular fibree
maa bl the surroundiiii; lymiih of carbonic acid, and eepectally of the
*T'Tf which give rise to tlie acid reaction.
When the frug'i heart ia thus " fed " with various substancee the InterM*
tng fiwt i« brought to light that ■ome Hibelonoes, sueh for instoDoe as verj
:.■■!■•. such I'lir iiisiitiii.'
■-..I-- '.■xjuiri^iiiii, thiti i^,
Wiilllll U|)[H'!II' [Imi
!■ .p rlivrliriiii' i'iiiitr;tr--
!■: Mill- lIllV iiri' Inll^i-r.
■■■: illv ililiili's lii'VMinl [hi-
'he veiiiricli'. w'nU t\i>-
Miiiil. Fiirlli'-r, iri ili.
• - .riiL'tlii'in'il tliL' rfl;i\;i'
.'.■i]«irt.'nlly thi' vi'Uirirl--
; "i.-It'il wliiii ;i new lii-;lt
•>- - I '.i: w.' shitll ,-[ii'!ik lit' ir)
. anil [he aiiLniiiit i>t' [iii-
■ M.::v lit' llii' vriilrii'li.', varii->
- -t7. 1-vilk-IIC.T tluit illlllliitiil V
"a.^k' tltU Ionic cuiitriKiinn.
'■! with siTiirn iir cvi-ri uiih
i-'i h_v -tiiiiiiliiti'iti, lu'i' ;i|ii Ili
-. s ^(ii L'rlUl[l^. This inU'i-niii-
_ uniiljk' ii>c'iiiTV on iiiitriti"ii
st'<|Ui'nl |ii'<Hliii'ti<>ti uf aliDiir-
a" fiiriliai' inliTniiiii'in't's >t^-n
,iii>iu- chi'inii'iil siilij'iuiK'i-- in
■|U' lu-ni'['s ln'at l)v aulid;^ "ii
-. 'I' liiilh. anil [liai [iriihiililv in
^ ih*.- rhythm, nv thi- iiiiliviiinal
-. ■ ■- ■[' ihc hcarl alsi> alli'cl its Ixai ;
aiiiLiiiiit III' th<- <li»ti'ii(i<i]L 111' it.-
. -. '. , liki' ihiJM' i)t' nriliiiaiv nuisih-
■■,ii Ivy llif ii'.-isiarice whii'li thcv
-.iii'i' lhi(i;;s ln'in^' tijual. vndiraii
. , ■. a.- ill ixiliiiary niiisck'. tilt' limit
.i-*i<i, am! an Dvcrl'iill ventricle "ill
. [iriilialily tiuiptics it-dl' ciiiii[ili'ii-lv
■ ]nanlily nt' 1>I I in the vciiiiiclc
' ivs: lilt' i|naiiiity ilirown mit w^iiilil
« 'iihl III' ('ifc'icil «iih ;:rcatct' I'nicv.
■■;rii'lc i- .at the cinnnicnccnu-nl .if
;-■ aiiiii'iihii' systiilc. the smrk nC the
1 ■ '.•.■'■ is ill a measure triiveriieii liy thr
•■■. nui'lianical ili^iis anil ilulirect iitr-
■■. iieait's Inal til 111 i-]ircssnrc. When
- lie resi-liiljce I'l the Velltlienlur -vslnle
:(!. iiiiin.' Kliiiiit lliiws I ill the iiianiina-
. ■.■.■\. I'.LTtli these eveiil,- iviiuhi itiere.i-e
;'i; ex|ii'el tlial the itierease «iiuhl lie
i^ «ell as ill the I'lPi'ei- «i' the iniliviilnal
»e i| I li;ii| ilii-, < In the ciiiitrarv
- ■■, may hi- |iiii alinnst in the liirni i>r a
■.'.Verse laii'i In ihe arlei'ial [iresffUie : "
'.'- 'IV a iliitiiiiiili'in. aiiil I'all nl' iires-iin-
I'iiii*. li'i»tvei'. I'lilv hiihl» I'liiiil if ihe
VASOUOTOR ACTIONS.
20S
ra^ (ig iuuu'U If (Imw )>e previnusly dirliled, then io irhaUvor wny th«
MtMiH-praMU* be nUsed — whether hv injet^tiiitc blood or clam(>iag the Horift,
or iatnamtg the iierlpheml rtsiauuice. ihrouieh thut uoii»ii m' tb* vonomotor
Dtrres wbien we shall harv to clfvcrilM.- ilim;(lr — i>r io wbktercr my it bo
lawcKd, wtaudi clear and il4ft<l(yt invcmr rclniitjii betir«en blood-pretnare
■sd pube-rate u olwerrcd. li 'n< Infvrn^i. iharvfove, thHl iiicrrnsol blmxl-
fiw ufe cauM* a riowtni^ of th<- pi>l*r, wlton lh«^ vagi an- ititncl. bi-ctiiiw: [he
fvnit'i- inhibitory c^ntro iti thr niviliillii U xtimulnltHJ by th'* Wt^h yrvMorr,
cittwr dimply by ihv prrwun' obliiiiiing in tb« bloodvexmile of iIm- mriliilln,
or ia aoOM indirect nianiMr, a»d this hrart in ooosmuence to a «crKiin extont
toKibtted.
CMA^IEB IX THE CaLIBRK OP Tlie StiNUTE AkTERIBS.
Action I*.
V.\BOMOTOK
f 163. W» hav« seen r§ 108^ that all arleriM coolnin plain muacular
ftbm, for tlie nivot nnrl circularly iliiijHinL-iI. and itiwt abundant in, or M»ne-
timr* alin(»l rntirvly oonlint*! to, ibe iniddlu otiut. Wl^ biivc furtbtirseen
that a* ibr nm-riea beeonM imnllvr the nimciiliir rlumcjit. tu> n rule. Innioaiu
nkorvand mure prominent » Gonipiini] with tlu- olhvr Htrmciit*, niilil, in the
Bunuir artvrieii. the middio cont onsiota almost entirelr of a series of plain
Biw^br fibres nrAp|)C<t nrniind tht- iiilcrnnl coat. Kervc tihn-*, of whote
aaiiin' and muree we abtill prcatnilly »|>cnk. are distribute] Innn-'ly to the
atVno>,and a]>pi.iir (o end cbicHy in line ]>Il>xurgs around thci nmsnilnr fibre,
Wl dirir exact temiinatiunit have not a* yet boon clearly niiuii; out. By
ntcfcaaical, el«clrical, or other uimulntioii, litis muscular coat may, in tho
livBS artery, W iniule to contract. l>iiriu^ thin cnutraotiou, which has the
tlinr chamcier lx.-liiui{int; to tbe contructioua of all plain luuade, the calibre
U thr vomoI is iliraiuiiihHl. The veins also, a« ue have wen. poasen min-
(alv «temeDt>, but ihnr vary in iimoiint and dlMribution very much more
iatbeveing than in tbe anfrir:>. MimI vdoit, bovrercr, are oontraclile, and
■avrary in calibre acconliiij; la the cunitition of their muscular oleoieuts.
Viki are also mpplic*) with ticrv«s. It will l>e of adviintage, however, bi
omUar aeparaloly the liltlc n<! l(iH>vr cMicerning the chiuigca in the veins,
aail \o ODDnQe ouraelvn at pm>ciit to the change* in tbe nrtcTie«.
If the web of a frne's foot be watched under the micmmnpe, nnv individual
Mallanary will be found to vary coDnidcrnbly in calibre from time to time,
Uv ■oaMtinwa narrowed and aometintc* dilated ; and tbrso changes may
Hhl place without any obvious changes either id th« heart-ltcat or in tbe
rnl circulation ; they ar« clearly changes of the arterr itaolf. Diiriog
Darrowiuit. which i* obviously due to a cootnwtJOD of tbe muscular coot
<f the artery, ihc cHpillarics fed by the artery and the veins intonhicb th«S
Ind become leaa filled with blood and paler. During the widening, which
aSRqwnds to Ute relaxnlMXi of the muscular ooat. the tame parte are fuller
rf bUisd and redder, (t ia obvious that, the pfeesure at the entrance into
■aripTai artery reDisiiiine the uiue. moro blood will enter the artery when
iwutioa takes place, and consequently the rcfttatanoe oBered by the artery
iidiBfailalied, ana lees when contraction uccura and tbe resiaiauce t» conso-
ipmly incrensea) ; the blood flows in the direction of least reaislnnce.
TW extent aixi intensitr of tlie narmniug or nidcniiig. the ooaatriction or
Alin which mar thus be observed in ibe frog's wel>, >'ary very Inr^ly.
VuiMiooB of Alighl extent, either more or less r<^lar and rhythmic or
fal^plar. occur oven wIm'u the animal ix njiparentiy ■ubjected lo no diaiurb-
fag liases, and may be ojwken of as ^MuUneous ; mrger changei may follow
26<t
TlIK VASClfLAB MKCHASISU.
events occiirriii^ in v&rioUB pRits of the bodr ; while ns the mull of exp«ri-
meolal iiit«rfeivnci? llie srt«ri«B may become either eonslrictml, in eonie ca««e
almoflt t'> obUteration, or dilaled until tliej* tic^uire double or toon tlmti
double tbeir normal dinmeler. Thit^ conBtriclioii or dilation mmr be
brouffbt about not only by ireatnieni applied direotly to Ibe veb, but also
bv cliaages aDecting tUe nerve of the leR or other parte of the body,
lliua, Mcliori of the sciatic nerve is (tenerally followed by a wideuin^ wbieh
mi^ be alight or which may be very marked, and which is sometimes [>re-
oeded by a padding eunatriction ; while stimulation of the [leripheral Mump
of the 4)ivtdeil nerve by an iut«rrupted current of moderate inteusit^' gener-
ally given rise to conslriolion, often so great ns almost lu obliterate aotne of
the miiiut« arterlM.
Obviouifly. then, (he contractile mii^rular elements of the minute arteries
of the wch of the frog'ii fool are capable by coutruction or relaxation of
caujting dfcrrnw nr IncrMuwof thi? calibre of the artcrie*; and this cnudltion of
Con«tricti()n ordilatton may br brought about through the agency of the uerve*.
Inilci-fl, not only in the frog, but hIm), vud utill more so. In warm-blootlvd
iinimnK have we cvi<U-ncv Inat in the CB»e of nearly all, if not all, the arU'riM
of the bo<ly, the condition of the muKCular coat, and fo the cidibn- r.f the
futery is governed by means of nerves; these nerve* have received the gvo-
general name of mMimolor »(nw,
S 164, If the car of a rabbit, preferably a light colored one, be held up
before the light, n fairly conepicuoiis arierv will be seen ninning up tli«
middle linear the ear accompanied by its broader and nwre obvious veins.
If this artery be earefiilly watched it nill be found, in most inataacM. to be
undergoing rhythmic changes uf calibre, constriction alternating with ililatMO.
At one moment the artery ap|>enrB ns a delicate, hanlly visible, pale streak,
the whole ear being at the same time pallid. Atler a while the arter}- slowlr
widens out, becomes broad and red, the whole ear blushing, and many smab
vessels previously invisible coming into view. Again the artery narrows and
the blush fades away ; and this may be refteated at Humeuhnl irregular inter-
vals of a minute, more or leM. The extent and regularity of the rhythm are
usually markedly increased if the rabbit l>e held np by the ears for a short
time provioui to the olwervatiun. Hiiutlarly rhythmic variations In the
calibre of the arti-ri*^ have been oliserved In iwveral placM, t.y.,ln the
vcanls of tile mesentery anil cloen'bere ; [irolmbly they are widely spreml.
!y>motim<» no snch varintioni' are seen, the nrUrrv remain^* constant in n
condition intermediate between the more extreme wiifening and extn-me nar-
rowing just described. In fact, we may Rpeiik cf an artery as being at nnv
given time in one of throe phaioa. It may be very constricted, in wbicli
case its muscular fibre« are very much contractoi) ; or it may be dilaud, id
which case its muscular fibres are ralaxcd ; or it may bo moderately con-
stricted, the maseular fibres being cx>nlraeted to a certain extent, and remain-
ing in such a condition that they may, on the one hand, pass into stnuiMr
oanlrsclion, leading to marked constriction, or, on the other hand, into dis-
tinct relaxation, lending to dilation. We have reason to think, as we shall
see, that many arteries of tlie body are kept habitually, or at least for long
periods together, iu this intermediate conoilion, nbicli is lire<]U«nt)y spoken
of as tome conlraetion, or (okus, or arleriat lont.
§ 165. If, now, in a vigorous rabbit, in which the heart is bouing with
■dequate strength and the whole circulation is in a satisfactory condition,
the cervical sympithetic nerve be divided un one side of the neck, remark-
nblc changes mar be observed in the bluudvessels of the ear of the same «de.
The arteriw and veins iriden. they together with the small vein* and the
capillaries become full of blooil, mmiy veuels previously InvUble coiuc into
TASOMOTOn ACTIONS.
267
"View, Um <rhole ear blufthM, mikI if ihi- rhythmic chnngc» described above
««t« pcvriondj goio|; on, th>«o dow oeaae; unl, in cmiicqu^ooe of the «sira
mppl^ of wmnn bloc^, tlw whole eur beooinm <li>tinctly irnrm«r. Now Oieee
rliu>g«a uhe plsc«, or may take place, witlioiit nnr ■Ittrmtion in th<^ h«art-
Ihsi or ill tbe |;e»cral circulation. Obviougly the arlcnr* nf llic viir have, iu
iiiMmUllliliiii of tb« wctioo of the oerve. lost th« tonic c»ntrnninn whioh pre-
riaadj exuced ; iheir muioular cohIji. previously fomewbnt. cotitraclitl. httve
bMMlW quite relftXei), anil whatever rhythmic coDtmctions wi-rc pre-
Tinodj going on Iiave ceaseil. The miire rnnrked the previous tonic conlrac-
ttnn. UM ibe more vigorous tbe beArt-benta, hi that there is mn ii<lii]uaie
Mpplr of blooil to Git tbe widened clianD«ls, tbe more striking the rMiilU.
StHnettiDCi, ■• wh«n t)i« heart is feeble, or the preexist iufc tooic conUtaction b
•Iwlii. tb« nection of th« nerve produce* no xm obvious chanee.
If, bour, the upper Mgnient of tbe divided cervical sympaUielic aervo —
tbM k^ tbt portion vt the nerrc pniinng upward to ibe bead and csr — be laid
apaa dw eleeirodM of an induction machini.' amlaifcnile tmerrupled current
b« Hitt through tbe aervc, new chanms lake phice in the hlixidvewela of th«
tmt, A ■hiiri liniv after tbe application of the current, for iii this efTect there
k a html period of very appreciable duration, tbo car grows [>aler and
eeolar, nany nnall vcskIs previoualy ixmipicuous become again inx-isible,
tba maJB artery shrinks to the (hintmt i)ir>tid. iind the main retna become
eami|MiDdinglr sniKll. When rhe current is »hui off from tbe nerve, theae
«BhcU dill last some time, but eventually pan oR'; the ear rcddvnN, bluahes
ooet mort. and indeed may become even reilder and bolter, writh the veoMla
Bwn Blled with blood than before. Obviously the current hfu generated in
tba emical sympatbeiic nerve impulses which, passing upward Ui the enr
aad Boding ifaeir way to tlie muficutar coals of the arteries of the luir. have
tbnea tbe niit*clM of thow^ omIk into forcible conlmctioDx, and have thus
hna^t about a for<rible narrowing of the calibre of the arteries — a forcible
tnmuietioa. Through ^h^^ narrovri.-<l vi>iu>lrivtctl arteriat lesN blood tinds its
wvj, aad bcnor the [wlcntw and colitnm iif the ear. If the impulsee thus
C'aied be vi-ry strong, the constriction of tbe arteriw may be so great
Uw iniallMt iguantiiy oiily of blond can make ila way through them, and
lieiarmay become almost btoodlen. If tbe impulM* be weak, the coustric-
li<« bducM may be slight only ; and, indeed, by carclnl mnnipulatiou the
■MTfl may be induced to send up to the ear impultiv only juici xulBciently
Mn^ to restore the moderate tonic conetrictioD which existol before ibe
aim wa? divided.
We infer from lh«ee exjieriments that among the various nerve fibres
■akiDtf up ibe cervical sympAtbetic, there are certain fibres which jMwiing
apwafil tu the head become connected with the arteries of the mr, an<l that
line tilirva are of ruch u kind thiit impuli^es };eneraled in them and jnmiug
gfwsnl U- tbe ear, lend to marked cuuiraciion of the mu*cular fibre* of the
ancrie*. and tliu.i produce eoiMrictiun. These fibres are vaaomotor fthn-a
^ iw the bloodvewels of the ear. Fn>u the loc« of tone, ho frequently follow-
H Bf section of tlw cervical iiympatlieiic, we may further infer that, normally
H wiag life, imptibet of n giiitle kind arc eoiitinuall^ (lassin)- along thceo
H Ara*. upward through thr r4;r\-ical aymjiathetic, which impulses, reaching
P tk arteries of the ear, maintain tbe normal tone of thoee arteries. But, aa
••■id, iheestMenceof this tone is not*o oointant, and tbfse tonic impiitses
M not fo conepicuoits as tbe artificial constrictor iinpubea generated bj
itimlalioo of the nerve.
1166. Tbe above nviilts are obuined whatever be the region of tbo
Mrrienl aynipatUetic which we divide or itimulate from Uk> upjier cervical
268
TIIK VASCULAR MUCIiAKISM,
^n);[1ii>ii In Ibe lower. We m«v. lh«^rcfor*, tlc«crib« ihwp viwomotnr im-
pulse!! ua [iaBsii)g upward Irurn llic loner ccrvicnl ^ii^jlion nlone the cervical
Myiii|)iilhei!i.', to the upjiirr mrvictil ganglion, rrom irhich iliev iwue br
liniiK-hcA which uUiiimiely tiiid ihelr nav to (he car. Itut llitM« impulBes do
not mart fnnti the Imkt cenricol gnnglion ; on ihe coDlran\ by repoitting
ihe i'xnrriiii?i)t« »f ilivEjilon nnd HtiiuiiliilioH id a aeries iif iiuiiualh, nv mar
tnict itic nnlh ol' ihrse imiiulHcs fntm the lower t-ervical ^iigtlon (Kig;- 97)
ihroiigti iho annidu^ of Vietiswos lu ihe gaii^liuu lullatuni ur firM thoracic
tiu«HA> lU.usraAtixa 710 Patih trr Vjv«<«<an«XToa Kiiiaai
AixiKit TirE ctHrtcu STHfittHma a>i> (rin w) Tua ApommimU'
9ria*aiMC
Ami., >il<T)' ol«ar. 'i.L'.» . nipetlor orrrlotl puulWu; XIABpl..
ii[>)n' nuuurbni] |i*rt of al-lomlDii) a|iluioIiulc uarm: V.M.C «■>»-
nuilor ivuiivtii iuviIiiIIh. 'fliv oilw raltmicn *n (h* mmt ta in
PIr, W.)lin. TliB t«''» urihe oiDnriclor niitt* an ibowa hj tk*
armwi. Ttu itoiiail lliiu Iti iliD tjilnal cofd. 8f^C ii to tndkaM !&•
roiHBe (if tiiiuLHctDf lin]iul*a dona Ui* O0«il AolD Uie <
(<uiilr«iii Uic iHMlulla.
ganclion. aiid theuce either along ihe ramus commuuictuis (visceral branch)
to ilifl unt4-rti>r rniii uf tbc second darutl nerve, and thus u> Ihe HpinnI cord,
or lower down along the thorncic Hyinpaikeilu cliaiii, and lliciice by iXher
rami com nniiii I'll iit*ii lo sonic other of the upper doraal nerve*, and ibu*
to lh« spinal cnnl. The pulh talien by the*e vatiomolor iinpulw^ for ll>c
Mr b in fact very Mniilar to that uf tht: ntignK'nior tibren for the hrart icf.
FiR. 9(i) from the spiual cord up lo iho niniidiiH of Viciumnit and i« the
lower oerviral ganglion ; hut there th'-y part company. \Vv can lhn:< tracei
tlwoe iiDpulaes along the cerrica] svmjuithetic to the anterior root* of cvrtaia
doiMl nerves, and through lh«ae to a particular part of tbo BPUial conl.
vhfK we will for lh« present leave ihem. Wc may accordingly speak of
vaaoiDotor fibres for tlu- car as pawing from tbo doraal ipinal cord to the
ear along the track just marked out ; stimulation of iticw fibm at ihair
origin in the spinal cord or at any part of their course (along tlie anterior
raota of the second, third, or other upper iloreal nerves, viseemi brancbee
of lho*e nervftv ganglion stcllatuni or upjier part of thoracic syiupatbetie
cbain, annulus of Vieuasena, etc.) leads to constriction in the b'looavesseU
VASOMOTOH ACTIOKS.
309
of Um cat cl' thai nda; tand ■ection of ihne fibm at any part of tli« mme
wnm uod* to nboluh nay prcviotitly txbtiii^ tonic ooiwtrictioti of the
UoednsMb of tbo *ar. though tt)i« rllKt is not so coosbint or iiariliiog as
tlwt iif rtliDakUon.
6 167. W« nUMt now luni to another caae. In d«alin); n-ilti fliK«etion ire
•ImU have to itudy tli« Mibmnxiltury salivary gLaiid. We may for tLe
pCVMt ainply aay that this is a (•laudular inafs well supplied with blood-
riaib,aad powcaJng tt double nervous supply. On the one hand it reeeiree
flbm ftom the eervical ayinpnlhotiu. l-'i;;. Ot5 v.ei/m. (in lh« dog, in nhidi
tlkc Hfecta which we are abotit to deacribe are bat wen, the vafcus and cervical
rni. w.
*«■>>■■ iTur Rtnaonittcui nr tiia BrnmixtuAav Glixd or m* Doa, vim tn KmvM *ki>
BlJMITVMUA.
TWiUmc^Idx ku tiNd mult (HI ■■Knliiul Ijlnf on it* tack, tint iIdcciiII IhB puti iIidwd tn Qtc
afB*«adai Mon Ihnnuir oacfulntof Tt«r, ibeBKiuvlaet iioi kIv* Uie ciarl anuonlio] rvK-
*««r tt> n*«Hl MnwMN*.
»t^Hi«— lm»lltMy«l»jna. lnu>tbBdiici[—. Jml which aunolA bu bran Unl llwntb-
MaHl (iHid and duet uo nutibuoii. aJ..!!/. Tlic Uiik'i'I tiniiicli i-rtlwMli oorvo. tl»«(*n sj.
• fM^lattalcafw^ <*,f..rAf .rb r Thocliooli trmr>»l. I'bi (•!( (A. I", li rxxmllnc Roni Ik*
tiM Mn«, •> (ft. r tt btcona Rin>)IiiL>d wlih Ibc tinituil i>.r., uiil ancnfard illnnclnc j«nii M
At l> Iha lUiht alonf tlniliKt: ih* «nilliii»riuii of Ibv nirvptn nuDistiy wilh Ihr llninuil. n.l..t>
■» (J Tim wlw I in> IT »"■>""■*'" I* "' — J-™' «— ^- a. cor "na ouiMUl krurj-. Iirn
■bvof wMi4i, ■.•!•■ «.u«l r, ■u.p., IWB (o (be ■nlorkii luid powwlor t«n> of lb* yUnd,
< ■■ tts Milwtot and ixMarlur Tdn* frum Uw«land, Atlllus lata r./.ttui Jugular t>Iii. r.i9iK.Tti«
I i«(iB and (n>iF*UMU<i iniok*. g, frf, i. Tbe uppii nrrlotl aangUon. l*o bniiialic* of
■iiK ■ ptdn (oj-t onr Ihs llulal utcrj. an dlilrlbutnl (a.ivKni.i alont tbo Iwn
I Id Ih* aoMsm and (oU^or ronlcn» of Itivsland.
Ha WW «a todinaw i>a dlwwllwi ukvit by iLeaEnKiiii Myulta dsrlni mztx iilaiulatkni pIU*
nar nwuid (a IJic loaln br Um Ungual aiid dcaomd b; Ike cborda ljnii«iU.
)*tBfaihrlic aro encJoewd in a c<>niin<iu sheath »o as to form what appean to
Maiinclv trunk), which rea«h the gland in eoni|>aiiy with the arteries nip-
plmgUivglMid (N.nwm. fm.). On die oth«r hand, it rcceivoi fibnv (W>ri a
^alTBervv callnl ine chorda tympani (cA.fJ, which, springing from the
VTWih cranial ( iWiid) nerre, crown' Uh- tympanum of the tuir (henoo the
Mat) aitd. joining tbi^ lingual branch of the fiAh norvr. rum for «ome ilin-
Uan m compaDy with Ihnl n" rvL-, and then ends partly on thi- tougtie, and
270
TUe VASCULAR mkcuakish.
partly in k small nerve which, leaving tlia linriMl nwre Itefore rcttchio); tht
longucruM alnng tlie duct of the tubnudTwry gUod, mkI is lost in ihe
ButMtaoce of the glmM) ; a small branch is also given 09" to the Bublioi;iial
jllund.
NoTc wiieii ihe clitirdu tympnai ia siniplv divided no verv rvin&rkable
cUuiii;eti ulce place iu the bloodveesets of me gliind, but if t}ic peri|ih«ral
eet;i>ic<il of the divided iiervi?, llinl still in counection with tli« ulaud, )»
atiiuuliit«*l very luarkt-d tchuUs fullow. The suiall arteries <if the gluoA
become Tcry much dilntei) and the wbole'glond becwoea Btisbed. (A»ki>
tball ne Inttir ou tli*> gland ai the jiame time secretea aaliva conioudy, but
thia doea not cuiicerii ua just now.) ChiiDijee in the calibre of the bloUiro-
wIh are of cniirac nut *<> reinlily mcu in a compact glnnd a« in n iliiii pjctentled
but if II fim- tiilit^ \>c [ilticeil in one uf tliv small veini> by which the blood
rnr
rettirna from the gland, thv eBeotd ou iho bl(ir)ilvi>wi*lH of iftimulutin}; tbc
cborda tym|iiiini Ikciihic very cbvicu?*. Before (ttmulation the Wood irickla
wit in II thin nlow nin-ani of a dark rcnoua color; during atiniulation ibe
bluod ni»h(9 out in ii rupid full streum, oilcii with a distinct pulsation and
frc<|Ucnlty of n color which is still scnrlit and arterial in spile of the blood
having IravcnK-d the capilluries of the glnnii ; the blood rushes so rapidly
through the widened bloo<lveiwtels that it has not time to undergo cnrnpfetely
that change from arteriiil to venous which normally occurs while the blood
it travervitig the cKpillaries of the ginnd. This state of thing* may continue
for SOBM time alter ihe sriuiulation haa ceawd, but before long the 6«w from
the vans slaokeRs, the iHuiiiK blood liecoaiea darker and veooue, and
eventually the circulation beftmea uonuul.
Obviously the cborda tynipani contains Rbrea which vre war ii|>eak of aa
" vasomotor," since stimulation »f them produces a change iu, and bring* about
■ movenieni In the hliiuiive(«ela ; but the change produced ia of a cbaractvr
the vm* opjMMtili; to that produced in the Moodvewivlit of the mr by simula-
tion of the cervical sympiithelic. There stimulation of the nerve cmtued
contmctiou of the niu«cutar fibres, constrict ton of the small arteries: here
stimiilalion of the nerve cniist^ a widening of the arterien, which nidening
is undoubtedly due to relaxation of the musailar tibra*. Hence we most
distinguish b^wcen two kinds of vawniotor fibre, fibrea the rtimulation of
which produces constriction, KOAo-eonttriclor (ibrus. and tibree the atimulalion
of which causes the arteries to dilate, va»0'dUali>r fibres, the one kind being
the antagonist of the other.
The reader can hardly fail to be struck with the analogy between theae
liro binds of vasomotor ^hres on the one hand, and the inhibitory and
augOMntor fibrea of the heart on the other hand. The augmentor cardiac
Hmta increase the rhythm and the forc« of the heart beats \ the vaso-oon-
Kttiotor fibrea increase the coDtractionsof the muscular fibres of the arteries;
the one works upon a rhythmically active tissue, the other upon a tiaiue whoM
work is more or lea* continuous, but the etTeot is iu each case similar — an
incieaM of the work. The uihibitnrr cardiac tibr«« slacken or Ntop th«
rhythm of the heart and dEminish the tieat* : the vaso-dilator fibres diminish
the previously existing coiitructinn of the muHciilur fibres of the artcrin so
that these expand under the prewua- of the blood.
We must uot attempt here to di*cuwi what is the exact nature of ihe pro-
ctw by which the nenoux impulses poising down the fibres thus stop coo-
traction and induce relaxation, but wo may wy that in ull prdhahility ibe
proceee, whatever be its nature, is one which tnkc« pliue in the muscular
fibre itself on the arrival of the nervous impulse, and that (hen^ is no need to
pn-wppoM the existence of any special terminal inhibitory or dilating
nervous mecbanism. We have repeatedly insisted that the relaxation of a
\
VA30MOTOB ACTIONS.
271
I
}
r
fibrv i> a* tuuch a complex vital procns, i» u Irulir th« reeuit of
DoUmu of iIm muHGular AubstiiDce, as llie ooiitractioii iMelf ; and
tbwv ii A oricti tio rawoa wlir a uervuu* iiiipuNo >hoiil<l not Kuverti tlio
liinwr u It doc* the lulter. AVl- tuay, jMrhiijw, ^i I'lirtlier aiiS M\y ilint
rcUxatNOi need not b« cuuHd<er«() lu tbe iti«rc iiutloiiij^ of a citutrm-lina ;
tb»t %h» actioo of dilator fibru in ool nMHwnrily Itmiie'l Ui ihc rrniuval of
B prvvioudj cxipttuj^ cowiricliuii. We miiy iiiuigiiic n tiiuimilur tibri' lu
—bjacl toUirnctiuool'tvo <>[i|Hitiiiig l<im» — ttiv uite vioiigntiiie. rvinxirig. or
dOatiHI^ : th« otlicr vbortCDing, ■.■nuimcling. or ouiulrictiiig. When nviilhtr
i» in acUoo, or «bni thv too ati; («ii]i{i(ill»it, tlio fibn it at rCHt, Deilhaf
nlaxiDg RorcoDtnu-ti»g; nlx'u iiiieacUaluiiv, orwh«a one acts more pontes
fullr tluui tliv othvr, itirii rrUxntioa, elougnlkiii, dilatiou. or otiivrwiae coii-
trw^tun, sboru-uiii);. LviU'tric-tioD, u tli« mult ; wu liuvo )>rob»bly a* muctl
riKht tu (uptww rt'ltixaltou tt> bo a iiitvasiry nntvi'eilcot of oonLriiction aa tO
■uppMc couuacliou to be n uvccaaity uitocnJvnt of ivIaxatioD.
( 168. Uut we rauBt ralurn to th« vaMunotor twrrm. The oerviod sjiDpa-
tbAir (XiDtaitM vaaoconelm-tor fibres for the ear, and vk may now add for
•tbtT rafioBtr alao of the head aod t'licc. TIiuh the brundiea of the oervtcal
i|MtliMic. ijoinji tu the «ul>mnxillary inland of which we juat epoke (Fiff-
n. *jtit. am.), euntsin vaeo-eonslriclor libnn for the Tewejs of Uw gland ;
•llinuUiiun of tbeae fibres producw on the vessels of the jfland on elftcC
exsctlT tbir opporile of that prudticed by BiimulatJDO of the ehorda tjrnipani.
But l(> iki« paulicnlar point ire aball liave to return H'h«n we deal with the
ylaad lu connection with dJ);ettiiou. A more importinii fact for our present
parpOM b that the ecr\'ical evnipulheiic appenni U> ooniaiu only vii»o-4.-i>n-
auiolor fibrea; if we put oude m exceplioiiul and doubtful the rvxult uf
certain oheerrem who obialtied vaau dilator cdccLn in lliv nimilh and fnvc,
v« any aay that in no re){iou lu nbich tht? libm of the crrviciil aympathctic
•na distributed can any vavo-dilator avtion l>c observed a» the result of
•tlinalation of the nerve at any [lart of il« course. In the chorda tynipani,
Ml the other luind, the vuwniotor tibn^ are exchisivelr vnsudilalor fibres,
and ihts is true both of the part of the nerve eudin;* in the Bubmaxillary
■nd *ubtiD|[ual gland* and the mtt of tlin ending of the nerve in the tongue.
i?(iliiulatioD of the cliorda lympani (a» far as the vosoiuolor lundiona of
tba narre are iionccrned, lor it hoa, as we flhall »ee, other fuDciions) at any
patftof lt> courae, frum ii» Icnviiig the facial nen-e tu its endings In the
nogn« or gland, producee only vaso-dilator etleet». never vaso-constrictor
Wkh many other oervee of the body tite case ia dilTerent. In the frog
dhrliiMi of the sciaiU! aarn teada to a widening of the arteriui of the web
«f tbt Cmx of the aane tide, and stiniulaiion of tlie peripheral end of the
MTT* fiiscs a constriction of the vessels, which, if iht; Miinulatiou Iw strong,
naty ba ao great that the web appears for the time licing lo Ih- deruid of
blood. Abo ill a maromal diviHiou of tfai^ Kialic iutvc causes a Himtlar
widnting of the small arteries of the akiu of the leg. Wheni the condilioii
uf Um escalation can be readily examined, on, for iiutanee, in the linirloB
Ulboftbe too. especially when these aro not pigmi'ntml, thevesscJs are seen
to be •litalc«l and iitjecteil, ami a thennoneter placed between the loot shows
a rise of teiii{ierature aiuouuting, it may Iw, to •vvemi degrees. If. morv-
«>r«r. the peri|tli«nl end of the divided nerve be stimulated, the vcwels of
the skin beootnv cvnttrictvd, the «kin growi pale, and the Iciuperature of the
flat fclU- And very nmilar mulls are obtainctl in the fore-limb by diviuon
tad •ufaMqacnt stimulation of tlw nerves of the brachial ]>lexus.
272
THE VAaClTLAK UKCIIAM8M.
The iiuunliiy of blood prtwot In ibp bloodvcaiela of the mamnal, thongh K
niaj AODiclira^H be oburved directly, tutu rm)Uontly to be d^lcrinined indinctly.
The (ompcrHtiirc nt' pasnivc i>triicturn> lubifCt Id cooling inllucnc**, ■u<^ll ut the
nkin, i» tnrKply ilependcnl oii ihr Miipplv iil' bUtod ; the inotu nbunJanl the Miiipl*
the wftnner i^i? pnrt. Ileiice. iu tl]«ae purts variutiuiis in the qiianiiiy <if blixKl
may be inferred TrotTi Tmifttions of temperature: biil in dealing wilh more ik-iIvi^
Hi (ucturos there nre i>bviou»lf •ouroos of error in the imuibility of thr treatment
adopted, ouch as the ■UmulBtion of a nerre, KiviiiK riito t'i an incrtuuc nf tcrnpert-
lur« due tu increiuwd metaboli«m. indepwident i>f variHilooii iu blutKl-supplv.
The <|uaD(iiy ut blood mav also be determined by Uie pktIiyimoffTttph. (o Uila
InatTuinent a pari of the body. «u«h m the arm, is introduced iuto a cloaed cham-
ber eilwi with Hniil, rx. gr.. a Inrge glas* tube, ihi- (ipeninK by which the arm ia
inlroducL-d hthia securra with u ultiul cuniitcbnuc uieinhrnne. An inprtAao or
d«crcac« of blood aeni Into the arm will lead to au iucrearie or dcicrcBM of the
volume of the arm, and this will make ilacif fett by an increase or dimlnutiMi of
preMUK in the Huiil of the ciosed chnrobct, whioh may be repimwed and meaa-
urrd In the unual wiiy. Wo •hall have to iipcHk again of n luodilicntionof thla
iiMlrumeat «'b«u w« are dealing with tile ktdiiey.
^ far the reeults are quite like llioae obtained br division and atitnulation
of the cervical sympalhetie, and we might infer lliut t.bu M.-iniii- ner%-e aiid
hrachini plexui coDtain vaeK^eonsirictiir fibre* for the vectebi of the sktu of
th<^ himl-Iioib and fore-Hmb, vnMi'dilaUir fibre« bdng abnot. But «oni^
tiiitra u iJi&Kiit revolt in obtninerl ; mi ntiniuliitinf; the divided acinlic nerve
thv vcaMila of tlie fo"t an- not t-unatricttfl, but dilaluil — |>crhniionttielv iliUted.
And this vn:«ii-diiiilor ailion is alnuttt sure to be nianili.-»l«'d when tlie nerve
is divided, mid the pciiiihemi stump Bliranhiled Ajnu- dny» after diviiion, by
nhich tim« comnicucing degeneration has l>egnn to interfere with the irrita-
bilitjT of the ucrvo. For exiimple, if the sciatic be divided, nnd some days
afterward, by which time the Hushing and increased Icmporniure of tbe foot
foUowinji upon tbe aection bus wholly or largely passed away, the jKripheral
Mump be atimulated with an interrupted current, a renewed Hualiiuj; and
riae of teroperattire iit the result. We are led to conclude tliat the si-iatic
nerve (and the aaine holds ^ood for the brachial plexus) oooiaina both \-B0o-
oooatrictor and vaao-dilator fibres, and to interpret the varying rmilt a* dut
to varialit'NH in the relative irritability of tbe two neu uf Hbrat. The con-
Mriclor fibrca ap|)eur to predominnce in thuao ni-rvea, and henc« vonittrictjoo
is th« more common nMiult of stimulation ; thv constrielor fibm also *PP*>r
to be iDore readily alft-nled bv a tetaniMng current than the dilator nbrce.
When the nerve after division coinmcnciv to d<«cneriite. the cnnclriptor
libm lose their irritability earlier than tlit; dilator hurtv, so thai at n certain
stage a stimulus, such ns the interrupted current, while it fails to atfect the
OODStrictor fibres, readily throws into action tbe dilator fibres. The latter,
indeed, in contract to ordinary motor nerves (§4-1), f\-tain their irritability
after seotiou of the nerve for very many days. The result is, jxrhaps, even
still more strikiuK if a Dteehani(»I rtimnliia, <uch as that uf " crimping " the
nerve by rejieated Miips uith the teimon, be employed. Exposuic to a low
temperature nj^in aeetua to deju^eas the eonstrictore more than tbe dilaioni ;
lienoe, wbeu tbe leg is placed in ice-cold water stimulation of tbe sdatic, e\'en
when tJie nerve lias be«n but recently divided, throws the dilator only into
action and pniducea Hushing uf the ikXa with blood. Rhythmical Mioiuta-
tlon, nioroovcr, of even a freshly divided nerve pmducta dilation. And
there are other fticta which support the aauie view that the iivtatic itcrve
(nnd bracbial plexu») ontnin^ Wh vaao-ooustriclor nnd vaao-dilator Bbrttf
whii'h are diflcreiilly nflbclcd by diHereiil circumatani^c*. Wc may jmint
out that tbe caae of the vagus of tbe frog is a vcr^- analogous one ; in it arc
l>o(h canltac inhibitory (true vagus) ana cardiac augmenlor (tynipnthclic)
TASOIIOTOR ACTIONS.
278
ttbm. biit thi' funnier, like llit; vtw^eonHiicXor fibrtv in tbc scintic. arc
fim)<(iuinBDl. nnJ vfivciiil nuuiDS nie raciuiretl h) altow Oto prcMDcv of Ihe
KlU>r.
In ll»o fplanchriio iMTre i iib'lominAl uplnnchnic) which tiipptu* fibrw to
tlie tiliMxIufocU of wi targe a piirt of tho nliiloiitiniil vioccra, tbpra is abiiD-
4mDl •vidvnn- of ihp preKiicc of vasn-comtriclor Hbres. but ibo prceencc uf
TMO-ditnior fibn'c liiu uot yet been #hoiin. Division iif this nerve leads to
■ wulroiug of tbe bloodvnwls of tlie iibiloniin&I vi*cern^»tiin illation of itie
ncrvr In n con strict ion ; and, aa we shall we. lince tbc nnioiini of bloud-
finib ibii* fpivenieti by lliis nerve is very large indeed, interference eillicr
lo lJi« tine ilin<cti«n or llie uiber witb its vHBoniotor funetioiie produeea very
rkoi nntlte, out only oii the cireulaiiuii in the abdomen, but on ibe whole
»]»rtytUm.
In Dcrra jcoiog lo muadea voao dilKtor fibres preitomiiiate : indeed, in
Umw the ^neeace of any raso-ootutrictor fibres ut all has not at pTesciil lieea
Mtirfaetonly eaiablubMl. Wli«n a mu«cle (.'uutraclti there b alwayn an
iBcnaaed flow nt blood Ihrou^ili tlio muacic; lliia oiny he in imrt a uvcre
■Mchwiical reault of the diunge uf form, ilie ^liurteoiuj; atkd tliiekening of
tbe 8bn* opeiiiag oui tbe mtnnte IiIoucIvvkwIh, hut in not wlirilly, luid prub-
ably Dot nvtn lar;g«ly, tliim prirduct^l. A notable feature of va»onaotor
flbiW M ibai, iu rerv iDnnr c*m« nl all «vrJilii, their action id not nflocud br
■OmII or modcmti' d<Bc» of nrari *uc-ti ii« r<'iii]«r the niolor nervca of HtriaKa
mwde |iowerl«». ThiJ». in a frog plaotx] niitlcr the inflnenct! of n motl«nit«
■■iniiiH of umri, ftiinuIiUion of K nervv gi>ing to a miitcic will proditc*
T— pmcior »0tc1« iinuccotnpiinifil ami iitioUcured by any eoDtraclion of the
■riatvil fibm. By plncing a thin miiwrlo of a frog, auci as the mylo-liTokl,
oiKkr tbe micnaconc, and witiching the calibre of the Kninll ariericK ano tbe
draahuioa of the blixHi t]in»ij;h them while tbe nerve i^beinf- atimulated, the
wUeniBg of ihv bloodvewveU aa the reault of the fltimulation mar be actually
ofawmd. Tbia ex|)eriment appe«Ta not to aucceed Id a mauiuial ; and it hna
bean anggeoled tliat wlivn a niniM-k- euntraeta soiue of the chemical pmilijeln
ai tbe laMaboliam of the mmclc may. by direct action ou the minute bliHxI-
apart from any mTviiuK ageiicv, lead to a widening uf thuae blomt-
: tltia, however, ia doubtful. \^ ith regtird to vaxi-ciHiiilnc'tur tibrot,
tba onlv •videnc* that thryexiot io niuitela) in that when the nvrvo of a
nittcle It iltvidH the blooilvmvl« of the mutcic widvn, fouiewhat like blood*
va«rk of the car aAcr division of the cervic*! ayiupnthctic. This euggcvta
the pmonoe of vafir constrictor fibres carrying tnc Kind of influence which
we cwlJn) Ionic, leading to an habitual laoaerste eonntriction ; it cannot,
hamvnw, be ragarded by itself as conclusive evidonco; but we must not
JlBBil ibe (nalter here.
Smakiog generally, llien, most if not all the arteries of the body are mp-
pJiu Willi vasumoior libres niniiin^ in this or that nerve, the fibres being
«itber Taso-MoMrictOr or vaau dilator, and Mime nerves containing one kind
«f flbna only, some botli in varying jiropiirtion. Alinoai every nerve in the
body, tbnvfitre, muy tie lu»k<.'d upon aa inllueueing a certain wt of bloix)-
TOirb. M goterning a vaaeuliir area, the area heinj; large or small, and the
|ov<fn(nmt being csehiKivcly roiutrictor or exclusively dilator, or mixed.
TAe Orvne iff Va»o-fomitriet»r anti VatihdiMor FSirf*.
i 189. Itoili the vaso-cou>triclor ami the va«r-dilali>r Sbrra have their
urigin in the cvntrul nervous system, tbe «pinnl ct>rd, or the brain, but the
I of tba two *ets apiwan lo be very differcot.
274
THK VASCL-LAR HRCUaKISI
111 lh« mnniniiil. bb far n« wf know iit jircscnl. nil ihe v-n*o<^>Mt rid rtrt
JibrM fur tin.' whoK' Ixxly liikt^ liicir 'irigiti in t)ie miiMlf re^ii>ii of itie ttpioal'
cord, or rather Iwivc ihiy spinal cnrcl hy llie iioivw Iwlongin^' In tLU nii<ldl«
regioD. TiiiM in ilic iV>i Ihc vus» cinmtrifUir filtros nol onlj for tlip trunk
but ri>r tlio limbs, iiciid, liicc, timl tail, IcKVc thi> fpinat coixi by tbe itnterior
root« of the spinnl tiprvc§ rcnchini; from about the second doriul li> the second
lumbnr nerve, bnlti inclusive. UuRiiini; in the cose of eneli nerve root to
the mixed nerve trunk they ps«B hIod^ the vieceral branch, white rnoitis
comniunicnus, to the L-haiu of Hplnnchotc Rnii|>liu lyini; in ihe ihorax utd
abdomeD — the so-called thoradc and abdominal sympaltielic olicin (Fij;.97).
From theee gSQclia they rea<^'b their deelination in various nuys. Thus^J
tboee (foing to the head and neck paee upward through tbe annulus of
Vieusaeus to the Imver cervical gatiglioti, and then<-c, as we hnvi* «eeii, up
the cervical dyniiiaihelii;. Those for tbe abduminal vSaoera ihu* ofl* 'm bj
■iotilar way to tht- abdomintil sphmuhiiiv iiervea, Kij;. 97, n6'/. 171/. TboMl
dMiineil for the arm lake thdr way by thu retmrrent fihrei< (^rnv nini! cini*
inunii-unli's) (Pig. 4S, r. v.), and so reach the nrrvcM nf the bracoiiil plesuN;
wliilc lli(«e for the hind leg joiue in a itiinilar way ihrongh wini*- portion of J
the nhdoininat )iym|<iilhclic be-iore they join the nerve* nf ihu ticJntic plt^xuiL-^
And the conslriclur fibre* of the skin ol tho trunk probiibty mtch thu ■pinnl
ncrvw in which they nUiimitdr run in n niinitnr niaiinpr. All the vaso-
constrictor Bhr». whatever iheir destiiinlion. Irnvc iht' spinal cord l>r the
anterior rool« of spinal nqrvet. and then pae<iny: through the np|iro|irint«
visceral branches, join the thoracic or nbdomtnal chaiimf spUnchnie ganglia.
In these ganglia the fibres undert'o a remarkable change. Along the anterior
root and along the visceral brancn they are medullated tibrcs, but long before
tliev reach the bloodvee^ls for which they arc destined ihey become oon-
Diedullated fibres; ihev appear to lose their inednlta in tbe »\g[em of
uilauftiiiii- ganglia. We may add ttiai in the anterior roots am) along tbe
viiceral branches, white rami ciuumimicantee, theie fibres are invariably of
small diameter, not imin^ than 1.^ » to :i.6 u.
^ 170. The courye of tb<- viu>o dilator fibru appears to bo a wholly ilifl^rciit
one, though the delaiU have nj> Vfi been fully worked out in the caw of few-
of tbe fibres only. It is cbieHy in ihv nirrves bflunging to ihe craiiinl aod
ncral regions of lh<' central nervous system uiivnce, ns wc have nen, no
vaso-conslrictor fibres are known to iwrn-. that the ononw of the vaso-dilator
fibres has been succcsffully traced. Thus the vasodilator fibre* fiir the sub-
maxillary gland running in the chunla tymnani may he tractvl, a* we bava
seen, back to the facial or seventh iieri'e: and the continual ii>n oflbe chorda
tyinpani along the lingual nerve to the tongue contains vasodilator librca
for that organ; when the lingual is siiutulatciJ. the bloodvo»vU of the tongue
dilate owing to the stimulaliou of tbe conjoined chorda tympani fibres. Tbe
ramus tympanicua of tbe gloaso-pharjrogeal nerve contains vaso-dilator libr«B
for the parotid gland, and it appears probable that the trigcniinal nerve
oouuiiiis vn.*o-dilator fibres for the eye and nose and po<eibly fur otlier parts.
Fu the anti-rior raoia of tbe »ec"nil and third sacral nerves run va«o-dilator
fibrw whirb \>»s» into the wi-called ueri-i erl^tnlts, the nerves, stimulation of
which, bv leading to a tvideiitng of the arteritis of tbe peuis brings about
tbe en-rlion of that organ, the effevi l>eing uMisied by a simultaneuus bind-
ranee to the venous untAow, Though vasodilator fibres are, as we have
Been, present in tbe nervM of tbe limbu, and probably al^) in thuie of tbe
trunk, the investigation of their several path* r» renderfti very difficult by
the oiucumitant preeeiicc of vaso-eoiislrtclor labn-si. There are some reaaniis
for thinking that the vaso dilator libre-* in tliiw nervev putKue a direct GOUne
from tlie spinal cord throu<;h tlf lUitcHor sjunul roots, and thus afford a
VASOllOTOR ACTIUN'tf.
■275
tnn with the coDilrirlor fibn.« of ibe nme n«rrw, whinh, at ire bare
iBon. Mko > rmiiKblxMil niiinw.-. piiMiiti}: into the »pl»nchuic sysieni, l>ef<tre
Ihty JMB tbv norve trunk. Our inl-irnniliuii. Iionev-Qr, U loo imfierl^i^t (o
bUow an?' Vviy poeilivc HtmeOK^nl to be mu'le. Accepting; thU viow, how-
*v«r, w« may say that while all the va<ri>'Coii<(riclrjr fibres, a.4 I'ur iis vic
know, come from it iiarlii'iitar, lliiitigh consiilerable, pari of the sjiinal ruril
•IhI pBW into ibe splaiirhiiio system uii their way to their Mveral deitiiiii-
tioaa. ibe vaao-dilaior fibres arise from all parts of the jipinttl conl as well
■a fr^Mn the inealulla obloo)^(a, aod puraue a mure or leu ilirecl omw to
Ibeir (le4inKti»n.
Kuribw. while the vaso-diluior libra, a* they leara the central ncrvnus
antem. are. like the va*i-ci>itMri(.-tur llbriM. tint; inmlullated libr<», unlike the
vaao-conHrietur* iher n-taiu Ibeir luvilulla for the ynuiler {Wrt of tlu'ir
«oanv, and only Iom ii m-ar thvir lertuinatiou in the tiHiie wliow blovd-
WHlt ibi'v Kri[)ply'.
LaMlr, while in« riiM-con«tricl»r fibreii, a* in ih« cam nf the cervical
■TRipalbvtic, of the ntdlDiiiinal >|iliinchuic, aui) of the ncrvni of the skin,
ax»i] |irotMi]>ly tn nil iiwrs. utv mirinully in a ulato of moilctatv activity (ta
Imw aa liwy mninin in i-onni-ction witb ttie oeiilral in-rvout nyslcni), llie
Boomu aclirily maialainini: that nKHlerate coni*lriclioii which wo spoke of
abnrv a« "toiic," the vavodilntim nppear to fioaiMi no such continued
activity. i>ccttun of vaR»coiisi riclor hbr«-« lead* to loss of tone, diminutinn
of cimrtrictioo, lA»ting, as we shall eoo. for some coniirleruble time: but
aecliun of vaso dtlaion, nccordiu; at all events to must observeni. doe* not
iimti in aDalot;ouD eonslrictioii or iliniiniilinn of dilation ; all that U obierved
ia a tnasieoi increase of dilation due |tfub,tbly to the section acting as a
tnuatnil stimulus tn the nerve at the plscv of section. But before we study
tb* uaa made by the ceiiiral uen'ou^ syitem nf va»imotor ncrvt^ii, it will be
bcac to cnn«ider briefly some teaturea of
Ci 171. A <rery Utile coiuideration will show that vasomotor action is a
■ Inpoftant factor in tbe circulation. In the lirst place the whole flow
bloiNl in the body is nflnptGd to unil governed by what ire may call the
fiMH-al iotu of the arteries of the body at large. In a aamal oooditiua
of iIm budy a very ^*T\fe Dumber of tlic minute arteries of the bmly are in
a state of toaie, i. «., of modwate, coiilrii<.-tion, and it \a iht narroniug due to
thia eontraolioti which form* n lar^e item of that peripheral raei«tanoe which
mm bave seen to be one of the k<'c^<» factors of blood -preasure. Tbe Donnal
Munl b!(Kid-i)rui»ure, and, ttiert-fore, (he normal flow of Mood is in fact
r Smodent on tbe " general lone" of the miuute arteriea.
fe In tbe SKcnd p)ac« local vaitotnoior chauKea in the cundition of the minuta
V sileriea. changa, i. «.,of any particular voneular urea, have very decided
fflccts on the carviilation. Th«Ae changes, thouj^h lut-ul theuutelves, may
bave HiecU which arc both local and ^neral. as the fuUowiug consideratiotu
will elmw :
Let IIS suppwr that tbe artery .4 i« in a condition of normal tone, is mid-
way between psirvme conrtrielion and dilation. The tlow thr>Hii(b A is
dm I mil lid by ilte rtvistanci' in A, and in the voseulur tract which it luji-
plios. In relation to the miain iirienal preuiire, whicbaznin U dependent on
tbe way in whit.-h the hi-arl in bentimc and on the poriphenil r<t*i»tanoe of
■11 tbe small arteriet and capillarii'v, A included. If, while the heart and
tbe rwt of tbe arteries remain unchanged, .1 bo const rtcted, tbe peripheral
The ^«U of t'oMiiHoUtr Attioiu.
276
THB VASOULAR UBCHA<t!«».
Tn>!»tunr<' in .l.will nirri-ii:>r, mxl tbiti inrraiJN- 'if rtnttaDM Will Imd to kU
iiicnni'c of lln- gcm-nil rirlcriiil )>r(-.'<i'urc. Sitiw. nt w« h«T« seen, § 119, U
!r iirkTiiil |>ri.'N>iin- wliii;)i i% ilic imnicilliiti? cmwc of ihe (low from Xhv unprk-e
to llic vrin*, thi» incrcnsc nf nricrial [irwmrc will lend to drive piore lUood
from llic nrliTk-» inin th<f vtiiw, The c<>tistricUni] of A, hotrevcr, bv in-
crcaciug l.hc r»iiiliint-<\ oji|)')Ki' nny increase of the flow through A iUvlf, in
ftct will mitkc the tiov> through .4 lew than bcibre. The whole increase of
difchnrgc from Ihr nrlcriul into tbt> venous FViilctii will take place through
tbc iirtcrics in which the rreistance rcmaioe udl' hanged, tbut is, througll
channels other thun A. Thus, ns lh« resiill of the constriction of any artery
there occur (1) diminished flow through the arleiy itself, {1i increased gen-
eral arterial preeeure, leading to (^) increased flow'tbri>ugh the other arteries.
If, oo the other baud, A be dilated, while the heart and other nneries remain
unchanged, the peripheral resistance in A is diniint^bed. Thin leads to «
lowering of the general arterial preasiire. which in luro lends to drive leaa
blood from the arteries into the veins. The ditaiion of .1, however, by
diniiniBhiug the resiaiaucu, jjerinils. even «iili the lowt-red prei^ure. mors
blood to ])ai« through A itself than before. Hence the dlniini^biMJ flow lelU
ail the mure on the reiit of the flrteritrs in whirli iht- ret<i«tance rcniiiim
unchanged. Thus, aa the ivkuU of the dilation uf any nrtvry. tliorc mu-ur
(1) inert-used flow uf blood through thi' iirliTv it>elf, (li) diniiui»hi-d gciiemi
prwiiure, and (3) diiuiniiihed IIoir through Ihr ni'ier urliritw. Whi'ro the
artery ihun cunitiricted or dilated is --mall, iho Wal cllecl, tin- diminution or
incnMe of tluw through itself, is much nu>rit jmirltrd tlmn the general cffi.t'lf,
the diangu in blond -premurc and the tlow through other nrtcrici<. Wbcn,
howdver, the area, ihi; arteries of n'bich arc alfcctcd, i* large, the genera]
eflcclit arc very striking. Thus if wbilr n tniniie •4' ibc blood- prra* lire it
being lukin by nienns of u roiinomclvr connected with the carotid artery,
ifafl abdominnl splanchnic nerves be divided, n conspicuous but steady tall
of prenure is obeerved, very similar to but more marked than that which is
Men in Fig. ^if. The section of the ubdoniinul splanchnic nerves causea
the mesenteric and other abdominal tirteries to dilate, and these being very
nuuerouH, a larse amount of peripheral resistance Is taken away, and lli«
blood preeeure JallK accordingly; a large increase of flow into the portal
veiiis takt-a jilace, and the supply of blood lo the face, arms, luid legs ia pro*
portioniiUy diminiBhed. It will be observed that the dilation of the artrriea
Is not insianisneouH but somewhal gradual, as shown by the pretaiire sinking
not abniplly hut viith a gentle curve.
The general etlecls <iu nlmid-pressuie by vawimotor rhaDge» are ki marked
that the manometer may be used to detect vasomotor action*. Thus, if the
stimulation of a narticular nerve or any uther o|K-niiion leads to a marked
rite of the mean iilood-preft^ure, uniicrompiinii-d by any changes in the he«n*
beat, we may infer that constriction bat> taken place in the arteriec of tKxnc
Conriderable vain'ular nrt-ii; and similarly, if the eHi-ct be a fall of blood-
prewure, we may infi-r ihiit constriction him gtve:i nay to dilution.
Vatomolor funtiieit of the Ventral Jicniwu Sutton,
^ ITS. The central nervous Kysteni, to which we have i.niced the vasomotor
OCTves, makes use of theeo nerves to regulate the flow of blood through tbe
Tarii>us organs and parts of the body ; by ibe loeal elfecta thus produced il
artiitB or otherwise influences the functional aetiviiv of this or that tisane ; by
tbenueral eflfeota it secures the well-being of the body.
Tm use of the vaeo-dilatur nerves, which is wore flm{ile tliou thai of the
VASOMOTOH ACTIOS*.
nwMUlriclofs «nw it ii|>pi-iLr» not in )i« c»mplt<-ntc<l I>r (lie |>r»jeiice of
htUtiwI Utaw infliMiK-cs. in friMiiiitilly (.-unfliiii-umis bh pAit of n reilex act.
HiB^whi'ii ISkh) is plneeH in lliif miHilti. atf«rciit itiiiKiliM?^, (*euorat«') in thai
■Kn* of taste, give rite iii tW cniitrnl nervous system ta eflerent iinpiiUM, j
■tich dM<wn<l tne chorAn iytD|Miii and other nerves tu the salivary i^landfl^ij
ml, hr diluliii;; tlie tilnoctveescls, secure & copious How of blood ihrnuKh the '
|Iiadt, nhile, na vre sli*)! see Uler on. they excite ibeni to iH>r'rele. Tlw
mlnof thii reflex actioo njipears to lie in tho iiieiliilU oblootcata, and tiiiijr
b throVD into acliviiy not only by irupulites reaching it aliin^ the s|)ec-i6c
w&imot Uste, but nbo by imputsoi puasiiit* iiliinK other chumeU: thiu,
■DMkni ftartad tn t)>e hruio by the Bitrht of Ibod or otherwUe may cive riw!
k iwfiiliM pttninit dowD kIohk the oentral aurvou* Byfltcm ilMlf t^ thai
■daAi oblangHta. or eveata in the atomach nuy send iranutnes u|> the vagiNiJ
Mr*«,orUiniulBiion of Olio kind or another may Miid impti1^4±i itp nlmoftl
UTMBtiwit nerve, nnd the^^ various iiupuUoii n^ai-hin^ the mi-ilullii iaiiy,hfl
MnMtkwi, thmvrinto aclivitv thvrttiiii-dilaior filinaorilicvhunla tyinpunlj
nd (titer Kualngou* nervesi, »nil bring ubnui a l1iiT>hin);'if thrr aalivary gluudi,
«hilt It tho (Uiie lime ihry ntUM- ibu ^Innds In KWreiv.
Ibe irMu^ilator fibr<» i»i' the nervi crigenirti may !>« throwii into aclUlt^
hianitsr reflex way. the c^nlrv in this cn*i' l>cing ptac^l in thu himbiir or
Inrwilorvl txirtion i>fth).< spinal wrd. though it is wwily thrown into luitivitj
h iiiipuliMw d«Kending donrn the spinal i;i>ril from the brain; that such •
nwie doce cxit>i. in shown by th<^ fact that wh«i in a dog the spinal cord it
ONDfiliiely divided in ihe doreid rvjjioii, erection of tho penis may remlily bft
t(M^ about hy Blimiilntion uf the sentient surftces. And otlior instancw
■ijglit be ijuoted in which rnsi>^ilaiur fibres appear to be connected with a
'tntn " atMin after their entrance into the nervous ayBiem.
If. a* Mcm« pnilKible {$ in' I. the bloodveMftls of a tnuacle dilate hy vaan-
maUtr action whenever the muscle U thrown into conlraeiton, either in a
ftSex or voluntary movcniful, thr va»>-dilatur Bbret of the miiitcle would
(MB to bo thrown into action by imjiiilM^ arising in the Npiiiiil cord not Air
ftom the origin of the 'inlinary motor impnlxtt, and accompiuiying thuM
Botiir IfnpalM along Uie motor avrre.
i ITS. TheotMoftbe vnAo-conMrietorlibreallMfnewliaiinoreoninpliiMtm),
(D account of tile exiKtrncc of tonic influcmcea; sincis the sam«! tilirra may, on
iWitnn hand, by an incrtwMT in the imnulM* passing along them, bo tlie
■wan* of ronftri<.-t>oD, and, on tlic other hand, by tho n'moval or diminution
of the tonir inllucncw passing along them, be tJw incunK of dilation. We
ka** alrvady trau.-<) all the vaso-canstrictor lihrM from the middle region of
the spinal cord to the splanchnic system in the thorax and abdomen, from
■beacr tbey {was (i) by the abdominal eplnnchuic and bv ibe hypogastrie
aWTM to the vsMCra of the abdomen and pulvis (oonceniing the vaaumotor
■■rvea of Ibe thoracic vi»oera mc know at present very little): (2) by th«
wrvicnl sympnlhelic or cervical 'Splanchnic, a^ it might be called, to the skin
id* I he liead and neck, the •alivary ^lauda and mouth, the eyee and other
|Mna,and probably tbe brain, including ila membranes: \'-i) by the brachial
md •ctniic pk'xuaea to the skin of the fore- and hiud-limlw, and by vanoua
nther nervea t<> the skin of the trunk. Tlie chief paru of the body tuppllad
hr ira«o-couurictor librcn ap|>ear to be the «kiu. with its appendages, and the
aiiaKQlary canal, with its appendagai, glanduhir and other: tlw great maaa
of akeletal muanle* npjioara to raoelre an iniiignifieant kupjdy of Tiiao.eoo*
Mrietor llbrcf, if any at all.
If, DOW, In aji animal, the «]>inal ooni be divided in the lower dorwal region,
iba •kin of tJlO leg* bcoomM Huxhcd, their teniiMmtnn: rrc<]uently riw»t, and
tlMT* la a certain amount of fall in the general blood-pruMure aa inea«ured.
278
THE VAi!Cl'I.AU MBClIANtKU.
f»r instJiiico, in tlic rnrotid ; iiud Uii» slnt« uf things miiy lui for tomt oon-
tidcnible time. Ol>vii>ii»ly, tliu Melton of the »iiiniil «>rd lins cut ul)' lite
utiiul tonif intlucn<y« ilccpiiiliii^ to (In- InwiT limlM: in c»nMi|U(^tn> the
bliunlvowls liBV(! bccomi- ililnlcil. in eonwjuenct' ihp gcnerni periphcml
rreisluncc lin» bccpniv pro|i<>rtioii7tl«lT (limiiilehod. nn<) in con»ci|iieiivo lira
^ocriil hlodil-pirwiiri' hfi» fnllcn. The tonic v a fH' constrictor iiupultes lor
tlie tower limbs, lliorelbrc, linvc their origin in tl>e ccniral nervous sjsloni
hipher up llinn the lower tlorwil rofjion «f ihu i^pinnl eord.
tf ihe cplnol cord bedividti) between the rooiiof tiie tiflli and sixth dorsal ■
nerve; i iJiat is to tay. nt ihc level where the path of ibe Hplanchiiic fibre*]
from the cord eeenis to divide- — tee Fig. ill — iliose iK^iiini: above iMniog^
UDward to the fore-lioibe and liend, and those isauiiig below jiawtn^ to the
aodomen and lower limhe). the cutaneous bloodveaseU uf the lower limits
dilute, as iu the former case, and on exaniiiiali»n it will be found that the
bluodvesEeU of the abdonien are also largely dilated ; at Uie same tiine tba |
Miiiid -prewure uuderjfoee a very marked fall ; it may, indeed, be reduced ut
n very few niillinielres of mercury. Ohviounly, the Ionic viuK)-c<«wtriclor
■mpuWii passing to the abdotnen and to the lower liniliti lake origin iu the
itnlral nervous ayetetn higher up than the level of the lil\h donal nerve.
If thenectioDof the.iphiul curil be made above the level of the M-cund lUinwl
nerve, in uddilJon to ihe a Imre- mentioned rt-sull* the vcvscU of thi- head and
fact- alMi liccume dilatctl : but. in I'unsequence of the fall of general bloixl-
prcwurc just m<-nli<ini'd, these vt^sel^ never liecome mi full of hlnod, the hiKsof
tone H not to ol>viou» in ihem iu< after simple division of the ecrvical syniim-
thfflic, since the bitter opcTation [irt)duce7> little or no etH'ct on the general
blood -nrnvH re.
Ol>vinu»ly. then, the tonic va«ii-coii»triclor tmpiilees. which pDs*ing to the
skin and vioeera of llm body riiiiint:iin ihnt (onir niirrowing of so many small i
arleriea by which the general [Kiiplieral renislance. nnd so the general bloud-
preoHire, is mainlaiucd. ptoceeil from some part of the ec-ntml nervous
System higher up than :be upi>or dorsal region of the tipiiml conl. And,
since exactly the aaiue results follow upon section of the »pinul eord in ilie
ccr\-ical region right up to the lower limit of the mnlulla oblongaiu, we iuler
ihal the-ie ionic iuipul&es proceed from the medulla oblongaia.
On ihc olher hand, ne mav remove the whole of the brain right down In
the up|>er parla of the medulla, and yet imiduee no HuBliiug, or only a slight
Iranpicnt flij>liing, of any jinrl of the bu<]y. and uo lull at all, or only a xlight
tn<ni>i('nt fall, of ihe general blood-prei<snre. We, theivl'ore, Mem jiiMifii-d in
astuniing the exiiiience in llie mediillii oblongata of a ucrvouii eenlre. which
Wt may e[K-ak i>r»ii a I'lmointitor fmlrf, or the inrtlullur^ ruAnnwlof emlrr., from
which [lojcnil tonic vBKo-consirictor impu !»':>, or which regulates thcmiiaBioa.
and dittribulion of Kiicb tonic viixo-eoti strict or impuleca ur inlhiences otef'
various purls of the hiwly.
S 174. Theexislem-e of tliis vawmotor centre may, muri'over, beshowit in
another way. The extent ur amount of the tonic eonslrietor impulses pro-
ceeding from it may Im iucrcawd or diminished, l\w activity of the centre
may be augmented or inhibited, by impulses reaching it along various alfereni
aerves ; and provided no marked changes in the lieart-beat take place at the
Him« lime, a rii^e or fall of general hloiHl- pressure may betaktu as a token of
an inensnw or decrease of the activity of the centre.
In the rabbit there is found in the neck, lying side by side with llie cervical
symiiatliniic nerve and running for some distance iu conifiaDy with il, ft
slender iiervo which may lie ultimately traced dnwn tu the heart^ and which
if traced upward is found to come olT ^lomewhat high up from the vagus, by
two or more roots, one of w hich in geneiully u brunch of the «u|ter)or laryn<
TA!)0irOTO1t ACTtOXS.
^nkl aCT\t. Thti wrvv ithv Hl>re» ooiiMiluling which Air. iti thsdcig, bound
up mixh iIm> vn^iiD soil il» nut loriti nn indcficiiili-Tit iivrvcj npptUi to bo
rxrlunrrK on ulTeifnt Dcrvc; when, ufter division of ihi.- nerve tlm pfri[il«'riil
rml, ibr r'nd Alill in «>nnLvlioii wiih llie heart, i» HlimuUttx). nu iuKrk«J
nwuli* fi'lliiM Tho hnriiinius)! uf ibu uervc in the heart nre lli«rvfiin- (juilc
tUdaratt fniui ihi- ouilin)^ of lh« inhihilory librw of the vngue, i>r of ihi;
■anMntor Rhm of thu >>|>lniidinic (i>vm|>alhelici syBlom ; ihc nerv« hiM
OdthinK I" do iiilh (h« nvrvouii n-gululion uf the heart [tee pp. '240, rt. m^/.)
If. n.ni. while the prewure in »iii iirtery ituch lu the carulid ia being rvgislurwf,
iht nrulral eud of ihe ntri'e — i. *., tliv one eonnecteti with the brniit— bo
Biituulaied with the intcmiplnl eurrviil, a gratlual but nmrked fall of pr«»-
atm {Fig. 99} in the carotid is oliM-n,-cd, tttnling, wli«n (he period of «timu-
rro. ».
/'"^■^Lr^
^
TKMna Saovmn t«b Erncr on DimiIkphmmtu or SnmuTtMn Tin CRnK4i. dft or ma
DU-*B*nK NUtTK IH mi KlBOIT.
!)■ ttm Ume writer t»etow the Iniwvl* iiomiponA to letiinila. AI'ialnwnvploloDrKMwai
tfercnim iMlii Uio nelrc
latinn is sbort, tatae linte ai^er the rviuoval of the ftimulus. Since the beat
of the heart ii not markedly changed, the tiill of presMire niuat be due tn thv
•lininiition n( ji^riiihera) reMslauce occsuoDed by lite dilation of ■onw
ulcnr*. And it ia probnble that the arleriea lliiis dilated are cbiefiy, if not
tmludreljr, thn»e nrieriei of the abtloiuiiinl viscera which ar« jcoverued by
tha abdonilnal iplaiichnic i)i'r%'t? ; for if iheM iierves are divided on botb
lUt* prnt'iova t'l lite ex]>i'rimt'ni, the fall of nr«aBur« when the nerve la Miinii-
lalrd !■ very »mnll — in fad, almost insignificant. The inference we draw b
a> fidlnw*: TIr- alR-rvni impiilxra, pamns upward along the nerve in ijuesiion,
hare mi nlfu-<-li-d Nimu part i>f ihc centmT nerirou* xYSiem ihat the inducnces
•hirh. in a U'inmil comlilioii of thing*, pajsing along tW nbdominal
tplanehnif iiitvc* keep thi'^ minute* artt-ricM of thv abilniiiiiinl vtKX'ra in a
NaitT of Tnmh-ralv liuilc conolrirtinti. fail altogether, and tlii<w arlcrieo in con>
wqaeorv dilnt<' ju:>t an tlii-y do nhrn th« abdontiDiil Miilanrlmie nervM arc
ifivided, ihe elR'cl Iwiiig |H>r»itity iiivreaM>d by ibe tinnlar ililatlon «f ntlwr
TMTuIar Hrraa. Since vlinulalion of tlie ucrw of which we an- niiraking
il«ay* produce" n fall, never n rise "f bloo"l-prfMure— the amount of fall, ot
eoarre. l>cinF: di-|>cndent on circii to stances, luch a« the condition of the ncr-
rmn ayetcm. i^in of blood -premirc, etc. — the nerve if known by tlie naiiw
••f the dffr'j>M>r nerve. At we »linll point out later on, by means of thw
■Ami n«T\'«- from the heart the peripheral rHislnncc is, in the living body,
lowered to euit the weakened powers ><f a laboring heari.
Tbbgndnal towering of bluod-prasure by diminntion of peHpberal reabt-
S80
TBE TASCULAR ȣCUAKISM,
Moe nnunln n ninrkM contrast tn thv eii<ld«n l<tweriiiK uf bloinl presuira bv
cardiiii- inliiliilioti ; i-i.ni[niri- Fig, Hfi wiih Fig. fl.*.
S 17fi, But tliF gi.-iiernl til(io:)-)>rPmiiri? inny be moHilier] by afl«r«(il iriipulMfl
paaio" iiloiig lilhcr iicrvc* than the clepr*?aKir, the mixliHi'aiti'iu taking on,
aecdruiiic (o cin-iiMifiluri<;oe. the fttrcii eillier of deiTEUue i>r of iiicrviue.
Thus, if ill un iminial jiliii.'e<l iiuilvr the iiiniience of iiran (triine nitiNlIietic
othor thnn L-hlural, eu'., beiug uwd) ihe ceiitrul stumj) of llie dlvideil M-iatic
iKrve be eiiiuulaled, an iiicreaite uf hliHid-|in«aure (Fi|(. 100) aliuoat (.•xuotlj^
Klu. 1(0
JW.VywWWW*WV'
,;,^r^ww^^
^ww^^v.----^-^^^
X
EmcT 0.1 BUMihritiMnniK Crnvs ov Bnm*i.ATiini Smi-nr Nuitk i'nuru I'niiii- (c*t.)
(Durlu ibc moDiL-nl 111 which Itiv iiimtil wvlbmwn liiUilUt iiurvc. AninrinI n»(iliBllaa «ra*
carried un.anil HiBiimal napmitniT tiiiduUIIixivacwsbHiil
ihe reverse uf the decrease braui;)it. ulxuii by »tiniiitnting the d«pr«mor, \n
oliaerved. The curve of the bloo(l-pn?siiire. iil\er n lnt«iil period diiriDK
uhit-h HO (-bim);i-'» are viaible, rises Eteadily without any corratpuitdiug
L'hauf^ in the heiirt's heat, reaobes a maximum and afler a while slowly falb
ugaiii, the full winK-liini-'n beginning to apfiear her<ir« the stimulus haa been
rviDoveil. There can hv do doubt tlinl the rise of pressure is duo In the
coii»tri<rtic>ii uf certain Hrteries; lh« arleriea in (jueslion beiii^' Ihuse of ihe
■bdoininal «pIiuichQic aivn ccrlninly, anil ixmibly thiiW! nf other vascular
■raac lu well. The cffi^^t i* nol coiilincd to ih« *ciatii!: niiiuiilation of anv
nerve oontminiue afferent fibreit may ]>roduvf th<> nunc rJSL* of prcMure, and
so conilanl is the result that the expi^rinicnl Iiiik Ih'cii mitdc use of a* ft
method for determining the existence of ulH-n-iit tihrc9> in nny given nerve
iind even the pntbe of eeutripct.il imi>u1s<u through the spinal ennt.
[f. OD the other band, the tinifnni be under thu inHuence not of ururi Itut
of a large dose of chloral. in«ieiid of n rise of blood -pressure a fall. <{uite
similar to that caused by «tiniij!iiliiig the depressor, i« observed utien ao.
alTerent nerve iMbtimulatetl. The condition of the ceutial nervous system'
seems to determine nheliier the alfect of afferent impulses ou the centrnE
Dervuus system is one leading tn an aiinmeDiation of vaso-coniitricbar impulne*
awl so to a riwe, or one leniliug to a diminution of raso' constrictor impuliUM
and so to a full of UioiHl-prauure.
5 176. Wi- have uwd Uie words " wntral nervous eysiem " in spoakiitg ofi
the above ; we have evidence, however, that the part of the ventral ncrvout .
ayaieni actiil on by llie nflt-reut impulses is the vitaoniotor i-eiiire En U19
medulla nbhmjfnla, and that the eflivlA In the war of diminution (deprawor)
or nf augmMitatitm (prewior) are thf reaulisorttfiereut inipuUw iubihitiugnr
tuzinenting the tonie ar-tivitv of thisi et!Ulre or of a part of thin centre spe-
cially oonni'Ctcd with nbilomnuil splimchnic- nervea. Th« whcili> brain mnj^
bo rrmovnl right down to the medulla nblonitata. and y<it the elTucts of stimu-
lation in Ihe direction either of <liminMtiiin or of aui(meniaiiuu may still b«'
brought about, [f the medulla oblongata bo removnl, iIkvt effects vanish
tou, UMMi^h all (be rest of the nc^rvoiiH svstem be letl intact. Nay, nn.irei. br
|iWtiaJly inicrferini; with thi- iiio<lulln o)tlongiitii, w<i may partially diminish
tJMW enbcta and thus mark out. «•> to sjh-uIc. the limits uf the tvntre in ques-
i
VABOMOTOlt ACTIONS.
•281
tlno wllliin lbt> tiHoliilla iUi'ir. Thus, in an tntiici lUiiRinl iimli^r tirnri.stimu-
hiioH of ihp KJiitit- m.-rvo with n flimtilii? "fit i-i-rtniii ulrvtigili nill iiivliim
m rW of bliuid-|ircsHin: iip tii r rtrtnin ^siciil. Afitr rcnionil nf tlir wlt«l«
hrain right down to the medulla ubloDunta, tho snnic »tiiiiiilnrion will firo-
■lutT lh« nm* Hm u before; the vasurmitor centre has Dot bran initirfcrod
witb. Dliwetljr. bowevor. in proi-oedine dovrii<van). th« n^ion of lh« cwntn
fo qutMlon u FMched. itimiiliilirin (t( tho ecialii' ;ir<)diic«8 \tea and Uv» rise,
untt] at Ian wh«M lli« lower limit of the centn? ie arrives) at no cDVcl ut nil
oa blood-prcMu ri' iiin be pTodiiced by e7«ii atrong BtiiuulaliLiri of the m'iiittc
or othvr atTereDt nerve. In ibia way the lover limit i>f the medullary vnw-
BM>i"r centra liaa been determined in ibe rabbit at a liorizonlat line drawn
■bout 4 or 3 ntm. above tite point of the calamus seriplorious. and Ihe upper
limit at about 4 mm. htxher up — ■'. e., about I or *2 mm. below the corpora
JaadriMinina. When tranavene «ectioiid of (he brain are carried Eucoea-
irelr lower and lower down, an efRwt on blood-prewurein the way of lowering
it mmI abo of dimiimbin^ the riai! of blood-preeaure n«ulliii|{ from Alimula-
lioa of Ibe Miauc. i* &nt otiscrveil wlii^ii the u)iF>^r limit in readied. On
tmrtyitig tbe aectionH ittill lower, the- effect of utimu luting lb<j! idatic be-
CDOW Im> and Um. until when the lower limit ii readied do edectd are at
■II obwTTvd. The mntre a[i|H-an lo tie hiliitLTal, the halve* bcin^ placeil not
in th* middle liite but morv nidcwayn nnd rather ni-an>r tin- anterior than tike
poMeiior ■urfao''. U mav [lerhnp* lif- mure eliwly dciiinfl iw a xmnll pria-
matir ii[uuv In the forward pmlongniion of the Inlrml coliiniuK atler they
haw civi-n »|1' ihvir fibre* to the dccumttiii^ iivriimt>l.«. Tlii* Hpiice in largicly
otntiMd bv ■ mnn of gray matter, rnired by <^'inrke ihe nniem-hiternl
WmJmi. ami oontaiain^ lareo mtit(tt"ilur cell* : but il in by no iiicaiii certain
tkai tbia Kmup of nerve cell* rc«lly net* nii the centre in ipics^ion.
)[ 1T7. Tbe above es|kerimeiits appear to afliird udcmiate evidence that, in
amrmal ntaie of ibe bodv, the iute»rily of the medullary vasomotor centre
iiiweiiliil to tbe imMludlon and difttribulion of thiwe continued oinatriclor
(■polMa hy which the ^oeral anerial tone of the body is inainlaJDcd, and
tlal aa iunraue or d&t-reutfc of vom- const rictor action in particular urleriee,
orlnihe arteriea (cenerally, in hnm^bt about by means of the name m&lullary
nHuiMor centre. But we rauM not, therefore, conclude that thi.t xnuill |Mr-
iniof Ihe metlulla nbliiDgala u the only iiart of the central nervoun ^yitem
*Ucfc can act aa a i-euire for vn.-w> conittrictor tlbnat ; and, aa we have Men,
tkm II no erideiioc at pr(»enl that the va^i^dilator fibre* arc connecteid
eilfc eitlier ihin or any oth<-r one centre- In the frog reflex vnaoQiOloredecta
■ir be olitain>-il by xlimulnting variouii atlfir(rnl nerve* alter the whole
■Mlolla hiiii lH-i>n reiniiveil, iind, iiideiil. even when (inl^ a ivim[ianitively
Miall portion of Ihe uniiml cor<l hiu been ]i:t\ intact and connocleil, ou the
Mr band, with tite nnerent nerve which i» being (timuliiuil. and, on ifae
(thar, with the efltreni nerveit in which run (he viL'«>m'>inr tiliren whom action
a beinir »tuHie><l. In the mammal such clil<-ct>i di> not «i readily appear, hnt
My with care nod under s|>ecial conditions be obtained- Thiu in the dog.
*^n (he »pinal cord is divided in the doraal region, the arteries of the bind
&■!« and binder part of the body, as we hare already eaid, $ 172. beoonte
dilated, llib one would naturally eipect as the reanltof their severance
frnn tbe medullary vaaomolor centre. Uul if the animal be kept in Rood
nadhion for aome time, a nornitil or nearly normal arterial (one is after a
vhiU recvtablbbed : and the tone thus re^nined may. by atTerent iiapubeo
narhing lite c«rd l>el<>w the aection. be moililiexl in the direction certainly
•f itiniinolioii — i e., dilation, and noasibly. but ibis is by no meanit at> certain.
of IneraaaH — i. r., const riciinu ; dilation of various cutaueou vcttieU of the
282
TUK VA8CCI.AR HECOAXISM.
limbs may be readily [irMluced by atimululioii of the oeutnil Mtimp of one it
aootber nerve.
'Th^ia remarkable resuliji, nliich tli'm){!i (hey are im*t stHkioK i" euattec-
lioD with ilic lawvT |ian of ilic »|iiiiii1 cunl. hold ^Mjd a|)pHi'eiilly for other
perlii also urilu; ^|iiiial ri>ril, niiltirutly »ii]|f);i^l a ilinibt. iv'iirlbfr the ex[ilumi-
ti>>ii juxt gtvi-ii aliiivo iif llif rtTi.Tlii !•( scsrliiin of the inHlntla nhtoiigata b a
vflliil Olio. Wbi-.ii w(,' citmt^ IQ «l<iciy ilii;! centnd uitvouh nystrm, we nhall
again aod iipiin ^t-p ihnl thp tiiiiiicilintf uHect of opcrativu iiiUrrfercDW wilb
lh«o dolic'jilc i-lnicturoi in ii Ivrnjiuniry iiiii>|>cnKion of nvnrly nil iWir fun»
tions. Thii^ is oflcn npokcii of ns "«ho<-k" niid mnv be r<^irdi.il »■ ao
cximnii; ibrm of inhibilioti. An rxnmiitc of it. u<>4:iin> in the nbow «x|>eri-
mcni of section of the dui-Kil curd. For eonie timo nlt«r Uie operation the
Tnso-dil&tor nerri erigcnipa ( which, as far a« wc ktion-, havp no speciBl ooamo-
tion nith the medullary vaeuiiioior centre) cannol be thrown into m-liritris
part of a reflex action ; their centre rcmnins for trnne lime inncliTi?. Aftor
It while, however, it recovers, and erection of llic pent! through tlw nervi
erigentee may then §lill l>e brought abont by suitable stimulation of ernsory
surfaces. llen<« ihe (|uealion may fairly be put whether the etiects of caV-
tini; and injuring ihe structures which we have spoken of ad the medullary
vasomotor centre, at* not in reality aimply those of ahoek. whether the va»-
cular dilation which fullows upon pactions of the so-ealleil medullary vaso-
motor eenire, iti.eM not come about because section of or injury lo this region
exerciser a Uron^ inhibitory inlluencenn all ihe va.ii<ino(or centra attuAted
in [be Apiual rsird below. Owing tn the :^pmal function of the me'lulla
oblongata in carryluj; >m the all )nipi>rtant work <>( migration, a roatnmal
whnse medulla ban been divideil ninnoi be kept alive for any letigrb of time.
We cannot, therefore, put the matter lo the Minple ('X|>eri mental lent of vslir-
patiug the eupportd medullary vniwmoUir i^ientiv and nx-in^; wlint hiip^-iis
wbi-n tim animal has completely recovered from iho etfrcts of the D|wnilion ;
KC have to be guided in our decision by more or kt* indirect nrgnnient«.
AihI against thti argument that the effects are those of shock, we nwy put
the nrgunicnt. evidence for which ive 'hall meet with in denting with the
CCDlral nervous system, that when one part of the central nervous system ia
ivinoved or in any way placed hort dr cr/mbul, another juirl imiy ricarionsly
lake nn its function ; in the absence of the medullary vasomotor centre, its
fimction may be pcrfirnied by other parts of the spinal ooni which in iu
presence do no sucli work.
And we may. in coiuieetion wilb this, call attention to the fact that tlie
dilation or loss of tone which follows upon section of the cervical avnipatheile
(and the same is true of the abdominal sptanclinic) is not alw«yt>, though it
may be sometinies, perinuuent : in a certain number of caaea it ha.'i I>r«-n
found lliat al\er a while, it mar not he until after neveral days, the dilation
disappears and (he atteriiv regain their uMial calibre; on the other liami, in
sooK- cases no such retani has been obwrved aflrr monllm or e\'eu yuam.
This rei-overy. when it ix'curs, cuniiot nlunyji be attributed to any regenera-
tion of vasomotor IJIire* in thn nvmputlK-tic, for it i* slated to'bavv bm-ii
observed when the whole length i>I the nerve including the Mijienor cervinil
sanglion had been removcit. When rccorery of tone ha" thus taken place,
dilation or inereuM'd couMriction may lie occiisioncil by local ircatnient; the
ear may be ninik- to blush or palo by tlic application of heat or cold, by
gentle stroking or ntu&h handling and the Mice ; but neither the one nor the
Other condition citn be brought about by the intervention of tltc central
oervoUM system. S.> aluy the spontaneous rhythmic variations in the ndibre
of the ailerie^ of the ear of which we spoke, though tbey cciise for a time
afler diviaionof the cervical syoipathctic, may in some cose* eventually reaji-
VAfiOUOTOR ACTlOXit.
288
|<ra r «Btl tliat evt u if (lie superior eorvicnt eo'ifllion ^« retoovin] ; in otlwr
raw* Ibrj- ilo uoi. Aixl t]ie aiimlogous rtirinnnc varialions of ihc vdn» uf
ibr l«t'» winii liave htta proveil es|wHmeiilally (o gu on vigoroiiBly nben
all rMUMflivD willi the <«ulnil nervviia ivstcm lius l>eeu severed; thi'V mnjr
«(MliBa«. in f*a, iu iwiUte^l |>i«ces of the u ing |)n)videi) that rhe vimcIk nre
«iIr<)(MitrIr filleil aiti] ilUtciiJt-il with UIcimI or iluiil. From llicse nml I'lher
(bi1», rvcu afttr iiiakiux allimumx' for thi? mifative casee, we may coiifliiilu
that what we have (poken of as llif (one of the vei*«lB of ihe face, though
influeticwil hv am) in a ni«aMire ilcjHMidoHt uii (he t^entnil iiervoiii syetciii. i»
ant aimplj l^e r««ult of no c-irirl uf that KyMi-in. Hie inusciilar unlla of
Uh* art«ric> art uot mtn ]wiviv<- iiirtruinriiu wnrkcil hv the (vulral ncrvoiiB
ajMcui through the vn^oiiiolrr fihri-3<i liivy app-ar l<i finve an ininueic lone
oTtbclr Qwu, BDil it ocrmit natural to (U[i|>i^w llial nhrii the t'etitral nervous
■fMcm caom ililation or con^lririion of the vi-n>cl» of thv facr, il rijakea use,
ia wi ili>i»K, nf thu inlrianic h>ral tone. It ba* h?«ti »iip{i(iii(^d that this
iatniisic li>t>« ia <k'|>eii<li-iit on vomer htcal iivrvou« iiiechiini»iii ; in (lie ear at
kaai no luch mocDaiiiani has yet been found ; and, indeed, as wc bavc said
aboT*, I 167, DO nidi |>«'nphenil nervoiift niechani«ni i» really neceMary. In
Um cbm botli nf a vMael govcroeil hy vaso-dilnlor fibrea ami one giiremed
bj TBM^oMtricIor fibtti, we may suppose a ccrlain natural enmliitou uf llie
maactibr Kltrc« wbieh w« mar i-all a condiiion of equitibrium. In a vetsel
(ovencil only by vasodilatbr libres, if there \w Hich. thi« onditiun of
c^DJbrium ia ibo {wrmancnt condilion uf the niuseular libre. from uliieh it
bifiMurbwl bjr va»«i-dilalor impulse*, but to nhicb it i<|ieedily rclurns. [n
tnael govenH-d by vaao-mnAtriclur fibres, and subject u> tone, lh« niuncular
Bbf« a habituallv iv\>t on the eciiHtrJcior t'iile uf this eijuilibriuin, ami, an iu
1 iW raar* iiuoteii ahuve. may strive uf itself IuwhkI (ame amount of active
L (Mitrkltuu even when xparati-d fmm the ci-nlrttl nervous syatem.
H Dm Io retura tu ibe rui-dullury vaariniotor wntre. Wiliiout atteniplin);
V tDdiKUas the loatler fullv. wu mnv wy L)iat, afler all due weight haf been
aUsrliei) to the play «f in^iibitory inipulnw nr " aluK-k " a« a result of opern-
i"* iDterfervnctr, tlierc alill n'niain* n balane« uf cviiUiu-e in favor of ibe
nnr that tbc tvtnun of tbc medulla uf whioh ne arv K|Kuiking does really
■rtM a f^nvml vaaomiitur o-nlrr in the manner prevtouiilv explained, awl
{■knan imrairtaiit pari in the vnoimulnr r^'ulalion of iht) living Ixidy.
Ili* Dot, Wvitvcr. to be r<')(arded a> the MUfjU* vawmiolnr crnire, uhenoe
•hoe can i'Piue (onicoiiutri'-tor tm|>iiU(-« or whither aHcrt^>t inipiiUes from
*ll parts of the laxly mu»l alwny* irwvt-l hcforo ihey am atlwi the vaso-
•** UupulM-o |>«oi>iRg along thL» or that nen-c. We are rather to wuppoee
>W lfa» fpinal iHinl alung il« vrhojc length roulains, interlaced with ihe
itiaaiMl lAht-r mechnnivRH by nhicb the skeletal iuii<'(.-les arv Kuvemed,
nmmotor eeniren ami mecbaaisma of varied eomplexity, the <lot«ils of
■biat funetioiw and lo{>o|^pliy have yci larj^ely to he worked uul; and
■kougli. aa mr have m«u, the medullary {vnlre is essentially a centre of iui-
pdtta bmioi; nhmg vnso'«oi)sl rietor librea, it ia [Ktssible that there arv tiea
BMarrn ii and va^i-ilitatiir Kbm also. Aa in tl>e itbsenoe of the «inus
■Wmus, the auridcs and ventricle of the frofc's heart may still conliniie to
•MiMt in the altsenL'e of (he nie<Iulla obloutcata thcM spinal vasomotor
noim provide for the vascular emerjieiuiea which ari»e. As, however, ill
tW onrmal entire fru^'u heart, tlie sinus, m to sjieak, ^ivea tbc aord aiwl
l^nms the work of the whole orjcua, tti the meilullari- vasomotor rent re
rilcB ai>d corirdinaUa the I«mmt centres of the eord, and ihronich them i>tv
adt* over the chief vaacnhir ureas of tJie body. By nieanii of these vbm-
■Mor c«^iiral meehnnisnin. by meau* of the head centre in the mctlulla, iind
Ik auUidiar}- ceuimin the spinal conl, tlw delicate machinery of the cimi-
284
TlIK vasculab UECITANISH.
lation nhich dotcrmiues tlic blood-etipiily, and tm the activity of eHcli tiMiie
auil or^an, in ubie lo nxpoiid by nnrritwiii;; or wiilenio;; iirlcne« i<.> th» erer-
vttryiog iJenutDils e.nd to meet by vonipeontiiiK cliaof^ tite sliocltB siid
Mraina of daily life.
^176. W« niuy sum lift tbe hift»>ry of va»om»lor Holioiu turaewbal as
follows;
All, i>r nearly ull, or aa far as we know, all the nrteriea of (he bnily are
coniieiTlMJ witK t.bc coatrnl nervous sv^tiem by tiervc Abrai, called vuo-
motor Cihrti, Uie iictioa nf wliich vurieti tlie nmimnt of coutradluii of (h«
niuiteuliir coiit^ of thti srhiric^. iiri'l no leml* t>] chuugM iu calihrv. The
acli'iu of these ragomotor tihres is more maQit<M aad prolmhiy inotw
impurlant iu the oue of smaU imd miiiuie artcrin than In iho com o(
Inrgnr ones.
Thwo vasomotor fibres an- nf tvro kiiirln: \hr one kind, vago-vonftrictor
filirif^. arc of such n niiliir? or hnviMHich connectiiinH nt thdr central origin
or pcriplirml ending tbiit sliiiiiilation of (lu-m produces nnrrowin); con-
etnctinii of rhv nrt<Ti<^ ; and •Itirioj; lilb llipec Hbrcs Bpp«Kr to bo the ■□««»«
by which the cpninil ncrvoii* »y«icm pxcriB a cunlinued tonic inHiipnec on
thr nrtcri<« iinil mninUiinK an arC^irial "tone." The other kind, ihc vato-
dilator librn. arc of HUch a kind or have such conneclions that Mimitlalion
of tltcni priidiica' widoniiic. dilntion of the arteries. Tliere is no adet)uate
evidsDOe that these vaso-dilalor fibres serves as chaonelfi for tonic dilating
tDOpulMe or influences.
The va£o-coii!ilriclor Hbres leave tbe spinal c«>rd by the anterior rooU of
the nerves eomint; from middle re^ioua of ihe spinal eord only (in ihe doj(.
and probably in other mammals, thim about the aet.<ond dorul to the aecooa
lumbar nerve). pasH into the ^plauchnit- ;^uf;lia connected with tliooe nervea
(tlmracic and ahdoniiual chain of ayuijiatlietic ganglia), whera the libres loee
tfai^ir medulla, and procec<l lo llieir deKtimilion im non-mi'diilbilcil fibre*,
cither Htill in mi-cullcil nympulhi-tii; nerve*, such iw ■plaochnic. cervical •ym-
patlietic, liy|>ogaiiCric, Ho., or along recnrrrnt lininclitv of the splanchnic
SjBtciD, to join tlir »pimil niTvcn of the arm. W. nnd Innik.
In the inta«a orvnnii'm Ihr emiwion and diMtriliiition along thi.-«e vaM-
(wnslrictor l>brw of toiiioconrtriirtor impulses, by which general and loca]
arterial tone is inninUiincd and rcgnlalrd, is governed by a limited ponion
of the mcdidia <ibl<itigal» known as the mcidnlliiry vasuniolor centre ; and
when «)nie i-hnti);e of con<lilionE or olh«r natural stimulus brings about a
change in the activity nf iIki vaso'cansirictor libres of one or more vaacular
areaa, or of all the arteries siip|ilied with vaso-cuiistriclor fibres, tltis same
nwilullary vasomotor t:enire uiij^ars In such eas«t to play the part of a
centre of reflex action. NeverlneleM, in caaea where tbe nervous ooniMelionk
nf ihiH medullary vasomotor centre with a vascular area are cut off br an
operation, as by action nf the <.-()rd, other parts of the spinal eonl may act
as centres for the vaao-ronalrictor tibruti of Uie area, and puMibly tbvac sub-
ordinate centre* may be lo a certain t-xteiil in action in the intact organism.
Tbe vaiffdijator fibres apfiear to taki? origin in vnriou.i puria of the central
iKrvtMis iiyNtem nnd to proceeil in a direct niunte (« their destination along
the (anterior) roolnand ai juirt of the trunk* and brancbitfior varinuM cprebro-
•pinul nerves; tliey do nut hm- their medulla until tlicy nppniach thoir
terniinalion. Thry do not nppcnr lo sorro as channel* of tonic dilating
toflu«nc«s; they are thrown into nction generally as part of a rvflex action.
Mid their oonire in the rdh-ic ncl appciir* in «ach case to He in th<- e«nlral
nerroiu eystem not fur fimm the centre of tbe ordinary motor fibres which
titer accomi>any.
'f he ellectd of the activity of the vaaMlihOor tibm apjwar to be essentially
VASOMOTOR ACTIONS.
285
I
i
lormt iu nature; wbrn any set of ilivni coroe iiilo actiou llie ruscular iirva
«kirh tbrw lEOTtiii i« dilnred. Ami the vavniliir «Te«i w> governeil are
rrUiivvly to Btnill ilmi rlinn^vs in tli«tii jirinlure liiile or no effect on the
TitcwUr natMti io general.
'llivrfr«clii iif rbuDM in ili« Kdiritynf ihevii»i>rAti*trict»r libira arc botb
toctl and fti-timil. iidiI iiiiit tto nlM> doulile in naliiti'. By mi inhibilioii of
ttDic-t^iinlrirtiT im|iulH-ii i> txTCHiii iiouiimt of itiliilim) loxy \ir cflwtrd : by
n tOfEiM-DtiiUon of coiwiririor ini|iiili»«. CHiiftrii-lton, it ninr Ik- of miwiikr-
■bircxtmi. may Iw bmu^lit about. Wlmti the viuK-ulitr arL-n w> iiHrclr^ it
oiiiU, Uia eflccia arc local, mom or Ick< blnotl i> itirlrihulrd llin>iigb ib^- una ;
•btatlM raMmUrarcn nHcdnl !« large, thv inbitiitiim of conHrictioii nmy
bad la a marknl fnll, mid an augiDentalion of consirictiun tn a marked tim
tt rmtral blood-urtnure.
{1T9, Wc ahall liavs occatinn lalcr on ngiiiii Uld oniin to point out
bWucn of lliB tStvtf of vaMmutor action, both local an<i ecnemi. but wc
•a; hen tvavXt vov nr t«o rbamdcrimic nnn. " Itliiabing ' i« on«. Ncr-
1M« iaipu[*n starird in khic {lan* of ihc brail) l>v an cinolion [irodiii-o a
(■arrftil inhibiliikn of that )iart of ibc niE'dullary vasumotor centre nhich
funtni* thv viiKiilnr areas of the head supplied bribe cervical sympalbr-iic,
•arf btncT hji» on elTcct on ibe vaeoniotor tibrea of tbe cervicnl sym pathetic
■InatMlOliy tbe rn»>v n» ibat produceii liv MctiuD of ibe nerve. In mnse-
^omnilM niiiiH!ul»r wall» of tbe arleriea of tbe heud and face relax, the
ttirrir* dilalr, and the Khiile region beconiea iulluBed. Kuiuetiniea an eino-
u»ii ^ivra rise not In blu»biug. but to ihe oppotite elfecl, vix., to juillor.
In a Kn-ai number of ea»«t tbift biin <pitte it i^tTereni CAiiae, being due to a
Mililea diminnlion or even tein|H>rarv urrenl uf the heart'* beam: bnt In
MM oMo it Riaj (K-ciir without any elinngc in ihe bent of ibe liearl, and ia
iImi due to a raiulilion the very convcme of that of blmhiitg, ilint i*. lo an
iMmwd arlrrial miiatrietion ; and this incmscd coofiridion. like tbe dila-
■in nf bluahiog, i« eflretcd through the agency of tbe central iierTousaysieia
ut the eerviou ^mpathetic.
The vaanitar ci>udiiion of tlu «kin at larg« aflbnb another insianco.
Vh«D Ibe teniperaiure of tb« air is low the vraeels of tbe skin are eon-
•cHrifd and the »kin is pale; nhen ibe teinperulur« of tbe nir is bigh tbe
T^Mb of the ikin ure ddaied and ibe xkin is re<i and tlurbed. In both
Ikne raM-« tlie eHirci is mainly a reHex one, it being the central nervous
iTtfrm whieh brings about augiueniatiou of coDelrictiun in the one cttue and
(iUbillon in tbe other, tbotigh possibly some alight elfet'l is reprociuced by
ibt dirtet adton of the colli or heat on ibe voaels of the »kin dimply. More-
erer, lb« vucalar ehangm in the fkin are aceompvinied by corn?ii(K>ndi»g
nacnlar ehanges in the vnsoera < ehieHy abduminal) of u revorae kind. When
die rwtia of the skin are dilnTtnl tbuve uf ibt! viwecrv ar« con«lnc<ed. iind
iwe arrait, so that a eonsiderablt! jiorlion of the whole blund ebb* and (Iour,
» (o speak, aeronline to circuuitiliinci^ from akin l» viHoi-ra and fnim viscera
Mikln. By ibeae cnutgea, an we nhnll see later on. thi: nmiutonance of ibe
■anaud tenil>eralure of Hie Wly i« in large metmiirc fet-untl.
Wben food >a |>laeeil in the mouib ibe TdondvwMdN of tbe aalivary glaniK
■a we have tee». are fluibird itilh bluml an an adjuvnnt In llic fccretion of
4iKt*ttTe fluid: and ns Ibc IWl panm along the alimentary canal, each
■aeiian in turn, Milb the glandular apix-ndiigiii Ix-lnnging to it, wclotiineii iu
Mltent bv tlusbing with hloixl. the tlilniion U-in^ souielimea, lu in thu COM
u( the aaliTary gland, ibe rcxtilt of Uii' aetivity chiefly of vneodtlator fibres,
bataonrliniea the rntult of the oeniniton of constrictor imimUes and tone-
iban tbe mtill of lb<^ two oiinbin<-<]. .So also when the kidney eccretts
wfiw. ilamwls bemme dilated, and in general, wherever fiin<-lii>nal activily
see
THE VASCCtAH MECHAirrSX.
oomn iota (iliir. the iii(!tu1)»!Uni nf liiuiic vrliic')i in tlii^ liiuin of that nclivUy
is AMiMMl liy n more geiit-'routi tluw i>f bl<MHl ttiiviiijili iho liiwui-.
§ 180. rii*r;iii'((ur itermv »f (hf rein*. AUlHuiyli tW vriiis itiv providwi
vrith itiiim.-ulnr tibrc* itnd Mre (IwliiiCtlT coutnii'tilc, nixl nlthough rhythmic
rariatiiiti» of ciilihn.- ilue In coiitrnctioo* tauy Iio vtxa ill the f^rmt vdat
oi>ening iiitci llic heart, in th(? voins of the bntV wing, nn<l elaevrhera, aod
tiiuiilnr rhythmic viirintiou*. nl«j poeniblv tlu« to avtivo rhyltiniiv ooninu;-
timid, hill powhly ttlitif of nn iinlircly pm^ivc niitiiro, hnve been obeerved in
ihv portal vein*, verv littli; is kiiuwii of nny nr rvous arrangemeoU gorcm-
iiig the vein*:. When in the frog iho lirniii and epinHl corH are deetruvet).
very little blo'id comes back li> the heart us compared with the norninl
rii|i[]|y, and the heart in conacr[ue[ice appears almnel bluodle^ auil heats
fcehly. This iia; been, by some, re^'ardoti as more than can be accounlfil for
by mere loss of arterial tone, ami arcurdinh'lv interprelcil as indicatJn)! the
esiitieuoe of a normal lone in the veins de|>eQileiit un the ceiilrvl ncrviw*
syMem. Whoii the Utl«r is desiruyed, the veins become abnormally dtMcnd
and a tnrj^e (luantity of blood become lodged and hidden oa it were in ihen.
TUR CAPII.LAltT CiRCULATIOll.
|S 181. We have already some t^mc back (§ 117 1 meolioneil amue nf ih«
tulieut leatiircs of llie circulation throU{>h the capillariea, viz., the ditGctilt
Iiawage of the corpuscUe (generally in single tile, though someliiii&' in the
nrger ehanneli two or more nbrenst) and plasma thrmigh the uarrair clioti-
»el«, in a stream which though more or lew irregular U titettdy and ev«i,
not brukeo by pulsations, :ui<l slower than tliat in either the arteries or th«
veins. We have fuilher fiei^ii (^ HXi) that the cjipillarie* vary r«ry much
in width from lime tu time; luid there uiu be no doubt that tlie ehongM in
their enlibre ai-e cbielly of a piuuivt- nature. They ni% oxpandcil when a
hirgc "iipply of lilooil rein'hpM them lhr«iiigh thp •upplyiitg arteries, and. by
virtue of their elniticily, »hrink iigain when the supply is lessened or nilli-
drawn : they may also become expanded by nn ol>staclc to the venoaa
oultlow.
On the Other hand, as nc have also sialuil, there is a certain amount of
evidence that, in youn^ ntiiiunli> at all ex'eat». the calibre of a capillary
e«nal may vary, iiuile indq>en()eutly of the arterial supply or the vcuouV
outHow, in consojuenoc of changw in the form of the epithelioid eelU, allieil
to llie changes which in n mmx-k-fihra or muscle-cell constitute a conimc-
tion : and though the mailer re<iuirea further investigation, it is po^ible that
tiieoe active changes play nn important part in delermining the <|uaiitity of
blood parsing through a capillary area ; but there is as yet no Mtiitfaciory
evidence that they, like the corresponding changes iu the arteriea, are
gurernHl l>y the nervous syBtem,
Over and above theae obaages of form, tbe capillariea and niiuule voacU
are subject to cbansee and exert in6uences by virtue of which ther piny an
important pari in llie work of the drculation. Their condition >k-terniiur«
tbe uni'mnl of rejialance ottered by iheir channels to the flow of blofxi
thronjrh 1 1 lode channels, and deteruiiues the amouiii and character of that
interohangi! tx^tween the blood and the tiaaues which ia the main fact of the
circiilaiioi).
If the web of the frug'a foot, or better still if aotne tnuuparwnl ti«siie of
a nuimiiial Ih; watched under the micrusicojK.-, it will be obaervtil that, white
in the small cnpilbriiw the corpunohw arc prcMi^l through the channel io
single lilc. imtt nllcr the utiter. each corpuMlc as it panes uocupying the
TUB CAFtLLARY CIBCI'LATION.
287
«h»te bur« uf iIk* mnillnry. in tbo Inrgpr capillHrim fof the luatiimali, aur]
•ptdally ill llif •iiiBll (irlmr* aiwl \-mbs nlik-h permit the piUHiee of more
IMB iiDtr mrnusolc' nlirtNut. the rvl corpuicle^ niii iu the miildle of (he
(lBnBi>l. romiiiig n cnlonil mrv. tiehtceti uhioh and ihe Mde^ of the vcMeb nil
II alBllwl u n <'<>h>rl<w layrr. ru[i(»iiiin^ no rvii ^jrpiisclra. oulkil t)ie '~ |ilitiiriiatic
■^^b" or " [M' rip III- ml t"ae." this divi.qiou inlo a |H-ripheral tone aix) mi
^^^Blllfmiii it ilitc to tti« fact lliai in aoy atreaiu pai«iug tlu'nu;;h a rliuetl
Vrimail Um friction !• };rcuile«t nl ihe udca, and diniioiabe* liiwani iIk- axi^.
VTh* foqioarla paM n here tli« fnclioii is least, in the axU. A iiiiiu^ ximilnr
ftiia) Rin- is wen nbeii any liue parliclev arc driven nilh a siitlicient vvloi-ity
ia ■ unMiii iif fluid tbrou(;h a narrow tul)e. A» the veJucity in diniiiii*lie(i
Uie axial core Itecoiuca kn marked and dixap[M>an(.
' In tbe peripheral zone, (speciallr in that ui' the veiitf, iire frequently teva
' vbiM coqN»cl», Mmeilinca cliiigEog to tJiu «idei> of ihu vvnh;). siinieiinie*
I folliMS »lowly aJonp, aiul iu general tnoring irtcgiilarly. (topping for n wbilc
■lid (ben audilenly moving ou. The crearor lli« vehidly of ihe flow uf
Und. the fewer Ihe white LifrptiitclcM ia th<- pi?riplier»I loan, and with a very
npid flow iJiey, a> nell a» the ml i'i>rpii!tcic«, inav lio all confinnl t<> tb*
aial itrHun. The pn-m-nw of the whiu- i-orpiin-lci in the pcripheml ztino
hMhrea attributnl lu (li<.'ir bcinj^ »peeinlly li;;hler thnn the red corpusclca.
met vben fine jMrticlot of two kinil*. one lighter than the other, are itri\'eii
ttwwgh a iinrn)«t ttitie. the li<-avii.'i' parliel«« How'iii the axis and the lighter
bifc* tDure peripheral |H>riioiiii of tlie stream. Btit. besides this, the white
MnHMck* have n greater teti<lcn(-y to adhere to surfaces thao have the red,
~~ M mn by the manner in which the former become fixed to the ^l''^
and cover »lip when a dr~>p of blood is moiiated for luieroicopii-al ex-
■lioo. Tbey probably thus adhere hy virtue of the amwhoid move-
WBtaof tbdr protoplasm, »o that the adn«jiion is to be ci>n«idereil h[jI no
I a nierv i>byi>ioal as a physio] o)^ical procrau. and hunoe may be expecieil
■•nnr irilli tli« varying nucrilivu (vtndllions of the oorpusclus and of llie
Uwlruwh. Thus irhllc the am>i-arani>e of tbo whita' corpuscle* in tliP
pnipbanO aoiui may be due to their lighiiKMt, their tAinpomry altnchnienl
la tM iiilai nf Ibc vtwels Bn<l ehurnntenslic progrewion U iho mull of their
p>««r Co adhere ; and as we shiill preirntly see their amntboid raovem«Dts
mn atrrv them on beyond men.- nil(ir»ii)ii.
f 18t, TlirM" are the phenomnia of tin- norrnnl circiilution, and may be
nganlril ii^ indicating n i^tnle of niiiiilibriuni between the blood on the one
faiail and the blootlvesrf.-le with the li>>«ii<» on the other; hut a diflereat
Ale of things setx in ivheii that «iiiiilihriiini is overthrown by causes lend-
Off In what is railed inflaniination or to allied conditions.
If an irritant, such as a drop of chloroform or a little diluted oil of raus-
tsnl, be a|>pli«ij to a small portion of a frog'n web, loogue, mewnterv, or
■■■ other tRaiu|<aTvnl lisaue, the folloiring ebnnges may be observed under
the nlrmseope ; they may sbo be seen to tbe nie«enierv or other tmnnpsrent
liMH of a nuuouiAl. llie Arst eD'ect that is noticed i^ a dilation of the
iricrica, aoennijianied by a r|uiekeuing of the stream. The irritant, probably
W a dirwt action on the muscuUr tibrea of the nrleries, ha* led to it relaxa-
Mil of ibe muscular ettot and heuee U* a wid»iiHg; and we have already,
I lift. eKplaiii«d how *ueh a widening i» a «inall iirlery may lead u> n teni-
fnnry ifuirkening of tbe stream, lu coQse>|iieuoe of the greater flow thmugli
lh« arUrrim, Ihe eapillarieri become filled with corpuscToM, and nuiuv ]hw
•gWi pfwiously invisible or nt«rlv so on lu-ciunt of their oiniiiining no
'Wpwit*. n»w oonie int'i view. The vein* at the same time appear eu'
Wftd Mod full. If the Nlimnloi Ih- vt'ry slight, thii mny all [MM away, the
gaining their normal eousuiction, himI the oipillnrint and veiiw re-
888
Tnr TASCOLAIt MKCTTANISU.
tuniiutc tij l.lidr normal I'omlilum : iti mhor wurd-t, the eflet-t oftliu irtitnul
Id such & ca»e it simply u tcni[Hintn- tilimh. UoltM, Iwiwcvn-, t)i« chlor
f«>riii or niiutunl ha ap»li<'il willi tKjM-ciiLl cnn.' the ei&cU »T0 much tuort
proroiiiii], aiid a ^-ricn nl rfuiarkublc cbunjctv tei» in.
In the iKinnal circulntioti. un no hnvv jurI mid, while oorpuHcIc* may bo
Men iu itif jirnjilieriil, rihiMnulio xiim; but tbcy itri^ scaiilv in numUtr, and
eaob nno aflrr nluyiiif; for ■ liltl« time in one spot guctdciily g«ls ftvc. «oa>fr
time* nimniii by n jerk n* ll were, mid ihrn roll* on i'or a grealw or k« di»*
tuncir. In Ihc iirwi now under coneidcmii-in ii lurgc nurabrr of while cor-
pititclce doon gnllii-r fn tbc iMrinlitni! /ohm, cupccinlly of tbe veina and
vcniiiw c«pillnric». (that is of tin- inrgor oApiUiirlfi' which lire joining to form
vein), but nieo. to u lew cxlent. of th« nrtcrice; and thi« tnkee place
nlthongh the vcescia still remnin diluted nnd the streiim still coniinues rapid
though not HO rnpid at at tirst. Kach white corpuscle appears to exhibit a
grenier tendeoev to slick lo ihe sides of ihe vessels, and tnough driren away
from the arlenes by the Hlron^r arterial strenm, becuiDes lodged, so to
speak, in the voiiu. Since new white eorpuades are coiiliuually being
brought by the blood atreiim on to ihe scene, Ihe number of them in tlie
peripheral Kouee of the veins increases more and wore, and this may go on
iiDtil the inner surface of the veins and venous capillaries appear* to be
lined with a layer of while corpusclefl. Tbe Ainall capillaries too ooniain
more white corpuscleti thiinusual. and even in the arteries these are abood-
aut, though uut forming the distinct layer seen in the veins. The white eor-
miscle^. howi^ver, are not lliv nuly IxHiies present in ihe pertphenil xoneL
Though i» tbe oorinal circulation blixid-plalvU-l^ (see ^ 3^) cannot tie act-n id
tbe periplieral none, and hence luust be ciinfin<H) (on the view, which has ihc
greater support, that tbcMC bt)di<« arc really prcvenl in i]uit4> normal bliHid)
to th« axial stream, tlioy make th<>ir iipp«irnncc in that zone with th«
changes which we are now dc«criking. Indeed, in many cases Lh«y are far
mon: ahiindiuil than the while corpuscles, tbc Inllcr np)icaring imlwddcil nt
intervals in ninsees of the former. 8oon alter their appearance tlie indi-
vidual nliiiclcls lose their outline and run together into fonntfw miissos.
^ 183. This much, the appearance of numerous while corpuscles and
filatelets in ihc |x^ripheral zones, may take place while the stream, though
en rapid than at the very lirst. still remains rapid ; to rapid nt all eveiila
that, owing lo the increased width of tbe passages, in spile of the obsiructioD
ofFered by the adherent while corpuscles, the total t)UBnlity of blood llowiog
in a given time thn^ugh tbe indained area is greater than normal. But soon,
though the veascis fatill remain dilated, the etreani is observed most distinctlj
to slacken and then a remarkable phenomenon makes its appearance. Tlie
white corpuscles lying in conlacl wiih the walls of the veins or of tbe capil-
laries are seen to thrunt pri)i-e.-o>e« through the walls; and. the process of a
corpuscle incrwii-iug at ihi- ex|K-iitK- of the rest of the body of the furpu*<,'le,
the whole corjiuriclt^. hv what nppi-urs to be au example of amoeboid more-
ment. inake:^ its way ihniugh thi^ wall of the Te«el iiilo the l^mpli spue*
outside ; the [wrfomtiun ap^H-nn lo lake place cither in Uie oenieul sulwiance
Joining the epithcl)oi<l plnt4'i> ton-ilicr. or, pomhly, by an actual bn-ach
through (he subiitHiiee »f a plnl(^ Inr bri'iich being rcjHiired immetlintrly ufivr
ihc (xiiiHigc of Ihc enrimsclc. This ix the migrniion of llii; white corptiM-lm
lo which we nlluili'd in ^ -12. nnd takfs plnct- chiefly in tht- vcinn and cJipil-
larim, nut at all or lo a very slight extent in the arl<;ri<-». Thntttgh this
mignition thi- lymph spaces around the TwnU in the intl.imed art* bocoffi«
crowded with white corpuscles. At the nme time the lymph in the samo
Kpacn not only increases in amount kjit changes somewhat in its clietnicnl
ehanicters ; it becomes more distinctly and readily coagutable, and ts some-
times spoken of as "exudation fluid," or by the older writers u«"coiigulBbie
TUK CAPILLARY CI BCL'LATIOCT .
289
'D|ih.'' Thit tur;:«ciccnce of llie Irmpli spnt'ei', loc«ttier witli ihc liilaied,
(n«ikil cooditiou of tbe blo-idvcteeU, t:ivo8 rise U> the Bnelling uhich m nuo
tt U« faatora of iollaaiiuatioD.
1/ the inflKinnialioa now ]«mm oS* the wbiu rorpusoles onue to emigrate.
Htm to ttirk fur anj length of time to ihe sides of the veesels. tke mkiiiii or
bltod llirtui^b the v«aMb quivkeos XKaiu. atul tbe veuela themselves, tlioiigh
UicTiDkT rfmniii for m ioog time dilateil, i>v«ntiially n-guiit tbeir oatibrc, hikI
a»iriul I'troitntion (• roiMablithed. The niigruled eurj>iiH'Ii-.H move iinnv
ftMD tbe region, ilon^ ihv liibyntiih of lymph ipoeea, and llie Hurjilu) \ympa
ilu MM Rwajr algni; the Irmph zi\tnitM uud lymphatic veM>)ii.
i IM. The copilitiun of tiling, however, instead uf |itu«iug nff may go on
t» ■ (iirtber stage. Morv aim rnvrv while ourjniaclux. arnwk-d in thdr
ftm^ crond th*- chaonvis and block the way, wi that tlioiigh tht^ vtwels
ivnain dilMed tbe strmm b<-eiiniM flower niici «lowcr. unlit at la«t it stops
aJloKtiher and " stn^ation " or " stasis " «el« in. The red corpusoW are
•Irirea in, often in manee, among Ihc white corpusclci and platelets, ihe
(latiiiction between axial Mrcam and peripheral tone becoming lost ; and
ffUriak reioe. and eapillarie', all diatendcd. «nmetimee enormously so. are
iDed will) a mnaof niingW red and while oorpUBclcs and niatelets. When
tdul tto^ation occars the red eorpuH-les run together so tlmt their outlines
tr« no loaser diatiBUuiBhable ; they api>ear to Become fused into a homo-
pMMH red mass. And it may now be observed thnt, not only while cor-
!■«)(• but also red eorpusolee make ilieir way through the distended oiul
ikfftd walU of thf cupillari<». ehielly, at all events, at the junctiona of the
enjtheliaid plaits, into the lvmj>b apaces bevuud. This h spoken of as the
^malmi* ia the rvt cori>usGles.
This Utt«r "etagnalion" stage of inftammntioii may )>r tlie prelude to
bftber niuchief and indeo<] to the death nf the inflanie<l tiiwue, but It, too,
Eks the earlier stogm, may pas* awny. As il pasKv away the outlines of tbe
iwpuscles l)ecoino once more difiinct, thuee on the venous side of the block
(rMually drop away into ih«' neighboring currents — little by little the whole
doCracUoo is removed, and the current through ihc are« is reestablished.
TW alnwinj; and final arrest of the blood currcni described above is not
^ to OUT lessetiini; of the heart '» beat ; the arterial pulaations. or at least
Av ortcrtel flow, mav be aeen lo be continued in full force down to tbe
iHrtlii area, aud there ■» cease very suddenly. It is not due to any eon>
tuidioa of iIk- flmnll arteri<9< increasing the peripheral rfsialnnw, for lh«se
eminiK dilated, nomuiimt^ eievedlngly so. It must, thervforc. he due lo
«■!• now and unusual miMoncfl occurring in the area il*clf, and there caD
Wnodoubl thatthi* iKlobefonnd in an increased Icmk-ncv of the eorpiisclve.
apoeially of the white corpuscle*, to Mick to the tiidui nl' the resMls. The
(ecifsa of adhenivciiiw is not cauited by any (Jiange conHnud to the corpus-
riM t&etnseUes ; for if, after a tcmnorarv delay, one set of oorpusclee bos
■aaoced to pass away from the am-ctcii area, tlic next set of corpuscles
W«apit b> tbe am in the blond streiiin is subjected to the same delay and
ihs iorae apparent fusion. Tbe cause of the inenaMcd adhesiveness must,
dnsvfere. lie in the walls of the hloodvessds or in the ti>eue of which these
ktm • parL That the increased adhesioD is due to the vnecular walls ami
•SI primarily to the corpuscles thenuwlvee is further shown by the fact that
if fal tbo froc, an arlilicial blood of normal saline solution to which milk bm
UlB added be sul)stituie<l lor normal blood, a stosi* may by irrilauta Im- in-
ia which oiU^lobules play Ihe pari of corpuscles, and by their Oggrs-
bring about an arreat of tbe How.
W« am drivt^u to conclude that there exut in health certwn relations
u krtveen thcbIouil,on tlieuneluind.and tbe walls of the vesseUon the other,
L ^
290
kR UGCUAI
by wliich ihi" teodCDcy of ihr oirimsclw to mJlKTr to llwr Monih'tw*:!* i.* kcpi
itilhiti ci-rti>ii) UmiU; tht^- rcliUtong ro«iic<|upiilly di.-UTmi»o lli<' iiormnl
flow, witli it)t axial strewm ami peripbrml zone, and ihr normal flmount pf
p«riphcriil rnnsCaiios: in intlnmiiiiitionN these relstioni, id a manuer ve
canuuf M yet fully expltiin, nrc <lif>turl)e(l io that the tvodencv of ih«
coriHiBcles tu adhvre Ui the 8i<IeB of the vcm^I in larKely and pro<;T»8ively
ilicr«aw(l. Hence the tarning of the corpUBcIes in spi[« of tlie wideutiij; of
llieir path. ai>d finally tJieir a^lomeralioD and fUBioa in L)i« dtUended
<ihanuels.
We may add that the chan^ occurring in the Ta»cular walls also at least
facilitate the migration of the corpuscles, and uuxliry the pASMge from the
hliHHl to tlie tissue of ibe llui'l parU iif the hUnnl, the lymph of inflan>i'd
Hri'iM being richer iu prDteidv thuu normal lymph.
W« tuiiM not, h(iW'L*ver, ptintue tluii i>iihits:t of inStimmutiun auy fiirthor.
W« liav«Mtd enough tonhow thai ihi- iM-riplienit rminltinuc' laml (x)npH.i|iiently
alt l)mt di-jx-ndti dd that [i<-ri]ih<>ral rmtftaiicv ii> niii wholly d<MmiiMv>l by tba
varying width nf the minute paiwagct, but is al«o dependant on ihv vital
ooiidilioa of the tinue of irbich t)ie walU of tbo paangn form a fart. WImq
tbi tiffu« v in hcaltb. a certain mi»[atiee ii omred to th« passnBQ of blood
thiYiiigh l]ie cnpillari«» and othvr minute y^meiU, and the vrhoT« vascular
mcvhanidm \* aaa)>led to overcome this TMUtance to Buch no extent that n
Dornial circulation can take place. When the tissue becomes aflect«<), tbe
diaturbaooe of the reUttions beiweea the (issiio and the blood may, as iu tlw
later stages of ipflammation, BO anLfmenc the mislauce that th« poaw^ of
th« blowl beoofDes at first diflicuU and ultimately impossible. Aud it ia
quite open to us to suppose that under certain ciretimstanDee tbe reverse of
tlie above may oecar ui this ur that area. ounditiouB in wbieh the nesistaace
may be lowered below the nunnal, aud l)i« circnlation in the area quiiJteaed.
Thtis tlur vital conditiitH of the tiwue becomes a factor in the roaintraaiioe of
the cJratliition; and it is possible, though not yet proves), that tJi«M vital
eondiliono ar<' diri'ctly umW (he cloniinion of the nervoui Kyxtem.
§ 185. ChanfTw in the pcriiihcrul miistaiice may «lw> t)e brought about by
chanjjpit in ihi- character of the IiI'mbI, (ipfciitlly fiy diaiiKiv in the rvlntivt
aHMHiuc of gases present. When ii slrmm of .K-fihritiartHl blood if nrlifidallf
driven throtisb a perfocllr frt^li i-xrixivl (irgiin sucb as tlie kidney, it is foana
that ibe reeistanoo to lfie tlou- lyf IiI.mmI through the orgiui. measured Au
instance by the amount of outflow in relntinn to the prasure exerted, rarka
considerably owing to changes taking pine.' in the organ, and may be incroaned
by inoreastDg the venouti character and diminished by increasing the arterial
CDsraeter of the blood. Kemaikable changes iu the renistaJioe are also
brought about by the addition of small <junntitieB of certain drugs such as
chloral, atropine, etc. to the blood.
These ohanges have been attributed to tbe altered blood acting oil the
wnllK of the vessels, inducing for instance constriction or widening of tbe
small arteries, or it nuy Ixi afii^iting the capillaries, for it hsH be<en aa^rled
that the epithelioid itlaiea nf the capillari«H varir' in form aotMinling to the
relative i|nuntiii49 of cHrboEiio add and oxygen present in the IiIimkI. But
this is Dol the whole explanation of the matter, since similar varisttions in
resistance arc met with when blood i^ driven through fine capillury IuIh-d of
inert matter. In nuch exiwriment^ it in found that the resistance tu the flow
increase* with a diminution of ilit- oxyg<-n carHrtl by the red corpuscles,
and is niodiHe^l by the a>ldition to the bIi>od of even small •juanlitius of
certain dru^
It is <d>vious then that in the living hcxly the |N:ri|>h<-ral n-)ii.ilance, lieing
tbe oatcoDie of complex eonditions. may W modified in many ways. Ex-
CHAKOCS IS THK (iCAJfTITT OF BLOOD.
291
meure leacben ua thai, ev^n iu dealiiif; with Dun-ltviug iii^^rl miUlt'r. l)i«
low of Huid Uirougli npillary luhet may be modifiei) ou tliv «■■« liUDil hy
IcbtDgee ill tli« subetaDoe of wbicli tli« lubea are CLunpistd, iiml on lira
'tuber bawl by cliaoices in tli« chemical nacure (even imtejitradnnt af the
specific ({rnvily) of l)i« tluid wliJvb is umhJ. In Uie living body both th«
fluid, tlie blood, aud the walU of iJm: minute reawls, being both alive, a ro
inveHMntly Mibjcct to •-hanfri:; the chuii^ iu the oiie moreover are Mpidilc
of neeliDg upon uud iudinnnx clmngifti in tlic Otli«r ; and, la«tly, the dinii^t's
both of tbeonv and of ibt^ oltn^r may hv. priiuiirily mt going bv cvoute litkiug
place in tome rmri of tin: body far amty from ihv region in which these
ebutgOB are molifying tlic r(»i»Umcv to tiiv How.
CUANGEK IN THE QUAKTITY OF JIlOOD.
^ 186. In an arttficifll Bcb«ni« chan>^ in llie total nuautily of fluid in
cirvulflliim will have an immc^liate and direct ell^ct on the arterial praature,
iorreai« uf tlie (juanlity heightening and decrease diiuiuiAbintr it. Tbu
elfevl will be produced parity by Ihe pump beiu|{ more or lew iiU«4 at each
Dlmke, and |iartly by the peripheral reaimaiice being im're&ieil or diminixhod
by the greater or lew fVilneM nf the small f)«rii>herul chnntulK. Thi- vi-nmia
praaure will under alt ciroumstanoea be rawed wilh the i f tluiil. but
Ihe arterial prnwurc w ill be raiaed In proportion only m- . -. _ ■■■ ijte lEnvtic
wnll* of the arterial ttiben are able to exert their etaxliinty.
In the natural rircutation the dirvi't rwulta of vheiigi! of cjuantity are
nimliKed by iximiKywatory arraiig<;mcHlM. Thu» experiment Hbonc that
rhea an aiiiRinl with normal bloiHl-protcure w bled fmiu one carotid, ihe
RUT« in the other carotid Hinks m long a» the blct-<ling is going on,' and
'^laaios dt'pTtwMd for n brivf jteriod iider the bleeding nae ccsmmI. In a
aliort lime, however, it regains or nearly rcgaina the normal height. This
re<»vorT of blood-prowure. after liemurrhagc, is witae»f«d so long ns the
Ima of btood doM not amount to more than abuul ^1 per cent, of the body-
weight. Beyond that a large and fre'juently a «uddcin dangerous pennaneut
deprmsion is obeerced.
The restoration of Ihe pressure after the ceBsntton of the ble«diD(t is too
rapid to permit us to suppose that the (juantity of fluid in the bloodVQSMt*
is remured by the withdrawal of lymph from the extra-vasctilar elements of
the tuuies. In all probability tlie result in gained by an increa^ action of
the vaw>inotnr nerves, increaung the peripheral resistaiiee, the vasomotor
centres being thrown into incraased aetion by the dimbutiou of tbdr blood-
■upply. W^en the loss of blood has gone beyond a certain limit, this vaso-
motor action ia inaufllcieut to compensate the dimiuiahetl tjuautity (iMwiblj
the vasomotor centres in part become exhausted), and a consldera^>le de-
preuioii takes place ; but at this epoch the loaa of blood' IVequeiilly causes
anwmic roovulnions.
Similnrly when eii additional quantity of blood ia imeeted into the vewMrts.
no marked incresKt of blood-preBsure ia observed so fong as the vaw^motor
centre in the nieilulla oblongata Is intact. If, however, the cervical spinal
corti be divitlcd prttviouf lo the injection, Ihe preaaure. which on account of
the riMiioval of llie medullary vasumotor centre IS very low, i* jicrmniieiitly'
raised by the injection of blood. At each injcelion tlie prvmure ri"(», falls
sODMwhat afterward, but eventually rvnuiins at a higher level tliao before.
> Cbldljr In <Diuca'wi(* of il>o ft™ oip*nlus In llis vcwel fttnn wbloli the bleedlnaliflolnc on.
cuauiic sft a f tcki ao*.! of ibc pcripbcni mltiaaca and to InMlliis lo ■ sDueisl loveiuia at Iba
Hmd-fftMiire.
I
290
THE VASCTLAR JJ KCH AN'ISM.
I>r nhirli tln! Icntlcncy of the cuipii^clcf to adlioro to the bluodvcxcls U kvpt
wiihiri (vrtHin limit*; ihese rolHtiniin c(>fi«e<|iie>ii(Jy <lv4«rmLi)e the normsl
tlon. willi its axi.tl ilrcam and |)crij)hpnil lunc. nnd the nurRinl itmount of
periphcrul rct^istance : in iufliiiiimationi', thnt< rrla{ioii», in a maiinpr ire
caonot us yet fullf explain, are di§liirbc<l to ttisl the lendcDcv of the
corpusoln to adhere to the sides of the vcMei if luridly ond pru;^reMiveIy
increaMd. Henct ihe tarrying of ibe oorpUBdw in sjiilc of the irideniae of
lh«ir jmth. and lioatly ilieir aggloiuentioo and fuMon iu the diateaoed
cbsnneU.
W'v may add itiat the ehangee occurring id the vasoular walls also at lout
fueiliintc the niigralion of the corpusclea, and tiiodify ibe paasag« tmm the
bliMjd lo iht- tiMue of the fluid paru of the blond, dm lyinph of inflamed
anrtii hdng richur in proteida than uonnal lymph.
We must ni>t, however, puntU(> ihia subjcvt of iiiflRiaraation any faiihtr.
W« have mid enough tonhowrthat the [M.Ti|)heTal reit'istauov (and oouaaquoDtly
all I hat dcTH-iids on tliut iH-ri])lientl rraLxance Lt not wboUy dcteRoiiun by IM
varying nidth of the minute irnxNuges, hut is al«o dcpODdoni on tW vital
eondition nf the tinnw of wbieh tht- walls nf the pMBitgos form a part. Whra
the liwiic i* in health, a ecrtain reoirtiince is onbnd to tlie pMnW* of blood
throii^'h the cumllariM nnd other minulo vends, and llie whole vafcularij
nieehiiniEim is adapted to overcome thi? resistance to such an extent that t-'
Dornml circulation can lake place. When the tbeue become* aflboUd, the
dUturhuuce of the relations between the tissue and the hlood nifty, as in tfae
later stages of inllHmmaiiou. so auL'nieol the resislance that the paeeajce of
■lie \tUioA becumes ai Bret dltlicult and iiltiniately impouible. And it
Suite open to UB to AU|))Kise that under certain circurostanoea the reverse
le above may occur in this or that area, conditions in which the reibtauee'
may be lowered below the normal, and the circulation in the area <|uickeuc<l.
Thus tlio vital condition »f the tmuf btsMineit a tiwtor in tlie maint<-nancu of
the circulation : nnd it w poMtthle, though not yet provei), that thc«e vital
oondilions arc directly under (he dominion of ih« nervous system.
^ 18A. Change in ihe perinhornl rcislanoo may also be brought about by
ohaofea in the character of the blooil, especially by chanra in the relative
amount of gases present. When n stream of den bnn Bled blood is arliticiully
driven through a perfectly fresh excised organ such as the kidney, it is founds
that ibe reeiatance to the tlow of blood through the organ, measured tot
instance hy the amount of outflow in relation to the preMUre exerted, varic
OunMib-niblyiiwing to changes taking place in the organ, and nay be increaw
br inereiiBing the venous character and diminished by increaaing the arteria
cnarncler of the blood. Remaikable changes in the resistance are aim.'
br«Hight aliimt by the addition of small ijunntities of c«nain drufrs such ua
chloral, atn>{>inc, etc., t<> the hloMl.
Tliew chaii^ bare been attributed to the altered blood acting »u ilw^
walls of the viwela, Induelug for iuataooe constriction or widening of th«
small artertet, or it may be aHecling the (uipillarieg, for it has been asserted
that the epithelioid plates of tlie capillaria» vary in form according to tbe
relative quanliliw of cnrbimie acid and oxyjiteu |>reiienl in the blood. But
this is nut the whole explanation of the matter, since xiniilar variations in
rtwlanc« arc met wiiJi when blood U driven through fine capillary tubes of
inert ntalter. In such ox|M:irinicnla it in found that the resixlanoc to the flow
increases with a diminutinii of [Ik- oxygi-n e»rri<'d by the red cwrpiweles,
and b modified by the addition to the blood of even small (jnantitim of
eertain dru^
It is obvious then that !n the living body the periph<Tal reustanoe, being
ibe outcome of complex conditions, may be mollified in many ways. £x-
CttAyOKB IX TttK QUASTITT OF BLOOD.
2»I
Mriuiet toaobtfl at thai, «Teii id d«aliiij; with iiMii-tiving ioett matter. tl»e
Bmt of fluid throuvl) cApillorj tul>e» may be ninililiis] ou the on« hutid by
changTw it) lli« substance or which the lubut tm roiupoMxl, and on tJ»e
olbsr hand by chaoKee in llie chemical nature («tcii indviH-ndcuI of tfa«
•psdflo gravil}-) of tli« fluid whic^b is uw-hJ. In thv living hoAy both lii«
fluid* lb* blood, and the walls of tli« minute veaMl*. hdng both alive, ara
iooaaaastljr aobfoot t» i'ban^ ; the diau^ in the one moreover nre capnble
of rvarliD^ upon and inductnic chnnji^ m the otlivr; nnd, liwtly, the channa
boll) of theon« and of the t>lber mav bciiriiuarilywt going bv events taking
plaee in mme mrt of the t)ody far nnar fntm the region in which ibeae
dwagsan iDoaiQring the reautuuce U) llie flow.
CnxnoFst ix tiie QuA.TrrtT of Blood.
i IM. In a» arlifinni M'hemi? ohaii^eB in the total •luantity of fluid in
dnolatiou will have an immediate iind direct ciToct i>u tlte arteriiit pMMuro,
iacraaaa «f j,b» quantity heightening iiod deoresae diiDioisbing it. Thtt
»IA«t will be produced partly oy the pump bang more or leas filled at each
■tmke, antl partly by the jieriphoral re^isLniice bi'ing iticruii.-ieil or diminished
hy tbe grvatcr or less fitliiewj uf llie muall [wripticrul diaiiUKU. The venous
p€vmun will under all cireunKtancea be raised with the increaae of fluid, hut
Um arterial pren^ire will be raised in proportion only *o long aa tliv clastic
wmlla of tbe arterial tubes are able to exert their ela»tielty.
Id tbe natural drvuUition the direct reeutta uf ehaua* of i]aRntll]r are
tncullfied by iH>m|ienaatory arrangemeui*. Thus expenmeut ahowa thai
wbea ao animal with normal blood- prexiu re iw bled fmm one carotid, tbe
preMare iu the oilier carotid rinks ao hmg u lliv bleeding a gulag on,' and
Wiaaiaa denraaod for ii brief poriud nllcr the bleeding has ceased. In a
•boft Ubm, Dovever, it n-gnint or nearly rcgaina the norma] hdghL Tbit
ncanrf of Mood-prtaiure, after hemorrhnge, is uitneMed so long aa tbe
loai of bUml doVH not antounl to more than nb<jiit :i per cent, of tlie body-
wai^t. Btyond that a large and frc<|ucnt]y n Mudden dangerous jtermaneut
dtpnnioo is obser\-ed.
The restoration of the prewuiv nfler the ceasntion of the bleeding ia too
Ttfid to pennit us to suppose that the (juantiiy of fluid in the bloo(TTaKlB
IB nutued by tbe withdrawnl of lymph from the eslra-vascuhir eJemento of
the tiSBBeii In alt probsbility the result is gained by an iiicreanetl action of
the TMomotor nerves, incneong the [>eriplieral reBiataooe, ibe va»omotor
cenme beiiig thrown into tooreaaed action by the dimtDulion of their bloo<l-
•apply, ^^*len tbe loss of blood has gone Iwyond a certain limit. thi» vaso-
laolor action u insuflicjent to compensate the dfmiutalied ([uantity (iioesibly
the rasomotor centree in part become exhausted), aud a consiiiferaule de-
prearioo takes place ; but at this epoch the luaa of bloud' frequently causes
aa»aic convukioDa.
Simllnrlv when an additional qunntily of blood is injected into the vessels.
IH> marknl incrvuse of blutHl- pressure is observed xo long as the vowmotor
lBntn> in the tiKitulIa ublongaln is intact. If, however, the cervical spinal
tarA l»e diviiled prrvicm* to the injection, the prenurv. which on ocoouut of
the reinuva) uf the medullary vasomotor centre is very low, is permanentlr
raised by the injection of blood. At each injection tl»a pressure rises, falls
rhat oiUrward, but eventually remiiins at a higher level tluin before.
'1 rmm which ib« bldedliw li ai>lhc ftu.
K leailliic Iu a («(uit^ looarliiii of III*
> CUMr la H»i«»uwnB» of Om lim opcatM In Uio *«■
mimmm ■ ci«* mi tf Uw K*)pb">l»bU*M and
294
TITE VASCl'LAB MECHAXTBX.
which ascending from the mucous inembrane of the stomacli alou^ c^rtnin
afferent librcH of the VBgui to the raedulla obloD^la. so augmenl the actiou of
the card io- inhibitory ceutre as to stop the heart for s beat or two, the etop-
ptti^e beiaif frecjuently followed by a temporary iocreosc in the rapidity aii<l
force of tne beat. Huob a passing failure of the heart-beat, in iij^ Mdden
onset, in ila brief duratiou, and in ilie reaction wbidi follows, very cloodjr
reacniblcn i)jc temjiorarY inbibiiiou bruugbl about by artitirinl xtimulation of
the vmgiu. Itut lhft<e Aararter* are not eweutial to canliac inhibition. Fur
it Diuvt be reiurnibert'd that tbe oentral nen'oiia aptem potMMM, id the form
of natural ui-rvoim impuli^s of VHriouti origin, n niMina of ■timtdatioti far
finer, nioru delicate and niorti varied ihan anything we can effioct br our
rough niean^ of induction cmln and clcclrmt*^ ThiiK in nmnv nun of fnint-
IDE. lh« henrtb.-jiis, inotvnd of nloojiiiij; nbriijitly. gnKlualfy die away or
fade away it mny lie to tin nhiiilotc brief arrcsl, bul more frcijocntly invrely
down to n fcel>lf.-ne«s uhich in inxuflliciciit to «upply the brain witb a '(iiajitiiy
of blood adei^unie to mniiiUiin conxciiMiNKw. nnd then in many cnsefi, at all
events, arc resumed, or recover strcnt'th gradually and quietly without any
boisterous reaction. In all probability all cas«B of uniple fainting from
emotion, pain. dige#livc> trouble*, etc., ns distinguished from the syncope of
actual heart disciue. are inslancca of vagus inhibition, and though we cannot
accurately repro<luce their varied phases by direct stimulatiou of tbe va^ua
trunk, we may approach them more nearly by producing rellox inhibition,
n> br mecbanical irritation of the abdomen, see ^ I5U.
^^hethe^ definite temporary tri'egularity ia ever brought about by nieitna
of the auguentor fibres, we have at present no dear eviileneu; hut cilms do
oocur of palpitation without previous stonpage. caies iu which a few hurried
strong l>eata come on, para of, and are followed by feebkr lieat* : niid ib<w
may possibly be due 1» «ume troiuieut influence of augmentor fibres thrown
into activity as put of a reflex act or otherwise. And though we have no
direct expenniBntal cvidonco, it is very probable that the acceleruiioD or
augmentation of the Ix-itl. or a combinnlion of the two, wbich so often follows
emotion, is carried out by augmcnlor tibres.
Iti all probnbilitv. however, irregularity in the heart-beat is much more
frequently the result of inlrinsic events, or the product of a diaonlered nutri*
tion of the cardiac substance. The normal nutrition sets tiie pace nf tbe
iwmal rhythm. We cannot explain how this is nflbcted ; nor can «e exidaiu
why iu one Individual tbe normal pace ia set aa low as 50 or even 30 beata
a minute, and in another as bijfb ea 90 a minute or even more, while iu uHmt
pemna it it about 70 a ralnute. The slower or the quicker the pace, though
not normal to the H[)e('ies. must be coDtidered a* normal to tbe iudiviilual,
for it may he kept up through lung vean in an orgtiniHin capablu of ciurviug
on a normal man's ilulicn aiici wurlc. So long m nv cannot explain tl)u«e
diiTcrenccs wc cannot h'ipp (o ex[i1ain how it i.* that a diwrdercd nulrilion
bringt about an irregular hcart-bctil, rithir the more r^ilar irregularity of
a "dropping" pulse, ihut is. a failure of iM-tiuence rather tnan an irregularity,
or a tnorcBctivdy irn-gular rhythm, such nn that aceompanyiDg a dilated
ventricle. We may, liowcvcr, dislinguixh Iko kinds of irregularity; one tn
which, in spite of all litvorable iiulntive conditions, the cflrdiac substance
cannot secure, even perhaps fur a minute, a steady rJiytlini ; and another in
which the rhythm, though normal under ordinary circumstances, it, so to
•peak, in a condition of unstable eouilibrium, ao that a very alight cliange in
oooditions, too much or too little blood, or aoroe aiuall nllemtion in the com-
iMsition of the blood, or tbe advent of some, it may tie sligfat, nervous
impuW. augmenlor or inhibitory, devehipa a lempornry irregularity.
J 19L No out thing, perhaps, ooncemiug the heart i* more atrikiug than
SOME reaTL'RKs or cinci-LATios.
20S
r
the &ct lluu B bniTt which lua gon« on beaUug for manir y«H», will) only
tctdpormry tn^gularititt, KD<t thoce few and far lwtwe«ti. u liiiurl whirh muxt,
thambiv, haw axeculcd n-ith long-continued regularity, mimy niilliim* uf
bcAU, afconld sudHvnly, appHrendy without warDiog, utlcr a brief Hiokorioz
•tnintt, oeue to biMit any tnor«. But we niusl remember tliiu «u-h beat m
■n eOuK — an eflort noKOTer, which, as we bav« *e«n (^ 1J>&), ia th« bett
tluu the heart can make at the niomeni, the accompliwhaienl of mch beat
ia. a> to apeak, a hurdle nbich hsit tu l>e le»pe<]^<ine of tbo long Mriea of
hairltea which wake up tin- t«te«:ple-tbn«f uf lifi^ At anyone leap failure
may ocear; m loo^ aa failure doea not occur, w long a* the Ix-tit is made,
ma<\ m lUr proportiou of the ventricular conUtnla are di^chargmi iiilo the
graat rawl*, the chicJ* cmt i* Kttinol, and whcthrr the leap is mn<te dumaity
or well in, rvbuivcly oonaidereu, of w:c(iadnry importance. Uut if th« beat
b* oat nuitli-, vvxrything alnwat (providt^l that the miss be due not to vtgaa
tnhitMtiai) Irat to intrinno eTenia) is iinfnvurable for a succeeding bent; the
myatMKMia moloculor change*, by which rhe actual occurrence of une beat
pfwpana ibo wav for the next, are missiop, the favorable iril)ueuoc« of the
extra hmJi of blootl through the cornnarj' arleriea due to a preceding best
mwm mimiag aito, and even the distension of the cardiac cavities, at fint
fltTntaUSt ipeodily (>•*» the limit and beooinee unfaTorabl& And tlieie
aolowanl influencea accumulate rapidly as the Unit miaa is followed by a
•■eowi. and by a third. In this vrav a heart, which has tie»i brought into a
Mate of unstable e<|uilibriuni by disor^lered uutritiuD (at fur iniitttDce by
ltu|ierfp<.-i curuuary ciretdatiuii, such an tn^im tu ncoum[>any diMiaMf of the
aofflie Talres leadiDK to rcgurf^ilution fmm tlie aorta into the ventricle, in
whiUi CMC* sudden death is »i>t uncommon), which i* ablv ju.->t to accoinpliob
••eh beat, but u» more, which hiw but a scanty saving stun- of energy, uniler
Mn« etrain or other untoward inDucncc, misses a leap, falls, and is no more
•Utt to rise. Dwibtlen in such vki>c:< i-outd adc^uole ariiflcial aid be
pfOOMly applied in time, could the fallen heart be stirred ereii to a single
good Mat. the favomble reaction of that beat mi^hl briu]* a successor, and
M once more start the series ; but *uch a jwriod of >{race. of potential recovery,
is a brief one. Kven a coarse skeletal muBcle, wlten out oB* from the
drctdation, soon loses its irritability beyond all recovery, and the heart cut
otf from its own influence ou itaeif runs down so rapidly, that the )wn(id uf
poMible recovery is measured chieDy by seoonds.
S IM. Tbming now to the minute arteries and the peripheral resistaoc*
wUcb tbey teguiate, we ntay coll to roiutl the exislenoe of the two kinds of
iiirhanlsin, the vaso constrictor mcchanisTO, which, owing to tbc matntcnanc*
hj tb» entnd nervous sntem of a fmic innuenoe. caiiljc wnrknl \mth in a
pOiids* comicrichir, and in a negative dilator direction, and the vii«i>diUtor
OMeluuiEan, whieh, as (iir as we knuw, cxcrt« its influence in one direction
oaly, rU., to dilate tbc blomlvrMcIic Tbr latter dilator mechanism wems.
■s m hkT* Itaa, to be used in fpceinl innlaooes only, an cocn in tbc cases of
th9 chorda tymiMUii and ncrvi crijri-iitiv ; the use of tlie former couatrictor
MSghanirai appears to be more general. Thii; the relaxation of the cutaae-
ans arteries of the heiul and nock, which is the cwcnlial feature iu blushing,
se«BW due to mere loea of tone, to the removal of constrictor influences pre-
rioasly exerted through the vaso-constrictor fibres of the cervical sym|m-
tlkctia Though probably dilator flbres pass directly along tlie roots of the
esTTical and of certain cranial nerves to ibe nerves of tiie head and neck,
we hare do evidence iliai the^e come into play in blushing ; as we have seen,
Musbinc mav be imitated by mere section of the oen'ica] syrofMtlictic. So
■ho tl>e " icfiiv> " and rednos uf the skin of the whole body, i. e., dilation
jmwally of the cutaneous arteries, which is produced by external warmth.
296
VASCITLAII
ie probiiblr nnothcr iiiMniice i>f ditiiiiii^licd activity >if tonic c»nMnrt<>r iiiAu-
enns; though th? rcmill, that tho <)iUlion produced bv irarriiiug an tiniiumi
in an oven is ervatet than that pniducpd bv §cctiun o( nerrcs, tMtns to poa&t
to the dilator nbre* tor tlie ciitHneou^ vcetels which, as we have seea, probft-
l>l;r exist in the Hciatic and bmchinl p)exua«e and possibly in nil the spitinl
nerves, alito taking pan in the nclion. A similar lues of cuuBirictor action
in the ciitaneou« vessels uiav be the result of certain emocioDS, whether gmitf(
Ml) far an actual bluHbinij i>{ the body, ur merely produdQg a " glow." Timi
elTeri i)^ coll), on the tither luind, and of certain eiuotioiu, or of emotion
tinder certain eunditiuna, U tu increase the uonetrictor aetion on tlie nitnuiy '
oar veAwlH, »n<l the *k!n grown pule. It may be worth while to point out,
that in lioth thr :iK<v > ,lii4, wliile both the I'old and wannlh {iroducv thrir
eDect, chiefly m ' . i'tiI.'> thrnugh the central nL-rvinii Ht'sLeni luid vi;ry
slightly, if al all, by dimcl action ou the ulcin, their action on thr c^ntr
nervous svftv in i* not oininly a gvnenil augmentation or inhibition of
whole vacomotor centre. On the eontrary, the cold, while it oonstricts the
«utaneouH vcMels, so act* on the vnsomotor oentrv a» tu inhibit that portion
of the vasomotor centr<- which governs the abdominal splanchnic area;
while \vm blood is carried to the colder skin, by tlic o{)oning up of ll
apliiuchnic area more blood is turned on to the warmer regions of ttie bodv,
and the ri»e of blood-pressure which the constriction of the cutaneous veM«M
tended to produce, and which mii^ht be undesirable, is thereby prevented.
Conventely when warmth dilates the cutaneous veeaeU. it at the same time ,
cotiatrictn the ubdominal aplanohuie area, and preventa an tindeeirable
of preMtnre.
The warm and flushed condition of the akin, which follows tlie drinking
of alwholio tluidw. U probably, in a dimilur manner the re>ult of an inhibi-
tion of tliui imrt of the vneoinoior centre which governs the ciiuneotu
arl«rie» : and it is probable al*o, thni except for the local effect of the fluid
on the gastric mucoutt manibrai)c, whereby some amount of blushing of the
niBtric bloodvefeels takes pliii-c iim a rcHex act, thii> eHcct on the vessels of tfafl
akin is aoconi|)anied by nn inverse coiiMrictor action in the splnnchnic anti
This last point, however, has not been provci.1 experimentally and may not
occur, since the intluence of the alcohol is nt the same time to increase the
heart's action, and thus to obviate the fall of preMure wbioU would certainly
occur were the cutaDeous and splanchnic vascular are« to he dilated at the ^
Mine time. This elfeci of the alcohol on the heart may be a direct action 0(4
the alcohol on ilic (-iirdiiic Hulwtance. being carried thither by the blood ; btit^
the ctlvct , in being an aiignienlAiiuu of the force and acoeleratioa of the pace
of tho hcart-lKuii of a temporary cliaracter, followed by a reaction tn lli«
direction of fceblvnem and xlonueM, »a atrikingly rvMtnible tlicefltMla of arti-
ficially stimulating tin- •anliac augmentor fibrts. thai it is at least probable
that the alcohol diMv net ujmii the cardiac an^tmuntor mechanism.
S 193. The inrtiicnci- on thi' l»dy of cxcrcis".' illunlmtat both the maiuiePi
in which the two \-awiilar fnclon, tho hcurt-beat and the peripheral rasiiM
aoce, are modified by circutiislancra. and the mutiml action of these on each '
other.
When the body riawee from 11 condition of comparative rest and (juiet to
one of exertion and movement, the metabolism of the skeletal musoloa (aud
of tlie nervous dystem) is increased imd more heat is generated in them.
We know for certain that the increased metabolism throws into the blood of
the Tc^iiu coniinv from the muscles an increased amount of carbonic add|i
and It U probable, but not m certain, that it also loads the blood with lacliol
acid and other metabolic ]ir(>iluc1s : at the same time, there is an increiiMvl
miisuniptioD of oxygen ; the blood of the body leiKJs to become km arterial
80JIK FEATtFKBS OF CIBCULATIOK.
297
and man T«noii». In dMiing i«i(li nvpiriition, ne vtinll tm tlinl tW iiiflii-
«B<» thtu «[i*Tl«J OB llic bl«Kl IfKil* to aa iiifTnuw in tb« mpirnlor}- move-
nteijU, BDil «<■ ihiilt furtlior rv thai Uui nimv vigorous working' of lli«
rMpiniinrT Dump, «inoo il [immirfpN ihr flow ol* l>)on() to mkI through tlw
bmrt «Dii liiD^, iiiiKkvns ntiil xtrrngthcRu itic livnrt-lxrale. l'<«»iblT this
tDwi nK^-tiniiicnl rneet of th« more vigoroiM bratthiug is sufficient bj iiaclf
to aocuuiii l^T ihr JiicrMM in the freqnenoy and vigor of th« fa«nrt> srtiou,
ImiI it i> nion? than jmilwble ihni il i* Inechatif^ condition of (he tiloivl,
whirh. tihilv it hurries on (he respiratory pmnji. al*o stiiiiiilates tbe vafcutur
paritp, rilh«r )>y n direct HctioD on the ttardJac Mibe(ance, or thn>UKh iJie
wv<liuiii of the ceD(rKl iier^'oua «n1eni anil (he nuj^inenuir tibivH. If, na
vxprnmenla Mem to obow, the ioereated vi);or of die reaptnitor}' inoveiiienti
MMapeoatcs, or even (wei^eompeoMtefl. the leiidenrv of die whole blood to
bvoM&e more veuou*, >o thnl during exervae (he blood, vhioh ts ilUlribitt«d
bjr Ibv wvta, bMumIIv dont Dot oondiin more carbiiuic ncid itnd Icm oxygen
tkaii the rest but even iIk' rvvAne. then ihete eflbctti uuK be due to aome of
UMiiroducti of muMMiInr wctalxilititn otWr iban carbonic >cid.
The Mme cbaDgr<l coiKliliiin <>f hlnin\. while it tliUR excidw the henri diUU*
like ftitiuteoni rnvclv, m if rlmrly shown by thL- wnrni Ilunhed tk'iu, nnd at
tb«^ Nimv linx^ ihniwi into nciivity ihc pcwpimtory nicchnnisra viilh vrhicJi
WIT ahalt hrr«MA4-r have to dc»l. Tlier*' can bo no doubt, v wo uhnll tv
Inler on, ihnt the iwrspiratiun which acoompaaies niuxculiir cxcrciw i«
bfmght abotit by m4«n< of the central nervous tpAvm, luid wc mnr nlnxHt
with eertainlr conclude that ihc dilation of tlm cutaneous arteries i" also
bcoocbl about by means of (he c«D(rBl nervous system, ami most pmbnbly
bjr HMOi of an inhibilioa of that part of the vasomotor centre nhich nmin-
tatap aoder ordinary circunis(anc«a a greater or less tonic constriction of the
«alWMUUt arteried ; how far (hiit may \m lutointc^l by the tt)iec!a] acliou of
T«KMliUtor (tbivs we do not know.
Tbii widening of the ciitane<>U!> ar(cri«> dimiuishe* larjccly t)i<; jieriphend
raiaUDce:, and do (cmU (o lower (he blIllld-p^?0llr<^ Aforeuver, with Mich
«flbrt of each tkclclal niiut'lc thr minute artvri<« of tluit iiiu»cle ure dilnt<.-d,
•u thai during pxcrrise, and (fpccially during vigorous oierciac calling into
action nnnv slieleuil musclvs, there mtmi be in the body at large a very con-
aUifabk widening of the minute nrleries distributed to the various miucln,
aad fa eonafqncnoo a very considerable diminution of the peripheral resist-
aac*, TImw (wo dtrninuiions of peripheral resistance. cu(aneoUB and mus-
cvtar. vuuld tend to lower the blood-prenure — a result which would be most
iojnrinut, since the increased metabolism of the uusclee demands a more
rapid rircnUlion in or<ler to geit rid of the produfiU of metabolism, ajxl fur a
lapid circulation n bij;h blood-pressure is in RHiat cases ttMCenrv, and in all
tasea atlvaniageous. The evil is, in part at all events, met by Lne increased
brrr and frequency of the heart's beaii^ lor, as we have aaid again and
acatR. the mean IiIchmI -pressure iii the product of the licart-beat working
afaiiMt the peripheral resistance, and mar remain eon.tluut when one ftwtor
b inoaaMd or diminisbed, provided that the other factor be propoitionatelj
Jbafaikbed or increased. It is poanble, th<-ii, that tht.- mere increaae ia the
heart's Iteat^ are. during eien-tw, iiuDiHeul to ntiutndiM: the diminution of
pcripbenil resistance, or even (o niiiM> (he blood-prtwun; above the normal :
aad, indeed, we Aod, as a matter of <iic(, that during exercise there t* >ueb
an iacnase «f tlie mean bloixl-prrasurc. Hut il is more than jinjluiblc that
■Uteb valuable labor of (he heart it ccDnonii»-d by itetKrnlixinff ibe imminent
Ml of blood- pmmi re in luinther manner. Il would appear that while thai
Crt of the vaMimotor centre whicJi governs the cntanaous vascular area i>
tog inhibited, that jiart which governs the abdominal splanchnic anta ii,
298
THK VASCULAR MKCHANISM.
on tlie contrnry. Ik-Idj; uiiginunt«<]. Atitl in thin WAV ii Joiiblo «n<1 i* frnioed.
On ihtr cini- hnnd. thu mean Mood-Dnviure u ouiiiiuinml or incn-tuu-tl la a
Taorc ccunomioil ninimpr ihnn by ini.'ri.-«i>ing ihr b«Tl-bvnli>, anil, on tb»
uthiT hand, tlic blowl during tin- cxi-rciw i* iiirncd Rirar from ihr digtwlire
Orgnnv, vihich iil the time nrc, or "Ughl lo be. at r«»t, niia Uiercfure iV|uiriDg
COiDpiinvtivol)- little blood. Tlicee orgnn* riTtitinly, ill all events, niigbt not
during cx«n;iBe to be eogn^pd in the (n«k "f digmting am) absorbing (bod,
and tne nld saying. "af\er ilioner th awhile." may serve ns an illnstmtion of
the wurliiug of ID« Tascular mechatiiaiu with which we are dealing. The
duty which some of the digestive organs have to carry out in the vray of
excretion of nielabulic waal«! products is during exercise probably taken on
by the flushed and uere|>iriiig skin. It is true tbai a( the beginning of s
period of exercise, before the tkiu, so to speak, baa settled down to its work,
an increased Ron of urine, dependent on or accompanied bv an incrttased 6ow
of blood through the kidney, may make its appearance ; but in this case, aa
rre jihall see IhIit on in dealing with Uie kidney, the flow of blood (brougli
the kidney nnty he. incrvujed in spite ol cijUBtnetlon of the mt of tbe
splanchnic area, ftad, bc«idcii, Mich an initial incrtnse of urine wpeedily givea
vray to a decmwe.
i 194. The eflect of fnod on the vowidnr nKvhnniKin nflbnlii a market) con>
Inwi lo the effect of hoiiily lnb«r. Tlir mo»t marked rwult i* n widening of
the whole nbdominnl »platichnie nrtn, accompanied by to much coo«lriction
of the cuiAncoiis vn^culHr nren. nnd ho much increase of the heart's l>eat, a*
isBUlGcient to neutrnlize ihc lendenev of the nidcnin^of the splanchnic an*
to lower the meao pressure, or perliaps even sufficient to raise slightly the
utean prewure.
Any large widening of the citlaneous area, especially if accompanied by
muBcuUr labor and the incident widening of tne arleriee of the muscloo.
ntmld lend *o to lower thi; general bloud-prewure i unlew met bv a wasteful
use of cardiac energy) »» iiijuriously tn lamm the flow througli I he act ire
diseating vijtceni. A moderate conntrictlon of the cutaneous vtMiels, on the
Olner band, by throwing more bloixl on the ahiluminal >j>lanchnic area with-
out tiiiking toe heart, is favorable In dtgt«lion. and is prulmbly tlie jihytio-
bigicn! cxpUiutUon of the old saying, " If you cat till you're cold, you'l)
live lo be old."
In fact, during life there sccma lo bo a continual give-and-take iN-'twun
tlio bloodvewcle of the somatic and thono of llic xplanchnic diviHJoRS of the
body : lo fill the one. the other i» prop)rti»i lately emptied, and vice Jtrtn.
i 195. We have seen (^ 174 i that ecrtnin aHervat nitrva of the vngus form*
iug in the rabbit a separate nerve, the deprrasor nerve, air osMciatod with
tite vasu-coustrictor nerves and the vasoiuolor centre in such a wrsv that
impulses tiassiug ceutripetally along them from the heart lower Ihe blood-
pmAure by {limiuishing the peripheral r««istiuice. probablv inhibiting tli«
tonic constrictor Intluencea exerted along the abdominal splanchnic nerves,
and io, as it were, opeaiag the splanchnic Hood-gales. ^Ve do not jpoaaeaa
much exact information about the use of these aSerent deprewor fibres in
tltc living t>n()y, l>ut proluibly when the heart is laboring against the blood*
preasurc whii'li Is t<>o high tor its powere, the condilbii ot the heart Aorta
imnulsrs which, piiMliig alon^ the deprcMtor fihrfs up to the medulla
oblongata, temper down, so to s}i«tk, the blood-prenuro to suit tlic cardiac
strength.
Wo have, moreover, reason lo think that not only dom the hejirl thus
regulate the blood-preMur« by means of the depressor fibr«. hut also that
ibe blood-preesure. acting, nt it were, in the reverse direction, rrgula(i.> the
bean-beat; a too high prtiwure, by acting directly on tJie cardio-inliibitory
SOMK FKATl'ftKS OF CIRCULATION.
399
ectttra to tbe tnedulla oblougata (either direcllv — thai i», a* llie rasult of the
tvcnku cotitliuou or ih« toMiiUa iloelf— or inilirectl}', by imitulMS rencbiiig
ikit tD<«lulla mluHK ufl'ercDt nirves fmni vnrinut nnrtit uf the biiHy) aiay
mm) iohibilnrv iu>p4iltn> down tltc vngiix, ninl wt nhicki-ii <>r tone <luwii tliv
hran-bmti.
1b tb« riillowiiig wctions of ih'u work w^ shall mm; rcpvntt^ in»tMiw*,
railar til or rvcti more Mtriking tb*n tbo tibovc, of the iDBnnpt^nit-ot of tho
ra«cuUr ni««liniit«m by nxmns of ibo ncrvouB tyflem, and wv, ilicrcfui^, n«'<-d
4vtU BO Ir>ng4rr on the siibjccl.
Ve amy sini|tlr ropent cbnt at the centr« lie* the cardiac mtuciilar fibre,
nd al tite periphery the plain niusciilar fibr« of the lainute artcrr. On
Ant two p)eni«nli the central nervous By»l«m, directed by this or tbst
■Bpolw nachin^ it alonf* adereot nerve librw. or nflected directly by this or
Ikal iiillacoM, » dnrin;; life continually playing, now augmeDtin^, now
iakAkiBC, now tbe one, now the other, and so. bv help of ine elasticity of
ibruUneaand tlie luechaniau of tbe valves, direciiug tne blood*flow accord-
iif ID Uie needa of ibe body.
BOOK II.
THE TKSL'ES OF CHEMICAL ACTION, WITH THEIR RESPECTIVE
MBCHAKISMS. NUTRnTON.
CHAPTER I.
THE TISSfES AND 31E0HANISMS OK ULGEHTION.
f IM. The food in pswiofc alon^ tbc aliin«Dtary canal is Bubjeclcd lo ilic
I of certain juicM supplied by the secretory activiiy of the epilhdiuai
irhfck liue the o«nal itself or wliich form part of its glaiidiilnr appuii-
^ja. Tbew juices, vii., saliva, iputtrJc juice, bile, pancreatic juio.', and
ih* mcmImm of tl»eiruall and large inttstiuea, poured upon and minding
»ith tbt tooi produce in il sucb chaujtnt that, fmiu bein^ largely imwifiilile,
it liiiLiiiiiM Iwgdy Mluble, or otheniviiie niudify it in •iicti a way that the
iMftf put v( what IK ealeo paaMt into the blood, cither dini^ily by lacana
«f tbo eapillanra of the alimentary caual. or bidirei-tly by invAm uf th«
IftCUftl ttjWeta, while tlic :<ma]l<-r part u diycharged as i-xorimU'iil.
Hmmc pU\B of the food irhiuh are thus digi-slcd, ab»c>rbed, and made use
at by the body, are «poken of as j'ood-»tuff* (they hare also been called
mlummUuy prinetpU»i, aod may be convonjemtly divided into four ureal
1. fntrittt. We hare pr«vioualy (5 15) ^ken of the chief cliaraclen of
tbi* rlwa. and hare dealt with leveral memben in treatioK of bloiid and
■MM^. We uav twre repeal that in general oan]|)C6ition ihey contain in
100 part* bv weight "in round numbers" rather more than 15 part* of
■Itngtn, ratber more tliau SO part<4 of carbon, aboul 7 parta uf hvdrugen,
aad ntbtr mnre than 20 parta uf oxyj^en ; though eeMotially tlie nitro-
gtoooi taxlica of food an<l of the bi>dy. tbey are made up of carbon to llie
yrttn* of more than half their weight.
TIm nitroocnous bwly ijflalin, whicb otN'iin largely in animal fmxl, and
•OOM albtr Dodic* of Uv imjxirtauce, while more cliMcly alliixl In jiroteid
bodla tbaa to ooy other claw of onauic aiiliKianixs, ditfer considerably from
pMKfla in comporition and cq>emlly in their W Imviur in the body : they
am Mt nf lofRciant im|>-frtnno(- to form a clnx* by tbcniwlvi,'?<.
2. flat*. Avquently but crrontim-ly calUii hydrocurh'inx. Thi>c vary wry
wld<^ly in chemical compMition. ranging from >uch a comjiarativcly »imptu
fat a* liulyriu ta the highly complex Incithin I^Tl); they all poawn, in
virw of tbe oxidation ot both tboir carbon and their hydmgen, a Uf:ge
iBt of iMtential energy.
(.itr^k)dral€i, or lugare and ftarchn. These ponces, wdght for
rdativety leas potential energy than do (iit«: they already contain
Melrea a large an>ouiit of oonbined oxygen, ami wbcu completely
WlMbtd ^TS out, wei^t for weight, lees heat than do fat*.
302 TKL TISSUES AND Ub'CHAX]
■I. Saline or miiitnl h&dui, and tuatcr. Tbeae Balu ore for ibe most part
luorKaiiic huIib. Hnd this cluits dill^n Arom the three [•receding cImhs in lu
iiiDcli &* itif iiTc^riiliieM of ita memben to llie body lieia not so much in the
iitiiouut i)f eiiLT)cv wbich may be given out by ih«ir oxiduioii lu iu the
rnri»ii« iiiAuenoes irbioli, by their pmencv, cliey exerciw on Uic metabolic
CvrnU of tlic liudj.
Thaw (wvcml f»iid-Hiuffi» are vunouitly uctud uprni in the Mvcr&l parts of
the alinicnlHry cunul, and we mny dlMtiiiguiidi. ai^ tht' fixid yeuMt along th«
digMtivi- tracl, three inuiii singea: dizestton in the mouth and tlonuich,
digestion in ibe unmil iute»Une, and dtgcvtion in tlte large intcsliiM. In
many animnU thi7 lin>l singe is. lu a Inrge extent, prcparolory only to tbc
second, nhich in nil the aiiimnlx w the ctnge in which the food iindervoes
lli€ ^'rentcsl change; in the third stage the ohangea begun in tbc preruMU
siugee are ci>m|ileted, and l)ii» »tagc is evnecinlly characterized by the nbeorp-
lion rif tliiiil from the interior of the alimentary canal.
It will be convenient to study tbeee stages more or less apart, though no4
wholly so, and it uill also be convenient to consider the whole enbjecl of
digesiioti under the following heads :
Firsi, the characters and properties of the various juioea and the clian^
which they bring about in the food eat«n.
Secondly, the nature of the proocaaeo by meam of which the epithelium
i:t]\A of the various glands and various trncta of the canal are nble to niauu-
finctiire so many various jiiicea out uf the common wurve, Uie blood, and the
manner in tvhioh the secret<iry activity of the cells ia n^lated and nibjecled
to the needs of the economy.
Thinlly. the niechiiubiu», here us elsewhere, chieHy of a muscular nature,
by which the fooil is pa.'wed along the <!anal and moat efficaeiitly brought
into contact witli thi- wvprnl juices.
Fourthly and lastly, the nienns hy which the nutritious dimtcd malArial
U separated from the undigested or excroineiital inatcrini and absorbed into
the blood.
The CHABACTEttS kVVi PBOf-BRTIES OF SaUVA AMD GaBTBIC JuICB.
iSb/i'vu.
S 197. Mixed mlira, as it appears ui llie moutb, is a thick, giniry, gvn-
ernllv frothy mid turbid fluid. Under the micrnfcopc it is fteen to contain,
licsidcs the niulf'Ciiinr d^brin uf fuod, )>acti>ria, and other organisms (fre-
■uently cnr'pt'ininiic i>iH>n.-«), eji it helium M-'ulen, muc(iu»KMrpuscli?, and
rgraoukts, and tin- "ocalUH] salivary corpiuM-lest. Its n-nctton in a healthy
subject it nikniino, specially uhwi the wcretion in rbiimlnnt. When the
saliva is scanty, or when the mibject xuflcni from dynncp^ia, the reaction of
the mouth may be ncid. Halira contains but little solid matter, on an aver-
age probablr about •'■ p«T cent., the s[>ecilic grnvily vnrying Irom 1O02 to
IfMIS. Of iKese solids, rather ln« ihnn hall, nbout 'l per cent, are salts
(including at times n minute ijuanttty of potasAium sulphocyanatei. The
organic bodies which can Iw recognised in it are globulin and serum albumin
(see ^^ m, 17), found in sniaii i^uaniilies only— other obscure bodies occur-
ring in minute (pianlity, and mucin ; the lotter is bv far the most conspicu-
ous organic constituent, the glairinces or ropiness o^ mixed and other kinds
of saliva being due to its oreseoce.
Jfim'ii. If acetic acid be cautiously added to mixed saliva, the viscidity
of tiie saliva is increased, and on further addition of the acid a semi-o{ia4ue
SALIVA AND OABTIttO tVlCB.
of the «ativa liiii|iic). Ttiil ropy
niili II ):liu» i\mI, utiriiikf, hcoominff
o|i«r|o«, i-liop %•• (lie ^liiw t>Hl. iiuil iiiny be lliutt nrmnriil rn)iii iho flutil. If
■MAiaiM QUI, Iwvtng ihc nvt
wbieb u Biticiii. ifuirrvd onn-l'iillv
^
th« '(UUilJty of Hindu K* ■mali and the mIivm bv violnilly «liiikcn or *tirml
whtiv ibe ■<;»il iv Mn^ ■•lik-<). ibr niuriit ■* npl to In- prrcipitacil in HnkM,
aiu) Bwy thra b« w-iwrittvl br tiUrntion. It may be fi<lilcd thai tbc |>nxtipi-
tettoB of miiciii by Hci<l is ^raaily ioflu«noed by tbc prracncc of otidiiim
cbloride uid other mIih ; thus, ifUtr tbu uddition of sodium chloride mc«lM;
mtrid. even in coD*idernblo exenw, will not cuuso a ]>reci|>iljile of mucin.
Mucin, thus preporeil utd piirifi«l by wiwhing with aeeiic ncid. Biralk ooi
in water without actually dissoUini; ; it trill, bowi-vor. dissolve into a viscid
fluid readily iu dilute (l>.l per cent.) solutions of {lolaniuni hydrate, more
slowly ill tolulions of alkaline salts. In order to l31ler n mucin solution,
grrai ililution nilb nsier is DeceeearT.
.Muciii it pre4.-i|iiiaied by siroDf; alcobul and by various metallic sails; it
nwy aha be precipitated by dilute miiisnU acids, but the precipitate It tlien
•olabla in exeeas of (be acid.
Macin KiTea the tbree proteid raaolions roentianed in $ 15, but it iii u vrrv
mtDfdex body, ntore coni|>lex even tlian proleidii. for by treuiment witli
flilutv mineral acidti and in other vays, it miiy be coiivertet) into tumx fonn
<if prutvid ( acid -albu milt when dilute minenil acid i» u^), while nt the
•an* tin>c then is formed a body which apjtmni to Iw carbohydrate aod
rHMobto a sugar in harinj; the power of r^ducinii; cupnc sulphate soliitioiw.
Salatiou of mucin, moreover, ou mere keepiiiK are ant l» lose tlielr viw^diiy
umI to becoroe converted into a proieid imt unlike the body peptone, whicn,
•a «« •liall me. is llie result of gantHc di){C«iiton, and into n r»lucin)( lH>dy.
iMTvral kinds of mucin appear to exist in variouM animal bolin, but (hoy
HNB all to Agree in the ennraoler that tbcy can by appropHnle treatment
be split up into a proteid of some kind and into a carbohydrate or allied
body.
il0t. Tbo chief poipoM Mrved by the saliva in dilation is to moiateu and
soAui ibe Ibod and to aansi in mastication and def-lutitioo. In some animala
Utia is its only ftindioD. In other animals and in man it lios a B[ieciiic milveait
■etsoa itiiMiDeof the foo<l'«tut&. Such minerals as are suluble iu slightly alka-
Ub0 flaids are dissolved by il. On fata it lias no dEtct save thai of produeini;
a rwTf Aeble emulsion. On proteida it lias aba no speciHc action, (houftb
I will M of meal, cooked or uneooked, apfwar greatly altered al\er iliev biive
DMB ma)4lcateil fur M>me time; the chief atteratioD, liowevvr. whicli thus
tskaa niare is a cbunp> iu the hjeuto^'lobiii and a genernl wifleiiiuK of tbo
Huaealar fibros by aid of Ibe alkiiUniiv of ilte saliva. Of oounc, wh«D par-
tidaa of fond ar« retaiued Ibr a ifmg time in the mouth, as in the intonbcas
or is cavities of tV teeth, the bacturia or other organtsm* which are always
preaent in (he luuiitli may pnxluce niucli miirc profouiKl chaiigm. but lli«se
ar* Dot (be li-yitini»ii> priiducls of tite action of saliva. Tbe charaoteristio
pruuerty of saliva i* that of coovcrtiiig starch into aocns form of sugar.
AH'on of taiira mi tlarrfi. ]f tn a quantity of boilvd starch, which fs
aliraya more or lew viscid and somewhat opaque or turbid, a small luniility
of luiva be added, it will be IbuiMJ after a short time that an important
cJmuhK has taken pliicr. inannuch as tbe mixture has lost its previous
viacMUtT and become thinner and more transparent. In order lo umler-
tfand tais chanj^ the render must bear in minu the esiftteoce of the follow-
in bodies, all belonging to the claH of carb>jhydratee:
1. SIoitA, which ibrnu with water not a true solution but a more or Icfli
viscid mixture, and gives a characteristic blue color with JMlise. Tbe for-
aala is CfH^Ot, or more correctly (C^U^OJ' since the molecule of starcb
SOi TUK TIS8UI8 AND MECHANISMS OF UIOBSTION.
i« some multiple (n being not lose than 6) of the aimpler formula. A kind
of March known aBMi»Af<r«Mi-cA.irhile giviD};kbIu« color nit )i iodine, fornia,
unlike ordinary etarch. n clenr Boliilioa.
3. Dexirins. diflerinf; from slitrcU iu forniiu]^ u clear eolutioa. Of these
Ihnv are al least two ; one eryt/irodextrln, otten xpokeD of simpl}' as dextrin,
giving a port-wine red color with iodine, and second, aehrooilfrlrin. whk'h
pves no color at all with iodine. The formula for dextria ia the Mm« u
Ibu for starch, but has a smaller molecule and might be reureaeuted by
(C.H.,0.)-.
:t. IJtxlTon-, aliM> called glucose or gra[ie-Mgar, f^ivin^ uo coloration witli
iodine, )>ui vharacteriiEed bj- the yavivr ofreducinj; t-uprio utid other metallic
Mlt« ; thus, when dextrose a boiled with a tluiil known an Fehlin^V fluid,
which i* a smlution uf hydrated cupric osidc \a an escoM of caustic alkali
and double lartmie of nodltini and potaaviiiui, the cuprio oxide i« reducMl
and a red or yellow de[Miiiit of cupronw oxide \* thrown down. Thiii miction
»ervnt with olhen na a convenient l«»t for dextrow. Neither riarvh nor that
(■omm(in<-«l furra of iiuf!;nr known ai* cnnc-HUgar, give the mclinn; wlH'ther
the ilvxtrinti dn U doubtful. The formula for dexlnwc Is C,H„0, : it is more
niuiplc tliiin thutof rtarch or dvxtrin and conlnini an addition H,0 for every
C,. Cnlike Klarch iind dextrin it can be obtained in n cryttsJline form,
either frum n<]ue(tui> iJiliitions (it being rcadilv ttoluble in watori. in which
case the eryHlals contain water of crvetal ligation, or from its solutions in
alcohol (iu which it n sparingly fi.<lubfe), in which ca/n tlie crrstals hare no
iralerof cryetalliitBlion. Solntions of dextrose have a marked liex I ro- rotatory
jMwer over rays of li^hl.
4. Mallote, very similar to dexlr>>«c, and like it capable of reducinf; cunrie
salts. The formulu ia somewhat ditl'ereni. being C,,H„0,|, Besides this, tt
difl^rv from dextrose chielly in its smaller power—/, t., a tn^'eo weijihl will
not oonvvrt w> much cupnc oxide iutu ciipriiUH oxide as will the name ncieht
of dcxtriHtv, and in havini; a ntronger mlatory ai'tlou on rays tif light. I.iks
deztroMi it can Ik crvntallixed, the eryitals from ui{ueou» itolutioiM conlainiog
water of cryxtaltiuition.
Now. when a ijuantitv of Hlarcli is boileil with water we may recogoiu in
the viscid imperfect ikilutioD. on the one hand, the prmenoc of the *<«rch
by the blue color which the iidditlon of iodine give* rinr to ; and on the other
hand, the abecnco of sugar (niBltosc. doxtnm). by the fact that when boiled
with Fehling's fluid no reduction takes place and no cuprous oxide is pre-
cipitated.
If, however, the boiled starch hv submitted fur a while to the ucliou of
saliva, especially at a eomewhat high temiiemture such as '.f't" or 4*1° (.'.. it
ia found that the subsequent addition of iodine j^iva no bkie color nl all, or
very much less color, showing; thiit the starch has disappeAr«l or dimiuished ;
on the oilier hand, tbe mixtun- readily gives 8 precipitate of cuprous oxide
when boiled with Kehlin^'s fluid, allowing tliat maltose or dextrose ia present.
That is to Nty, thi; mVivu has converted the «Ukrcb into maltose or dextrae.
The prewncc of the iirDviuimly ab^ut Mijfar may also be shown by fermenta-
tion and by the other tcstM for sugar. Moreover, if an adetjuntely larg«
ijuantity of rtarch he Mibjected to the char^, the sugar formed may b« uo-
loicd, and tts cluirat-iero deteruitnetl. Wheu this is done it is found that
while tKimc dextrnic Is formeil the greater |iart of tlie sugar which appear* is
in the form of mattow. As i» well known, starch auir, by the action of dilute
acid, be converte<l into dextrin, and by further action into HUgar; btit ibo
auptr thus formtil i« ulnar* wbnily ilextrtwe, and not maltose at all, Tbo
action of raliin in lhi# res[iect diHerw from the action of dilute ncid.
While the conversion of tliv starch by tli« saliva it going on the addition
I
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SALIVA AKl) nASTRIC Jl'IOR.
SOS
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mT KKliiM> fm|iiFDtU give* rise lo a red or vinlel color iustead of u pure blu«,
but when \ht- tonvenioa b compile iio oiitanUion bi all \» oluervciil. 'Die
>)i[inirM)i-« i>{ this red color indJcaUa llie prak'Hi-e i.f ilextriii (uryihrock-x-
Xiin) ; the rMel color b due lu the r«d beini; mixed wilb the blue of still
oaeliMi|{«d Btarcb.
Tbp «ufieannee or dextrin ahowsilial the aotioiiof the saliva oii the Mtarch
is M>iiienbat oum|ilex ; ami tht* h Hiill runlier prutrcd hv tbi; fai-t tliat even
wbca ilie mUIvk has eoRipIeiea) its work ibe whole uf ihe #tnrrh doc« not
nsppCBr m malhac or H&itr(«e. A ooiuiderable (|UuDti(y of Ibe uther dei*
Uin («bn)od«xinD) alwiivx ai>p«-im nii'l rfriitiinn unchanged to the end;
•Dd tb«rs are pnihalily mv.thI iitbL-r hmlies also rnrmtd out of tlit> Mlirob,
tbenlatim pN)>>rti<>ii» van-iiiK ucv-inliu^' to cirL-umRUmcM. The ohango,
thfrfdW, thiM]{li )icrliu[i* lie may :<(Huik nt' it in a gencnil way lu ons of
hjrdratiiHt. onuot b* •ihibilrd undfr H :<iniplf Ibrmula, aud vo may rv«t
«aaiait for lb* prtMnt with the Plalvnieiit lliiit Mnrch Mhen fuhjccted io tbe
Wlion of aaliva is roDvertrd nbicHy into the Mignr klxiwn as mallow kiiIi a
coQiparaiivelr snisil •^uaiiliiy uf dt-x(T\«c and lo some exl4.iil iiiio nchroo-
dntriD 1 «r\'thnH)exinii apjicantie tempontnly onlr in ihe prooes'), other
bodica CM w'hioh we iiee<l not dwell kcinc formed al the same time.
Kam iiiibuik-d »Mrch undertow n similar chants but at a much glower
nuv. Tbts is due to tho hu-t that in iIk- curiouslr formed elarch guraiu tbe
Mm Staivb. or ffnuuJtft, is invested nith coals of rrHuloe^, Thi^ latter
aaaloria), which requites previous Imatment with sulphuric aeid before it will
R'<M ifav blue rrwiioR on the addition of iodine, is apparently not acted upi>n
' isliva. Hence the saliva can only j^t at tbe Kraiiul<«e by iraverdin;; tbeJ
euatsof (fllidivie. and the conversion of the formi^r lit thereby initi'h hlndere^
BIMJ delajred. m
S IM. The cunveraiuii of starch into Mifiiir, and this we mar speak of Ml
Um anrlulytic adi^m nt niUvh, will ^ on at tin- unliiinry lfiii|i<'ralurc of ih«
aUnorpneTe. The lower ihir icmperalure ihi- kIowit the chan^*, and at alHHit
U' C. ibe iV'nverwkiti i* indetinilL-Iy pn>l»i))t:ed. AAcr exfHniorc to ihix culd
fur V¥«u a ooDaiderubk* time the action rvciimroetKi^ when the tempcratiira
is again raiaed. Increase of ten)|>i;ntture up lo almut 3>'>**-4(>°, or even a
littlii lit){ber, Avon the change, tlw gresiU.wt actirity being «id to be mani-
ImUd at about 40\ Much beyond this {luiitt. however, increase of tempera-
lurv becnnwa injurtou*. ninrkcdly so at 60" or TU": and saliva nhicb has
barn boiled Tor a l<-w minutm not niih- hits no netion on »tarch while at that
UmpenUurp. IhiI iI'nw not rc^in its puwenon cooling. By being hailed,
lb* amylolytic activity of saliva i* permameotly deetroyed.
Tfav action of saliva on starch is mi>>t rapid whcfi the reaction of the mix-
tun is ncutnl or nearlv so; it is hindered or arrcMeit by a disliuclly acid
futioa. Indeed, the preseoce of even a very small <piaDtity of free acid,
U all erents of hydniCDloric acid, at the temperature of the body not only
•MpMids tlw action but speedily leads to penuanent abolition of tite activity
«r ibi JQim. Tbe bearing of thiit will be seen later on.
Tlie action of saliva is hamperol by the preaenoe in a concentrated Mat6
of tbv pmluct of its owu Kctioo^-ihat is, of MKnr. If a small (|iianiiir of
s*liva tie added to a thick man of boiled Ktarcb, the action will alter a wliile
•lacken.aud eventually ouroe to aliu»!ii a Bt«ii<Lsiill long before all lite starch
hw beeo convened. On diltiliuK the mixture with water, the action will
RMMiiineDoe. If Ihe pmduots ■>f action lie removed tu lOon aa they are
Ihmed, dialysis for pxatn|ile, a mubII ipiantilr of saliva will, if .-lufliuienl time
be allowed, convert tnio uitrnr a verv lnrK«, one might aliixnt miy an indeR-
Dilc. i|uatitity of stamb. Whether iW particular cou'litunnt on which the
S06 TBE TlSSl'BS AXI> UKUUAMISMli OP OIOBSTIOX.
iictivity »f uitiva (lv{>eHi!it a nt all oouauMed iii ita acllon luu not at pmeot
bK'it ilditiitcly iH*tiIt-il,
On wimt runnrittioiilH iln the iiiiivlolytic- vinuo»of nnlivii ilepeixl?
If Hilivu, lilti-rcii mill iliu^ (rw) fiiiiii timob uf iu mucin ami fmu o(li«r
formci] c(>ii»tilueuM, be ticiitcil uiili irii or liAevu lirni.-* iu bulk of alf»Ut>l,
It prcei])itiilc w forintKl I'lmiftiiiini^' Ih-ki'Ii-ii "liivr tiilMtarioi-H ull tlio [inalfii
tntittor!. L')>on »l-(iiii)iiig iiixlf r tlic ntciilml for mmur. linn- i M-vcriil duyn), Uk
protciiis thus prccipilsU'd Iwc'iiuc i-'Higiituii-'l nini iii«»liib!c in wnti-r. Hcut-*,!
an aqueou§ cxirurl of tho pnvi|HtnU', mmb- nllor iJiis intfrviil. oi>nliiiiui vrry
Utile proteid DiiiCcrial ; yd it is cxcccibn^ly nclivo. Mnix-ovrr, by oiIkt
more elabornte iii€lhuil» tWn nitiy be ubtainvd fmrn mlivn mlulioiis wliich
appear to be almost onlirvly firv from pmidds and yet an> iiitcnsi'lv amylo-
iytic. But even tbcw probably CDiitain other bodici* beeide* tlie nallr active
CDoetituent. Whatever the active eubstniice be in itee'f. it exists in #ucb-|
extremely fimall quaatilies that it bos never yet been satisfaL'torily iaiflattJj
uud indeed the only clear evidence we bav« of ita exiHtence is ibe manifeaia-^
lion of its |>eculinr jKiwere,
The ialieni fenlureaof this body, ibis aniylolytic ageut, wliieh vn may call
jilyniin. are then ; let, ita presence iu minute ami alm<:iit inappredable •|Uiui-
tity. 2d. the close dependence of its aeiivJIy on lcin|terature. Sd. ita pernia*^
n«nl and total destruetiun by a liit;h K^uiperature and by various clteiuica'
reafceiitit. -1th, the wanl of any clear pr<>i>f that it iioelf undergoee an]
cbaiifce iluriiiji the omnireetation of ittt ponen — tbal w to ?ay, theener
neCMMiry for the iraui^forniatiuu which it eflecta i/om not ramir out of
if it iit at ull uoed up in iit> action, the low t» mtlier that <>f nimple vrtar i
tear of a ninrlune thiiu tbal of a «iib«lanoe exmodcd to du ivork. 5tli, tfaa
action ubiih it inducM in probably of tucb a Iciiid (Kpliltitig ui> of a molfr
ail«n'ith iimuiuption of wnier) ns is elTcclcd by thai particular cuiM of agmU
calkd " byilrolytic."
These features mark out the amylolytit^ active body of saliva n« belongin
to the clasB of ffnamt^ :' and we tnoy henceforward speak of the amylolytie"
ferment of NilivD. The tibrin-femient (^ 'M) is so called because its action in
manr ways reeeniblcs that of the ferment of which we are now speakiof;,
§ «00. Mixed nliva. whose properties we have just diseuned, is the result
of the mingling in various proportions of saliva from tbe jmrotid, submax-
illary, and the subliiifrual j^laDos with tbe secreiiou from the buccal Klaad&
Tlie«c constituent juices have their own tjjieoial character*, anil thee© are iMt
tbe same in all animals. Moreover, in the oaine individual tbe aecretioo
diR^rs in composition and propertiea accurdiug to circiimHtaiicu>; tbiia.aswe
ahall see in detail hereafter, the nliva from the subiuaxillary (;l<uid aecTMcd
nnder the inlluence of the chorda lympanj nerve i* <litfen!Ut from that which
b obtained fruui the same ^land by ^tiuiiilatin^ tlic syjii])ulhetic nervK.
la man piire parotid saliva aiaj- easily be oliiiiiiied by introducing a fine canala
Into tbe o^ioiiing of the Stcnnnmn dud. sud -iil>cm>xillAry saliva, or rather ~
mixture of luhmaxillary and lubliiiKUiii MiliTH, by •iinilsr citheterixation of (ti
WhnitoDian duct. In aaimatii the duct uiay be dimtucinl out iinil a canula iodo-
duced.
I Parauoumajr. fcf 'be j.mcuIM loul. bt divldwl liil«t*D olMWi. oonunun Ir (aQart orpaalial
■od wMraoManf. I>i ibo funiwr, xca>i ia»} b* wkni «■■ trpll'kiiiwTn MsmpiB. The IbrmeaiMttv
MaTtljroTTMutirMohlMdsiallipfloiivmlnnof lawlnio alcohal. Ii derauileiii • >ri |^F ll(ci>rilM
joM-all. I'nIvB itie jriaatorll be Urlna and raDitU^ual. ftrniaiituliin iltm iwi inkt iJiirv -. mhta
tbii TOUI-cttll iU*> IbruiraUtlDn p«*m : aiul iici tiilMancp obWIniHl Irnrn Itwilulit r^rucil laM. bf
piMluMloa Htlh ■Iniluil or otliucwli*. •■■II th<v rtw tn altMhnUe (irniicTiMllaii. The atirniry fer-
meat Mow to tbi- Inl(«rrlu«: ti l>o nilnlinm. nul aUiliie uriaiuiiii II Ve >tmn. U iiiay Iv sJtItil.
Ii0«**tr. Ilwt iJO^IiiT (lie iimnlin] rcnucni. tbu ittrt fit Tiiiuuim. [initintB it> vilvct lir nxwni at
■a grttaMf nuutiaalSBd KfMtnt wlilcli II swivnieK but wlilcb li laiinullftiely MiBiteawBT Willi.
BALIV.t AND flASTRIC JDICR.
807
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IHuutiil lalivft in niito in cimr anil liropM. not vucii) ; lli« rvRition of llie
fifM drops (vctrtetl in oAcn acid, the sncerriltng portiuiu, at nil eV«nl« when
tlw Sum ia at all copious, arc alknltae; that ia to ay, tbp iiMura) wcn:t)<in
h alkaltae, bot thin niiiT W ubectirvd by ncid cimngc* taking plnco iii ihfl
laitl wliich hflB b««n rciain^H in tlie iluct, pof«iblv by the tnniiatioa of an
«ScnB of csrbouic acid. On »lnnding the clear tliiid becotitcti tiirhid from u
pnapltate of t-alcic carbunalc, due to sn escape of carl>oiii<- ni-i<l. It ciiu-
taiiu clobalin aiid some other foruia of albuniiu, with little or do mucin.
I'iXjuuuu Milphocyauale may aitio soiuetiiiie^ be delected, but stnicUiml
■Amtota are aoaeni'.
Babnaxilbu'y tsHva, in nmn and in mort aoimals. difl^re from parotid
mUt* in beinr more alkaline, and from the preH'Oce of mucin more viscid ;
it flMtalB* Mllvary curpuscltv. tliat is ImmIics cIimcIv resembling if not iden-
tits] wilii Uueocvii-H. and oAeu iu abundance uumrphouH roaaaea. The K»-
calM eborda mlWa in the dog, tliut b to Miy wHva obtained by atiniulatiog
ibe cborda tym[i«ini nerve (of ahich ne *liall preeeiitly fti)ettlci, it uiidvr
ontlMuy circuiu«ianc(« thinner and Uvn viM:id. cvntaiua tew niucin and
tovar Mruciural element* ihaii lite (u-calle>l tiytn pathetic •&]]», which ia
nMrltmbU) for it* viacidily, its •trvcuiral elemtjita, and for It* larger total
oTWiib.
Sabtingual Mliva !• mor« vl»cid nnA oontalna more salts (in the dog about
1 DV nmiL) than the »iihniaxil!nry i>itlivn.
The action »f milivii viiri<« iu inlci>»ity in diflbrent auimnlti. Thu« in
nan. tke pig. the piim'W'pie, and the rat. Ixitli parotid and lubnuixillary
and tnixvil MilivB arc nniyrolyiic: the tuhninxillary Nilivn hdn]; in moit
tmm tDore active itinn the pnrolid. In the rabbit, whik- the siibmnxillvy
aalira baa scarcely any action, that of the parotid is cticrgctic. The snitva
ol th* eat b much lew active than the above ; thai of the dog is Mill )«iw
acfive, indce*! is alnioei ineit. In the hor»e, shecii. and ox. the ani^lolyiic
jnrnm <if either mixed naliva or of aoy one of the constituent juion arc
titnntlj feeble.
WWre the wliva of any glatul in active, an a<]tieouH inAiaJon of the Mimo
glanil in also adive. The im|>onancc and bearing of this Matrmrnt will be
aiiMi later un. From the ai]uroti» infuciiiu of ibt gland, a« from Mdiva ilwlf,
ihm frnneiit nay bo^pproximately itwiatcd. In norae eaao* at leiut •one
fcrmrat may lie exlrarted from the filand even when the secmion is iuelf
naciive. In fact, a n-ady n>othoil of preparing n highly amylolytic liquid
totnaldy free from protcid and other imparities is tn minec fioely a gland
knunn to liaw an active Kcrction, tnicb, for imrfance, as that of a rat, to
dehydrate it by altouinp it to stand under absolute alcohol fur some dars,
and then, having poured olf moat of ibe alcohol, and removed the remainder
br evaporation at a low temperature to cover the |Hecta of glaod with strong
gijcenii. Though tome ol the ferineni apjienrs to be ooMrtqred by the
alcobal, a mere drop of such a glycerin extract rapidly converts stnrcli
iatoMigar.
0<utrie Juift.
I 901. Tbn« is 00 difficvliy in obtaining what may &irly be eonitideTed
as a normal ralivK; hut tlierv are many oMtaclta In lb« wav of dctermiidjig
tbe normal charactrtv of the H-CnHion of the rtomach. \Vbeii no food u
lakrti the stomach is at rest and no n-crriion take* places When food ie
takes, the eharactera of the gs*<ric juice Mvrcted are obscured by tlw food
vilb which it is mingled. The ^atlric membrane may, it is true, be aril-
ftcially ilimulnled, by tuucb for itutancc, and a sccreliou obtoincd. Thb
6m
TIIK t'lt^SirBS .KSV MKCHAS'ISMS of rtlOKSTIOI
we may ei^eflk nf ii» i^utrk- jnico. but it tnnv Ik- (liuihtnl wlM-thtr it uagh
to be L-i>i)ai(l«rL>d ns imrmat ;;[isiric juice. Ami iiiile^'l. ii> <nt; Elinll wv, ev
Uie juice, wliii.'li is poured IdIo iIic stomach duriog n mcnl, vnncs in i»nii
siliua aa digetitiou U- goiii); ou. Uenoe titp cbaractcru wliich vie slinll git
of juiSBtrio juioe must be L-i>D»iiicr^l as having a j^noral vulue only.
Uaslric juiM, obuined in m iioriiial ii (.Miidittoii as puaaible from the h«tillhy
SKHnaeb uf a lusting dog by uieaiia of u gastric l!etula, ia a thin, almost culor
let* fluid with a iMiur taste aud odur.
Tn tbp operation fur gwtlric RKiiIb aa iacision is iaad« tbroavh th« abdominal ^
wall*, alonic tbe linta alha. the •Ciiiaai^b in opvucd. nnd ihc iip4 of the gaatrli
uiiuud Kvcurely dhwii in tliuau <i( tUu iiiditioa io the alxbuiiiind waIIh. UiiUi
Huoii LaktiH [iIhcc, «o Ibai a ii^rmancni opi-nin); Trvin ilie eiterior into iho liiud* i
Oip ■tomncb is Mlabliihrd. A tiibi.' of proper c'ini>trui-iir>n, inlroducvd ai the limi
of Ihc operation, becumra lirtiily ■ccurvi! in pUci- bv Ibe conlmciicm of bonling.^
liirougb tbe tub? thi^ cuiiLeiiUof the ntuuiNcli can 1i« received, and ibc niuMual
menibfun« sllinulaied at plesHiire.
When obiiiined fniiii a natural fistula in tiiau, il« »|>eeiliR )(ravit,v hat li««u
found to difl^r little I'nmi that of vrai«r, vuryin>: frum 1. 001 bi 1.010. aiid
Ibi! ■moiiMt iif solids pn^cnt tii b« ci)rre«ii<iii(HiiicIy amall. In auiinalii i>ur«
giwtric juice Hcem.H to be eijuativ iMiur lu Molidei, the bij;h«r Gi>liuiuta> wbick
~Ww obvervcR' lutvc ubtained l>eii)g probably due to admixture iritli food.
Of thv Mnlid matters prt«mt about half are innrgiuiic itnltN. t'liicHy nlkiilin^
(snditinO chWidtv, tvitli xmall igunntltiiv uf iihtHphntm. Thr cirtninii- tiia-
terial coM«iM« <>f i)('}Hin, a bndv In Im- dcvcnbei) inimivliiilcly, iniird withj
other snlwtanciv ul uiidetcrnitni.-d nature. Iti a hcnllhy ■t'iniacb gastrir juioa
contains a v«ry amall (|iiiiiility only of mucin, unlcM some subniHsiTlary
Miliva has been swallowed.
Tbe reaction is disiinccly acid, and the acidity is normally due to free
liydfochliiric acid. This is shown by variouii pronfa. ainouj; which we la^y
mention the conclusive fact that the amount uf chlorine prvaent Id ftastrie
juioe )t> mure tbau would atjflice to lurm chlondra with nil the haaca preaeot,
ftiid thut the cxceoB, if regarded as extHtiuK in the form uf hydrcichloric acid,
cmm'sjHiiidH exactly to the <|uauiity of free aci<l pretent. lactic aud butvric
aiifl iXhir undii when present are teoondary t>ro<luctH, ari-iinft either by liK-ir
ratpcdivc ferment Hlioii A fmm articlt^* <>f fu<><l. nr I'rom the ii«comp<Mtioii uf
tlwir alkntinc or ctiber mllH. lu man tbi? amount of free hydrochloric acid
in hfntihv juice may Ih- vtnteil to be about 0.2 |)er cent., but In aom« nniinala
it ii^ i>r<ibiiblr higher.
^ 202. On Marrh gastric juice ban no amylolylii- aclion : nn the ennlniry,
wbrn B<divii i# mixeil with guHric jnice any nmylolylic fcrRwnt which may
bo prrocot in the former i* at onoc prevenled from acting by iho aeiilily of
tbe nl)Xlur<^ Mons)vor in a very xburt tima, »p«cially at the lemporaiiir*
of Ihc IkkIv, the aiiiylolylic lerinent \f dnttroyed by lli« acid, so that eren oa
neutral iiaii<Mi tliv mixliin.- i» unable to convert starch into sugar.
On d«itroM hcHltby gn#tric juice has no clfoct. Anil its power of invert-
ing cane^ugar seems to be less than that of hydrochloric acid diluted to tlie
eame degree of acidity as itself. Id an UDbcaltby etomaoh. however, con-
lainiDj; much mucus, the gastric juice is^rcry active in converting cane-
BUjjar into dextrose. This power seeoiB to be due to the jirvnence in the
UUGua of a apecial feroKnl, anato);nufl to, but quite didlinct fr»m, tbe
IrtyaliD of ntira. An exceaaive <{uanlity of ('ane-.iugar iniruducol into
iIk stODtKclt CBUsee a aecrctiou of mucus, u>d heiK« providat for its nwo
ccponnioD.
ivl fnv. I hougli llicrc )■ ps]x'ririK'n(nl rviik-iK-c ihiil <:'mii liion <>t riiU la a ;
rrrtBin rxtritt liim lirkr |>]ii<ri.- in (ht- filtfinncli. ibi- trrcnl niHM iil' lht< fnt ftf .
kI i* iMit Ml changed.
Siirh luinernU an are soluble in Tree hydroi-liloric aci<l arc fur the nicMt
|af1 dlMolvvil : ihriii^h there is a ditrerence in (hia anil it) kiiiio other ro*
ifMcIs bMwef'D ipiftrie jiiico and »im|)l« free hydrochloric acid diluted wilh
«atrr tit tbe ratne de^i4« of ncidiiy as the juice, itie praenee either of ibo
pv|«tii nr u( (tllicr Inidies upiiareniiy mMlilyiDg the solveut action of tha
■dd.
The r^wnlinl [iroperty of gastric juic« b tli« power of diMolving prot«id|
maitm and of o-nveriiii); iIk-id iiiln a suhMnni-e ca)le<l |)epli(ue.
AHi»u of yuirif JHtee on protfiih. The Kttullit are t«>euiially the Bame,.]
wbellMr natural juico obtainw by tn«aiiB of n Halula ur ar(iticb) juioe, i. e.,
ui arid inAtuoii uf the tnumtis nwniknn* of tlio Nloniarh, be uned.
ArtllMal (Oniric Juice mij be prepared Ed luiy of Uie followlni; ways;
I . The inuciiu- incmhniiic of a pig • i>r ilos'« (tomacli u rcBiored tri.im tho ihur*
ttilv i-oal. Siirtv- uiiiicril. rubbo>) in n mnrtar witb pouiide<l g\f and ritrHcio]
with aialer. Tlie aquniun extract tillered nnd •cidulati.-d (it U in itaclf aomewhal
B«ri<l). unlil it hu a rr«« aciilliy ourmfKiiiding tu i).'2 i<cr cent, uf bytlrucbloric
add. niotaina but little nf the producia of ili^caliou, sucu at (H^jitone. bat is Ikirly
pMaaL
S- Tliv aiuciiUH Dirnibninc nituiUrU- prp|inrrd mid iniiu-ol in nllotrvd todijmt
■I S6* C. ill • lance >(uuiility of )iydrucliU>ric uci-l ditutcil lu 11.2 |<i'r c«nl. The
fnMer port «r the aicml-raiie dl<Mp|)Oiiri-, ^ircdn only being Ivfl. mid tlie soine-
arhal iifialfM-rtic ll(|uid ctiu be devanied and flttensl. Tbe filtrntt' ba« imwerful
dJKiatlve I |>eiiiic) iiroiiMtiM, but contains a tywsiderable amount ••( ibn jiroducta
M dlcnlloa (pe|>totie. tic.), aruinx from tba dig<«lian of tbs niucoun ntrmbmne
tMrif.'
3. Tbi- niiKiKm iiictiibnirit', .liinilurlr pri'iiutc*! and mliiced, b thriMrn ioU> a
mlB|<«rBtivFly tar^c iiunntity uf ccncetilniltH) ^treeriii, and allowed to stand.
The nirwbntn* may w prtvioudy •U'hydriitfil liy Wng iilliiwed In 'tand under
alrriti'd. bat ihlit ii* not necnwitr, and u lo" iirolonxod action nf the al«<ih<il injures
nr rvpu dniroy* the activity ol' tbe ]>rDilut.-l. The dccuiilcl cleat Rlyoerin, ill
wbirb a COO) pa rn lively sBiail auaatity of (he unliimry jirMelda of the mucous
■Mtaibrane are dli^~-l<ed. if added (u hydrochloric acid of 0.2 per cent, (about
I r.c. of tbr xt/ccrin to 100 c.c- of the dilut« acid are Hifbficni), makm an
arllfirlal juici- (iiierably free from unlinary pmtehU luid peptone, nnd of rooiark-
able ["ileni-y. the iirmeoce of the glycwrin not ioletfcriiig «'ilb Uie reBiillM.
Bi'f.'re procectlin-; to xludy the action of );iu>tric juice on proteid*. it will
hr iiM'fiil to n-vicH very hriclly the cliief obaraclen of the more iuiporiaiiC
iii<imtii-n of the groups
The more Inijioriaiit proieida wbich we linte tbu> far studies) aret I. Ft-
hrin. itiiiolubte in Hiilcr and nut reully w>tuble i ■'. r., without clnuiKc) iusnliita
•oluii'inn. 'i. Mynnn, insoluble in niil^r bui MiUiblo in «liiw «iiKit>n<i#, tifi>>
v]d«l ihnp are not loo dilute iir trio coniiiitmied. 3. Olohulin (Incliidiiig
pKrak'lobiiliD, librini^tn.etc... inudiiblc in «ali-r, hut rondily mdiihlo in evca
vrry diUiir valine Mibitionii. 4. AHiu>nin,tfr\im-all/utniH,»n\iiMti in water
in tW ab«rncr of nil ra\\r. •>. Aciti-ulliumin, i»tii nhicb vlohulin* and
mrriMn arc npidir ronvi-rtrd l>y (he action of dilute acidii, llie [nirtii-iilar
ami albumin into uhich ilir niyirin of riukIi.' in ohaiigtd, Ix-iii^ cimriiinM
railed 'y'cNin- If tlir rmirciil UK'd be not dilute acid hut diliilo alltall
th> prt:<tuct i» called nlknli-nlhumin. Tbe tno twdics, nc id -id bum in and
I
Invvm. aw) l« tenoTed tif rcnnntnUnn *l N* C Mrf MlbivtiwviH liluittt
J
TI1K TtRSirifl ATfD MBCRAKtSMfl Or DldlilTIO!?.
slIutlialbuaiiD. are very parallel ia ibeir clinraL-t«rd, and may readily be
ooQverleil, the one into ine otbcr, by the u^e of dilute alkali or dilute and
nMI>edively. Their moat importaot common uharoolersare iowlubilily in
WAtAT and iu saline aolutioos and ready solubility in dilute ncida and
alkaliea. 6. CongulUed prateidn. PiS we have teen, wbcii fibrin suKpeiided
in water, scrum -albuniiR in noliilion, tirid -albumin ur alkali-albumin fir«>
peiulo'l in watvr, or para^lobiilin »U4|>t.-nded in irater nr diMtilvtid in a dilute
ml ine solution, ore hmtvd l4> n li^mjierature ivfaich for the whole group may
b"* put down at ubnnt "■'>' to Si)" C, each of them bcuomiM coagulatnj, and
after tbe chnu^- i« in*(i]uble in water, saliue solution*, dilute acids, Mc iftj
fact iu everything but very strong acids. Myosin and libriuogvn undergo ft|
similar change at a lower t«mpernture, vix., about -^li" C. We may. for
prasMit purposM, spojik of all these proteid* thus changed under the one
lem of coagulated proloid^.
To the above list ne may now add two other proteids, vix : 7. A kind of
albumin which forms tbe gr«at bulk of the pruteid matter prewnt in r.iw
" while of egff," and which, since it differs in minor character« from the albu-
min of blood and of the tisfues, ia called i'jy-alhii'nm, H. The pwuliftr pfo^
teid eaiein, an importaut conslitnent of milk. This may perhaps be refardedl
as a naturally oocurriuK alkali-albumin nince it hiu many resembUncea to
the ai'tilieial alkali-albumiu ; but for several reasons it is desirable to consider
it a.t an iude|iendi.-nl body.
Eg;;-albumiu like serum -a I bu rain becomci coagulated at a temperature of
about 7o° u> SO"* C, and though eas^D as it uaiurallj esiais in milk is
not coagulated on boiling, whun iieparated out in a special way. andiiuspended
in *water in which it i* insoluble, it becomes ooagulatett at about I'l" to
80' C,
It will b« observed thai all these proteids form, as regards their solubilt-
ties, a dwoondiiig series in the tbllowing order: cmtgulated proteids; librin ;
acid'albumin wilh alknli-albumin, nnd casein: myofliu, globulint; eeram-
albumin with egg-nlbumin.
We must now return to the action of gastric juice.
If a few shreds of (ibriii, obtained by whippin;; blood, after being thor- I
oughly washed aud boiled and thus by the boiling coagulated, bethrono \
intoaquanlity of gastric juice, and the mixture be espoied to a temperaiure
of tnm 35° to &" C. the fibrin will speedily, in some coses in a few
minutes, be disaolred. The shreds firxt swell up and become transparent,
tlicn gradually dissolve, and finally disappear with the exception of some
granular dL-bn:*. the amount of which, though generally small, varies accord-
ing to circumstuuoes. If raw, that is, unboiled, uncoagulHte<l tibrin l>e
employed (be ssune cliangea may be otuwrveil, but they take ])lace much
more rapidly.
If small raonds of coagulated albumin, such as white of cvg, be tmted
in tbo same way, the same solution is obserted. The piece* uecom'^ trans-
partinl at their surfnoes: this i^ ejtjxtcially seun at th(> edgft*. which gradually
iMOone rounded down ; and solution «teiulily progru&tes from the oubtido of
Uie lueca inward.
If any other form of coagulaU-jl nlbumiu {.e.g., precipitated acid- or alkali-
albumin, suspended in water aud boilvdj be treated iu the same way, a simi-
lar solution tJJtoa dImc. The readiitess with which the sDlutiuu is etfeoted, J
will depend, «et«ru parihtu, on tbe amollncis of the pieces, or rather on the 1
amount of surface as compared with bulk, which is prewuted to tbe actJon
of the juioe.
Oastrie juioe then readily dissolves conguUted proteids, which ntherwiee I
■re insoluble, or soluble only, anl that with didiculty, iu very strong acids. '
wnior or in diliil*
with gMlrii: jiitn*. no visiljlr (.'himgo inkaf |)liu^v; tnit nt-vcrlht'lum. it it
oo «zai»inBl»<>ii tliitl ilip aoluttoo* linve umliT^oiK' n rrmurknlile cti«n^, iha
•Blon ><r ntii';)) i» oti>il.r seen bv otnl nmtiiig H with lh<r chim^'c cHV-clcd by
diluii- mil) bIoiic. If raw nliiic of egg. lurgcly (liliiieil wiili nnter nnd
•tniiiKi). tw ireiilMl with n <utlicienl '(uaniitj' ol'ililutv Imlrochloric nciil, the
uiMlnt-piKxi or liirbi'liiy whicl) a[)|>enn?d in the whit« of egg on liilulioii (nnd
Which t± due tu the prvnjuuitiiio of variouH forme of globulin accompanying
ihv ei^-albuinin in the raw white) ditappearB, and a clear mislur« rcsulta.
If a {Mrtioii of the mixture be at once noiled. a larj*e dep'nit of coagulated
albumin occurs. If, liowe^'er, the mixture be expoaed to 5(1^ to 56' 0.
tot 9om« time, tlie aniouui of coaKulatJoii which in produced by bmlin^ a
•Mriniea bveoiuM leMi, and. fiually. builin;; products no col^■ulalion wbaUver.
By Ofutraliitiition, however, iliu wh»lu of the albumin (with such reMrictiuos
a* the preveooc of certain neutml »alt« mar cnuiie} may be obtained in th«
(ana of Acid>albuni!n, the filtrati: alV-r nvutraliiiution containing no ]irut«i(b
M all (of a ref}- small quantity^. Thii* thu wbolo of the albumin urKwnt
in the whiltf oJ'vgK i^ay be. in time, conrurteil. by the nimplv ncliou nf diiiit«
hydracbloric ncxl. into ai'id-albumln. S^ruiii-alfaumin nimilarly trejitod
UMletvocs, in eourtv of time a »imiliir rntivernion into ncidaltnimiii. and we
lwT« luraaity M<en (ji o9 1 that volution* of mro^io or of any of thi; globulins
an) vriili rrmnrkablp rapidity con\Mrlcil into ncid-albuniin. Thu« ciinple
dilute bydrochloHr of the aame dirgrrv of nciility at gastric juice, merely
coDvvrtB ibeae pnM«i<ls into acM'Slbiimin. Ilie rapidity of the cbanga diflsi^
iog with the diflEtreot proteide, being in some crm« very elow, and requiring
» relatively high tem{)eratuiT.
If the awe white of egs or serum-albumin be Ireate'l with gastric juic«
hmfwd "f simple dilute hydrochloric acid, the events for some time socm tbo
aaaic. Tbu» after a while boiling cuumm i>(> ciMgulatiou, wbile neulralixation
pn» a cMuiderablc precipitate of a protad bodv, which, being insoluble in
waUr and in todium clili>nde »olutiiiii.^ and Mluble in dilute alkali and acids, _
ai leaK cIomIy nwmbW acid-albumin. Etut it u found that only a ponioaB
of ihe pnicid originally prmcnt in the white of egg or »erum albumin can '
tkna be n^iwl by pn-cipitnlion. Though thv neniralluilion he i-arriud out
irttli Um graateit care it will be found, on Rltering off the neutralixation pre-
cifMtAt^ (nat i«. ibe actd-nlbumin, ihnt thv tiltratc, a*«bown on cmplnying
Um Tariqu? teets for prolcid (mo ^ !■>) or on ailding an adciuatu i^uaiitity oT^
MinO|f alcohol. Mill contains a very considerable quantity of pMttud matter ;f
^ tinri
■ cha
and, un the whole. iIm.- lunger tbr digestion is carried on. tbc grraK-r u the
pro|>>irti"n buriH- by ihc proteid remaining in solution to tJie prm^iiiitatu
ibrown down oii neulralizntion ; indeed, in some cnscfl at all events, all tite
(uiileid maltcr ori^'iniilly present remains in solution, nnd there is no nMi<
iratitation orecipilulion at all, or at ntiMl a wholly ioMguilicant one
$ 203. 1 he proietil matter. thu« remaining' in <4ilulion after ne^ilralixnltoa,
diflbti from all the proteids wbich we have hitherto studied in a^ mncli as,
Uiourb exiitinjt in a i>eutral solution, ii h not coagulated by hest, like the
effg-<lbiimio or serum-albumiu from which it hts l)««n produced : tbo solu-
tinri, after the iieutratiiaiion precipituie hiu been fillers) oir. remains quite
char when )M>il«d. The only other noluiiivna of protei^b which do nirt ooagu-
~ itr on boiling are Aolutiuun of aciil- ur alkali-albumin ; but these aolulious
iiu>i Ix- acid or alkali rMjieetively : the acid-albumin or alkalt-albumin ta
'fatoluble in a nnitral uilutiuii, and when niintdr tiuii{>euded in water it
raadily eoagulntc*) at a tcmjivfnlur^- of 7V ('. Thu new pr^iteiil matter of
which are arc viieNkiDg is •■>luble in neutnl tolutiouji. indeed, iu diMilled
water, nod vn under no circuOMUnew be oongulalvd by heat.
312 TUB Tissues AKii MECUAMSUS Uf UIQKSTIUX.
H VpoD ^xHininslion we find that ttie netr protoid ii>Mler tUua left in anlu-
Hon ciJiifii^E uf Hi k«Bt two iMsiincppruteKl binliEit. Iflo tli« aolulinn ainino-
■ niuiii sul|ihHle be ui!i|«il, pari of iho ]in)tci() luaUer U precipituted while i«rt
is lilill Itlt ill eoluiioii. Tiie prot«iil litidy tlitia tliruMu dnwa is called a/fru- '
vine (ttiere are several varielica ol' albunune but them- ut-L-d not aavt ili-tniii
na). It iipproacliea albumin ia nature by n-VAoii nf th>- Inci tbai ii will tint
diflbfte ibmuKb meiubraues ; tbiit it ilitrt-r« hunuvcr widely fmm that pmirid
is .hUcjWU by it» nolulioitit uol cimxulaiiiig on builiiij;. Thi- boiiy wbicli i* not
thnmn down by »fiini<iiiiimi .Mdphalu ia culled wjAunf; it ililliTt (rmii allHi-
DKiM' iu beiiif( clilliiKiblr, fur it ndl iiiisii tlirnugli iiii'ii)braiK'«. Tin* dill'iitiiin
is not iicorly Ml rapid nii ibiil nf ■nliii. HUgar, aad olht'r nimilnr imltrtAiice^ :
indwd vobitioDN uf |ii:|)|iiniii niiiy lie In^'d from nilU by dialysis. But it in
very ninrk<-<I ak cnnipan-il witb ihaC of other proU-idn; tbo»p pan tlirouph
ni«ndinuic« wiih the ^mu-:<l dillieiilty, if at nil. I'eplone i« in«ulubl<> in
slevbol. and may Iw prei-i piloted Ihim il« lobilioiin by tJie «d<litiwi of un
fltkiiiiaii' i)iinnlity of (hi« mgcnt; but for tlii» purpoM a wty lur^ excess
ofnlcoliol IS needed, utherwixp niui-h of the peploiie remains in solution. It
may W kept undor aleobol lor n Ion); time wiihoui underf>oin)r cbange.
wberms olher pr»(eitfs are moro or Uh slowly eiMi}>uhited by alcohol. A
UBcfiil l»t lor peptone is furnisheti by the I'ncl tbnt a solulion uf p«pt«n«.
mixed nilh a slroii]; snlulioD of caustic iKilush, jjives on nddilion iiT a mere
Iraco ol' eupric sulpbatc in lb(> cold a pink eulor, wherctis other proteiib give
a vioUt color. Id iipplvin^; tbis leel, liowever. cure must be laken not to add
loo much ciiprio wilpliaie MUce in ibat case a violel color, deepening oo
boiling, tbnt is. the ordinary proteid reacliou (see § 15), is obtained.
Thei'e are cvasoiis lor ihinkiii); llial ihere are several kinds or at learil;
more than one kind of peptone: but "e may for ibe presenl re>,iird ihe sul>>
stance as one. For a \ua^ time albumnae was confounded with |>(.'p(oiK', and
many of tbc coninicrrial formn of '* pcpIuQC" nmiiiiit Inr^ly ot allHimiiM>:
indeed, thu two arc clowly nllicil and havfl many n^iictions in common, iliu
mu«l ulrikiiiK: diffen-nce* being that jie|>ton« b difrtis^ible, while albuimiw to
not. or hanlly at all, and that. |)epl'>np is not liki- albnnic^e precipitated by .
amiDuniuin sulphalc. The nmoiiiit uf albumiwo np|K-nrine in a dij^estion
ttxperimeiil, relative to ibo amount of true peptone, denends on the lu-tivitr
of the juice, and other circunislAnees. Wo may regard albumow as a leas
oomplcle product of digestion than peptone.
liw precipitate thronn down by neutraliintioD after the action of gnslrio
juice on egg- or senini albiiniin resemble, in its general characters, acid-
albumin. I^ince, however, it ))rc<bah1y is distinguishable from tbe body or
hu<iiefi jiTodneed by the ncliuu of simple acid on muscle or white of ej^g;, it la
bcsi to reserve for it tbe name of fiarapi^Ume, which was originally applied
to it.
Tbun the dii^'t^iion by gastric juice of solutions of e^j^albumin or serum-
albiimin rt-sulU in theoinveraioD of all tbe proleids present into jie{>tone.
alhumoie and narafieplunc, of iibich the tinit mav he considtirtMl as tbe final
and chief pro'liK't, and the other Inn tL* iuiermeuiBle prodiiet?. ooeurring in
varying iiuantitv, |i<in<iblv not nlwayx fori]i<<<l. and [indwbly uf (ccvimlarr
im|Hiitan<'e. wIm'H librin, either raw or Ixiilcd, »r any form of coagulated
proteid is itiwolved himI seems to d!sap|H-ar under the intlnenct- of gaalrio
juice, the same products, peptone, iilwimiwe, and |Mini|>e[>tone make their
ap[N.-amncc. The same biMlice result when myosin or any of the globulin*
are subjected to the action of the juice : and acid-albumiii or alkalialbumiii is
similarly converted into albumow and peptone.
It is obvious that the edect of the action of tlw gastric juiee is U> ohango
the Ins soluble prolcifl into a more soluble form, the eluinge lH:tng either
SALIVA Ah'U OAATBIC JCICE.
818
CDiuiiWinl U[> lo ili« ixaev of miioik. tite ni'isl sululilc (if all pruteKls, or
hnoK It-n in [Hin tnc»<n|>l«li>. Tliia will life smn from ilio fdljowinf; tnlwlar
mmiijnrtiirui iif [tniteiili nocordiiift to tbeir MilutiilUiiM.
titinih •!• dUlUM iraltr.
AintK-u* •nlutton* DOt cja^Uted on boiling :
I' P*i>lon».
^ 'ilo AlUuiDMe.
AfUfiBi »'<iuiion*ctiaguUted on builliig . dlliuDiin.
InKiliihl/ in tliilUM teller.
ttfittf wlable In ilUuie saline mitutiotio (NaCI t
Mrnrnt.) Globulins.
Solnbl* (Milv in sUonger ■•lino wiUitiim* (XaCI H
lu 10 p«r MtiL) Mymii).
ItuolfAU in itilnit m/inr toliifkmt.
Rwdity mttilite li. dilute -eid (IICl 0.1 per «•«.) ] JfJli"";!!^!";
'"'*"''■"''' ■ • jOMcin.
t^>lulilF with <lifllcultj lu (lihil« add, ibal is at
bicb Ifioprrntur* (UO^C) Mtid iift4>r prolongrd
trrUinviil 'inly Fibrin.
Inviluhlc In dilute adds, ttfilublff only In Mmng
Hiila Coafulattd |>tnU-i4l.
Milk «Wn tnvtv<l with gnriric juice Ia Am nf ull "mnJIeil." Tliis is the
Trmill pnrtiv nl' llir ni-tii>ii of lliv tt^c ai-lil but rhirtly af the *|>erial nctioo
»f H particiilnr oii^liUivDl of pi»lric juiiv, of which w« ■hull ■•(x-itk her»-
afbr. Tbr run) c(itiflisl#nfHfinrticuliirj>r<)(4.'iilmntlvrmixeil with till ; iiikI tbbi
IpfTitriti Buitter a Mibtoiuenily diMolvti) wiih lh« Mine nppMiranco of pi-p-
Incir, allMinoae and iMrapeplone aa in ibi? nw of other nMtciilB. In favt.lhe
di|t«9tiiin by icutric juice of all tl» vaHolica of protd<l« c^irsmU in lliv con-
vvnion of ihv |>ni<«t<l iolo peptoo*. with the cimoiniitanl np|>caranc<.' of K
certain rariahh- amount of albuMnwe and nnra|>eplone.
tS04. Cireiimtiiinfri nfffHinif e/iulrif 'li'jffiioiu The aulvenl action of
OMric jiiit« on proteitU ia inmlifi^l hr a vari«tv of circumstan (.■«■». Th4
■Uun* of th« prolrid llMlf raukcn a (lid«ren<«. tliou^h thid in tk-trrmincd
profanbly by phyiiical rath«- than by cheuiit-al characters. Hence in mak*
hig ■ arriM of cuniparalive triab tiic aauK^ pruiett) uhouUI be U*eii. anil tb«
fcrm of prolciil mo*l o>fivctiiL-nt for the pur|KW Ja fibrin. If it be dcairrcl
■linply t« aca-rlaJn nhcihrr anr giviin iifMx-iinen has *ay digealtve |xmer» at
alt, it ia Initt to iw lH>il<fl fibriu, ainoc rati' fibrin in weutually Oiiuulvcd by
tlilule hyilrtwhlorit- arid nl<>n«, prohnlily on ucfiuiit of vouiv |ie|>?iu pre-
vboalj pimai in thr IiIixhI Iwonniinic eutauKkil with tbe fibrin during
PelMtinic- Bui in <.-alimalin^ 'luaiilitalively thu jicplic power of two tiieci-
ntw of euirii.- jui<v nnder dirti'^-nt coniliiliHu. niw Rbrin preparen by
ffniunrr'i nsHhuil ia niD«l coiivviiionL
Ptirtimwuf wrll-narbcd fibrin arc atainwl with camlne and wtaln waahed to
ranofC ibe >uii»llii<>ii* coIaHuc niatlt'r. A fraEini-tit bt lhl> culorcd Ahrin thrown
^^ lotu an acli^e juie* on beo-niini; dln"olvo>l, tivni up ll« cilur lo 'hi- fluid.
^H Hcm^ if Ihr iiHdti- atiirk <i( coturod fibrin Im- uofiI In n M'Hirs of cxprriini-tii-. and
^f tb« aamr bulk* <i{ fibrin and of fluid be urcI in «iai'b case, the amount «r ilbrin
^^ itWdtpil Biay bi> fairly nlloiatcd if tbe dGnth nf lint ^iven to tlie fiuld. Kilitla
tbti* cnlfltFd wiih carniitic may be i>re*«rveii in eiber.
814 THK Tissues XSt> USORANIStfS OP DIOESTIOS'.
Cilice, if suflicieiil time be allovrei), even & emnll '(Uantitr of giu^tHc juice
irill (liiwulve at lenat n very large it' Dot an iiidelinite ijtiaiiliiy of dbrin, we
aiv kiJ Ic iiike, af a measure of tlie aclivily of a epeciniea of -gastric juice,
not the iiiiaatiLv of fibriu which it will ultiniateW uiaM>lTe, but the rapiditj
with which it tliasolves a giveD quantity.
The t;reater the surface presented to the nctioD of the juice, the more
rapid the anhitioo ; hence minute division and constant movement favor digee-
tiun. And this iit prohably, in part nt leaat, the readon why a fraRmeot of
sfiuu^v lUameotout fibrin i^ more readily di.iaolved than a solid clump of
bdiliKl white of egK of the earae aiie. Neutralization of the juice wholly
arrets digf«tiun ; fibrin may be »ul>iuitt«d for an almost indefinite linie to
Ihc action uf neiitnOized gastric juice witliDiil being dig«*ted. If lJ>e neu-
tnUiiuid juioe be properly acidified, it may a^ain becomr antire; when }ifi»-
trie juioe, however, Iiaa been made alkaline, and kept for «iimc time at a
temperature of 35°, ita solreiil jiowen are not only nuKiieDded but actually
dettrovcd. DigMtiun in miwl ra|iid with dilute hydrochloric aciil of 0.'2 {le^
rent, (ilie acidity of nntuni] i^iutric juicii). If the Juice cotitnins much more
or much lew free acid than thU, iu activity in distinctly impaired. Other
iicidi>, lactic, phwph one, etc, may be sulwl I tutted for hydrochloric ; but (hey
are not so fiDwtual, and the dcgn^c uf acidity most useful varies with the
ditTcreiit acids. The presence of neutral lalta, »uch lis sodium chlimdi-, in
exocsi is injurious. Tne action of inuniniiilinn gantric juice is most rapid at
8A''-40"C. :at the ordinan' temperature it is much slower, and at about
V 0. ceases altogether. Tiie juice may be kept, however, at 0^ C. for an
indefinite period without injury to its powere. The gastric juice of cold-
blooded vertebrates is relntively more active at low temperatures than that
of warm-blooded mammals or birds.
At temperatures much above 40° or 45° the action of the juice is im-
paired. By boiling for a few minutes the activity of the most powerful
juice is irrevocably detitroyed. The presence in a ooueentratetl form uf the
pri'ilucts of diici^iiuu liiniters the process of solution. If a large (Quantity
of Hbrin be placed in a .imall iiuauiity uf juice, diguMtion is soun arrc»t«d ;
on dilutlun with the normal hyiirochloric acid (l).2 per ceuL), or if the roil-
ture l>e submitted to dialvftfi to retuuve the |tcpt<in«s formed, ami iti> acidity
be kept up to the normal, the action rciwminenccs. By removing the pro-
duciM of aigntioD as last u tbcy are formed, and bv keeping the acidity up
to the normal, a av<ia amount uf gastric juice mav he made to digest a wry
large quantity of protdd mRt«rifil. Whether llic qusutity is really un-
limited it dtspiitcd : hut in any case the energies of the Jui(.-e are not rapidly
exhausted by the act of digcjlion.
$205. yalinr of llic artion. All these fncts go to show that the digestive
action of gastric juice on proteids, like that of saliva on starch, is a ferment-
action ; in other words, that the solvent action of gastric juice is eestiitially
due to the presence in it of a ferment-body. To this ferment-botly, whiL-Ii
IIS yet has been only approximately isolated, the name oi'peptinaaa been
given. It is present not only in gastric juice but also in the tthuidi of the
gastric mucous membrane, especially in certain parts and under certain
conditions which we shall study presently. The ulyi^erin extract of gawtric
tnucous membrane, at any role of that which has bt:en dehydrated, contain*
a minimal <|uantity of proleid matter, and yet is itjtemely peptic. Other
methods, such as the elubonite one of Brin-ke, give u* a mnlerial which,
though contaiaing nitrogen, exbibiij nune of the ordiuttry pmtcid reactions,
and yet in concert with normnl dilute hvdrorhluric ncid ix peptic in n very
high degree. We itcein, tht-Ti-forc, jiiitlbed in iMwrting that pepsin is not a
proteid, but it wutild Ixi Imxantuus tt> make any dogmatic statement con-
3ALirA AND GASTRIC JUICE.
815
I
ilngflBHBwicii', olftalned in an tauM m (|iiaDtilT nt a limo chat iu cxsct
eitmuiait AttlMtn ItaTv npt yet Ih-mi a>M?i?riniiii?il. At prewot the luxiiirea-
Utioct of pcfitic powcn in nur ouly mIV- tral »f the pr«wnc« of pep*in.
In (HM- iruportnjil m>p<-ct prp«in, ilic tprmciit ol' gMtric Juice, didera from
|it_\uliu. ilir fi-nwitl <ii' wiliv*. J'nliva it active in h pert'eotly oeuiral
■nMltijtu, nnd there vvrmt to \jv iiv epocinl cotiiiectiun beLweeo llie ferment
Mill any alkali ur ndil. In gutric juice, however, there is n nrouj; lie be>
t«fHi !!»■ Brill an'l the ferment. »<> ttnag thai some wriien apeak <it peiisin
■nd hyilrurhluric nciil »;• forming togelner a compound, pepto-hydroL'hforic
>od.
Id the alMenoo of exact knowled)^ of the conittitution of proieidi, we
^iiiiil state tltstinctly what is the precise nature of ibe ehan)^ into pep-
tooe; ibo vanou^ proieMk difler from each other in elemeiitan* compuaitioD
•laHe u widely as does pe|>ione from any of tbein. Jurl^ng ftom tlie
MMlofEy with tbe action of saliva on etarch, we may fairly ituppoie that the
ptwcMi ia al bottom one of hydration ; and this view i-. further nuggetiod by
llw fact thai peptone closely resembling, if not identical with, thai obtniii«d
by putric <lixi«tioo. may be obtained hy t)i« artiitii of the xtroug acii)*, by
tM pniloDf^ action of dilute ttt-id* eK[)eria]ly at a high tcinperaiiin', ur
•imply Etr aigcMioD with super- heactsl waivr in n Pai>iii's digester, that is to
say hr hmoik of a^nt« which, in other com* ]>r<Klu<« thoir eSccIa by brine-
it^ abnut bydrolyttc changea ; beyond tht« we cnniiot nt presoJil go. Vr*
nay idd, however, as supiHirtiujc the t>ame view, the statement of flom*
obaarvin that peptone when In-aml wilh dehydrating agents or when sim-
ply boaud to I4ir-170' ('. ix in [lOTt rt'oonvcrlcil into n body or bodies re-
Mnbling acid albumin or globulin.
i 806. All proicid;', sn iar as we know, are eoovorted by pcpin into pep*
IMM. CocHerainK the action of ^i^ric juice on otiier nitnigenoua sub-
■r«am more or leas allied to proteidd hut not imty piuteid in nature our
kaowledge is at prenent imperfect. Mucin, nuclein, and the cbemieal boaia
of bomy liKVee are wholly uuafTected by ^a^tric juice. The gelatiniferona
lisaues are dissolved by it; and the bundles and mem b nines of connective
tiaue are very speedily so far aSected by ii. that at a verr early xtoge of
4i|ntioa. the lyundles and elementary fibret of U)uscle< wfiK^b are hound
laystfaer br coiineotivi;? tiMuefall aitunder; moreover, both pnijuirvd gehilio
•od U)« gMaiinUerous bagia of connective tissue in lis natural condition, that
ii wflbout being pr«r1ously heated with water, are by it ehangeil into a sub*
■taacvso far nnaliii^'ni^ with pepinne. tlmt the chanicicrisltr property of ^la-
lloinlloi) is riilin-ly hitt. ('h'>iidrin ami the elastic lisiucii uudrrgn n similar
cbug*. It in ti '■''!. liow far thla cbaoge is due simply to tbt
acid of gaatrir - i: .> i My of the i>cp«io.
(207. .1 ' jnir.- -,n milk. It lias long bean known that an
iafaninn ol : i i n-h, citllnl •rnnrt, has a remarkable effect in rapidly
enrdlio); milk, ami thiif pr»|icrty is made uso of in the m.inufHi.-turie of
eheae. (jastric juict> has a similar ofleel; milk when iiihjei-iml to the
aetkio of ^trie juice is first eiirdlol and then digmted. If a few drops of
gasirio juiee l>e added to a little milk in a losl-lul)o, and the mixture ox-
ptanl to a temperature of 40'. the milk will curdle into a complete clot in a
▼err sbon time. If tb« action l)e coniioucd ibe curd or clot will be ulli-
DBtaIr dlMolved and dig«at«d. Milk contains, beside- a peculiar fnnu, or
peculiar form* of albumin, fats, mitk-^ugar and variotis salines, the peculiar
tarutrid n»riii. In natural milk casein is present in solutiou. and " curd-
iag" conaiaia easeatiailj in the Mluble ca^iu being converted (or more
pf^bly na wc shall aee preKntly, split ujm into an in^ilubie m'xiilicalion
af cosrio, which as it is lieiux pr«cipitated carries down with it a great deal
310
of ihe bt ati'I *o funns tlic "ciinl," Now ciUL-iH ia nwlily jmN'iptutcd
fiKtm milk upmi the mMiliuii ul' it Hniutl 'iiiatility nf ncii], mid il uiii,'lii b«
Biippi:«ed tbnl tlieciir<llin)^ (Hl-ct (it'Eiistricjuiw wns due to it« lu-td rcacliun-
But thii is nol tbc niac, lor ticuiraliu^d gastric juice, or neutral reaoet, ia
e(|ually ufficiK-lou*.
Tbc t-urdling nction of ivonet is cloHely dependent ao temperature, belrif;
like th<; [H-]>lii' HClion of f^tric juice favored by a riw of (oiuperattir<> up tt>
aImuC -IO". Mnrenvcr the (.-urdliug m-liuu ia deetmyed by pn>i'ioua builinx
of lh« juico or nMiticL. These lacU aaggeet ihat a ferineui is al (he bodom
of itiu niali«r: and, indeed, all ilie features of itie aciioii support llm view.
Mor«i>vcr. as a mailer of fad. a vurdlioK ferincDt may be exiracled by
jrlyoi-fin and by the olber metlioda iiwil for (ircjiarin^c feriiienia^ Tbe fer-
nK'iil. however, ia unl jwpalu but »<ioie other bmly ; and ilie iwu may W
»cpiiTiik-d fr»ni eaeb other. If mu^nie^iium i;arb>iii»lu in ]>uwiler In- i-aii-
tioufly added to ^'astrc jiiiec or to tia infiiiiiiin of calve?' alomuch n cu])ii>ua
precipiliile ie formed. If thi- adiliriou of niogMiium oarboaat« be Uoj){inl
aa soon nit uuy fiirilit^r pr<-ri|iiijitir<ii ct-aiKg to he roiiivd by It, and the mix-
ture be ullonei] to Aland, the cli.-ar fluid li:!! abovf the predpitalv will Iw
fouuil to I'urdli- milk reuililv, but r.v<'ii when iiciililiinl to have no iH-plic
action oil pniU'iili'. .iliowin^ ibat thv^ nrri'ipiiiili- euui^i^il bv Uio addition of
Uh' majcnoiuui <-nrbiiual>.- hn» tiirrird donii all thu iK-pnn (lut M\ W'hiiid at
lca--i| a -^uxl di-nl 'if ihf " curdling" or r«n net- ferment.
It nii^hl III- (hou^'bt, thai tbo rcnnei-tiirmeni, rmuin «i- may mil it. actcH
by inducing » iVrnictitnlion in the sufrar nf milk, giving rite Ui laolie noiil
irhich pn-L-tpitaCeil the casein by virtue of iu Wog an acid. But ihl* view
ia dispruved by the foltowiit); facts which show that the fcrmettt prixliiixv ita
nirttting effect by acting directly on the natural coeeio its«lf Casein may be
precipiiated uiK'hiiii);ed, that is, capable of rediaHolvint; in water (the pres«m«
of culctc fihoiipiiiiit.- WiuK axitunud) by «atunUiiig milk with neutral »altn«
biHJii-^ I sui-h It* uiiHum chloride or magnealum niiTpluilit.) ; ami by betn;; jifo-
cipiialvd anil redi^iolvi'd more than once may W obiniuvd lurgidy fnv from
fat and wholly fnc from niilk-:<ugar. Such milutious of iwlutvil ciiaein fretil
from milk-augar uuiy \ie made to inirdlc like natural milk by tbt- adililion of
rvnnin. xhoiring that the milk-Ku^r hiui untbing U> do with tlio matter.
Morvovcr the precipitate thrown down from milk by dilute acidn, Imnii^ arid
included, is itself utmllercd or very jjightly alt^1r«l <-iu>eiu. nol etinl. and
with care may br so prepared as to b<t redioolvvd into solutionv which
eiinlle with rennin, like solutions of casein prepnrvd by means of nctitral
salts.
When iBolaf^l casein is curdled by meaoH of reniiin two (>mt«iiW. !t u
slated, make tbeir appearance, one which is soluble and allied to albumin,
aod anotlier, which is iimoluble and forms the eunl. Curdling, therefore,
Bcoonliug to Ibis result apiveura to he ibespliltins up by a ferment of ■ more
complex bmtv ; oiid it a iutereslio); to olwerve. as ]ierliH]» throwing light on
the ?iiiii<-wbat analoKouH formation nf the librin. that this cunlting action
will not laki- plutn- if i-al<-ic pb<>i|)hate be wholly ulMeut from the mixture.
The calcic [itmni'hiiU' apin-uni to pUiy a jwinliar part in deterniiuin;: the
Inwcduhility of the <-unl, lor tl»r« ia evidi-iK« that in the abwucc of catdc
phiwphali- ihi' Itirmcnl haa [hiwit Co attack itie ciuein and »plit it up. but
that iMilb proilurin n^maiii in »>iluii<iu; if calcic phudpbate be pr«aeut, Uto
one. vit, the curd,' U-conuMi tn-iotublc.
Itonnin i* iibundant in the kfO'tric juice and in thi- ciLilrie aiucoiiH mem-
branv of rrimiiiaul*, but it also found in the grwlnc juio: of other animal*,
■ It nilRh* tig uirfiil. Ill iin)<-r [» <lL>iitu.-iitili ll>« -^unl fraa Ittx TUIuia) «diililDta«ctn,IOGklllba
BmuH Itrtfa <it^. •^Iiww. aa>) lu rvwiiv Uir uuim ot iu-vIb I<i( itir luiiir.
flAblVA AMI ISASTItlr Jl-ICB,
SI 7
■I
it. or wJmt we ahull pre^iiiily liave orcostoit lo speak i>f hh ihe
•f tlit-rcniiriit or :ir""»;'^''.iit |)ii«eiit ulan in the inuooui iii«iiibr»iie
riinfli nr iiKiM iiiiiriiiil.'. A verr Biniilar if tigt Idtfultcnl leriti«nl
<i-D r<>uD(l in niaiiv |>Iiiiit(i.
Tmk &TKL*(-rrK». i>K Tiir. Sauvaky Glakik*. Tiit: OAi^ntK; Mirooo
Ml:MllltAXK. TIIK l'A^CItt:AO. AM> TIIK lK-«»'IIA()t^.
$ 906. i)«roT« vm etuily the iintiin? of llie proiTWcti liv uhloli ihe sluoincfa
BDii ibewlivmry glnnila are nble ta Mcrclo ihe >ra'<triL- iiiice nii'l giiliva, whiMO
rrroarkalile pru|ier1i«a we have jugt dwrribet), il wit) be desirable [o mv a
few tHiirxia ou the stnicturo of bulb the above or|;aDS,
ThfXNigbout the rmter {lart of il« Ieii]>th. rroni ihe curtHao end of (he
aanphvus to oear the aiius. the alimentary caonl is eonslriieiiil on a corlutn
gHMralplan. Thi* pari of the alitDenlary eaosl is liirmed i>ut of the mid*
jntt of the embrj'D, aiid the e(iil helium whii'h tioes it is of hrpul'laittic origin.
tbr mi'iiih and the anua hsv« a diflereitt bHjfiii : they ure n>rnie>) hv im-olii-
tioiu of thi? external skin, the epithelium uf which in of f;)ililnHtii; origin;
moA th* ptau of ttriielure of the motith ami lerniinitl fiortioii of the n-cluin
Ib in iBMne Napeda ditfL-rent from ihal of the rent of the nllraeniarr i-anal.
TIm tranriilon Atrni ihv vjiihluiiiii- to the hypoblaatic ninul on-urii in Iha
fvrlan at the ani». but at ihe ulhor end i» at P>mv dixlutK-e from tho mouth
drar U( ihi' jiinrti'iu of tlx- <i-H»phagu« irilli (he ittamarh.
'rtw plan uf Rtnictiire of tbc nypublaittic [wriion uf the ctstti is soiDewhat
M r»tkNn :
A "iW^ ^*7*f '>f cjliadricnl, columnar, cubical nr upheroidal " proto-
plaamic cells, that is lo «ay cells which art not trniMfoniicd into Battened
I'M las, forms the ininxslinte liniiij; of the niviiy. Th« celh re^t on a con-
liMtlTa-lisme basis, irhieh if tioe, delirale. and often of a [leciilinr nature
ianwiliately uiwler the epilhelium. but becomes more open, loose, and coane
•1 Mine litlje dimnnoe fnun the cclk This ooonectinvt issue basis b richly
pntvided with bloodveueU and lymphatics, and also eonlains a eertnin
numborof nerrca. The bhxxlvcMch reach up to. and fine capillary networks
an eapecially ahumlant immediately beneulh, the bases of the eellK, but mmo
fm» l>ei«een ihe c«lb ibemselvM; the whole of the epithelium i.-< cxtm-
*aacnlar. The connective ti«ue wliere it louche* Ihe eelU formx a more or
Urm vontiuuouB slieei ; this is nflen «poke» of as Ihe baaeiiient inrinhnino
and may be refcarded as the demarcation between the extra- vascular cihiImv
UutD and ihc vaKular e>jniiective-( issue basis. The two together, th«
•|)itlwtium and the coniicctiv«>iisMie ImuIs, form what U known as the muenv
At the faMM of tho crlindrical eellj, weilgcd in lMMw<«n ihi'm an<l die
lwia*n>cnl meDibrane, may be Mcd, in <'ertain Hituatiou* diitinetly, in other
ailuBtioos less disiinctlv, small oetI«: thai is to Hay, tvlU the IxMly of nhicb
b small nUtivelr to itic nucleus. TIhsw are nupjvMM.tl to be young cells,
brld ID merve to replace any of iW larscr eylindnnO <<clb whicJi may from
time to tinN dbappoar : if «o, the epilhelium dom not strictly vumist always
of a Moel' layer, tbough pmdically il nmy )» an rugarded.
Oulaide tbo mucnus nH-mhram- or miiomK coat Ih place*! the ihick mutrut'ir
coot. Tbb consists of two lavcr* of pUin mux-ular Hbm. an inner thicker
layer, in which the fibre* amf bundU-s -f librcs arc disposed cimilarly mitiid
the lumcD of lh<' alimcniarv canal, anil nit otiler, thinm-r one in which the
fibre* are di«p«s<.il I'lDgituitinatly. The bnndh-* and ihtrteof librcs (see Sd9)
aio bound logrtber by oofineclive ttnue carryiu)^ IiIoimIci^swU. lynipliatict, and
S18 TUE TISSt'SS AS'U MECIIAMSMS Or UIOESTIOX.
nervM, and a tlitn jliopt of conneclive liuiie more nr l«« diatinctly Mpamteta
till.* lliickt-r imit-r cirt'iilur mur-ciilar vutil rnmi tliv ihinni^r mj(i;r lou^tudituill
iiiiiivutar Liiiit.
TIh' iou'i-r (irmilrr narl of ihc iiiticouii mcinUmiio where it lircrini(» ntlnched
111 tlio muscular cunt ix foniKil nf vi-ry Iihw coiinrolivv ttviiio. ihi? inlvnpsoci
of the hiiiidU^ Iw-iiig lurgu ami 'Hifn. Tlii* it tpakta nf lu th<- miimucQU4
tijf*ur nr tulimufitin roal. It i» »■> Iimiik- that the iiiucmw coiit mn raiilr move
over llio miJSviilnr cont, mid along il tho nnc can ciwily br (om away ironi
the "ihcr. nmrc cBBily in »iinif pnttn «f the caiml tbiiii in utiien. It carries
thf larger arU'He*^ mid veiii«. wlKwcgmallrr tiniachcs and cnpillaries |H(m into
■nd front the niiioux mendtranr. Lyinf; in lln' nuioi>iJ» membrane bl soma
lilttc ditiaiice IVdiii tho c|iilhch[im is liioiid » tliin layer of tilaio muBCoUr
fibres, culled the lunirn m»»fulari* nmconr. It ie niDrc eoiiapieuous iti some
eitunticiiis thnii Id othere, and when eomplete consists of au iiinf r siii^fle Jayer
of fibres dieposed eireularly and an out«r single Inver of fibres disfioeed
longiindiDally. The conueclive tifeue on the inside of th« mtiscularia
mucMie, between it and tbe epithelium, is generally i)f a aoniewlutl fliirei*nl
chaniL-ier fronj that ouidide the muscuUrls mueosie, and man y jilaovs U of
the kind called •idenoid or retieuiar tissue ; of f hib we thail hereafter have l«
speak.
Lastly, from the ^tonmcli to tli« rectum the inuacular coat of tbc alimeuiary
canal is covered by the viitcenil layer of the i)eritoneiiiii, Thi* cuiuiiiiit «f a
single layer of ijolyjconal tlatleued nucleated epithelioid cellit (beloHgioE in
reaJity as we shall vev tr> the lyiiiphntic syniem) retliiiK tm a ihiu coniivctivc
tissue basis uhich ^cimrutb them I'mm the Imiuitudinnl iniinciiUr crjat.
The jjetieral ^iliin of Riruciure of tliL- ulimi-nlary canal, then, in iu hvpo-
hlastic jiortinn, ■» a compact niuocidar cual iH.-|iarut«'d by n l(iii«c, more or Tea
muvahli-. Hubmiu'nU' coat from a fiiirly roiupiul. nmcotiH c»al. Thv mucmis
coal ci>nsii>i» of a viLHMilar ci>niicctivc-li^uv bivis, in which ii imbetlde«l a
thin "pecinl miiK-ntnr sheet, and of a tingle layer iif frpecial hypt^laistic
emthvlial celli. The mnscular coat eon«iels of a thick inner circular aud a
tuin oilier loii(;iltidina1 layer of plain muscular fibre*, and the whole ia Cov-
ered with an epithelioid peritoneal laver.
§ 209. Oliiu'lt. The Burlhee of the muciius membrane, however, ia Dot
even and unbroken. Jt dips down at intervaU— thai is to say, it is involuted
to form pockets or deprewiuna sunk into the underlying connective liMue,
and diOering in site and form in different parts of the alimeotary canal.
8ueh an involuticn is i-alled a y land. The most simple kind of gland ia a
eylindrical deprcHsiim with ii blind end. somewhat of the form of a ttwl-tiibe,
lined with a sjngic layer of epithelium cells, cuntinuous at the mouth of the
^land with the rent of the epithelium of the mucou* membmnc. The wall
of the ;;1uud outside the epithelium in siipplit-il bv the connective Itwun of
the mucon» membrane, which gt-iierally forms a diMinct bma-mi'nt membrane,
and is jti-nerally also richly Mipidied with capillary l>toodvr«»eIi. Hcn<«,
when two >nch glamU lie side by aide, a ccrlaiu rjuHntity of I'uniicctivc tivuo
carrying blucxlvntu-U ninj" ui> lulwcrn thr-m to reach the epithelial cells
which cover the surface of the itiueous membtmi'; lielween their mouths.
Such a 4nipl<' tubular gland muy hnve the wamc diameter throughout, or
mav %-arr in diameter at diflii^reni distances from tJi« mouth, and tbe epi-
thelium lining it mav be of the same chnrnttcr ihrougbout and similar to
that on the surfne^-s lietween the ni<jiilliH of the glimds ; very frujueutly, how-
ever, at the louer part of the ^land the ciMlholnini is mndilied, and takes on
certain special charaders which we shall speak of presently an thoM of a
"secreting ■" epithelium. When this occurs the upper part'of itie gland.
I
I
STnrcTVRK or the stomach.
819
I
the p|iitli«)ium it not ■o modifiod. u <>lU-ti spoken of h " the duct " of
\ «tj fnojumtly tkc ^Innd u not tiniplc. but l>rnni-ltc<), nod tlw branching
umy b« ilwlit <^r vxcVMvt. 'iach bnuH-bud glnnds, eapFci>lly thoM in wbi«h
tkc brmnchiug is consi^k-nible. an cAlled eomp«wid gUnda; and in tJiete
ttMn ia Blwayi a wry marked distiiictioo b«(«reen the temiiiuil portiona of
tlra MTViml umncbiiip< vrhcrc ihe epithelial fiella have w«retiDg obaracten.
Mid tW |>rosiinal porttona or ducts wlicr« the cel)» tiave not theae secminfc
dnnwUtB. In eucn a oompound ^laiid n mbubar maiii dud i,n'li(iBu luoiilfi
o(NM into the interior of toe aliaiciitarr canal, and whoee epithelial lintDg
h eMtlnnoiu wilb the smcral epithelial lining of" the csnaJ) dividea, dicbolo-
mooalr uroth«rnwe, into »ec4>ndnry ducts, which agaJD divide into tujoller
dncta, aod iliie divtuon may be r«{tc-ftied ajrain ana a^in ; ultimately, hov
•Wi each duct enib in a part in uliich the epitbeliutu laked un M«reliiig
rhaiWlfri and fuvh lenniual )>i>rii'>n9of ducta which ure j|[vnerally wider,
mora awvllrn a* it nere. ihaii the diieta leading to iIh-ui. and nut infroiiuentlf
flaak-akaDMl, are >puk«u of aa olvtoli. Tlwac ulveuli, tsjit-ciidlj when iUwk-
ihapiil. bear a certain, tboucb br no mean* clone, rrsvmbtanoe to the Indl-
vidoal berrioi tm a bunch or gra|ie#, the duels being tbu branching (talka;
baace, than conpound i^IsixIh are si>okcit uf nx " niccmcaa." 8oineuin» the
glukd io ditidlns apremU <iut luow-ly ovrr ii widv snrihoe — that it to Mjr, ia
"difiMC;" aooMUtnea the ducU and nlrcoli, with all the oonncclive tSmat,
blobdvanb, etc.. belonging to them, are bvntid np tiehlly into a more or
Im etnbalar oum — tiiai i» to mv. >'<>na a " cumpnct ' ghiDd.
Ounda. in fiu't, vary widely in *txo, foiin. and oomplenity, but thoy nil
hsva an* feature in c-ominnn, Itiat they, being involutiotiaof the mucoua
i—nhiiiii,i II ill! iil'ii I 111! f I mil mini eonnecliro (iaaue lined by efHthelium.
aaJ in the luajority of glamU there it a distinction in the chanifltera of the
«|ritWium betireeu u terminal eecreting ii»niuu nud a proximal conducting
portlMt.
When, aa in the Mo<nach and iutettioe. a nuiuWr of ciKiipitntiiveJyaimple
l^aadi are doeelv pa«koi] t»ge(her fide by ude, the whole niiicnun luembnine
■cquiraa proponioDatcly iucraodi-il thickneM; iuM«acl uf l>ciug un alleiiuulrd
.fetnedof a wngr^ layer i>f i-i:IU on a ihiu conncctirc-tiwuc boais, it
BW a naai whotu- thiclciicm i« deiermiim) by the leagtli of the glaDdi.
It Mtv be adtled that genorally, but not ntway*, the gland in lU whole
bwtii Iwi above >ir uoKioe the mUKuhiria mucoan, to that wl>cn a vertical
MtUoo ii made of a mucona membrane the aiiiM«lari« mucuw ia aeeu running
in aa even line at eooM litUe dialance below the thick layer nhich ia |>n>-
Hated bjr the longitudinal sections of the glands.
Beariag in mind these general chiimcton of th« aJimeiitary canal and it*
glaadat we nay now prootM to atudy aome of its Kpecial diaracten, and it
will be coDveDieat to bc^ with the rtruciurc of the atonucb.
Slnuiurt of the Stomach.
( £10. The Homach in \\a uructure fuUufra Ute geiiemi ptau ju*l duwribed,
ami nAtisIa of a luufcular c<>iil and a inucouo nieiiilirauc. «e))araleil from
Meh other by looee Mibmucous eonnevtive tiiwie. The musculur ooat, which
baa ronsMerable thiokneM, conuiirs of au outer, somewhat thiok, limgitudinal
nal, awl an inner, still thicker, circular coat, the innerntost buiidlen of which
take an obliijoe dintitinn and fuini n more or loM dtMinct thin oldiiiuo liiycr.
As wr shall Kr. the nh>vva»ent« of ihe utoniach an tuon eitetiaive and com-
{■(vx than ihusv uf the rvM uf the aIin>outiiry cwud. TowBr<l Ihv pyloric
H As wr I
320 THK TlSiiVtiS AM) MKCIIANISMS OF UIOEf^TIUN. M
t^A. in uhflt W soiiietiiues ciilleti ibo anlrum pyfori, the circular liiver increaws
ill tbii-kni?M, nud at tli« jiylDrus n di^veloftvil lutu u iliirk riiijc. cuilvd th«
8|>liiui-icr of ibe pylurUB; a les§-marked circular agtliiiic-ter it iilio |>rvH-Jit nt
tito Clint hko oriHce.
Tli« eitv of llie cavity of tUe sloiuuch varie* from limo to tim*. nvofling
to llie )iulk of n^nieiiLH [nvdciil ami llif u<iii<liti»ii of tW niiii'ciiiitr |]l>rc«.
When the MUiDiacb is empty, tbo niutctiliir filnts luv in ii ilutr •>( ionic cnn-
tractioD, urn] the cavity ■• »niiill: vbeu ilii> »t«mat'li in Cult, ihv muscrilar
iibro, t1iuu|j[li carrying out, as wv^liall x'r, niorr or httf rhytbtnicnl ni»r»-
menu, are, ua a tvhol«, rvlaxci] and tixtitnileil, <«> that Ihr cnvity \* Inree.
The niucotid nienibniac in Its uaturiil coiHlitinn, sii lo #pcnk, i» of iitch a eiae
tliat it lurnia a smnoth, even liniiiff li> the niiiw.'tilur cont when this is extended
and relaxed and llio cavity dI iTie MtoiriHcli distended. Hcncv, when lli«
etoDiarli ih empty, nnil llm niufculnr cual oiitntcled. tlit- mucoiif mriubiaa«
t» tbniwn iiili) folds or ru;;ir, which, on nii-imni of the ini.-|ionderaiice of ihe
circular miiAculitr ccnil. lake a Wigiliidinal coiirra, the Imse iiibmuojua
timtuc alhiwitij; thi« movement of llic mueomi over the niueculnr ciiac.
The uiiiciiiin membrane is reintively very thick, the thicknew being due
to the fad thiit ihc membrane over ils whole extent it thickly rtudded with
gland*; it may, in fact, be «aid to be almost wholly comjiufied of a number
of shorl, comj>nralively "aimple." elsndE p1iK«d vertically aide by »ide
and bound together bv juHt as mucn connective liesue as serTes lo nurrj
tbe liloodTcwelB and lyniphaiiwr. These glands vary iu aiite, ahane, and
character in diDerent parW of the Btnmacb. and Ihe atomaclu of dtfft-Tciit
aniinnl* jin^wnl in these rea|>ects very considerable dilferencea; but, I'^r
preetnt pnrjiui^a, we may consider them as of Ino kiiidn, ihc glauda at the
cardiac rnd of the alomacb, or "canliao gUuda," and the glawh at the
pyloric end, or " pyloric glands."
§811. Canliae t/hnd*. Thrse are lubnlar glnndH, ubntit O-'i mm. lo 2
mm. io length by hiii' to UKlfU in width. wIhm; [ittirKc t« nut wholly siniigfal,
but wavy '>r gently lortiioii», and fn^qiieutly curvnl or Iwnt nt the Idind end.
fFi)£. 101.] Some an simple or unbniuchttil, but others divide inlo two,
three, or oven more lube*. They are packcit liig^lhcr si<le by side in a ver-
tical tKiMtioo BO closely ihnt in section* of hnnlcncd and prepared stotnocha
in which the bloodvessels arc for the most imrt emptied of blood and the
lymjib vpacCH of lymph, each gland seems to he separnicl from ila neighbors
by Dutlimg more thun an extremely thin she^^t of connective lissiie seen in
Mctions an almost a mere line. In the living stomach, when the nanwmua
bloodTCHclt ill thiit oinnective liune ore tilled wilh blood, and the Ivnipb
•pace* nr« dinti^ndeil wilh*lymph. the glands are separated from each other
by a oonsidcrable Hpace, eifua! probably to about their own diameter.
Tbe outline of each jj;luDd is defined liy a distinct biiseiueut membrane,
which apjipar* lo be formed by a number of tiat trauspar&nl coniuictive-
tinue corpuN'lc* fuaetl tuKCtber into a sheet; iu a seciiou of a gland, longi-
tudinal or iransvcrw, Mme of the nuclei belonging i>i the cunatituciit cvlla
may be seen imbiuldeil, an it were, iu the baaemant luembraiK.
£ach g)i>nd may \k divided into a " mouth," by which it ojwna into tlic
cavity of the stomach, and which rmchcx alKiiit a third or a rtnartor down
the length of tlte gland and into a " bmly " which forms the nwi of ihn gland,
the Jnnciion of tlic two bring calliil ihe " Deck," Thmc two paf1»dif)br
fumlantenlally in slriiclure.
The motitii has a wide, open Iiiichi, and u limil with a single layer of
lone, eleuder, conical cells, enlliil " nmcJiOH wlls," The lower two-tliinU of
eoon mucous cell, including Ihc piiiiucd or hlunC or sametimc> ^tightly
brutofaed end reAtiug on the undi^rlying bascmeol membrane, is comi>ueoi) of
322 TlIK TISSt'Ka AM> MECHANISMS ftr lUOKSTluV,
Tlii; iij3i«!r tliinl \* moiw i-lriir itn<) lmnit[tnrt.-iit, A'X» Dot «uiin r«ilil_T. nn(f
dtAi.-re in ii[if)r-nrniii-a iit iliflrrciit tiniM. At nuc time this part iil' the c«II i«
oocupicil l>y tniini*; at amttlivr tttni; t)ic iniii'iu hiu liwii dUchKrgiil bv n
mplurv of ibr outer fiiw- or lid of th« ™ll, k-aviiig a hiiiaII cup8hft|»H) csivitv
(I'ODtsiiiini; Huid mid u irniiiiuu of iuiii;u« i. the liiirly distinct vaUa of whiili
lire coTilinumm vilh iW protopliumic lower two-thinlB of the ct^U. We eliall
bhortlr liuvc to di^'iiw nior« fully itiv naliire of iiiucnu§ i.'elU in coiinectiou
with itie KalivHry glnnds, and niny here »iruply say ibac in llie tipper tliinl
of the oell the cell-siibetaiice of the cell, exi^epl fur a puftiun which reniaiiit
lu the i-ell wall of this part of the coll, is iraDfifunued tiiln mucus, aud that
the inucuH so foniied ie sooner or later diachartfed from the l-«I1. Iu |dnoe
it^ug \o lime ocoupied by new cell subeluice, whiuh again lu turn u ■.■on*
verted into iiiucub,
Thi»i* Diucuus celU not only line the mouth* of tbo gland*. Ixvnniiug
•Imrlor where the mouth joins ihe neuk, but n\m cover ihn ridgia> l>i;(ni'i;ii
the ^liindi, uiii) *o furu tbe immediate linint; of the iiitrrior of the ntoniiu-h.
Th<- fri'u mirfai'e or lid of each oell is men; or le«* huxot^onal or |>olvgiitinl iit
outline, ut)d in .^ectiunii of hardened Hlonmch tbt; hiinlrnod ccll-viills of the
lope of the d^lln n'lve hm to the uppeuriuice of a moiuiio of hcxa^mnl or
liolvgoiial ureuM where the wcctioii |>r««cnTa a iiiiinlivr of lline ccMt ecvn on
vnd.
Lying between the buw of the miKionit cilli' (which, from the conical form
of Uio cells, diverge from each ••ther), above the hiurnx^nt membrsue,
BUf beSMn in TPrlica) »c<-tioni< u i-i'rtnin number of amnll fell»,i-ach coiiaRt*
Ing of « nurlouB »urrounil«<d by a vcllbxly, whidi, though email, stains
deeply, and hence lioconies coiDpieiious in stained soctioDs. These, as ir«
previously Bai<l. have been regarded as young reserve cells which will, upon
tJie destruction of any of the mucous cells, grow ui> to take their place.
g 218. The hoily of tbe {(liiDd is not only in iueif distiuctly leas in diam-
btcr than the mouth Cso that a larger amount of vascular connective tiasua
li» l>aween (lie boiliu than between the mouths), hut has a much uarmwor,
indeed very narrow anil tortuouD lumen, atid Is lined by cell* of a wholly
different character. Thcte are of two kinilH.
Tbroughotit its whole K-jigth helow the moutb the ghuid i.-> tiiieii conlinu*
ously with a lingle layer of polylir^lnil or I'uhieal or nt times conicftl c«llt,
the outlinvH of which are rr^murkiiUly indixtiucl. The cell-body of each of
these, which contain" n ^phi-rii-al iiucIi'-um jiliii-eil near the crnln- of the cell,
but more outside loniird the hiwriiienl niciiiliriuie. varie*. lu we shall see later
OD.very niLicb in iip|>v«rance according t-o what hiis been taking place in the
atomacii. and to the mmle of preparation. In scirtioni of a elumach hard-
ened and prepnnKl in an urdinarv way ihe crll-liodies fre<|ucnlly pivscnt a
" iaintly granular" ap|>earnnco. Cells of this kin<l are spoken of from their
|naitioii aaeeHlraf cells, or Bomc4imM, for reasons whidt we shall see presently.
mtekief cells.
The cells of the other kind do not form a oontinuoiis layer, but are Mat-
tered along the lenvlh of the bivdy of the gland, being moat numerous (but
gntaller) in the region of the neck, and leas frequent l nut larger) at the bot-
tom or fundus of tbo );laiid. They are. moreover, in the lower part of the
eland, aittl indeed over the greater [>art placed outside the central cells,
being wedged io IwtWM-n tliese and the basement membrane, and freijiiently
caanng tlie latter lo bulge out: they therefore in luoil cases do nol abut on
the lumen of the ^'bind, an>l ihcir only direct connection with the lumeu is
through »iuic(« Ix'twcni ibc ccuiral ''elln. In lite neok of the glaud ihey uniy
however bound Ihe lumen. Kueh cell in ovid in form with an niitliiM
which, ill Contrast to thai of iIk' iviilrul cell*, is sharp and well defined, aud
STBUCTCRK UF THK STOUAOU.
828
n onttd mK-Imn j>l«oH in tho miilille of m mll-bodr which, like
tint oTtbr »ncnil oi'll, vknn in iippmrnncr niri-onlitig lu nrennuMooes, but
wUefa. til a fwclioii of Ktnniiich baMMiMl sii<l [ircparM in Kn ordinnr; way,
b rrcqucnllr " ooatMly " f;ninalnr. Crllo of this kiwi nn called from their
ptiaiUiin ftnrifUtl celU, or from their shii|H-, oi-oid wMf,. ^len tlie smaller of
tbcn an- ItirgtT ihiiti the coiitra! cells.
Acfakriii'icri^iiic " ^nttrio i^IiukI " thrnof (he cnnliMc ref^on of tli« Moraach
i* a labuUr il<>|>r4«Moii, otlen Hittlght am) tinijile. but at tiitiw bifurcatiajf
lovmrd the lower |iart or otherwise ilivirliiiu. llic eods rret|U«Dtlr curling.
Kaeh d^irasion GODBJata of a luoulh. with a hnrnd lumen lined by sleuilor
moawa cell*, a neck in whioli the imicouH eetl» Buitdenly (.'lianxi^ to ceuiral
fW» witb iiunierouB ovoid c«lla lyins aiuone them, and in which (he
lutnm tMM»nw4 oarraired and Inrtuoiis. and a bodyendiiif; iu a blind fuorliKi,
with the luoMtD Mill narnin' winding biKwoen the ceiilral («IU ontdidu which
■rw jilacml avoid evils loM iiumerciu* than in ihe neck. Sneh K'liniU pliiccd
aidr )iy aiito fund the thickiKM of (he mucriuti membrane, nm) below them
at a a^rt diflaiicc run* iti a Uderably e\-eu line (lie thin muiteiilnria muci»»
wHb it» tingle ioiier circular and milt-r lonjciUi'limit tavent of iitain tnuiculAr
6bna.
i Sia. Th« *pM* b«t«Mn the level of tin; bottom uf the glands and the
otaaeataria nueoNS w wdl u> the vrrlii-al njinciM belwMD th« gland*— that is,
all tlw •)!«• bMwwn the raiiel) foldciil ba»cment mom bra ne aba ve and the
■HMeoIarw muouMP below is occupi«xl by d«licati> oitniTlirc liseuo (he meali-
wofkAf which, foTRMd of thin narrow shoetd or himinic rnthcr than of flbrea
or bvudka bMom«» «p«ci«lly clow »ct imme<)ijitcly nnder thi> basement
Willi a nil In the spaces of ihe tncshwnrk » rvrlnin number of lymph cor-
paaelaf or launtcylca may Iw seen. Small aricrir^ ptwin); upward from the
aabwuwa ihnMi)>fa the muHLtilaris muPos» break up into capillnrieii encir-
diof ibe );lai"l^ <'■ >bt^ ^'^t™ '^f plexuses which are especially cIom set at the
Mnrait* tii line spaoes between tlie };landa. thai is to uiy, ut the placM where
tba cnan««tiTe tiKiie Ilea nearest to ihe interior of the stonioeh. Small vvioi
rrin^nj; from these capiltariex. eniiecialty from thuM last luunoil, running
wnward pierce tlw muacitlari» mucosie and form the larger vcii» iit the
•abmuiMU* cout. Lymphniic veMnU and .-itniclurM eallcd lymphatic " gland«"
arv prr*«nt in the muoHi» ciMit. but of thrM.* we iihall ^penk tatvr on.
jl 814. i'^/Urrie ylmid*. At tbu i>vluric end of the Mlomadi th« elaiidti are
Isadaselj packed than at (be cnrdnu' euil,aiiddiMVT from thecarniae ghiniU
fai iter, altaiic ■n<) stntcturr. [Fig. loi] A Irpical pyioric gland poeHwea
a nouth wnidi t« murh longrr nnd gpnemlty hroaoer with n wiiler liinieo
Iba mouth of a cardiuc glnm), though th<- walls aiv limnl with mu-
< esll* like those of thr* cardiue end. The hmlv of the glaixl instead of
bainc. m !» tiM- canliuc gtnnd. »ft>.-n lobular and unhranchol. frec|ncnlly
■livMMa into two or nion- branches c)o>e to the nock, nnd (he»o branches
which %tr rrlnlively cliortcr than Ihc bnly of a cardiac ginnd and have
a iBurh wider lumen, may again flubilivid'- ut thiil the whole ghiod is most
distiactly branched. The whole Imly with all its branches from the mouth
to the several blind ends w lined throughout with one kind of cell only, which
la vwy similar to the central cell of a pyloric gland, inasmuch as it is a poly-
bsdral or sliori columnar cell with indialinci outlines, a spherical nueleiia,
and a adldwdy which in a suetfimen prepareil in the ordinary way is faintly
Kranular. The " ovoid " cell so cliaracierisiic of the cardiac gland is niMent.
Tbe arrangement of tite eouuective tiaiue with its bloodvessel.'i and lymph-
atita asd of the muirulari* muoKue is nmcb the aaiao as ai the canliac end.
Tbus the rariliuc end of the «toinarh eootaina gland* which are tubular
1 oAen simple, which have a very uurrow Inmen, and which potiess con-
3-2A
THK TISSUES AND HKCHAMSMS OP DIQBSTtON,
tTsl uoi} ovoid cella, while th« |>ylorio end cxulainitglaiKlH which iire brniicti^),
ithich have h relntivclj* ducp mouth aiid wide luweii, mid which pueww ihhs
kind of ccIIa unly, <!eiurul vtlh i<r udl* v«ry Uki
f^'^ '^ l)i<-«f. Id the nii'liUc rtviixi of the ftoniHdi ibl
iini- liiiid ol'gluitd ^mdunlly iiicrgrs into tbcothvr:
ill iiiiMiti}; rniiu thi- curiliii to ihv pjlitruN the civoid
cpU» Wi^orh- I<-iu iiuiiivniui' uuil nl hut diMijijiviir,
the niouth bi-ajiiiiif luii^r, th« lunmu wider, uid
the body of Uiu gliiud iK-oamu* more und diok
brand I CI 1.
Thv ithuv« niipplifg n f^nenil (leMriptiuD of the
gHstric glttiidg, but thcM vury in minor characten
utid to n cvrtain t'siuut in distnbuiiun iii diller«nt
animnla; tiiid u* wi- »hall pniiFijtly «cc in all
the glaDd]> vnry in ii>ndiiiun lutd sl> id oppcAnuc
according us dig\'»tion in or hue been goiof; on in
the etoinaeh.
The Salimry Olands.
>i 21fl. The Diructiiral diflereucea botweea tbe
"mucous" uells lining ihe omuth aud the " c«ni
tral " and " ovoid " eella lining the Iwdy nf a ga^i
trie gland lead ua lo inter tliui the fonnor diflrr
fruiu the latter in limcliun ; and wu hare oth<T
evidence Ihat tWu i» so, thai it is the central ami
ovoid cella which aciually oecretv the giwlric juice,
luid that a» far un the gaAlric juice is cuncenied, the
tnonUiM ol' the gUindx Hervt: diieHy (though liie^
niuciona ooIIh bnvv a pnrjxise ul' their own) to coii-*
diieC lo ibe interior of the ntomach the juice
crated by the li'idy of iho gland. We may tlief
fore speak of the t><>dy as the eecreling (inrtiou awl '
tbo mouth as the " duct " of the gland.
This dialiaction betneeii a secreting portion
a conducting |M>rtiun. more or lees obvioiig, aa «
bai>~e said, m must glands, is esjiecially Mrikinj;
in the case of tbe salivary Klu"de. Tbe»e are lovif
lutlons of the (cpiblastic) inncona metiibnuie ol the nioutb as Uie gastric
elandft are involutions of tbe (bypi'blaslic) miicous niumbrane of tbe stomach ;
but, insleail of being ciimparatively simple they are exceedingly brnndic'
mcemoBc glaudH, uud tbe secreting porlion of the gland iii remov»l Ui a |_
diatanoe frum tbe epithelium of tiie mouth m that the conducting |>orlia
is of very great length. Moreuver, nut unly the epithelium lining thi
secreting {mrlioii, but aWi that lining the conducting |K>rli»n ditlers m> cob
pl«t«ly Iriim Ihe epihlasiic e|>iihcliuiu lining the inoutb that wc niay siudj'
ib9 Mmctur* of the gland iiuile apart frutn tlie stnietare of the lining of the
moutb. wbuitc senMir}' fiin(rlii>ni>, in the way of laiclr, fur iuHtiiDce, are so much
mom imporlaul llmu il» digestive t'uuctiuUN ihul we may renurve the study
of its fcttlurw until we comi- l» diul nilh tin- KcnM».
A snlirar)- gliuid, such nx the ^nlimaxiltury, consists of n long main duct
which pUT*uc« lui uudividt'd course backward fur several centiiuetrtrs trom
its onening into the cavity of the moutb until it rencht^ ibc body of llie
E' nii, when it rapidly divides and subdivides into a number of smaller due
oh oi' Ihe ultiinale divisions uf the duct at last ends in a " secreting "
Al'iu-i.ii uuxn nMM A
eiiCTi03'0MiisI>c«"i*ToJl»nr.
m. mouib ; n. nK'lt; tr. •
dMp I^lUOO of > lUbUlV CDC
iniwntMly.]
vbich ia liitnl by n " »wr«ltrii; " opilln-liiiin, ttifftrrent in cliinirter from
iW •ohlMliiiiii liuin;^ llt« diict«. i^iicli n leniiinBl sccnrliog portion J; call<vl
«a imKtiiu. Hataotimtt » duel Urmiiuilcfl in a »ins1v alvonni*. vihwh then
■pprati u It swollen or aom«irhat flack -Ah npi-ii lorminntion of the duct dis-
tintciiiehr)) fmm the iliict by the fliz« and elianicler of its colls and bj the
Okrm«ti>eB» of il« lumen ; but more commonly n duet ends in Beversl alveoli,
whiHt then spiMwr ■• a number of xhort curred Bonwirhnt ewollen tubra.
br«nfhin|{ otF from the end of the duct. AJI the duels and the alveoli iit
which ihej eud are bound up by coonecltvc tia^ue, carryiuK bloiidveweU,
ncrrea and lyn]|>tialica, into a eompact, rounded bul somewhat lobulaled
mam, the kLumI |>n>|>er. Each alveolim, or eaL'h ];r»<ip ^(^ alveoli, and the
saiali duct of which it forms the blind end is aurrounded and sepnrnleil from
tta neiichbiirs br a certaiu araounl of i-oDtieetive liaaue. A number of alveoli
with tAe duel* ^eadiui; to tlieni are bound toxether into a lobule by a rather
larjrer Amount of connective tiMue. (imupi of these smaller lobules are
Iwtiud MtrHber bv onutsi-iive It^uf and env«l<>|ic't by a more distinct ooal
'if that tiatiie. aixi thus form liiricer i<r |>nnmrv lobult^ ; and these lar^r
1)>In>)m are Iraund up to form tliv glund itsulf liy a i|uanlily nf eonnwtive
liaaa, nrhich al«a fonm ii nrufipiuK <>r itheath for the vrhole gland. Hen(«
■ ihln KVlion taken ihrviugh lh« (ilaiid i* w.-eu, when cxamin«<l under a low
powvr, to bedividi.ll by scpla of [;i>iRK^-tivc tlitsue (.continunuiwilb tlii' Hhealh
of lb« tcland. and currving btoodvMnI*, cic.) into irregular arMu. vrhii-h are
■MlcnUTy angular from comprarion. ThiiK- arm* arc w«tioii> of tho prinuiry
liibulea, anil nudi may In' M'cn to l>e nimiliirly but lew distinctly Nulidividnl
lato aUnilar omaller areiu. the »mnlliT lobulot. ICuh of ihetc xmiillcr lobules
will in turn b«eMO to be for the most (xtrt niailo np of rounded boiliex var}->
ing aoraewhai in rise ai>d #hapv. but on the nhole very much alike, bound
tDi^lber bysBtnall anvount of conncctiv* tU8U«; then an the nlveoli which,
Mug diapnaed in various directions and being frequently more or )e8acurve<l,
an out in various planes by tiie section. Where the section cuts the alveolus
UmBSTenely the outline of the alveolus is ciroular, where obliquely the out-
liiw is more elliptical ; a aeciion, moreover, mnv pass through the mere tip
or sido of the alvvolus and so misd the Imueu allo^^ther ; ami indeed many
rsri«d appearanoei nay be prx^wnled. Am'mg these alveoli are seen other
b<idi«>*ora •oniewhatdiHerrutiiiiiWft.ctreular.elliplieal. or cylindrical in out-
liap,orbour-gla»Hba|)tHl,uri^ven irregular in furm. Thene are the small lubulsr
duda cat in variou* pbtnca. Se«lti>ii> i>f the larjter ducte uf various siue mav
also be se«u in tl>e Miiia between the liibul««. Even with ijuitu a low power it
ia«My(a distinguish ueiwecn the alveoli or i«creiingelemeuta and tlte duels.
MhI whtn we cddm> Io ezamine them more elnsely wa find that ihey <li0er
naritadly in *tructurv. Mnrrover, when we eiuniine the three glau<ls. (wro-
dd, nbraaiillnry and (tiblingual, ami tvix-'cially when wi- employ fur the
purpoM diflcreDt kinds of animals, we lind that, while iIk' diicta have nearly
ihaMiRiw structun' in nil ca«««. two kimltt uf alveoli nia^ lie diatinguisbed
diShring from each other in the charactem of the cells lining ibero. In the
one nu«- ihu (.vlls, for nMSoiM which will prcseally ttitfwar, are called mMCmu
trii*. in tlw other serow («Ui. or. perhajw btlWr, owMwinmit e*Ut. In one
([laod all the alveoli mav be lined with niuMua cells, in which case it is callcn)
a " niucnus );Iaiid." or with albiiminouK e<-ll». in which cjuc it ia callol an
"albuminous gland," or some alveoU may be " mueoiM " and olliers "olbumi-
D»iis,"il>c gland Itcin^- a mixed uue: and this distinction between mucous ami
alliuniin'MS olitniiMi bIai> in glands uf Um mucous muinbnine which are not
il(«iin(ily Mlivnry, lor instance in the small " buccal " gUixtn of the mouth,
and in (hr v'lnnds of tjw pulinonarv paatngM
$216. J/iiMMi ^ndM. The submaiilbiry
and of other slruciurok
gland uf the dog is a
fiurly
sae
TtlK TISSUES A.\'l) MEc-HANISHS or tilOSSTIOK.
trpiral mucouii ghaA [Pifr. 10.S]. Tho alveoli of tht> iclantllmrr k kwmI
clml in Jiumi-tdr, but um uii nil uvcru^' iihuiit ^A^. Tnti 'xitlinr ijI' t-noh
tilr«oltia i> ili'-fiiK'il by n itiRliiicl liiiM^nK-iiL nu'nibrum- runiicd nf ii ■lunibvc nf
tlull«iKil onini-clivctiwui- corpuncltv fiiwil Ingellicr inio nithn-t; in n «colioo
the long oviil iiiiclvi of the oiiiiRtitupnt crlU iniiy be wun Wn' mill thi^rc im-
lM'<Idc<I, nn it nrn.', in ihe nmmbriinv. Oui«iilo tlic bnwini'iit nicmbmiic li^,
va i-lxcHbcrr in n mucous mcnibrnne, ihv Ivmph cpnccs of the fiop i-oniiecli\'«i
Umuc
[Flo. 103.
ftmAXiiUHV Oi.txii OP 1 Dno.
■. fBunHM'cflh^ h pmoiilannlp celli : o, ileuilIniiD celli : < trumtne lecllon oraii cicrMorF ilDct
wltb lu pMuUkr Qolnuiiur qilttwIUI n>IU.|
Tbe tjpnce defined by [lie basemeDt membrBoe Ih nmrltr wliAtly fi\M. a.
vury siuall ceuiml luiii«n only bang left, bjr c«ll» arraiigid for ibit lunvt t>urt
in B tiiu){)<- layi-r. Tli<- rclU are large relnlivek to the rIvciiIuk. mi ibiit in ■ '
iniUHVi-nK- si-i-liuii of nil iilvenlun about firo or nix eell« irill be awn. KncEl
crlt In luoru nr i(^ MilKridd or riilhiT coiiickl in t'orm, with it« broiuU-r Inw,
which i* VDiiiotiiucii irregiiliir in outHiie, miiiig on the bavfinunt menibnne
and ihe narrowtr a|MUC abutting on tho. liimrn. Thi- ehnrttclcnc "f the cell
difltr aedirdine to thi> condition of llic gland. If (be ^Iiind hn», prrvioiia
to ili prvnantion for exiiminntion. lu'l hi-en nctirely recrt-ting, lh« crll* have
certain ciiariict«n, nud innv Iw i>)ioken of a« " loadctl" or "ebarccd." If
tbe gland bas boon flclircfir ^ecreling. theto i-hnmclprs are replaced by
nilKn. and the cells nmy be tpokca of ns " iinlnaded." " dlKharged." In tb«
" loaded," or as it !■ often called the " resting " pba^, the cell, in hardened
Bpeoiniens, is as a whole trnneparvnt, and stains verv slielilly with the ordi-
nary staining reagenU, Tbe nucleus, whifli in haiaeued specimens npiwars
ditw-eliaped and eoineliinea curved or bent, but in the fresh living coll is neeo
lu be splierical. lies at the bare of the cell not far from tbe hiiseiuent mem-
brane Arotiud ih« nitcleus is gathered a email i|UBntily of Drvliuary pm4u-
plamic cell -Hubeiauce, Maining r^mdily uitli tbe usual dyva; tbe reit of tite
cell-body eonabia of a tramparent nialerial. which dora out stain readily,
and wbicb oceupiea the anavea or mt^lit^ of a v«ry ilelicMtc me«hwoTk c<>n-'
linu'iUf, apiwrciitlViWiih the (tlaiuiiig pnit'iphiMitiir <M.-ll'MilBtance around tlie
nueU-UK, and with a thin *be«t nf >iimlar miitcrial forming tla* wall of the
THR SALITARV GLAJtDS.
327
I
I
cell. Tliia tnotfittKnt inaterinl h cillier luiiviii, uliicb wc hnvo teen to be
« n'iw]McuauBConHilu«ni nf viilxuaxitlarv «nliva liii llin >l<ig), om •uhaUncc
whk-h Mn easily bv ciinvcrt«<l into ncttiiil miicia, tbal i» to nv, an siilccodeul
hF luuciu ; lienve tbL> oatuc " mui-ous ciOl." A nttling nr Imdvd mucoua ooll,
UwB, coDUiU [ar}[«ly nf mucin (or ii« nnu'coili-nt ) li«)j.rri] in th« mwliM of
t^ piniot>laiiBiic cvjl-*u))MtNnci.- vtliiel) nvvr iW grciitcr pnrt of tfao ccl] dxiita,
in ■ banlriiiHl ^liiuil iii mir rate, a* ii lii-licnu- m«»hworK or roiieiilum, biit '»
gktlwml inii> 11 i-i>in|)Mrl iiihi' in n ^nialt nrc« immt^lintvly nroiitKl the iiucloun.
Id manv alveoli n moro or l<9« triKiicuIar »pnec k-fl l>et»«%n thv diverging
IwMi of l«<> of lliv mucwii <.iOIh nii'.l [no boM^mcnt ni«inbnine niuy bo wen
tn be <ic«upi«d bv irDi> or hjr two or more pecnlinr small cells. Thf«« on
vzuniDtttioD an <ouimI tu W irregular in form, but oileo half moon -sh a pod,
and >rr Imimm called liemUune cell*, Kocli consbts of deeply eUiiDiDg ccll-
Bab«t«DC« with b Bpherii-nl Ducleus, From their tiie and their slAining
dMplv. M veil M from their position, these demilune cells cuntrast sirongl}-
wkn ih« mucous rells.
In the "diKbarf^cd," or as it ia often called the "active" phase, the
nacoiH mII has a different a|i|>euranoe, aipecially if ibf activity of the
gfatml has been irreat. The i-ell in i»ow amalW, ami Ihiu fr<ves rioe to a mora
oiMtnet lOBieB in the alvoi'lun, n liLr;:i-r |>orlii>n of thu cell (tiiiiu, especially
Ml iho miUr side, attd Aui»vtimi.-» l}i« wholt? itII i>taiii!>; the iiucleuH, now
spherical even in bardeiiLtl ■(lefimtiis. iK.-cn[ii('9i a mtirv Miilnil poHilioii. The
irBiMfiareDl, nuu vtainiDi; mucin hiu in Inrgr part tir wliolly ili.->ap)Kuired, its
ptaw luu b(«ii taken by onlinnrv Ktnintnu; prut'iplniiniic c<'ll-i>nhilance, aod
tba diatiuclion lH:t«<^-n the cicmilunti cvll< iind the pri>i>er gvIIn of the alveo*
liw it nueh ln« diiiinrt. Wi- »hnll prtaeiitly have to aiacuai the iiatura and
aw atag of lhi» chungr frnm ihr Inatlcd to the di'chargad cell.
S2I7. A (mall dun of the «iibniiixillary gland, even when cut tnioRvcnwly
in the section bo as tu pmeiit like many alveoli a circular outline, has an
a|ip«aninc« very dil&rent from that of an alveolus. The duet is lined by a
mjtla layer of epithelium, but thew are slender, narrow, columnar cells,
laaTiDfc in tli« centre a relatively wide lumen, and the outside of tlie duot is
BOt so sharply defined by a coiispicuoue liai»emeiit membrane ss is the case in
an alveolus. Each c«ll, whieli iK-am an -u-al iiui'kuH plat-ed vertically in
the cvW at sImui the middle, but milicr near the huitc, consists of a prolo-
rilBitiiic cvlI'MibMauce which «n th« inner side of the iiudeus towaixl the
umi^n has no sfwcial fi-alnr(?>. hut on the outside, toward the basement
nwnihniue or ooiinectivc limue Iwvi*, haii fn-qticnily a i<digiuidiual striaiion,
as if niB'te up of a nuuilx^r of rmlK or narrow jiriMiis idaoeil xide by Md&
Thr InrEer ducts running' bi'lw<x-n (be Iot>iili-i> iHtb-r from .lucb a Kmall
inira-loUilnr duot vbiefly in the gn'atcr thirkncxs of l.hi.- (.-i>unt-clJv«-UNMe
ba*i». which in tbew is develojH-il into a iliilim-t coal coniaining In the caw
of the larger brnnches and the main duel niaiu muoniliir tibrm. In the
Dkaln duct and it* rhief branches the single layer of columnar cells is
rrplanril br two »r three lavvrs of aibicnl or soniclimcw ttatlened cells iKrt
marknl wilb thcslrisiion spoken of above. When a small intni-lobular
dud is about lo end in an alveolus or a group of alveoli it beooincs nar<
mwn), the cells [iwe their striaiion, from being slender and cylindrical in
form become sbori, cubical, and st tlic very end of the duct chnnge inhi Hat
•iiitidU- (hDiivtl plates, the tranntion front which lo the cha met eristic cells of
inr iiheidiis is in ihe esse of meet animals <)uite abrupt. Such a UMKlilied
lerniinal [Hirtion iif a duct is sometimes spoken of as a " ductule."
\ 216. Albuminuua 'tlaniln. Thoe differ from the mucous glands in the
r>iii>tiiutvin «f the cells lining the alveoli, but the structure i.<f tW duels and
iho gvneral aiiaiigemeitls of the gland are the imiue in both ; indeed, aa we
I
S*23 TUE TIBSUSS ANIi MKHH.VNISMS «r UIUBSTION.
Iuiv« already aaid, !n the snme gland some alveuli inmy be albumiDoii
othere inucouB.
Id ai) ttlltuminouB alveolus the wIU are ratber smaller than thow tu a
loiuli'J muci>iiB Klaii'I. a"d their oiitlitiee are ratber mure angular. In each
tell the uucleua. which U spherical, U placed near the centre of the cell, but
rather uear the basement ineinhrune, and the eelt-Aub^tADoe, which has the
general appearance in an urdiDary nrvparation of aciuiewhat densely granular
pnituploHni. Htaiiis readily and uniloriuly all uver. \o cells <yirrea ponding
to the demiliinea of a iuucouh alveoluit are nrmwut. In Tiuit, an albtiminout
c«ll dwa not at 6nt night apftoitr tu ilifler niarkedlv from a iltwharged
mucous coll, and iloea nut Hhnw the .lamc tiinrki-il (liHvrenoM between a
loadei) and a diitchargei) oinrlitiun an dcjnt a niucou* cell. There arv, how-
ever, difli'renout betwoen t)ie loaded and di^iHmrgod albuminous cell, bnt to
lliMc we shall n'tiirn prratc-iiily.
The parotid eliin<l nf mnn, and indeed of all mnmmalM. is a wholly «lbu>
niinoiis gland, though in the dog n fen* cells arc mucous; the siibmaxillary
of mau is on the whole a mucoii* glutid, bnt some tobulea in it are albuminous ;
tbeflubmaxillary of the nibhit is an lUbuminouB gland. The sublingual may
perhaps in all mammnle he regarded as a aiucoub gland, though it diifbn in
aereral respects from other mucouB glands: the L'ellfi lining the ducts are much
shorter ana lees distinctly striated, the alveoli are more obviously branched
tubules, and llie cells of some alveoli contain no mucin.
The Braall buccal glands which lie in the nuWance of the mucous meia-
brwie of the mouth, and whose secretion eontributej lo " mixeil " saliva, are
formeil on a small scale aAer the plan of a salivary glan'l— tlmi ia to say,
they are cunipused of a duet (or ducts) and alveoli which in ntructure are
ximilar to thine of a salivary gland. They furiher rt«emhle the sulivary
glands in that some of ihein are "albuminou*" and some "mucous."
S 219- The salivary gland:" have each of them a "pccinl nervous supply of
which we shall spcalx in detail in the following section, and will here simply
say ihnt the librcs pawing into the ^lnnd» are both mcdullatod and nna-
medullated libree. and thai the terminations of the fibres have not been as
yet exactly made out : for, though it htis been nmintained by some oWrvcrs
that tome of the nerve-tibres end in (ho secreting oelli^, this has not been
fnlinfttotorily proved. Numerous nerve-cella may he seen scaliered along
the nerre-fibrea, where they paaa into the glands at the " hilus." whence the
main duct iiuuee.
Of the n<TVOiis supply of the stomach, derived tiartly from both vagi
ncrviM and pnrlly fr>iiu the solar plexus, we ehall also have to a|wak later
on : we may here ximnly say that tnc fibres end for the most part in a (ie«u-
linr plcxuR Iv-tween the circular and longitudinal muHcular layeni, and Jo
anolhvr {n^culinr plcxu* in the submucous wM., the two plexusuct oorrc*]M>iMl-
ing lowhni we ^linll dc*cribe in the small intiwline a« the plexus of Auerbach
and (he plcxu« of MeiRinej-.
§220. The stnicture of the pancreas » so similar to that of a salivary
gland that, (hough we shall not deal with the pr»pcrtie« and characters of
Uie juancreatic juice until later on, il will be convenient to consider the
hisloln){v of ihe gland now.
Wlietlier as in man, in the dog, and in most other animab it forni7< a
compact mail*, or as in the rabbit is spread out into a thin sheet, the pan-
croiu is in all eoMa a cumiiound racemose gland. coDBtsting of <luctA and
alve(di arranged in lul>eA and lubul«a. [Fig. 104.] In man (he smaller
A
339
daeu Joie one bmuii duot. nhicti. runoinii leiii-iliwiM- thr>iiK)i llic iil>'")<
pi»fw dw OOMla of tbe dutMlmiiiiu in cimijiimv nilb hm'I ojiL'tui inU> the
wUrioroTlhe iotcstine hj an oritirc L-onim»n loit bikI id Ihf Inlc >luct. Nut
iafrai|Uetltl<r ■ Moond bui Hnuiller main iluot ivirDiHic rr<iiii llii' tiiwi^r piirt of
lb* Mad uf the f;laiKl join* the int(«tiiie luutr ilnun; in the dog mvh n
«*t»ilid dptt is* i»uhI •M-nirrvni.-v. In the rabbit (he miiiii duct dot* nut
join tin inirttinv nilh lli« l>il« dtiL-t. but «l ii ninnidcrahlc dUtCDce, •H^vt-nil
<«alttB«lfT*. la«or dnwii. »o ibnt in ihiK nnimnl iIh- hibt nnd iiKncmiit- juivn
■r* iKit putiraJ iif]^ilM-r iiiiu ihi* iiit<^ini*, btii ihc fix»i in for a diNiunct- expiwicd
to llw KtNn of tlit formtr bwfun,- it uiii-iii iriib ihi- Inller.
{Vm. IM.
Sn-niw V* niE r*>caKu or iiii Inn.
4. Itnnlniulaii at ■ ilati to Ihv lulialai *Itm<I. ■>'•■]
li« HtrucluiT nf the ducts m. in all ew«niial p(>inli>. similnr to that of Uw
vbct* (if a salivary );hind. tjive that iho Mriuiion of ihi- eipilh«liiil crils i* lea
4i«iatl. Ai in ike cue of ibe Mlinuy );land. Ihe ductule, or unrrtxr Icr-
■inaJ nortioD of tbe duct, jou m it JINM the alveoli is liiwt bv llul »pindU>-
ihuvl cells.
tImi alvtoli also are •iiuilar to tlioae of a lalivary tcla^d mv« perhaps that
they an nUtivcly lonni^r and more tubular: tbe Inmeii in all eaaea ih vcrj
nartnw. A* cuniiinrcd with n salivary glaud tbe alveuli art- relatively mom
Daneroua tlian the duct», *<i that in a neotion of Ibe gland relatively fewer
ducta are area cat acni«L l-^icb ulvcoluit in liued with on« kind uf cell only,
which >* much murf «imiltir to an alburoiii'vut than to n niurous cell : there
wa no ilemihine rellsL The more minute luaturM of the alveolo* differ ac-
ennling a* lh<' iflatid has In-en "rctiling" and an in " taa<)<<d." or han beca
''active" niid •■> i* " dMrhargvd." The wIIm lining tlte alvouluik are more or
lea* polyUnlml in fi>rni, and each cell cwumhIh nf a olwr tnuu|>nreni cell-
badr.in which mx-nra niim)>er i>f n-fraelirr divcntte "gnnulM;" a sphorical
•acltu* lie* at alKuii the oiiier ibinl of the cult. In a " load«d" oell ibeaa
gnuHilca ar« vvry abtindnni. and reach froni Iha narrow, iiicunffiiiiio^u lumeti
M Bear ih« oalrr mari^n "f the call, au as to leave only a narrow dear traaa-
parent f><ne imiiiediuiciv bordering nil tbo baa>Di«nt tnemhrniw; tha c*ll-aab-
KanM- is so ihicLly 8liiil<ltil with tbne " f:T«iiiilfa " that the nuclmM b eona-
ptelelr bidden, and th^ grmlcr jiart of tbe cell apfwara quiti' ilark. In a
" dtBcnargtd " cell these granules are far km numercMS, and arv Urgrlr con-
Inad to tbe inner pan of Ilie cell abutting on tlw Innx-n. so that liu-m la
«alablMbad a clear diMtinctioD betireeu a narruw iniHr " gninular " ton* aiid
330 TRK TIBRrEB AXn «BCI
VlOKSTIOS.
a dt*r traodiiarent outer xone, free or nearly free ttom gnmulee. The widtli
of the frranular loat; varieti in (nut with the condition of lii« gland ; when
the ^Inncl hua bem very nc-iivt? llie Kmiiular zone is very oarrow. when mod-
vrnli.-l;t' uitlvc, it i^ broiiilirr, iiml nhen the )(lan<l has been for amne time
whiill)' vt n*t nod i* Lhi-itifnro lomlt^l. th«! ){runii)iir tone inuy eDoixMoh on
ueurl}- the wholly Cull. But wc»liall have to return to then; inalten pre*-
ently.
In thd paticmu 'if lh« rahhit and 90mt other animal* sroupa of edia of a
pocutiiir nnturr may he nva iiiii'r(-aliit«d at tnter^'iiU in the midtt of tlie true
glitndithir giiluitniico. Thoiw arc rounded or pulyhixlrul in form, aad have
a cU-ar ■.-Hl-'iibstaticc with a relativclv large nucleus ; they do not fortn
nlvc><>li and lh«v have no (hict«. Racli of ihcw; gn>uf)8 i» mipnlicd with
hIootivvM>«ls formint; n caniltrirA' nelnork more closely net than i-WwIutc. The
exHcl nature of thcw cgIIm if nt pruent n matter of douhl.
The |iiincrci»s in supplied with tierveB coming tr^im the solar picxuv, and
coneisling partly of mcditlhted and partly of non-med ill tilled Rhrva. An in
the ca<e of tlm salivary glands nerve-cells are Ibund in cunncction irith thv
nervp-fihres ns these pnss into the gland.
The Strwture of the (Knophii^/iu.
S S31. In the general plan of it» xtniclure thn (fsiipI]H|.'itEi re-iembled the
real of the nlimeulary canal, for it consists nf a muc»u:i iiiemhraue. with a
niunciilarii) miucomd and glanili*, a loose xubniucotie coat, and a muscular ooAt
Cunipriviiig an inner circular and an outer longitudinal layer. But tbe
tfiithelium, ooiblcietie in origin. \f very diflerenl from that of the ri'madi or
intevlino, and hoth circular aiul longiludinat muscular layer* ar«i ooinpuaed
U) a large extent not of unstrialed but of striated lihrt« like thon of th«
skeletal muscles.
In a veriif-nl aeetJon of the «tH>phat;us it will be seen that the epithnliuoi
is aot arranged as a single layer of cells, but is several cells deep. The
lower oelb near the basement membrane, which is not very distinct, are
cylindri<:al or spheroidal cells with granular " prutopWmic" cellsutwtanoe,
hut tliiiae nearer the surface are more flattened, and the uppermost cells are
mere llattene<l nucleated waU^, the bfidies of which are no longer prato-
f'lasmie but have become changed iniu a jieculiar material. Such an epitbe-
ium is called a "Airiiiiritnl" epithelium. A nimilnr ejiithi-lium lines the
greater part of the pharynx ami the month, and \k conlinunu.H with the cor-
Rfponding epithelium oi' the Kkin or " epid^-rmiit " of whi>'h Me .-vliall fauve
to speak later on. At the cnnlino nrificr ihure is a unddrn tnitiiiitioo frum
this xtraiifiei) epithelium to the gastric epithcliutri previoiisly rlc^crilied.
The loii*en««< uf ihr ■ubmuous cnat rKtmiitx the mucous nicnibmiie to be
thrown iniii temjKirurv longitudinal folds which disappear when the caiuil ts
dinleiMled. But lj(«i(i<v this, the line of The bnsement monibmur, of the con*
nedivc tiwuc liunis of epithelium, "dermis" or " cormm " as the correspond-
ing part of the skin is called, is raised up into a number of permanent coni-
cal clevulions or fmj'ithr. in which the conneclivc tissue is (Specially Hue and
and which are nchly provided wnth bloodveseels. The surface line of the
epithelium docs not follnw the inc(|ualiiiee ot the dermis proiluc«d bv these
pRpillav. but remains Airly even. In the presence of this papillw the mu-
cous membrane of the (esophagus also resembles the skin, but in the latter
Blnicturo the papillie ar« more atuiudaut and more regular in form and siKe;.
The dermis, or connective ti^ue basis of the epithelium, is a network of
Gbres and fine bundles of euunectire ti.-ctue, with connective tiMue corpuaclea
ami a cuDbiderable number of flitc elantic lihre*; tbe number of leuoocytea
m
TIIK JiTHL'CT
tiinbtv of itip iH-in<itlc u ix'lutiv<<lv >cnniy. A fw >ciit(orcil nuiMM
f'lrni ur niltriiiiiil limiii*. of nhidi no oUitll ^iciik liitor od, ooctir li«rt'
■ndlbera.
TW nneotw iii«'nihriiti« proper is defitwd from lli« undrrljriDe ruI>riuixiui>
time l>* ■ iniiM-iilurU miicoHP of pUiti anMrialed niuM-'ular fioree. Iviiig nl
•rail* diaUnci- from itie ejiiiUeliuin. Tbne rouwutnr fibres are alMciit «t t)H>
Bpfirr i«rl of the cEaophaeus, a|ijJi-iir lower down in uulated longiluiliDs)
MDclln, «ii( nronlusliy &rra a <Iiatii)rt Inyer, whiuli, bonvrer, is ni>l to
r*gnW H in tlw rat of ibe alimi-titAry <!aiiiiI. an<l rotiiitU of lonj^iludiniil
Ghrrc only, cir^-ular fibres beittg nbuniu'
Id duui « few liu( iu ullirr aiiimsia n ciuifirl era lilt- iiiimlier of small '* mti'
cmh" snd "fllliiimiiioii*" gland* nrt Cwiad ill tlw subinutious tissite : their
duals, (wnciniiini; tbi> miwi-ulsrie miRinie vbcrr pmmt, opeo on to the
•urfcer »f the niunMi» mrmbmoet. In mnn iind maiiiinnlia ibeoe it'an<k
■pp««r Itt »ervc nuly the purpose of kecpinji ihe iiitemiil mirfaee of ibo
mopbami* moid ; hut in Mine aniiiiale. a« in the fro|;. i» which th<^ epilhe-
thm nf tlw (nophnL'iii is not the mnny layered almtitied enitlicHuiii JuhI
4aRri1>rd, hut n ciii);1e layer uf eoluninaV eijinii'd cells iiiii<Mi with muctiUS
ealW. uf the kind nhicli w^ shull later on deMrribe m "goblet " c«ll*, there is
a Urgv dnvlopim-nl of ;;]nnds at th« lower pari uf the oesnphaguv, nod the
eHb of lh«w glandf nMiuuruciun! pvpin.
.K» in 'ither parts of tlte alimentary iiinnl the *iihim)cotis tiam« ciirri<9 the
larscr hloiKkeMelB whose smallL-r bmHchc* riippiv the raueous tncmbriine ;
aM lymphatics, beginning in the lotirous memlirBn<', fumi considtfrsble
plexuNv ID tlie Bubmueoua coaL
i 222. In man Uiih the ihtcker inner circtdar nod the oiit«r thinner loii^-
tndtiuil muscular laver <»nfin in lh« up)>or part of the <ewi>nhagus eielii-
iiifly of bundles or strtiiiMl lihreu. wliich in their main cnancters an>
idc«t*r«l <*ilh nnliiinry Rhres of skeletal tiiUHeles. At about the end of the
opprr third or rauner, bundles of plain uii^triiitcd fibre* make (heir appear-
kbce amoDg lbs bundles of alriat«l fibn-x. and a little tower down the
striated flbna disappear, to that, In the li>wrr half or nior« of the tube, Wh
rirrular and li>nf[iiudiiial Isyerx are compownl idmoil exoluiively >>f plniit
ni»lri*[<i| lib res, a few itmy bundlni of Kriat«d muscle heiag loond hvre
•nd ibani. The relati>Ki of the «trinl>.il and umlriuli-d fibres dilleis. Ix'W-
rwr. In diflS-rrat animals; in sfmie the st(tiiie>l tioiue raaehesduwH ncjirly
to iW Mnt»Bi-h.
Above, both iMigitiidiiwl and drcular Inyere merge into the inferior con-
atridor of ibe pharynx; Iwlow, the lon^tndinnl bundle* spread out in a
fwttsl ftshioD (o joiD thv corrttspondiuf; Ionj;itudiuiil niuKulnr iHHit of the
stosnaeh, aad the circular fibres are also continuous with thi' circular and
vbOqat layen of ibo stomach, more especially with the latter. Before the
fimtUr fibres idw apreml out over Ihe ttomaeb. Ibey undergo a tunievrbst
iDpri»td development formiuK >> *"rt uf sphineter of (l>e rurdiiic nrifit'e.
OnCsidc the longitudiuul niunrtilnr m>«I of the aw»pha;;u^ iher^ is a oou-
Marable develupim-nt of ciinn«^-ti\->' liwui- fimning what is tometiroes spokeu
of aa a fibruu* slteuth. In. or mtln-r |MThii]iH »n, this sbeath in thr |f>wer
part of tba CFa(M>ha)[us run the two i-stfi niT\'<«. with the miophaireai pU-vus
■ hieh is furnml by branchi-* ruiming from llw one to the otiter. In it also
nin the larger bliMKlrnwIa.
t 2S3. It in obviouH ihni (Im ivsophaetis is much more a muscular than a
Hotting slrui-ltire, und I'liiiher thai a Ji»tiui?ti<>n t* to lie modi' hi-twfi-u ihe
apfKr [art "f the (i-*"jplingun wIhtc ihi' niuEcnliir tihr« an* »triiil"i. and llie
lowrr part wbeif llwy arw unflrinicd. Corresponding more >ir if" clearly
to this distinrtkon, «c find that though the «hul« osuphagun it supplied bj
i
SS2 THK TISSFE8 AND JTECffAyiSMS 07 DTOBSTFOJT.
nerve libres from ihe trunk of ilii; vajiua (n'tiicli, however, it must be remem-
bered coiitainH besides fibres of the v>i|;im propur, Rbrea ttata the apinsl ac-
eavoty wcrve and from other source*) the aiip]tly to Ihe upper ]iari takes a
diReifiii courae from ibe supply to the hiwer part. Thiu id nuui tlw upper
part is siiii]>lied by branches of Lb« recurrent liiryngeiil nerve ta it runs up
between the trachea and OMUphoKUK, wbilo the I'>w<t {wrt derives iu nerve
fibres from the iUKi|ihaKe»l ploxu* formed br tJio two vi^ri. In varinua
aninialK tbi? supply uf the upjirr purl viiricn, eiiminK in tu>nie cuea chiefly
fraiu the phtirynK^iml hrnin-h nf thv viijeii*. atxl being in tbn rnbhit a dittincl
branch nf the vnpi*. hi all vn*v*, however, it notild M-eni llmt the Inni^r p«rl
of thv inDphagiis, the up[>er limit bein^' plitced higher <ir lower in dilfiTent
animals, is supplied I'min ihe (pnophngiiil plexus. Il mny be ff^nmrked (hat th«
fibmi in this pkxne arc for ihf mom purl non-niedulUteil fibres, but wc -hall
have to rvtiirti tu tbeiie nerve;^ in !>]>eAkinguf the movomeni«of the oimphuguM.
Thi: Act op Sgcretiok op Saliva a:cd GAffrnic JotCB asd tbk
NkRVOI-^ MEDRANIHMit WRICR ReOULATR IT.
!; SSI The saliva and gastric juice who^ properliist we have studied,
thouch »o dilTereiil from carJi other, are both drawn ultimately from one
common source, tbu blotHl, and they am rKiiircd into Hic alimentary eanal.
not in a ountinuuus flow, but intermittenLly a» uccaaton may demand. The
epithelium cellii which nupply ihem have their {leriods of ravt and of activily,
and the amount an<l <pialitv of the fluid* which ihoto cell* aMrete are
determined by the need* of the'eCDUomv n» the food pasws along the eanal.
We have now to e.on»ider bow the epitWliitm cell maniifnctum' it* vpecial
Mcrctioo out uf the maleriald supplied to It by Ihe blood, and how tlio coll
ia called into activity by the presence of food, it may be aa in the case of
saliva nt some distance from ilaelf, or bv circnuuitMiOM which do not bwir
directly on itself. In dealing with the^e matters in connection with tJw
digntive juices, we sliall have to enter at some length into the physiology of
aeeretion in general.
Tbe queslion which prceeiilfi itself flrel ia: By wlial laeohanism b the
aclivtiy of tbe tecreiiug ccIIh brought into play?
While fluting, a iinall ijumitity only of saliva is jmured into the oioutb :
the buccal cjivity i* just moiat luid nothing more, When food Is taken, or
when uny napid or slimuluting ftiibmaucv. or indeed a body of any kind, is
introduced into the mouth, u How i* induced which may tw very copious.
Indeed the mianiity necreled in onlinary life during '24 houn hai been
roughly ealculattyl at lu much a« from 1 to 2 Hitri. An ahuntlant McretioD
ill the nhitenee of (bod in the mouth mav be called forth by an emotion, aa
wbeui the mouth wniern at the «ght of fooil. or by a nmull. or by events
occurring in the stomach, a* in immc cumw »t' naus<-a. Evidently in iheae
inataneee some ncrvoiii m<.'ehani«m i» at work. In studying the action of
this nerruiif mechani*m, it will be of advniitiigv to confine our attentiua at
fint to the aiihuinxilliirv gland,
$ SSS. The uibmaxilliiry gland is supplied with two set* of ncrvM. These
are represented in F'tg. lOt, which is a very diagrammatic reuderitig of the
ap|>cHninc«s presented when the aubnuixillary gland is preparoit tvr au
expt-nmuiil in a dog, the animal being pliiced on \l6 back and tbe gland
exposed fnni tbe neck. The nnv Act, and that the more imp<>rtsnt. belongs
to tbe cAordu timpani nerve (ch.l." ). This U a small nerve, which branches
ofl ftooi the facial or seventh cranial nerve in the Fallopian canal before lh«
nerra isBuee from tbe skull. Wivitiber it really betougii to Ihe facial proper
AEUBITIOX or SALIVA ANIi O.VSTBU' JITICK.
uarlio
oomilig
Art
•a (kmbtvtt : in nmn iIk- fihrm which r<'rin
m tb» root* »)' ttii: fkrwl pmpvr btit Croin
m, nr. «coonJiD^ Ui maw, lilirc^ which tbixigli joining tliu t'iK'inI in lh«
FKliolMJin {«ii«l Bra ultinmiflv Hrnvcd tnim aii'ithcr ^(liv Kllh) i-rnniiil
metyr Lr«Tiii|! the fm-inl nirvc, thp chonlii lynipiini pants through llw
tjiuiiKnic ixviiy or ilnini of \hv rar (h<^ncv (he onnie) Ui<l joins or mthi-r
rani in 00Ba|Httiy irA.i') wilh ihe lin^'tiid »r j^iieliiliirv hniich of tlw SAh
Mil n Somu of the fibres run on with ihv liogunl riuhl down (u ihe toogne
(tk«n Ut tMit »bowD in the ligur«), bul tunny Irave llio lingual as a steniler
ti«m (ek.t). which reaching Wharton's duct or duct of tnosubiDuillary
fl*aA Imt.j) runs along the duct lo lh« gland. Ab tli« nerve coursea along
llw dun nerve cclh make their appearance omong the Hbree. and these are
iwcially abundanl just aAer the dui.-t enters the hilus of the glaud. 'I1ie
fibfw mar Iw traced into the jtland fur eonie di»ianoe, but as we have Aaid
tbilr vllimals endiDK baa out yet been defiiiilely made out. Along icn whi>l«
eoniM up to the giaud, ihe fibres of Ihe churdA are very tine medullated
fibfw, but they li»e th«ir medulla in the gland.
TW oihvr wi of nervi^librea reaches the gland along the small arteries of
tba ^atid. Tliesc are uu«-ni«du Hated fibres mixed vilh a few medullaled
fibro add may Ik imreil ImicIc lo rhe nuiieriur c«rvii.-a[ itauglinn. From
Ihsnn tbcy may be traced »till further buc-ic duwu the corvioulayitiiiiitbetlo
M tbesi^DBl mn), rtillimiug appurcnllv the *ame tract as the raiiu-cunatriclor
ttres.tnaivduf in M)i4>.
I tt8. I( n lubv \tf pluc-L-d in ihe duct, it is teen that when mpid >ul>-
■bUKVS arc idaocd hq ihc lungue, or ihe tiiuguc is Mtiinulated in any olbvr
way. or the finguitl ntrvc i> laid Iwrc ami «limulntcd with nn inlcrrupted
ct)ttt«(, a <>>)>tuu* tlow <jf Mlira tuko phic«. [f ibe syniiiatln-ttc be dividi-d,
atinulatioa of lh« tongue or lingual m-rve niill produces u flow. Bol if the
■nail clKirtla nerve be divided, stimulation of the tongue or lingual nerte
pradttcn do flow.
Ciidently the flow of saliva is n nervon» reflex action, the lingual nerve
HT^ing sii Ihe irhaiinel for lh« adt-rent and ihe small chorda nerve for the
tffuCDt iniiMilscs. If the trunk ut the lingual be divided above the point
■ban Uie eboTda leavM il, as at n.f. Fig. lUo, stimulatioD of the (front part
of > tongue [irodncee, under ordinary circumstances, no flow. This shows
that the ccnttv of the reflei action h higher up llian the point of section ; H
lias in bet in tlie brain.
In tbc angle betwcoi ih«- tingnal and the chords, whero the lattsr leavve the
fiWIBSV to pass Ui tbc ^liind, lira the oiisll Kubmsiillsry ipinglion (rc)ir«^nled
dli^^BiiBalically in Fig. I<r> an.yl.). Thia ciinsUtji ornmall nmises of nerve culls
Ivinf "n ib« niiall buadin uf nerve fibren which apreail out like a fau from the
ItDcual anil chorda tyin|>anl nervM (cA.!.) lowsnl the diictAof tli« Hibcnaxlllary
and sublingual gtalxu. Il biu> bevn much ilvbuicd whetbrr thi* gnnffltoD can avl
■a ■ <'rDii* of rrSez action in cxinei'tiim with Ihe aubtoasillary gliuid, bul no
eonrluuve cridcDce that il does wi aot bte as yet been abown ; It probably belong*
In ri-ality to the MiUmeual gland.
Stimulation of the glofaujtharyiigonl is even more cflectual than tliM of
the lingnal. Prol>ably Ibis indetxl ii" tin- chief alB-rwit nerve in 'irdinnry
arcrvtiou. Stiniulatiou of the mucous menibriuie of iIk; Momacb (m> by r<H4|
iatrt>(luc«d through a gastric fislnln) >'r of th<> vugu* rosy aim' pnxluiv a ll<>«
ofBallvB.!* iiidwd may ftimulalioJi of the sciatic, and [itoluibly "f many
other afferent iwrve*. All tlieso oosos an insiauu« of reflex acliun. Uiv
oerebro-spinal syetem acting as a ceatn^ Wc may further define ihe ccnirv
as a pait of tbe medulla oblongata, apjiarentlr not far retnoveil from llM vaso-
motor centre. When the biain is removed down to the medulla oblonguta,.
884 THE TissrKs AKD UKcn\irtsvs or digestion.
thntor^'flii li^iiit; IcftioUcl,* dgWofiMiIiva inavaiill l>col)Uii>ed byid«quBt«
iliiiiiiliilioD of rnriniiii nffi'roiil iMrrvn ; when (Im- moiiiilla i^deMroyed no audi
aciion i» p')wil>K-. Anil « How of mlivn niiiy be jtr<Kl««t) bv ilirvct slimul&j
tioii or lti« (ikhIuIIii icmII'. Wlico it Dav of «iUva ii oxoiKNl by i'teaa, itr b^
emotions, t\w norvous procwen l)egiD in the bigliur |>iirt« of tiw brain, ami
deMond thenoc to the mcdtilln bcforo th«y pin riM Co di«tiiictlr cflcreui im^
pulM>: Mill it u'oiilij a|i)wtir llinl lliew hi);'i«r parts of the brnin are C*ll<
into actioii wh«D a Bok of mlira is excited bv distiuct MiuMitiuna of tiiM«.
Pn. Itf.
. W-MnA
K.t}f0,f
nZ
it.*m.m.
Jdt.dt
:?5s*
nW'
-*m^
. mX
V^
-1./^'
DtiURAiiwiTW KnunRfTATioi' oT ni HuaKixiM.tHi UWNi> ur THB I><M. wtTii m Nun* 4yi»
Th* dUMVlKHI hat bcvn niatle eu an animal tyliis nn lu hack, bHt dnw all Ih* pan* Bhtmn Hi I
ncnncouinaltvmn ftumauyoDfiDlniof vIcK, ibcncurRiOoaTiat ginUicuaiHaiiaiiimtcaliali
IMn or Itw tnranU nmotun*.
w^fM. Tlw nlbmaitllaiT Iknd. IiiId ■lioiliicl(Bi.if.i<il wlilnli a lanula ba* lc*n Ileal, TMiuIh
IMisual viand an-l •liiit art uot ikimn, ■J..H.r Tho IliiRUal blanch of iho Dltli nunv. iliv tan aJ.
It cti'KS U til* ■"■iRUg, rA.r.fA I* .cA.r.Ttui'notda l;in|aiil, Tbn lortiA.I'', Ih imnKiliiiii m<inllie
IWtMl iwm : al rk I* II Ivmni** ronjolimt Hllh Ihp IlniiiwI B.l.and anerwanl dlioiglnc |iiiiaia aa
r>.(. tolbeiilaiirl alunic iliciliicl ; Itw CDDtlniialJuo of ib< ni-nc lu oiiutany vMh Iht UiiHiial . ■< , ta
D-il»linwii, m. y, The nuliinixlllary (anrtli'n wUli )t> •mral Rnit, u.mr Tbe arellil uMty, la
(iiiall branvliwoT lahltfb, n, is.a, uiit r. ni.p.. |>ui t»tt>( anlpniiran<l imlcrtor lamaof ItMgb
f m.TtuBnttriuralHl pcaUtli>r Tttnt (Rna lt>Fulaii<l. falUiiK liilo t J., tbc JiimitaT ntn. ■>««M.1
funjiitiml t-AfXK anil »}riii^llivUc Iriifik*, (r rrr, r TIw h^il**! ti-n'IniJ gaiiftlun, tW4 Tiran^'li^ t
wbkli, FijnDliis a piriu* In./ 1 nvn Ihp (Mlal anvrr. an dlilntiutol la iyiii.n(.i aluiig ilw la
(UiMlvlar annlef lo (iLe anurior and pxlcdiii iiur(li>i» •<! ilie Ktaml.
Ttwamnra lodlnta tha rllfucUfiii laivi) by Itir Titrrvun* liiiinLltn* ■liLr^iif nilex tUinulalloii nflll
Klaod. TlMT ammd to Ih* hnin t^r the llUfual anil dcawiul bj Ihc chorda tjrmiauil.
Con«d«ring. thon, tlic Row of aaliva am a rofl«x ac^t lh« centre i>f wbii^J
lies Ul lh« iiimIiiHii oblongata, wo may iinHgin<> tlie efTcreat iR)i>ut»(« paMiig]
from tluit mitre to tlie gland eilber by tbe chorda tynipeni or by tli« ayni*
]>athelic nerve. Although il nould perhajK be raah to ray lliat iu thii
relation llie §yni[>alhetic nerve never acts aa an elfereiit channel, u a miillvr
of ftcl Vfe have no aal»r«clory experimental evjiieiice that il does wi : and
vte may, iherefurc. Male thai, praciic»lly, the chonla tympaoi la thir hoI«
e-flereiH nerve, .Seolion of that nerve, either wliere ihi- librea \m» fmiu the
liugnal nerve and ike submaxillary ganglion to the gland, ur where it nina
BKCHETIOK OP SAXTTA JiVV flASTRtO IVIVB.
8S5
in llxt itunti' nh(!«tb m tbe liiigtinl. or iii any jwrt of it* courac frniii tlx- iiinin
fncinl triiuk i» llio liiifriinl, pufn nti tiul, u» fur lu we kii»w, i<i iIk' [nMxiliilitj^
of nny tl»w being exrilMl by vlimuli u)i]ilirHl to tliu iHiLsiiry iii-rvw, <ir lo llw
aeniknl surfacra of cbe iiioulli or cl' otln-r [iiirU gf (lio body.
The iiHlunil reflt-x ftcc of »r(Ttrlii>n inav In; inliibiM^. like lb* rvllcx iirtion
of tbc vnsomotor nerves, nl iu cvnlrc. 'fhiiK when, nx iii tbi> old nr« urdeAl,
fear p«rchce tho mmith, it if |>mbiLblo that ihi; nflm-ot iinpulmv cniiscl by
thfl pnBeDcv of food in thi; muiilh cnw. ihnnigh emutioiml inhibiiion of
tbttr r«S«x cMitn. to give Hso lo fHornit impuliKv
S2ST. Ill life, iheii, th« tl»w of Knlirii in bnuigbt iibiiiit by tbc adveut lo
the gland along lli« cbords lyinjiniii of ctTpri'iil impulMx, sUrt«d chiefly by
reflex actions. The inquiry thuit nitrrow* itself lo ibc ipit-slton ; In what
tuuniier do ihme erterenl im|>iiliM% caoM' the increaiw of flow f
If io a doj; a lube be iotroduped into Wharton '» ilurl. niid tho choriU he
divided, tlie llovr, if any be going on, is from the !ack of elfcreiit inijniliea
arrtttteil. Ou |>aaeing an inlemipted I'lirrenl ihroiigh the peripheral puriion
of lite chorda, a iiipioua Bccreiiou ni oiieo lakeii plat^o, and the mHvs lios;inM
M ri»c rapidly in llie lube: a very short lime after the applicalt»ii nC the
current the flow reaches a uiaxiiniim which is maintatoed for some lime, und
ibiii, if th« ourrent be long continued, gradually l«weiis. If the current Ittt
applied fi>r a short time ouly. ihv Mention may but for soiue time after the
current haii Ixien shut off. The «livn thu) ohuined U bul slightly viKcid,
nnd iiod«r the iiiieriMciijte a rery feve Halivary eor|iuscle>4, and, oecuMionaUy
only, uatorpbouM Iiininn nf (itxciulinr luaterinl, pritbably iniieoUH in naturo, are
eoei'i. If the gtiind ibvlf Iw watched, while its activity in thun mused, it
will he see:i ^a* w<- have alrvadv ituid, ^ l«'>7 < thai its arUrlea nro dilntod, nnd
its uipilltiri<« lilli.-i'I, mid (but the IiIimkI flows rapidly thnm^h tho veins in a
full Hrenni anil of britrht arli-rial hue. friviue:illy n illi iiviUnlltig niovoineiita.
If a vein of the ginnd In- o{)r-iii-<l, llii* largu iiiereatu! of flow, and (bo learn-
ing of ll»e onliniirv d^-oxygt^iiiitioii ^tf the blood coiuo<|ui.'flt upon the rapi<)
atreaui, will be dill more •.-ividenl. Il i* clear (bat cxiilation of tlie cborda
lart^ly dilates the arlerio : tbe iiervr acW energetically as a roao^ilator
nerve.
Tbns etimulation of the chorda hringK about two events : a dilation of the
bloodvMwb of the gland, and a flow of ealivu. Tbc 'lUMtiou ai once ariaee,
la the latter siinnly the result of the former or is the flow caused by HOiae
direct aclion on ihe secreting cells, apart from the increased blood-supply f
In tupjiort of the former view we might argue thai (be activity of tlie epithe-
lial Kocreting <^-ll. Iik«^ that of any ikiher IbrDi of proloplanni, is dependent
on blood -Mupiily. When tlie smalt arteries of the gland dilate, white the
pnMurr in i)ie arteriuon the side toward the heart is i as we have prvviouitly
jiMD wbtrn inraiing generally of blood-pretsure !i l:fO)corr^j)oQdinj{ly diniin-
bkad, tilt: prvwure un lliv far side in (be capillaries and veins u umr-aned ;
brae* tlw (uipillarieii become fuller, and more blood pas«ai tbroti^Ii thera in
a ^ven time. Prom this we mi^ht infer ilmi a larger amount of iintritivc
nialttrial would \>nii» away from lh« capillaries into the .lur rounding Ivmpb-
«plH(M. and Ml into thi* epithelium celU, (he resiill of which would imlurnlly
M lo <|iiickcn the priKcwivi goiuft on in tint ivlk and t<i stir Uhsm up to
Sealer aolivitv. But even iiiinilLlini^ nil ihi.i it iIinii not ncoemnrily follow
at tbe arlinty thu^ cKi-ikHl should take on the form of Kerrftiou. Il is
quite puwible to conceive [lint the inttnmird lil<H>d niipiilv thoiiM |«ad ontv
lo the aooumulntion in thi' ii-ll of ihe const i (urn bi ol the saliva, or of ilic
raw nuileriala for tk-ir nim^tnu-lion. und not to a dimbnrgc of ihc MiTctioii.
A man worhs hotter for iieing fed. but feeding dots not niiiko him WDrk in
tltc abeoenoe of any stimulus. The incroMwd blood.«uptily, therefore, while
ZBd
TDK TISSL'KS AND HKCRANISMS Of DIGESTION.
fitvorabli* (o nctirr wcirtioii, aixd tiot DOccnMrily briDv it Rboiit. Moreover,
the following liu-U ilifiiiiclly thow thnt it ami not. WWn m cunula >• tkd
inlu (he ihict imd ihc L-hoi'<l:i kcncrgrtieiillr HlimiiliiU^, ihc prnmirc acquired
by llie saliva ntciimiil/tteil in tlic cuniilii uiid iti (lir liuct muy citvol li>r tho
time beiiiK the nrlcrini blDod-pmsiirc. even tbut <if tliu ciirutid nrt«ry : thnt
id lo «ay, Ihe piv(>euiv of tluid in the glaiid uiit»ide tho bl(>o<lv«M(>)8 w ercntvr
ihau that iif tli« blomi iiiftide the blooilvemcltt. This inu«t, whatever Ixr lb«
exai<l luoiie of traneil of nutritive nialcrial ihroiigh the vascular uhIId. lend
lu cheek tbut transit. Again, if the head of an animal be rapidly eut off,
ami the obordu immediately eti initiated, a flow of saliva takes place fnr
tiHi copious to be aeeiiiinled for bv the emptying of the iialivary ehnnnels
thruUKli any *iip[iotM-d c:i)nlrauti(iii of their walls. In this ease wcreiion is
excited in the gland though the bloodaupply is limited to the small quaulity
Htill miiaining in ihv bloiHlveaselti. I^anlly, if a small ouaulity of airopiite
bi; injected inlu llie veiiu. stimulation of the chorda produces Do secretion of
saliva at all, thuiigh the dilation of the bloml vessels takes place as uxual : iu
?iitc of the greatly increuited bloiHl-«u pjily no secretion at all lake* plftc«.
hciH! fiii'ts prove that the seeruiory activity is not simplv the result of vas-
cular cliaiigni, but may bo ealk-il forth indvpciidenlly ; they further l«ad as
to Kiip[H»o that ihv rhonia o<iiitaiui> two m-T* of fibn«, otie which we nuiy citll
SGcrt'tory lihm, acting <lirr('tly im the MciTciing flrueturv!) only, and tlto other
vtUD-dilntor tilire*, seling on the hloridvnaoln only, nnd further that atrupiiic,
while it hiw no cHi'Ct on lh« latter, paralrn'^ the former jum an it paralyses
the iuhibilory fibrvs of the vnEU». Hence when the chorda is rtimulHtc<l,
there |)as<'I<iwn the nerve, in addition to impulmw aRccting the bUMMlfiipnly.
impiilecf nKectiug direelly the protoplasm of the Kicreting cells, and calling
it into nction. just as similar inipiiUcx call into action the contnictility of the
subslanoe of a niUHciilar fibre. Indeed, the tno things, wcreting activity nnd
CODtraciiDg activity, are very parallel. We know that when a muscle con-
tracts, its bloodv»sela dilate ; and much iu the same way as hy atropine the
tecreiiiig uetiou of the glaud may be isolated from the vascular <lilntio«i, so
(iu the frug at all evenUi bv a proper dtjee of urari muscular voDtm<!tion
may he removed, aiiil li<tive dilatiua of the hlniKl vessels as the only effect of
stimulating the mu.ienlar nerve. In holli runtA the greater (low of IiIockI
may be mi udjiivant to, hut Is uut the «xeitiug cause of, the nativity of ihe
structure*.
Since the chorria nets thuK directly on the secreting cell*, wc should «xpccC
lo find an anstoniical e<>unecli<in between the edls and the nerrtt*. and UIM
Bulhon hrivc nminlaineil thai the nervc-tibrcs may be tracwl into the «.dls.
Bui. *nvv ncrbaps in ihc cbw; of ccrlnin glnnils of invertehrat« (si>-nilled
salivurA- glamls of Ilhtta). iho evidence as wo have Niid is as yet not
CouvinrinE!.
i S&9. \Vhi-n the cervical eympnthelic is stimulnlcd, the vascular effectB,
as ne have ulrendy Hiid. ^ ItlX, are tho exact conirnry of tliutw seen when
the ehiirda is Mimithiled. The small arteries are eoustricted, and a small
auaiitity of dnrk vcuou* hlowJ escapes hy the veins. Sometime*, indeed,
»e tlow through the gland is almost arrested. The sympatlietic, therefore,
acts as a vssa- constrict or nerve, and in this sense is autagonistic to Ihe
cborda.
As eoncems tlie flow of saliva brought about by stimulation of the sym-
polhetjc, in tite ease of tite subma.\illary gland of the dog tlte eAbda are
very peculiar. A slight ilow results, and the mdiva so secreted ia remarkably
viscia, of higher specific gravitr. luid ri<'ber in ourittiielcs and in the above-
meulioned SNiorphous lumps tWn is the chorda wlivn. This action of tlM
STui|ittthetic is little or not at all aflecl«d by atrt>]ri»e.
SXCBKTION
Attn 0A8TI11C jnici.
WT
^
I
Ib Uw tnbnuailbry glnnd uf the <k>g iheii tliv ctinlnm between the elfecta
of ehonbt tUiDiiUlioniini] thoecof eympnlheiic »tiiiiiilatian are verr nmrke<l ;
tlw forawr civn riM to Ttt#ciilar dilntion with u co|iioii» dow of fairly limpid
■■livs poor 10 •olids, ihe lalter to vasculnr eonalrictiou with a tcaniy duu' nf
viacid mHvt riclier id aolMs. And in nllici- anintals a similar coulrsst ni«-
VBib, tbuugli with minor diSerencw. ThuB in the rabbit both chorda Mlivft
mad tTrnpfttbetic aaliva are limpid and free from mucus, thouj-h the tatt«r
oDolauM man proleitb; in the cat. chorda aaliva it more viaeid tliau xympa-
thelicMllva; but io buth tlMse cases, as in tb«dog,atiniuIatioD of the chorda
cum n cnpiouH flow with dilated bloodreneb, and stimulation of the aym-
palbetic a acanty Son- with vascular couairiction. We ahall return Uf^ain
prcMDtly t(> iheM different uciiuna of the two nervce ; meanwhile we linv«
Men enough of the Iii«tory uf the aubmaxillary gland to l«atn that »evretiun
ia tfab inotance ia a rettex action, llie elTerenl impulsea of which din-ctly
sflkct tlte ■Nreting cell*, and iliat tlie viisculur plienomena may awiat. but
an Dol the dinct caiue of, th« flow.
( SS9, W* have dwelt long ou thin j:\unA l>ocau*e it ha« b«en more fruit*
filU; attidicd iban any other. But the dltvoua niet-linniama of the other
MliTftn clanfU are in their mtiiii fmturm similar. Tbii* the •ecrtrtioti of the
pwvtiil gland, like that nf the Hubnmxillary, io governed by ttvomitiiof libnv;
uov of rerrbro-cpinul ori^ii, niniiiii^ along the auriculo-tvuijionil branch of
tba llfUi ocrvp but originating iiOHsibry in the gto«Mo<ph*ryngcal, and the olh«r
nf arm pathetic origin coming from the <^vrvical »ympathetic. Stimulation of
tba cmbro-tpinal fibrrt nroducn a copious How of lininid wiUva, free from
luuena. the tevrvtioii reaching in the do^ a pnwurc ui 118 mm. mrrcury ;
ttinmlatinn of ihr cervical syni|wthctic pvv» riM in the nibbit to a fccrction
aiK> fnv from miicui> but rich in ppiCcKU ami of greater amylolytic mwcr
than the ccrcbrciHoinal McrctioD ; in die dog ItttloornofocrotioD is produced,
UMMrh, aa we shall »ee later on, certain chanKca are brought about in the
riaao itaelf. In both animala the oe re bru spina 1 fibres are vaao-dilator, and
tM ajmpalheiic 6bm raso-miutrictor in action. Stimulation of the central
end of tbe frloBM-pharrnmnl produoee by rellex Mlioo a seotetiau ftnm the
IMratid ghiod. but tliat of the linjfual iit said to be without eBect.
{ CM. The tfcrHton of goflrie Juice. Though a certain aniDunt of gastric
jtuc* may •om«iiraes b« found in the utotnachs of faaiing animals, it may be
■Caied genenlly that the stODiach. like tbc Milivary );Unds, remains inactive.
rieldtng nn wcr«iion, so 1m^ aa it is not stimulated by food or otherwise.
Tbe advene of food into the stomach, however, at oi)c« causes a copious flow
of gaslriojuke; and ibr i|uaiiliiy itecreied in the twenty-four \unm is probii*
hiy very eon^dinrabh-, but we have no tnistwurthy data for calculating the
vxact amount, So alto wli«n the gaalric mucouii membrane in stimulutvd
nccbanicallv. a« with a feather, M-crt'tion i* excitci]: bnt to a very small
amouiiteTni when the whole interior tturliicc-nf the nlooimli ia tbu* regicatcdly
Slinillaled. Tbe moat cfBcivot alimuluK ii> tbu natural stimnluM. vit., food;
iboagh dilute alkaliv seem to have unuiunlly nuwerful ■limulatiiig cIRvta;
thus tbe nwalloving of xaliva at oti<« pravoIcM a flow of gaMric juicr.
During fnRliiig the gastric membrane u of a pale gray color, Muncwliat
dry, covervd willi a thiii laycrof niucui, and thrown into fold*; during digea-
tinD it becomn red. HiU'hrfi, and tumid, the fnliU dimppeiir, and minute drops
qf fluid ai'pearing at the mouths ol' tin- ulnndx. s|HHiliiy run together into
S0iall ■irrains. SVh^'n the seciction if viry iictiw, the bltHxl flows fWim the
cn|dllariea into the veins in a rapid stroiim without lotting il* bright arterial
hoe^ Tbe secretion of gastric juion is, in fact, acoimiwnird by vascular
dilaMon in the Nicne way ns is the spcn^ioti of Niliva.
(SSL r^ing tlmt, unlike tlie case of the salivary secret ion, iood is brought
888 Tim Tissriis anu mkciiakisms or oioestion.
into tbo innuvHiatc nfigh))i>rli<vxl uf (h« crcrctiti); cells, it ts eiraedlnely
nrobablo ihul n grunl ilrni i>l' ilio s«creiton is tW nwull of tl»e working uf a
local mtx'hatiiBm -, iin<J this viow is m|>|Kirte(t bv tliefnct tlinl vrbeii a OKclisni-
di] etiniulu!^ K ttpplidil lo one 8[>ot of ilie },'iiB[ric tnciubniiit! llie wcrelioa is
limited to the n«i),'bborliood of ihal f>pot aud is uot excited iu dUtiiui parts.
This Will lueubaiiifiiii lunv be nervous iu nature or the eflect of ibe stimulus
may |)erlia|w be <<i>nveyed directly from cell to cell. tVoiu tlie luouth of the
f;liui«l Ml its extreme base, without the iutervciitiou of any nervous elements:
lUt the vascular cbau||;ei at least would seem to Imply the pruaetioe uf n
ncrroua meehanidm.
Tilt' sliiiimch ia suirplieil with ueri'e-fihrcs from tJie two vngi iivrviv luid
from the ^olar plcxua of the iplnncbiiif! system. The two vug* nl^<-r lormiug
tbe ivsaphngi^iA iilcxiiv ou ihe cusophagus are gnthi-rcd together n^in a* tiro
main tniiik* which run uhing the orvophagns, the left in thr fmui the right
at the back, to the stomanb. The left or autcrlor nerve i» <IUtribute<) In ibe
sruailer curvntiirir anil tht^ I'n^nt nuriiice of tlie stomacb, fonning a plexus to
whik-b nerve-cell* are pniu'iil ; luiil branchof pass on to the liter and proba-
bly to the diiodeuiini. Thi- right, or potterior nerve is dbtribuled to the
binder surface of the stomach, but only tu the rxtcnt of about one-third of
its fibres; about two-thirds of the librcs pam on to the ralar plexus. The
fibres of the vngm nerves thus distributed lo the stomach ar« for the most
|iart noD-medullatcd fibres; hy the time ihc vngus reacheij the abdomeo it
consists nimiiftt exclusively of nun -medu Hated libres, medullaled tibre* being
verv few : the targe number of niedullnteil tibres which the nerve contains
in t'be upper part of the nock pow oH' into the laryngeal, cardiac, and other
brnnclios.
Friim tlie solar plexus nerves, arranged largely iu plexuaea, pass in company
with thu dividions of the cteliac artery, coronary artery of the stotniwh and
brnnclM> of tht^ hrjwiic artery, to the .itomurh. Though the two obdoroioiil
splanchnic nerves which juin the !>oltir plexus '.itimiluamr giuiglia) arc chiefly
composrd of meduUiitcd tibn^i, the nerves which jiujw from the plexus lo th«
stomach arc to a Urge rxtent ci>mpowd of iion-mcdultiitc<I lihrcs. iVll these
nerv<«,both bra^ch<.^)> of the vagi and those from the solar plexus, lie at first
in compnnv with the artcri<.« on the )>urliiec of the stomach beneath ibe peri-
loueuni. l^rom tlicncc they jiitss inward, still in company with arteries, and
form on the one hand a plexus, contnining nerve-cells between the longitu-
<liual and circular muscular coats corre»])onding to what in the intestine we
iihall have to speaik of ns the plexus of Atierhach, whence fibres are distrib-
uted to the two miiseular coats, and on the other hand a plexus in (he sub-
mui'uus coal, also containing iter ve'<?c lis, corresponding to what is known in
ihe intestine as .Meiesner's plesns. From this latter plexus 6brea pass to the
mncou.i ineailtraue: some of these end in tlie niUBeularis niucoste; whether
uny are connected with the gastric ;:laud% aud if k> how, is uot at present
known,
Th«re are no facts which alford satisfactory evideuoe that any jiart of this
armngenwtnt of nerve* supplies such a local nervous mccliniiimu iv waa itug-
gtytcif above. The importaucn. however, of such a local nu'chauiitm what-
ever it* iinture, and the aiilHirdinnte value of any connection betwwn tha
gastric mi'mhrane and the centnti itcrvoui) srstt'.m, \n further *howu liy tha
fad tbut a secretion of r|niii' nonual gartric juice will gn on after both vagi,
or the nerves fmm the solar jdcxns going to iho siomticb have l>een ilividcd,
and, indci^l. whsn all the nervous couiK'Ctions of the stomach nrv n* fiir a*
possible Slavered. And all attempts to provoke or modify gastric secretion
liy the stimulation of the nerves going to the stomacli have hitherto fnilwj;
On tlvp other hand, iu cases of gastric fistula, irherv by complete occIunoii
CllANGBS IX TOE flLANHS.
S80
I
*^tR a»u(>lift^ii« fli rim latino by lliv dract-nt of Rilivn b^* )>**n avoidisl, ibe
uM-ra N^ht 'ii miit'll of f<HHl linn beoii Mh-ii (u [>ruv»ki; a lival^ oucrolion of
gMtrk juiott. Till- mutt linvi- W-ii iluv tu Mniu nervoiu actMHi : ami ihu
iKOM iMftT bv HiiJ ol' tlH- vnK) wWrr inintioaa uf gripf or niig«r •inldi-iiW
ailwt iIm- McrvtioD going on or prevent tin- tfcr\-li:>ii irliioli wniili] utIiorwiMi
bare lakni {>lac« m tli« mull of iho )inM-tiov or lliuil in ihc sl'iinach. 8j
tKftt miicb bus yH tit be lenniad in ihu mnltar.
i 932. Tbr c'tDtrast (jreMnteH between iha taiiUy Hcration resulting fntia
Birt-hanical stinuilalion and tlu> cii[H4UB flow which oi'tiial I'ucm) iiii)u(M« U
interatinK becaiue il sceinB to shoir tbul thcr •ecrelory Molivitr of tlie cells is
bcMiiaiM br tbe Bbaarption of ccrtiiiii prodiicU ileriveil t'rotn thi? |)i>rlKms
of fbnd fim dif^otM). TliiH h well illiutrac«d br tbe fiillowing expcrimtnt
of HetdMiloiu. Tl)» oWrver, niiogititi^ the method employed for the iiit(»-
iiw!, of «hicb w« thall apenk later i>ii, sucoeode*) iu iwluiia^ a purlioii of the
ftmdus {rata lb« rat of tli« tiioiiiiicti : tbst in to say. lie cut out a porliau of
tb« fiindiu, Mwe<I tnj^tber lh)> cut M'^ of the luiio iitom:ich, »j ait to forni
■ anuller but otlkemae coinj>lete 'ir^'un. whik- by liiitured he conrerte^l tbe
'I piece af fiiiidui iuiA a »iuall iiidujienilvot sluinBch opeuiri^ no to (lie
. I >r by a fii^iiloun orifioe. Whfii fiiml wai> iutr<»luc«it into the itiniu
*i-mt*fh Momion alsci took [iIsik in the ixi>Ut6i] luailuH. This iit fl»t HJKht
nUgbl Mvm tbo mult of a nrrvuio n-th-x ima -. hut it kah olueri'ed Iha: the
MCMkdarr wcrvtion in the Inuilii* niu dt-|)cri>)<Mil on netuiil ilij^lioti takioe
p)M9e in tbo nuin nloniiioh. If tbir mntvriiil iiilrodiiCL-il inlii iht; main aiomich
wart istUgoMiblu or <li^<«ltil nith ditlicitlty, »'> thiit Hltll^ or im [iroducts •>{
dlntMioo von ((>rmrd lunl alMxrlii-il into the blond, ouch c^. 71-. ms |iie['«a
•tf ligamcnlnm Dticbee, very- lililp^ivrelion look \t\iux in thv iMiintttil fitnditit.
Wo ijuot« tbit noK ns bearing on the iiiiMtioii of u [xjMible iivrvou* iiKictiaii-
hm u( gaaitic wcMioti, but irc shall huvc !■> rvtuni to it uudcr aiivlher
Th* CUtanyrt in a fliand OanMiltHinj Uic AvI o/Sterttmt,
ft S33. n'c Iwve lion' to coaiider what are the chim;;ed iu tbe glandular
oeOs tuid tlirir KiirnHimlinj^^ whiih c^\ne thi.i t)<>w of fluid |iu^MHilliE sfiecific
dnnutvn iato the liimvn uf un alreolus, anil au into a <lucl. It will be OOD'
VUMOl In iMEgin with the pancruiLs.
T1m> thin axteodoi) juuicms uf it rabbit may. hy menu* uf special prei-au-
liom, be (prcwl out uu tbe Ha;^ of a mii;ro*(.v>p<! luid examined with eruti
bt);b iioni-r*. while ibi; aiiimal it ii»l unlv alive hut under iwch cmditi'ins
thai till- gUnd n-mains in a nuirly nonnHl lUali:, L'a|mble of Morelinh! vttjor-
outly. It is |HMuhlf umkr llxae dnniRulAiK-i^ to observe eren minut^'ly tlie
HpearuiDM prr«onto>) by thi! yland when at iwt and loaded, awl !■> wiil«h
toe changes wliiuli take plai>- during twcretion.
Wb«i tbe animal h« nut l>oen digesting for (nmc IJttlo lime, and the ^laod
ii ihereruro "luadol." ihe oullincs uf the individual ocll*, a* we hav^ alrvady
■aid. i TM, are very indiftinei, the lumen of the alveola* it invisible or vi-ry
ineitiitpicuuu*. and each coll ig crowded with »mall, refnietive spherieal gran-
ulca, fonniu;; an irregular granular man* which hides the nucleus anil leaves
only a vt^rr narrow clear outer £i>ne next to the biuemenl membnuw, or it
mav lie hanlly anv sueh xone at all. Fij;, lOS A.
Tbe Uoafl-aupply. moreover, it §eantr, the small arteries being omstriclod
and the capilUries imperfectly tilled with curpusdes-
If. Itowevef, llie same pancreas he exaruioed while it is in a sute of aciirity,
^tber from ibe preKnce of fijod in tbe stoniadi, or from tlie injeetion of some
MiiBntaittig drug, such as piloearpiae, a very difleiwu ttale of ifa'mgs is seua.
840
Tlie riPSt'KS AND MKCHAMSMS UK DIQCSTION.
Tho indlvidiml ct\\» (Fig. 106 Jt) Imvo Wcomp sniHllpr iintl much nmre dis-
tinct in outline ttml the contour ul' the nlvcoliia which [ircviou^ly wiu evcu
is now wnvv. the hioMmfnt m«mbrniie being imlcutcd nt the junction of Um
eel Ik ; nlso (lie lumen of lh« nlvcolus i; now wider and more omspicuOQl.
Tn cnch c«)l iho grnniilee bnvp hecoiiie much fewer in number and. as ii were,
have retreated to the inner nitir(;in, eo that llie inner granuliir zone is muclt ^
nanviner and the outer transparent zone much broader than before; thi
latter, too, i^ IVcquently marked at it« inner part by delicate striie runDing*
into the inner xone. At the same time the hliiodveaBels are lar>t«ly diUted
and the stream of blood through the capillaries h full nnd rapid.
no lot.
A rttiTiotf or TKI fxici-KKw Or Tits Kjuiiit, iKiuxn *utl siumiiAi i.iu.i
i(»lre«. OlnxUlvnrnrllvlir. a Ilio iDncrKruimJungniiv. «htvli In A it laivvr, khiI noK i
ftudil«d Willi tiUii Jrnnuli'f iltiiii In Jl. Ii> Hlilrli Ihr kishiiIitf b» I^Har nJiit coaratr, ftllM tmU
tfaiii|*i«nl Jtonp. uuiiU Id A, laricr Id A. ■mltn [be laiicf iii*rlK>l irlili Max ttrtit. rlli« 1«ni*ti.
TSnr otirloiu In 0, bal iii'lMlniUn^, it an lUdnilHilun at iiii< juiictton or iwooetli.wdiin B. hm
not ooourrtOE In A.
With care the change from the one state of thinip to Ibe ather may bo
watched under the microBcope. The vascular changi's can, iif couwc, be
easily n|>prrcintcd, but the granules may aUo he seen to diminish in numhcr..
Those at the inner marj^in seem lo be diHtlmrfccd into the lumen, and lbof«^
Dearer thf uulcr niiirffin to travel iuward ihr'>UKh the ccllnultetancc lowarrl
the liinicii. ihr Itiint Nlriie epoken ol alxiv^, upjinrentlr, at nil rvvnli. Iicing
the niarliN of their jitithii. ObviuUHiy. durinjc Aecrelittn, the granules with
which ihc cc!l»uh«l"iii*e was " loaded" are " disohHri^inl " frt>m the cell into
the lumen nl'ihc alveolui^ What chatigeii thott (^runnlcn mity undergo dur-
ing the discharge ue xhull consider nivseutly.
Section)! iil'ihi: prepared and hardened nancn-ax nf any animal tell nMfly
(he same tale n» ihat thim liild by tht living puncri'Md of the nibliit. In
sediiins for inrtancc, of the piincreaii of a dog which has not been («l, and
therefore h»* nut hrru digesiing, for wimc hours i twenty-four or thirty), the
cells are seen lo hr crowileil with granutce (which. Iiowovcr. «re tiaually
abninken and irrv-gular owiii); t» the inHuence of the hnrdening agent),
leaving » very narrow outer wine. In similar MCtiontt of the pancreas of a
do)t which has been rcccnlly fed, six hours helore far e\nmple, and in irhich,
therefore, the gland ha* been for wnie time actively secTeiiDg, the granules
are far less nuinemue, nnd ihe clenr outer inne iiCLordingly much broader
■nd more const'iiuous. With o>niic acid these granules stain uell, and are>^
prrMTVed in their splirricnl fnnn.so that the cell thus stained maintains
maeb of the appcsmnoc "f a living cell. Hut with carmine, htnoatoxaline.
He, Che grauulf* do not Main ntnrly so readily as does the cell-BubeUltce of
t)i« C«I1b, h> ibat a discharged coll Rtaint more deeply than does a loaded Mil
CIIAN'OBS IN THE GLA.VDS.
84L
I
I
bnqmie the Maininji; of (lit- " )iroui{ilii«niM.' " ocllxLihMUiuee is uoc » much
obMBrcd by iW uiiMuiiitrJ uniniiln; baules wbii-li, Iiowovit, the ac-tual cell-
•nbttftOff »tittti» I'Ttitimblr mxui wliiil more <lrq>lv in tliu dUcbargeii cell. It
m»y iit mIUmI tliiit iii ihi- iju('hnrg<:il (vll thv niioli^i^ !« coii»picuoiu aDd w«)l
fanned : in ib« IoiwIhI ocII it io ^ix^mllv in prrpurci) MCtioDit, more or Iras
imftitar, poMiblr bonuMr iu tbcw ii i* (<!» dvnw Bti'l more watery than in
Ik* diMharg«d evil, and an ahrinks under the iiiHucnoc of th« reoieetiU
Tbrae M>ver«l i>WrvBtiom «ugg«t tbi.- cuiicbiMon thui in a gland ut raet the
ocll u ncctiptoal in ftirminif bj mnttw of ihc niclaboliaiD of itn cnll-itubiuiiiice
SD(I Imlging in itn-lf ($ '.iOt oertaiii gruniilw of peculiar suhttance iuUnded
to hv a pan ami |in>b>iblj an importanl pari of the wKretion. Tliia gnca on
until ili« n'll i« RKin< or leM completely " load»l." In sucb a evil iho amuunt
of actual liviuK i<ell-#u beta nee ib n^lalively auiall, its placo is largdv uccu-
PmO by granuTa, and in itwif bns boeii [lartly constiniod in fornifiij; lh«
gnUHM. During tbc net of Mcrction ibe ^lanules are dL8cbcirgi-d to form
|Mrt of tbe ttcretion, other iuatu>ra iiicludii^ waier, a«i ue ithsll *•%, ninkiiig
Dp Uw whole MCKtion: and tho cell would be prnportiunalelv rcducixl iu
w« werv it DM that tbe act of tbe discharge teetaa to itiiuulato ibc <-cll-
nbrtvicv to a new activity of ^mwib. «o that rhc new cell-«u balance it
fiivaad; tliia, faowerer, 'a in lura aouu in part cunaitiuied to order to form
Dew ytaiiulea. And wbal is thus aeeji wiib conxidcrable diitjuotnew and
bCBM IB tbe pancr«a«. b »een witb more or lcn> di«iinctn««< in olber glaixU.
(234. W'bvii ue study au iilbiiminoun elund, tbe parotid gland, for in-
atmotx. in ■ living .-uiv, no lin<l tbiil tbe (Pangea wbiclk take plaoe duriDg
•rtiviiy are ijuite comnorablc to ihow of Ihc pancmut. T>urit)j; rest (Pig-
1107. A), tbi" cellg are large, tbeir DUtlint^ very indistinct, in fact almost in-
rbiblr, and tfae roll-Rib«tancc a studded witli granules. During activity
' Fig. 107, U » tbe cells became amnller, their outlinea more dIsUnol, nud the
ilk
Tlwanp^sIMb I* tanmitat dlaaBmoiaur. npnMaW iba niicroHOpIc (kuicis wblchuuf Im
BtavndlDlkalMVfllBd. A. piBlltf ml Tlw etwvnr outllna oT tile calli arc lnlr»lv»l lo
UvMiUI** dHof lb* •vCb: Uiei- o»ild noC be mdUjr awn In Um f|«daiin ilvtf M. Atlsr
•aniiilaUnn. C Attn pmXiuaml lUuuiUUuu. Tliv qocIbI an dlacmniniUo. «ud lutni.
l>*lwwia«lrap|iWB»eEMid ]a*l(lnn.
graaulM disappear, eB|)eciallT from the outer portions of each «cll- After
pr»l«ageic) activity, as in Fig. 107, C. ibe celU are «ull smaller, with their
rt». K/s.
nnlltnea Htill more diittint:!. luid the gnuiulca have diaapiMarad almrxit
Mitiraly, a few only Iwing lelt rtl tlio exireBM inner margin of anob cell,
ntnittiuj; ufiun tbt' conitpicuouB. aln^Ml gaping lumen itf ibo alrenlu*. And
upxn #|>m:ui1 examination il is found that tbe nucli-i arc large aiiil round.
In fart, ite mi(;bt almoH take tbe parotid, ax tbu« studied, tn be mure truly
typical of M-rretory ebaDgea rhan eves the paiicmui. For the demarcatioD
of an inner and ouli-r lane i» not a necenarr feature of a secreting cell at
mi. What 11 eweutial is tbai the oell-suwtance manufaclum ruaierial.
S42
THE TlSSfKB AM> MECHANISES OF T>ieESTK*N.
wliicli r»r n n hilr, ihtic in diirhiE rT»i, i» depnfiiril iii ihc veil, genornlly in
Iht! form nl' griiniilca hut not iii'C'<^'»»rily «», iin<l timt diirinc iirtivily ihtt
tnnlrrial iti iiM'il up, llie ili«n])|M'iiruii(f ■>>' (h<* ^rnniiii^, wltcii thmt nre
vitiilik*, Iipiiif; nainrullv fiirlit-^i na<\ niiml niiirki'i) nt ihr tmu-r lairtioiis of
each cf'll, and progrcwJiig hiwnril Kiwiird tliu lumcii, ihe nliole cell becoming
itnallvr iiiiii uc il wen- ahninkrn.
lo iIk' cell* of tlic pnrotiil ^luml nnd otlicr iilliumirimie cells the gninti'r*
Bccn ill (lie living or Inth cell ditlcr from ihi- gninul^ Been in iho pkncrmltc
cell, itin[>niitch tm they lira tiiMly dJKolvpd tir broken up by the action of
nicohol. chromic Rcid, mid the olher iiFunl linrdenin;; reajjenU, ond lieacc in
hardened specimens hare di»appenrpii. In coneefjuence, iu aectioOB of h.ir-
dencd and prepared albiiminoua glands the difli-rence between reetint.' ''T
loaded niid active or ditcbtirged cells may apjiear not verv coii^icuuiis;
and ihis is especially the ca^ iii the jmroiid ^latid of lh« rnbbil when rhe
activity lias been calleil ioin play by ^.tiniiilalion nf thu iiuriculo-l«m[mnil
ii«ve. When, however, either in ihe rabbit or the dog iho cerrical sym-
Intthetic i» atiuiulHlcd. tboucb the Htimulntioii girCM ri»e in llic rabbii to
itlle »(crelion of -laliva, and in tht: doj; to muKt nt all, n marki-d vllr<-t on
the gUnd ia prodiicvd. and chan;{«« in ilic auiuc direction iv ibfHc alnuidy
dtfCrilH'd niny he obsiTvcd. During rwl thv cells of the piifolid ai" rtrn in
tectionn of the jjtnnd hardened in alcohnl ( I-'ig. lOM, .1 1 jirr- |>nt<*. lranM|ia-
ri'nt, sininijig with ditrn.-ultv. ami iIk^ nuclei piKcc" irregului' outUiH-* a* if
»hriink«'i] by tht^ rengi'iitii employed. AIUt Hiinudation <<l' th« sympalhetic
the pnilop)a«m of the crllx hccomn turbid (Fig. IU8. B}, ■»<) stains much
Fm. 101.
^n-iii>.v or TiiK i'tnuttii »r ih^ KaUkT. iJinti lluiicMUiit.i
.1. AirtfL S. AHut itlBnUUui or llie rarvl(iil«pnp*.|li«]i:. Unlti tccUUu afe IRiia MnlnnT
■tend.
more rendily. while the nuclei are no luuj.'sr irreKnInr in outline, but round
and large, irith conspicuous nucletdi. the whole cell at the muuc liim^ at
le**l afl«r prolonged Hiiuulatioti. becvimtng di«tinctly smaller.
^ 235. Ill & mtirouit gland the changed which take place are of it liko
kind, though Bp|>ar(!ntly »onieu'hat more cnmplicJilcd. ntvin;; prulmhlr to
llio peculiar dianiden of the mucin whtdi i« (o cunapicuous a (.■uiwlitupnt
of the MiTclioii.
Tf n piece of rf»ting. Icmdcd tiihmasillary glaml bo teased out while frwh
Mul warm from the IkhIv in norninl sniiiie nobition. the cell 'ubelance of tiie
mu(xiu» eclle < Fig. 109, a) i» seen to tie crowded with grnnulos or ephcnilev,
which may fairly be compiired with the granules of the pancreas, though
perhaps leM dense and solid than these.
If ■ piece of a gland which hae been secretiDg for eonK time, auil h
therefore a dischargerl gland, be examined in the same way (Fig. 100, h),
the granules are far less numcr^ius and largely I'ontineil tu llie ]Mrt of tli«
ell nearer the lumen, the outer part of the cell aniimd ibeuucleufi i-(iti!<t-iinfl '
rpr onlinary "protoplasmic" cell-subsUince. 'flie dtMinction. howcvt
CIIANii£S IN TUB GLANDS.
U9
no. lat
bvtwnii Ml iuiMT "gntiiiUr zone" ntrxl to the Uinwo ttid in niitcr "dnr
flooa" nrsi III thi- hMrini'Dl mvmhmnt? tc li-w iliMiiict tliiin in iW pimcrcmM,
|«rU7 bwniur itiv granntn ilo not diNin-
jmmr in t» rq^lar a nmnnpr ns in llio
t»anmii. umI |<«ily bc<-»i)w lh« oiitrr
of tli« Diuruus cell, ne it fornM,
BM boiDOffeDMHifl ihnn that of th« pan-
■tie cell.
Th« "i^ranuln" or "Bph^roleg" of iho
nmeoui e«]l are tunn^tver of u peculiar
natun^ If tl>« fmli c«ll, mhuiviug (;nui-
alc» (eiihtrr man}' aa io iho ca»e of a
loaded ft few aa io the case of a dia-
char^ml ccllk, Iw irrigaiett with water ur
with iliUiif acliU or dilute alkalies, Uie
gratiiilr* *«a'1I up ( Fi){. 109, a', b'} into
s traii*)iBiri>l niaK>, giving the react iou^ of
marin. lniv^r»r<l l<y n network iir"pm-
Ui|>la>niit- " cpllfulmUnce. lu lhi> miy
i* producvil aii apfK-umnro von- limitur to
ibM aliowii in sfCtionii of niuroii* v''i'n<U
hardcfxd and lUiinctl in the ordinnrr
*L
_l««e havo alrmdv snid (£ 210), in
ibe loaded tnucmic ^'cll in such biink'iH'd
ajdd Mainvd pnpanUioiu (Fig. 11 n. a)
ihon 1* IMA « nnall qiiuntitv of proto-
plasmic i.-tI1 wlttinncegntlRTed round the
ngcleu* at tbo milcr part of the coll next
U> (!>«■ bawtiMint nK-nibraiie: the mt of
Um nil rante^i of a uelwurk of i-ell-Bulwtance, iho inlervlicm Itciiig lillvd
whb tnuMpnn-nl material, which, unlike Ibe network iioolf and Ibi- ninw I'f
UBV QbUlU oi Iw. lLj--«>LtT.)
■ Hiiil b bmbitfil 111 t ivmnt. Mil tUn-
tiuii: a. fRmi |m|ed (Isiid: b, ftwii dbk
rlinfiTTd kI«jii1 ilbc sucM lira ii'wll)' mum
MbM'iircil by (mnulpt Uuii !• btir npre-
•mlM> on ln»Ic« mil • mtli (V*Kintal
ill I tu 1 im miL Mil K-1uUi«. lUn nils
luiully brcnino linAm ui'tulliii Im^IhwiI
rall> ft» rkfvlf obiainal tultv: kHitiM
cell* ■!« (muiicin iruwiliml Iw Ml lu llic
tnl)- Inr • Oijr allK dMtb. ■'. V. trnlRl
■ilh ditiiw ariil: ■', ftonkaiiloa: V.tnM
lUwIiiupd rUnid.
Ainou <w I'M'* A'MittiiuBV CiuAii lUu>mr ik Aurnitoi. ako fttinai wnu CiHm.
lUuiawnt-i
n«aM«««tto>llw>unmiW. a,Avia«lmleiltluid. Ufra)na>llMh«>v»4sUiMl: tlMtbanli
(jnitBDl lw*lna ••«« atUDUUMil at (bprt UiUml* ilartiis live InHIix
niK-lett*, d<iea not r^iain with oannine or with corlain
r^d nil in wmilnr preparations (Fig. 110, hf difKira
844 TUB TISSUES AN'1> MKCHANISUS OF D10E3TJ0.V.
from the Insded cell in the atlll>un^ of lrHnH|Hireiit non-stainiiif; material
bung much leee and cliieBy caallaed ii> chi> inner part nf the cell, while the
pntopluMnio i.<en-iiul)HtaiR'e around ihe now larve nod well-formed nucleus is
DOtonlj^, both relaiively and uhaolutely, ^r^itter iu iiiuoiitit. liut stains still
more dwidy tbiiu in the li^aded ivli.
It wuuld nf>|X!Kr, therefore, that in the luucoiia cell, an in the pMicreaiic
cdl, the <<v))-»>ibH(aiicc forma atid d^-ixisitd in itself certain material in ihi-
form of gmniilc-!'. During secretion ibo^e ii;rni)ule* iliwippMr anil iin^^uni-
nhl}* f>>rni part of the M^i-retiun. But ihi- crumilvs of u murou< wU dilli-r
t'rnni thiwi' of llic |iiiiicrrJitic ci'll. iniisjniicTi h* llivy iiri- iipt, under the in-
flticnci' if ri'iismlf, t'> hv trnn#fc)rnii;il, while Htill wiibin ihn cell, into the
Irniit-pnrerit vwciii malvriid which we call mucin; hrm.* tlio appearances
prewnlcd by sections of hfirdcncl pliint!^ It Mcem' natunkt to infer that
the grunul<« consist not i^f mucin ilsclf, hnl of a forcrnnucr »f mndn, i>f
some siibettimT! which can give rise to mucin, and which wc might call
mucij^'eD. And we might further infer that during the act of secretion the
;;ramiles of muei^ii are ImnsforniMl into mnwcs of mucin and so discharged
from the cell. Lnd«ir this view the appearances presented br the hardened
glands, &» distinguished from the living glands, might be interpreted aa Ui'
ilieatinK ^''"l under the iofluencr of the reagents eaiployed, tbe mucigeo of
the li>a<led cells hnd undergone the I runs forma tion into mucin tiithoui being
diacharged from the cells. Up to the preeenl, however, it has not been found -
poauble to isolate from the gland any deBnile body capable of being oon-
vert«d into mucin, and there are aume reasons for thinking that not only
tbe gninulea, but part aUo of the substance between them. cuniTihuiai to
tbe ^rmation of mucin. Apart frum this comjilicnttoii. bow<^ver, the geii-
cral courae of events iu the muci>us cell ^eema to he the Mime m in the pian-
crciilic cell ; the cell -substance oiaitulacturea and loads iUelf with u ^jincinl
prwlnct (or s[)ccinl products): during the. act of secretion ihi^i pr^xluct,
undergoing at the lime n ccrtniu nniount ifl' clinngc, is discharged from tbe
cell to form nnri of the secretion, and the cclt-eubstuncc, Mirred up to in-
creased growth, subsdjucntly nitmufnctnrun a new supply of the producL
§ 236, The " ccJilral" or "chief" cells of the gastric glands also exhibit
Nmilar changes. In such iin animal ns the newt these cells may. thotigb
with difficulty, bo examined in the living state. They are then found to oe
Mudded with granules when the stomach is at resL During digcfltioil tlieae
granules become much leas numerous and are chieHy gathered near the
lumen, leaving in each cell a clear outer zone. And in many mammals the
Name abundance of granules in the loaded cell, the same paucity of gnuiulea
for the nioHt nart restricted to an inner /.one in the discharged oell, may be
demousLrated by the uae uf osniic acid (Fig. Illy.
When the ttComach is hardened by alcohol thttie ehaugM, like the similar
change* in an albuminous cell, are obscured by the nhriuking uf the
"grsnuW," or by tbeir swelling up and becoming ditl'ii^ed through the rr*i
of the cell-sub«taaoe; ao tliat, tliough in aeolions ao pri-imn^l vt-ry Mrlking
dilTcrmcci' arc wen between hmik-d and duchareed cells, ihi-sr iirc unlike
th'wc scrn in living ghindn. In >|niciraens taken troin an nninirtl which baa
not l>cen fed f'lr ngmi- ttnn-, the centnil (■i-IIm of the gastric gland* are imk',
6ncly granular, and do not stain readily ivilh carmini:' and otli<-r dyes.
During the corly stages of gastric dig^'Ation, the same cells arc found some-j
what sirollen, but turbid and more coarsely granular; iheyxtuin much mor
readily. At a later stngc tbey become sni.illor and shrunken, but are '
more tnrbid and gmniilar than iHrforu, iind stain still niorr deeply. Thb ta
true, not only of the central cvlli^ in the ennlinc glands, but also of the cella
of which the pyloric glands are huilt u[i. In a loaded cell very liltJe staining
CtlAXOBS IN THE 0LA.VU8.
S45
I
I
l-i.> III
:\
<B»
take* plaoe, becanse the nmMint of living sUltiiinj; vtrll-mbetince b Rtiuill
niatiwiy to tli« mmiot of malcrial with whioh it i> loaded sod which
dooi not Main rtadilv. In tli« cell whidt iil\i.-r
Ifnmt Mtivity hu <lbobufed ilMlf, the cell
MHBdl«r. but «h«t remains ia Imn^elr living
eall MihsUum, Bume of it new, and afl ^lain-
iafl rautUj. li would ap|)ear, also, that dur-
iajt Ibc activity of the evil flume «ulMlauc-€H,
capable of beiiifc precipitaleil I>y alciiliol. make
Utwr appwraDoe. and itiu pre.-i^iu-e "f i)ii»
■atfwial add* to the lurbiil and icmnular aspect
of tbe cell; pii»ibly. nUu, this niiilunal con*
tribaiM tn ihi- *taiiiiiig. A siuiilur niulcrial
■MM! lo maki- ilK appearand! in the oelb of
albaminoiu glandji.
In th« OTOid or lxjnl«ir c*ll» mi vtry chnrao-
tariitia dtangn make llK>tr npix^ranw. Duriii)c
di^lioQ Uwy b4«onw larger, niont vwolU'n, lu*
it mtre. and in oun»«iiiwicii bulge uul thu luiw-
mrai iu*:iiibrantr, hut no cliarncU-riotic di>>n{>'
prNnnrt' of gmnul«» can tie uhi«rrol. In tti«
living ftatv, [h« cell-siihstancc- iif itiov ovoiil
rvlb app«an finelv gniiiiiliir, but in hnrdi-nvd
awl prepared N.-cti^oni hv n cniirMdv ^'raniilar.
" rv(irulat« " ]i»ik. which is prrhii|» lin* innrltvd
ia ibe ivullen active cell» than in tbc reeling
nib.
I S37. All ibcae vamua aecreiin); cella, then
— puKreatic cell, inucouii c-ell. albuminous cell.
MM eanttal fastrio cell — exhibit thenuieaeriea
of Mcala, tnodifted to a certain exlent in the
•wml ouea. lu eacli caitv the " pro^jplaamic "
«*ll4uhatanoo maHufaciurei and lodra in ilaelf
nMafial <teMintd to fonn part of the juice
■ai rnmit In iJw tnA veil this material may
MMralljr be recogsiMd under the micrnjm|ie
DJ its optical obaractore nf gmnul«^; tbaae,
bomtirr, an> apl to beoono allied by reamita.
Bat «e must guard oniwlvM against tne lu-
naptioa that the ninli-rinl which can thus be
rffinpiif' is thoofily matprial thus stored up;
we mar. in future, by ehontical or other means, be able U* diHVrentiaie other
parts of the eell-b<ily as being also nuiU-rial similarly stor^t up.
During activity, while the giniiit tiiM'cri^tinc, tht« mntcrinl, either unchanged
or alter uuder^gving change. i« wholly or partially dlscharge<l from th<t oell,
Tbs cell, in oomequenoe of havini' thim got Hd of more or less of ita load,
(onaisU to a lareer extent of actual living eel I -substance, ibis being to mnnv
amm inoreaaed by rapid neW'gT<-intli, though iho bulk of the discharged cefl
mar be Ina than tliai of tlie loaiied cell.
"hiia adlvitT of erowlb oontinuee al^er the net of secretion, but the dis-
ehargHl cell soun begins again the task of loading itself with new seeretion
tnatrfial hir the next act of secretion.
Thus in most cii^m tb<?re is. corree ponding lo the iniennttteoee of secretion,
an altemniwn of dl^-hiirge and loading ; but it muil be bjrne in mind that
Mwli an allemaliun ia not abaolutely oeeauwry. eren in the ease of inler-
liVttlK IlLAHU (ir U*JIX41|1UT)
DCKlini ACTITITi:. ILUIOLKT.)
t. tli« motrth uT 111* Klftod wtlli ll>
tXlliKl'lotl sella a. Uio nwk. turn-
UUUllUI CaiI>>Clll>IU llTdld (Bill.
wlih tliotrcnano p(«M|,lH>nk iiot-
wurk. /, IhlF botjr iir llie Kland.
Till' r™anlw •>* (Mti In tbuntn'
tiKl ivlb to br iTinltnl lii ihf liintt
luniuiu uf raeh mil. I he rouiiil
i>iw]Viu at It hloh U co(i«plouartB-
840 THE TI8SCBS ANU UBCIIA!fl8MS OP DtCESTIO.V.
tnittotii accretion. Wc cnn cn«i1y imnginc ch»l lh« tliM'linrgc. »nv of
" jiiraiuilcB " (lurin)> wcnl'mn. fihuiili) stir up the coll to iin iiii'rcasml Rctivity
ill romiin)^ ^'cnntile^, ninl ihal tho formnliro activity flioiili) i-eosc wlncn tli«
Kcrclorv activity I't^nrteii, In such a cftee tho number of nvw graniilM furnied
might alwayc be eijual li> the number of old grHiinlo; uhfH itji.and th« active
cell, ill spite of its discliarce, koiiM prwees an miiiiy granules—that i$ to mv.
IIS laree n load— aa the cell at iwU Ami in the ceniral pastric cells of »ome
niiiDiah it wuiild nppear that sucb a cMiitioued balunciuj; nf luad aiid dis-
charge does ai-tually take phtce, to that no distinction in Knioulee cao be
ubwrved between rettin;; and active relU.
§ 238. We s]x>ke juat dow of the material stored up in the cell and
destined to form part «f the secretion ii« undergoing i^fmnge before it wm
dischni-gied. In the iiitiiiius cell we hove ^leen that the maleriHl deposited in
tlitt liviuf; cell has at lini<i the form uf ^raiiul«». TliMe granules, bou'ever,
are en&ily i-onvertcd Into a trHii.-']inrent matea'al lotlged In the sjiaeea of ih«
cell-Rubstauce, nbtch miiioriiil, even if not exactly iileniieal with, at loojl
elowly rtseniblcv, the muriu found in the lucrt-tiou; and ap|>Mrentlr in the
act of secretion llie ^annh-s do uiider^M «ume nucli change. In Uh' <.■■»
of tonic other {;land:<. iiinrvovi'i', vre hav<' chemical ns well as optical
i>videii(% that the inaicriid ttored ti|i in the ci-IIm u, in part at IcuM, nol the
actual »ub«tanc'c apitcnring in the wcretiim, but nn antccck-ni of that sub>
■taiioc.
An imimrtant cnn»tilueiit of pancreatic ^oici: v, iw vrc Hhnll M.-e later un. a
bodv callwi fr^ipna, a Urnivnt very similtir to pcp»in, acting "ii |irotviid
bod'iM and convi-rling thcni into [wptonc and other »iih«tunci». Though in
many r«pcct» alike. prpMn nml tryniin arc quite dintinrl l)ndi». and liiKvr
markedly in ihi>, Ihnl while an acid medium in nci'iTMnry for the actimi uf
ppeio. au nikalioc nicdiurn is nec^iwary for tho action of trypsin ; an<l accnrd'
iii^ly the pancreatic juice is alkaline iu contrast to the acidity of i;aFtrio
juice. Trypsin can. like pepsin i,S 2i)5). be extracted with glyc^riu from
Bubeitancea in whieli it occurs; glyoeriii enlracf^ of trypsin, however, Qee<l
for the maniftfilation of their |><>wer.« the pre-ence uf a neak alkali, sucb aa
a 1 jier rent, anliition of sodium ctirl>i<nate.
?ton', try[uiin is pr(?«nt in ahii udiiiuv in normal pancreatic juice ; but s
loadetl pant-reun. onii nhtch ix ripe for ^-crelion. and which if excitnl ta
wcrcle woulil inmiciliiili-ly pour out n juicct rich in try|i»iu. cniain* no
tryiKiin or n mere [nice of it; nav, even a pancivitt which in en^itwetl iu tbc
•ct tif w;i'r('ting eonlnin* in il« actual itIU an iuKignitii-aiit <pianiity only of
iryiwin, ii« i* sliinvn l»y the fMllowing cKiM-rimonl:
If the piiricn-n» of nn animal, even of itnit in full digrftion. bo imiied.
whilr itill u«Tm/roin thr liody, with glycerin, the glycerin eximcl, n* juilgol
of br it» action <>» fibrin in the prcfencc of wxlinm carbonate. ii> inert or
lioarly ko w regards prolcid bodies. If, however, (he same pniicreaif be kept
for tncitly-four houre before biing treated with glycerin, the givceriu extract
readily digests fibrin and other prolcids in the presence of au alkali. If ihs
pancreas, while still warm, be nibbed up in a mortar for a few ininutca with
dilute aceik add. and then treaietl with glycerin, the glycerin extract i«
strongly proteolytic. If the Hlycerin extract obtained without acid from the
¥rann paucreos, and iherefore inert, be diluted largely with water and kept
at 35" C for rtorae lime, il hecumes active. If treated with acidulated instead
of dmilh-d water, iLi activity io nuieh souiier develii|>ed. If the inert glycerin
extract <>f warm piiucrmn be precipitated with alcohol in excew, the precipi-
tate, inert as a pnUooiyiic Icrmeni when freab, liecomei active irhen ex|)ose<l
fur some time in an iKiueoun folulion, rapidly su w hen truited with iicidiilateil
water. Tbew fads show- that a p;tncr«as taken fn-sh from the body, even
I
hI tillle mad^matlf femunt, lliitiiu:)i l)ior« b
iruo kiu'l of ucoumpcwillAO, jri'i'"' oirt/i to thv
nL Wp RiBif rrmark iiiciik'tiiiitly, ilmi lh<>iigli ehp prcw-in'i' of mi nlknli
!• wtBlMil to the jiTvitrolylic Kctuni of tbr ncliial ferment, the fori imt ion of
Ibv ftrnient out of il* ri^rrruiiDcr i« favorol tiv ihc prvwtnco uf ii vmnll ■(unii-
tilT of acid : llM-aciil muil he uwal n it h Litre, «iiico the Irypein, »iiee fonixnl.
M de»ti»rril bv acti)!. T» lliis tHiilv, this mother of the formcnl, which hiu
uA U pi>«ent been MlUfActorilir iMlatcd, bnt irhieh «i>pciini to be (t coRi|iIex
body, Fpliiiing up into the fernteni, nhich, as ire hitve »ea, is iit all ovcuie
m4 cmainlr ■ pr<>1eid body, and into Hn undeniably proteid body, the natne
of 9«Mi^«n niu be«n applied. Dm it is better to reserve iti« term zym>oeeD
m ft K«Derie name for all suoh bodiw aa, not being tliemKkee actual fer-
n«al«, IB«T by internal chanjfes fpve rise to fements, for nil " molhere of
fcrmtrnt " in fart : and to fj've to the [uirticiilar inoilier of the pancmiic
l>ni(ei>lnic ferment the name liyp'iuoj/en.
Cvideoce uf u aimilar kind nhoui» thnt the Kiuirio Riaiid*. botti the L-;ird!ac
and tbe pyloric jflaudi, nhilt^ they cotituin ccim]mratiirely little actual
p«pNn. csMitsin a considenihle (|utiu[iiy of n lymogen of pe[nin, or pri-tiiw
ym : attd there can be little doubt but that thia pcpnuojfeu is lodged in th«
evnlml ceJIti of the canlinc t;t))ndi and in the aomcwbat timilar cells which
line the «h<d« uf (lie pyloric ^landi^
It ia fkirtfacr inlvrMting to ntiMn-e that, iia a g«neml nih*. the amount of
tryf«ifK>^n in a |iancri-aa ni any givwn tim« rttc* and txwVr pnri pattu with
tbe nmniilnr innfr /one, '. r., «'ilh the aumunt of gninulcs in the cell. The
aidrr the inner tone and the nion- ahtindant the gmnulm ihn laricer tbe
unuant, tbe namiNcr ihe xonc ntid the fewfr ihi* gntnulcs the smaller the
ftMMiut. of trffwinoLfn ; and in the nufe» of old -■■« tab limited Rtitultv, wIm'TO
lbs (vrri'tion m wholly inert on proleiilii. the inner grnniilnr kioo u alnonl
fttMB the eelli. AimI the same paralleliain Uiu Wen olidervcd between tJio
aboadance nf ^ranalea in the central celh and the 'juantity of pepsinogen
prvarnt in tbe gitslrtc glands.
Tbe [uarallelisni, botrever, at all events in tbe case of the iMiHtvm.
ftppMUV nut to be ab^ilule, for it ia Maietl that in tbe paucrenH of dng« af^vr
InOK lUrvation there i;) little or no iry|(>iuogen in tbe gland and yel (he
odb exhibit a niarktil inner n>ne of i;nimileA. Moreover, ue should not. in
•nj eaiM>, bejusiilied in concluding ihnt the )i;mniileM of dte pancreatic cell
ar* whfdiy coni[>oM:<l uf (ri'|Rinogen : for, lu we ihall pre^otly aee. the pan- ■
rreaiic juice coiitainii l>eittde!i tr^iwin not only other imniirtant fernwrna htil '
a In I certain {irulcid conslituuita; and the granules, which are of a i>mt«itl
uUurr, prolwbly Hujiidy (hrw prolcid* of (Ih' juice. Henoc the [Miriilleliitm
between gniuulm and tr^'pinflgni !it at Ixvt an incomplete < me. Itnt even
•ueb an incomplete pMrallclbm \f. of value. Th« granule*, whatcwr their
tuklure, are product* uf the nictalxdium of the cell, lodged for a while tn tl»e
7«lliib*tancv hot ewntuallv dtfcharged; and ccnain of (he c«>n>tiiuent«
of ibe M-vcral tecrction*. «uc1i m mucin, (rypvin, pc|iein am) ihe tike Bp|K-iir
to he in a •imihir uiiy products of the mflubotitm of the cell, l<Hlir<rd for a
while in ibc cell Mitntiinc«. not in all cftim exactly in lh« condition in which
tbcf will be diKhargcl from (he cell, but in lui antece^lcnt phow »ucb ax
ayogeu or the like, and iu all ciues ultimately ejected fri>in the cell, to
•nbpl*- part aixt ^'nenilly (he imp'irl8n[ pan of Uw secretion.
t 8i9. Tfit itH of ttrriiitin ilt'lf. The above dtscutvion prepam ui at
ooM tot (be MairnH'ut that (be old view of fecretion, acoordin); to which (Jm
l^laad picluont, separatee, secretes (hence tbe nan>e secretion), and to filter*
•■ t( were from the common store of (be bloo<I the several oonaliluenU of the
nice, ia uoleiuible. According to that view (be specilic activity of any on*
I
i
I
I
848 THK TISSUICS ANU UlfOnAKISMS OF niOKSTION.
gliiiid WAS conliu«d to the taak of letting VLTljiiii mngiitueiita uf tlie blood
p*B8 from tlie i^upillarita Hurnftindiiifi: ilie alveoluM lliri>Uf{h the cells tu the
cbannelaof itit^ ilnt-Ui. while n-l'uaiiiK n imwiiii^ to otlierit. We iiuw kiiuwr
lh*l certain iiiiix^rluut i.-ui»(itut:iita of tiacli juiee, tlii' |Hnwiu of ;;adtrir jtlioe.
the luuciu uf iuili<.-ii unil iho liku nre rurmuil in the i^i-li, atnl i)i>t obtAJned
ready made frum lli« blooil. A mintit^t i|UHOtily iit' [H'juiii dii«> exidt !t b
true ill the blood, but th<'re nn: n*»"nt for thinking! ihni ihl- liu* tniule iu
WKjr b«<:k into thi.< blixul. ).-ithiT \».-'m^ nWirln-d from tbt^ liitt^rior of the
Btoinai'h i>r, u« xn^nu innti- priil>iible, pii'krd u|> dinrctly IVoin lli<! ]|^trio
Cluida 1 und «m witb mime of tliv other iDtiiitititciil" of other jiiiitts. The
t'hief or siwelfic eon»tit>ienl« of wieh juice lire formeil in ihc wll itMlf.
Hut tht! iuioc Bocrotcd br any glatid ooiu>i«tfl not only of tb<- Mpi^ltic Mib-
stuticu* Niion ail mucin. |)cp6iii. ur other tcrmciil. or other b>»ti«. found in it
alone, but also of a larg^ i|unnlity of water, and of vuriuuv other Mb-
nianotw, chiefly inline*, cotntnoii to It. tn other juices and to the bloo). And
the ijiii9'lii>u ari»e.f. Is ibe irater. are the saltA and other common Bub'tauco*
fiirnmbcd by tile same act ns that which supplies the specilic GoD»tiliteHt« *
(V-rtjiin facts au^roat that Lli«y are not. For instance, m meiiliuiied fomt
linw a)^i, in lh« auDRiaxillary ifland of llie dug. stiiuulatiaii of the chorda
tym|»ni produces a copious (low of saliva, which is usually thin and tinipid,
while Klimutatioiiof the oervicaJ sympathetic produces a scanty llow of thick
vikfid Mliva. Thai is to aay, atimulation of the chorda has a marked cifed
in prorootJn<; the dboharge of water, while Blimulation of the sytnpathelio
has a marked effect iu prumotinK the difcharKe of mucin. To this w« may
luM the caaw of the parotid of the do^. Iu lliii> K'*^od Btimulati»u of a
cenbro-sptoal nerve, the uuriculo-temporal, prodtiees a copious flow of limpid
•alivK, while Mimulaiion of (he sympuiht^tii- proilucea itself little or no secre-
lion at all; but nhen the nymjMtiliutie anil the oerebro-spinal nerves are
Htiniulated at the name lime, ihi' mlivu which Aam is much ticker in sulid
and eftpecially in organic mallrr than when the oi'^rebro spinal nerve is stimu-
lated aloue. And we have nlrcjulv iwcii tliat in t hi* gland the microscopic
dianges foltowio)^ upon »ympathi.tie !«timulatii>ii arc more conspicuous lluui
tboee which follow ujion <x>rehro'i'piiial ncimiilnlion.
These and other facU have led to tha corici^pltoit that tlie act of aecretloii
cuiuiata of two ]>arls. which in one oiw may coincide, in another may take
place apart or in diKercnt proporliuiu^ On the one hand, tlicr« is die t)i«-
cliarge uf water carrying with it common eolublv substancH, chiefly "aliacs,
derived from tlie blood ; ou the other band, a metabolic activity of the oell-
aubBtaoc* ^vce rise to the speciHc constituents of the juice. To put tbo
niait'T hronitly, the lallcr proce« produces the Spocilie conxtiluentis the
i'liruier wa«)ics ihe^e and other matters into the duct. It has been further
ippoaed lliat tno kioda of nerve libree exist: one governing: the former
•nd, in the case of tlie submaxillary aliuid for inilance. prei>on-
cratini;. though not U> the total exoluMOu of the other kind, in the (chorda
tympani; the other ^nveming the latter iiroceos and pre|>onde rating in the
branches of tlte oeri-ical syinpetJietic. 'Iheee have been called respectively
" tAcretory " and " tnipbic " librea : but these terms are not desirable. It
may be here remarked that even the former prue«u is a <iistinct activity of
tlie gland, and not a mere iriRltration. For, mt we have seen in the caw of
the salivary j^laiid*, when atropine U given, not onlr do the Mpeotfie cunfititu-
enta ccttse to 1m: ejected iw a council ueiicc of Ktimiilatiou of the chorda, but
the discharge of water, in spite of tlie bluiHlvtwM'ln bccominjE dilaUrd. in alMo
arretted : no Niliva at all leavco the gland. And what is tnie of the salivary
IcInmU a* n-gards the ihixjiilfuue uf the (low of water ou sonielhing cIhc
liaidcs) the mere pretMirc uf the blond ia (he bhwdveMcU, ap|ic4tr« t^i hold
CHAKOES IK THK OLAKPS.
i) wit}i oilier ^■IniidH aW Indeed il bua beni ■ii^'trf*I('<l tlmt i)>i- wry
(liflcburge of wuler \» due lo nn aclivitr of (he cell ; llio hyjxittiesis liii? \>vrn
put forward (bat i-liiini;eit iu lli« mU give riwe lo tb« fnrniiiliaii iu tbii crll of
aubelances nbich nlvurb water fnim tbe hloiMl or 1yni]th on lh« mu: nido
and p\t it up on (he other ud<- into the liinM^n uf the iilTeoluft. (•iii-h no
b]rp(>tbeniii cantiut h« regard«il a* nrovccl ; but th« Di«re ixitliiig it forwiinl
nilsea doubt* t\» li> lh«- validity or tlu) distinctioD on uhich wo haw bcvn
dwellino;; ami other rtiiinidi'rationK point in th« itanie direction, For in-
tiaiuv, if the common Miluhlc )«lls jinycnt in n juirc, nn <ti(>liiiguishi-'l from
(he ■]i4-cific (itnntitiiciili', wrm iiifn'ly curried into ihr jiiicc by th« rusli, «o
to H[icak. of wut<-r, wt- nhotilii expect lo ttiid llir pereentiLg« of tbcM) ults
rillur nninining llio wimo or perh«]M> dccn^iwing when the juice wjw ttccreted
more rnpidly nixl in fuller volume. Kut under (hew drc-utnEliincev (he per-
C4-nIiige ven>' frcr|uently incmutn: and in genrrnl uc lind cbnt under various
drcLi»i»tiiiices tliP proportion of ealta Bccretetl to (he quantity of iriilcr
M-erctcd mny vary constderably. Obviously, nhilc sumething determines
the 'pinniit}- of wnt«r pusing into ibo alveolus, something eloe determines
bow tDHcb of common »oluble enltt that water contains, and Still something
else determin^B to what eilent that water is also laden with 8p«oi6c coostitu-
eni« and other organic bodies. Tbe whole action is too complicated to be
described as conebting merely of the two proeeeees meniioued above, but the
time has not yei come for clear and delinite statements. Everything, how-
ever, tends lo show that the cell is the prime agent in tbe v bole biiainew,
iJiongb ue cannot ai present define the nature ol the several changes in ihe
cell, nor can we sav him these chenge« are exactly related lo each other, to
changea of the bfood-prcMure in ihe bloodvessels, or, we mav add, to
chnnges taking place t» the lyniph-9pace« which lie lietweeu (fae^locHl aud
the cell.
We inajr perliap* add that, vinre in certain culaneous secreting glands the
alveolus, or what eorre«ponda to the alveolus, is wrapped nmiid with plain
musctilar fibreK, the contraction of whith upixnra to force th« secretion out-
ward, the idea hiiti been suggcMed that in glands, such nn we are now con-
sidering, Ebe oeU-*ub*tance making uve of " pixKoptaxmic " contraction in-
stead of actual mui>cular contraction, may force part of the cell contents
into (be lumen of the alveolii*. Such a mode of wcrelion woviUl lie ci>m-
parable to tbe ejcclion of iindigtvted maierial, or " vxcrvtion," by nn nma-bu-
Kut we have no witiafuctory evidence in favor of thin view.
S 240. Throughout the above wc have wjokeii tu if the secretion were fiir-
Disfatd exclusively by the ceJls «f the alveoli or secreting portion of the
S'and, as if the epilhelium celU lining Ihe ducts, or couducting portion of
e gland contribulcii noihitig to the act. In Uie gtutnc ghtmU tb« slender
cells lining Ihe mouths of tbe glands (which correspond tu duels) and cover-
ing the ridges between, arc mucous eclls sccroling into the stomach gener-
ally a small, but under abnormal conditions a large amount, of mums,
which haa ils ueea bat is not an essential pari of tbe gaxiric juice. In the
salivary glamU we can hardly suppose that tlio long sirelch of ehanicler-
istic coluuiuar epilhuliun) which reaches from the alveoli to tli« mouth of
tl>e long main duct serves simply to ftiraish a smooth lining lo the conduct-
ing Mssages; hut we lisve as yet no cleflf indications of what tbe function
of tills epiU>etium can l>e.
§ 241. Before we leave the mechanism of accretion there are one or more
mrcMHiry txiiiils whi<;h deeerve attention.
In In-aiing just now of the gastrio glands we spoke as if pep«n were Ihe
only iin|K)rtaiit conaiitueul of gastric Juice, «fa«ms, as we nave previoualy
seen, the acid it etjUBlly esseulial. Tbe fonuBlion of tbe free 8ci<l of tbe
B
TlIK TISSCES ANU Ml
KWlric jtiioo U i'«ry uWure. nuU luaiij injftniouB bul tiusalMlactorr vi«irt
luive bo«n |nii foruanl lu «xplaiii it. It tteema uaturnl to sujifHr^e tbat it
nriaeH ill aonie way from the deconi)i<>i«iti<iu nf Hoitiuin chloride ilrnwu fnia
lti« blood ; ami tait h xupportMl by iL*:' I'ui-t that uUkd the aecretiim i>l' k*^-
trie juice i.i aL-tlvi-Iy tciAii^ on, tlic niii'iuiil of clilnridM leaviti); the Muoil
by Uie liiilnvj^ u [iru[>orUi>uHldy tjiniiii'igilieil : liiit it>ithio^ deliuitv mil M
prMenl be MMtxl ui in (he luechiiiiiKni of ihnt il(!i-(iiii|H:ditiou. An<t even
■dmlttiiiK thxt the wdiuin chloride of' the hvly ai tarjic i* the ultiriuil«
■ource c)1 the chloriiKi eleiticnt of ibc aciil. It uppi^nni imire likely tliut that
element Kitoiild be t*l frcv in the Htumacli liy thn tli-coiuiXHiiliiHi i>l' miiot
liighlv complex and uiuilithle chlorine compound ]>rcviotiily 2fncrnt<.-d. than
llmi it should arise hv the direct splilling-iift i>l' w (tabic a Wlr n* Kinlium
cblorido nt the very tiino nhm the acid ie nx-retcd.
In the fn>g, while iKpfiiii troc tVoin aciil if i^ecn-ttil by llic g)iin<ls in the
lower {xirtion of the irMijihatjus. an acid Jliim- '» uffinled by gland* in ili«
atoiwaL'ti ilBcIf, which have accordingly been cidlcd o^gntic (^'Vi-o to sharpen,
ncidulaic I glands ; but ihrac oxyniic |>lim(U appear also to secrete pepsin.
In the niainmal tlie Uolnlcd pylonit «ecrct«8 au alkaline juice; in tact, (be
anpearnnce of an acid juice is limited tu tbo»e iMrtions iif the stomach in
u^iich the glands coDtaiu both "cbici" or"ceDtml," and "ovdd" or ■■l>i>r-
der" ccUb. Now. flrom what lias been previously Mid, tliere can be ik>
doubt (hat (be chief cells do secrete pepain. On the other hand there is nu
evidence whatever of the formation of ne|aia by the '■ border " or "ovoid "
eellH, though this was once suppoaed to ne the case, and thcae oelU were un-
fortnnntely formerly called " pe]>iic" cells. Hence it has been iufern^ thai
ilw bonier oelU fti'vreie acid ; Imt the arKUiiieiK is at ]>rewn( »iie of exclu>
itigii only, (here being no direct proof that thaie cells actually numufitctun
tlic acid'.
The rennin an)ieani to be formed by tlio same oellA which manufacture thi
|Mip*ln, that is, by the chief r<-ll.-> of thd fuudun gi'iicmlly. and to »omo ai
tent !>;■ the cells of lh« pyloric glands. We may mhl that we liav«f ovidMlD
of the cxiKieiK'* of A lyinogcn of roniiin aiinlogoun to the lyaiagi-n of [lepaii
or of trypnin.
The muoiis which is prei^ent as n thin layer over the surface of the fiutiog
fltoitiach, and which eitpccially in hcrbivoroiw animals, is increased dur~"*
digestion, comes, as we havr> siijd, from the niuoiis veils which line
iiiiHitli^ iif the Hcvcral claniU and cover the intervonitig eurfac«S.
!; 243. We previoufFv callc^l ntli^ntion (o the fact that in the cose of the
oraach the afasorplion of the product* of digestion largely increased the
elivity of ihe iwcretint; colls. This has led to (he idea that one effect of
fixnl h to "eliari^" thej^slric celU with pepiiiogen.and that eerlain arliclw
of fiKHl might be considered as eipecinlly jieptogenous, t. '., conducive to lli«
fomiatiou of pej«in. .Such a view is templing, but needs as yet to lie more
fully siippiirCed bv facts.
§ 243. i-^N!tng the gtvat solvent jtower of both gastric aud pancreatic Julc
the tpiitxlion is naturally fluggmtml. Why does not tJie stomach dtgett it«clfr
AA*!r di-nih, tbo stomach is tretjiieutly found iiarcly digeated, vii., in case*
when death has taken place suitdeiily on a lull »toma4'h. In an ordinarf-
diiilh, the niemhmne ceasea to aecret* before the circulation i." iit an enttj
That ihrrc is uo niirctal virtue in living things which pn-vent* their beini
digi.-Kt^'Kl is shown oy ibe foct that tlio leg of a liviu^i; t'rog or the ear i3
IV living rabbit intrixlucetl into the atomach of a dc^, through a ganrioj
fistula, is readily dig««tvd. It lins been tngEOslcil that the blooil-current'
kwjm up an alkalimtv lulKcicnt to iicuinilTxe the ncidiiy of the juice
in Ibe irgioo of the glaiidn Ibemselvtv; hot this will unt explain why the
I
Mtii! jiiici.'. whirli in iiiriivi' ill nti iilkaltoc neilitni), iIim-* iiiit ■lii.i.'wt iliv
ills o1' lin- jmurrcii* it»":lt', or «liy tHo diMattvo wIU »f tin- lituodlcss
uciitiu»Kiii or hy<iiv»"in <ii> not Higcrt llK-nw^nri*. \Vi- mijrht ikIiI. it iliwn
not exriinin nhy lh« iiinirbii, whik- diMoIving the |iri>tn| ilium or (lit- dvrnU
lowed iliiitoiu. dim niA dissolve it« own )ir»U>put>ni. We mniiot Amnrvr lliLt
question u alt tit |ir«BeDl, itiiv mnn.^ thnn Ihe iimilnr one. wliy Iho delic«l«
lirntoplasm uf the iniwba r^wiste during life tlw entninc-e into itM.Of liy uemout
of more water tlinii it re*|uiree to carry on its work, while n Tew nioineot*
after il is dend natereDtenfreelr hyosoioel^, audiheedectsof that eiiimooe
l)«vome abuiiiIaDtiy evident l»y the fomialioii of bullw and the brenkiuK u|>
of tlte i>r<>io|t1iigtu.
Toe Propkrtiks asd Charactees op Dile, I'axcrbatkt .K-ice and
SuOCUfl ExTRlilCTS,
I 844. In (he Hvin^- hmly the titotl. Btibje<1ed 1o the artinn first of tlw
mltva and then «f lh<.- niutriv jiiir«, underKovn in lli« stomafh i-)mu)!;i,ii whi<rh
wf ntiall (irvwritl)' ('i>i»id4-r in (U.>tail, and tin- foud mo cliiiii^'^l i.« iiaAMed on
into the iminll intn>tiiii', nhrn^ il iii furlhi-r iivihj«i<t«d li> thi: B<'tii>ii iif tlK- hiln
iM-crotwt hy the livi-r. of iNincrvatic juice .■xn.tvsIi^'I h^- the piincreiw, and prw.
sibly to »»nK- rxli-nt, though Ihiit hy no nicium certnm, of a jiti'r ««rrcti.-d by
the intestine iiM-1f and cnlTed luwu* tntrriftu. It will be oonvcnionl to iftudy
thv ininiile«lriii-iiire of the liver in cuitneet ion with other fiinctinm of the
liver more inijiortiiiit. perhup*. than tJint of the M-crotion of bile, nnnivly,
Ihe fbrmali'in of glycogen, aiid oth^r nietnbiflic eveni« occurring in the
hepatic eellt; we have already ttudied (he structure of the rancrens: nnd
the etructiire of the intt«linc will betil be i-onsidoivd by ilselr. We. there-
fore, turn at once tii the ])ro])ertiee and characters of the above-nanicd juice*.
Bitr.
Tboujfli bile, after secretion in the lobules of the liver. It puted on nlon^
tlie lK-|Nitic duel, it in in the case of uinat animals not poured at oncp into
tlH- dumlenum but taken by the evstie duel to Ihe i««erv»irof the j^uU-
bladder. Here it remain*, until suco time ai it id needed, when a ijuiiiitiiy
b tioured aloii); the common bile duct into the intestine.
The 'jualiiy of hilo vuritn much, not only in dilfbrent animalit. hut in tlie
name animal at diirvrt nl titiico. Il i-" moi-covi-r airecli-d by the li-n^lh of the
•iijoorn in thf gnlMilnddi-r : bilr Inkc-ii direct from the be]Mitii: duct, <v|M)-
cially when «ci'ri'lril rii|iii|ly. ivininiii* little or ik> tnueiu; that taken fr>>ui
tite gall'blailder, ati of ulinif^htcri-il oxen or i>h(^'[i, \» loadwl with muciu.
The col'ir of the bile of caruivoriiiiM mid omnivorouii animals, ami of man,
J [s gem-ratly a brigin i^olihn red ; of herbivorous aniniul», n yellowish gr««n,
or a bri}:ht green, or n dirty gravn. according to circum«tnMCw, being much
nii-dili< <1 by n-tenlion in llic gall bindder. The reaction in nculr>il or alkaline.
The f-'lJifwinL' niav 1h- taken as the average composition of human bile taken
from the gairbbdiler, and ilicf<>rc containing much more mucus as nell na,
relatively to the AoIid«. more water tliUD bile from the hepatic tluct.
Ill ti>ni«n>.
Water ,-...., IWli.a
Ho)>iU:
Ilile.^uii» ....... 91.4
Fai».«lc. , , , . . ».«
Cliolwleriti . . . . S.8
Mnciu nixt rt|ini>'iit 20.8
Inorganic IIniIi> , . 7-^
S52 THK Tiastjj
lOKSTIOl
The vatin abMticv of prolcids it n mnrkcl luilurc of kil«; (ttincnnitic
juicv, ii» «« ehall era, coJitaiDs a coDiitlcnibte ijtinnlUT. mlJva, m wr havu
Been, d small <|uantily, normal guetric jtiioe probnl>ly «iill lew aikI l>ilc noiia
■I all. Evt'n the bile which has been retained eonie time id ihc gall-bladder,
tliOBgli rich in niucuB, coiitains no proleids.
The cooUitueDta which form, apart from the mucue, lti« great bulk of the
Rolids of bile, and which deeerve chief alieolion, are the pigmeoif and lh«
bi1»sRlta; of ihcK we eball apeak immediately.
With regard to the iiiorgauic aalta actually present as auch sodium salts
K» compicuouM, flodium chloride amouutiu^ to 0.2 or more per cent., sodigia
pho^hate to uearly u much, t]>e rut beiug eartliy phosphates aud other
imitt<-n in unM i|uniitity. The presence of iron, to Ihe extent of abom
O.OOB |icr c«nt., U iDtereMin^, since, as we !>liall tee, them are reaaoos fc
thinking thiit i\u- iiigtiient of bile, itself free from iron, ih dfri\'e<l Trom iron
holding hii-iuoKlohm ; woine. at Wst, of the in>u net free during the cmver
MOB of bBaaoglobtD into biln pigment, wliic^h pmbiihly takm plane tn the
llTer, finds its way into llit- bile. Bile ik]«> ariiH-an to ixuilain a •inuU •|uan-
tity, at all cvcntx oc(7i»i<>nBlly. of »lhcr m<-Utl«, such as mansuicse and cop-
per; mrltiU iiitnidiiccd into l)ie hoily art' apt to be retained in tile liver and
evenluiilly Icnve it by the bile.
The imuil quantity of fat pruicnt consiata in part of tbc complex bodj'
lecithiu-
The peculiar body thotffterin, which though fatty looking (hence tlie Dimt
" bile tat ") is really an alcohol with the composition 0^11^0, is conspicuouLi
by its quantity and constanev. It forms the greater pan of most giUl-MoDes,
though some are composed cbieflv of pigment. IniMliible in water and eoM
alcohol, ihouf^h soluble in hot alcohol and readily soluble in ether, cbloro-
forni. etc.. it is dissolved by the bile-salts in aqueous eolulion and hence la
prcwnt in solution in bile. Its physiological fuDctions are obscure.
The ash of bile conuslB Itu-gely of soda, derived partly from the sodiun
chloride stid partly from the bile-salts, of sulphates derived chiefly if not
wholly frum the tsiter, and of phosphates partly ready formed, and iu part
dvriveil fnicu the lecithin.
§ 245. I'itpnenU uf bite. The natural ^Ideii-red color of normal human
or oiniiviirous bile is <Iue to the j>re9ence of bilirubin. Thitt, which ia alao
ibc chief pigmeniary eoDHlJtiieiil of gall'iitoue«, and occunt lar|^ly in th«
urinsuf jaundice, may be obtained in the form either of an orBn^oolore<l
aroorphoas powder, or of welt-forniHJ rhotDbi<^ tableti* and oritinis. lu-
•olublc in water, and but little iwluble in I'tbi^ und nicciho], it iti rcAiUly
soluble in chlonjfDrm mid in alkaline fluidx. Its coiniiiwilioii i» C,tH„N,0|.
Treated with oxidizing »£«»!», mch m* nitric acid yellow with uitrouv ncia,
it di(|)layifainiccnsionof coloTH in tho urdor of the xpectruiii. The relloaitih
golden-red becomes srran, this a KrcfDish-blue, th<-n blue, next violi^t, after*
ward n dirly red, iind Rnally u pale yellow. This cbn meter i»tic n-action of
bilirubin is ihc biieie of the K-callcd (imeliD'ii tott for bile- pigment*. Each
of these Mngvs reprcM-ots u distinct pigmeiiturr vubslance. An idkalino
■ululioM of bilirubin, cxpuwd in a ehallon VMsei to the action of the iiir,
turns green, becoming converted into bitirerdin (C„H„N,Oi or C„H„N.O^
Maly). the given pigment of herbivorous bile. Bilivorain is also found at
times in the urine of jaundice, and is probably the body which givt* to bile
which hns bceu exposed lo the action of gastric juice, ss in biliary vomits,
its chiamclcTielic green hue. It is tlie tirst time of the oxidation of bilirubin
in Uuielin's test. Treated with nxidiiiu); agents hiliverdin midh ihroueh
tlie same series of colors as bilirubin, with the exception of the iniiwl
golden -red.
BILE.
358
$ S48. Tkf. bile-so/lA. ThflMO0iuiM,tn nauaBd UBOy animaU, orWi'tin
g/tfroekotttf: and taio-ofJiolaU, the proportion of the tiro varrlDj; in (liilerent
lUiiiiiklit. In ninn bolh tlii! total t|uaulity of bile-Maltn and the prftjiortion
iif tli« niic bilt^-Kult to tliu (ittivr mh.>iii lo vury ii^^d dcjil, Wt thc^lTc^icliolale
n im<\ to bv alKnya tbi; muni iihiitidiint. In ox-^iill tuilium glyi-ixTlioUti.- in
alwDdant, nnd laiirochQliUv Kcnaty. Tho bile-mlts of the il(>f(, mt, boar, itii'i
other i-itrnivora <y>n»iMt rxcliiHivcIf of the latter.
Insoluble in ethor. but mlublo in nlcohni mill in wittvr, the Mqueous nulii-
tiuiM having a decided allcAlint? ronction, both wilu mn^ be obuined by
cnittiilliution in Rnv nciculnr n«edle4. They nre «xoee<lin};lT ib-liquiiicent,
tdv Mlutioiu of both acids havi< n dextro- rotary action on [Hilnrinil light.
AyporafMM. Bil«, m)x«d with animal charcoal, la evajporatod to dnrnrw and
•xtncidJ with aloihol. If not coIorlcM. th« ulcohniic nitrate miwt bo fiirtlict
deootoriiccd witli ntiimiil charcoal, and the alcohol diatilled iT- The dry reaidue
ie treated with abiwluie alcDhol. and to the ak''>haUu filtrate anhydrous ether la
added as long as any precipitate Is rormed. On standing l)ie cloudy ]>rocipiut«
bocooiM traiMfermed into a crystalline mas* at th« bottom of tlto ve«vl. If the
aleohol be not alMolute, tlin crjoliilii are v«jy apt to Im chanmd Into n thick
syrupy fluid. Thi* maas uf cryitaU baa been often Hpolcen of as iUi'a. Bi^th »alte
ar« Uius predpflated, so that la such a bile as Uiat ol tLe ox or man blllD coa-
■istH boib of eodiam f^lyoocholate and sodium taiirncbo1nt«. The two may bo
Mi-lnKiiliil by |)rcrii>itauon fmra their aquoou* solution* with luirar of Icuil, which
thfiwA dnnn the fonner much more r«adUy than the latter. Tb« acids may be
se|>ur-jii(-ij from tliclr raR>eutive uilla by dilute sulphuric acid, or by the action of
lead Ac^tlflte and sulphydrlc acid.
On boiling with dilute acids (sulphuric, hydrochloric), or caustic potniih
or bar^'ta water, glycocholio add in split up \nu, eholalk' (cbolio) acid and
f^lycin. Taurocholio acid may Himilarly ite itplic up into cholalic acid and
Uurin. Tlius :
4R)rroeliciU(i a(M4. CboUllaaoU niyidii.
C>H„NO. + II,U = C„H„Ot + OH,,NII, ia).OH).
TkaroGhollv Mill, CbodsUcatdd. lUiiila,
C^UuXSO, + H,0 = C«H„0, J- cn,.N HrSO.H.
Roth acids contain the aBine oon>nitrogeitous add, oholnlio acid ; but this
Mid \» in iIm! firu case awociatcd or conjuMtcd with tlie impnrlnul nilro-
l^uous body glyoin, or amidn-aoetic aciil, wliiuh is a conipuunil formod from
■unmonU nud one of tlie " faitv acid" iH.'ri4!e, vix., acetic ; and in the second
MM with taiirin, or am ido- bar ili ionic acid, that ix a rarunound into nbioh
reprcacntflliviw of ammnniu, of the ethyl gnmp, and of nulphurtc acid enter.
The deoompoeitiflQ of the bile aci<la into chololic acid iiaa tiiurin or glycin
rvefwctiToly take* place iinlurally in tne intcetinc, the gtyciu and taurin
Lbving probably absorbed, m that from the two acids, aRor they have e«r\'cd
Itbcir purpott in diireaiiuQ. the two amnii^nia coin|munds are nturued itil'> the
IiIoikI. FmcAx of too two acids, or cbolalic iieid alone, wlion Ireatett ivith
sulphuric odd and cnne-sugnr, giveaamagnilicunt purple color { Pelceukuler's
IMi), with a cliarocUjrislic spectrum. A HniUnr color may, however, often
be prodncol by the notion of the same bodiei on ulbumin, amyl alcohol, nnd
eome other organic b'.H}ic«.
$ 947. Adion of bile on j'ocni. In mate animnb at least bile contains a
ferment capable of con verting starch into sugar; but its action in ibis reapoa
is wholly Hibordiuate.
On proteids bile has no direct digeetiro action whaterer. but being, gen-
erally at lea&t, alkaline, and often strongly eo, tends to ueutrolixe (he acid
354 THE TI88UK8 AND KROHANISUS OF DltlBSTIOir.
oonUuta of Ibe slomiicb aa tbejr ftm into the duodeuum, and, u «• thall
•M, » j>r«pai!«« the way for the acUon of the inncnealic julc*. To peptic
notion It is disiinctly tiniaf^iiistic: ibe {iresence of a Miflidirnt <|ua»tity of
bik reudere K^^n*^ juice iuert toward [irotddti. Morcurer, wh«n btk-, or a
eolutkin of bile-8ait«. ia added to a fluid oontaiiiing ilie jiroduviii of giMtric
digtiriJon, a prvdpitate take« place, voDiistins of ]>arnpi'i)tMiin (wh«n pn^ent),
p«pto<u% {M-iKiiii, ajid bilv-nultA. The preoipitatt! i» mbiuuilvn] in an cxcos
of bile or wolmiou of Mlcnnlta ; but the pepain. tbou);h n-ilimolvcd. renuaina
inert toward proteidn. Tbw |>rvcipiiaii<in nntiially <liiii> lake place in tbe
duodciiiini, aiid yiv ^linll ipiuik of it ngiiiu Inli'r on.
Wilh regard In l)i<- action tif bilv on liiU. tl)*- following statemonte may
be innd« ;
bile bnti a xliglit wolnrnl itctioo mi fntx, ns seen in its use by paintera. It
bail by ittelf a ilight but only Flight HDnlsifying power: a inixiure of nil
and bile sppnmK^ after »hakiiis rather leas rapidlj liian a mixiur* of oil and
water. With fattv aciili> Ink forms oonps. It is, niorMver. a solvent of wilid
RiMijM, and it would aiijicnr tlttil the emulsion of fnts is under certain cirftuai>
UMKea at nil events lacilitJited by the preseooe of soapfl in solution. Hvncie
hil« ia probably of much i^realer »ae aa an emulsion a£ent wben mixrti nilh
paiicruilic juice tiiitn wben acting by it«lf alone. To ttiin pnint we sliall
rcluni. I.aiitly, tbe paiwagc of fats through ntcmhraiiea in asuiMiNl by welting
thi> mcnibrnueH Hitli bile or witb a solution of bile^uilta. Oil will |>aiw to a
certain extent thmuxh a 6llcr'paper kept wet with a wiluiion of bile-enlu,
whcreiu il will not paaaor pasae^ with extreme dilHcuIty through one kept
oonatantty nc-t with diMilled water.
llitc pijiwwi's Koiae autiKKpiii- ((ualities. Out of the body its proMiioa
bindera various (mtrffai-tive [iroetai«e« : and wli«» it ia prcvenlwl from flow-
ing into the alimcnturr cjuinl, tht^ coniitnta of the intwtinc undergo cbaueee
diflerenl from tbow wdicb tnlcc \>]»w un<lcr normal eondiliona, and leading
to the appaanuwe of vurioun prodiictji, ffiiecially of ill-emeltine gaaea.
Theae various actiotm of bil« tf^m to be i^a^ndenl on tbe hile-solts and
not on ibe pigtueiitnry or other Goii»tiluciiu.
Pauetealie Juiee.
^ 248. Natum) benllliy pancreatic juiee obtained by uiettna of a (emt
pnnrrealic tiatula differs fn>m tJie digestive juiew of which we bavc al
H|H>k<'U in tbe coinjttimtivctv large quantity of proteids which it cuntaiii
Il« i^mip'wition vanca at'ctirdiiig to the rat* of w>cretion, for, with the mo
rapid flow, the increnso of lolul wlid.-i dm-M not kiwp jincc with that of the
wai«r, though ibe ash rcauiiiDt rcniurkahly iiui:>tant.
Ity an inciaiuD through the linen allui Ibr jiancrratic duct («r ducts) can easll;
be loand either in the rabbit or in the ilug. uod u canuln Hectin-d in it Thate
no difficulty about a temporary llatula: but with permanent fintuliu tliv oocretio.
ia apt la breomr allt^rcd ni nature, and to lose many of Its cbaracteriBik proper
tJM. t^nie. liiiirrcver, have mocceaed in obtaiaing pcrmanonl fistula- witliotit an}
impnirmviit i-f llic secretion.
H<-althy |iaui-ret)tio juice is a clear, ROnewhat Tiacid fluid, fVothiog whe
shaken. It hiu a very decided alkaline reaction, end contains few or ne
structural ooustituenla.
The avenigD amount of itolids in the pancreatic juioa (of the dog) obtaiuedl
from n lemiiorarv fintuta la about 8 «> 10 per oeot.; but in oven ihorouglilr^
active juioc obtaiuwl from a permanent tUtula is not more than al>out 'i to ai
PANCREATIC JCSCZ.
3&6
per ctai., OlfltlMing inurgmnir matlor; and thi>t i« probnbly the normitl
Binount. TIm imp»rUii)l ron«litii«DU of quite frceh jiiioe are ftlbumin, a
prciilinr form of prot«)d allied to layoRin. giving Hm U» ■ sort of clottin);;. ft
Mimll nmountof lnt« nnd eaitn#,anrl ttoomparativelr large quantity of eodiiim
<»irb<>niit>?. to which ihe nlknline reRcti'>n of the juiw is due, and which SMnu
to be peculiarly associated ^vilh the pro[eid<i.
Since, as we aImiII pmenlly see, pancrefttic juice coDlnins a fernient autin^
eni-r^liiitlly on proteid matters in nn alkalioe niediiini. it rapidly di),'«atH
its own proteid conelituents, and, wlieu kept, epeedily cbau^.'eM in character.
The luyosin-likc clot is diasoU-ed. and the juice aoim contains a jteouliar
form of alkali-albumin (precipiiatable by saturation uritb inagn«tauin itui*
I'hale), as well as »niall qiianliliea of leucin, tyroaiu. and jxtptonc, which
fteem 10 1)« ihe ^rMluuIn ot Mlf-di^tion and are eiiliroly alwcnt from the
perfri'dy fresh juioe,
3 248. Anion on food-ttiijfft. On slareh pancrailic Juice acts with great
mefgy, rapidly converting it into niKar (chielly niallonc). .\ll Uiat hna
been Naid in thb respect conoeraing MUira nitght be rv»oat«d in the i»w^ of
paucrtatic juicv, except diat the aotivtty of the lattt-r ia nr grcuhT thaii that
(if Uiv foniier. Foncreatic juice and ino aqueou* inliiaioD of the gland are
n1nii» cupabtc of converting atarch into •UEar, wbcthor the animal front
which ihcy were lak<:n bo utarving or well fwT. From the juice, or, by the
glyoeriu nielhixl, from tlit- glniiii it«clf, an amylolytic rerment may be
■pproximatcly i]>oliitc«l.
On pntei'U pnncnulic juice also «xcrciM« n Bolrctit action, so far similar
to thai of )^Cric juico that by it proteids are ronwrted into peptone. If a
frw flhreds of librin are thrown into a small quantity of panrrciatic Juice,
they speedily disapi^ar, e8i>eciully at a tenipcruture of 'A't" C. niid the mix-
ture is found to contain peptone. The activity of the juice in thug converting
protetda into peptone is favored by uiOKaae of tem]>erature up Ui 40° or
tbereabouta, and hindered by low tmnperalurfe> ; it ia permanently destroyed
by builinjf. The digCitive powers of the juice in fact (te|>end. like tboee of
j;astric juice, on the preseooe of a ferment which, aa ne have already add,
nay be isolated inuvb in the same way as pepnn is isolated, and to which
the name tiyfuin bus been given.
The apji«arwnce of Rbrln undergoing pancreatic digeation is, however,
<)ifl(!!reiit from tliut untlergoing )>eptie digestion. lu the former ciuv ilie
fibrin doca not swell up, but remains as upaijue as before, and ap])«sira ta
sulfi-r cornwioii rathtr thiui wilutiim. But there is a »till uiok imfioriaut
di»(iiicti«n lictwiH-n [iniii'n-ittic Bn<l jH^ptic dig<qciun of protcido. Peptic
dwasiioa ic etevnliiilly un acid ilig<»ti<jii ; w<' have seen that the action only
lute pUce in lli<- pn^scncc of an iK'id. anc) t* armt«d bv neutral ixntinn.
Pnncr«atic digmtion. on the otitvr hand, may Ih- regarded iw an alkaline
digcstioo ; the action is most energetic when somi' alkali i* pnmnt, ami tlie
activity of an alkalioe juice i* himleml or delayed br nciiimlizntion and
arrerted by acidilication al lost with mincntl acids. The glytwrin extract
of paDci«»B 18 under all cirriimatancaa m inert in the presence of free mineml
acid as that of the stomucb in the preseooe of alkalies. If the digestive
iniKtur« be supplied with sodium carbouale t» tbu extent of 1 per cent.,
digestion proceeds rapidly, jiitl ns does a peptic mixture when acJdulatod
with hydntchlurlo acid to the extent of U.'i per cent. Sodium carbonate of
1 jier cent. Meois in fact to play in try-ptic aigeetion a jnut altogether ci^ni-
parable to that of hydrochloric acid of O.'i per cent, in gastric digestion.
And just as pejisin la rapidly destroyed by being heated to about 40^ with
• 1 per c«nt. Kilution of sodium carbonate, ao irvfisin is rapidiT deatroyed
by beiag similarly heated with dilute hydrocliloric acid of 0.2 per oenL
;jr.4
■in K Tl.'i.'rh-
oiiit<'ul>' I't' tilt- stmiiiK'li ;
M'c, sii jiri'iMinw ilif "uv t
iii'tiiiti il is 'listiij<-tly iitti!
I)ili^ ri'iiilfr^ jni^irii' jiii'i-
HilulillU fll' Ilill'->!lll-. i^^ ;i'.
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t> carried on io llie |>reseucie of aalicylio acid or thj'mol, irbiok prevent the
dffV«lupciHnt of bBAteria tind like or){&niani but permit the aclion of tha
trvnmln. no odor is peroeiveil ami nu indol h produced.
-I'uT loiiKK^ontiiiiitid digMiitii). especially when accompanied by putre-
' " ')i.-4^>impoHiiicii), ihe amoutit of pn>tei<u which are oarried b«y«iii<l tho
I '^ge and bri>l«ii up may be very Krvat.
:no digmUon such a profnuDd di-»inicti»n of protcid material ocoiint
Lii a inDcli tft«i extt'nt »r not at all ; noilhor Inudii nor lynwii can Hi pnxent
b** ninttiilcryxl nn nuttiral product.-' of tlic acti'Xi <>f pepsin.
On tht^ eelntiDiferoiin ulemenU of the liwun u* lh«y actually t'xiitC in the
tiiviK pr«'rioii8 to aiiT irwiinieni, pnncmttic juice nppoira to have no «iilvonl
aciroii. The fibrillir am! biitiillcs of librillR of ordiiiarj- nnlouchcil con-
niirtivo tiMiio arc not digntlcd by pancreatic juico, vrhich in this respect
airor<li> n ulriklng rontraMt to gn«tric juic«. But when lh«y hare been pre-
vio«i»ly trait«'l with acid or boiled, so w> to become canvcrt«d into actual
eelatin. try|win i« able to dissolve thero, apparently chauf^og them much in
Ine Mirae way as does pepsin. Trypsin, unlike pepsin, nil! dissolve mucin.
Like pepsin, it is inert toward niicleiii, horny tiasuce, and the sn^wlled
amyloid malCer.
) >n j'tlt pancreatic juice has a twofold action. In the Grsl place it emul-
aiti«« fats. If ho^'s lard be gently heated until it melts and be then mixed
with |>ancrealic juice before it solidiliea on cootiDK, a creamy emulsion taating
for almoat an indefinite time is formed. 80 also when olive oil is shaken up
with jiaucreatic juice, (be separation of the two tluids takat place very slowly,
and a drop of the mixture under the microacope allows that the division of
the fat is verr minute. Au alkalinti aqueous infusion of tho gland hiwi
similar emulaitying jiowcm. lu thf w^iond place pancreatic juice splits up
iMUtral fata Into iht^ir n-«[H-cMivi- nciiln and Klyceri"- Thus, f>almitin (or
tn|ialmitin)(C,H„.('0.0),.C,H,iawith the aMumption of :{H,0 nplit up into
thru! RiolecuU* of palmitic acJd H (CnH^.CO.OH) and one of iilyovrin
(C,H,) iOH,l ; anil wt with tho other neutral fill*. If |K'rfoctly iti-iitral t'al
be trattwl with pancroutic juice, cvpcfially at (be bodv-tvm|M;rature, tho
emulsion which i« formed Kpcnlity (akei* on au add reaction, mid by sppro-
pfiate means not only the corresponding fatty acid», but glycerin may bo
ob(uited from the mixture When alkan is present, the fatty acids thua set
free form their corresponding soaps. Paiicrmtic juice contains fat« and is
conser^uently apt after collwtion (o have lis alkuliDily reduced, nod an
atiieous infusion of a pancreatic gland 1 which always contains a consid-
erable amount of fat) very speedily becomes acid.
Thus iwncreAtio juice is remarkable for the power it poesessee of ac4iDg 00
all tlie fuod-etuflfc, on starch, fats, and proteids.
The action on March, the action ou proteids, and the splitting up of neutral
fats appear to be due to tl>e presence of tliree disliuct lerments, and nwihode
have wea auggeeted for isolaliog ihein. The eniulaifVing power, on the other
hand, is counec[ed with tbegeoeral oompoeitJonof the juice (orof the ac)ueou»
inl\i«ion of the gland), being probably in large measure dependent on the
alkali and the alkali-albumin present. The protemlyiic ferment, trypnii, aa
onlinarily prepared iMiina to be proteid in nature an<) capable of giving riaa
by digt«ti<Mi to pept'>iie ; but it may he doubtetl, as in the uaso of porwin and
other fi'rnMtntx, whcttxT the purv (ormetit haa yet 1)een iwilattd. Then- an:
no mMuiH of dii>tiiigiiiHhing the amylolytic ferm«nt of the pitncrenn frifin
(ityalin. Tlw term paHirrrittin hiia lieeii variouslv ep|)lied to iiimiy ditlemnt
pnparationN from tho gland, and ita use had, p«rna[w, I>ett4:r Ih- avoided.
The action of pancniatio juice or of the inluiion or uxiruct nf the inland,
on starch, is seen under all vircumataiiem. whctlier tho animal Ik- fa«ting or
<
us
THE TIS81
iSU MKCHAS
gbiid WM eonliued to tlie Utsk of letting certuiu coiutiiuenu of ilif hluud
pan from the ciipilturieB liurruuiidiug the iilreolus tliri>U|{li the c«lls to ih«
channel* of ihe ducts, while refusing a paisui^ to iitht^ra. We uomr know
thkt ceri&in ini))ortunt iiuialitucnld of each jnu-c, thi- peiMin uf KUtric juice,
Ow mucin of salivn iiml the like nre r<iniierl in the cell, uiil not ohuined
Tandy made from llie hlood. A minute (lunniiiy of )ie[>«i» does exUt it b
true in the hlond, hut there iiiv reaunn* fur thiukintc thiil this bu tnnde it«
Wny back iiilu the hlom], viiher heiiig a1)sorbiril from the intvrior of tie
etoinnoh or, an teem* more prohuhlc. picked up directly from the (tastno
gluids; and ito with some nf ihc iithi-r I'oustitiientK of other juices. The
chief or Apevilic consljtuent* of <-nt.'h juice nra fomieil in the oell itself.
Btit the juice vccrv-lcil hv any gland conKi»l« not only of lhe»pecilio tub-
Mooee* audi ■« mndn, [>cp»in. or other ferment, or other boiIi««, fouod in it
■lone, but aha of n largi^ iiunntity of miter, and of variou* other tab-
stances, diietly »iilint'», common to it, to other juic« and to the blood. Aim)
the <jue^iim ari*iT», Is iho water, nre tltc Milt* and other comiiion BubitKDce*
furnished bv llie wime att at that vihidi supplies the spmlic conttiluenta T
Certain faafM »ug)nMt that they are not. For ioiiancc. a« mentioned wirat
lime &gi), in the submaxillnn- gland of the doe, etiraiilatian of the choirla
lympuni produce* n copious How of saliva, nbicn is iisunlly thin and litaptd,
while ntimulatiou of the cervical synipnthelic produoes a scanty flow of thick
viscid wliva. That is to say, stimulation of the chorda has n marked effitot
in promoting the dischari^e of nater, while stimulation of the syiu{iathatic
kaa a ninrkcd efleet in promoting the di»charf^ of mueio. To this ire may
add the case of the parotid of the dug. In this gland aiimulation of a
cerebrospinal nerve, the au He ulo -temporal, prodiieee a copious flow of limpid
saliva, while stimulation of the ^ruiiutlbolic producM itaell' little or no secre-
tion at all; but when the aympatlietic and the cerebro-tninal nerves are
MtimulaCed at the name time, the nuliva whieh flowi iti much ridier in aoliil
and eepcciatly in organic matter than when the cerebro-Mpinal nerve i* stimu-
lated alone. And vt- have atreadr -leeti thai in thtx gland the niicroeoopic
chauj^ fiilliining upon nympitlhetie i<timulntion are more von.-<picuous than
(hose wliich follow uimn cercbronpiiial ttiiaulation.
The«e and oilier facl^ have led to the conception that the act of secrettoii
conugta of two purt«. uhich in ono ease may coincide, in atnnher msy take
place apart or in diHercnl proportions. On the one hand, there is the dis-
charge of water carrying' with it eoninion solubkt snhetAnu«Si chiefly «idinee,
derived from the blood ; on the other liimd. a metabolic activity of the (vll-
subitauce gives rise to the specific constituents of the juice. To put the
matter broikdiy, the latter pioccs produces the specific eoiutitueuts, the
former washes these and other matters into the duct. It baa been further
auppofcd that two kinds of nerve fibres exist: one governing (he former
ptocm and, in the cose of the aubmasillary gland fur instance, prepun-
aenting, though nut to the total exclusion of the other kind, in the chorda
tymjuaiii ; the other governing the latter pn>oea9 and preponderating in the
branches of the cervical sympathetic, liicse have been called reapeclivelr
" secretory " and "trophic" flhre^: hut these terms are not desirable. It
Biajr be here remarked that even the furnier proceai is a distinct activity uf
the gland, and not a mem infiltration. For. iw we have seen in the cane of
the salivary jflamU, when tit.ropiue it given, not irnly do the npuuJGe cousdlu-
enis ceose to hi- ■•jecleil lu a conM-iiucnce uf utimulntion of the chonl.i, hut
the diiicliarge nf iraler, in spite uf the hliiudvivnelx liecumin;; dilnt^il, i» also
iu-r«>ted : no siiliva at all leaves the gland. And what is tnie of the salivary
viands as regnnls the dc|K'ndene4- of the flow of water on something eMi
beside* i1h! mere pivMiiTe> of the bhiml in the hloodviMseU, npixtar* to hold
iijiseslisl
wuier n ilup lo nn iictivily of ihc cell : llif hviMttI
pot forwaH iliai rliiinKn iu ilie cell give ri»e lo (lie ft-ntiatioD in Ui« cell of
■qfalUmwa which alworb nnler from ilie blood vr Irniiih on the one >id«
«ed fpvt it up on ib« other aide tuto ilie lumen of ilie itlveolua. Bucb an
bjpolbeua csnout be regarded as tirave<) : but ihe mere putting it fnrward
T«bn doubt* as to iliv validity of tlie diatiuction on which we have been
■)««llta8; and niher cviuidenitioiis point in the Mme direction. For in-
•uiKV, if Ihe ntnimon aoluble mUh pievent in a Juice, as diatinguishcd lW>m
tbe •(■ecilic «iii*>iiiueii(», were merely cnrried into the juice by the rush, m>
u> tfrak. nf water, we •huuld ospe<-[ lo timl the |ierc-enin^'e of tbei»e aolta
eilhrr rrmaiDiiig the Mtntc or |M:rhap« decre&Klng when Ihe juice wila cccreted
atun mpiilly and in fnlkr volume. Bui under these circuniMtnuci'" the i)er-
rmtap very frcjucnlly )n<'re>i9n>: luid In general we find thiil under vnnou!!
drruBiftaan* tbo pro]i»rtiun of wits tecreieil tn llitt rpinntity of wnter
•rmtnl may vary «iu»*ieT»bly. Obviously, whik- something delcrmines
tb« quantity of vnter pacing inio ihe uIvcoKib, soniething rise livtfrminrs
boo much of coniRKin MiltibU- wiIIa Ihnl wuIit ciiiilaiiii>, snd i>till KinielJiing
aJaa rietcnninvs to Khst extent Ihnt wnicr i;i rdso hulen with ■pccific conifiiiu-
rat* and oth«r organic bodiro. The whole iiclion is loo compliciited to be
dracrilird as coiwisttng merely of the two procesMV mentiimcl aliove, but the
liaM has not r«t come lor clear and definite statemenlf^. Everything, boir-
snrer, teii(b> to ebow that the cell is lb« prime agent in the whole business, _
though we cannot at prM«nt ddioc the natnro of th« H-vpral changes in tbo ■
cdl, iNr can we sav how thuse changvo arc exactly related to each other, to "
rhangin of ibe bfood-pmEHre in the bloodvessels, or, we mar odd, lo
thuugm taking place in the Irmph-spaces which lie between thetilood and ■
the «elL I
We may perhaps add that, uuce ia certain cutaneous secretiof; (''■'■tils the
alreolus, or what ci>rr«t>|H>nt}H to tlie alveolus, is nramied round with plain
musniUr fibres, the contraction of which upt>ear9 to force the secreiinti out-
ward, the idea bas ticen miggeetetl that in gluuds, such as we are now voti-
ddning, the ctJl-subsianee niakitig ii>e of "protoplasmic " coDimctiou tn-
M«ad of aetna) muscular ci>iilraciion, may force part of the c«II contents
into the lumf'n nf the alvooluH. Such » mode uf seorctloD would be I'om-
Mrable tx the ejirtionof undiKCvtcd malcrial, or "excretion," by an nni«ba.
ilut we hove no MitUl'nclori' vvtdrnce in favor of ihi* view.
I WO. Thnmghoiii the above wc hnvc spoken as if the secretkm were fui^
nislted McluHVely by the cells of the alveoli or stKircting portion of the
C' D«l, Bsiftbo cpiihcliura cells lining ihe ducts, or conducting portion of
flaad conlribdtpd nothing lo the act. In the gastric ghinds the slender
cells Itniog t)>e moulhs of the glands (which comspond to ducts) and cover-
iw ibe riages between, are mucous cells secreling into the stomach gener-
ally a Btnsll, but under abnormal conditions a large amount, of mucus.
Nbicb has its uses but b not an essential part of the gadric juicu. In Ihe
salivarT {.dandit we ean hardly suppose that the lone stretch of character-
istic ruluinnar ejiithetium which reaches from ibc nU~eoIi to the mouth of
tiM ioag main duct wrves simply to furnish a smooth lining to the conduct-
iag I WSj^i s . bnt we have as yet no clear iodication« of what the function
of ibb epitbellum e»a be.
I ML Before we leave the niechanisin of secretion there arc one or more
acvaaory noinia which deserve atleiition.
In tniitiiig ju*t DOW of (he gastric elands we spoke as if pepsin were the
only ImporloDt couliluent of ^iric juice, wbenaa, as wc have previously
, the acid is e(|uat[y casenlial. The formaltoa of the free add of the
I
I
1
I
85(r
THE TI»St'EA A.VI> MECHANISMS OK l>IOKSTtON,
KMiric jiiico is vtry olMcurc, nud maiir ingenious but tiRMtufKclorir v'ltvti
n*v« liM-ti [xit f'lrwiiril ti> cxptitiii it. Ii tvum* uutiirul f > •ujiihml- ilint it
siriscc ill M^ino nuT frum the liecompOBiliou of fiJiiim chloride dntwn fn^ia
ibi.- IiIoihI : tind this is 8U|>purt«d by the fact thsit whcti the eecr«livn •*)' jn*-
tric jiiico is iictivoly going on, the nmotiiU nf cliiondm lenviog llie l>loud
by ttw; kidney it proporiioniklcly dimiiiLghod ; hut nothing de6nit« cui nt
ptTOHUt lie elated as to ihe ntechnnisiii oJ' ihnt deeoni position. And even
ndlDiKin^ thiil llie sodium chloride of the body at large is the utlinistf
anuixMi of ihe chlorine element of the ncid, it appears wore likely that thai
elemenl &liould be set free in the stomueU by the deoom position of aonie
highly coiu|>les und unstable ehloriiie compound previously eeoerated. Ibon
that it ahould arise by tlie direct tiplitiiu^'-up of eu stable a body a^ Miliun
chloride at the very time when the ncld id secreted.
lu the iriiii, while ])e|isiii true from acid in secn-te^l by the glaud* in the
lower iMtriiun of the (t.^>[ihii);uii. an acid juice it utTorded by ijlaudn in the
ittouacli itself, which have accord iu;i:ly been culled ujrifiifi'r (iiit-nr lo *hur]>co,
acidulate) glaudn; (nit thrau oxyntic ghinthi Hp^war nlgo to wcMte [K-jKan.
In the inanimal the Uolnliil pylorus secretes an nikniine juice; in tnct, lb*
nppi-araiiee (>f un acid juice is limited lu tho^e ]>oniuii» of the itomMih in
nhich the gluuds oonluin boUi " chief" or " centrni," nud " ovuid " or " bi)r>
der" cell*. Now. from what has been previously wiil. there caii be do
doubt thiit the chiuf cells do wri-rcte i>w|i«iii. On ifie other hand lhen> is no
flvideoce wliatevir of the formation of i>e|»in by the '- border " or "ovoid "
cells, though thi» wns once supposed lo no the cose, and these oelU nere un-
fortunately formerly culled " peptic " cells. Hriice it hns been inferred that
the border cells secrete neid ; but the argument is at present one of exclu-
sion only, there being no direct proof that these cells actually niauulacture
the acid.
The reuuiu apiieais to he fonuetl by the same cells ivhtch maiiuradnrc the
pejwn, that is. by the chief celU of the fundus geiiertUly, and to totuv os-
tent by the cellti of the pyloric ^lauds. We may add that we have cvidencv
of the cxistenvc of a xjidc^u of renniu analogouti to the cyniogen of pepsio
or of trypsin.
The mueu» which it prevent ns a thlii layer over the surface of the fasting-
(toiuach, aud which miH^'lally in hcrhivciriiu:t animals, is iocreoMid during
digwtioti, comes. iu> vr hare «aid, fmui the mneous cells which line the
mo(ith:< of the si'vcrni Kb'udi and cover the iuterveniug surfaces.
$ S43. We pr«vioii»fy cnlleil iitlenliun t" the fact that in the case of the
stoinnch the al»iiqitioii of the )mNtu[.'t.-> of digestion largely iucnasod the
activity of the secreting cells. This lins led to the idea tliat one efleci of
food is to "charge" thegootric evils with |>e|isin[igon, and that certain article
■>r fend might In- considered as es]>cciidly pcptosenoite, i.e., conducive to the
formation of pcjisin. 8uch a view is tempting, out oeedfl u yet to be mom
fully snpjiorted by facts.
S 843i. Sc«in;; the great solvent |>om>r of both gastric and pancreatic juini.
the queetion h ualurally suggctited. Why dom not the stomach digest itiKilfy
After death, the stomach is freipiently found partly digested, viz.. in cases
wben death has taken place suddenly on g full stomach. In au ordinnrr
death, the membrane ceases to socreie before the circulaiioa U at an end.
Timt there U no inncciul virtue in living ihiaKs uliich prevenia their being
diMsted ix shown uy the laet that the leg of a living frog or the ear of
n living nihhit introduced into the stomach of a di*;;. thr^mzh a gastric
tbtula, is rvadilr di){usted. It has been »U{U!eate<l that the blc»>d-<-urreut
keep* up an alkalinity »uffictent lo iieulndiAe the acidity of the juice
in the region of the glnuiU themwlv<i' ; but thid will not explain why tlw j
A
BILE.
691
fOcmilc jnico, which !* netivr In nn nlkiiline mcclliim, don unt dt|fc»t iho
f«Mnli uf tho (Kiucrm* ilH'If, or why ihc diiitw'.ivt' cwll* of lhi> IiIuimIIi-m
MfiDMOon or hy<lni»<oii il<> iiol iligwl ihfiii--"'lv<«. \Vv mi^hi mlil, it diwn
Mt ripiMn «hy lli<- iinid'hii. whik <liiwilviiie iho ]>r»tc>iilH*iii or th<.- Kwnl-
l-tnl ilintDin. Jx* mil ilinwiK-i- il» oiru ]>n>t»{>lsKin. Wt <-itnih>: nii«w<T ihi"
yxttiiM at nil aL ]>ri-*<-iil. niiv nuitv than tbi.- piiniiliir oiir, mhy the ilclicitlo
fflAaplaamnribv niiKiOin rifUUiluriiit; lilb thortilnirK^^ iiili> ilaclf l>y immiwit
Umm WBtvr than it rit|iiirM lu nrry on ile work, whik a iVu luomi-iile
idtr It bdeiiil water vnlvn freciv by oNinxiK. bikI thcolTccUot'ihitt ruiraiico
taCMM abuiutatiflv vvidcut by lli« fomutiou iif biiltn nitd tliv brankitig ii)t
0t iW |iru<u[iUsnL
TbK PaOFRKTia AXO Cba RAtTKKg »»' UlLK, PaXL-RCATR' JUICS A9IO
I M4. Id the liviiij; btHly llie food. Hibjectei) to the Bction rirat of ibe
mJIvh niul ih«u nf tlie ^iric juic«. undergoes ia lh« Muiiiach ehanK«d which
•« •hall [m»etitly onDSKlcr in deinil. and th« food so rlinngcd » paeaed no
into ibv wndl ini«»tlDp, w)i«rv it in further iubjc<eteil lo Ibe action of the hi)«
MctMod by tht liver, of |i«ni;n.-atti- juic« i^«crcti.-d by the ]>auereB!i, nml |iib-
aibly b> •otne «stenL, tl^ouuh thi* by ao nieoiia cvrtaiu, nf n juin^ wcreuil by
tb» laicatiTH' ilicllt mtd mlh'd mccu* enUrirut. It will be c<>iiveiii«iit to (ttidy
tb« Miliut*.- utriK-inrc of ihr livor in ixiniie<-ti»ii with oth«r functloD* of the
lir«r omrv imporlanl. {irrhii|u, ihini that nf ih« Mcrction of bile, rinmely.
tb* femuUiau of Klyoi'e*^". i*"d itlhiT nu-tabnlic «vait« oecurriiig in the
b^Mrtir C«ll«: «« nave iilrvitdy •(u<li<r<l the vtnicturD uf the {intKTCiw; and
llwMrtlcUiraof th« inlwiinr will \w*t bi^ 4-unridetvd by ii#i-)<. We, ther(»
Lfbn, turn al once to the pn>[>cftivs aitd chnrnciere nf the nltowiinmcd jiii«ci.
I
I
hilt.
,Tbuu^fa bile, after Mcretioti in ihe Ivbulea vt the Ii\'«r, in inuved od »loag
IwfMtic duel. It ik In the cnnc i>f iu<i»t aiitniiiU tiirt poured at uncc into
lb* (Itwlenuiu but liikcn by ihe cv»tie duel to the ruwi^'oir uf tliv gnll-
bbwider. Ilrre it rt-inaiiiK. until tiicli lime lu il ii i>ecd«d, irhen a ({liKntiLy
M I ' irig ihi' r>iniriii><i bile duet Into tlie inusiine.
1 . ' iiy i)f bib- vnrii? tiiiich. ivii only In diHt-ronl iiniiimls, but In %ht
Mmc animal nl diDl'nnl time*. It i- ni>it»i)vi-r alferled by ihe tcn|>th of the
Mij-tum in the gnllbtii'Mer: bik liikvn direct fotni tin- hepittic duet. n(w>
rially wliro KCi-retrd rapidly, omlitiii* little or no miicun: ihnt inl>t>D IVom
ihe ^nll Madder, na of »Iaughtere<l oxen ur ahe«p, i« loaded irilh mucus.
Tlie eolof nf the bile of camivoroiii' nod oRiuivorous noiniak. and of nuiii,
U jrrnorally n bright gidden rvl \ of herbivorous animals, a yellowi-ih grc«n,
or a brii:bl griTii, or a diriy green, accortling tn eircnmslancc*, beiiij; inueli
nxidifird by rclenliun in the gall bla<Id«r. The rmi-linii in nouirdi or alkaliur.
Tbe full(iirii)i[ itinv Im taken as llie average coiii|u)«iliijn of buniaii bile takflO
C! ^it-blailder. and thefire c.iniaiiiing much more iniiciu m well ta,
y to iIh- N.dids, in<in> water ihaii bile frixn ihe Wpaiii.' duel.
)u vm lanr.
*r . ... «.W,2
Ida:
llc^lt. ... m.-i
U^ftc . . . !>.J
bidMlMln . . .2.6
ueiit and Piicmiiit SIM
•nrpulc Soil* 7J*
IMS
852 TBK TISSVE9 AND MKCaANI3HS Of OIOKSTIOK.
The ODLire absence of proteida is a inarkei] feature of biie; {WDirreatie
juice, as we eball see. cimttLins a conuderable tjuantity, aaliva, ii» we liave
seen, n eniol) quaiitity, normal ftniitric jtiit-e probablir still Ivm mm) bile unoe
Kt ail. Kveii tbe bile wlilcb bas been retaiiicsl «onie liiii« tn tho gall-blnd<kr,
thoiif^b ncli in mucus, ctniiaiu-t uu [>riitri(l».
The ciiuetitueuu wliicb toriu, ttjmrt iVom tbe tmicuM, tbc fircntbulk of ibc
Di>li<l» of bile, iiud which ileitcrve I'liicf utlcntioii, are iht pigmenlf ukI tli«
liilc'Kulldl of ihue we tiball *)irnk i mm<'i]inlv]3'.
Wilb regard lo tbe liiiir^iiic mill* imuully prwwiil ii# nuch wnlium »nlt»
are confpicuoiis, aotlium chlfiridi? anitiiintiii(; to V.'i or more pt-r cent., foilium
pbocpbale to iienrly nw nuicb. (lie nt\ being cnrlhy ph<wphalti( ami other
mnlters in small i|oiiiility. The iir(»ciicc of iron, to llie extent of about
0.006 jwr cent,, lis intcn^tinf, tiiia.; n» wc shall Bee, there arc reiuons for
thinking that (he nignient of bile, itself free from iron, is derived from iron-
holding hirniDKlobin ; snnie, at least, of tbe iron set free during the cotiver-
sion of btemoglobin into bile pigment, which probably takes place ia the
liver, tinds its way into tlie bile. Bile also nppean to contain s small ifuaii-
tily, at all cveniB occasionally, of other metals, such as manganese and cop-
per ; melaU introduced into the body are apt to be retained in the liver aud
eventually leave it by the bile,
Tbe small quantity of fat present consists in part of tbe complex body
lecithin.
The peculiar body eltoU^erin, which tbou|{h fatly looking (beoce tlte name
" bile tat ") is really an ak-ohol with the com|K)(iiii»u C^H^.O, in coiupicuoas
by ila ijuautity and i.-ouslancv. It furmd the greater tmrt of moiit gall'MoDca,
thou^'h »ome are comiMiited i-iiietly of pigment. luiMilubk iu water and col<l
alcohol, though soluble in hot alcohol and raudily woluble in ether, chloro-
funn, etc.. it la disaolved by the bile-aalts in aqueout tolutiim and bcnre is
p^c«^nt In Milution in bile. Us |>hysiolo|ricnl functions arc olucure.
The n»h of bib' ci>rigials largely of uraa, derived partly frum the H>dium
chloride and |>nrtly from the bile-salts, of sulphates derived chiefly if not
wholly from the latter, and of phospbatoi partly ready formed, and in pan
derived from tbe lecithin.
§246. Pigmenlt of bile. The natural gulden-ret) color of normal human
or carnivorous bile is dne to the presence of bilirubin. This, which i» also
tbe chief pigmentary ranstltuent of gall-stonei:, and occurs largely in the
urine of jaundice, may be obtained in the form either of an orange-colored
amorphous powder, or of well-formed rhombic tablets and urisnis. In-
soluble in water, and but little soluble in ether and alcohol, it is readily
aoluble in chloroform imd in alkaline fluids. Its compo&itian ia C„H„Nj(V
Treated with oxidizing agents, *uch as nitric acid yellow with nitrouit acid.
it displays a siicceteion of colors iu the ordor of the spectrum. The velluniih
golden-red becomes green, ibia a i;reenisli-hlue, then blue, next violet, after-
ward a dirty red. and finally a pule yellow. This chnracteriBlic reaction of
liilirubia ia tbe basis of tbe so-called Onielin'!< tett for bile-ingnientH. Each
of ib(» Biases repreeenla a distiuut iiigmeutarv >ub«tance. An alkaline
BulutioD uf bilirubin, expnaed in a shallow Tinsel to the action of the air,
luriiB green, becoroiufc converted taio bilivmiin (<'„H„N,0, or C,(H„N Oy
3laly), the green pigment nf lii.'rbivor»u» bile. Bilivenlin in aldi found at
timts in tbe urine of jaundice, and in probably the body which givtt lo bile
wbich hiu been cxpwcd to tbc aolinn of goxtric juici;, ii( in bilmrv vomits,
itM chiiracteriittic green hue. It is the lirAt atagi* of the oxidation of bilirubin
in GnHtlin's Icst. Treated with oxiditing agents hiliverdio runs tbrmigh
the Mime seri» of colors nn bilirubin, with
golden red.
the exception nf tbc Initial
BILE. 858
I 9M. TV bite-m»lts. Thow coimst. ia mnn tad nifttiy nniniRl*. ottodium
fhftatMaU uid loutvfhotaU, the proportion of tho tiro vamni; in dilfcteiit
■aimmk. In ouin bolh Lbc total i{iiBnlity of bik-ealta and the proportion
■if iIm utw bile-skU torhe otWr swm to vmrv n ^'oi^d deal, but the glyoochoUto
kftMtid to b« always the niorv nbundant. In ox-gall eodium glyeocholate ia
■fawidwDl. and Mtinicholalc HcaDty. Th« bil«BalL» of the dog. cat, bear, and
athar evuivora nxtMst exchi»ivolf of the latter.
Iimlubte in ether, but soluble in alcohol and in water, the aijueoite sola-
OM havJDfc a dei-iOed alkaline retiotiiin, both tuilta may be obtained by
crrrtallixatiun in fine acitular needles. They are eieee<liii(;ly dcliijiieacent.
T&c -wlutioui of Ixith aci<U have a dextro-rotary action on polarized li);ht.
,- '»M. Bil«, mixed with animal charcoal, ia evaporated lo drynew and
•xtnuTird with alcohol. If ogt coloriest, the alcoholic nitrate must be fiirlber
(l«<Ri((iriccl with soiinii) charooal, and tho alcohol •littillFd off. Thr dry r^Miduc
it trcaud with absolute ali-obol. and to the alcoholic lillratc aiihydroui olhiT U
added •• 10D( aa aor precipitate in formed. On vtaudiug tlie doudy precipitate
haODBiei tiannibrmed into a erTMalline tnaa* at the bottom of the vea»el. If the
akohol be not alMilute, ibe cr^Hala ore very apt to bo rbanK4wl inui a thick
■yrvpy fluid. Thi* mmvi of erTntab hw b<^n ollon apoken of nx Win. lioth «alta
ar« thu* pKcipiiatod. wi Uiat in surli a bile lu thai of the ox or man biliu oon-
»bto both of B'^Iiuui glri-ucholitie and soJiuiu taurocbolatc. The two may he
•rjiarated by preci pitta Ion from their aqiiMiua solution* with vnicnr i>( lend, wtiich
larows down tho former much more readily than the lultet. The acids mny be
■ayakratsd from tlieir mwclivp salta by dilute aulphuric acid, or by Uie action of
md aeelaie oad aulphydric acid.
Oo bofllng with dilute acids (aulphuric. hydrochloric), or caustic j>otneh
nr baryta water, glvoichidic acid in »plit u|) into cholalic (cholic) aoid and
glydn. Taurochofic acid may similarly be split up into ch«Ulic add and
uuria. Thua :
I
K
(HreoctwHc uU. Ctaobllc aetd uljoln.
f,HaNO,+ 11,(1 = C„H„0. - CH,.Na, (OO.OH).
TMunehoUe urtd. lliulallc add. IViulii,
Ci,H^SO, f 11,0 = C„Il«U, -i C,H..NH,.S<I,!I.
Berth aoda contain the aame non-nitrogenona oeid. cholalic acid ; but this
acid ie in the lirai caae WMocialed or conju){ated with the importnut uitro-
^enoua body glycin, or amidoacetic acid, which U a compound forme<I from
anHDonia and one of the " fatly acid " scriee, vii.. acetic ; and in tlie »ccoiul
can with taurin. or amido'i<elhionic acid, that is a coranound into nhicb
wnfMenlativcB of ammonia, of the ethyl ([■^'■I'- ^o^l ^^ aulphuric acid mter.
The dev(»mpueilioa of the bile acidi into cholalic acid and taurin or glycin
napectirely takea place naturally in tne inteadue, the glycin and Iiiurin
b«iiK priibably alMorbed, <» that from the tno acid*, afUir they liavo wrved
llwir purihtw in dinuion, the two ammonia oiimjiouu<b are rvIurniM) into the
MoimI. Kacb of the two acid*, or cbolalic aeid nhme, w1m-xi treated with
*ul|>huricaddaiid mne-dugar, );iveHaraa)(iiihi;eut purpk color (Pctteakuler*
Ual), with a cluiraoterutic Bpectruoi. A aimilar oolor may, however, ntlen
W pruducet) by the actiou ot ibe aame bodioa on albumin, amyl alcohol, and
ium* other oi^anic budiet.
$ M7. AttwH of bift on food. Id aom« aitimak at leatt bile cootaim a
fnvH-nt ca I akble of Converting atarch into mgar; but its action in this reepect
it wlmlly MilinrdtDAtc.
On proteiiU bile hu no direct digwtiftt action whatever, but being, |;eo-
fntUy at leaat, alkaline, and often atroogl; so, tonda to iieutraliM the acid
2i
THK TIS9UK8 AND MKCUAKI8M9 Of U1GB8TI0N.
oonteuiii of the stomach km tbcy jMtt aHo the duodenum, and, as we attall
Me, w> jirvjiarctt Uit' vikv for the nctioD of the pnncreulic juioe. To peptic
action >t i» dUtiiiellv Buiti^Hiiittiv; the prcaeuce of n suflident i|uaDtiir of
bile rendera Ki><>tHc juice inert toward proteida. Moreover. wh«n bile, or a
BolitdoD of bili-'MiltH, is niltkd to a fluid oootaiDtag the products of gaeuic
dig«etioD, a pR'cipiiute tiikes place. eoDMitin^ of [Mtrapeptoiie (w1i«d |ir«sefil),
peptone, |iep«in. iitid liile-ults. The precipitate ia redueolved iu au ezoeH
of bile or sohiiinn of bilc^sfilt«: bill the pepsin, though rediaaulved. reauiD*
inert lownrd proteids. This precipitation actually do«B take pluc« It) the
duDilenum, mid we »hall »peak of it again later on.
Willi regitrd to the action of bile oo fats, the followinj; stmXemeata mar
tw Riadi? :
Bile haa a slight solvent a<.-tiou on fats, as seen in itii uat by painter*. It
ba« by itself a elJjilii but only aliiihl emulsifying iwwcr: a mixture of oil
and hile separate after shaking FBtner leaa rupidly tnan a mixture of oil aiid
watrr. With fatly acida bile forms M^a])s. It b, more<;ver, n mlvoot of solid
euHpi>. and it would appear that the emulsion of fal« if under ccrtAio cireum-
Blancc* at ttll events facilitated by the proienoe of MWfw in solution. Henoe
bile i» pnilmbly ut much grenter line an an eiuiiUiDn aernt when mixed witb
panerentie Jiiii» than when aclinj; by ilnclf alone. Ti> this poitil we $hall
return, Lnntly, the pastui^ of Ints thnni^h niemlimncs I;" ii!«ist«<) by weltini;
the menibrani')' witli bile nr wirh a solution of bilcsnlls. Oil will poas to a
certain extent through a tilter-pAper kept wet with a iohitioD of bile-Mils,
whereas it will not pa.-« or naaeea with extreme difficulty through one kept
constantly wet with distilled water.
Bile pnmrmni some aJitiiieptie qualitiea. Out of the body its preaeuce
hindea vaHous putrefactive proceaK* ; and when it is preveut«it IVoin llaw>
ins into the alimpntarr canal, the content? of the inteatine undergo changra
diflbrcnl IVom those which take place under normal condilioiiK, and leading
to the ap|>earanoe of various products. ei[>e(-ially of ill-timelliDg euee.
These lariuus actions of bile seem to oe dependent on the bn»«alle and
not on the pigineiitary or other constituenis.
Pancrralic Jutw.
«
§ 248. Natural healthy pancreatic juice obtained by means of a temporary
paocruitic fiittula differs from the digestive juice* of which we bate already
moken in the ooinjmratively large i|uantity of proteids which it cuntaina.
Iu conipocition vanes according to ihe rate of secretion, for, with tlie mora
rapid flow, the increajte of total solids dues not keep pace nilh that of the
water, though tin- uvh remaiuit remarkably conslaul.
By an incision through tlic liiiea alba the paDcreatic duet (or ducts) uan easily
be lound ailher in the rabbit or in the dog. and a caoula secured la It. There b
no difficulty about a lemjioRiry (Ixiula: but with ponnanont fisiulie Ihe secrelion
b apt to lieeome altered in nnVure. and to Imc many of its characiflriatic proper-
llCH. gome, however, have mccredcd in oblaiiilng permaaent fiatulie xritbout anjr
Impairmeni «r (he tweretiou.
Healthy pancreatic juloe is a clear, somewhat viscid fluid, frothing when
•baken. It has a very decided alkaline reaclioii, and coniainN few or no
ftrudural ooustituenls.
The average amount of solidi^ in ib<- piiucrealie jiiicc (of the dog) obtained
i'rotd a tan)]M»an- listula is nboiil H to 10 jkt cent. ; but in even tJiorougbljr
active juice obtained froin a permanent fiMula if not mors than about 2Ut a
no juicK.
886
*
\
per «HiU, 0,8 lK>iiiK iitor^nic mulicr: nnd th» i< probablr the iionnal
■moant. Tbe iniiMtrittDt conviiiudile of quii« Imh juice are albuoiia, a
ptealbr fonn of nrotegd allied to mirosin. giving rise lo a son of clouinj;. a
MMilI atnoaDtornta andsoape, andacompnratirely Inrge quaDtil}* oraadium
oarbocMte, lo whidi ibe alkaline reMtion of tbp juiiie is due, and which seenu
In b« pecaliarl}' aaaodatcd with the proteids.
Sace, a» we shall piewDtlr see, pancreatii.' juice eontaiofi a ferment acting
*ser|{clicallv oa )>rt>ie»d luatten in an alkaline medium, it raf>iiily dig^la
H» own proWid miwliluenu, attd. wbi-n kei>l. speedily chauKea in cliarai.-ter.
Tbc mjraain-likf Hot b duaulved, and the juice soon couiains a peculiar
Hirm fif alkali-albumin ({irectpiiuiablt- bj- HUiuration with majKneaium nul-
pb*lv). a* K'idl R» omnll qiiiintilie* nf leucio. tynMiu. and peplune. vrbich
ttvio Ut b* tlif nnxluct* uf neir-di^'Mtion aud ui« entirely atment from tbe
perftvUr liMb juice,
f MS. Aelion ON faod-Mu^. Oo «IarcA ]mn4-rentii- uiioe acU with gmU
rapidly oonTertiDg il into NUgar <chielly malloae). All Ihat bu
in ihi* nepect ciini^rniii^ Milivii might Ixi repealed in ibe 4!iue of
pUtcrfWie j»i<«, execj)! that tlie iicttvitv of tbcHattrr in liir gimilrr than that
td iIn feraM>r. i'nncreBlic jnicr and iW aijitepuK infunion nf tho gland are
•twars capable of mnverUng starch intn Diigiir, whether the animal fmm
whidi Ihey were taken be iianHne or wcilt l«<T. From tbr jtiioe, or, by tbt
gljnariii method, from the gland ilaelf, an amylolytic fi-rmeat may be
•pMWlimat«ly isobited.
On f/rot^iiU pancreatic juice also exercises a Nilveiil aclion. so far similar
Ut that of {[astric juice ihM by it proteids arc I'ODVerted into peptouc. If a
fr^ Anib o( fibrin are thrown into a small iiuantity of pancreatic juice,
tbey apeedilr diaiippc«r. especially at a tempcralure of 35° C, and tho mix-
tare it found toaMitaiu |W|>{^ne. Theaclivity of the juice in lliua converting
proUMb tDto iie|>tijnc ii> lavore<l by iner^une of tfimjieratiire ui> to 40° or
iTmtilliiUlli. and himlrred by low u-tn{ivrikliin«; it in jiermaiientiv deMiroy«d
bv boiling. The digaitive imwt^ni of tbc jiiiu'! in fact de[Mnil. like tbi>M of
IfHCric juicv, <>n tbo prvciire <>f a fernieiil uhii'h, a* we havr already oaid,
nay be boloittd mueli in tbi? «anie way lu {tepnn i« inalatvd, aud to nhiob
tbe tume trjpnn bns been ^ven.
The appewmDoc of fibnn umlcrguing pancrcatii' digc«lion is, however,
Jiftimt fron that undergi>ing [wpiic <Iige«tion. In Uie former nur the
ftbcia doei not swell up. but remains n« opiujue as before, and appears to
Miflhr eorroaion rather than solnlinn. Itut there is a still more important
diitituAJoD between pancreatic nnd peptic digestion of proteids. Peptic
ilimlioa is toKutially an acid digestion ; we hare seen that lh« action only
laua plaee in the presence of an a<'id, and is arrasled bv neutralixnlion.
PWwnatM! digeatium, on tl>e other band, may be regarded as an alkaline
iljljllliili ; the action b nmet ener^tic when aomc alkali is prceent. and the
amrkj <^ an alkaline juice is hindered or delayed br neutralization and
■iiiti by acidiReation at least with mineral acada. The glycerin extract
of paDcnaa ■ untb-r all circuniotancea as inert in the presence of free mineral
■M ■ that of the uttiinacb iu tbe presence of alkalies. If tbe digestive
■ixnm b« nppticd with sodiuai carboDat« lo the extent of 1 per cent.,
dteatlen proceed* miiidly.Jitat as doee u peptic mixture when acidulated
wUb kj^rodiWic acid to tbe extent of O.'i net cent. S>x)ium cnrlronale of
1 per omL aeemi in fact In nlajr in tnrplic alftestioD a part altogether eom-
pniablr Iu that of liydnichloni- acid of 0.2 Mr cent, in gartric diMstion.
And jiist as pe|isin is rH]>i(ily tkviruyed by being Itcntnl to about 40° with
a 1 perceoL aolulion of mHliuni oarlHinalc, so iryiMin is ramdW d««K>jed
by heUK similarly bcatnl with dilute bydiw:hloric acid ut O.X par eenL
S&6 THE T1S3UK8 AND MEmANlSMS ()>' DiaKSTlON.
Alknline bil«. whidi nrrmu jieptJc dtgesliuu, seeraa. if xnylhiuK. GiTorable
to Irriitic digution.
Cornvpouding to lliix dilTfrence in tliu lietpmale nt Out fernwrn, ih«re to
in the two ciua ■ diMVri-ii<-« in thv niilure of the producU^ In both CMm
peptone is prwliicyd, wni itiich ilifliTca (■<■-■< lu can be dcb«Uyl betWMin iwn-
erratic and gnslric pt-ptoNtv an.- rvlntivi'ly kiuhII ; but in iiancrcntic di);(slion
the bye-product is not, n» in giwtric digcctinn, a, kiod iif ncid-albiimio, but,
OS might bo expected, it budv hiivint; more: analogy wilb Klkali-Klbumin.
Moreover, before the iilk)ili-nl!i<imin is iicLiinlly Ibrnted, the fibrin becoinee
altered and takes od eharnctcre iiiiprnicdiiilc between those of alkali-albumin
and of ordiuary albumin ; ntiii when fmti raw, i.e., unboiled, fibria is acted
upon by paiierealio juice, one or mort- (ilohiilins appear a» initial products.
Further, there are evidences that <lilicrenc«8 of even a more profound
iinlurc than the above exi^t beln'ecii puncretilic and gastric digeetioD. One
o( lbi9« is the appearan<% in the pancreatic dit>es[ioD of proteidi of two
roninrkable nitrogenous crystalline bo<lic8, leitcin and It/mtin. When fibrin
(.i>r other prolcid) is submitted to the action of pancreatic juioe. the anioont
of peptone which can be recovered fVoni the mixture falls ftr ihort of the
original amount of proieids, luuoh more so than in tlie caae of gnMric juiw:
and the lunger the di^tive action, the greater is tht.« apparent Inn. If a
pancreatic digestive mixltire he freed f>om tlie alkali-albumiu hv neutmliu-
tion and filtration, the fdlrate yields, when CDiiceniruled by evnponuion, a
crop of trj-atnls i>f tyriisin. If these lie reinoved ibt peptone may bo prc-
ci[>i|jit<'d fnim the on cent rated fdtrate by the addiiinn of a largo fixccoa of
alcohol anil M-puratcd by liltration. The necond liltratc, upon being concco*
Initcd by evaporation, yield* abundant cryatala of Icucin and Imcea of
tyroein. ThuK, by the action of the pancreatic juice, a ooneidernble amotint
of the prolcid which \t being iligv«l>.^l i» *o broken up ns to fpve rise lo
products which arc no loncor proteid in nature. From this breokiuj; up of
the proteid there arise leucin, tyrosin.and probably several other bodies, such
as fativ acids and volntile substances.
As ir* well known, leucin and tyrosin are Ihe bodies which wake tbotr
appearance when proieids or gelatin are acted on by dilute acid«, alkalies, or
various oxidizing agents. Ll^ucio is a body which, in an impure state, cry»>
talliusi in minute round lumps with an obscure radiate xlrialton, lint when
puri' forniB thin Kliltering flat crystals. It has the formulo <',H NO, or
VjH.j.NH, I.C0.O1I1, and is nmido-eaproic acid. Nnw, caproic acid is one
of ihe " fatty uciil " series, so that leucin may be regardeil iw a compound of
ammonia with a fatty acid. Tyrnsin, t\II,,NO^. on the <ithcr hand, be)ao)>B
to the " ari>matic " 9vti<» ; ic is n phenyl c>rupounil, ami hence idlied to betl-
xoic acid and bippuric acid. >hi that in pancreatic dimttion the laice com-
plei proteid incileciilr in nplil up into fatty a(rid anil aromatic tnoIeculM,
Bome other IkkUw of ItM imjKirtance making their appearance at the same
time. We infer that tlie proteid nioleculei are in some war built up out of
"faUy acad" and "arcmatio" molecule*, logothor with oilier componenia,
and we sliall later on wc additional rmsotw tor this view.
Among the HU]i|iknii-ntary nniducls of poucrmtic digestion may Iw men*
tioBed the body imtol (C,H,N), to which iip|>nrcnily the strong and pocii-
Uarly fecal odor which someiimt* m[ik<;:> it* amx'nrance during ponorcaiio
digeUion is due. Indol. however, unlike the leucin and tyrneiu, b »oi It
product of pure pancreatic digeclion, but of an accompanying decompoailiiMl
dUB U> the action of organized ferments. A pancreatic digeUive uixtiira
•OOH beoomee swarming with bnclcria, in spite of ordirinry preeautiont, whon
naiural juice or an infusion of the gland is used. When iaohiterl ferment
to ined and atmoepberic germs are exdtided, or when |>au«reativ digestion
PANCBBATtC JCKE.
M7
I
I
I
I
m eanied on in tb* pfceencc of inlicylic acid or tlirmol. which prevent tho
(WvdofNnmi of baclfrin anii lik>^ organUm l>iit permit tho action of thu
trjotio. no odor is |tcrc<'ivccl nnd no indoi n produced.
Aft«r lonp;-c»i)tinticil di);c8tioi). cs])cciullT uhen R«.-ompanied by putre-
iMStfn deo>n>p'>«iti<<ti. iho ftmoiint of pmccids which nro carried boyotnl the
fwptoiM tH^gf nnd broheii up may be very i^mil.
Id f^utric digCBtwa such a profound destruction of protcid nititcrial occurs
to « rauch lem extent or not at nil ; neither loucin nor tyrosin can at prflseot
b* eonmdered as natural products of the action of pepsin.
On lite |elaltniferoii8 elements of the tissues lu they actually exist in the
timoK preTHMi* to any treatment, pancreatic juice appeHra to have no solvADt
action. The flbrillie and bundles of librillse of ordinary unlouched con-
DxtWc tissue are not dif^ed by pancreatic juice, which in this respect
aflbrds a strikinf; c»uirtii<t in trastric jutce. But when they have been pre-
viooilj Heated with acid or boiled, i»o ai> to becunie convi-rieil intu uciuul
nIatiD, tripsin is able in diwolvt.- thcrii. itppareiitly chaDjpnK tbem much in
Dm mMot way as dom pepiuii. Tryjwiu. unlike ]iein>iii, will di.-oulve mucin,
Lilw pepiin, it is inert tuward nucleiti, horny luwues, and the so-called
•inykad matter.
On /aU pnncmalic Juicr hii* u innfotd action. In the first place it cmul-
aMn btik If hci^t'* Urd Ix- gently liciitMl until it melt* and lie thun mixed
with paoon-alii' juice hefi>ri< it imlidilitv c>d cooHng, a criiiniy cmnUinn liMing
£»r almoal an indi-tinitc lime is jV>rnii-d. 80 aim; when olive oil is iiUnkeJi up
with paocn-atio juice, ihe wparniion of the two fluids takes place very slowly,
and a dmp of the mixture under the micmecope shows that the division of
lb* &i is venr minute. An alkaline aijucous infusion of the gland has
aimilsr emutsinrnK powers. In the second pince pancreatic Juica splits up
ncutnti falfl into their respective acids and glycerin. Thus, palmitin (or
tn|ialmttini(1-'iiii».C().U),.t',II, is with the a^umption of .'Ul.Udplil up into
tbrr« molecules of |ialniilic acid :i (('„H„.CU.OH) and one of tclycerin
(C,ll«) vOH,) ; and so with the <>ther neutral fal&. if |)erfeolly iH-tttral fat
be tnated with panoreatic juiee, eiiiie<-ially at the body-tum|R.-rature. t)i«
■mullioa wfaioh is formed sptwdily lakiv on na acid rtacton, nnd by appro-
priata means not only tbi' ci>rri-)i{ii)ndini; fatty uddM, l>tit g^lycy^rin mny be
obMimd ftwB the mixture. When alkali i* preacnt, the fatty nci<b tliux Kot
fna form tlisir oorrfspowling soaps. Pancreatic juice conlaiit* faUi aiul is
OOnsaqOMitly apt after collection to hiiw its nlkulinity rcdu<yd, aitd an
•qnwus innision of a pancreatic gland 1 which always contains a coiuid-
•table amount of fat) very speedily becomes acid.
Ttius pancreatic juicv isrsmarkablo for the power it possesses of acting 00
all the foodstuffs, on clareh. fats, and prolcids.
Tho action on starch, the action on protvids, and Ihesplitting up of neutral
fiua appear to be due to the presence of three distinct ferments, and rn«tb«la
have DMO suggested for isolating them. The emulsifying power, on the otber
band, is nnnccted with the general compositionof the juice(orof theaijueoua
influion of the gland), being probably in large measure depemleni on the
alkali and the alkali-albumin present. The prole«lytte ferment, Irypbin, as
ordinarily prepared seems to be proteid in nature and capable of giving rise
by digestion to peptone; but it may be doubled, as in thecaseorpepeia and
otbir fmuents. whether the pure fermeui has yet been isolated. Tnere are
no mewu of distinguishing the amylnlytic fermeut of tlie pancreas from
piyaliu. The term pancr'--itin has li«ea variouslr ap]>lied to many drlTerenl
prvpaimlinns fmni the glaud. and itn use bad, perha|i«. belter be avoided.
Tbe acti»n of paucr«atie juice or of the infusion or extract of tlte gland.
I aUn-h, is seiEii under all circumstaiuns, wbether the animal be fasting
iHB TUK TlSai'KS AXD MECHAMISUS OF DIOESTIOK.
nul. The miue niav probably l>e ^id of the action no f»U. 0» prat<
the DiittinU juice, wliuu ULt^reled ui u normal state, U alwayn active. '
glycerin cxtrani or anucous iiirunJoii »!' the gland, on the vomniry. aa we
nave alrea^ly Gxplaineu (^2:t>*), iit active in pfoportioo aa the tryiwinu
b«« been (<»iivcn«il into tryjiain.
Sufoat Eiitcrieiu,
re IB Itefl-
§ 200, When in a living animal a portion of the small intotine is It^-
lure<l, nil ibnt the eecretionB coining down from above cannot enter ita canal,
whik' yet (he blo()cl-i4iip|>ly is maintained aa uHuaJ, a small amount of secre*
tioit colleutA ID it« inieriur. Tbis is spoken of as the menu enUrieu*, and b
•upposed to Im furnished by the glands of Lieberkiihn, of which wre shall
preaentl; q>eak.
BuccuM enterlcus mav be obtained by th« following method, known at that oi
Tkiry modified by VpIIh. Tho umall inmtiiiv i« divided in two places ai some
dlatancv (3U to GO oiii.)apait. By fincauuires the lower end of tliouppofMClioa
ia carefully united with the upper end of the lower auction, ibua, a* it were, cut>
tine ^"^ " **l>ole pleci- "f the small iuioiliiie Irom the ulimeiitnry tract. In suc-
eessAil cases union between the cut surfaces lakes (ilnce. and a shortened bui
otherwise satisfuctorv canal in r«^'siablislied. Of the liuilsied piece, the two ends
are He pa rate ly hrouent through incisions in the nbdominal wall, anil tfaetr nianlh*
carefully fanlcDed In such a manner that each ujouih of the- uiece open* on \a thr
cxlerior. During the prooeas of healing two llntulie are Idub eMablUied, one
leading to the beginning of and the other to tho ori<l of a abort tneca of IntMtine
quite Jsolaled from the mil of tli« alimentary canal ; br nirsn* »f tbcH Openings a
small (|ijantity of lluid can be obtained.
The ijuaiiUly seeteted Is said to be considerably increased by the adminisiratioa
of pilocarpiu.
i
Huccus eutericus obtained from the dog by the above method is a cl
yellowish tluid having a faintly alkaline reaction ami conlAiuiug a certain
quantily of mncus. It is said to convert siArch into ^uj^ar and pfoteids into
pcpI'Kie Uhe action being very similar to that of |Htticreuti<' juice), to aplit
np neulrul fat<>. t>i emiiL^ify fuls. and K> curdle milk. It is also aiud to con-
vert nipidly cane-.^ngar inl» graiie-sugar, and by a feriuuutative notion to
convert cane-sugar into lactic aiMd. and this again into butyric acid, willi
Uio evolution of carlmuic nci<l an<l free liyilmgeti.
According to the above rMtili.'> .■>iiccii!> enlericus is to be ngardedaaao
important nvcretJon acting on all kindu of food. But even at its be«t iia
actions nro i'l'>w and tl-chlc. Moreover, many olHN-rvrr* have obluincd nega-
tive results, so ihnt the various stalomcots tuv conflicting. Besidus, w« havo
no exact knowMge as to tho amount to which such a tccrotion take^ place
ander normal circumstnncc* in the living body. Wc iniiy. thcrefbn?, con-
clode that, nt present at all evcnb^ no have no satislnctory reasons for sup-
pcning tliai the acinal digestion of food in tlie intestine is, to any great
extent aided by auch a juice.
Of ibo powible action of other secretions of the nlinientarr canal, as of
the oecum and lar^ intestine, we shall speak when we come to consider the
obaniEee in the alimentary canal.
%w\. QaU'rimtti. Conerelimw, often of coD§>idembIe size, known oa gall-
MOneaarenot unft«quently lormed in the gall-bloilder, and smaller concro-
Uons are sometimes formed in llie bile naiHiges. In man two kinds of
gallistonea ore ounmon. One kind cwuists almottl entirely of cholcsiorin.
sonietimes nearly fVee Arom any admixture nilh pigment, somoiimos
more or liiw discolorcil with pigment. Oall-stonra of thi« kin<I have a
crystalline structure, miMl wlwn broken or cut show frG<|ilvntly radiaManii
SKL'ltKTlDX or PANCREATIC JUICE AND OP BILE. 359
nooMilfic Rwrktngn. The otber kind «io*i*u cJiiefljr »f bilirubin in com-
Mwitlun Willi cmldiim. Giill-«tODC« nf tlii» kind urv <Inrk-TOlorcd uid amor-
jtum, L<n Doninii>n thnn (lie «buvQ nrc «mnll, dark-oolorcd DtoDC*, liavlog
■ nalbnry shape, k-misieting not of biliruliln tiMlf. bul <^r one or vth«r
ktite of bilinibin. Gnll-stoncB coiLsisting nlmml cnlircl)' of inorgtuiio
ailcic c«rbonata and phoafduites, rnn nlso ocvii»i<>nitlty met wilh. Iti
lower ttnimab. in oxen for iosUaoe, bilirubin gnll atoacs «ro not unvoni*
mota . btit cl(olat«rin gall Btaoea »n tm.
A. pUI-stoiw Kpp«ara alirayv to contiitii ft motv or Ion obvious " niiclou*,"
uound vliicfa ibe mauHal of tbe stone hat becii dop(wit«I, and which nofty
!■* VVaided oa the origin of ibe «lone ; the rcnl cauao of the formation of ihii
*lii*ha1iei, bowever, in certain chanj^ee in the bile, by which the choleetorln,
•f l*ilinihiii. (ir ulbi-r cooalitiient oeaaea to [«niain diaeolved in the bile. Dut
*■ caouot dbteust this mutter here.
TUK SeCKETIO!! op rAXCKEATK; JuiCK AKD OP BiLK.
I t SH. The tffrttwn of panaytfir Juie^. Although in Hunio cmos, m that
k^ tbe p«raljd of the sbvep, the Bow of mliva in con tin nous or nearly bo, in
^^Pm ■Ainuls.u in man, tbe intortnitt<-M(.-i- of the «ecre<ion is very nearly
ibolule. While food ia in thr mouth mIivu dowit fnidy, but helneeii meala
only juM aoliimtnt is srcrvted to ket'|i ihi- mouth mniHt, and probably (he
rwtar part of this is supplied n»t by the l(up;r nalivary hut by the Amall
Euooftl ([lamia. Tbo flow of pnncretitic juice, '>n the other hand, is much
BOre protonged, being in tbe rabbit continiioos. and in the do;; lastiuK for
twMitT bours after food. But thi» eontrnst helwcT'ii the secretion of tuliva
aod taat of pancreatic juice it natural. «iucc the ftay of food in the mouth,
una during a protntcti^ feast, in relatively short, wliereot the time during
wblcb tbe Dtalenal of a meal is able in aouie way or other to afiect the pan-
trma h ^•rj prolonged.
Tba flow thouBh coDtinuoua, ur nearly bo, i* not uniform. In the dog tbe
fluw of pancTMlic juice b^ns immediately aHer food baa l>ocn tiiJton, and
tias to a maximum which may be reached within (he Rm. or as in the caae
Aimi«liing the diagnoi given in Fig. 1 1 2 tlie Nceund hour, but which more
caninwnly ia not reached until (ho thinl or fourth hour. This rise ia tlien
fall»wtyi by a Imll, aAer whii-h tlHTt^ i* a xi-ronilary rise, reaching a eeoond^
mazimtim at a very vitriable time, but gntemlly between tbo fifth and eevmtil^
hours. Thix second ninximum, however, i« never m high as the Rral.
Tbe Beoond rise may be doe to ninlerial alimtbed from the inteatiiies beioff
CBfried in the circulation to iho pancreaa, and >o direoUy excitin); the glaiHt
to activity, much in the aame way at. in toe cnae of tbe ttomoeh, the iilu-jrp-
tlao nf digolad naterial promotes (he flow of gastric juice, see $ '2^2 ; and
a aioiilar ahaorptioa may contribute \a> tlie Hrst rise also, but it \% more
protMible that so oiarketl and sudden a rise as this is carrieil <iut by M>me
MTVoua raecbaniHm. The details of this raeofaaiuBm have, however, not m
y«t been aatitfoetorily worked out.
Tbe pAncrea* derires ita nerves, which reach it along its bloodvtMeli, friMn'
tbe K>liir plcxut »f the nplanchuic ayelem, but tl»e ultimate; origins of the
flbra have not been imccd out ; auroe of llieni, however, oortainly ooiM
tbrMgh tbe plextt* fttim the right vagus.
Stinulalion of tbe m«<lulla obloa^la, or of tlie spinal cord, will call fbrtll j
wcratiun in a ijaiMcont gland, or tnercaiw a »ecret)on alreiuly gi>ing OB.J
Prom thb we may infer lli>' oxiolciice of a reflex tncchaniam. though wa
onant as yet imx out aatixfaetorily the exact path of oitJier the aderent or
360 THK TlSSt^ES AMI UECHAXISMS Or UI088TIOS'.
the eflervut impulses ; all we cnn say is, that Uie latter do not reach the pan
creaa by the vagus, since stimulation of the medulla is effective after :)«c(ion
of both vagi.
Pn.112.
S
z:;
^
;gl3H|S|6|7la|9llO;itll2!l3lt4llM'6lll^l3l4l5l6i?l8i9[l0
OutitAH iLLiBTacnNu TMK l^pML^ts^T or I'ooii □!■ tuE SeuitTioji or Paxouttc Jc
IN. u. BunmiN.)
'niB»l»otn» rBpHtlilbnurTnHtfrmilnj; Ibnd; Ihe oidliiaiM irfimenl In e n. Uio aiiwont or ■
UoD In Ion minulci. A markiMl riic li Hvn bI B linmgdlaulr nRct Ibul hu Ukcn. wiUi a i
im hflBiyiillhu founli niul tinh linun iiftvFHinli. W'hnv Ills llnv U i1i>ll(>] Itia otmtrmHon ■*
Inletmiilcd. tin fno4 Mug nmin rItdii al i~. ■noincr rite U foca. fnllowcd In tiltn br ■ i1li|ii ii^iw
and m wmnilu) rlw m tht- nnii liaiK. A rory liinUar ciirvu waiilcl n>|iramit ilin Mervdnn tt tot.
A Kcrvtion alreiiily going ou may be KrrcMcd by stiiniilstioD ol' tJte ceti-
trnl cikI of the viigun, and Ilio «loppngc of the x-cnaioii, which haj b«en
ohM-rvcil ail nccurrinc during and nttcr vomiting, iv prolmbly brought about
in lhi« way. Thin ftlWi. whirh, howcvor. ii not roRfinwl to the Tajfus, gtiniu-
lalion nf i>ihcr atR-rent nerTct>. «uch at thtr eciatic, producing the nme efl^t,
may bo regarded (in th« alMciiee of any proof thai the result is due to reflex
oonstriclionof tbopuioreatic bloodvemeels unduly choking the blood-supply)
M an inhibition of a reflex lUGrbanifm al ils centre iu the medulla, or in
wnne other part of the ceulml nervous system, much in the same wny u
ftsr inhibiu al the central nervous syslem the secretion of saliva following
food iu the mouth, S, ^2ti. Bui if so, then we must regard the secretion o?
pancreatic juice as closely resembling thalof salivii Id as much as it is wiled
foitb by a rcllex act. Yet it is Hlaletl that, unlike the ciisc of Miliva, the
secretion of pancreatic Juice eontinuts after all the nerves ^oing to the gland
bave been divided, an openilion which would do anay with the piMMbility
of reflex action. Kuch an cxtifriDienl, however, cannot he n.-ganli-<l ax de-
cisive, since it is almost im]H)mildt! to be sure of dividing all thi^ ncrvm.
No evidence has yet luxa brought forwnnl to j)rove the uxiftcncic of tiny
double nervous mechanism nmilar to that of chorda fibrts and tympatJielic
fibre* In the saliviirv gland. All that can l>e Hud is that, whi-n thv gland is
stimultitt^ to .-^crete, the blootlvcMQls ar« dilated as in tlie salivnrv glaod;
and wo have already, § 2^9. dwelt on the hislolugicjil changw wbicn accora-
pnny accretion. Vt c mny add ihai when the gland is otiniulated U) incroaed
secretion, the increase is not nieroly an increase of water, llie discharge of
SKOKBTION or PANCREATIC JUICE AND OP MILK 331
wMda If inercMMl fives mttn than tbe diachKrge of water, so tbat th« pe^^
tmt»0r of tolids in the juice iiicrensea.
TIm iiiMiilitjr of paDCTMtio juice eecreied, in the case of man, in twenty-
knr hoan hna been eiOeulated at WO cc, but such a culcninlion is of verf
Mcgitain valiK.
%Vr Invc Been, $ '227, tliat in tbe snlivarr inlands lli« pressure, nhiolt may
W mxttutl bv the fluid in the duot». is very considerable, exceeding it nmy
fc« •Ten the biood-prwaure in the carotid arwry. In this respect tlm pwi-
tftmm^&n from the salivary glandH. When, In a nd)bit, a eauula iwt-
■McCvd with a vertical tube or a nianonierer la |ilat«<l iu the puucrealic duel.
*^ ColuiDD of fluid doe* not r\m> alxivi- ii beitint correapondinif to a ptCHura
>>( aklMMii 17 nm. of mercurv. But al lbi» pruMture tbe Riand becoin«a oAe-
iMftt^Hu on account of the Juioc wcrded ptuaing buck tbniugh tbe walls nf
**^ dnota and alveoli into tho conncctiro tiamic; a much higher praMure is
■^•airf to raider a nlivnrv jrfnnd ind«mntou* : niid whether the low pmaure
'^"•ajiwl In the pancTOU i» due to the en*c with which micnm takes (ilnce, or
** *. Ill- actual wcrvtion not being able Id reach a higher |in»»iire. cannot be
•*^*«il with certainty.
S» US. Thr Koriion of biU. Tho act of eecrotioo of bile bv the liver,
*? ^^M nol be conlbaiidcd with llie disL-hargc of bil« from tho bil« duct into
^^duodaaum. When the acid contenls ul the rtonmch art- |H>ureil over the
'^T^^ lire of the btltsry duct, a gush of bite takes place Indeed, titimulation
tkL ''*'* '*?*''*' °^ *^ dno<lenun) with a delule acid al onoe calls forth n flow,
^^«ugfa alkaline fluid* ao applied have little or do etfect. Wlicn no Rich
^_^?id fluid ia paMiug into the doudenuni no bile in, under normal oirciim-
^^VMMt diaobarxed into the intestine- The diBi.'liur>>e is due to a contraction
^Y ibe muacular walls uf tht^ K""-bladder and duc-tii, accompanied by ■ ro-
'^^tation of the ^tphlnvter of the iirillct.-; both acta are probably of a reflex
*^siun-, but the detail* of the me<.-bgint»in have not been workol out.
Tbe M-crttinn of bile, on the other band, as ohowii by the rcautta of biliary
^atnUe, i» continuouit ; it nppenra never to wa»c. When no food ia Utkeii ibe
\aHn p«Mn fr<>m the bver along iht: hcpalir and then back along tbe cyatio
«3act (the flow being aided pr<tbnblr by peristidlic <«ntniction» of the niu»-
«alar fihres of tbe duct) to the gnll-bliidder, where it i» tem|K»nitily *(»re>l;
kcoca in alarving animaK when no diH'hari>c is excited by fixid, the gall-
tilsdfltr bccumcs greatly dinlcndcd with bile. Itut the Kccrctiou, though
oootioiiaiM, ii not unilorni. Tbe rate of secretion varies, and in ivfiecialty
■afloeon-d by food ; il is seen to rise ramdly after meats, rraohing itJ> maximum.
tBiloga,infromfourtoeii;hl hours. There eecma to bo an immediate, sud^len
riw when food is taken, then a fall, followed Bubse<)uentlT by a more gradual
rlw up to the maximum, and ending in a final fall to the lon-cnl ]>ninl. The
enrvig iif teeretion, in fact, reoemblea that of the secretion of jmncrcatic juioe
in having ■ double rise ; and as in that eaae ao in this, it is venr praMhle
that tbeflratriK ta in part the reHult of nervout action, and it is also pomble
that nervous influencee i»len,-ene in the aetiMid. more laaling rise : but, a» w«
■hall !we prcaently, even oerAoaa influeiieea may aflect tlte liver in a very
indirect mnuner, and our knowledge ns to any direct action of the nervous
•yvtem un the liver la at preaent very iniperfeet.
The liver teceive* il« chief nervouw nupply from the aniar plexus, and to a
^H pmt extent throu^ that part of the totar plesun called the hepatic plexus,
^B whjieh enbncea iho portal vein. be|uitic nrti^, ami bile duct, an tfaeee plutige
^H into tho liver at the ]>oTta. The M>lnr plexun if feil hy the lw(> abiloniinal
^H fpUochnic norvm, nnunr nixt niinrir, by <ilbur »niallor nervea fri>tH the lower
^^ parta "f tlie >|ilanchnic (aympatlHlic) chain, and by tlw tcrmitmt jmrtion of
ibi right vagua nerve, timul bnncbca from tlie ieft vagus, mnn hcfiatici,
I
a
I
THK TISSUES AVT) XBCnA^CISltS OP DIOKSTIO.v.
>lao pius directly In rh« lirer froin ibe leriniuatiom of (hat nerve on tbe
ttomnch, fiudioc ilieir way aUu Lhrougli cbe (torta. The ftbrea thus enteriD|r
Ui« liv«r from the several mourmh are, for the aum lurl, non-meiliillniMi
fibr«a; with these, however, are nixed a certain number of roedallalod fibres.
As to the funutitim of iheM uerve* iti n-rereiic« to (he aevmion of bJle.
n e mar say nt once (bat no «Hti»ractorj or cxu-i outemt^nt can at prewnt ba
luailf.
$ 2&4. It muitl be ruinumbered, however, tlutt the liver Is »> i)ecullarlr
reluti^d to th« other orgaiwof digmtioo, and its va»cular arntii^ii>cnt« w
api^ciul that, with rmrd to it. as compared with mauy other organs, an
inl rinHii; iiervoua mechaniam must occupy n more or km subordinate poriliun.
Th4- blood-supply of the pnucrens for inatniice is dependent chieAr on tho
width for the time beiu);; of tbe pancreatic arlerie«; it will be afiecled, ol
eouiao. by the ^neml arterial jirra^ure and by any circumstances whicl
tflbot the outflow by th« pancreatic veitis, and therefore by the condition 01
the portal venous system of which th(«e veina form a part ; hut iu the main,
the amount of btood bathiiif; the alveoli of the pancreas will depend on
vrliether the pancreatic arteriat are coii^lricted or dilated. The quality of
the blood reaching the iwncrea*. being arterial blood drawn direct IVom the
arterial foundation, will \te mtKlilied only by «uch cirenmstances as modify
tite eeneral ma« of the blood.
Very dilTeruit iti the ca»« of the liver. The fiipply of arterial b)
comtni; direct through the hepatic n^t^^^y is small compar^-d with the n»
jtourinji tbrougli the vttna pfirtie ; it moreover, as we shall see, i* dirtribut
in capillaries among the small ioterlohiilnr branches of the vena portie ai
haa beoome ve:iouK, indeed mcrgvd with tho portiil blood, befbra it reaol
the artunl lobules. Tboaupply of blood for the liver i* mainly that (hn>n|r'
tht- v<inH [lortic; and this supply is not, like an arterial supply, ii fiuny
uniform our. ntodilied chieSy by tlie vasomotor events of the orgnn itfM-lf,
hut in de|K-ndent on what happeoa to be taking place in the alimentary ennal
and in abdominal organs other than the liver itself. When no food ie hv'mii
dig<!H(ed and (be alimentary oanal is at rest, the reaaels of that canal, nx we
have already said in speaking of the stomach, are like those of the paiicroiw
and salivary glands, in a stale of tonic constriction ; a relatively small
(juantily of blood paasea through tliem : hence the How thn>ugh the vena
Dortffi is reUttively (Dcoiwiderable, and the pretaure in llial vessel is low,
When digestion is going on all the minute arteries of the stomach, intesliD<
»pleen and iwiicretn are dilated, and geiicral arterial preMure being bv sooil
meaiu or ulhiT maintained <»w- i I9i), a relatively large quantity ot blond
rushes inUi tlxi vena portte and the pressurv in that vewel l>Momea mueh
increaHcrd, lliongh of courw remaining tower than the g<eiM^r)kl arterial ptea-
sure. Moreover during digestion, peristaltic mnvi-oMnts of tho muacnbir
ooata of tho alimentair canal are. as we have «i.-«ii. active ; and theee oun'e-
mentfl, serving us aids to the circulation (see § Til), help to incrauo the
portal flow. Further, the spleen, as we shall »ee in epuaking of that organ,
ii in CDOny animals richly provided with plain mii«cutar fibres, and in saeh
CMW HOOnis, especially during digestion, to act as a muscular pump driving
the blood onward, viih lucrensed vigor, along the splenic veins to tbe liver.
So that even were tbe liver not ooni>e«ted with the central nervous rretent by
It single nervoti!) tie, the tide of blood through the lirer would ebb and flow
Mcording to tbe absence or preeence of ftK>d in tbe aliinentxry oanal.
An incresne of bl<iod-»iipply does not of course neoeasarily mean an
iocmse of secretory activity. As we have seen, § 327, in the preuence of
aln>|Hoe. the secretion of nliva may stand Mtill in ipite of diUted bluod-
vevela and the consequent niali of blood ; but we may safely aiueri that.
I
okriUag* bring equsl. ii fuller bIn<Ml-«upj>l}- » IkvornbU) to nctivity. Ap-
pnallf ■ mwr* cbiw)^ in thu qiiuntilv ol nlmKl bathing ma alvmii* will
Miaui Is tfa« odU tbc ohanK«* urhidi comiitiite the act of sovretion, anj
Mnihan aa tncttaM in the bliwd bathing a muKCiilar fibre will noocmirity
Mmaic n coiilractidn : liul iintma tUorc bn ttoniccminUT-iictiiig infliu'iicc at
ntLa tullrr uid rirbcr lymph Hriiiiinl it cn-M wilt niitiiritlly W'l to ihr ex*)!
likiqg up iDdriK nHMrial fmni thv lymph, iiiid »» will in(Tcii>o t\n- ccll'ii «lon-
^mtfj. HcDM, Mpaeially in thv hvpntic cell, which Hpp(mr» lo be nlwnys
Mnrii.alwav* tinilrrgoing mHaboliim of iiiich n kind a* to givr rtim !<■ bile,
■t nit^t liiirly expect the greater flow through th« portal v«in to quicken
iJhrmw through (ho bilo <)uct.
Aai) at a ninltcr of lact we do Bnd vaao-conetriclor action dominant orer
licBForvtinn In (be varioui experiments which have boen nmde to Rsecrtain
ifae arliikD of the nervou« system on the secretion of hile, it luis always been
fnuiMl (liat rtimiilB(ion of the metlulla obloogata, or of the spinal cord, or of
tJi« abdamioal splanchnic nerves, bIods or at least checks the flow of bile.
XtTW tlic eflect of these slimulatioae W, aa «« have already seen more than
ooc«. a powerful cnnstriciinc action on the abdominal bloodveesels ; by such
AiiDuUtion (Jie blood-supply of (he liver is materially diminished, and in
eDU«n|iivtice tbe aacretory acurity i» slackened or arrested.
Ilut iht^re It something bcaidea the nwre quuiiuiy of blood to be oonudered
la ikia relalMMi. The bloml which poasea mim tlie alimenlHry cunal at rest
ia vtSoBTj vanout blood, laden timply wiili cnrbimic ucid and the onlinarv
pradoMiaf the metabolism of the iiiu.->cular and iiiucouh couts of (he canal.
wImb dEmtien b going on (ho purlul IiIimhI i.-< ]udL>n, ua we «hall h«, with
tntut at all evcnla oTlhe produce of dig«>iii>n. with lugar probably and wiih
VkfioM nfotcld bodiea. And il ih ijuiie puwlhle (»r even probable that some
of then Dodica in tbe portal blood reachiog th« hepatic cells stir ibem u]> lo
■acnton* activity ; indeed this view maybe regarded as supported hy the
beta that protetd Iwd increased the (|uantity of bile secrolcd, wheniis its
fitlty feud, which as wc »hall see passes, chiefly if not whoUv, not by tbtt
noiul vein hut by (he lymphatics, and which is probably larveiy dispoml of
m Mne way or o(her before il can reach (he liver, hns no such etfecl.
Heooa we may infer that at nil events the second increase of the flow of
bil« which oocan during the la(er stages of digestion may be to a large
aait4:n( (be direct eflbct of blood, laden with digeatire products, pasaing from
t stomach and tDtestinea, especially the latter, to tbe liver by tbe portal
Bh, quite iudepeudeat of any direct nervous action on tbe liver itaelf : nud
\oAt*a It is fjiB9thle chat the first ri.w also may be partly due lo tlie increased
flow of blood I'rum tbu stoiniu-h, aidud by (be abwrptiou from that ot^nn of
k cartaln aaaoant of diicuile<l nmterial. Since, honerer, there la no evidence
uf any decrease in blu(Hi'jiiip[ily, or :u thif rate of abaorptlon, oorrespondtog
U> the fall b<-tween the twu hms miiih- luduences otlier tlinu those which «re
are discusung must lie ut wurk in the matter.
( SC6i Tbe blood-supply of the liver beang thus, cjulle apart from any
Btrmas supply of its i>»n. ao cloesly dependent an what Lt going on iu tlw
■lisWBtsry canal, it will be coavenivnt t» *av a few w»nii more connrnlDg
ifae nMoootor nerves of that canal. As wc liave alrcaily i>ui<l in speaking of
iba rascatar system, S 1*>9. the vns»-constrictor tibm lor the sloniach anil
iutertinsi, large and small, imuing from whut we mav call the vaMCon-
ttrictiir regioo of ihv mril (Hun for the most part throtigb tho two ab<l»minal
mlaocfaoic nerves, major and minor, n small numU'r only passing nut below
Uke roots of those nerves. When tlicw aplancbnir nerves arc dividnl the
Eof U>e cnnal are dilaKnl, when they are conirifugalty stimulatiM tho
■re oonelrieted. \Vhc4her there l>e any distinct vuso-^lilator fibrM fiir
364 THE Tissusa and mkcmanisms op DIOKBTIO.V.
all iir any imrl. uf the citnal, nnd if oo whnl course llicy take, u iwt knnvo.
Wlici) iii> itxiH hug fur sonic titno been taken, (he mucous membrane of the
stonmch ns Mftn Ihrough n giLetric listula is. p»lo : the blotHlrmsfIa are ooo-
Slrii;lc<l. Am) ni for iis vc know a simitar condition oblniDS Lbrougbout ibt
IIDbII nnd liirgv intestines. When food is taken the mucous membraiK of
the stomnch becomes dtished : its vessels become dilated. This nppcan In
1)0 the result of an inhibition of the previously existing tonic constHclioD;
at least we have no evidence sujiporting any other expltmatioo. Appareiitlj
ihe presence of food in the stomach starts in the mucoue membraoe iDRuenoea
which, ascending to the central nenous system, inhibit llie vasomotor centre
for the abdominal splanchnic nerves or such part of that centre as unrerns
the vaso-constrictor fibres of the stomach. !iy wliat path such aflereiit iin-
pul!>ea reach the central ner%'«u» srslcni i» nut oa yet defiuiiialy settles) ; but
poeMhIy by the vagus nerve, if it tie true, as slatird, that ceiiirijielal stiuii-
laijon of that nerve, vrhile it raises the jct^iieral blucid-jireiHure by iiicreaiing,
in a retle;if tiianner, VHso-constncliou in other regionf. IcaiU to a dilatation of
the K<>^tr!c vesaeU. 80 alwi it in probable that a* the food n-ache* succeeding
secliiinit of the alimontnrv canitl, thcw in turn in a similnr manner liccome
flunht^d with htiHid. In ilic frog (bent is some cvidonct> tbnl vai«>-coniitrictun
leavruji: the »|>iiial cord bv consecutive spinal nerves, govern the bloodvotwl*
of coUMculivi' M-cliiins ol the alimentary cniial.
All thi^i Hushing of the canal with blood lend*, wc repeat, to an incn-a^cd
flow of blood nt aliijrher pressure through the portal vein. Whether besides
there bo any additional mecbnnism set to work, such as, for in»tance. which
tome ohmrvations suggtst. u rhythniiciil peristaltic contraction of the portal
rein, l>y which the blood is siill more rapidly hurried to the liver, and
whether the increased venous supply through the portal vein is a^icompanied
by a eorrespoDding increase of tne lesaer supply of arterial binod tlirough
the hepatic artery, is not known. It may |>erham be here remarked thai
that there is no need for anv iiicreii«e of nrlerial blood, since the blood fW>ru
Ihe alitnentarv oanul, owiufc to its more rapid pi^^ through the minute
probably like the oorrenix
uilivary glaml 1 though probably aluo not to ihi; tame oxtciil) leoH veiioua
veMcl-<. v probably like the oorrenixinding blocHl in the veins of an active
than U!<aal during digiwlion in spite of (he extra i]uaj)ti(y of carbonic acid
thniivn into it hy the increased melaholisni of Uic muscular coat during thi.i
|ierb>tallic muvemeiits.
^ 259. It is interesting to observe that ihe pre^wure under which the hite
Is »ecruiwi is rclativelv low like thnt of Ihe pancreatic juice, not high like
that of (Jic taliva : it is much lower (han the arterial prcMure in the name
animal, whereas in the case of saliva (§ '227) the prvMure is greater than the
blood- prcMu re in tlie carotid artery. But, in the caw of bile, since the blood
whioh flows throngh the hepatic lobules is, mainly, venous iwrtal blood, vrc
have to compare tne pronnre of the secretion not with arterial preasuro bat
with the venous prcaure in the portal system ; and in the dng it hu been
found that while the prrasure of the bile secreted stood at about 21)0 mm. of
a eolutioD of sodium carbonate — that is, about lu mm. mercury, the blood-
pnoure in a branch of the superior mesemeric vein stood only at about 90
mm. of the Mme eolutjon — that is, about 7 mm. mercury. Now. the venous
preraure in the mewnlenc vniit is higher, though only slightly higher, than
that in the portal vein into which these pour ibeir blood (the dilferem^ of
pressure being the main cnusc why the blood flows from the one inbi the
other), sud In. thvrel'ore, ecrtainly higher (hnii the pmsure in the i>orliil
capillaries of the hetiutic lohuhs. ."vi that what is true of the salivary gland
is also tn>e, on a diH<;renl «cale, of the liver, vix„ thai the ptewure exerted
^
IN'I'UKTlNKl).
r lbs Hontion is h^^ber than the pre«nre of the bloo)) in tbo wnels fMing
m »ttavX\aa eelU.
f 907. Ifitie presHire in the bil«-duct bo nrlilici.itiy incrcnsed, ns by poiir-
}t>g tlui'l iiili> the fiifHB tube or tuaiionwter with which the caiiiila io the ilucl
it nmttectt*), a ivtu>rptiofi of the wcrelecl bile takee pItu.'L- ; and resorption
«ill «bo tnke plftcv wilhii) the body, wheu the preuiire s«Qerated by the act
rf»«mti"ii iUelf reacha and h luaiutaiiied nt ii -luifii-leDtTy bigli lev«l. Thtn,
«1mii in iii« liviog body the bile-du(.'t h ligatureil, ur becomee obitruc-ted by
KaU-aUxna or Uh«r»iw, fluid is ot^cumulated on the ueur Bi<le of ihi- ligature
at a prwure which foea on iocretising until rotorption of bile takea phn,
bite Mit* and biliary piKinent-i are rhrovru tiacic ufoii Uie ■r9t«tit,nnd "jniin-
iita" n^ulu. It nouM ap[>ear thut in thcsie eauea mMrplioii take* placv
tfaroagh the intrrlobulnr tiileHJucta aiid nut tlinnixh ibe hepatic celbor otlMir
•tractam within the lobultu^ Th*< high prvavure in the duct* ijoex nut lead
U> ■ rtvana) of tk' current in tbo hepatic oc1l« Tat in«sl it dacken* or \xi$4-
Ujr ttOM th« currrnt) but ihv bile tecreted into the iatu-lobular dnuU wca])«
ftDn tttMD. It further ap|)car« that the eerajM; in noL into the hloo<)v«neU
Irnt into the lymphnlic*; the bili? (alt*, pignicnt*, and other conRtiiiiciit* ant
otftied iiitu tlx- thonu'tc duct, mid in an indirect nmniier only find their way
iDio th« blood itreaiii.
To oomplete th« history of the »ocrction of bile we oiieht now to ttirn to
the maout'acturc of the biliary 'onstitiicnli within the cello. Ilut einoe the
hefiatic cells are alM> cnffnged in Inliora other nnd uuitc important, perhaps,
Umn that of t«erc4tDg biFo, it will be convenient to defer what we have to say
oa tbu point until we come to epeak of tbo formation of glycogen aod of the
gtmml roetabolio e%-«it» taking place io th« livw,
TbS f»TRUtTURK OF THE IntEBTINES.
Thf SitulU Jnhatine.
I SM. The intesline, small and large, throughout ita length from U
pylonu to cloae opiHi tlie reetuio, follows in it£ structure the general plan
piwionair doeribed, S '^OH. A thin outer loni^itudinal muscular layer,
eoreml by prTitoaeaiii, is succeeded by a thicker inner ciruubir muwulnr
layer, and thia double nuacular coat is separated by a eiibniuooua layer of
loaat eonaective timie carrying the larger bloodveBBels, froni tbe niuooun
liMlwhriiii which eoDsista of au e|ii[heliuni lyiltK upon a eonnective-ti^MiC
buit of peculiar nature. » u elide veli>[>e<l mu^cularis niueosie of luugitudioal
•ml eircular HbrM marking uti' the luucoui membrane proper from the under-
lying >ubtiiui-ooii ti»ue-
In the uuall iutt»tine llw outer longiludinul inu^ular layer it evenly dis*
tribnlcd orer iho whole eireumfereiice of the tube and ia everywhere mnob
ifainoir than lh<> inner circular layer, which U tbe mor« iiugmrtant layer of
At two. Tbe indivklual Ghre-ccllii of these muM-ular layern of the inleslino
ai* large and well-drwioped. lu tbe thin Kheet of connective Ua>u« which
•ejianues indistinctly the two layer* li«i Ibe plexus of Auerbach, a plexus of
iMrT*-fibres, for tlte uK»t prl non nxidullai^^l, iit the nodes nf which are
KBtbend groups of very >mall nerv^cvlls, ihe sulwtnnce of each cdl being
eapecially scanty. This plexo* »up]di(M the two rausrular tnycni with nerve-
flbns.
The fubfOUCHis cml cimlJiiix, bnides hloodvaiMit and lymphalioi, u #»me-
wbal nmilar plexu»of iicrve-liliree, calle<l the plexuK of MtiMiier [Fie. 113]:
A>aai this plexiw line nervi-Kbtm pnicred to tbe blootlvvwels, to the mils'
' I inuoMK, and possibly Ui uthcr rtrudurce.
I
I
366 TlIK TIBSt'KS ANP KKCnANISHS OF DI0K8TI0N.
^ SAO. Tlic mneoHt meml/rriHe, Tbiii is tlirown iuUi folda wlileb ant tiot, u
ill the oue of ibe stomacli, tenipornrj' loiif(ilutliQKl foldA, rujw, but pnrnia-
IPm-iii.
^r-
Ifuxw or tinvMia nMui nt* tWsKVOoiv Omt iw m* timwnMi. |Carui.}|
nent trtnerene fbl<b, the ratniltr emtnivmbv, raichliiE hKlf-wfty 6r twu-ibirds
i>r the way rwutid tho tube. Kach fol'l u n fold of ln« whole mticouM mem-
bnne cnrrying wilh it « part of tbe tmbmueous liw«ie,
inu. iw ||,e liillor tluiit furniing it miHillc nbwl bctwwu ihc mu-
cous mcmbniiic uii tbc iipjwr mirrHon an<l tluii on the
lower Hurfncc of thi: fo!iI. The fi)l<K wliiih viiry in
eiu?, large aixl einsll fn->:|iicntly alccruntoly, Itcgin to
appear at a little ili^ttinoG from the pyloruH; (bey are
especially well developed just below tbe ojKDiof; of the
bile and pancreatic diictfi. and are coiitiuue<I down to
about tbe middle of the ileum, where, becoming aoialler
and trreguUr, they )t<^diial1y disnppear. They aerve
to increase the inner surfaoe of the intcaline and preeent
an obaUcle to tbe too rapid transit of material along
the tube.
Orer and above tbe ooaraer inequalities of surfoce
caused by these foUU. tbe level of tlie mucous mem-
brane ia broken on tlie one hand hr to»];ue-like pro-
A 014SU ur utDU' jectiuns, the t-i//i, and on the other baud hr tubular
■■.'■Hi dcpreniooB, tbe ;/laHtlt or erypU of LifbtrkHin. [Pig.
114.J Tlic latter arv vi-ry mnrh sunaller and aro more
numcroiu than the former, neventl cryixa iw'mg placed in the Interval
between two villi. Both are found Oiii tho pi^emng valvulie at well ws in
^
Tils STBCCTl'Itli or TlIK INTESTINES.
367
I
tbf raUr^ btint^D. aod both cstrud along the whole longth »r lh« intMtiiie
Ads) lb« pjlorus Id lJi« ile)]-<'irctil vnlvv; but while Ihv villi varr> good
dMl, betBgtbart ond frvr imnKdiulely ii«-xt to th« pvloru^, very DumeroUB
■nd UrgB (d Ihf <luo(l<'miiii iin<l iiptii-r ]>art of the mtrctinc. Icvs niinii^roUB,
■iDKller, ind more imrgulBr in (ho lo^rr jinrt. ihi; crypts hav? nnu-lv tltp
■•Rt« ehtnctcn and arc ntiirnituly ditlnhiilcd throiighouC Vorr mufh sh
in tit* cm* of lfa« ftiinmch, tht iuti»ciiUri« mucnent runs in an eveo Hue
<txcc|>t fur the svmiw of the valvuln' i-onnivi'iite*)) iit u litile ilUlaiioe
il**im iht bMM of lh« rlosely pnched crypts, anil at a greater dislHDce
(»it. tbe length of the cnrpte) rroin iho basta of llie villi ; as we shall eee.
bwmr. the oiuscubrw muooeRmidaup musL-ular tibr«« into each villus.
f MO. Before proceeding to describe tbe villi and cry|itd it will be cixive-
aim lo study tbe diaraetere of tlw peculiar conue^-ltve liwue Iyiu;j betneeti
(ie raitheliuiu above Bi>d tlie uiuscularie mucoue beluw. The up)ier nirface
of tbia tisua k defiued by what may i>e fpolten of an a buneniciil niemliraiii;,
■ hich, however, spfieiirs not to be here (at Imni over the villi i n» iu the
MaDnM-ha«nititiuou>tHbeel<<oiii|H)»e<lof Hut con n um ■ vc- 1 Iksu l^ corpdnc In filled
Mr, but to have a Mruclure, which we ■Imll pmciitly ilrii«ril]«. Thv
•larii taaiXMae con«iMa of an outer longitudinal and an inner circular
t of plain mufrulnrfibrea. Iu NonM places the one, and in other placxtf the
atlwr brine prrdimiiDnnt ; each i>hc«t ooiiti*ts in moet cntrs of a single layer
of fibm. like couatiiueat fibnw being cemented into flat bumlles nod the
iMiDdtaa uDiUd by Bne connective tianic Between tht^ Unt bundles veaaeU
paa* Ui and from the snhiiiueou" lissui- Wlow and the rut of the muooua
MMabrane above tbe muscte itMclf boint: iilso well provided with bloodnawls.
Tl« cvaoecttv* tiawe which oocupiea the whole of the narrow irregular
aoaa bMWMa tbe baaonHot membrane above an<l the luusculariii mueosie
balow, cxecpl for the space taken uji hv the hloodvea^la and delinite lym<
pliBtk VMBds (of which we Hholl oreAently «]ieuki,iH of a kind which, though
It ii not ijuile the same in tbe villi ns elsewhere, is on tlie whole doeelf allied
lo the kind known under tlie vnnouB aama
of rftihrm or nfwH&tr connective tiwue, (n*. \n^
arfmwrf liaue or lumphnid tiwue, and iu-
hwlwJ is oHen CRlled bv Mt« or otiier of
IhaeaaiDM. [Fig. 116!]
"^ptml adenoid tianic such iis vt met
viitl) in tbe lynipliatic foUicKii of the in-
tosuae, of which we shall presently have
to ipwk. in lymphatic gland* and else-
wbn*, pmMtle lot apptamnco of ■ tine,
eloMBM. and fairly regular network, nith
lawbte M araall as not to aBord room for
mure ihaa one or two leucocyiee in each
meeh. The bars of the network are deli-
i-wie Hbrea compoeed of material which is
■imilar to. if not identical with, (hat of
the ribrilla.- of ordinnry connective tbaue.
At the ntrtlal poinu of the network ihick-
minKS are freijuently but not always pre»-
ont, and wme of the n>on> ootupiououH of
tliMe ihlekeniDgi may contain nuclei uther
apherical in form or mora or luai mis-
•oapen : but such nuclei are not Dumeroun
AdtfoaJd tianie. in fact, u ootopoaed of
aaaHomwing branched cellx, (ba graaiar pan of the cell in most com*, and
iaihad Um whole of the oel) in •omc cases, having been tranformed into
S\j.V,-v(i
.■s-
t'V-
caoMMBiTiu!! or * «uu- PaiQingn nr
Ttll Hll~<ni> MUUMtlll lir TIIK IKTW-
TUK, INCI.IiMM •IHI KniuK t'lvn or
LiKMMKiiiiH iMDMinortuanvnita*.
(MafulSnl 4(u illanicmi. |
a.eatliyulUir iiilul*' fUiiilioarrfd:
^iHHctf it>rUiiliicti<lt>iUal (I'll*: r. lb*
loMfslanlular or ■4*0014 IMnw : 0. tpsph-
otlk.]
(
(
,168
TUK Tissrl
JiW\
THOKSTIOl
filfttaeotoiiB prooenee of a dit&renliiiled nntiirc, nhicli join Trwly nitli <
other and wilh the like ptrxxeaa of other cells to rorni h line ri^'ular act-
•mrk, a portioa only of the cell, aoinetimes with and >>ometimts wilfauui Ita
oiMleua t.this bavioK disappeared i being left lo form n nodal tliickeoin^.
It nay be regarded as a leas develo]«Bd form of GOfiuenire ttuue than the
white Rbroua or the ordiuan' areolar conuective liasue. In the earlier Mtagt
of Ita derelopment in the eoibrvu conuective tissue of all kiodit a r«nre-
■ented by a iiuiabcr of nucleated grauular proioplasmic oelitt lyiiiK in n I'uid
or nearly iluid matrix. The eell-bodia are braiicbed, the tirauchcM joining
togi^hcr ul inlervalfl tu form a network. In the development of onlinarr
c«>omH.*tivv tiwiic ibc outer finrtjoii of the i^ell-biKiv of lume of the cell* la
convertetl into or at knxt p^'t* ri.-<<- to (iliriUnr E>*1at iiiifitroiin malmal, or th«
whole of it may be >o enlivened, ihe riwl of ihu cpII or uthcr tTfllfl being
)*It m oonnuclirc liauiv cor|HiiH.~lm. In ndeooid tiaiuc Ihe edls rvmain as
brandiod ci-IU joining into ii network, and the cell-wibrtaDce is not in any
part transformed into huDdlcs of KbHIln-, tliotigh it )uu ttndcrpiDe. Ixaidea
an increase in ile branching, in |>«rt at all evv^ntu, n cltcmical Irsnaforma-
tion.^ince the material forming' ihv ban of the network is in a large measure
no loajzer ordinnry "protoplasmic" cell-aubstanoe. The meebts of typical
admoM tissue are always crowded mith, and practically filled op by, ku-
ocH^tea of various sizee: it is only with very great diflictilty that the network
can be obtained tt«e tram them.
Tlie couneetive tissue occupying the spaon between and below tlw glanda
of Licberktibn is very similar to adenoid tissue, inssniuH) as it preaenia a
network uf delicate fibres ; but the meehea are somewhat lar)^r and mote
irregular (ban thoae of true aden<nd tiasiie, and though tbev coulain, an
not cniwded with. leiU'UL-ytes ; the anvount of («t I substance left at aonio of
the nodal ■H>iutii in greater, nuclei are more abundant, and aciiiio of tfae pro-
fVNim uf ttii! cell.-< I'oniiing the bare of the network are Hut rxpaiuions rvtber
than fibrm. It ixoii the w hole, therefore, aomewlml dilferent fn>m the typical
aili-noid li«i>uc' of lymphntic ?iriic<.ure9, and though it \a oflen ajwken of
under the Mime name an ihiit tiswuc, it will be convi-nient to dtMingUt^li it
by some term : it might be calle<l ratirtilar tittttie.
The tissue which mis up the bmly uf a villus difflsra Kiill more from triia
adenoid tiwue ; it is formed of branching cell" whicli linvi; for the rouat part
ntlaiDcd their nuclei and n larg<-r amount of cell-substance round each
nucleus; the proccwea an partly mcnihninouH, jwrtly fibres, aitd some of
them exhibit a tendency to form minute bundUsa of librilliv. Il is iiiter-
niedinle between ndentnd liwuc and ordinary oonneetivv tissue, and may
perhaps he described as forming a loose somewliat open spongework rather
than a network.
Lying loose in the meshes of this peculiar reticular connective Itsaue, both
In the villi and elsewhere, are seen bodies having the general charaetera of
while blood -eorpuseles 'see )i 31), which, though they are probably not all
of the same kind, we may sjieak of under the term of leucocytes, Si>i»^
limei these are soauty, but often are very numerous. This reticular eon-
ueotive liame forms, in fact, a labyrinth of irregular naassK^ which am
nciiipinl by fliiiii. but through which leucocytes can wander toand-fnt. Sv«
kIiuII lau-r on point out thai thia labyrinth of passages is aasbciated in a par-
ticular tiiHiiiii-r with ih<- lyiupbfltic veasela, and that the fluid ocoupjing the
8JUICC8 is in rt-nliiy Iviuph. IikIcmI, this tissue ougfat perliaps to be regarded
as part of thi- lyinpluuc :<yi>li-rii.
The basrmctil iiiiiidiniiir 9|)<>kcn of above appears to be formed largely, at
least over ibi- villi, by the expanded ends of fiorea of the reticultiin, which,
reaching the surface from below.spread out laterally beneath ih« epitheliui
THB STBDOTPRK OP THK IKTKSTINK8.
869
mill bi-!n^ juiiiei) b\ a i-erliiin iiumlier »f cv\\» lyirij; Hnl on ilio Kurfaw, form
lu^tlivr a nlioi't ntiich U iii>l Cdnliiiti'iu:! btil <li:>c(>titiiiiii>iiK, bi-tug hrolcMi )>ir
o]H>tiiiig!t thn>iii{h wtiicli till* banc* "f thi; riti\» of tbu Qpithttliiini nre broughl
into timUct with ibi- fluid ocnipying ihc xpaiti' of ihr rttit'ulum Mi>vr.
5 361. The illli. Th«' viUi vary in mIto Hmi form in (liRpju-iil unitnuls. Mid
in diRemit partu of thu iutcKtiiM! id (lu-Ninu) aniiiiAl; each villus, moreover,
riirMB in form nt (litfprviit times : tbev may be generally i]««rrib«d as haviii;;
(he stiapv of a flHtl<-iiecl tingpr, but arc froniiently broader at rhp free eai
than ai tin- bust-: tbcy hav«, in man, a letigiii of about I mm. and a breadth
of from 0.2 mm. to D.S mm.
Kacb viltuH consists of a b«4g of reticular tissue, the outer surface form-
ing, as explained above, a bHsemeut membrane, nbich is covered by a single
layer of epilhflium ailt. [f-'iu. 111!.] Two kiuds of cells, ihal i», cellit prc-
aeutiDg two seta of characlere, make up this single layer of epithelium.
[Fu. 1141
A. Vrun 0* HNiar: n. Vilu to' Miit, (SIlBtiftr snaiod ftom Tnci«a*io(.|
Villi of laloillae Kliunlnt ntlDinait «Idtb»Uum corerfCR Ihcni ; *l>o (itnuM Ualto, iMtliii In Ita
iBBba (An duk tn— li. wlilrji an fotM wplUjult*, and ■ lacteal it|>|ifarfn8M* vliltr loni|i.I
One kind bn 4>»lumnar or nmioal cell [Fig. II?], with Ita broader «nd
foriDtnff part of tin- tnv nurliatw of tht- villun, and ils narniwer t^ml reslini;
on or lUliOg; up a enp in llie biMcinout mi-uibrniii'. The gn-uti:r jmrt of the
vell-body in furmi-d '>f the kiud nf "granular" i.>^l!-*iili>tiinr^(t i<p<iki'n nf aj(
[>roto(>laauiir, but iIiHctm in app-iiraurR and <^>iiditiou ncmriliiii; l» circum-
nIaiirM; tbo>o vnrialtuiui wc iihall Klu<lr >«parat<-)r. A.U oval niKrU'U.t is
plni'ed vertically at about the loner ibinl of the cull. At the fr«ie hrtnler of
eoch c<-ll lh« firnuiilar wll-milNilttncv clmngat to a narrow band of clear
byaliiiD rvlnKlive mat^Ttal niarkml, in many prepared spceinicnK and oI\pu
4;veti to ti\t fmh alatv. with fiiw vrrtical Iin« so as to appear siriatv^l Ter>
tiimlly or rmtbcr radially ; in a MM'tion of a villus, optical or actual, the
1'4
370
whole Tillu^ BeetDi tu be ■iirroundi-i) bjr & bUMl of lliu cloir rvfmrtivi-
iuai«riaL
A ciliated epiihetiuni t>carE. m ire bave seen (S ^3). a »imili>r hynline n-
flraciire border fmm wliicb lli« cilia project and wilh whirti tUvr an; limi-
ll«(:t«il, but wliivh dnea Dot share iu tlie muvemeuts of the citiu btilonging tu
It, reffiaining uu<.-bau^ iu form while these »re moving; its exoc* tiniure
b at present uowrlniti. The refractive border of a cotumnar evil i»f n villus
diffen fWxn Ihe xiiiiilar border of a cilisleil oell tn thai on the one hand it
DC^'cr, ill vertebrates. )>ears cilia, ami ou tlie mher hand dues under cvrtaiu
[FN. 117.
[Fm. ua.
voimiuu CituTtD imatuni Csua]
uoataiciu.)
dreumelant.-n' change its form. The Ktriation cpoken of above appeoni to
be due lo the lad that the bonkr U comitcttoil of ii number of rods imbedded
aide by aide in n iubclance which is MmetiiuM of the same refractive p-m-er
u the rods, in which case thi^ wh')lf ImnJor nppcam bonioKeoeoue, bat which
it mmeiimes of diflerent refractive poncr. in which case the atriatlon i> di--<-
tiDct. The rode, which are thought by some to be hyaline proce^oe* of the
unilcrlyiug «dl-«]bslan<:e projeetiiiji into the al}<ive-im-nli»ti<^l ceJiicnt-«ub-
stoncG. are someltnieH h>ug and Ibin. Boinelinieii short iind thick, the whole
border being in the I'onuer (-u>e narrow, io the lattvr hruad. Under the tn-
flucnoc of reagent* or of circumHancea iIk one oHidilion nuiy change into
the other, and the change wemi to be an active not n iMMiive prc>c««e, Hnc«
it will only litke place »o lonx ta the cella ar« alive. Tlii> n-fmctive boixler
of ihu columniir cell of a rilliiii ia obvloualy a pceuliiir nixt presumably an
ini|Kir1ant utriictun^
I 262. Mixed in varving proportion with Ihe columnar cella pooMBeiog
this chnmi't«ri«tic hyalincninrvItT are cells of another kind, the ^blel eelU.
[Fig. 118.] Thcw lire cmenttHlly nin(y>us celU; in nil their important ebar-
aciers they rovmbli! tbo mucmii' cella previously described (§ 'i'Mii, but
reeeive their special iiuinc bcmuxo in shape llicy usually resemble a gobl«t
or tiaitk. In a burdened and prepared xpcdmen of a villus numeruuH goblet
ctWi uiay be seen scallered among aud aurrotindet) by colutiinar eell^ Kadi
goblet cell hiiB a bow, often irregular and ai^melimes brnucbed, lying ou or
near the bu^ment membrane, and a top which reaches ihc surface of the
villus between the refractive borders of the neighboring eulumoar cella.
Near the base is placed a nucleus, generally disc abiijud, owing lo tli« action
of the reagent, surrounded by a small qunnlity iif xmiuing protoplasmic
cell-MihMuni-e. Above Ihii^ itie cell consists of a niaea of iranH[Kirenl mucin
lying iu the meahea of a delicate reticulum, and surrounded by a thin layiT
or suvcloiw which id |>r'>l«U);i'fl tipnurd from the ceJI-subatauce heluw, and
which on Ihe to]> or fivu itirfiice of the cell usually beai^ a diiiliuct hiuimI
oridce or mouth. The up^>er jiart of the cell in e<.)ii»et|iifnily a sort of cup
filled with mucin (ami reticulum^ oud 0[>eQing into the interior of iIm in-
teeline by a somewhat narruw mouth, through which the mucin in due time
escapes.
TBB 8TBUCTCRE OF TIIK TKTKSTriTBS.
871
I
Id ■ villus ciBinifinl i|iiil« frvftk iu tioruial uiline sultitiou nome of tht»o
(obkt <vlli Duj be nWn'«l iu a uuikIUiou wkk-h bna beeu ilemTihnl (^ 'j:{5)
■• ttw Doniial oumlitinn ol' « niuouus i>ell. The t-dl U then crliriilrionl or
oral nultvr tbut dulincilv fliuk-iiliupcd, ami tbu upjxT jxirt ur cbi; cell coo-
iiMi of <TU-aulMlilllCO Murldiil with grnatilu* auil ^pltcnilu*. thi? traiwuurent
nutein baog thttal nod llw inoutit nut viiiible- Bill in jiertVetlx rmiti villi,
■ludinl andervvto titc raoM ravoniblu conilitioun. moiiy if not tDtmt of ibit
gobbt ctUa will be mcu tn have become goblM-ahnficil, t» hnvn nlmuljr
aDdOTgaiMi tli« traiMfvriniiltim into U«lW]Mi[«Dt nmiti nml rrliruliim, nml to
hmxf Bcoiirad « inoulli. Iu sucU cm«c« the cWr inuiajiuviit bmly of it
goblet cul atwub out in strong contrast with IIk more dim (pvuular liodin
of tbe oolamnar rvWf wh!t-li mmxind It, Ivith wHmi they uro M«r on thplr
iide uid when tlwy arc looked nl rnim iihovc. In ibc Intter cfl», irhrii ibo
BtCTOKope is focuned for a [loiiit n little below the fre<' KUrtiiec ol'lhe villus,
tb« gobJ«t oella look lik« routid clear droptelJ- »cnl(ered in the dim gmuod
fclBul bjr tbe columnar c«lU. A iiniit»r ci>n(ni»l ix tiHorded bv nrepared
■|wi<liini» ■HiiK'i with carmine and eertaio other dyi», which leave ttic traiui-
DAreat mueiu uni^tained. Under certain methods or eondiliMis of bnrdeiiioK.
KtweTcr. BKd with eeriain dyes, as with bwmatoxylin, the mucin mav staiu
I a* dacply <ir even more deeply than its surroundings.
Obfiiiuidy ihwe Kobl«t c«lls are simply mucous cells somewhat modiBed
bv mwiin of their pueitioo. Tliey are nut hidden in iIm rectsMS of on
alvtolua lik« aalivar)' mucous cells ; they do not form a layer by theiDselva
like the ipMtric tnucyxis cells, but ai-e»callered anioii); other cell^ carrying
oa inportant funelidiis. Hence amiBr^nily their iliape ofa Ktiblet and Ibeir
wdl-delined mouth. A ;;<>hlei wll to sinrt i^ith i' a cell «l' a more or lea
flolnmDnr form and urdiiiiiry |>r)>|i>)diu)iuie i-i-ll^sitbxtance. Tbe cell-subHano*
nutufarluitii iind bv^-otmv pluddtil with gntiiuleH ur Hjihernlcs which varj
Ecdily give ri«; to mucin, the- cx'll nwulh-n nilh il* luail amumcs » );i>lilct
(H. and the lonnation <>l ■ numlh in ih<- Hjiacu bctwecD the converging
n-frni'tivr Imrdcrs of neighboring eolumnar cvlU aMMuls in tbe discharge M
the hiad.
Tbe culuniiinr cell« of the villus are. m vir ahnll mx, chiaflv occupied in
tbe reception of material from the inle«tini- into the body of the villus; the
goblet cells arc chietly occupied in secreting into the interior of the intestine
■Bucio awl possibly some of the constituents of the kuccus enteriCTia
Bckxr tiua layer of ooliitnDar and goblvt cells rxtvmU the thin boKment
nwnbrane, above whicli, between the bases of the other cells, may be aeeu
oull cells, that is to say, celU with a relatively smnll (iiiantity of cell-
nbbtanoe nmnd the nucleus: iheee have been taken to l>e reserve »r re(>lace>
mrni cvlU. But at timeo clearly recogniirable leucocytes may be seen be-
tween tbe columnar c«II»: these have probably wsiKlerM into the qiith«Iium
floin the bndy i>f tW villus; and it may be that Mime of the aniall cell* in
(|il«etioii are of au allied nature.
fSflS. Th« rcntre or ruihtr (he axis of the body of tbe villus is occtipicd
bjr a cluli->ha|>i-d *)iace, •umetiun bifurcate or even branched iil the dutitl
«kI. rarting imlrtd a great deal in diflereul animals. This is tlic ccntnil
Waphatic spoi-e or " hirtt-nl tadicele," as it has tte^-u calW. whii-h may lie
Bllea with uitty or nthi-r material, or, as niorv frequently in mvu in harrleni.-(l
prvjMUlttion*, may bv cm|ity and (-ulla[»>:d. It is lined nith ci>itlicliol(l
plates, and '» nt the base oi' the villus continuous with the lympliniic pat-
•ages and vreeels of the mucous menibrana. Il will be coDveoient to deAr
tkt further study of ibis lymphatic apac* until we come to deal with tbe
Ijnpbatica geocrally.
Between this lymph^pncc, niid tbe bnscineDt membrane, geoeimlly oinit
372 TlIK TIBStJRS AXn MECRAKISHt
UDderncfttli ihe laiur, lies a fairly dote-set nelwork of citpillar/ vemtAt,
MpociKlljr well developed towanl the upper port of ibe rilluii. Tliis network
is feil Itv gi-ucrully "uc iimiill iiN.rxy which •prin^iii^' l'n>in the arlori«i of
ihv mlimucoiis limtic iipliti> up into cai>illnrin> tuwanl the midillf of tite
villiw; nml lh« hloml of ihfi cupillurio piixKii into vcijw, geaenWy twti^
which in n Mniilnr miioiivr pnw duwn to ihv vciiit o>' the «aI>i»ucwHi* liivue.
liotwooQ the basoment int?nihrnne itnd tb« cnntrnl lymph-itpaicv Brc iIko
found II number of plain mueciiliir filin^t. some ruiioire eingljr. otli«r« fonn-
iiig small hundlo! of two or three libnw a1>rcn»t. Th<w varj muub in
number and diauoation in diUcrout atiimnl*. Sonw of thoni lie clow itixter
and end in the btiwnient membrane ; olliera lie nearer tbo lymph-Bpnce, to
which in hohic aoiinals tliey form a »irt of luusciilar sheath. Thew fibres
belong to Ihe luuHculariH iiHicoaie: at the bnse of the villus the 6brea of lli«
rauftcalaris diucobw lake an upward (.-outw, passing between the adjacent
crypts of LieberkUhu, and run into the villus, following most commonly a
longitudinal but aonietime« ii mure or l«aa oblique or even a transverse
direction. Hy the eonlrattion of these fibres the funii of the villus can be
changed; but we shall return to this point »'heu we cnnie to s{«ak of il>e
absorption of digeated material by nicuii» of the villi.
All Iho space intervening between the basement membrane and tlie ecotml
h-mpb-spaoe nhitrh i* not taken up by ihe hlutMlvenelB and Uie muscular
nbm, it oceupied hy thi; Kj)eeial kind of retieular eonnecliire-tusufl docrilied
above (§200), tbo ni<«h<» of which are lu a greater or leas extent oocupi<^
by leucocytea. On the outer surfncB of the body of the villu:« this reticular
tissue is connoclcd with, and indc«d iix wc have seen forinH the bawoKiiC
membrane: in the centre it furnis anjund the epitlieb'iid jilales of Iho
lymph-spaco the walls of that cavity, nml suppUc* a nimibir bed for the
blooi! capillaries; the fine oouoective-tiMues lielongin^' lo the small bundles
of muBciilar fibres is continuous with it, and iome of the muscular fibres
ai-ciu 111 c-ii(l in it; to it also is atlnclieil the (-onncctive-tiMUc of the "uter
nalii of the small artery ajid veins. The body of ilie villus is in fact a
OpotiKe work of reticular tisttue in which are evacuated the lymph-apace and
tne blood channels with thi-ir m)[ieciive linings, into which ihe plain niue-
cuhir fibrus plunge, and which is condensed on the ouUide into a basement
membraoe. The m<»h<4 of the ajkih^ work are further occupied, as we
liave said, wiili luuc-x-ytm or wiih uuciealed oella of an allied but diller^iii
nature; heoco in unliunrv "lained apeoimens, not specially prepared. il>e
lymph-space and bhvul chunneU being oollnpaed, the whole body of tin
villus ap[><!an> a cunfuw»l miM* of nuclei ; there are the nuclei of the mu»-
cular fibre*, the nuclei of the epithelioid plated of the Ivmph-siMce and
capillaries and of the other coata of the arieri,- and veins, tlie nuclei of the
leucocytes in the muhea, and lastly the nuclei belonging to the ivIictiUr
ti»ne itself.
The tbicknon of the body of tho villus, that Is lo say the amount of
TCticular and of the other tinues lyiu^ between the boM* of the ciHtbcJium
cells and the central lymph-space. vaniM in dilTorent animais, being for in-
elance considerable in the doc ami small in the rabbit ; in the latter nniinol
tlie inuseiilar fibres are ver}' k^uiIv.
$S04- The rrypU or 'jt/iiidr fj IjitbrrkuhH. These arc found cverrwliere
nlunt; the whole leu^th of the small intestine from the immediale vicinilv of
the pylorus to tlie ileo-oscal valve, except immediately iinderiicaih each
viltuB, and in the ^poU occupied by the lyuijihnlic folliclee of which we sbiUI
pK-nentlv sjwak. Tlw mueoits meuihraiie or the Hniall intestine is in fact to
n verr ^argc extent iiiiide up of a number of these short tubular ^'otids
|>lac«(l aMo ujr side luid packed cki&ely toother, though not so clueely as th«
373
iiilnrlj-
it cnnlbf glnnHt
(he
['h: th<
tcUnds
I
It
slonin
ncx> of tlio iiik'xlina) mucous memfoimaci,
and llw muHtilBm mucoe^r nitis in n Inirly even lino a( eame lilUe diEUiDce
bvltnt thotD, ihsl is oiiltiHe their bliixl eni|«. Knch ({land is a Btraigfal or
atmrlr Hrni^bl tube, rarely iliviHin);. nbout 400»< ioof; and 70u broad. The
■wtltDe is fiimisb«<I by a ver^' dminct bssemcnl tnembrane, in which Ducki
arm UiAteMtd at intervals, and this boBemeat nietnbranc is lined with a
MOi^ Urer of short cubical cells, leaviof; a small but digLinct Iiiraeu; the
oalbaboiild perhaps be ratlier detcrilied ta eomewlint conicAl, with a brotuJer
b«>r U the bsBenient membrane and a oarrower aj>ex abuUiu^ <ni ibu
luDirt). The cell body, aurraunding a somewhat spherical nucleus, iit faiotly
gnuolar «xcepl for a hyaline free burder. which, however. i» ni>t *n oon-
■picaoiu or ao consiaot as in the eoKimnar oells of the villi. Similar celb
Cttnr ibp ridifM iiiterveuioe between adjacent irlaniU. and where a villus
ennci orxt t» a clood llie snort cubical cells or the f;lnnd may tic tnwod
intii mlomiinr crlU of thi.- vjlluf, the hyaliii« border becomine m'irc marked
and ibr nuclcu* IxH-iitiiing oval. Amno); the cubical cells nf the j^hind iiro
In bv fcund, in varyinn iiuinlx^ri. j^ihlct ccIIn l{uil<^ liimllar to (hiwc of the
tIIII. Ii (•;tuv(imvi* hapiH'ttii that duriiit; the )>ri.'iiaralii>ii of a sperimc^i the
wtiola CDJthaliiim » s)ie<i en twure, the cell* being much mors adherent to
•aril otoer thati to the basement membntnv ; in such a case the features of
tbo bwinvnt lucmbranv an; well *een.
Out«ide ihe bucm«nl membrane, between adjacenl glands and between
tfa« bliix) eiid* of the gUnd:< ami tlie tioderlying mmkrularis mucosn. Is
tvtiraUr coBnoctive-tiHue, tincr and more truly rocicutar than that of the
rilli: it is perhaps len crowded with kucocytos. In this reticuhir lEssue
run. mctrcling the glnndn. cApillary bloodveeecls nipplied by small artfries
cxHRtOff from the Mibmucona (issue, aad pauriny their ciIoihI into currvapnitd*
iwrras, and with thi^ iviii-nlar tiwmc lytnuhacit^ are ennnected.
TliMr glauiLi of LielM'rkuhn are suppcisei) to funilsh the suocus ftntericus.
The ra*an« for thia rteu- lie iu their tubular form, which is that of many
verntinn; mlnnds. In their lumen beioK
toil namiw for tlw! puuigc of food
iulo it. anil in tlM fad ibst, at wesliall
tlwy unlike the oluninar cells
V th« villi are not concenicd in the
abairplion of fat ; otberwife lliere are
o<> dvliniti; fads to prove that the
cubtral cells are c^mci-rnwl in >«re-
li»a only »r that t1>ey may not alworh
BMitcr (iIIht than fat. The gnhld
oells in ihew glands ns in the villi
certaiDty sevrde mucus, and may w
mte also Slime of tlw oocistituenl* of
tbc Biin*us entericDB.
Besides tbme glaude |>roperlT an
called, that u to say involnlioits of Iho
epitbeltal (hypoblasticj mucous mem-
bnuta, there are fouod iu the niHcous
nnnbrane bodies bdoncing to the
lvn)|>hsti(- arstem also oftf-D calleil inlands, vii.. the colitarr glands and
tlie MtRiutoti^l Elanibi or pntcfaes of I'^ycr. We shall spealc of these U
lynpluilir fulticles, and it «ill )>e convenient to itudy them aepaniUly in
ooooortion iiiih tlw lymphatic system.
{SM Imntedistely Mow the pylorus in nan, but varying soroewluit la
[no. iiK
hsntua or cat or ttai ntiu'* ni.uita, raoM
las UvKin DtovcHnt.)
ST-1
THB
poellioii in difivrent animnli^, nre the j^lnDdH of Bninnff {Pig. 119]. Thno
ninv be reKarded as laodKicaiioiis i>f liie pylorit? glaiuls of lliv Momiich. In
cui-)i ^'Imitl u iluci, IiDed with short ci)luiiiiiar (•pitl>ciiuin c-elli leaving a
i]ii>liiir-i luuicti. exteoda atDu^Ie for eume dislnace. and piercing the luuscularis
iiiHiMMu (lividea in the aubinucuua liMtuo into n number of lubee, whirh
Kuliiliviiliug luke n Lwbteil coune and end in alighl entargemeDU or slvmli.
1 lit ci-Ile lining lioth ilie hranching tubes and the ulveoli »n abort eiibieiil
cvIU uiili lUi iiidiftlincl outline, ttimilar L'l, hul in a freah coudition, iu»re dis-
tinctly granular tliiui the cell* iif the guMtric pyloric glands. Uiindlee of
plain musetilnr Ilbrus, ntraggkra from the muacularis oiucueue, are ecaltensd
among iln- tuheti.
Th«wo gitinil* of Briiuner when traced liavk La the flt»Riach are fotrnd
lo pamgrHdiiiilly inlo the i>yli>ri<! rlanda; iruotnl along the int«Mine they
soon di!3it>|>ritr, tlx' dui-la iif thitti: gmiuU which n»u'h intu the du(Mh>iiuni ao
ftr u lo lie ll'iind in c'i>mpiiny with the glnndn ni Lieberkiihn nml villi, vpeu
into the luiuinH <il' the t>iriiir-r.
Il is not clear ihnl uny nppcinl piirpfwe it acrvnH by thcwe glnmU of Brun-
ner; an extract of the gbind« is tniil to digest Rbrin in tho prMcnce of acid.
Thf. Iiarifr Iiilatinf,
^266. The gi*ncra) plau of ^ilriicture of the large intestine is the wme aa
that of thti «niall tut«aiiuc. the aalieiU pninu »!' ilUiincliun being the absence
of villi, and a poouliar arrangeiuciit of the lougitu'linal coaL
Instead of being unUornily dUtributeil a^ a thin layer orer the whole
circumfcnnci' of the tube an m the >ma1l intt^iine, tbc l»ngiliiiiiual ooat ia
in the large inlctine chiefly gatbcreil up iuto three ihidcened handa or
bundles, being vt-ry thin oWwliTC. TliMiri biLn<J«, morodvcr. are eburt«r
than what may be calli-d the natural length of the ink^itine. «» that the
tube ini^tead of hoing us in the small intivtine uf Ikirly iinifurni boro, is
[tuckered up into "eacculi" iQor« or l«ai divided by the three bands into
group!! of three. This sacculated arrangenient anawcrs much the same piir-
pMc tut the arrangement of vnlvulic connivcntes in the smnll intestine. The
circular muacnlar layer is thicker in (lie middle or bellies of the eaeculi
thau at the puckers, where it is very thin.
The villi, as we have Jual said, are wholly absent. Id the lower part of
the i>iimll iiitetiliae lliey become fewer and smaller, and none at all are found
t)er>iml llie ileo-cjecal valve. An increase of surface is provided by longi-
tudinal ridjcei, but these, like the corresponding ruga? of the stoouiob, involve
tho nbole mucous membrane, including part of the aubmucous tissue.
The glaiidii uf Lieberkiibn in the large intestine are in the luaig like thoee
uf the «uall iutefline, but larger and better developed, being both deeper
and broader, and owing to the ahseoce of villi are more easily atudied. The
cells of the gland* have the same diaracters a» in the amall niteatine. except
that th« bvaliue bonier ta rardy preneot ; goblet cella arc, perha|M, more
abundant (hnn cvt'n in the .irnall intenine, especially iu itome animal*. On
tlie ridgi^ lirtwcen (be glandii the eelU become longer ainl thinner, and the
byiiHnc border, frei]iiently striated, mak<« il« B|i[>carancu. The marked
dcvelopmoDt of thou; glandii in the large intO'tino a noteworthy sinca, a* h*c
ahall SCO, nlwirplii-n of mjilrriiil, nnd not tho secretion of digortive juice,
U the chnrarierii'tie W'>rk Dl'llie largt intmtiiie. Il can scarcely be ima^ned
that abs>:>rpli'>n takes place only at the ridges between ibn gfands, and not
by the inmx-iisely larger amount of surmce which is presented by the
iDteriors of all the i;InniU ti<i:ether : but if these glands absorb in tbi; l«rg«
intestine, ifaoy probably act in the samv way in the small intestine.
TUX HtiHCl'LAK UKCIIAN'IHUS uK DlflKSTIO.V.
375
liynntuilic follieta tat abuiMlant in lh<- Inrgc iiitrvtino, t1i>- fjcaiin iiml
Miaaulr tbe spfeDdix TwiDifarniu being cromlcd wiih N>liiiiry folliilcs.
Ttn* iMtirh<v of PoycT arv Hbacnt,
^S67. Tht rrfluM. Anbe agtaniA nrxxtn jwm inlothr n-ctum tlw ibree
bftDtb of ibc lunijitiHliRnI iiiu^eiilHr cmI ■(■rtuul ou( aiii] becoio* onoe nwre >
onifonn Uy«r. and with ihu chnni^ itie Micciiltitioti dMnppcan. Tiis (onxi-
todinal coal is niat>Dii«d to Ibe aous, vhvrv ii coil* uhniptir- T)i« cimibir
euU M it« temitoaliun at tbo anuB is d«Te)o[>ciJ inio n iJiMiDct ring, lh«
islMiwl >|iliiiict«r.
Th« D1III.-UUII iDeiobrane ■■ thruwii inio numcrouH folds or ridga urliicli
bcluoarv longitudinal, bill higbcr up oMii^uc or even iransversc iu dirM-tiou;
to fmtmit the fcrinatlon uf ihete fulds, wtiicb lire oblilcratwl u'li«n Uw re«tuiu
ia fatlT diitvnik'd, tbc riubmucouB tisrae i> more nbundoot and more IoumIjt
dvTvbiprd than in tb« t(*i of the inUsline.
I>a«n ft tbv margin uf tbe iioud tbe uucoiia membrane retaiiu Um cbar-
mdan ftf Um larso iiilwtinv. jilnnda beinjr still pr«aeut ; it iben abrnjrflj pitta
« tk« flpilihuuio diarnrti^rn uf the epiilcrniin. Tlie rv<-luni bw ii »|)ocial
mirvaaa wpplj. but uf tliLt wc ithall >])«uk in ci>iuiecti»n nitb ihv iDOT«neata
of tbm alunenlarjr' mmtl.
I
Tbe Muscular MKCBAimm of DioEffriogt.
f M8, Prom iti etitrance into tbe monib nniil buoIi* retunant of it ns ia
uadigoted leave* tbe bmly, tli« fund ia oontinunlly iiubj«i-ted in inoTomeuts
bartufc for ihnr objwt the triluralioti of tbe food M in Taimtiailion, or jta
mora nuiintvte mixture wttli tlie dige«uve juices, or it» forward prarrefla
tbmvKh tbc alimentary, csna). Tbuw various inorcnicnta uiny briefly be
oaa^£nd in dciail :
Mattintion. Thi* in ninii consists cbi«fly of an up-and-down roovement
of Ibe knrn- jaw, combined in tlvc grinding artion of tho molar teeth, wjih
• evuin antount of latornl forv-at)d->n moratncnt. The lower jaw i* ntiitod
trv mmat of the temporal, masMt^r. and internal ptcrygoiil roiiAcl«t. Tbe
Mgbtweflbn ntdcpnarioo brings into artiuo cbiedy the di^fattrir miiM-le,
tltfMtsh (be mTlo-hyotil and i^eniohyoid probably share in the roattrr. Coo-
tnMtuHi of the etlrrnal {iterygoi4ls pulls fomard the ouiHlyln* and thnuti
1^ lover tenth in front of the upper Contraction of the pterygoiids on one
iidia will aUu ihn^w tbe teelb on to tlie opiiofiile Hide. The lower ItoriiMn tally
plaMd fibrti* of tbe temporal atrve to reirai-t the jaw.
Puriog inaniieation the food b nwred tu ami (W> and rolled aliout by the
navMMOta <>f tba- tongucL Tbeae are aflk;ted by tlte niuvclce of that or^^
gemntA by iIh- liy|xiflawal nerve.
The act of ma«licBliijii u a vuluniarv one, guidnl, b» ari- so mnnv volun-
tary a«-l«. not only bv muMnilar aeiiw, but aUo by cifutact MnsationA. Tbo
iDtMiir fibres nf tbr tit\h cranial iien.'C convey rootor impulaca from the braifi
to tbe above-men I ionod muiwlca ; but mrolyiui of tbe wnsorr fibre* of Ibe
■Mna aenre reiMler< nuutioilion difficult by depriving the will of tbe aid of
llw usual aeDaalMHu.
jS89. Dtylulilioit. The fnod when sufBnenlly mariicated is, by tbe
mavnoents »f th« longriK-, gtalhi^ml up into a bolus on iJw middle of (he
DpfM-r aurfiice of that cir;[aH. Thf IVunt of the tongue being raiwd— partly
hy it» iniHnMi' muscles and nnrllv by tlie wtylo-gl'wwe 'the bolus is tbrutt
hack betwprn the lon)C<ie and ihi- |inlat« through tbe anterior pillar* of the
liUKea or isthmus laut-ium. linriiifli.iidr before il nrrirot there the soft
b raised by the lovauir patali. and m brought lo touch tlie pusleriur
87lf THK T18SUKS A N H M KC II A X 18 Jt8 OT DICIKSTION.
wall or the ph.iTjrix, which, by t)ie contraction or the iifi[<^r niiir{(iD of iht
superior conetriclorof the phiirynx. buI)-CH somewhat torua'tl. l1i«elevittioo
of iheaoft palate CAUSM a (liBiiiict riae of preaaure in lJi« luwal cliumbort ;
tliitt can be Bhovfn bv tiilruduciug a tviiier niaDomecer iuto ooe imttril ntid
oluaing the other just orovioua tn swallowing- By the conlrai-licm of the
palato-plisriTDfceal muscles which lie in ilie pimieriur |nllan of the fniin« the
curved edges of those pillan are muilv mraiithl. aud thun lend tn aic«t in
the middle line, the small Kap hetween llit^rii hein^ llllcd np hr thr uvida.
ThrDiich tlit-it! ninno-vrnt tlie eutrnncc Intu the iii>«tm<>r nnrec is hlttckcd,
while lIic ti'ift pulnte if fiirmvd into a .iloping rour, Kuidiiig tbo boltu down
the phnrvux. By the cuiilrutniuu of the ntylo-pluirrDgaiu uid palato*
phiu-vngriiA>. the runn''l-i'hnpi.-<l hag iif ihc pharynx u Drought up to meat
the (l»c<'niltng nion*ct, very much u-* n glove may be drawn up over the
finger.
MeaDW'hilc in the Inrrnx. a» shown by tlie InryngMwpe, tJie arirUowd
cartila|,>es and vocid corus are approximated, the latter being also reiaed ao
tJial they conic very near to the lalse vocal corde ; and the cushion at tiie
base of the epiglottis covers the linin glottidis, while the emglottis itself is
depretaed over the larynx. The tliyri>id cartilai,-* is now, Dv the Bctioii of
tJw larA'ngeal luiisclee. tiuddeuly raised up behind the hyml liooe, and thus
aaiiatN th<f eiiigluttls li> cover the (;loltia. Thia uiuvemeul of the thymid cud
easily be felt on the outside. Tlius, bulh the entmnoe into the [luyturior
uiirtw and that into the larynx bvini; chuted, tlie impnlw given to thi! boln«
by the tongue nui tiave no other ctl'iH't than to jirojiel it bc-ni-ulh tht sloping
wd palnle over the iticliue formed hv thv nitit of llu- tou^uu and the epigluUi*.
The pHlal» ghuni or cimittrir-toren wtbmi fauciiim, which lie m the anterior
pillnrv of tho fiiuctu by contracting clunt tJiC door behind the food wbicb liM
p«»cd them.
When the bolus of food ia large it in received by the middlo and lower
conslrictors of the pharynx, which, contracting in eequrnce from above
downwar'l, Ihniftt it into the (ceophngus, along which it is driven by « similar
lerie* of succi?eaive couiraclions which we shall speak of iiuiuetlialelT as
periylaltic action. Tlii; eoiiiparalively slow deec(<nt i>f the food from lh«
phnryux into the stomach may he readily ^een if animala with lonf; necka,
anrh as hor«a 8n<l dogw, bo lAati-hed while swallouing. When, however, the
DMmi'l is nut large, or when tlii? .xuliBianee swallnued ia Uijuid. the movement
of the txick part of the tongue may be sufficient not merely to iutrixluoe ibe
food intx the gra*p of the couslrictora of the pharynx, hut even lo pru|)el it
rapidly, to »h<3ot it in fact, along the tax naopfaagua before the mn^cUv »l that
organ have time to omtniet. In Much a mode of swallowing the middlu and
lower conntrictort take little or uo part in driving the fo<i<T uiinard. lh»ii>.'Ii
they and the (rwphnguK upiinir to cuiitraot from atHive donuwiird at\cr the
food has pimcd bv them, as if to ci>mplcle tbti act nuit lo iniinre that nothing
haa beeo left bchmd. t><-glutitinn in tht» favhion still rcmninn powihle after
tbeae oonalrictore have heconw paralyzed by lectinn of thvir motor nervtw.
When a second act of degliilitioii tuccetMl^ n fint with iiiilltcicnt mpiditjr,
the nervous changes which itnrt the phnrynL'ral mi>rcmviits of the fccontl
Upct appear to inhibit iho teeonhngenl movcmenlji of the lirel act; and whoa
Birallon ing is repeated rapidly several times in lucccwion, tho taaophagua
reniaius ijuiet and Ian during the whole time until immediately after tbo Uist
•wallow, when a iierii>tallic movement cl'Kfcs the series.
Wlien the atethuM'0|K- is applied over the cesophagus, at diflbreat regioua,
a MWind li lieanl during deglutition ; situeiinies two sounds are heard. The
flnt and motit conaiant in coincident with the pauageof tiie bolus. and is due
to Ihi* and to lliv mtisculur <wuud uf the oontraetiug inuAolea. TIte later and
TUi: UVBVVUAtt MKCIIAXISMii Of l>H)K8T10N.
377
coiMUnt KKiad appi-ara to be caused hy a (juatitity of aii^bubbla with
>b>ch tk* bolua waa Miuigled ludgod at the cardiuc etid of lh« meophagua,
: Ibroad tii(n tlte stomach by the se<|ueul periBtaltic coDlrnctiuu of the
I
I
It will ht 4*en fntm what haa beoii suiil that ilvglulltiuo. though a ron-
tlniMufl act. tiiiit' 1>e re|[ar<kil nn divided into thnw sitiga. : lUw tint &ta^ is
tha libnulinff of the fiwd through the iathiuiiH fatinum ; this may bo either
of Ionic w Nwrt duratioo. Tlie Mcood tinjje ).i the paMage through the
up|vf [lan uf thv iiharyux. H«rv ibo fuuil triiviTMv ii region cuinmon hnlh
la the loud anal to ret|nratiua, and ia coiiMijuaico thv movotntMit in a* riipid
aa Bonible. The tliin) Htage U the detoeot through the gnup of the con-
■UMOca. Here the food hiw pataed iJie rapiratory oriSce, aiul in oudm-
qaeaoa ita punge sgoin beconua wrapamttvely mIow, except in oud of fluid*
wad anall non^. wh«n, » ire hnre fetn, it may continue to be mpid. The
BMM* along the owopfaagu* may. pcrb>|i«. be regarded na contUluting a
■mrtb ttBfa; but it will he more citnvcfliieot to coosidor tin- nMophag«al
mowmaim by thmiKlv**.
The Gnt atoge in thia complicated procom it undoubtL-illy a Tplunlnri' act.
Tb* raMM of Ibo soft palate and the approximation of tlic |>otilpiior pillara
aB>j aim fw at tiniea voluntary, since they have been teoo, in a ave where
iIm pharynx was laid ban- by an operaliou, to take place before the Iboil
had iiiat'ti«d three parl«; but the movement may take place without any
rjii^'iic of the will and in the aWnce of consciousneaa. Indec<l. (he second
atu«, laken as a whok-. thmi-^h some of the earlier componeul movemcnia arc,
■•It were, on the liurdcrland lietween the voluntary and involuntary kingdoms,
muat be re^rded as a r^ex act. The iliird aii'i last stage, whatever be the
ouct fimn which it lakea, is undoubiedly reflex; the trill liaa no power
whatever over it, and can neither originate, stop, nor mwlify it.
Drchililion in fact n> a nholu is a rcAox act ; it cannot take place UDlen
■(MB« Mtniulu* 1m> nnplivd to the mucou* niembraiw uf llic faiicn. Wlioo *•
V«luiitaiT bring aoout fwnllowiiig movcjacnta with tliC mouth cm]rty, ire
Mpplv lae iMoeaMry atimulun by forcing irith the tnngue a »inall ijuautity
of MU<ra into tlie faurm. or by touching; iIh- lau«.-ca nilh the Inniine itwlf.
la the redex act of deglutillun. cauKil in thv< onlinary way by the ftMxl
eowittf in contact nilh ifie fauces, the Rtfercnt impuiHa originalod in the
6mmh u« ctrried up to the nervous c<-nlre by the gloeBo-pharengcal nerve,
bjr braacfam of the fifth, and by the pharyngeal bnndies of the nipcrtor
Iwjugcal divUion of the vagus. The latter seem of special importance,
ifawe ibe act of swallowing. i|uile apart fr<im the presence of food in the
nonth, may be brought dui by i-t^ntrifwlal alimulation of the (superior laryn-
fltmi nerve. The elTerent impulaee d^-end Iht' hypov'loMal lu (be musclea
of ibe tongue, anil pau down the glwau-pliaryiigeui, the va;^iis throii^h the
pb«ryti):v«l plexus, the fifth, and the -iiiinal aooeaory. to ihi- mitM-les of the
nima ami pharynx ; their exact patlu being as ye< not fully known, and
erubably rari'ing in diflerent animals. The laryn^al musclea are governed
J iha larragcaJ brancbca of the vagii*.
Hm OMitre of the ruHex iK't itm in the medulla oblongata. Degliitilion
BW be exdted. by tickling the fnucm. in nn animal rendere<] uncoiucioiin by
raowival of thv brain; provided the medutla be left. If the mt^dulla l>e de>
Mnijed, deglutition is mi|XNMible. 'I he centre for deiHutitiim lit* higher up
ibaa that of in>|>inU»on, so thai in the disMMe* or iojuricd involving lh« upper
|«rt nf ihr UHiiuIIn oldongatn, thi- former act may be impaiml or rendered
■ mp<«Mble whiU* the hitter reuiiiins untoiichot. It ha* lioen wkid to furm
part of the ■uperior olivnrr ImhIIiv, tail thi< vi^-w in luueil on anatomical
>niU uuly. We shall have to dvnl with tliui and tiuiilar matien in treat-
378
THE TlSSrKE
inp "f tlm iviitrnl ii«rvout lyKlcin. Il i* |>ri>biiblu tliat, U ut the caHi in W
luiiDV nilicT rt-lV-x ii<;u, the whole niovcnieiit cun bo caSld forth l»y MlimuU
RRccling ihr crntrv iliirctly . iiiid ii'il tiding on lh« u«ub1 altercDl oorw*.
§270. J'rrii'liiUi^ mif-mruln. Piitliiig iwiilff the Komewluit complicated
plinn-ri^oAl pnrt of Hcslntitioti, Hnd Itikini; ihc awipliugcnl mi>vi.-inr!nU by
tliemsclvra wo fiiti) lh»t thcso, tojpthor ivitii the luovemeni* of the etomftcb.
and of the small luiil Inr^o intwUnCB ri^bt down to th« anus »n inoro or Itm
alike, and mar be deHcribed under tlie fteiienil nnnie of " [tehstaltic " mave-
neota. We liave already in § 92. epokeu of th»e, biit it may be well to
oonaidcr tljeiu briefly at^iii under a ^neral aspecl, before dwelliDg uo ths
Bpecial niiivetiienis of the severul purla of the alini«uury canal.
The Riii8(-ular coat of the allmonlary t-aiiul cdu^U, aa we have seen, of
two layers, separated more or leai diHtiiicily by a .ibe«t of ciBnectiTe tiane,
ui nuu-.T ihiiijuT limt^itudiiml layer, and uii inner tliicker circular layer; nod
aninibtr iirrun^c-mi^nt uliiuinn jti nearly all ihc luuiieular hollow tul)i« of tbe
body, esocTit llie arteritr*, in whirh thu niumudur (•k-mvnl^ are jirtwiit uot m
much for the piir[Mifli- of driving' ihf blciutl onward il> for the sake of rCK^iU-
tin^ the irrii^aliun.
Thu Aclii'ii of the circtulnr coAt it fairlv itiniplo. A c-oninclion ularting at
any jinrt, travels onward in (he Niini.< dirivlion, generally dnwnwaril, thsJ it
III nay. from a part nearer Ihc numlh [o n part nwin-r the roettim, for ■
greater or lew di»t»ncp, the eircularly dispomnl biindk« ciMitractine in
tequvntx. Tho result i* a narrowing or constriction of the liibc whidi,
travclliog more or tcsa slowly along the tubu, drives the contcnl« onward ;
when a butcher emplie« tho iiiicvliiie of a slaughtered animal by eciueexing
it high up with hin hnnd or with his thumb and finger, and carrying the
ec]ne«King action downward along the length of the inteatine, he makes tbe
passive intestine do very much what the circular coal doea actively, by cod-
traclton, in the liviuK aiiimal.
The ai^tion of the loni^iltidinal ooat ia [terimp* not to clear ; but a contmc-
lion of the luui^itudiuul coat takin); pluiv iu iinv negnienl of th« tube would
tend to draw Ihe tube over tho contenta lyiiiti immediately abovf, or botow.
ibe segment, vnry luucli i\» a ^dovc in drawn over u finder. And a suoeemion
of aucii ciintni(-lic>n» travelling alonj: lh<; tubu would lead to a movcmeHt of
thec»nti>i]|.'>io till' Mine direction. WerethecireiiUrcoat alw-nta liingiludimtl
ewat might by it»elf jioMibly suliio' to jiniiwl the enniente along the tube.
In the proflcnoo of tbe circular coat, tbe action of the longitudinal coat Id
any wginNit of the tube, if taking place immediately before the circular
ooalmclion, would, by Ulliug the segment with contents^ render the squeezing
action of the cimilar coat more emcicnt; if taking place imoicdiately after
the circular contnu;tion, it would help in quickening the return of tbe tube
lo its normal enlibre, for tbe contraction of Lhe longitudinal ooot tends to
■borten and widen the nement, and ihna would pr«|iare it for new oonteoU.
We can hardly imagine ttiat the two coals would contract at the same time,
■anco they would lend to neutralise eaeh other's action. Indeed, we may
(irobahly go farther and aftjunie that In each j^eguieni of the oanat first tbe
ongitudinal cuat ooniracts while the circular e'lut i« relaxed, and that ihea
llie circular coat eontraeid while tbe longitudinal rulaxiw. Wli«n we come
to deal with rc«jiirattuu we shall meet with a MUiilar double anlagvmiftic and
MCManve action belweeti iiis|>)ntt.»ry and exniratiiry muxclc*; we tihall
fbrtlwrsee reason to think that the proccwioa which >iart the expiratory act
tend to check or inhilni tin' in^pir-iiiirr act and vitt vertm ; ami very posubly
a like aee-Miw of stimulation and inhibition obtains in tbe muscles of the
alimentary canal.
Il must be rem«m1)crcd that Uic drcular coal ia alwaja maoh thicker
TIIK MUSCrl-AH MKi-HANISMS UP U10K8TIM.V.
:m
ihu (bv laniptiKliDal cost : aixl we tuny infer tbitt wliile lii« chief work of
(triiiitf tbe oontcnU onward falU on itic; fumier, thi: lailL-r ntuhiB ihv work,
ttlbrrm Uk way ttlik-)i ut have stifcxiatlcil or id wine otlicr way.
ImhoamaU int(«<tirM- (he iul« iiiliuii^ looiicly anrl much Inisteil. ^< that
■Ufkiopa arc formed; tli« roDl^uU, moreover, are liirgvly tlniil. Hvuce,
iktmwlj uDWanl nxivciiicnl, ini(.-li a* h aoeii wWii more ai lid conuniii pi\M.
atom lilt Mmighl and uititvwlini lirmly nlCncln-il 'irMi|>liiigiu(, is ean)[>ti<'nt<H]
bf avMBUtnUi dao to ■ loop lioiag proj^'cCcd forwdTil by (lie entrance of fluid
fii« a)iuT«, or tw-JDi; driiggt'd dnirn by rhr wtight of it* tivw <miiIphU, or.
a ihp ndirr hand, duo to a loop being rrlntct^il by thr driving onwnnl nf
iu niDttatm and iIh- fmntying of itwir, ami tlw! like. Tn iliiit way a prtruliur
vrilliing UHiTrmeiil <■<* the bowrl n hronglit about, and thv pliniM " pcristultio
■nimirnt " is gfn<'rally uwi) to dcTiolv thia total otfcet of the (SHilroetioli
oftlw miiKTular chiI»: it will, however, be b4M lo r«Mrtet the raeani»gto
Ifca tm^ rmirv cnntration of lh« circular ooat aaiiBtnl, in mmt ctwH, by n
nnilar prograaivo contraction of the longilixlinul coat.
( 271. }tortmeaU of tlio tttophagtu. I'haii.'. n>t we have just said, are
fclrlj uniple. Th« cir<'idar conlraetinn hogitn by the constrictor* of iho
phvyns is codIidiiinI along the circular iiial of the a«o})hagii«, and iisiisted
DT M) accompanying cMilraction of the longitudinal coat, the direction
bosir always, lave in the abnormal action of vomiting, from above dowo-
wanl.
It will be remembered ($ 'ti.2) thai the luuaoular bundles of the ceeophagua
ai« nwBpased of HtHat«d 6bres in the up|)er purl, iind of plain uiisirLated
flbreoiIlR in the low«r part, the tran^iLion occupying a difllTeni level in
<lilTrn-ni animalA. Nevertht^lcis, ai* far a^ the [leHataltiu woveraeot is uiu-
cenxd, tbe two kimb of <ihri» behave in the ttame way, except that the
periaultie wave, if we m«j bo call it, travel* more rapidly in the Mriated
Thaw peristaltic movemcut* of th« a«'>phBgii« may. like thoM of the in-
llina, be aMn afUr removal of the organ from tbe body ; and, indeed, may
oootiaiM to appear u|Km (timiilniion. tor an iinuiuiil IcngtJi of lima. They
tmj, iWabre, be carried nut by the mufrulnr dements, witli or witlKHit the
balpof tfat ncnroua olemcni* embeihteil in thcni. apart fnim any action of
tb* MSlnl SMTOna ^rtem. N«vcrthel«H, in the living bo<ly, the movemcnta
«f the naopliagai anm lo )w tn a tpcciul way dependent on tbe cciilral nor-
vmiaflTat»n; use contnicltont are not started niid carried out bv the iralU
af Ibe tube alone, and so transmitle'l from section lo sfwtion in tbo walls of
tbe Idbe itself; but aHercnt impulses starte<l in the nliarynx, and paseing tn
tbe nieilalla oblongata, give liso to reHex etlereut impulses which decceod
■loQK nemtu tneta to auooewive poriions <}f the organ. If the ti^ophagus
be cut acrui wine way down, or if a portiim of the middle region be excised,
atimnbtion of the pharynx will pr^xluce a iterialaliic ooniraetion, which
tra*«nipg downward will not Miopalche''Ulor excuion. but will be continued
on into tbe lower diaeonneeted jxirtiou by meana of the central nervoue
vpUm. And it is stated that onliuary j>enMaltic conlractioDH of the lover
part of tbe owiiihagu* can be reulily excited by Ktimulatiou of the pliarynx,
bat not by ntimuli applied to its own mucout mcinbrane. In the reflex act
which thiM briogfl MMUt the periatallic contraction of the (Mophariu the
alTi-rvnt nervm are thooe of the phar>-nx. viz., the Nujxiriar lari-iiKeat nerve
anil pharyngi-al branchex of the vagus, brnnchis of the fil\h. and in lome
•aimaU, at Iceat, brancbe*of ibc gludBuvpharyngml. but childly tlip ttrtt ; and
OBOpbageal mofeinenui can cu.'uly be cxciml by centHjHtal •limulation of (he
anperiur laryngeal. The oenire li«t in tbe ninlulla oblongata, iHiiiig a part
of tbe general dcgliititioD centre; and elTervnt iiupiiUif pa*« abmg librv^i of
380
THK T[S!fL'R9 AM' MRCIIA.VISUS OF DingSTIOX.
the vagus, raacbiii;; the ujiper (mrl nf the cuophngUA I>t the recurreat 1
geal nervei. and tlie lower (inn. rhrcmgb tb« aetophngealplcxuiia of tbo vagus
iFia. 70). Section of the trunk of thr vngiis n-n'lore ilitBciiIl thf pawage
of iOD(] along the ONtophiiguH, imi] stiiiiiilution nf lh<: poripbcnil «tiiinp caiisu
ffisophaEt^nl coin met inns.
'i*)e liitct cif ihix inovemiiiil In ihc (Mophiigiis U considtrriible : thua iu ihe
dog K ball pulliug liy iiu-uns of n inillcy iigitinal » weight of 'JoO gntamm
hiu bccii friunil bi ]w reudily cnrricii ilowii from the phiirrnx to the Momaeti.
At th« juitucioii of the OMOphiigtu with the itomach the circular fibres
usitally Kmniti in n more or a less norniaDcnt coadtliou of tonic or obscurely
rhythmic <^>nLriiclioii. more particularly when the stomach is full of foot), and
tbtis M;rvc as n ephinctor to prcveut the ivturn of food from the stomach into
tho a-sophagiiK. Upon the arrival of the bolus of food at the end of ifae
CMOpbagiis, thv centra for Ibis sphlDcter is inhibited aud the orifiM is thus
opwad up. Possibly the patency of the orifice is still further secured by a
cnntraetion of [he longitudinal muscular 6bres which radiate from tl>e eud
cf the CMophatn>s over the stomuch.
§872. .VovemciiU of flic stomach. Wlillo the obje<t 'if the (noohagesl
movement is simply tu carry tlie swallowed bolus wjlh all iluc a(>een to ttMi
stomach, and nhile the iutaitinni tnovemeiit has. in like muiiuer, simply to
carry the Jutt«tinal eontenU onwiinl, the twistLvl courw uf thi- li>uiit;(j pmb
ensuring all thu mixiufcof thr c<>n^>tiliii-iit3 of the; ontt-nt* ihnl may ui; nucca-
Gary, the niovvmunln ai thu »t<)inach lmv<^ n double nlijvi't : on the one haixl
to pivividv an ad«'<piati- rxpiMure of tbt- ('huNtiI* nf ihc ililntrd chamiii-r to
the iulluenct' i>f thi- giiMric juice, nnil <m ibi other (n propel the panialljr
digeuttil food, when ready, into the diindcnuni, Wc may, accordinglr. di»-
tinguiih between what we may call the "churning" and tlio "propuUive"
niovvniciits of the etomoch.
When the stomach b empty all tlie muscular fibres as vre have said, loufti-
ttidiniil. circular, aud obli<jue, fall iiitc a condiiioo which we may iierhapa
speak of as an obscure tome contraction. The whole siooiach is sioaU and
eotitracied, its cavity in nearly obliterated, and the muooua membrane, owing
to the predominanet' of the circular c»hi, is like the lining membrane of an
empty artery, thrown into longiiudiual MAs, As more and more food enten
the stomach all tho enam bucomu nlaxed, with tlie exceptiofi of the pyl.ino
aphincler, which remninn at lin>l [wrmaneutly closed, and the lew marked
dudiac sphiocler, which nn'rely rtilnxiat from lime to time at eucli act of
swallowing. Xo sooner, bowrvrr. do the coats thus become relaxed ihnn
iher M.-t un olwcnre rhythmieiil [lerinlaltic conlnictioni, givinv; rii>e to the
"churning ' movemenl«. Thc»e ninTemtint:< have liccn dcacribral m of such a
kind that the eonienl# flow in h main current from the cnrdia along the greater
curvature to the pylorus and back to the canlia alone the lemor curvature,
•ilbndiary currents mixing the peripheral portions of the contiinl^ with the
atttn («ntral ; it may be doubted, however, whether luty such regularity uf
flow i« marked or constant, and it is not cany to see by what ooiDbination
and (c<iuence of contractions in tho three coats, tongitudinat. circular, and
obliouc. such a regular flow caii be produood. But in any cniH', by such
rhyltimical ooniniciious the food aiid gastric juice arc rolled nttout and
mixeil to]*ether. Tlie^ churoiuK movements are li:t)ble at first, even though
tJbe stomach be tilled aud distended b^ a lar};e menl rapidly eaten; tfaey
becooie more and more prunouuoed as digestion proceeds.
Before dignttion has proceedetl very far the " propulsive " movomenta
begin. Thiiw occur nt inieri,-Hls, and are repeated at lir>iC slowly but aller-
irnnl more rapidly. ICach movement ooiisisis in a conlractioa of the circular
Tim JIC8CUI.AR UBCHAMSUS *>¥ HIOKSTIUN.
381
aMNmltf 6brw mora powerful than auy taVIng y^tt in l)ie dmmtitg move-
WWWi, uti UaAiug to m ctroiiltir ooiistriciitin wmoli, bo;{iuniug nnpareiillr at
■lMiatllioobteurel^<I«fi»«d gnxivonhich tiiark^ t\te iteji'mmti)! or iheatilmin
pylori, Ifmwb down toward the iiylorii*, pmjx-lliD^t ihr fnoil utiwanl. Tlitx
toiDVQBtnl tf act'omiiniiM or nitticr prvixitiril l>y n rulaxiiliuti <if, ihiil \» to
mf, in all |ir>ihal>ilitr an inhibition uf tho [H'riniiai'iil n>olriii:tiiin (if, tho
iMaaUr pjhm ilMilt*. in ontir tlial Ihc gulric conU-nU nmy piut* into ibo
•MiMWtD. Itiit the urcurrfiici- of this rolnxiuion it (ldi;rin:ntvl by tho
OMare of tb* gastrin: contrnc* : Tir if tW [iropuUivo moTelnl^nt <lrirc:i Iafg«
aafawltd |>i«cn tovunl the pylnniB, ilio >]ihin(rt«r i« apt lu clow a^in, the
MbR of wlii*-Ii U that tho utidigntcd tnonwis are carried back intg the
am bodj of the alonuurh.
Tba oorabiiiei) eflect then of the churniog iim) of the propulsive move-
MMMa b, after n certain pari of the in«al htu lieen r«diice>l to n thick Buid
oamUlioa aomcwhat rtaemblint; pea soup and often called ohyrne, to strain
rtfflhb n>are fluid |Hirt into the duodenum, and to submit Ibe remaJDing still
••tlid piooe* to the further acliuu of the Ktulrie juice.
Am iligatioil proceed), more and ni'ire material leaves the stomach, which
» ibiM gradiuUIr emptied, the last portion!) whicb ure ciirri«<l thn>u^-li beiu^
tbom part* of ibe fooij which are leant ilij^tible, ainl any wholly indij^lt-
bW forvign bivliiw which happen to have been swallowed ; the IntiiT may
pwrh»|i* never U-jivc tbo atomacb at all. The ]>reaeDce of food lead* to tho
dvTcl<ipn«il of the moTetnenta; hut evidently it is nut the mere mechanical
ri-|ili'ti4in of tbe orgw) which i* the cbum of the movementa. since ih«' stomach
.» fiilltal at the banning when the movemcnta arc slight. an<l becomes
oa ibvy grow more forcihlf. The one tbin^ which dors iiicrcoao
•owu with the moremenla is the a(-i<li[y, which i» at n minimum wheo
tbe ({^nerallv alkalino) IVxxl has been fiwallowcd, and increnMS sl4.-n<lily
(ttiward. It baa not, bowever, been definitely shown that the increaaiii|[
acidity is Ibe efficient stimulue, civin^ rise to the movements.
Tbe mtireiDenta of even ii Kill stomach are said to cease during slecjt.
The ttervoas mechanism of tbe f^triu movements had belter he L-ontiidero)
ID coaneetiun with that of the iutrntinal movements.
( S73. lomiViNjp. lu a conscioiu individual this act is preceded bv leclliigi
of tiait*>«, durinif whivh a copious How of saliva into the mouth talea place.
This beio)£ swallonnl carriea down with it a certain quantilv of air, the
pMHOOe of which in ili<- Hioniach. by OMsistintr in (he ojienin); ur tbe cardiac
tpWfltff- Mihscfiurotly fui-ilitatcs tht- diHcbarjce of the ifiulric CKinti-nl*. Tbe
tMOMa ia geDenulv luooeedcd at 6nt hr iiH-tlectnal retcbinj in which a ilcop
ipaplfllQiTT ellbrt U made, ao that the diaiihraghni is ihruKl down a* Inn- ns
pfmtPrfff agaiaat the stomach, the lower nlu Inking at the hmuk time fiircibly
4nvn in: since durinj; this inufHRitor^' cD'ort. tbe gludi* ii> kept chwed. no
•iron enter into the Iuuk*; but some i^ dniuu into tbe pharynx, and thence
prahnbly deeeeods by n ^uallonini: action into [he »t<imiich. U'hcn retching
Canon to actual vomiting.' (his in^pirauir)' cflnrt i« nun-crdnt bv a sudden vio-
(•xpiratar>'oontructiou oft Ik abdominal walls.tbc glottis slill being closet],
•o that the whole forcn of the cflorl is spent, as wo shall see it is in deiecaiion,
in preMure on tho abdominal contents. Tho stomach is, (hcrefbro, forcibly
eonprsaaed from without. At tbe same time, or rather immodialely before
Iht ei]dniIory HTorl. by a conlmctioa of its longitudinal libres tbe oMoiiluifTiis
is sborK-nMi and the cardiac orifice of the stoimich brought cloec under Ihe
diaphraj^m, while apparently by an inhibition of the circular sphincter,
■idtsl iNrrbapa by a coiilracliun uf the fibreti which rwliate from the cod uf
tb* OBOphagn orer the ttomach, tlie canliac onlicc, which is normally cloeed.
Is ■BBM'hat nddaoly dilated. This diluliuu opens the way fur tbe cuutenta
TU£ TISSL'BS AX1> MKCUAMSIIS Of DICBSTIOX.
of the Btomnch. wliicfa, prewed upon 1>r tbe conlractlon of the BtKlometi
Ui n i-erlnin but jirubuliK' only to ii aIi^Iii extent by llie c»iitr>cti»u i>f tba
gastric walls, iire driven forcibly ii|i iIm: a*)phagu*. Tb« nioutli b<Hng
widely ojjt-n, aud the iicck Mrvtcbeil lu afl'unl lu Mnuglit « coune a* (Kwiblu.
the vi>init i.-> rjvt'tt^l IViiiii ibc body. At thi» monieot there Ik mi tiddilioii«l
cxiiinitticy cllort. wbicb nvrvcn to pnivi-iit tbi' vomit pHsnng iat4> ibo larynx.
In uuMit iiiMa t<)u tlir jiiixti-rinr pillnn> iif ibi; liiucw urv upproxitnatt^t, in
onb'r to cln«! Ihr niwal piii^n^ ngniiiHt llio n«c<rnding rtnwiD. TbU, how-
ever, in n:vvn: vomiting is Trcijucntlv inclH-ctnttl,
TbiiK ill vomtiing [hero are two ilialini-t ncU: the dilation of iIm citrdinc
ortti<-v und ibo rxiriiisic pr<«UTv »t' ihr nbitoniiiial n-nlU in iin expiratory
cBbn. Withoiil ibf furmt-r iIk- Imlcr. even wrhrn dislrcMtngly vigoruUB, tt
invflbctual. Without Uu' bitter, u in nmri poisoning, the inirinsiT tuov«-
iuen1« of the eloniiicb it<clf nro mrcly otbcicnl to do ntoro than i^eci gas,
and, it luny be, n very sniiill <|iiantily of tbod or fluid. I'ymeie or naier-
brasb i«, howevAT, probubly brought about by this intriiifiic action of the
alMDiteh.
During vomiting the pylorus is gonemllr dosed, bo that but little material
eccapea into the diiodenuiQ. Wben the gall-bladder is lull, a copious How of
bile into the diKxIeuuiri aeci'iuipanieH the act of voioiting. I'art of this may
lin<l its way into the Btoiuucti. lu in bilious vomiting, ibe pylorus then having
evidently beL'u ogiened,
Xbe uervuuii uiecbaniaiu of vomiting is eunrjilicsied and in niauy asjiects
obecure. The dferent iiniiulsE's wbich ennite the expirniory etTurt umat oome
from Ibe respiratory centre in the lui'dulbi : with these we shall deal in speak-
ing of rcHpinitioii. The dilation of the eanliae orJIiee h eau^. in inrt at
at leott, bv inipuNts descending the vagi, sini-e wben tbcDe are cut real vomit-
ing wilb lii.irlmrge of the gitstrie eunients, if it talteo plai-v ut all. lieeoia«a
dLmeull tlimiigb want of rrailinnv in tlie dilution. Such inlriiiKit: n>ov«-
menlM of the •tomach w do tnlie |>lKec, and the inoveJnoilH of the u^ipha^s
■]ip<!tir lo he rarriwl out by the usnul uervw. The i-iren.'nl impiilMi* which
cause tlie flow of wiliva in ihe introiluelorv nauwa alio doecnd nUiiig the
u»iiid nerves siieh a» the chorda tynipnui. ^hetc various icapulxcs ntny l>eat
Ih- i-ouNderei) sh eltirllng from a vonuliiig ctmtrc in t^c tnudulla, liaving ctuae
relations wilh the resi>irntorv centre. Tliii' cenlre may bo oxcitod, may bo
Ibronn into action, in a reflex manner, by «timnU applietl to {teripberal
nerves, as when Toraiting is induced by tickling the fiLiiccs. or by irrilutioD
of the gnsttic membrane, or by obstruction of the interline due to ligature,
bemia, etc. That the vomiting in the la«t inetimco is due to nervous action,
bihI not to any regurgitation of the iniextinal coulcnls. lA shown by the fact
that it will take place wben tlie iutestine is perfectly empty and may be pre-
vented by secti'in of the meaenterio ncrve«. The vomiting attending reual
and biliary t-alculi is apparently also reflex in origin. \ oniiting in fact
u rule i^ a rvHt-'-v nclion. ihe atlt^rent impolses passing along one or oti
nerves, but muet fre'iuenlly along th(«e connected with the alimentary caoal.
that is, long aflerent fibn-s running in ilie vagus or in the splanchnic nerves.
Tim centre, however, may be iillt-cieil directly, as probably in the eases of
some poisoBS, and in some innlmice!) of voniitiug from disease of the medulla
DUoiig*t& Ltutly. it may be thrown into action by impulses reaching it
fironi porta of the brain higher up than itself, as in coses of vomiting pro-
duced by siucllii, tostw, or emotions, or by the retwilection of past eventa,
and in some cans of vomiting due to cerebral disease.
Many emctica, such us tarinr cmutic, appear to act directly on the wntro,
siuce, introduced inb> the blood, they will prt)du<« vomiting after a hlnddor
ba« tM:en sulMtitiited for the whole utomneh. Olben again, such iis mustard
m
TDK Mt'SCt'LAB MBCUaNISMA Or UIGESTION.
■Ml wmln. act In > reflex maoDer bj- irritation of the B:ii*trio uui'uns mem-
branr. With ulWn, aj^iu. vjIikIi niu>« vomiting by ■Ivvi.-lrijniig u iinii»euuB
tsM«, lb# MFtion tDVolvts |mrU nf tW brmtii higher tliim ibo crntrr ileclT.
% 8T4. ihtt^fiiU »/ (Ac tnuiU inlr^inr. 1 tivw.', ne w have nlmidy said,
ktv thr lfpi<«1 p«TUialtio iiiQVcaMrnt*, nimpto except in s» tar lu they ar«
evniplicsiiil by tbo niatvoc* of tbo iiemicnl Imipn. Ihc [>cciiliar oecilluiitig
■M*Miiiriitit nCnhii-li appear to he proclucod cliiclly by che hnf^tudinuJ fibrea.
TW prri)>iallif m<ivi'iiivnt», n« a, rulv, take phoc fium nhovf^ downwnrd,
anJ K nve liegintiing at llie pylorus may he Iraeoil n Ion;; vrny ilunu. Kut
eantisniui* uiny, aiiJ in all proliabilily occiu^ioitally <lo, begin at vuriou*
puiate aloog tlw lengtli nf the inteilicie. A movement Glaned liy arliticial
MtnulatioD aom« way dunn the iulesline, may travel not oulv ilonuwani but
•bti ii|i«ar<l : it hat been disputed, however, nholber in the living body any
iHlural haekwanl (teriataltic n>oveni«Dt really lakes plaee. In thu living
budy tlw tniNtinee have i>eriodB uf rest, alternating with periodn i>f activity.
tW oixurreuce of ih« pertoda depending ou various circuuutauve* ; tlio
■Mcnrity nf tbe inoveinenta uldo varies very con»i(lcnib]y.
iS79- MttnUftU of Ihe l-irife intetliiie. Tliew." an* fundu men tally the
MBM >a ihoM of tlto •atall tutcalinf. but distiin-t in wi far il-< thr hitler ceuM
•I tl)« ilvn-opcal valve, at nhieli itjxit tb« Ibrmer Doruially bceio; titty arv
Mmp[«r, ioa*niuoh ■» thu jiendeiit hKifM are abcent, and not m> vigorotia, in'oeb
rvlalivrly ti tl>r dimiiclvr of the tub<-, tlio ninount nf muwciilnr fibre is Icat.
Along tbif itikifi *\\rrv the Mix-ul! ar*- well ilevelopwi (§ 266) the mtivrRient
nwjr prrhapa br deM-riU-d iim aliuoat inlermillrnt Irom wccutuf lu Mccuhis,
tlw cooteiiU of unr NUTuhie In-iiig driven hv the pc<ri«tatli(r eoninieliotis of
ilt riroilar lihie» itilo lb<> next riixuIu», wliicb preparm to reeeivo ibetn by
■ ntumtiou »r ila eirailar ami a eiiiiiracrtJoa of its longitudinal fibres.
fiiacv Ihc li|« of the iltn-ewcal vulvo are placed transversely acrOM tJie
moum, nut ouly doea duteusiun of the cnN^uni. by &ir«tcliing the ruUe along
ibe line of the li[i«, briu){ them into ap[KS)tioD. but the pressure exerleil by
tlM periMallic muvenMiui has the same elfect. In this tiay any return of the
caoieiits fram the large to tli« small intestine is prevented.
Arrind at the u|{inoi<l tiexure, die contents, now more or lew volid feco,
an MMMirted hv the bladder and the tacrum, to (hat ih«y do not pniM on
S 8T8. Dotation. Ih'a b a mixed act, being MuiiuTficially ib« result 04*011
•tfurt of the will, oiul yet carried out by mentis uf an invuluntary owdianban.
Pari of the vnluDlnry elforl (viusisls iu producing a jircfBure-elMct, by meana
of ihe alnlcimiiial niuacloi. ThCM arc contraett.il fnrrilily an in ex|iiratioii,
but thr uhiitis Iteing cluaed aod lh« eava)K: uf air fnm the liiuif* prevented,
(he wb<i)v liim- »f ibe premiire in hrnught to Itear iia iht^ ab)h>men it«elf, and
«i driver llii- ei>niciil--> •>!' the drM-t-ndiiig <<idoii ounaril loward ihc recluin.
The i)>[moj>) Hcxure it> by il« )>"*iiion vlielKirt'd fniin this imssure ; a hotly
intmduivil per snuni into ihK i-iiii>ty r««tuni is not nUhctod by even foreible
eofiirsriimif' of Ihi- alHloiiiiiial nails.
The arms is guanled by the spiiinvtrr nni. whieb is habitually in a sinte of
aoraial tooic <x>ulracli'>n, rapable of tieiiig inervused or diminished by a
KJmutiis applied, either iiitenially or externallv. to the onus. The tonict
CutUradion u in part at l«a«t due to the action oj a nvrvoas centre silunled
in tb* lombor sninal cord. If the ttervotis connection of the sphincter witJi
ill* spinal cord be broken, relaxation takes ptnco. If the spinal cord be
divided somewhat higher up, for iiuttanee iu the duival region, the sphincter,
•Aor the deprewing elfect of the operation, which may lost several days, bas
pawed olT, regains and «ibM^|uently maiotauu its tonicity, allowing that the
tMitre i* not placed higher up than Itic lumbar regioD of the cord. The
884 TIIK TISSLES AND UECIIANISUS OF UlOBdTlO.V,
incrawed or iliminixbi^il crmtriicUao rollnwinft an local slimiiUttoo U |ir
ibly diMJ Ui ri'tlfx iiu)fin«ntaiioti or iuliitiition of the nt-tiou of tlii.i tviilre.
Tbeveiilrp in hImi xubjeot to iiillu«tice6 proeccdint; from hivWr re^ona of the
cord, Kiiil from th« anlo. By tlie nciioii of iIk- hIJI, by einoUon*. or hy
Othrr iK-rvouK ■•vfiil*. th« luiuluir »>p)i inciter i-eiitrc nmy lie iiiliiliit«<l. iind thus
Uie HpluDctvr ioclf rdnxoi) ; or atignic-iiU-iJ, oiid ihun ilii.- sjiliinuriu- li^liitued.
A Koond ilcm, Llivrcfure, of l)iv v«]iiiiiary proem in d«r«catiiiu i» llw
inhibition of the liimlxir aphincttr cHitn-. iiq<I conte<)Uent reliu(Hlii>ii of the
ejiliinctpr muffle. ■Since lh<' himlmr I'l-iitrc iiiny ntiiiiin whcillr «ffici«iit
when Deparatvil from iho hr«iti, lh<; imnity^w i.f the "pbiricttr which ooeun
in cerlfliii cerobrni diiciu<c» i» iirubiibly due I<i inhibition of thi> lumhnr
cesirp. nod not to pKnily>t(t of nny ccrtibrnl centre.
Thus a voluntary conlrnction of th? iilMloniinnl wiUK nccnmjHuiird bv •
ivlasalioD of the sphincter, might prcw the conlvnla of the di^>««ndii>g colon
into the rectum nnd out nt the iinii». 8ince, howoTcr. lu nr« htiTC B(«n, iho
pr«t8ure of the abdominal nnlls ie warded off the cigmoid Hezure, such a
iniide of deJecatioD would aiwayi end in Iciiviog Ibe eiemoid tloxure full.
Hence the DCceasity for thcee more or lce» voluntary nc[» beiiic ncwmpanied
br an involuntary iiiif;nienlalioii of the peristaltic action of tne l&r);e inue-
ttncsiffTDoid flexure, and rectum.
In the movements of the reciuni we cad trace out more diBlinctly than in
other reginnii of tlie alimentary canal the wparate actions of the loo^'iiudinal
and drcular fibres. The former, by means of coniraciioas travelling from
■bove downward. Hhcirteu the rectum, nnd aiiiee the auu* aHbrda it more or
l<n flxul HiijijKirt pull the rectum uiid ltd iwitcnta down ; the latu-r, by
incani* of c<iiilnti.-ti<'U!i tnivi-llin^' from above downnnrd hut taking place
•umi'whHt Inti-r, inirrow tho ri'cluin and »> Miueew thi; conti!3)t« onward and
outward.
X)efc(ratii>i) then ni>i>cjirs to take plan* in the ibllowin^ manner; The
lar;^ intestine and »ii;moid (Icxun^ becoming more and moro full, stronger
and urouger peristaltic nclion u cxciled in their walls. By this means ihfl
feces are ariven into the rectum and m. by a eonliniianee of the movemento,
increasing in vigor, against the sphincter. Thruu>;h a voluntary act, or
sometimes at least by a simple reflex action, the lumbar sphincter centre n
inhibited and the sphincter relaxed. At the same time the contraction of
the abdominal iiinecles presws firmly on the descendin): colon, and thus,
tjoulxactious of the levator ani awiAting, the oonients of the rectum arc
Reeled.
It must however be remembered thai, while in ap|)ealiiig to our own con-
gdoufnen, the euntnietion of the ahdumiual wallH aud the relaxation of the
irphiucter t>eem purely voluntary' eltort». the whole act of df fccittinn. including
both of llieM; KM-mingly *a \-«liiiilary eouiponeobi, may tnko plan; in tha
abwnev of cunaoiouonn*, and indt«d, in th« case of the dug at IcaiM, after
tbc complete acvenuKC of th« himlmr from thn donml cord. In auch amu
tbc whole act mn»t be purely reflex, excited by the presence of fecM in ibe
rectum.
S 877. The nmviu meehaHimu tf i/attrie and intentinal moivmrntt. Uotb
the (tomach au<l intcMinn when renitived from tl>e body and thus wholly
Hjvralod from the central iier^'ou" MVHtem may, by direct Btiniulation, be
roidilr exeite<l to movemeuis; aud ind-.-ed in the alweoce of all obvious
otimufi, movements which teem to be fjHintancous may at limes be obaerrod.
The movements of which we nre irpeithing are or^lerly movements ol^ a
periataltie nature, not mere local contractions of a few bundles of [»Ia[d
muacular fibres. Tlie alimenlitn- canal therefore, like the heart, lh»uich to a
Ims degree, poaseMes vithiu itself such mechanisms as are re(|uiAt« for
TIIK HtlSOtrLAK MKCHAN19HS Of DIURSTION.
S85
I
mrryixtf oul tii owd iDoverae»ifi : itnd, hb in iW case of the heart, lli«re is
M> •d«(|iuiu e?iilenc« tlint tlie gaugliii scsllem) in iis muscular walU, tbwe
■HDtljr formtDg die plexus of Auerbadi, play onj prini« part to (I«velopiuK
UtCM HMTemeDU.
On the othrr bBiiil. nuui'Hul movement of a perUialtic kind inajr be
inducnl, out oiilv iih wr Imvo alrouily Men lu the iMuphuf^ux but alio in lli«
MoBUcb, in tbe sninll inicwline, uul eveii in (he large inteAtlue t>y mimulatiivu
at lb« nwiM iwrre.
^W chwf and u«tiit) c«tif« nf the movcmcnla of tivK Momacli unci intiiftincH
ia tb* pnaaooa uf fi»Hl in ih«ir iuu-rior. But vm do nut know ilttiuitely the
•XMt manDer in which the food prtxiuctn tbu niiivemenl. It ina^ be that
the food, bv stftDUlatiog ihv niuous mcmhrniic. iwndit up BfTorciil inipulcc*,
aai) that ihne give riw by tv^vx action to rfr«ri-nt impuW» which deaccnd
th« <raff>M Sitnt to MK-ccwive portioue of the canal, in a niamicr nmiliir to
that alraady deacribcd in icIVrciicc to lh« u«ophagn«. If ihi* be *o tht:
cflereat Impalan rncb ihr alonmch aud upper part of the duodenum by tbo
termioal ponions of tlK' tuo rn^^i, Pig. 120, R-V., L.V., und rmch the irilw-
tinea bjr Ute pottion of the right or portprior vagii». Pig. 1'20. It'.V"., nhich
paNna islo Itte eoliu plexus and tbcncc by tbo mesenteric Derrce. The
fefinvat itnpulafS from the stoDiach travel also a|>parcDtly by the vagus; ti»
{■lbs of tfaoae from tJ>e intatines have not yel been determined.
Itut that !ucb II reHex action tbrou^ih vagua fibres ia not the only raeana
W which the jmecnce of food brin^ about the tnorements in (lueatioo, la
iMVn hy the Act that thcee continue to be developed after seistion of both
ngOB nervca. Probably ibe whole action la a mixed one which we may
pkUm to ourselves somenbat as follows: The alimeuUirr coiial poMeaaee a
M««r of Bpunlaneuus n»venient, feeble it is true, verv Inferior U> thai of the
bnn. and vvn- apt to bt: Ut^'nt, but still existing. The prfwcnoe of food in
•aoMi way or other, by M>m« dirrct action quite apart from the central nervous
^auoi. u able to inereaM; this |>owrr mi that, without any aid from the central
nervous system, as after M>clion of the va^, adequate peristaltic movements
van. uudvr favorable circumstances, Im carried out. Nevertheless, in the
Dcirnal roorre of events atisfactory movements are dill further secured by
the retlex sction through vagtii fibres just described, llius, in tlie dog, the
•M of swatUiwiuf; fiKid or even the mere smell of fo<id has been observed to
imnaaii tiie niovemcDt* of ■ piece of intestine isolated from the rest of the
aliOMBlarv coimI but retaining il» coniiectious nith the central nervous
^ntMU. tinder this v'kd the perintallic niovenienlf produced by centrifugal
■dnulatiou of the vagu^ in the neck are comparable not an much with the
eenuaotion of a skeletal muscle wWn its motor nerve ia stimulated as with
tbe beats which may be called forth in an inhibited or otherwise quiescent
bout by stimulation of the cardiac augmeiitor fibres.
Indera. we may, perhaps, call ibe ra^s fibres uhicb pass to the stonaob
sod lottatines (and time, we may remark, are, like the cardiac augmantor
Hbre*. non-medu Hated fikr«* along the urtUtr part of their coiine) aaa^
iB«Dt«r flbrea rather Ihsu motor Dbreit. We have all the more rvasoo tu do
ID sliwe tbcn exist conniauion but untauionifttii' iiihlbitiirv fibres. If while
lively peristaltic action is goii^ on iii the t>oneI* the i'plnnclinic ncrvt* he
Miinulatrtl, the l»wclii are brought to rtsti often in u very abnipl nnd marked
BMincf. Inhibitory fibn-a. thcri'tiirr. run in the splanchnic nerves ( Fig. 1'20,
UpL maj. aud mrN.). paM<ing nlotig ibi-ni iVoni thotpinal con) to the abdom-
iaal pleKUseS. and tbcncc to ibc alinif-nlnry canal : probably some of tbo
tiM Bitdlilleted fibres which may be obecrved along thin track ar« of this
Dalnrs-
It will be noticed that tbn splanchnic nerves, while cootaiuiug vaao-oon-
886 THC Tti^Kl-KS AND UECII ANItiUd OF DIQKSTIOX.
Rtricl'>r. I. P., augmeDtor, fibres for the blooilveaaela uf the itilestinua, cmrrw
inhibitory fibres for the luuscular vDat; aud [trobaibly the rogiM. whiU doo-
Initiing niicnientar fibred Tor the tuuamilar coat, nrries iohibitorv dilator
fibrai' for the bluodvmiieljL It mtiv further be rumurkeid that ttw vagns,
vrbile eupplying an^tiK'nt^ir fihruo f()r ibi' niiUK-ulnr iiienUttniain§ of the
aJimontarj' cnoitl, eiirrits, tt» we so wttll kn<itT, inliibiturj- fibres for tli.j
cardial- niusaular ii]cc.hiu)i«in.
KI6-1W,
RVi
LV
-pi
"0*.pl.
i-^D,6
..pi. hyp.
Lr
Rst.
DlAOSlH R> ILLISCUTI Tn> NKRVW OF mi AWOonutV CtMAl, » Tin DOD.I
Tb* l>ura !• fur Ihe wJib of midpUcKjr maAQ u •lluicminnuitk u |iciMbI*. ami itaa KM rqin«W~
Uc amila«iiq< rdnKnu. i~>v. bi I(c4. TliDftllinanUryMriiLl, >i-»i.hiu(ii>.>l"iiuicli.aiiull lni««U(ict,l)U|*
UiMUd», raMiuL LV. I.enii*8»*ii>-rr(i. •iiilliiir<iii rr»>lnr>lovuu<h. rl. leciirmul UrjrnvMl Mr**
ninl)rlncBpi«Tt«neCaaophBfni. tt V. KIbIiI TOciuJolnliielcA nKiwIii <i>->ii)i>Bi«)pl«Kiu, oe.
tl-> niiJilrUiB tbe poattfrfot |«tt nt •loitiadi uiil contlntinl *• K'.V tojiin the mUr ptaRHi. Stn
npramtnllvaiditMiniWlMnan'IrnnnwiwI wlili Uw Infrrior innHicnc iruiclliiti{iitplri(ui m.al.
— «. bnuwtiM ftiMn UiOKibu pluiuloiMuuicliitadnniLll lalcMlno, and fluiii il;v mivgnlalc m^ioa
Ulha lam IntoUne. !^pl. nu)< Ittw ■I'Iui'^bIob'** *'t*'"ll '''ooi Itic thorBrtf caiictlt ami ninl
MMSMiiiicaiila, r.CL, Iwlonalag lo dniml u*m* from ine4lh lolbeSiliior lliilil .^jil, mln, «awtl
■pMachnlr lunxr tlmllarljariilBf Itom IMImnd llili ilonal iierv**. TbuwtullitaintlieHilttr |>kia>
■ad UirfKD make Iboli «a]> lo (li» ollmanUf} ouiut, Cc NtrrM (Tom lli» (■nulla, «(f., bclan|ln(
MllikuMl mil donal and |<| anil :U iiimtar atmt. |>nxvt>Unc MibelaJbrtat nwviiMrir ta^Ua
lOcptHOi). n. s>., anililicncatqr UMbnoCuMc nerve ii.hrp. anil lb* lijrponalrlo pl*iila |i. liJii
Is thi dKiOar niEiKl« of (ba iwciuni, 1,1, Ktmt fcim Uia Ud aad M matl cenat. »■.£. aU invrl
M||»nuu, i>n>ti*tdlu( tr ibo nn>i««*rtc tAtxm to (be laiMtHndlual Budaa «f (h« rMtna.
In the abovp ttmcuii'iit wp hnvt purjimii'lr ii»oit thp general term, pori-
etallic HMVcnM^Dt; but, m vk hart •oou, in the muvemenU of lh« aliiDenury
I It SH DM ntwnai niitll loo laio Am Iu ■b«<lU<niiBol ih» nuTMnr iha allUMiilarT euial In
<!• d«i, Mm dorwl noiTM bad lucn npMnaM. TBo Os"'*. ■« tuwd. makoa nu pxctciic* M
■aauMlnucmMnaB: but li would tiara taa&bKWr id rc(irsaiit^iIiurUiUtteaurafii«a|im|11Q
TUK HL'SCUI.aR MECilA.VlSU8 Ol' DinKSTlON.
S67
^^ AHL two Ma of uusoles are coacerned — the circular iind the lonj;itu<liiiaL
P lfW|b tbe rectum we are able to recoguiie that tlie iwoacUuf inusclM
r bvtwlu tliMiDct iii-n'ouB aufipliw. Tbe looptudinal coal is goveniwl hy
I iurT»>fibmi «hk-)i. ill ll>c Acyg. leave tlie anioal cord iu the anterior riM>u in
I cJi« arcuod anil tliinl Mcral ncr\'<at Fif:. I'lo. ■S2,>$3).paaB along the branobea
I of ihuK oorv» frL-iiuetilly ximken nf aa the ii«n'i erigettiet, I. r., to tli«
I ImngMtrie pleiw, {pi. A^j.>, and Ihciire to the rectum. StimulatioD of
I IMM nmta cnuaoa contmciion* of ibt: n^clura. which are conIiDe<l to tba
I loagtttidinnl ami aud, w vm huxc .-aid, pull ibc n^'ttim duwii. The circular
SMI U piTVrTH'd liT ti)irc9< which Ivavo Uw opinal conl by lh« anterior roola
of Ibv lower dur»al and lirrt twn lumbar nrr%'i«, Fig. 121) (coming fWtm the
luwwr |«i1 af that »)iinal nngion fnm which, lu we have *ec», §169, the
Tan>-c(>iii>lniHor l)tin« take origin), and, nirly Iwin^ their incduUn, pwv to
iiim r«eium by ilio inferior incsicnteric sanglia. Ihc liypii^iric ncrvva, and
''fP'^KUtrir pIcxuM ( Fif;. 120, iit. i/L. n. hi/p. pi, hijp.). Stiniijlalion of th«ae
'twGM give rifv to roftlfacti»tis which are confined lo the circular cimt and
^I'li !■ mil Ibe oonieDUof ihc r«cluni. A Rimilar double nervoiii tiipply
ivotkably goverot ibe lunj'iludinal and circular cont* along tho whole
■IllavnUrr caiial ; but the delails of »ich a supply nrv at present unknowu.
Otir konwIedKc, nioreovfr, oouccmint; the details of niiy special nervous
^^^lianienw, bv ineaiiB of which tbe more complicnted movements of tbe
*(»ntiirit. tnt.-t(idiDK the cloeiii;; and opening of the »nhiDct«re are mrried out,
" •*! j>reti«nt very imjwrfevl. We cannot add to what we have inddeiilnlly
*'*J m speakinjc of vomiting.
1*lie moTements of tbe rectum, including the fei(;moid Heiure, appear to be
''\'*<^b niure chwely dejiendeni on llie ceutnd nervous system than are tboM
?f the real of the aliraeniarv ouial. As me have Mitd, the moveinenta of
^**-t) btrge and nnall iotwliDC are rallter aaualed and augmented ibtB
f**ciMrily otllcd forth bv impulttcs dvc«ndiiig from the centra] nerrooa
T****! Mong tltc vagus fibres- A* ibc large intestine, however, pnssM into
~^* rectuiD. giivcmment by Ibe vagus is replatwd by govemmrnt through
^^^ lomlMir cord and tbe nerves jtwA previously meolioned : and this goveni-
I ^*^«at apiiears to be not so much mere augmentation as the actual carrying
I V^tt of the ntovenients through reflex iiclioii. Hence, this is tbe pari of
^lit«e|ina] tDOTcmenl which fails iQ dlMasca of the central nervoii* system,
^)ie failure leading lo obstinate cooMipation, if not lo nclual dilticulty of
^sfeotion. Tl*e presence of feces in tbe sigmoid flexure no longer etir* up
ife icAex inechanifiii for their di^liarge; meanirhile the more iodependGut
■OTeatents of the higher [wrts of tbe canal oonlinue to drive the contents
Mwanl, and hence the fcMti accumulate in the sigmoid flexure and colon
svmiling the delayed action of the imperieot reflex mechanism. Witli refpo^l
to the exact manner iu which the |>feseiKe of (bod acts as a stimulus, it may
bt> iioith whik to reninrk that, althuujth in tbe »lonmch, aa we have seen,
nnv fiiliKiw i* not the efficient ratue of the movenieiita, vince thete lieeomo
tDum active as digealiom proceei)* and iIh- bulk of the cnntvnls dimininhi^
»y(4 in the iniestioe distention of the bowvl up W cerlain timiln mu.i di*-
linetly inoraaMS the vigor of tl>e movotumitd junt ax diiitenti»ii of lh« canliiu:
tBvitira oitbin certain limits improvw itte cunliac Mroke. Thi* ii« wull ntn
in nhttruriinn of the bowcU, in which cnsoi tlie buwul dt*u>nde(l nhovc tbu
nlwt ruction is frciiiiently thrown into violent pttrirtnllic iuuT«ni*Dti>. This
tdMl i* in part nt lea** rlue lo Uic distention exlvn<ling the niucalar Hbr«,
bmI mi in a ilirect niDnner promoting thc^r contraction (■•0 $11). but may
be ill i«n due lo au-'iiK-nior impulses excited in a reflex manner. Probably
hi an inte^iufl isolated front the central nervous system f-xnl provokes jieri-
lUltie DKivenieuta much more by causing ilistcniion and »t elreivhing the
388 TllK TISSl'ES A.VO 11 EC II A N liillS OF DIOESTIOjr.
muwular (^^ais tban by acting as a slimtiluH to the mucout nietnbnitc, eiilter
ilifDUKli eliGmk-iil octioD or in May siiuilar nay.
tj 278. Next to the presence of food in the interior of tli« alimeuiarv cons
a d«llcieiit oxygeualion of the blood tiu|)tili«d to tbc walls of the cuoaf or ili
kudden cutting off of the aupply of bloml may be n-jpirded u Ibo mont i>uitfr*
rul pfovomtirea of peristaltic uctiuu. Wbeii tbu iiorta U clamped, or wlm
tbc r(«piratioii in ii«rioualy iuterfer^i) nitb, pcri«taldc niovemcnl* bccon
rery proiiouuccd. Thiin, iii dvath by HHpliyxIn i>r :<til}i»uUt»ii, aii tnvulunlai
diM-liar^ of IV-Cc*, vrliioli i» in part at lcu«l llit- midt of iacreoMd paririalt
artiun. IN not an unfrciueJit mult; and ibv niark<Hl peristaltic movcmioti
which an? to freiiuciitiy w-i^ii in an nnimni when the abdomen is laid ■
imniediftlcly al'trr di-alh apjtcnr In hi- du<- to the cuMntioD of the circtilatioa
and llio conttcniicni failure in the supply of blood to the trails of the aliruen-
tary canal, and U'jt, as hiu' been i>ugef«led, to the contact with air of the
giritoOffll stirfuec. Since it is bloixf which brings oxyg«n to the tiesuea,
ilure ID the supiily of blood is tanlamoiinc to failure in the etipply of
oxygen ; but the blood current brings other things besides oxygen and aUo
takes thing's away : nnil the failure of this action also probahlj', a.* nell as
failure iu the supply of oxygen, provokes the luoveincnie in question.
The movements iIiiie producer are to sonic exlenl the result of the defi-
cient fiupply of blood acting directly on the walls of ibe canal, tliougb in
a#jihvxiii at all events ibis etfect may be iucreaited by the ton venous itla
atititulaiiiig the central nervoua system, and thus sendtug aiiguienlor impul
down the vagus.
Wi[b regard to the mode of action of tlie drugs whidi promota periMullic
action, it will be HufBcient here to eay thai, while some, such as niooUn,
appear t« act dircellv on Ibe walls of the canal, others, such its strychnia,
prodiici: their etlecl chietly by acting thnmgh the central oervuiis syMeni.
Tbb Obakoes which Ttii^ Fuun L'kukuoue» ix tik: At-iMEMTARr
Canai-
$879. Having studied the properties of the digestive juices es exhibited
out^cU) the body, and the various mechanisms by means of which the food
introduced loto the body is brought under the itidtietice of those juices, wa
have Doa' to consider what, an nmtU-t« uf fact, are the actual changea wbich>
tliu food does undergo in passing ahmg (be alimentary canal ; what are tha
steps by wliji-h the eontenta of the canal are gradually converted into (temi\
The events which lead to this conversion are twofold. On the one hand th*'
digestive juices do bring alioul, inside the alinivntary canal, change* trhich
in the main are tbc nnie as thoiu! olxrrvi-d in laboratory cxpcrimcnU) outside
the body and described in previous .locliouti, though the refulU arc somewhat
modified by Ibe ^{iccial condition* which obtain ffltliin the body. On the
Oilier hand absorption, that is to say, the patwigc from tlic interior of lite
canal into the bltRidvcsscls and lymphatics, of digt^tcd mut«-rial in oomnany
with water is going on along the whole length of the cnnal, and (vjiccially in
tbe sn>all and large intcslines. It will be convenient lo conRnc ourM'lves at
present to the study of the first claas of events, the cbangvs eflectdit in thii
canal, merely noiini; the disap{}ea ranee of this or thai product, and di-fvr-
ring the dilftcuU prublem of how abeorjHion takes place to a sulwequetil and
separate discussion.
in thf nwvtK the presence of the food, anisled by tbo movements of the
jaw, cBusea, as we have seen, a flow of saliva, ily mastication, and by the
addition of muuous saliva, the food is bivikeu into small piocos, moistened,
I
389
I
I
I
■ad ntbrm] into a onvcRimt holti* for ilc^liilitinn. In initn Kimc nf the
Marea », even iIuHiij; lli« fUnn »liiy of ihc fo™l in thu mouth. (.■unvKrtctl
iota Migar ; for if ImiM rtnrch frer from mgnr be even inDitivnlnrily helii
to tb* mouth Biiri then rjcctcd into nnt4-T (kept lii>iliiig to dvstror ibe
fiiraHnt), it will Iw funnd to conuiin ti decided ntnonnt of siiiinr. Id many
utiaiAk DO (udi t'buag« takeit place. The viscid ealivii nf the dog «ervM
alnm solely to urist iu degl'iti'''"! i "ix^ ^v<!i ^^^ longer «tny which food
mrnknt in Ihe tnouth of tbe noree ii inBuflieient to proditee nny nmrkeal am-
fWlluu of Ibe March it may contain, Inuring tbe rapid inmeit through the
MmiAd^tu no appreciable chanee lakes place.
Tlie Biuouiil of iibsorplion of dignteii inatertal, or even of aiinplo n'Oler
flfmn tbe mouili or oasophaRUB, inust always be iofliKniBcanl.
7%« Chaa^ IN the Smnaeh.
i 880. The nrriviil of the food, the rwictioii I'f which in either nnturnitv
alkaline, »r ii made nikaliiie, or at lead is reduotnl in addity, by tho oddi-
liea of Mliva, miw* a flow of gastric juice. This, already cniunittnciog
while the tonl i« yet in the mouth. iDcnvises an the food nccumulstn in the
alnumeb, and aa, by tl>e chuniiiig giLi>tric moreniQiits, one part after another
of the firad is brought into contiict with llio mucous membrane.
Tbe chanctefs of tbe juice appear to change somewhat as the net of digee-
twm proceeds, The amount of popsiQ in thp gastric <xiat«nts increases for
MOW time after food is taken, aud probably the actual secretion increaMa
•Iwk Tbe acidity of (he gastric contents is at first very feeble : inilee^l in
nan, la some cases at leait, for some little time after the beginning of a meal
DO free acid is present, and during this period the convereion of starch into
■ugar may coiitinue. Thi» condition, nowever. i» temporary only; very
won tbe oontenu become acid, arrMtinc the action of and ultimately
dMtloyiag the amylolytic ferment ; and, *!nce the nit« of »ei-retioii of acid
appean to be fiiirly i-nnttant, the contents of thf xtnmacb, unlea fre^A alka-
line fitod be taken, bccnnie more add a* iligiMtion goes on.
The graa eBect of gastnc digestion is to hn-ak up and partly t» di<«olv«
the larger luBim of nHtttcated food into a thick grt-yiih Hiup-like liqtiK)
called fliynw, with which arc itill mixed in variable <jiianltCy Urnr and
smallar naans of len changed foml. This is the nwilt. partly of Uie aolu*
UuB of imXcid miilt^Ti, partly of the tolution of the gelatiiiiforoaa eonoec-
tin ti«ue hutiling the nroleid elemente lognlher. In a fragment of meat,
for iniitanee, the muscular fibres, through the solution of the oonncctive-
tiMue binding tliem together, fall asunder, the sareotemma is dissolved, and
tbe tibr« themselves split up sometimes longitudinally but most frequently
by transverse cleavage into discs, and are ultimately more or less reduced
|Mrtly into a granular mass, partly to actual solution. In a piece of tissue
ooDlatning (M, the counectivo-tiasue bto'ling the fat cells together and tbe
oovelopee of the fat eelU are dirsulveU, so that the fat, fluid at the tempera-
tore of the budv, is set free fVom the individual cells and ruiu tojcethcr Into
larger and nnialler roaaies. In vcg^'i-tnlile tiaaue the proicid etementM are iu
part diMolved and, though there it no evidence that tn man vellultwe i« dis-
solved in Ibe siomacb, the whole lUsue is softened and to a certain ettenl
disinifftated. Milk b curdled and the curd subeequeoily more or lesi die-
solved.
Til)? thick w<i|i-liko arid chyovc cuuisU acoonliugly parilv of HitwIanoM
which have entered into artoal w>hiiion, partiv of mere particles or droplet*
of proteid, fativ or nlhfr nsiun^ and partly nt manes imall <>r ifreal. wtilch
may be rrcogniwd under the inii*r»»ci>|>c as more or le« changisl poritoai of
390
TBB TI98UBS AND MBCHANISMS OF PIOESTIOI*.
niiiuuil or vc^dublc liiwiii\ Tlic muouut t\f niiiUTial ui'tuitllv iltoKiIrcil ui in
BUM flpAciDK'nH i>t' chyme vxcceilingly niinll. Wbcii ihv luiliil piirta ore
roinijvril by flllnuic>ii tho clair fillriilv c-uiitniiiii li(«i<l<« nlu, pcnnn uml free
hjr<lruchloric iicid (ihit <-i>mIt[iii.'iil» of the gulric juice), a «miill aiaoiiDi of
nigsr, of pantpoptonc nix) of ptrptono. The sugnr ui oftr.ii ndfCJit, tbc pnrm-
peptonc is nol alvcuvs present, nnd th« umounl of peptone (or ulkumuM) )■
alwny» ymM.
Duriii); gastric <lig«etton tho chyme thus forin«<l is fram tiin« lo tiaw
ej^clciJ ihrough the pvioriit, ncoonipnuicd by oven )>t^ morwls of sulid
let«-(ligr«ted matter. "I'hia may occur withio a fow iuitiitt«« of food having
be«ii laken ; huC ibe larger escape frt>ni the sloinach probably iJnes not in
tDan begin until from one to two, and lasts from four to five Iioim. after ths
meal, becoming more rapid toward the eud. and such piec«e as are tlie kut
broken up by the gaatnc juice and movomenia bein^; the laat to leave Ibe
stomach.
The time takvu up in gaatno digeatton probably varim in the rtame animal
not only with diflereiit artii-les of food but aUo with varying cimditioas of
the stomach and uf the IxHly at large. lu ditlerent aoimaUi it varies verr
const dcnihir. being from twelvL- to twenty-fiuir honn in Ihr dog after a ftill
nul, whilu ihf xtomuehii (if rahbit.i an; never empty lull alnrayN remain
largely fillt^l with tixid, itvMi iluriuj; starrtition. la nuui the (tumach proba-
bly becomes cmjitv bclwoi'a ihi: iinunl mcaln.
The total amuiinl af ohaiigK which the fnod iindergoM in the stomach,
ibst is the share lalci^ii by iho siomach in thi- whole work of (ligmtion,
Mcms to vary largely in diflcrcut antninU and in lh« same animal dinbn
acconling to thi' nature of the meal. In a dog fr<l on un oxclnsivlv meat
diet, a very large part of the diucstiun i* »nid to he carried otii \ty the
Stomach, very lillle work upparenlTy being lell for the intwtines; that is to
say. tho larger part uf the meal ib re<luced in the itiomach to actual solution
and a cuDiiiderable 'luautity h i>rol)ahly abxurlied directly from the Mouiach.
In such ctiscs rhe amount »f jieptone found in ihe Htomaeh during the digea-
tion of the meal in found li> lie fairly couBiaiil. froai which it may be in-
fltrrcd that the itepiuiK* is aliMirbed ua soon a^ it is fonued. There is also
evideuce that fat may ti> ii cerUiin extent undergo in the Hlomach chauga
leading to emulsion, .limilar lo lh<i«e which, as we shall see, are carried out
in the stuall inintine.
But such ciises ns thtw cannot be regarded as ty|>ictt! caiu'ji of guiilrie
dtgcfllinn, and in man, at nil cvcuts. living on a inixeii diet the work of the
•tomach appearp to be to a large axtcnt preparatory only lo the sul]icqucul
labors of the intestine. It is true that our information on this matter is
iiuperfecl, being chiefly drawn from the study of eai«s of gastric or duodenal
fistula, in which probably tho order of things Is not normal, or being in
large measure deductions from experiments on animals, whose economy in
this mpeet must be largely diRerent from our own ; but we arc probablv
lift in ooncluding ihal. in Kurselves, the chief eflect of gastric digestion is
by means uf the disiutegraltou i4|Kjken uf nbuve to reduce the tumps of tbod
to the mure uniform chyme and so i<> fscilitalc the cbangcs which take place
to the snuill intMiine. During the dUiiitegraUon some of ths nroteid in the
meal is convened into [leptoue; and the peptone so formea is probably
abcrorbnl at mvc -. but much |)r<fte)d remains unchanged or at least is not
convertol into |M-pl<>ne, and the fata and starches undergo in themselves
very little change indi-til.
In the act of sirallowing. no iiicunitidcrable •juantity of air is parried
down iulo the slomneb, enianglcil in the laliva, or in the f<HH]. This is
returned in eructations. ^VIlcn tho gas uf cruclaiiou or thai obtained
CUANOKS I.V TlIK SUALL IXTKSTINK.
891
k
t
tlv fmiti the MoiDkch r* oxnmiiifd, it it fouiid to coDsi«t chicflv nt
BtlnMKn iiDil cnrbonic arid, th« uxyet^ of the Btmoaplicric sir Imving iwon
iumSr mhm>Tbttl. In most cjms liie carbonic acid b drrivrd hy »iniplo
dininoa tnia Ibo blood, or ttvm the lumin of the itoinscb, wbk-h Mmiliirly
taliv np the oxyj^vii. In mnny oniK-e of fliitulrncy, linwcvpr, it mnv nrUc
frooi ft ftniwnlattvc dcrompr«riioti ol' lh« sd^ju- which hue been InVon na
■ocb in food or which bu bcrn prwliiced fn^ni the slnrch, the giU: l)ciiiz
atb*r femed in lb« ■tomacb or piwein); upward Irom the inlcflline ihrougo
itMpjrlonia.
Td* etwnnoufl quantity of ga? which is diacbarged through ihe mouth in
CMea of hyiMlical flatulency, even on a perfectly empty eloiunch, and wbich
Mcna to eousist Itrgely of carbonic 8ci<l. preeeiiu difficulties in tbe way of
•xpfauation; it is poasiW that it may be »imply ]iifi'uae<l from ibe blood, but
it » mbo poanble Inat in many cwwm iI m derived fVuni air which the patient
hu hy«teii«aUy swallowei), thv uxy^cen liaving been remuved, in the atoniaeb,
by amorptioa and replaced by oirhouic acid.
In liif Small InUttinr.
{ Stl. Th« ■eini-digc«ted acid food, or chyme, as it paMn ov«r tho biliary
ariUce, caiiBce a^ ne have H«n (^ '2AA) (lusbes of bile, and at the same time
the pancreatic juice flows into ifae iotealine freelv. These two alkaline
fluida, eapecially the man atronglv and ooi»tantly alValine pancreatic juice.
Cant) ti> oeulratiw the acidity of tlw chyme, but llie {■oiiieiilit »t' the duoile-
■DiD do not become iliminctiy alkaliuc iiutil Mime diMlnuce from the jiylnrua
i> Raeh(«l. Tbe rapidity with which the c.hanK<! <■■ the readion is com-
)4rU<d b not the in me m nil animnlH. and in tlic Mime animal appeora to
vary avcording to (be naiiiri; of tlie f<>»d. and vari<juf circumnance*. In
man. living ou a mixed diet, the content* have probably bceonw diatjlictly
olkalino before tbey have jiatwod far down th« duodenum. On the other
hand in doga. tb« ronicnte of the small intcslino baro bc«D observed V> be
arid thrmigfaout. and that, not oidy when fed on starch and fnt. which
migbl. bv an acid fermentation of which we shall prouotly ejieak, giro rise
la an and reaction, but even when fed on meat.
The coDTentoD of starch into sugar, which as we have seen is sooner or
Istar arrceted in the ftomach, is reaume^l with great activity nud indeed
completed by the pancreatic juice, po»ibly aMisleJ by the «iiccu6 entencus.
tbe nrcaeoee of bile beinf( asid to increaM! the activity of the p«iicreaiic
tBylolyiic fermoni. The conversion begins as won as the acidity of the
diyme is sufficiently reduced and ctintiriniw along the intestine: fH>nio(i».
bcnreTcr, of still utnligested starch may be found in the large iniestiae, and
even at tiroes in tlie feces.
Tbe poncreolic juice, aa we have »fcn, cmuUifles ftus, and aba splits them
(DIA iMir nspective falty ncids and Klyo-rin. The fatty oeids thus set IWe
faecmne converted by nu-nni' uf thi- alkaline contenln of the inlestiDt Into
■Mpa: but t» what exli-iit MpfiniK cation thiii> lnk<w place in not exactly
known. Undoubtedly M>ap navx to a small extent hvMi found butli in
piirial bl«(M) and in tlur thoracic duct niter a niMl : but there is no proof
that sny large iinanlitT of fat i« inlrtHlucvd in this form into the circulation.
On the nihvr band. if^M pn-MDcc uf oeiitral fals in the hKtMils, aitd to a
slight *ilenl in |t<>rtHl blood, is a cnii*picunuB mult of tbe digMlton <d' fatty
Biatten : and in all pn<t>abilily Hnpimitimli'm !n the intrvtine is a subsidary
iirocm>, llie eifecl of which is rather to facilitate the emulsion of neutnl
at* than to introduce soaps at- sucb into the bloml. For (he prmence of
sulnbte toapa bvore the emulsion of neutral fa(». llen<xi a ram-id fiU, t. €.,
892 THE TiaerKS and hkchaxisicI
roiSTioy.
■ fill ooniainine & oortain nmaiint of (no fntly ndi), f»rms nii emitlsiun witli
nn alkaltiM fluid vaon readily ilinn does » iinitc iiL'utrnl fm. A dro{> of
ruocid uil lot fall on the tiirrnor of nn nlknline fluid, such a» a solutioo of
Kidium cjirhonnto of suimble slrpiigth. rapidly forms a bruod Hag of emul-
aion, and tliut cv«n without the Toimt agitniion. As sapooUicstioD taket
place nt the juiicttoB (if the oil and alkaliac duid currents are Kt up, hr
which globules of oil are deCnched from the maia drop and drireo out in a
ooutrifugal direction; the iDtengily of th« curreota and the couBequent
amount of emulsion depend on the fou cent rat ion of the allcalioe niedium
and on the solubilily of the «aa})8 which are formed. Now Uie bile ami
pancreatic jutoe ■upply just BUcli conditions aa the abore fur einukiomung
fats; they both to;i{«ther a0brd au alkaline medium, the pancreatic jutoe
siviss rise to an adequate amount of fr«e fatty acid, and ilit; bUe in ndditioD
Erin^ into solution l)ie B0tt!>9 as they are formed. So tlist we cuay tpeak of
the emulsion of fata in the itmall intestine as being curried on by the bile
and paiicreftticjuioe acting in cDujiiiictiou ; and ana matter of fact ibebile
and pancreatic Juice <lo largely utiiuUltS- the coDteuta of the iiniall intcatine.
MO that the grayinh ttirliid chyme U cbajiged Into a creainy-Iooking fluid,
which hti.1 ix^-n mimntiracH culliid chyle. It io advisahlii, hiiwover, to rcMrve
thix name for lli(^ i-imtcntti of thi: Inctcalx. Many of llie fnts present ia food,
for iiixliini-o, linltiT, alrviiily unnlain viimi^ fntly acid* when VAten ; for thcMi
Gilo the initial nctiim uf this pancraatic juice is Iciv neciM»3iry.
Thi« mutual hrlp of bile imd pancnmtic juica in priHlucing an erauleioa
explain' Uinccrluin extent the controversy which long existed Dottroeii those
who niaintaini-.d thni the bile and those who maintained that the pancreatic
juice was necewarv for the dig«etio» and absorption of fntty food. That the
paDcreatic juice docs produce in the intestine eiieh a change as I'arorc the
transftreDoe of ucuItbI fatji from the int«etino into the lacteals is shown by
the foot tliat in diseaaee airectiug the pancreas much fatty food frequently
pawee through the intestine undigeeted and t^n>at wasting ensues ; but it
eanooi be maintained that the uaucreadc juice is tbe awe agent in this
matter, siuoe in animals in which the pauereatiu ducta have been aucceasfully
li>[atured chyle ia ttiil found iu the lacieals. On the other hand, that the
bile ia uf utc iu tbe digenliuu of bil h «huwii by tbe prevalence of fatty
•t«ob In ciuca of iihstructiou of the biie-dui U ; and though the 0[M:raiion of
ligaUiriog the bilc-duclx and Ic-ailine nil the bile extenially through a tuituU
of the gall bladder U opi^n to cibjt^'ttun, «iiK'« it in sopne way or other so
txhauata the animal lu indinvlly i» ull<>ci digu.itii>D, still the rutults of
ex|wrimom« in which the rtwjrpliiin <if fill wan diHliuclly leaaenad (the
quantity of t'ai in the lacteiils lalling frum IVl to Q.'l per oeut.) by tltc
ligature and fistula obviously point to the same conclusion. That in man
the euceuK entericus poescwes a wholly iiisuthcient cmulNfying power is
thown by the observation of n case in which the duodotiuin opened on the
surface )>y a ftstula in such a way that the lower part of the intnttne could
be kept tree from the contents uf the upper part containing the hilc and
pancreatic juic« and matters pnjccoding from the atomoeh. Fats introduced
into ibe lower part, where they could not be acted U]xin either by the bile or
by the pancreatic Juioe, were but slightly digested. Without denying tbo
poadble aawaance of the suocus enlerious, or even of gastric juice, we nay
ooacla<le that tbe digestion of fat is in tbe main carried out by tbe conjoint
action of bile and |>ancreatic juice.
§983. \Vc have seen (^247) that the addition of bile to a digesting
Riixltire giv« riac tu a precipitate. Thi^ is partly a coarse, floooulenl pre-
dpitate, GonsiMing of |>ara|)eptone with some amouut of bile acidi, and
partly of a 6n«r, wore granular precipitate, which is longer in falling down.
OIIANOE« l.V TMK SMAI.I. INTKSTIN'K.
898
I
nd ecOBiita chiefly of bite arids vilh h varinhlc amount of ])eptone; the
bavrii ndinolvied nn the fiinhcr nddilion of ttile. even thou<;ti the reaction
<J \he mixtan nimin tu\A. En (he oppcr pnrt of the <Ii)i>d«Diini the inner
wrfiKv, if esitoined whil« digestion is i;oing oii, U found to ho lineil by a
(ulond floGcairat and f^niihir malerial, whit-b is probably a precipitate
ihv Armed: (he rwrpuee of this [irec-ipiutioo ia nrobahly to delay the
pMMig* of the unaij^Mtod ptrapeptone along the duodenum. >Ioreov«r,
rrt (hMn thb precipitatioD. mle mtM* the action of pepsin, even while
muti'Mi of the mixture tlill reoiaiiis aeid; an<t as ttoon ai an alkaline
retKiioD a eMabtbhed the peneiu is apnarenily (teAtrored by the trypsin, »o
th«l with the flow of bile and iMuicrcai^c juice iuM the duodenum the pro-
rave* which have been Koing on in the atomach oome to au end. In fact, it
wouM »ero that the juicM of Ihe various dtstricU of the alimentary canal
are mutually desimclive ; tllu^ while pepsin in an acid Noiution deatroys lite
active oonaUluefiU of talira and uf imncreaiic Uiioc (pri>l>iibly nl«o thoM of
the succuB I'DierivuA), it i( in ibi turn anta^niied or d<-4truy*)d by tlie htle
anil the uihvr alkaline juice* of tin' iHteMine. Hi.-nc«, paniToilic Jtilce intro-
dscad through the mouth inu»t low it-i powers in the slnmaeh, mid can only
be of naa aa an alkaline nM-dium nonlaitiing (certain protcid miittcni. On
tha otbaf band, if, an vf haw n-aitoii to lic1i<-vi-, the conti.'^nlii of the Klomaeb
■* tbey iMue from iImi pvlnmii ulill cmUain a Inr^ r|iinntily of undigested
praliUt, lhe»c must be digtvlcd by the pancreatic Jtiicc (nilh or without the
■^Mbm of (he succus onterteux). thi^ nrtimi of which sreiii* to he iMtii^tcd,
oral laaM not biadorad, by bih^. And in dogn f^d through ndiiixleual fimila,
tn that all gaitric digcation is cxcludcl. proteids nre completely digcnteil and
gin riH (o iiuite normal fecc. To whnt slnin the puncrcntic digcMion ia
cmrrkd, whether peptone n pmcticallv Ihe only product, or whether the
imBeraatic juice in the body, as out of Ihe l>»dr, carries on its work in the
nan dostnictive form, wlierehy the proteid malerial subjected to il is so
bmkaa down as to give rise to appreciable quautiiia of leucin and tyrosin,
H al present not exactly known. Leucin and tyrosin have been fonnd in
Uw iatcrtjna] contenla,aiM] may therefore be fonned dunog noritial diewcion,
bal whether an insignificant '[uantil^ or a considerable ijuantity of the pro-
leid material of food U rhiin hiirned into a cry^iatliue funn cannot be
didaitely Mated. The exteiii to wliith the aclioit ii curried 'a probably
Aflfenut in dilTerenl aniniuU, and probably varies also acconling ti> the
tmtun of the meal and the condition <if ihe bcxly. Possibly when a large
mad tUiDeoeaiary ifuanlily of proieid malerial 1* taken at n meal, together
with other auliataiues, uo inronsidcnble anxxint of the nmu-ids undergo thb
[trofound dianga. and, as we shall sec, rapidly Wv« the Wly as urea without
laving iKwm iimnI br the tisMCS, their ooutribution to the energy of the hiHly
being limilcl lo lh« heat giTen out during the changes by nhicb they are
eonverted into un*. To this apparently waxtf^fut ui>c of protei<U we ohall
ratam hi speaking of what i» called the " luxos «>ii»umplion " of food,
{tSS, lo dealing with the action of pancreatic juice we drew attention
{( 849) to the dillorcocr bc<w<«n the rMulte of pure tryptic digestion and
those obtained when bacteria or other microorganisms were allowed to be
pnMOt- We saw that indol. fur example, was the product of the action of
tbaa oraaaisma, not of trypsin. Now indol is formed in varying ijuantity
dnriDg ue digntioo which actually takes place in the inteettne, »om« of it at
tima appearing in the urine as indii^o-yielding substance 'indiean). Moro-
ovar, baeteria and other microorganiama are present in the intestinal con-
MMa, H«Doe, we must regard tb cfaangee taking place in the intestine not
■• tiw pore reaults of the action of the several digestive juices, but as thaw
nsalti Bodlfled by or mixed with the results of the action of miorofirgiaiimis.
TUK TISSL'Kfl ANI> MKCUANlSMBor UIOKSTION.
Wo rpokc ftbove (| 247) of bile as being ftQlJae|itic, but thu miut be Hiid«r
stood na mciiHiu|{ not tlinl the presence of bile itrreil6 the action of all mirro-
oi^DisniH uitliiii the intestine, but tliat it moditieii their action, keeping it
within ccrlain liniltd anil along certiiin lines.
C^nceniinK the exact iiulun:? nii<] extent of the cbang«s thiu due to tnicro-
orKanisuis, uur kuowlcdgti In at jirc«eiit rvn- imperfeeL The prot«cb sad
the earbciliy it rat's m«ui m be the fnod-nutTii im wbii-h tlM«« orKanisme pro-
dncc ihtir chief ctrci-t. Out of the prolridii tlicy give nwe itut oiil; lu indol,
but to nevi-nil other iMninoiiuit«, iimuiig which may )m n>entloDed pbeuol
(C«H,0), iif which a ^intill (jiiantity may l>c recogninol in tl»e feoeH, Ibe rest
being alisurlH'd and npfwanng in the urine in the form of certain pbenol-
ci>uip(nin<lii, #iich ii« pheiiyl-i'iilphoric acid. Out of protcidt tliey may also
form ihi' [K-uuliar iKii»i>noue bodies called f)toma!ne*, which appear in ibe
ordinary putrolaction of protelds. But their moM conflpicuona eflbctt are
those on the carbohydnttes. As the foml descend* the intminf, the preatnca
of lactic acid becomes more and more obvious ; indeed, in romc COMS tlug
natuniUy alkaline reaction of the ini«atinal content!) may in the lower part
of the intestine be chanj^od into an acid one by the prcscncv of lactic acid-
Now, lactic acid may be formed out of sugar by means of a tpoeial orgaaiam
inducing what ta ft])oken of as the lactic acid feroieiitaliun. And we ham
every reason to believe that in even normal digestion a certain (|uaniiiy of
sugar, ^ther eaten an such or ari^in^ from the aniylolytjc convereion of
Btaroh, does uot pan anay from iJiv lul^tine into the blood as »ugar, but
undergoes this fermentation into Uetic acid. To what extent tbis changa
tak«» place we ihi not know ; the amount probably varies according to tb»
amount uf carbohydrates oitcn, the condiiinn of the alimentary canal, aad
other circimi.iiance*. It may Iw under certnin circnmstaDoes Bimply a part
of normal digmtion; under otlicr circunutanoe* It may be exeeesive and
give rinc to troubles.
That fermentative changes may occur in tho small intestine ia furilier
indicated by the bets that the gas there pteecnt may contain free hydrugeo.
anil that chvme. after removal from the intestine, continues at the temiiera-
turn of the body to produce carbonic acid and hydrogen in e()ual volumes.
This suggests the ])i«etbility of the sugar of the intestinal contents under-
going th« butyric acid fermentation, during which, as is well known, cwlMtnio
anhydride and hydrogen are evulve<i. By this change the sugar is removnl
from the carbohydrate group into the fatlv acid urmip ; it is thus, an tu speak,
put on iis way to become lat. We Nhnll m'e hcreaDer that sugar may be
somenhere iu the biHly converted into liil; thin euuversion. however, takes
place chielly ir not wholly in the tiwun', and i>uch change Bd mav take place
m tlie alimentary cniial is to be regarded nit suggestire ratiicr than as
imitortant.
TIk' Iiydriigen Ihim occurring iti the inti^titie may also arise &um the
protuid <lecqni positions spoken of above. However nrixing it may act as a
reducing agent— reducing sulphates, for instance — and thus giving riae to
sulphides and to sulphuretted nydrogen ; as a reducing agent it oauMs in (be
fornialinn of tho fceal and urinary pigments.
Thuc, during tho transit of tho' food through the small Inicstiiw, by the
action of the bile and i^ncmicic juice, and pi>»ibly to some extent of the
Buocus eniericua, assisted hy various microorganisms, tho proteid* are larg«lr
diseolved and converted into peptone and other products, the stnrcfa n
cbanged into nugar, the sugar poosibly bein^' in part further converted into
lactic or other acidH. and the fats are largely ciuuUified and to some extent
saponiRed. TheM productfi as they are fonneil t)as« into either the lactenU ^
or the porljil blood veMo In, so that tltc oonlents ol the aniall inusltnc, by Lbs
J
CHANOKS I\ TIIK riCrKS.
S95
liM tlwy murh lh« ilM-«M-al vtilrc. mn litrgolf but by no meaiK wliully
4iyriMd af ihdr nuiriliouft constitkieDtir. 80 far as water is concerned, the
(MMbu (if wxicf into lh« small inieatiiic niaioiaifti tuvh a relslioii to the
■bwMiMi fn-ni it thnt the inlcalinal oonlentit ai the end of the ileum,
Ib^p mach cbanm<). "re about la fluid ax id Um duoileoum.
I
I
/n ihr hartje Inle*tine.
f SM. Tb« roDt«ni>. itbcfbrr alkaline or not, in ihe ileum now bcooine
mon diMinctl}' acitf. Thi*, hi)wivi-r, is not <.-nu»cd bv any acid accntiOD
lh« toucou* mrrabrnnr : tb<- n-iit-lioii itf (hv ititrelinnl waits in the Inrgc
M in tiM unall intn>linr i* iilLidinc. It Tiiii>.t. ihercfurv. arise from acid fer-
BWDtftUona going on in the omlcnls ihemwlroi ; aiii) thiit rcrmi'iilaiion« do
go 00 is showD DT the npiM-aninoe of mnrah gas as well aa hydro^n in this
portiud i>r ibe alimentary canal. The ehamcler and amount of fermentation
pKib«bIir depend largely oa ihe nature of the food, and probably aim vary
ui ilifli'reni nniiiinls.
Of ihe particular chan);c8 which take place in the larf^ inteeline we have
no very dctiuile knowledj^ ; but it is exee«dinKiy probable that in the volu-
MiiRouH cKcum of the herbivora a large amount of digestioD of a peculiar
kind )foa OD. We know that in herbivoru a considerable quantity of cellu-
luTC dieap|>e«ra in iiaaiint; thniuKh the alimentary canal, ami even in man
■CNM i> ui([eBted. 11 •eeius probable that tlm i«llLilii«e dit^'etitiun takes jiloce
in the large ioteHine, and ia the result of ferrueiiiatiw chaiiKen carrie<l out
l» neanti of mtcruurganiams, marvfa gm beiu^ one of Itio jiruducts formed at
the same time.
Bo tbb oa it iiuiy, irhcthi^r dipaitioi). pni|>criy Miealleil, in nil but complete
■t tbe ileo-cnx-iil valve, or whether im|><>rtiint elmnge^ still await the cbynM
in the lar^ >n(o>litie. one great characteristic of the work done in the ndon
u abaorpltOD. By the abstraction of all the soluble constituents, and (»po-
«MUIy by tbc irith<lrnHal of water, the liipiid chyme becomes an it approaelMW
lb« rMlitin converted into the lirm solid fceoi. and the color ehil\s from tlM
hripbt orBDge, which the irray chyme gradually assumea atUr admixture
«)lh tnJv, into a darker and dirtier brown.
The Fe«M.
t S8d. Thew contfiitt in the first pltu-e of the indi^stibU' and undigcMed
etinrtiments of ibe n>eal : shtvds of elnxlic tieaiie. hairs and other homy ele-
mrnls, mucti c4-lhiloae and cbluniphyll from vegrtnbk, and some connective
twiie from animal food, fmj^ments of dii>inle){rate<l muscular 6bn>, fat'Oclls,
and lint nnfrequentty undigested starch-corpuscles. The amount of each
miiat of eouree vary very larf^ly according to the nature of the foo<l, and
the ditfcslive powert. temimrary or permanent, of (.he individual. In the
•eotad ]>lsce. to tbese moat be addeil sutMtancca not ditliuclly reoo^ nitablv as
parui of ll)e fimd but <lerived for Ihe moi^ ]«n IVoni the aeeivtiODB of the
alironitary canal. The fecea ratitain niucuu in vuriable amoant, oo«ne<im«s
albumin, choleaterin, butyric and other fulty acids, lime and umKntuia sonps.
rolorinK nwUen, aixl inorganic talis, Mfiecially earthy pltospbatt*, cryftala of
Rnimiatu>-ntagne*in phinjiliaitv Itdns very consplcuouiL The reaction is gen-
erally hut ool alwavii arid. They ahio ountnin n ferment nimilar in ila aettnn
to pe|«iB, and no amylolytic fenm-nl similar to that of saliva or pnncreatie
jukv. The bile Mlli arc reproMntcd by a •lUall ijniiantity of ehohilir acid,
or aome product of that body, and loHHiUmM n very imall iiuaiicily nf lauriu.
S96 THK TISSUES AND U KCH A N'liiMS Of D10K5T10X.
Tli« glycin and most or all of ilie laurin have beep absorbc! fmni llii- iiitt»-
tine, iin<J llie cliolalic ntid has li^n parlly ab^ifbiKl aixl piirtiv iJi-onii)|kH«il,
T]i« fact that the fecea become " clay-colored " wlieii ihe bite is cut on frotn
the iRtestiiH alion-a tliat the bile-[>it;Rieut U at least ibe mother of the TmaI
pinneiil : and a ujieeial piment, which Una beeii isctlated and called atcrco-
Diliti, ifl aaid lo be identical wtib ihu dulMtance callet) uroboliii, nhich may
be formed fTtna hilirubiii. As ntlier apecUl constituents i^f Ihe feca niny be
mentioned ercrelin, a itoniewbat complex nitrogenoun iMtdy. whoee exncl
cJiemical nature h ut present tineertiiiii, and n^-nraf ((^H,N), a oitrogeDOU)
body which like indol is derivcil from the denom position of proteids by tneaos
of micniiir^iiigmv, nnd whicJi i* ihc cbiuf ciinite of the fecal odor, since only
a small ijunntity of indol remain* in the I'eciu. These odoriferous bodies Kte
derived d>recrly from the food; nt the same lim<> it is <iuiie poenble that
other siwcific ocloriferuii* siih^taiiceji mny be secretMi diKctly ftom the inles-
tinol Willi, wpi-cially from ihul nf the large inlmtine.
Ths Lactkalk aKI) thb Lymphatic Sr«TRH.
§ 28fl. We hftvc »^>on ih&t nbsor[>tion does, or nl least may take ptiuc,
frvim the slomnt^h. We have also staled thai a large al)8ormion, especially
of waler. occurs along the whole lar<>e incestine. Ncvertheleiei. it ts durinf;
the transit of fowl along the small intestine thai the largest and most import'
ant part of the digested material passes away from the canal, parlly into the
Iucl«als. partly into the porial vessels. Tlie portal veasels are simply paru
of the itcneral vascular system ; the lacteals, mto which we may at ot'tcp say
the greater part of the fat paasea, are similarly parU of the general lym-
phatic system, being in fact the lymphatic veasels of tlie alimeniary canal,
and esjiecinlly of the .^mtill inievttnc. The only reason for the special name
of IttcteHls is that, unlike the tymphntic vcwels of other part* of the body,
the lymphatics of the intestine contain iit timc:< a fluid of a milky white
appe-nnince. Hence for the Iwlter imdcntnniling of nb»orplion by the lac-
U-mU it will Ik' iloirable lo study nl some Inngth tho whole subject of the
lymnhatic syKtom. .
Tlii;! lymphatic v<weU mny bo said to bctfin in rainuto pawiigw. pomm^^H
six-rial clinractcn, known as Igmjth-capiuitrif*. Ilroadly spenkine th^^B
lymph -capillaries are found, in lh« mamnial. in all paru of the body in
which connective tissue is fiitind; and they have special conocrlions with
those minute spactu in conneclivo tissue which we hare already more than
once spoken of as lymph-»paces. Of all the varied functions of conneolire
tissue perhaps the tuosl important If lliia relation to ihe lymphatie ayetea;
in nearly every part of the boily eoniieelive tissue serves as the bed or origin
of lymphatic veasels,
TlKse lympb- capillaries, which, as we shall see, are Oequently arranned in
pleiuses. are continuous witli other passages also minute but of a ditterent
and more regular structure, the Ivmphalie veasels progier, which are gathered
into luri;er am) lar;ger veneU, all running like the bloodveisela is a bed nf
conneeitve tiiwue, until at bu>t nil the lymphatic ve«seLj> nf the body join
eitlier the great thominc duct which o|x?m> hy n valvular orifice into the
Tcnouf srslvm at the innrtiou of the laft jugular and subcliivian vciiw, or
the small right lymnnntic trunk which aimiTftrlv opens into llic junction of
Ihe right jugular und »iihc1avian %Tin«. The liitler course is taken by the
Ivmphaiioi of thi- ri^'lit vide of the head and neck, the right ami, the right
Mile »f the duet, the right lung and the right side of tiie heart, as well ns
J
TIIK LACTEAL8 ANU TIIB LYUI'IIATIC STKTKU. SIC
h^ »omn ««kU roDiing froni pnrt of th« u|>)>er lurftoe of Ui« IJrer ; all tb*
it«t of iIm Ivrnpliatic* iiiclintiiii: tliu luctenlt fall into the thoravir iliiiM.
Tfaa lyupluuc vwwU, while likv tlie rciiw ther join in thoircoum into
Utkvt and uirger trunk*, rio uut iiicmsc in mlibro *<> rn{iii]|y or w reguEarlj
a* 3o tlio roiiu ; tliry inmy run for twmc <lii>tiuic« wiiWit gmitly innvMini;
(a *m; and funln-r tbey, unlike the vt-ine. freely aniutomoec. fonning plex-
OMB*. Moreover, during iWir roiino llicy ciitrr into pcciilinr reliitions with
Mr%xctnrM known w lynifihutir glnM(]]>.
It Mill Iw ndvunUgeiifiH lo consider M[Mraicly the lyiu]>hnlic vcomI*
•>cH«r ihmn tW' lyiD|>b-<-Apillim«a, tbu lyniph-capillikricti tliciiie«lvce, and the
iTSxipluUlc gisnds.
TA^ Lymphtlie VesKU.
% 187. On th(M wo ii«<hI nut ilwell iii lenj^h aiam their Minictniv, in n\\
ilkl raped*, rwcniblew tbut uf tbi- veina. The tiiDnicic duct, uhicb in
hft* it< lower vnd wbcrv it i.* widimcct into wbnt in jfaimctimai ciill(<(t tbe
'^■^CMtaTM/un cAjrfi n dtnnifti-r of xix or »ev-cn niillim«tmf. hut in niirruwer
*^9*rr ii|>, may be wtd lo poMtw thr«« conlA. Tbe inner coat con»i»t» of a
k^cr of fuKir'onu qiilhflioid cvll*. not unlikv ihnof in ii vein bnt more •lon-
kod with n tviidcncy to lio sinumiH in oullin<-, ami of a iil«ud«r olutJO
• nina aii whieb tbeso mt. Thv middlo conl ronnisiH of fine bundlee of
V^fcaia iDUfcuUr libnw, which nrt- fitr the motil pari (li»]>riecd L-irculnrly but
^^Im to a rertain oxl«ni obliiiucly and even lun^tiidinally. The spaoea
neoa Uie bundk* of muKrnliir fibres ar« occupied bv connective limie
Datworki if elastic tibre;^ Tbe oiitrr coal, whicli n not well defined
Itbrr fr<>ni the niiddk> co«l on the one side or the connective tiaeue surround-
thi' duct <ru like uibtr side. ciinsi«ts chiefly of connective tisue with elaatio
aU, ft few mnacular fibres being somctimea prtaooL The wall uf the
^bofaelo duct ia CMcntially nmitcular. and fiom tbe wantiDeas of c<>iiiiwtive
%iwae aod of elutiv eloncnta b more tender, nionn apt to be lorn than tli«
^_ 'vall of a \-«io of oorreapondlng ilxe. >'un>erou(i valvea are present, thiv«
^H Ilk* th« viUrn of the Teiai' bein^ fohtin^ of tbe inner coat.
^V Th« smaller vtMcl* rvaenible in Mrudure tbe thoracic duet, the coat* being
^P of cnuDH* mon.- «k-nder. In the majority of even ftinaller lymphatic vtwcJa tbe
^ BKiKular fibra are abuudnut. Valves are etpecially numeruu*. and in oianv
of ibe renels. as for in«tance in ihcM of tbe nieaeniery, iuMt aliove each
InlYv, where tbe tube ia acuDCwIiat *w»lleu, the intiiu^ular littm, which cUo
winr* are chiefly diapoead circnlarly, run in variuUN dinviii^iuf m ns to form
■ eontncdle network.
Th« malleat Tceaeb, Rprineing from tbe distinct lymph -caiul lanes to be
inBwdialely deecribed, oouut of hardly mure than an epithelioid linine
iwtfng on a scanty conDectiTe-liMiie basis. Thu epiih<-li')i<I cells are still
flirifbriD and TCf[ular in aluipo.and tbe cnltbra of each vesw-l i* fairly uniform
tlHNl|A> OWiuK to the valves whieb are exceedingly numerous, tlien^ is a
giml leodency u> become beaded. Tbeso smaller vessels like the others also
annffoinoae ftvely.
k
LymfJt-tityiUarU*.
_ f 888. Tbe •malhvt lyntphaik vcsadH just deecribed might. fVuni analooy
witli thn btoodvaaelt, alniMl lie oonBidered as capillars- vetaeh ; but tne
Muno lynijih-caplllariaak given lo veaaels which joining and feeding those
just dotcTtlxsl jHoMW rttj different cbaracten. They aiv ooi the whole
laffcr in calibre ihoo these, swi dbtinclly larger tluui blowl ca^llariea;
ihey are ezeeediogly irregtilar in sbafw, and in their junctions with each
898
TriK TISSL'KS AND M KCH A.Vl.SMS nr Dlr.KKTlUN.
oUi«r fonn irrogulnr InUyrintlis riUlmr limn formal plexiiM* ; tboj pomem no
vtilvca Kilt) thvir only ciitil ia iin e|iithc]iiiin of n wry sinking cbaracUr.
].ilie the blixid cnpillnncs Ihoir >!lritc1ure is rcvmlcv) Kr the tiction of rilver
nilrntr. When n piece of lissuo rontniiiiiig IrKiph-uipitliiriw. f". ff., one
tskcn from the IcmlinoiiF portion of the diaphrAgm, if exsmtncd itfior proper
ireatnieiit vrilh silver nitrale. luiiiiorous epscM, od the «h<ilc tubular but
highly irre^iilur in form, joining; into un irrcgulnr IsUyriDth, nn? »i>en to be
liivei] with ii liiyer of epitbelioiil plates of u peoulinr kind. Kauh plalv or
cell, which ia more or teas polyj>onal or at least not dirfinetly fuaiform. \»
inarkeil out by lint« which are not etraiK^l »n(t even, but very uiarkedly
niDucuit, the aeveral bulKiugBof one cell diive-iailiiig into th« depreasioiM of
iu iietghbora nnd vice versa. Such epitheliod plat«t) of sinuoui outliiM. or
Hucb u iiiniiouii ejtithelium, as we may for brevitjr'ft suktr gay, h characUriiitic
uf the lyiiiph-capillarice. A I yinjih- capillary ti \a fuct m«r(!ly u ajmce or
areola of cunueclivu tiifiue, aoiuetioiut more or Igm tubular but frequently
irn!|2»liir in form, lined by a ^ugle layer i>f fist, trauHparvnt, nticlmtM
vpillivlioid plated, each of wbicli potwcwca n remarkably oiiiuoiia uullim*.
Tlitr lymph -oapillariea onasturauiHj freely with utlicr and upt-n into or join ibv
MniilliM ri'^^^jliir lymphatic canals, which, many of ihrm »malIor innn the
Ivniph minllaiiea, are dixlinguiitlicd from tht^f^c by their mure regular dbpo-
sitiiin. by ihcir vpithelinid platen buiitg fiisifornj with very little sinuosity of
otitlinc, unci by Ino urotuuce iif valves.
Tbu lacUial riultcle of a vilhiH (§ 'Jti2) ia such a lympli-cnpillary, more or
less tubular in form, or |HTlin]w club-shap«d and sometimes bifurcnte or
branched, jiIdcik) by it«clf in the midst of the reticular timue of the villus;
ending. »r a;* wv should pc'rhn|)» say beginning', blindly ntuir the apex uf tbe
villus and joiniiii; hclow by a valvular mouth a re>;ulDr lymphatic caaal
forming part of Uii- network of rp^ular lymphativveHcU with which aa well
at) Kith lympb- capillaries the coiuieotire Ltuiue of the muoius membnUM i$
Aimiaheil.
Id oilier paria of the body where conueotive tuiue runn, lympb-capillarin
are more or lo« abundant, all i>aiuiiig their conleuta.oD to ilic inon n-giilar
^mjihatic cannlit. In certain parte, aa
lor tnslanoc in the c<cntral ncrvoot sys-
tem, the smaller bloodvcsHla are sur^
niundcil by large lymph •capillaiiea, or
by regular lymjthalic vemels, in tbe
shape of tubular sheaths. In these
casca the lymph-capillary fonns a aorl
of ballon jacket around the artery or
vein vhicli. covered with a layer of
ainuoue epithelioid plalee, liee in tlie
middle of a tubular B)>aoe lined with
amaller tdnuouu plates. The plaiuna
which exudes througb tlie walla of tlio
liluodvessel pnwee accordingly at ones
into the tubular dpacc or interior of tlie
lympb-capillary, wbcncu it U c-arriod
awav into tbe regular lymphatic canaU.
Such an armngcmvnt iit opokm of na
a " pcrivax'ular lymphatic. {Vig. 121.]
$ 289. The lympbcapillarim may in
one «e«aa be n^Hed a» the t>edn-
ningsof tliv lym[>luitic xystcm ; tbev arc the first lymphatic passagra deli-
nitely liiK'il with a continuous tipitfielium. But lympo exists outride the
triD.m.
Iiniviwrua I.TarnAnoL
A, annii «r <«V««- . il. »netj tmrn llw
limlD .!>. iciliu«i^Flyoi(i>>«|aociLiuiilLiVi
i«r«liiw. «dlL br 8U1IIUEJ.)
TETB I.ACTKALS AND THE LTVFnATir STSTBX.
8t>8
I
I
I
kpillariei. In U^atinjc of conneotive (iwue, $ 1'^. we ninra lluio oot-e spoke
of tbi> MjHicctt belweeit t)i« inlerladni; Ixmiliee of 6l)rillie as Iyiu|>li-Hpa(.'e8 ;
and indfuil lliejr are dtiriii); lile i)ocii))ied by fluid wliich mnr be spoken ol as
lyiii)>li. It in fluid nliirb has in Hume wav »r •^'tlier pawed uilo them trom
Ui« Mwid Ntream, tbroui;!) tbc walla of tlie <-a|>illari<» aud utber minute
bloMlriMclH. Wit Hhall H|>i-ak of thin [laatiafce an a procea* uf traaautlalicui
and vluiU muniilor itn iialurc later on. Many of the lafx«r of theno iipaoea,
th« Mwilie of areolar con HOI- li VI' lr»ue, are c»mplet«ly liiiei) hv enitlivlioid
plttra irith Htutioim <iiuHnn» ; th^tu) arc in faoL l}'iui>b-<'npi)lurir>. But
many Kpiicvn, twpt^'iitlly lliv mnaller onw, are nut ao lined ; thctii! liv out-
aido the Irmph Mipiltarint. NcrcTthdlw* tht'V conlaiu Irniiih, which nincli*
ing tbcm l>r traii»iidiui»n tliroiigh tbi* walfn of l)w^ lilooflvt-iucli, Mreams
(rota tliem in unnw; way or oilier into ihi- lyniph-enpillariM and no into the
other lymphatic vctacls Colored lluiil iiiji-cied by ni«ai» nf a line syringe
into tiKH speoM soon findi iw way into tin- lymphatic*; aod bmdce, in the
vast mnj'irity of caM*, a rcrhiin niiinbor ■>f tJieM anacw nlwitys interreiie
between ttie wall uf (he<-ApiIlHry or other imall hlooavesMl from whencs the
lymph cornea and the lymphcapiltary to which the lymph gov* ; the lyiuph
mual have some iiieuns or utbor of iiaMiiig fri^ni the spaces into the lyrapb-
capillary. It is of course ixweibie that the lymph transudes from the
lymph-«pa(W into the lympAcapillary ibroufih the oonliuuous sheet of
epithelioid plates, in the same manner that it transudca from the blooil-
capillary into the lymph space throiJ|;h the similarly cuDtinuouH wall of the
ai^>illary : but there are some reasons for ibinkin^ (hat. at placea the epiihe*
Itoid liniug of a lymphatii- capillary may be imperfect and 00 allow the
interior of the lympb-eapillary to i,\ivn out tnU> a ciinne<-ltve-tiMU« upace.
It will be remembered that, in the caae of Home uf thcnc tipnci'^. n con-
uectire-tlMua corpuscle may be found lyinKxm tliefacenf, or purtlv imh-rltlivl
in, <iue of tite bundles irhicb form the wails of tboajMice; au<l n <
the npaceappean iis it were iiojwrfet'tly liiieil nilh scattered fl.i: <
may |K-rha{M bi- regardcnl u-i imnBiiioiiiu forras iHttween an oriliniiry lininchMl
Conn4i!tive li.iHiie corpii»cle and a HinuntiK ejHtbiilioiil plat<>. We may per-
hap* iV|Enrd the epilhrlioiil pljiti^ as a diflihivn tinted eonn<^ctive-timuv eorpiiitclc,
«rho»e «inno«itii3< of giillinc nrv the rpmainn of ila pn-vioiMly branched con-
dition. If ihiN be «n we may cmiuder tl>e lymph-i.'«])illary as a difTeranCialiM)
CODiWotive-tiwiiv «pacr. and c<miMy|uenlly may fairly expect that tbo one, if
it doe« not IW HuggMtcd actunltv (>|N-n into, should bo at all events in oa«y
communication with thf otht^r. \Ve i<cem juMilkd at least in concluding that
the complrtely lined lymph -capillari<* draw llK'ir supply of lymph IVum the
iDcumplelely lined oonncvtiee-timic spaow.
We may probably tfu a step "till further. Many of the connective-tissue
oorpuacles are iiultediktl in. lie In PAviti<'i> exciivaled out of, the cement sub-
atanco which unites the tibrillie into biindlM and soinetimeA joiiid the bundles
togetbec; iu »ua>e situations the corpuscles are similarly inihtyldeil in a
hoouweneous ^'^und subBtanoe which has not become dilfei^ntiated into
6brilbe. The cavities in which these corpuscles lie are, like the eorpuactoa
tbemaelves. branched and K«nerally flattened ; they appear moreover U* be
geoerally lar^'er than the corpuscles so as t<i leave a unall space whicli oan
be oceujued by lluid, Wliere two oorpuaclee lie near e«eh other their Kpacta
niay, by ine«iia of the bnachea. commiiDicate ; and in aume situations, as in
the bixly of the (viruea when a number of flattened corpuaclus are imhodilod
in the lamina of jfronnil aubatanoe which uuiten each Iwo adjaoenl {wrallel
(or rathfv cmiicenirii:) larainte of filirilUled bundli'M, the MCricH of cavillM.
uniting hr their brnnchea may be reKUnled as constituting 11 lahyrinch of
ptUMi^e^ largidy but nut entirely Blled by tlie corpuscles, apiuy bang lelk for
400 TUB
riSHS OP D1GB31
MOPK HRiount of fluid. That fluid iro iim-iI liardlir My is lymph. And
tboUEh tlu: view in not nac ndiiiitltH) cm nil Imnda, tluire &re reaaoDA of some
Wflignt ><»r thtiikiiig tliut tbcw cavitir* lieluugintf Ui tbu corpusols open out
into till! ci>niii.'rlivi''ti'«iiir kiukv.'' jiiKt trviitc^I of ur cveu mora di racily into
the lytui>li-i'Hi>iIlnrin>. When n pii,'CP of CQiini-ctiv^ lit«u«, such Tor tiuUDoe
M tliat Iving WlHc'vii itic radiuiing iMindlfo uf llu* titndoii of tliL- dinphrajcra
on tbe pleiirnl »ido in irrAlcd in it luirliciilnr wny, the naiiilt u. what in cttlled
B " nogRtive stniitii)g : " tbu matrix is xtHiiK-d brown hut llu- connuclut viu)
cnvitim are left un»(Aineil, iind appvtir aa irrrguUrly liriuivliud clear jntdiet
»liinding out in contrast with the brown matrix. In Huch ii pr<;(iHration
many of theec clear Bpncc« arc »ceu to abiit upon and amMmiily to low
tlwmwlvcs ID a neijjhboring lymph-oapillary, which al«o iilitnyii «tanda oat
in contrast to lh« mntris, appearing as ti eloar »pao» markml wilb tbe ainuona
outlines of its plates.
Wiihoiil iofisting too loiich on tbe argument dmvn from tbis nvgntivo
Staining, and M«ting rather on tbe facts pivvioufly mentioned and oa goni-ntt
eonside rat tons, we may probably conclude tbal all tbo spaces of conneclivt*
tissue, including lh« cavities of th« corpuscl««, form a labyrinth of puswgea
wliich id to be constd(-red ba the r«al bei^'nuio); of the lympbalit.'s, and that
this irregular labyrinth is iu some way or other in fairly free mmmunicatioa
with the more regular bm Mill labyriiilbine lymph capillaries, tint-d by a
definite epilhelioiil lining, and that from thence the lym{^i pasee on to the
ngular ajtd valved lymphatic eanaK
All over the body wherever bliMMlvcueb go. coiioeotive tiwue and lymph-
RjHioea fro too. Cvrtaiu parts i>f the plasma of the blood pueing througb tbe
trmllso? the bloodvuMuls becoiuci Ivmph in thnte )yuipb-s|Mces. As such it
•oaks tbmugb not only the hundtva of gelntiniferous fihrillie of tbe con-
nective liiMue it«elf, but aUo the bawmeat membrane and ao tbe epithelium
of the miiCDUi' membrane and ito glauda, ihir niulriated muscular fibre, the
ivrcolcniiiia and munclu vubKlancc of the strinlcil tiliK, the nmrileoiiiia and
CODteuta of the n<rrv<^'flb^■ of n<trvLii, in fact the clementn of all tbe ItMuee
which an; :>u|>plicd with bloodvcstcU. ^lorv than this, Ivmph goes where
blotKtvnwIf do not go, and in th«« rituiilionK iht^ vahxi an lympb-iMMagea of
tbe cavitim of the corpuscles aeems most i>irilcing. In thi- i->iriiea for initanoe
bloodvoBeb and definitdy conatituhKl lyiii|ihatic vcwcU ccaao near the
periphery, and the greater part of the nutrition of the ooraea (beyond tliat
eflwtoti by what w may <-nll mcn^ imbibition, that is hy the jauuagc of fluid
between the molecules <.•( tbe actual subst^mce of the tissue) is earrk-il on by
the stream of lymph through the corpuecalar cavities. In a similar way in
bone lymph lin<l« its nay from the hloodvesaels of the periosteum, marrow,
and Ilavirsian canals through the very substunoe of the bone hy mmiis of
the labvnnlh of lacunw and caiialiculi. And in cartilage we hare raueon
to think that minute pa&sngos iu the matrix facilitate tiM- transmission of
lymph from tbe perichondrium through tbe body of iho cartilage from carti-
lage cell to cartilage cell, lar tnore efficiently than if its iirogrcw were left to
mere imbitition. The suniewhat peculiar relations of t^ lymphatics in the
oenirnl uer\ous ayf.tem we shall consider when we oome to treat of that
•yMem. Ueauahile we have said enough to form a general idea of the
anatigeinenla by meaaa of which tbe very eleawnls of all the ttasuea are bathed
with lympli, and by means of which that lymph is carried back from tbe
eletnenis of the tlHtiea along irregular and regular lymphatic channels lutclc
to the bloud fn>in whence it origiiiiilly cAine.
£ 290. Thr trrmu enritiet. In the mammal lymi^i-spac«fi ara for the
moat part minute and microwonie ; but in some other animals ihey may
attain cunsidf rahlv litu ; in tliv frog for instanoe iu which lymph-capillariea
THE LACTKALS AN'D THK T.VHrHATIC SYSTEM.
401
I
I
I
nmf lyinplindi.' vkswIr nr« scanty, tlio Inrgn 8ul)(ruU»«i>u8 ep«OM which ar«
diitcluMs) wImtd the «htn of i)i« back k <;ut through sn.> in mliCy lymph-
IpaoM liiK-d by »inuouB epithelioid plucs. Eloth id th« mnuiini)) nod otlier
Miimnlit oorUiti large mviliea, kaowa ns serouB i^avitiee, ettch as the pcritii-
iiukl. ittriviirdial, pieurAl, and olher oavitice, nitii^t be eonBidered us parts of
tbo giMKTal lymplinlic Hyaicm. aud, indeed, the "wroiu 6iiid " which they
COiitHiit is in reality lymph. The sulwracbnoid space nrrotindin}* the brain
wul spinal ooni may also perlm|i8 be recpuxted as a part of the lympliatic
system, btit this and the oonlain^^l cerebro-spinal lluid we shall cDnaider in
connection viih the central nervous wyMttn.
In the abdomen iif the frog, <m each side of Ihe Tei1«bra1 column, behind
orabot-e, I. r.doraal ti> tlie )ieritoucat cavity, I ivs a lur&e lyinph-Hpaeei »|K)ken
of as the cuttrnu niayita Ijimphatiea. the cavity nf whiob in separated fmui
the peril4Mi«al oAvily liy » thin mi'nibranoiM nhcet ooomBtinj; of a mi^diun
basis of coniKiitive tiwui; i-orcrvd on the [tcritoncal dd« by |>«ritoui-al t^piilie-
lium aiid on ihv elHlcrna »id(^ by lymphiUir epithelium. TIic lAtl«r consLtM,
as in a lynipbatio capillary, of tint epithelioid phtint with MJimouR outUnet ;
ttie former is made u|> also of Am epitlielioid plates btit thew,' are niorv or IcM
polvf^mal in sbnjM', and have outlines which are nut ilistincliy sinuous. If ft
piece of this narlilion, nl>er hein^ stained with sitvxT nitntv, be sprcttd out
and examined either with the penionea) or with the etstcnw aide uppermust,
it will he »r«n that in each case here and ther« a group of colli Ksuming n
triangular form appear to converge to or radiate from a centre which eomo
tinea, avpeeially on the lymphatic side, is a mere point but SDmotimcs is a
larger or » smaller hole, which, in other nonU, is an oriKcn or ttoma. some-
linua elmcd h<il sometimes more or lees open. On the peritoneal surface the
Moaia is surrounded and Kotrded by a crown of what appear to be small
granular cells placed at the apices of the convergioK epithelioid plates, but.
which are held by some to be the displaced nuclei oi the epithelioid plates
themselves. ^Vroand eanb stmma which is in reality a perforation leading
ftwa the peritoneal cavity intu the Q»ierna, the oonnective-tiasue baas
between the two efHthelioid layen ia art«ng«(i In a concentric manner ; tbe
whole arrangement serves oa a commiini<-ation from the peritoneal cavity
into tbe cisterua, and by these slomaia tbe perit^tneal fluid passes into the
oiatenia aud so into tbe general lymphatic syxlem. Owin^ to causes whicji
«re shall study pr«Aenlly tbe vMlcnu of the Kuutll Ivmpbatic ressols and sudi
scooes as the cisterna are ccmiinually li^ln^ draincti br tiie vascular syslaiu ;
tne ciflema is continually tending to empty il«vlf anti »a to draw fluid IVom
the |)erilt>ueal cavity thniiigh the etomata. In the fnmale frog tbe amall
granuUr cells encircling ihir ilomnta are, during the hrn^ling rwason provi<Ied
with cilia, the action of which incrmscs tbe current from the {H'HtoiK'um
ihnHigh ihr donia into the oistema.
In ihc niummnl Mmilar stomata place the serous cavities in connection
witli the lymphatics of the walls of those cavitit*. Tliey may be ntatlily
teen in (he tendon of tbe diaphrngm. The peritoneal memhrane of the
mammal ns of the fii>g oonaiata of a single layer oJ' flat roithelioid ]>■»(«•
lying on a connedire-tiasue basis ; the plates, smaller tlmn those in the ftog,
are polyfronal in form, and their imllino is not sinuous. On the tendon «
lbs diapliracm the epitlielioid platee over the radiating sjuiocs, or clefts
between tbe radiating Dondlce of the tendon, are smaller than over the bun-
dles thenaelTee, and along the lines of these radiating interteudinous spans
may be seen stomata, orifices guarded by small cells, similar to but smaller
than and letu conspicuous than tho«ie ju^t deecnbed as seen in the frog.
These stomata open into tbe lymphatics which are abundant in the connec
live tissue lying between the radiating bundles of the tendon of tbedin-
20
402 rna tissues anti
[OESTIO]
■ilirnfini, mill through thum this fliii<) of the pflritODMi] cavitr |M«W ftf
idUi (hr lyinj>liiilir« of tliu dinphrii^'iii nix) oi itilo th« eoMral iyiiinhftttc ! ,
t«in. The mor<-iniiiI]> of the ditiphnighm in brdittniie. of which we nltidl
have to tj>«tik prcwutly, ^rrnlly uHiel the tlow through thi< eiomata; itnd
ewn |in«MV« moremento ot the ttinjihriizm nre cBectiinl flir this |>iirpo«». If
a (|Uiiiitity of injection inatcrinl. »ueh no » filution of Uerlin btu«, be injected
into the peritoneal cavity of a livins animal it sotru euten into sod injccta
tite lymphatics of the diaphraf;m, and n similar injedion may be oblaineil
ID a dead but receutly killed animal by pladuf; the aDimal with ila bead
dflwnwurd. injeciiDt; the coloring msiter into the abdomen, or even pouriog
it into i)ie hotlow uf the diaphratcm, end then imidu^ii)); movemeois of tli«
diiphrajcm by a rhythini<'ally re|>eated artiBoial respirntinti. Not only ool-
OTM flaidi but eolored material raerdy ni*peaded in tluid and ounh ihinga .
the globtitea of fat in milk, or eveti reil hl<HMl'c:(in>u)K'loi may tbtis lind liieir '
war from the p«rll(»i«al eavity into the lynipbatiw of the duipbT«|[m.
IdoccjcI, if u \i\wt- of the dtaphn>i;m i<f a rM'enllv killts) nniiniil ))e $trecoTied
out Hi|<t milk p.iiiml npi>n it, the fill glohiikit »l milk iiiiiy b<- >wa with the
aifl of a leiut or inicnwcofw to dMipjiear thmiigli tUe i<tt>iaata in a tiumWr of
uinutv vorltccit-
By MDiilnr <t<>mata iIk- pleural cuvity w pnt into ciiminunication with ll
Ivmnhnlivs not only nf the diaphrnfoi (on ibi pleural enrrncc) but nim <
(W Iting*. and to n smaller extent of the tJioradc ntills, and during the m»vo-
nienl« of the chcet in breathing the ponleiita of the pleural cavity art) ooo-
tinunlly Imni; pumped away, partly into the lymphatics of the lungs partlyf
into those of the diaphragm and elicst walls. In a similar miinoor perinir*'
dial fluid passes away from the pericardial cavity, and the fluid in other
■mailer serous cavities such b* that surrounding the leatia, pMsws away from
the reajjeelive cavities into the general lymphatics. The quantity of fluid
in even the lar}^t of ilieee cavities i§ at any one time in normal eonditk
very Hnall, but that Suid appears to be constantly renewed, old lltiid pa
away to (he lymphatic system, and new fluid taking ita place. I'he ser
nivili(«. ili^refure, are to be re>,'ar^ed aa expanded initial rmervain tfov
which ttn well no frtiro the lynii>h-rnpillari(» and lymph »pa<!es of the timic
the lymph stream in omtinually being led.
TUf Slrudiirf of lofntphalic Olantt*.
S S91. Solitary fotlidv* a alt I'r<irr'gpolfiie». All along the small inteettoe
and at various points of the circuinferenoe are found, jtartly in the suhmu-
(VMi* tintue hut reaching up to the surface of tlie mucous owmbrane. small
rouniieil bodies, of (he nixe of a small piu's liead. more luiiueriKia ]>erl)ai)s in
th'- lower than in the upper [utrt of the bowel, uflen called " solitary glanda."_
!Pig. 122.] They are not glands, however, in the sense (^ 209) »f being'
iivolutionN of the niuenua membrane, and it is better perhagw f > speak
them as tolitnrif j'otlif/a. At the free border of tJie small iniMtiite, oppoaiM
to tlie atUtcbmentii of the rmwentery, tltc mucous membrane contains loi
OTkl patcliMi, Peyer's patches, phicwl lengthways, there being sonte twcotJT' i
thirty of tlicac; ihcy arc nhMt iiiirucrous in th<! ileiin) and dii>npi>ear toward''
the nuodenuni. Kach patch i« pructically a group of tulitnry folliclvit. and,
indMd, tli«M jiatches an- sometimes spoken of ns a'jminaled fi/tlielf*. In the
lanra int«liDf> <9p<%ially at the cncum, and in man particularly in the ver-
miform aptieiidix, solitary fblliclM are abnndant, but here iher lie tiholty in
tli« submucous titnuc below the rauscniaris muoosn. In the Kfininch iiUi, in
young pent>le, there occur in (he muooos membrane, geiieriilty betwe«u xho
TRB LACTEALS A»I> THE tYMPHATlC «T8TBM.
tivAHtlM of tliQ ^Inuils. strui^turee vhich itre very similnr to solitnrr folliclw
am) wliich are 8i>ni«tiiue« <tt\{eA " lenticular gUiuIe."
rt
, .*!-J-
^^*^^'
iiai-nnN av t»« Uxirs niEiiiiaii * SuuTiHr Pi>ti.ii'i.ii cm l.vspiioit< Nium I.K. K'tiiUT.i
«,ini>lilltorilMnoiliilDwltb tlic Irmpliaiil l!HUiit*nlT blltoxrkf rrom UwiHilari ; t% epitbelliim
til Un luMaitue; t. tIUI : Uk>1[ •pl(h*liuiM In (itrllT lin>l[«n away ; -I. envl* <■' I-lolsilKllio.l
A wiliury follicle cudnwIm nMCiilinllr of n upherica] mam of fioc mlvnoid
lifiii- till- m<wh<¥ f'f wliich are crdwilci wilh luiiciii^ytc In tin- interline ns
we linve wrn 'J "J6ll; llK^cnoHOciivctiMiir lying between the «pillidtiim above
Mxl the muwulnriK miicoxir beloir 1iiu> n rcticuliir arrangement and contaiaa
l«uoocvte«; but in lh« follicle the network i« 1in«r, clocer, and mora reij^'i'*''
than eWwhera. the me«he« are atinnst completely fillexi with teucootcf, and
the i>nhmcal mas breaking thro«igh the miiHculam mucosie reaches eomo
w«T uonn into the i^ubntiicouB tiwuu^ Over the surface of the follicle, which
bufgee somewhat into the iiilerior of llie intcetine, villi may he preKTit, hut
the ^'laiidn of Lieburkiihn ara pushed aside and are found only at its circum-
ference. Into this mass of adenoid tiaeue one or mora sinall arierim enter
Hitd break up into a capillary network the blood from which is cnrried away
hy one or more small veins. Around the mass there is placed a more or leea
well developed spherical lymph-spMee. lined with sinuous epithelioid plates
and ciinlinudtia with tlte neighDonoK lyinpbatic vessels. This lyniph-ajnoe
or /yKipA-n'wujr as it is called thus toraif- a hollow jacket filled with lymph
rouiiil th« spherieal aiass of adenoMl tissue, but is not cnmpleie, be iii); broken
by tite entering and issuiiiK bloodvcaaels, or by imperfect |iuriitton« psH-iuj;
Iroai the tissue without to the ade»i>i<l tiwue within. Thi? blo'xIvewniU mil
bridk* In uuuttioii are covered by a layer of ejathelioiil plnteri cotitinuous
with thai lining tl»e outer wall of the Jat-ket, ns also with the one which nmre
or luM <!oiuplelely iiiveMU the itiuer maax of adenoid tiwue.
The k^ticocytw whicJi occupy the iue«hee are of dilfbreiil Hixeti. Kumi^ are
I large or al'moM lu. \aT^ ait while blood -oorpiiac lea, from ntiich indeed they
' 'elly ditr<cr in the faet tliiit their nuclei nxhilnl a nuclear network which a*
have aeoii (S 'i'^) i* ap|inr<-ncly not prAtciit in the while eorpuitclf^ of the
blood. Tho majority, however, are much smalW than white hlonal-corpuMihv,
iheir emallni'w being cbieily due to tlw imnll amount of coll-MubntaiiMi sur-
4M TUIC TISSUKS AND MKCUAXISMS OP DIOESTIOX. 1
miiM)iBp[ the nacleus; io Bome only a inctv flim of oclI-ifibslRnm I'M) bt
(l«iecle<l so thai the nucleue appcare aliiin§t as a so-called " tree " niiclrut.
Many of the leucucytea nmr he seen to be uuderKoin); karyoinitueU. iiMli>
cniiiif; llial they are luultiplyiiig bv divisiou ; and, indeed, there are inmiy
rcANOUH for thinkint; that in the aiteimid imue of tliese foUielea and other
iiimllar ntruL-turef a very conttiderabic iuiilli|tliuati<in of leucooyt«s lakee place.
Many of the leucocyleA of Ihette rolliclt^ exhibit under favurnble eircutn-
HtanctM amieboid niovement*, and the nnaller leucocytes, indeed, even the
rinalUwt, iieHiii at tliuea m active a* the larger i>ij<«.
A tulitary follicle then may Iw contidercd a* coiiaiHtiiig in tlie first |>lac«
of a riiunclcd capillary network fed ami dniiiii-d by xmall arterie« and veina,
all ^uppiirtcd by a minimal anigiint of ordinary connective tintue. lit the
wonil jiln<% tbe intentticM of thi» vascular ni-lwork arc filkil up uilb ade-
noid tiMue the fine mo>hc« of which arc crow<lc<! wilJi IcucocylcH id' variable
but on the wh»]c «niitll sixe. I.iistly the nuimicd itinM thux ■■»n«iiiuu>(l »
aurrouniUxl l>y a lymph-sintjf. the fluid of which on the oav hand balli«i the
mass and on the othir hand ii frco to pan anny into the lu-ighboriag
lynnhslic canals. Ait the blood streams through ther capillary network part
of the plasma poning through the capillary walls becomes lymph in the
Dieshes of the adenoid tissue. HcDce, after probably acting on and being
acted ou bv the leucocytes, it paasea into the Ivniph-Kinus and to away into
ibe K«DeTal lymphatic stream. lu all probability the lymph-siDUs is chiefly
fillecl fVom tlie fluid thux coming from the ndeooiil tissue, bo that a inaiu
current tlows from the lymph-einuit int>i neighburing lymphaticfl in all diree-
tjona; but it niav be that the lym)ih:iiuus ii> partly supplied by the
lymnbatics around, ho tlmt wimc nf the lymph from adjoining slructurev,
while tlowinj; in the .liniis around the adejioid iiMtie, li aubjected to the action
of that tiiwue. In all prolmhility loo the Irantiil of niatiriul from the hluoil
to the adenoid tiuue iv accompanied by a reverse currant from the adenoid
tissue to the blood, so that the blood in poesing through the follicles not
oi>ly gives but also lakes.
Since multiplic-ation of leucoeytea anpeant to be continually going on in
the adenoid tissue, and since the folliclea do not increaoe indefinitely in size,
some of the lencocytes must disappear. There is every reason to think that
ifaey jHiss away into the lymph-sinus, and so joining the general lyiuph
Stream become the corpuscles of the lymph of whicn we siiall presently
speak. If tbo central adenoid luass is, as some think, inveated with a con-
tiuuous eoat of sinuous epithelioid plates, the leuoooyles which leare the
follicle must pass ihrou|<h the eout in the same manner that the white cor-
piieclea of the blood lui^-rate through the walls of the bloodvesttels ; but it
IS more probable that, an oiheni think, the eoating is discontinuous, Uie
s|)acesef the adenoid lUaue "jieuing freely at inter^'au into the lyniph-HQUK,
and thus atfordinn an eoiiy path not only for the leucocyto, but alao for the
lluid.
The lenticular ){liinds of the stomach a]>pcar to h>' only Ins condeii»xl,
let* completely arrangd) mantes of adenoid twsue . and, »« wc shall sec hero-
after, uuall miuM-H of adenoid timiue more or lew cimdmsi'd, more or Iraa
trauaiiirnied into definite follicin, arc met with in various parts of the body.
^202, .'I i'ev'i-'n juilek u, as the pbnue "ogminntHl ^'Imid" indicates,
merely an aggregation of solitary follicles. A well-formed Puyor'a patch
con>tsl« of a variable nundier. in man fifty or even a hundred or fewer, of
Military follicUw arranged in a single layer close nnder tlie epithelium, but
Mtrvichiug down into the submucous tiiHue, the distinction of which from the
tuucDUs mombnino proper is to a great extent li»t br tbo breaking up of the
muMularis mucoacB. Between the censiltucnt follicles glands of Licberkulin
THK L*CTKALS ANU TQE LYMPUATIC SVSTKM.
405
^
' fanid CBClroliug ihe (blliclM aod villi ptvjoct frgm the tuirfacc, whih)
WtwMo Mid bolow tbo Klandt l>lMM)vt«»el8 nnu Ivniphnltcn ntv nbunittint.
Ov«r cNcli rollicif iKitli gTniuk wiil villi an? absent, ko tKnl thn iipjwr mirfucc
of th* fullirJo If ill ootitiKt nitli ilii;. (rfiithvlium of the iDlo^linu. wtiicli is livm
•borMr ■!»] B»ai« cubiotl ihm olMwfasre.
Esoh follicle OOiMUtfl of n Komevrhst spherical vnscuLir maee of udcnoid
UaDt, forrouDdnl nii>rv or leu completely by n lymph -einuB; in fnct, the
tCratfara of tn/cb of thr»o aggivf^lcd foilicln r«ficat;« so completely that 'if
m Hiltimrr follicle ihnt tbc same dcscnptioo and diacossioa will »rTc for both.
i 893. KympJtatie gland*. If the strocturv of a follicle just described bp
born* in mind, ibkt of a lymphatic gland is niiiile more c»sy ; for. tu a
Pr7(T'» fmtch is a mere acRregBtioD of otherwi^ uDchanged follicles, so a
lympliatir (.'land is a ooUwtion of aimilar folliciee d i lie rent ia led into a com-
fmet aod stMncwhat complex or^^.
A tTpkal lymphatic gland has, though the form varieti a i^ood deal, tbo
ihapt of a kvdney [Fig- l'J31, in so far at all eveoU that n more or lufl
flMvcx aide can bi? di&linguijWi from a canca^'e Hi<le in which is placed the
hilna when Uie btoodTesaels enter luid mav ; from the hilua olao isue I jui-
fhrtJB veaiek ithioh, nnce they carry lymph amijr fram tba gland, are
calkd efbnDt lymphatieB. The afferent v««neU carryioi; lyuipli to Ihe gland
paai into the ([laod in a Mattered fushiou on the convex »ide.
Itia. vm.
PuwBjJWtif Sbctki.t or LtxnitTK (luiii-.
l.t.1. rm*M tiraptaMBt: C. Donlod mbiUMOT: JT. raUcatulag cmdioTMida)-
t: I. •.Irini^^noi: «, DbromcuM nadlBc uaibotalB, Ir.. Into ttia tutalaMVcf tk*
Th« ^land it inTcwti-d by a capsule of cODnectire liaue. containing in tbe
■MB of uiuny unimnl> a very consiilerable number of plain muscular Cbrei.
Two layui Biay M timn be distinguished in the capsule— an outer layer of
coMfBcr tad an inner layer of liner couneetive conneclive lii^ut;, it rich {>lexuB
of Imphatic veeaels bciof; placed betnecfi the tiro. Froni the oapMile a
nu&iber of partilJoua or tratiMiI'r, BtartiuK from various puinta of lb« turlkee
406 TB^TfBSTOnwrnfWWffrW^fnffBfWTW^^^^
anil oooN^ciug, like tli« capsule, of vliisuly iui«rw(iveii biiiiilii'j of conoeciive
lifMiv itiixiil ui> n'ilh » variiibli! numlwr of plain miiw-ulmr likrei. pwn Inlo
the g)an<l in niUrvcliinn convitr^'iug lownnl (Ik liiluti. In tlicotilcrorcirfum-
Ibrciiliiil jiart uf tliv i^lnml ihttc lrnb«culBo sn^ Inrgi--, run in a Mraigiii
dirvclton, arv Imt lilllv limiidi<'il, iitid iin- r»^ iirrimgi.-<l ihat they cat u|i thi'
jDiitiT port iif thi? gland infi ii i)iiinlM;r of c^hnmhuni, having more or lcs« th*-
pbrm of Iruni-atvil pyruinitlii, ci>iiv«rgineIo or nidiatiiiK from the ionrr purtion
*tif the glniid iioitr tlii; liiliir. TboH* cnamlMtiv have bc«D cull«d i/iw/i. ariI
CODfilitUie togeltivr the mrlrj: nf ihc glniid, iho iniMrr porlion being called ibe
mf'UiUa. On rciic4iing tbe medulla l\iv tralKvuliD, lliAoouneof wbieh, a>
Kc have just «aid, is in thp cortex, on the whoio straight and uohraDchul.
rapidly divide, brcmiing thinner and more slender, and. running and joining
together in all dirvctions, form an irregular o|)en network, giving nse to a
lulivrinth of pasNiges into whicli the alveoli of tbe corlex o()eu.
Tbe Irahecutfe. in fact. Blartiug from tbe capeule divide the gland iuUi a
Dumber of spacett which in the cortex are arranged in a re^ulnr manoer anil
have the form of cunvergiiig chambers or alveoli, comniunicutltig laterally
ttilb each other lo a small extent, but which in the medulla rajndlr diminiBli
in ai^e, and, opening freely into each oilier on all eidea, form a labvrinth.
At the bilus tbe medulla cornea to tbe aurtac-e of (be gland, but ^Itewliorv i*
separateii from the surface by the cortex. The number of and regularity of
diviniiin amimg the alveoli, and the sharpneKa of ditiiinction bctwwu tbe
cortex and tite medulla, difier in the glandn of dilfi^reui. animal*.
Each alveulns of the cortex eoDHiiilti in its itintrtil |uirt, oontitituting alxxil
two-lhirda <>t more of tbe whole chambtir. of ■ moiui of adenoid Iw'ue
crowdcil with leucocytes ; tbiM nuuu, which fiillow* the form of the ehnmber,
is wbotly like, in fact re|ieatit altuiut exactly the structure of the raaas of
kdmoid tiwiio of a solitary fullick of tlic iiitMline; it ia vpoken of as the
fittictil'ir or ifiamlular tuliMiinre, or moTV bricflly the/otfiW^, of the alve<>Iu».
Tbia follicle is sepftrated on all sides from tbe cni>sule and trabecule which
form tbe walls of the alveolus (or from the tnioecuUe atone, where, oa in
some cases, tbo alveolns is a small one lying between the larger AuperAoiitl
alre<ilj and the true medulla) by a space which is occupied aa a rule not by
true adenoid tissue, but by a coarser, more open reticular tissue, the meahcs
of which are larjfer and less regular, and the bant of which are mori^ mem-
bmnouH. liaving more the characters of being branches of nudeateil hraucki-d-
cclls than, as we have seen, ia the case with true adenoid tissue. Th<- mi^h<«
of thb reticulum, HIcc those of adenoid iissue, are oocui>ied by Ifucm-ytrs :
bill iImsc arc not .10 numeroua. and moreover more rciulily escape from this
nitustion than from the fullicleii. ao that when a section of a fn^sh gland is
bruHbtfd with a cnmclVhair pencil or nhakeii up in normal Hidini- i°olutioa.
the spacn» of which we are speaking arc to a large extent cleared of the
loueocytes previounly present, while the follictilar siibalancr .'<till remains
crowded with ibcm. After treatment with tWvcT nitrate it ia aoeii that tite
eur&ce of the tral>cculie (and eitpiiile) liorilering this spooa in ouch alveolus
b lined with sinuous epithelioid pliitcs, and a coating of similar plates may
sometimes be mode out on the surface of the follicular substance. In other
word^ this space between tho tmbeculai and the follicular substance ie a
lyiaph-spaoe corresponding to the lymph-sinus of tbe solitary follicle of lite
IntcMioe, and, indee<l. is spoken of as the li/mjiA-tiHut or lytjiA eAannrl ; iIh
lymph'sinus of an alveolus of a lymj>batic gland diHbrs from the lymjtli-
liniti of a solitary follicle of the intestine in its space being much hrokcu up
%y reticular lusue.
TIte imuulsr passages of ibe medulla are similarly occupied by a centtml
niaiu of follicular subatonoe eurroundeil by a lymph-siuus ; but, wherena, in
TBK LACTRAL8 AMU TUE LVMPllATIC SVSTKM.
407
thr nh-nili the miOH* of follicular e»lMUui«r Mke nn ibc torm of mor* or
\nm |>vmniiiliil l>li)clc», iu ibti iitiiliilLt tliu tiilliciiliir sul)*Mnoo it srranmd in
ibr fonu of brakiH-liiiij; und nii)Li|ii&ioziiig i-iir<l>, l/i-- nuvltiiftiri) tortii, tar-
roundvd br « lubulsr lintii«liiii^ and ■nnxtumo^iiiu jiu-krt or Ij'niiili noas.
At Um luactioB of iba c»rii'x ami mixlulla the folliclea nt' ihc alvtwli of the
fcroMT DrmDvh off Into un<l Iimi>iiiv ilia niMlullarv corIs vf the Utter, and tbe
Ijniiph-iintw of th* former mv Biniilflrl}* conlinuoug irilh the Ulitrrinlh of
lyawi-riniBW of the Uuier.
Tm gUnd, in fiict. may b« coDiidfircd as copsiMing uf three i^mrls— lh«
•fcalitlDn lupplicd hv tli« («pHil« and tmbecuia; and divi'linc tlio interior of
Um (claod into the r^ulur alveoli of the cy>rtcx and the Ttibyrinth of lh«
OMdnlla; the follicular eulutance occupj'ing the centre both of 0.w alveoU
•mI "t ihi! lubj'rinili and cootinuou? througboul both, as if it had oriK'inally
dlbd u]f the wliuk of the spaces of the skeletoo and lind BubsetjueDlly
■hnink awmy oa all sides: ana. lastly, the lympb cliaiinel uccupjin^' all ihie
s|MMa left Mt«eeu the fullicular aub^iance and tlie skeleton, ouu ibus form-
iiqc « labvriutb uf '\\s own ihniugbout ih« gland. Obvimtsl/ a lympbatio
cIumI is a contulidaied and dilTereiit idled collection of lyniiitiatic follicles or
Pij«r*s patr.b. In u I'ercr's uatch each follicle i* distinct aod iudcpendeut;
in a tymitbaiii.- tfland llie fi>[)icl« arc fuaed together, [wrtinlly so in ihe
oorttx, but (viniplHt'lvKi in th<- rawhilla.
Tbe attV-mnt lym|iliatic vi-mmU, nbich are mdihII- or niediuin-«iz«<l veiHcls
with thv utructiiri' dnuTilfd in ^ 'J^7, nflfr forming a plexim between the
two Ittrwi of lb« cu[i»nlc, ojirn out into lh<- lyinph-iiuiiMv <if the alvti)!!
btOMln tb« corttui : tluw lyniph-oinuiiea ar<^ pracliiMilIy tyniph-cnpiUario
inia wbicb Ilia regular aflrrcnt lynijihiuir vnneU brcjik up. The eflercnt
Itmpliatie vMsak are ■imiUrty connected with the lympli-rinustx of the
InlUnt the hiliM^; here \i\v lyinphoipillariM of the miidullu opitn into
form the re|>ular lyiuplialic ve£«ela uliicb inue from the gland at this
I piMnt. In the airerenl veaseb the lymph is flowing;, u* we altall see, at a
jcrftain rote and under a certain preamire: it cotitiuuctf to dow througb the
pjabyrinih of the lyinnb-Mnttwaof llie gland, bnthing as it (lows the fullicular
LaobMaiic«, its couiae being retarded by the reticulum of the lymph-siuusea ;
jii flnalty iMiia* by the dfcredit veMeln.
Tlie aniall arteHo* entering th« glaiid at the hiliu run along the skeleton
of traheculw, dividing im ihey go \ ut intervalu they iieod oH* small bnuichea
wbieb, leaving the trabecular hi pnort, traverae the lyini>h-:iiniu. and plun^ng
tnAo tb« Ibllicular subaiaiice brcjiic up into the cnpillanis. By fur tli« greator
part of ibe hliHxl *ciit to tbu gland thua nins iii I'upillary nclworku in (he
follicular MibstJUice of the alvi^di and medulla. From ihi^- capillaries the
,Uood finds its way back by veins tlirough the lyinph-«inus to the trabcculK.
kmnd sn baues from the gUixl at the hilu».
I (SM. Obviously U-re. ■« in the lymphatic follicle of the intcsiiue, the
^•deikoid iMnia, nr follicular salistaiiee, is Ihe seat of nn intcriK'tioii between
Uw btood and the lymph : here the I>!o<nI gives Kin>olliing to and takes same-
iMag tttm the lymph, or at lenel i* in some way chungetl ; here the lymph
■ (akA frnm and gives up to tlie blood. We may be oonlident that itieae
^changes tak« place, ihou^h our knowledge as lo the exact nature of l^KBt
iMOffniial present very limited.
Om event Uking place in the glaitd wems tolerably certain. The leuoo-
■ cyXn which occupy the meslics uf tlie l^^llieular suMtonce. and the vbar-
acten of which are similar to tliose i>f the leuoocyies of a follicle of (he
tnleiline, multiply in the follicular substance. Cell division appears lo be
particularly active in, but not excluMvely confined lo, ceitaia areas iu tlw
follicks vpttken of as lymfJi-ttioi*. In nuclear-stained seetMiis, that is in
408 TUK TISSUES AND MKCtIANI»U8 OP DI0B9T1ON.
PlWDuntiDiie so treaUHl thai while the nuclei are stained deeply Um oetl
Domes ftre very lightly ataiiied or not at all, there may bu frevjueatlv •eeu iu
■ follicle nil area (or more than one area) cuiiaialiiig of a very liKtit oentn
surrouuded by a doepiv stained ring. In ihe Hjthl ceutre the cell bodiea of
the leucocytes are. relatively lo the nuclei, larjjcer lliau the turruuiiding
eone; aud since the cell bodies are not stauied the cvotml tiortion u|i]>cars
lighter. It ia in the clearer ceiitnil area that uuclui uudergoin^ iiiitnwix, and
indicatiug cell divison, are efijieoiall;^ ahuudanL The surplus odl iMpula-
tion thus arising appears to poM, chiefly at all erenU, into tlic lympfa-sinus,
nai to leave tlie gtaiid by the effereut lymphatic rcasda ; on vxaniinnlion it
Ir fouud that IyiDi>li whicli has paiacd ihruiigh a iiuuiber of eliuxU » riclier
iu IvRiph corjiuMrlea than the lymph which in coinias to tbo g1and«^
Many lymphatic glanda contain a (luantily of ulack pi^ent whidi li
chiefly iIuiKwiti^l in Ihe hraiiclK^d celi» of tlie rccliculum of the lympb-
vinusiM. Tiiii" is iinihably, in many ca«> at all evunte. pign)L>nt bronght
to tliu ^lanit in ui« lymph vciW^ls, and arrolcd in ito cotirK^ tbrnu^'h [he
IjDipli-nniis; and in the bronchial lymphatic glands the pigment dimply
ooti»isl* of minute particle* of enrbon introcluc«<l into the brODcbtal pas-
sage by the inspired air, an^l carried from the bronchial paasaMS to the
?;lande. In some cnstv, however, pigment i>> also found in the bo£ee of the
oncocytes of the follicitlur substance, and ihis pigment has probably a dif-
ferent origin ; iU history and purpose are not. however, as yet known.
Tbk Naturk avd Moyeugmv of Lyupb (IscLUDUca CaYLR).
$205. From what has been said in the preceding section we are led lo
regard the mnltitudinouH itpacee, boili »mall and great, of connective tuuiic
all over ihi; buily, including among theiw thti "■unmii cavitiM," »* forming
tlic hegiiining or iiiuta of the lymplialic system. Into tfacH Bpacm cenail
partfi of the plnixna of the bloni transude and mi lieconic Ivmph (whetbtr
the cpith<dioi<t lining of the Inrgt! aemus cavities pbirs anv distinct part in
regulating the traiwidation of serous fluid, i.e., of lympfi into those cavi-
ties uc do not know); from these spaces the Ivniph is continually flowing
through the lymph caMllariea into the lymphatic vessels, am) so by the
thoracic duct nnd right lymphatic trunk hnck into the blood system.
Tlie amount of lymph occupying the lymph -spacce, tympb-capillariee,
and minute lymphatic vessels of niiy region varies from time to time accord-
ing lo circumstances. A hand for instance which has been kept hanging
down for some time becomes swollen and the skin t«nse: if it be raised the
swelling leesens and the skin becomes loose; and a similar temporary awell-
ing of the skin of the limbs, and of the skin generally, ia fre<|ueiitly tbe
reMill of active exercise. Suob a swelling is partly due to the bloodvuteli
being diluted, or ti> the return flow along the veins being retarded so that
the blood ca|iillari«» become distended Hith blood, but is much more laKcly
owing to tbe lymph-^pacea and lymplialic vessels of tlie akin ai»l under*
Iviiig stnicturea buing unusually Ulled with lymph. On the other hand the
SKin may buoomc shrivelled aud dry fr<im a defideuey of lymph in the
tymph-spanv and vceseU. Under even normal circunutanees tbe quantity
of lymph in tlie tissun may vary considerably, atiil under abnormal eir-
cumslnnces a very Urge amount of lymph may greatly distend the tpacca
of tbo connective tinue of the skin and otluT atrnctnree. giving rue tO
(Kiema or drojK^y. Obviously there are ageiicira at work in the bodv by
which the aiijienranco of lymph in tlic spaccf or ile removal thence along
tlH' lymph -cnauncls, or buUi, may be cither increased or diminished.
i 286. Ji£ it alowly llows from ite ongJD in th« tlaauee tu the rnoiiUi nf
the tlioracio duct (ire idb^ for Bimplioitj's sake omit the right lyiiiphalic
truuk) the lyoipli bmbjecud to the influenoe of the lirni[>)mtk- gUixU. and
IB iMasiblT aflbot«d hf th« wallit of the lyinph-vcaoels. Moreover the lymph
oonlng mta one tiaue difl^ts mora or lew id certain ohitmoten fr^^in the
Ivnifih atiiing in another tJMiKt, ju»'l un ilie v«nou* hloo<l of one organ diflera
fmiii the venoiia blood of amithor nr^nii ; and theie dtflttrcneca tnav be ex-
iiiigi-rac^ij by the aciivilv nf tfati i>iic or othor tinue. Of thwo diAerenoea
by fnr the niOMt atrikinff id that lM*twM'n the Ijrmph comine from the ulimen-
tary cmnal during activv digRMion and known n* fftifl^, ana tlie lymph n>m-
inii from other pnrt« of the bixly. When digtiation i» not going on, mid
wlivii o(>n«i»|iienlly no ounxidenible absorption of niiilerial fnxn ihci iilinicn-
tiir^' canal into the lacteuU u taking ]>lacu, tlie Duid Howing along thr Itu.'-
t«n1s M lymph, not differing from the lymph of other regions to any marktMl
degree.
The fluid aeoordlnglv which flows along the thoracic duct in an animal
which has not been fed for some considemblo time may be taken m itlui-
ttaciug the g«neral charactera of Irmph, The couteule of the thoracic <luct
mar be obtained by laying bare the junction of the ituhclaviun and jugular
(in the do); the iunclian oi the axillary and jtigulnr) veins, and inlrodiicing
a eanula into the duct as it entera into the venous system at thai point.
The operation is not unatlended with dithcutties.
Lymph, so obtained, is a clear tmiisparent or slightly opalescent fluid,
which led to itself soon elols. The dulling is not so pronounced as that of
blood, hut clotting a caused as in bluud by the apjiearance of fibrin. The
libriu which in formed though scaiitv, O-S |>er cent., is identical apiutrently
witli that of blood. an<l an likr u^ we know, all that ban been sud previously,
^$ 14-JH. eouoerning the natuns of clotting blood ap)>lie8 equally well to
lymph.
Examined with tlte miero«ci>pc lymph contains a number of corpiisolea,
lymph-oorpuHcle*, whiL-h in all their cliaracien' tu far as m at )>nweni known
are identicid u-ilh nhite hlood-curpuiicinf : lh<:y vary in nixe from 0^ to 15 f,
and the umaller eorniifclcs are much more abundant in lymph than in
hlixxl. LJkc the whilv blood -oorjiuacles of blocul they cxhibil amci'boid
ninvontcnts. Their number varies in different animaU, nod. apparently, in
tita same animal, according to cirnimstnnces; on the wbok- {icrhaptt it may
bft said that lymph -rorpuacl«fl ar« iibniit ai> nuniarous in lymph an whiih o>r-
ihiscIm in blood. Even when enrv care ie taken (o avoid odmixuire wiih
blood, lymph, and cspeciallv ehylc, ii»t unfrwiiiontly contains a citrtain
number of red blood-corpuwtee : eomvtimes tboee are stifflcicnt to give thv
lymph (or chylej a reddish tinge. Th«y have be«D observed within the
hTiDg lymphatic vessels, even within email onvs. and have prohnblv in
aom« manner or other tnftde their way from the btocl into the lymph elian-
nels.
j}2d7. The cheuiotl coni|iodtioD of lymph, even when taken in encfa cose
from (he thoracic dttet. varies a good deal, llie total aotids are much less
than in bkwd. amounting in general to not more than 5 orUjwr cent. Hence
the venous blood of a vascular area contains rather more solids than the
arterial blood of the uime an*, unc« tlie blood in giving rise (o tJie lymph
during iu paasage thn>ugh tbi; ua|iiltarin from the arteriat to the veins has
pan4,>J with relatively more waiter than solid matter.
Tlie ])r(itei<U amount on the average to about 3 or 4 jwr oeni., tltat is to
Miy, to about half ait much ai> in blood, tlw jMiniQular {iroteids pieaent being
TRK TISaUSfl Axn XKCRAKISMS OF niOKSTtON.
tlie Mue as in Mood, Tts., albumin. iiArsf;l<i))u]in am) anle>cedeiibi of fibrin.
In IvBipb. aa dietingub)ie<) Crom clirle, the iiuantity of fat U small, uni]
coDeiflla of tbe uhuhI neutral lata anil the M>a|« of their fatty acMla,
toseUier witli lecithin; cbulmtcrin may alaa lie pteaent. A osrlaiii aimniut
MSujiar (ilexiniM-) apjimra tu be aJvraya pnaeiit, am) wevftral ulKwrvi'-i*
hav« fouutl an a|){ii«ciHl>U (jiiaiitity of unm. The asli of lynipb like ihut
of bJoDtl aeruRi mtitaiiw ■ (v>ii«ii)i'nil>l« i]UaiilitY of sodium cliluridc, wlitla
idtoapliaieit and putttxh htw Kcaiity : it alit>> i:un[«iim iruo, aiiimiTntly in too
onat a i|uanlily li> l>c niviniiiltvl fi»r br ihc fuw n-<) corpuwlnt which may
oe pnwnt. From lymph a oertiiii) nm»iiMt of gu ciui be t-xtractvd, con>
aiating chiefly or slmaM vxcluaivnly of c»rl><>iiii: uciil, with a tmall nunntity
of niirogcn. ih« amount of oxygirn prwi-ni Ix-ing i-xiiTilingty •mail. The
iiniH'tiniR-o of this we nhall new when wc oome to rtiiily rt^ipi ration.
Broadly speaking no mar hit thai all thv sulwtaiiirc* jircwot in blood-
plafnia ara pnwnl at»o in lymph, but are uc«om{>niiied by a largor (|UaDtity
of water.
£ 398. Lymph may also be obtained from wpaTuto regions of the body,
ax from the tower or upper limlw, for instance, by introducing n fine canula
into n lymphatic v^iwe!. In iu ;^ueral leattirci the lymph bo obtained re-
semblm ihal taken from the thoracic duct. AiialyMV of the lymph distend-
ing the suhculaueous ctmoeciive tisane in cams of dropsy iliow that this
contains much }«m KtHd matter ihaii uuromi lymph taken from Die thoracic
duct or lar^r lymphatic voaela. From this it has been inferred that the
lymph normally existing in the lymph-spacea, lymph-capillariea atn) minute
veaseU contain* an exnia* of water; and, mdeed, it haa been a»ert<->l ihut
the percaitu)^ of tolids inoreaaea in paaain); from tbe itnialler to iho litr{^'<ir
reaseU; but this cannot lie reganlml aa diHtinctly proved. The numliei' nf
«orpu»cle«, however, tu wc have iiln:iiidy wii), appears to Im^ inenMuuHl in
[laning through the lymphatic glundv It haa alao been i>tjite<I tliui tlie
ympli in the 6ner ly'mph-vvevcli' ctnt^ even leas lirmly than that in tlw
ihtiracic duel. From this we may infer that wmie of the leacocytnt in the
adeuon) tia!>ue of tbe follicles of n lymphatic glanil find tbnr way in^> the
Ivoiph-sinus. and so into the efferuit lympliatiot, awl ibai •ome of the fibrin
ractors are added to the lymph, or at Icowt that aome changes farorablo to
clotting are brought about.
it 89fi. We showed in ^ -90 that the large serous cavities of the perito-
Deutu, pericardium, ete., were to be oonadercd as parts of the lymphalio
aysteni, and that the " serous ttuid " in these cavities was continually joining
tlie lymph stream ; indeed, pericardial or oilter serous Huid has all tlie gen-
ernl characters of lymph. We have aJieady laid, $ -'0, that these fluids,
nbeii taken fiwh from tbe body, dot (tbia is, at leaat, the oase in uoit
animab); the clot, when examined micmsoopically, is found to consist ol
oolorlc*! curpu)>cl«9 tike; thoae of lymph or of blood onlaiiuleil In the mealies
of librin. Both in tlutir proteid and otiier ohemicul cmuiiiut-nts thme serous
tUiid.i nwtiiilik- lymph. Anajysee of the aceitinulalions of lliiid iiecasionally
ucrurring in thtw i^nvitin Hhow that they contaiji aometimet Iuhm and ttom»'
tim'« mure r>ulid matlitr than ordinary lymph. The aiiiieoux humor of tJie
eye <'»ni:iina very little solid matter ; nn<l the oeTvliriiKpiiinl lluid ii' »o pecQ-
liar that il bud l>ettcr bo conxidered by itM-lf in coiioirclion with the nervous
syxtem.
|i 300. €tii/t«. In fitsliog nuimaljt tho duid flowing along the lai;teals, as
mny Im; M<cn by in«|>cclion of the mcvenUiry, is clear and transjinrent : it is
Ivmph, diflcrin};, ni* uc have tnttt, in no Mssntial r4>j)cet« from the lympb
ttoning along otlur lym|iliattc vvasols. Shortly aflsr a meal ouniaining fat
(and every menl docs contain some liu), the lympb becoints white and
THK KATURK AND MOVBURKTS OT l.YMI'll.
411
5r
Kke milk, Ibe morv w) tlie rklier the nieni U hi ftit ; it lit iImo culled
Owjnv lu tlK> relmivpl)- largo ciunnuty »f thiH itiilkv tliiiil which f'>r
'tins iilWr w nmil cu<itiiiit(» t'> Ix^ |ir>uriHl inr^ lh<< ifiumcir duct. th«
WmcnH of thu duct kIh Iwcunu! milkj:, uti<l nn alw) cittird dijlc In the
'^mdcdnct Vat chjrkuf ihdACUaU u more or Jen oilxoil with Irtiiph rroin
^^Htr IrnpbsltG v«web. but tb* former I* <k> propondanititif; thnt the I'nii-
'^ittM ibt dud BMy be tnkon iu> illiioimtiii^ th<? natiirp of chyle.
1 CKtU di&n from lymph In one in)i»irtBni rc»iicct, nnd one only ; whi-TM*
*ynpn utdinnrilr containR a ■mnll qiiginlity only "f Int. ehyli! coiiliiiii« u vi^^
*^m unount. The tidual Hinount of ftil preiwnl in ihu ctiyle ol' the Ihorncic
*4tj(t rxries. tic may l>c ex{)ected, very coiinderiibly. Rccordinif ti) the nittur«
^f ibv iiieal, the stap? of dif^tion. nnd vnri»iia circ'umBlnDo». Fiv<- por
Cw>t. b ■ very common amount : in the doi> it haa been found to vary from
2 lo 15 per cent. The iiiereaae in fat is chiefir if nol exclusively due to no
iacrnue in th« neutral iiua: ihoujfh nbetber tbo small •luanlUy of soape ami
of tecithin preaeni is greater tbau in lymph lias not been distinctly ascer-
tained. ChoiMterin ia probablr present in creat«r amount than iu lymph,
■oce it probably comes fVom the bile tMured into tbe iniettine duriof; diitee-
tiiN) ; but tbi* u not ccnaiu. How far llie nature of the fnt, that ia, tbe
|M«fNNtii'n of the veriouK kiiidii nf fut, of jttcuriii, etc, variw witJi the fltis
pnNBt in tbt' nrntl hnn mit lieon ilclinitcly aacerlnincd.
Tbe cunditiim i>f thu- fnt in clirle ia peeuliar. rinmv of it vsiiiDi, like tlie
fikt in milk, in ilic form of liit gtabulea of various sinv. but all i>nmll. A
xrty riin«idi-nibl>- ((UBniity, how^rcr. i» present in the form of exoocdiagly
licr tlmn any glob
Biilk ; ilu-tp exhibit active " ftrowninn movement*." The fat prcucnt in thin
niiituti' ■(ilirriilm or grnnuUv. far «niiiUcr tlmn any elobiil<« lo be Mon In
li>rm it Bpokei) ot at the " molecular basis" of chyle, nnd i)> very dislinclivc
of chyl«. In the emulsified cnnlents of the intestine, ollcn eitlled chyle, the
ftt b finely ilivtiled, ami lo a larxe extent into ^rnall kI"Ii>iI<^^ but there is
anthiug oorreAiHiudiux to thin molecular basis; tbo fut du^ not assume ihia
oonHitinn until it bus pUMed out of the inlaitiu« into tJie Incleula. Lymph
•saminni with the microacope shows bcBliK^ tbe white corpuscles only very
few oit-^lubtik«, and notlung of tbis molecular basts. Just as in IHct lynitiii
Is. bnMdIy T<pcnkinjt, blood niiaiu its ml corpusdea, SO obyle is lymph plu*
a very larp^ qunnlity of miniitely divined i>fulral Alt.
The liHal amount of lymph or of chyle which enter* tlie bli)o<l system
ihtouefa the thiirucic <luct, tlwugb it probably varies oonsidembly, in prob-
■Uy alp> alway> vcrv lar^ It Ma bwn culciilntcd that in n wcllicil uiimnl
M quaniitv djual nl loa«l to that of the nholc blood may pRM ihrwugb tbe
tltoracic <luct in Iwcnly-four hours, nnd of thi> it is supposod that about lislf
eoBSM ihroagh tbe hu-t^ls from tbo alimcuinr^' cuoal, nnd therdbr* to ft
larjl* cxunl num (bod, mid tbe remainder Invin ibc body at lar^ ThiM
caleulatjons are based on un4rertain dnia. and cannot, therefore, be taken as
uf exact value, but we may uw- them for the sake of nn iiluslmlion. Thus,
in a RUU) of avera^ weight, that is, nboul iri4 kitn«„ the •|uautiiy of blood
(t S>* I beinR -^ of tlie body weit;bt is about I'i kilod. Tbe <]uantity of lymph
or chvle. therribre, disdiaraed into tbe blood in an hour would lie according
to ihn calculatiun half a kilo., or somethinjr ietu than hall'a litre; and since
the duw nuat vary eooaiderably iu the twetily-four hours would be soowtimM
l««t. and therefore aonetimea even nwre, than this.
7%e MottmenU of Lymph.
i SOI .MakiDi* every allowance for the uncertainty of the calculation d«.
lailtil in tltc prcceiling paragraph, it is ohvioii* that the lymph nivn flow
rnx TISSL'Bfl AJttt MKOHAMISKS Of UIORftTIOK.
will) a utA incotiaiilerable rnpidity (if we Uke abaul half the above eaiiiuato.
the rate will Ik about •'i c.c. per miouCe) throuKli tlie iboracic duct, and
iberefAre muHt also be cootiuually MreamiiiK into that duct alont; the rarioas
lymphatic channolH fruai the inaiiifoM IrniuhspaoK of lix body. This
onward proKreitii o( the lyii)|>h i» deu^rniiiied l>]r a variety of circumilaDces.
Id llie finl |»Iho«, ibe remarkably witle-«pre«d preMoce of valves i$ 267) in
the lyra|>haiiu veiscU uaiiM^ every prewiire exerted uu the tiasuee in which
they lie bi its*i»t in the prapuLnon forwHrd uf the lymph. Itence all iuub-
nihir aiovemviiU increuo the Itow. If a caitula ba Inserted in une of tJie
Ull^r lyinpbalic Iruiikfl of tbl^ litnb of a d<%, the di>tcharg« of lympli from
tbo cnntila will be mon; dUliiictly iiicrMMed by inovemenls, even pasaive
mrivcmmt*, of tlic limb than by nnylhinit <'l»v. When we oome lo speak of
tlw entrance of chyle iiil» the liKrti'iil railiclcn of the villi, we sh«1l tee that,
Rt alt Rvonu according to out vivw. lh<- imiiu-ulnr fibred of the villiia act aa
■ kind of muscular ptimp, driving lh<' diyli: p^mi the valvnd «nd of the
Ucl«al nulicif into the lymphntic nnnl* boluw. [n niKlitian to tin- preMnoo
of vaJva along the course of tht- vtsmU, ihooponing of the thoracic duct
into the vonoui Byetem is guarded by n valve, so that everv i«i.-un<> of lymph
or chyle from the duct into the reins becomw itself a help to the flow. In
the Mcoiid place, we have already seen that the blood-prcmiire in the oanil*
Inries and minute veeseU ia considerably greater than that in the largo veint,
such as the Jugular; in fact, this difference of prtesure is ihe caus<^ of the
flow of blood froni the capillaries to the heart. Now the lymph in tbo
lymphatic spaces outside liie capillaries and minute vefseU undotibtedly
Btanda al a lower pressure than the blood inside the capillarias ; otherwise
the tranaudalioa from the blood into the lisstros would be checked ; but the
diflhrienee is probably much h»~* than the dilfereuee between the pressure io
the caplllarisi and that in the liirgn venoux trunks. So that the lymph in
t}i<! lyntjib-Hpucea of the tiMiniM may be c^onsidered as itandinjt at a hi{;ber
pruraiiri- than the blood in the vemiuH trunks, for instaoee iu the jugular
vein. That ii to nay, the lymphiuic reaaels asm whole form a system of
channeln leading from n region of higher preaure, vix., the lymph-s|>aoes of
the tiiwne*. Co a rcKiiHi of louvr pnMure, vU., the interior of the jugular
nnd subclavian vcnis. Thi# dil!erencc of pre«4ure wiU, a* in the case uf
the bloiKlvcufH^ls. aiitse the lymph to tl<>w onwanl in n continuou* stream.
Further, this How, caiucd by the Io¥mcM> of the ntciin venous presaiire at
the subclavian vein, will be iMiited at every iwninilory movoniml, since at
every iiis]>inition Ihe prmsure in the venous trunks becom«», as «* >luUl aee
in dealing with respiration, uenttve, and thns lymph will be sacked in from
ihe thoracic duct, while the increase of pressure in the enaat vein« during
cxiiiratioo is warded off from the duct bv the valve at m opening. In the
third |>lace, the How may be incrmsed by rhythmical eontrnctions of llie
walls of the lymphatics ihemseivcs, which, as we have seen, are remarkably
muscular; and the peculiar interlacing of the muscular fibres above cmcI)
valve suggests that the walls here act nl^er the fashion of a tiny heart and
by a rhythmical systole drive on the Huid, which by the action of the valve
below collects at the spot. We have, however, no experimental proof of
Uiii ; for, thoaj(b rhythmic variation* have been observed iu the Ucteals ol
the mesentery, it is maintained thai theae are umply passive, i. e., caused hj
the rhythmic perii^tallic action of the inleatine, each oontractjon of the in-
tetitine fllling the Ivrnph-chnnnels moK fully, and are not due to contraclioiu
of tlte walls of the lacteal vtascU th'mx'lvcs. In tume of the lower animals,
for tnstanee in the frog, llie rauseulur walls of the vcmcIs are develoj>ed at
placM into diaiinctly cunLructtlu propulsive organ:!. njiokeD of as lymph*
naarts. of whieli we shall have suinethiag to aay pnwenily. Lastly, it id at
TDK VATCRS A^'P UOVKUKNTS OP I.YMPH. 413
tariopn fiiT D«. oil the ilrcnutli of the snnlnny tliat nemosis may kito rise
ttlDcnuKi preasure un one tide of « <lifl\iiuon aeptum, to flU[t|M«e thol tlte
ntjfnttma wbidi give rise to the appearance of lymph in the lyinnU-
ifMaof the luiu«», tend theiowlTefl to prnmoU? l)ie tl<iw of lymph. We
hnu lm«, und«r »ll circ-iimsianoea, one or other of thene cauaw at work,
»rwtiUu|{ n ci>iiiiiiuul thm from the lympkalte nxit* to the great vein*.
ntyuv t<vi-{h<-r iiillii-ictii to drive, io mtui, llie lymph friim ilie lower
fabtml trunk, ngaiii.-t the eAevts of firavity, iiiln thv vi>lm of thu neck.
(a lb iipjirr lim)i iho iril1iK'n<t« of gravity, owing to the- vaHt'd muVL-inent*
■'iW limb, an' ai> iH\ta favorable to. «i oppuced to, the natiirul lldw of tlw
lynpfc; hut, M m have ulruiily Mitd, a loDc-cuiitinui-il mifuvonililv ai'lion
tfpavhy, Mpwtnlly in tl>e abneDce of tbe nul of niDVcmrntn in lli«r skoli-tal
aiveln. a» who« the arm bimfw dnnn ml>ti<>nll^M fur aunie liiiit'. leiiclo to ac-
'vmalatii'D of lymph at ilf oriKin in tin- lymph itjwce*. The ntivtigth of the
(tmn eombinin^ l'> drive on the lymph h Hlrikingly itbnwn in nnimnlii when
tlw UMiracic duct u li|CKturvd : in xtirh cawv n very grrat clulciwion of the
ly»iJ|>attc vcimU below the ligatiirr i:* nlitcrx'cd.
f 80S. Allbough ibe pheooDiena of divpiiM and, perhnpv, geDcral cnnridera-
nader it pruhiiblc that the nervous syglfni govenw in H>me wny the
ttnun of lymph, n-j^iiUtint;. it may be, not only the flow along thi; dctinite
Irnph-ranab, but ttlmt (ho ininsit n{ plasma into the lymph »pnct^ and the
%fm of lymph ihmce inl'> the definite camd». our knowledge- on thoae
(H^ti it very imperfci:!. We have no proof that the muiciilnr fibres in tJi«
««Ue of the lyuphntic ve«»).>is arc governed by nerves, or thai the lymph-
•pM*« are influeitecKl directly by nervous action ; and muet of the atteiDpts
to detDonsirate iiny direct action of the nervous syttem on the lympliMMB
Imtb hilJKftii failra.
It it very difficult to diasodate any lueh direct action fW>m an io<Iirect
infliKnee through vnaomotor ebunKM: for the mnditiou of the viueular
I •TMrm largely affects the forniatioii. and henc<- the flow of Ivmnb. Tbuti if,
I (n a dng. canuin having been plaee'l iu th<^ lymphntie triinitii leading from
•cb of the hind feet, thit »ciaijc nerve on one aide u divided, the How of
fruiD the f<iot DO thai oidc \» i:n«ter than on the intai't *idc, but i|
sisbod on Klimulation of the peripheral end of the nerve, the iliininii*
bottis followed by a iinliM.-i)ueiit increaMi. The Mrction r>f the nerve,
(iMimeer, Ifads lo arterial dilmion, the stimulation of the nerve to arterial
ooMlriction : ami until other reasons be xhoitn, we may attribute tko in*
onased or diminulMil flow of lyrapb to an incmtnol or diminUted Iransuda-
IMi from the fuller or cninucr bloodvasels. And thi« interprerlnliou is
sMfportid by the fact that urn'o stimalntiou of the nerve i* ki con<]ijclc^l us
Iio md to arterial dilation (^ KiS). the result is not n diminifibeil hut ao
liiiiiwiiiil flow of lympb. Acain, if the eorvieul sympaihclic in a rabbit be
dWldcd on ooe siule and a Hijution of the blue piigment »ulphindigotal« of
nd*. be Injected into the venous system, tbo ear on thai lidc betomea blue
tbe other, because the pigment pii>t>es more rapidly from the blood-
I into tbo Irmph-Spaces of the connective tissue, and the bluenece also
away MOlMir because it is foooer washed nwny by the subeeijueDt
oneoWed Ivmpb. But here, too, the iticmsed irstisudation uiay be regarded
Mtamply ifie result of the greater fulness of the bluodvcesels.
i 303. The pMtage of nuilerial. namely, of water ouotaiuine certain tnb-
•tanoes in solution, from the interior of tbe bloodveasd where tliey form part
• of tbe plaama into the lymph-capillary where tboy are allied lymph fonaists
»f twDSt«|is: tbe nnieafie from tbe bloodvewel into the lymph-Bpac«, am) tbe
pwvige from the lympli space into the lymph ranillary ; fur, as we liave sten,
ii u ■■oly in paritcubr plac(>4 thai tbe lymph-c«|>iniiry immediately surrounds
414 THE TISSL'KS A K [I MECIIAXISMS Of OIOESTIOX.
till- MoimIvi-.'wI. Once iimv(!i) in lh« Iyiii|>>i dipillnrr tbe lym^iti lincln ah
opon |iHili ■liiti^ llic n-fll of llie ]yin)ihatic iiy>teni. litil tab ooniKL-tioti betwwa
the lvni|ih')ipiice niid ilie U'lupli'diiiillury la. Ha we Uiive Mfti). |>ecu)itir aivl »l
lensl nol n tW-c niiil ii|)pn urn-,
Tho pHMHgc "I" nintcriiil fn)m the SIikhIvcswI into the lyinpli-tpaee we
sprnk o1"n» Irniwudnlinn. Whal i-nii w« «iy O" I" fhcimlumoniiis procCM?
Tlierp arc two kiiinvii {iliTviciil pniciwifii with which wc miiy comiMn it:
lUOueion ihroiigh n nicmDninoiis uv other porous iMTtiuun, ntiH iJllrnliua
llii'dUKh H siiniliir pHrtiliuii. Ilifl'iipion. though iiitliiciii.-c(l by fluid prawure,
IB not the iJiret'l result dI' Hiiii) prer^iin^ hut tnay, on the eimtniry. be tliv
cuuHe or diflerenem of preesuro on the two sMk of the parlilton, «»il may
ururk againBl Oiiii) preoeure. When n flrong solulion and a venk dilution
iif salt are bepnrntcd by a dilTiiMon septum, dittuaion lakes plaee nhelhrr iho
fDUimna of fluid be at ihe same level on tbe two eidea of ihe e«p<iim or at
ilitt'trc-ul levels; and if the columns bo at the sani« Iev«l to start Kith, that
of thL' stniuger aolutioD soon conies to exceed llw otlier in beixht, od account
«r ihti osninlic Aov. of witter t>om the weaker into tbe stronger anlution.
Filtraliou, on tbe other hand, is the direct result of prea>ur« ; nitbout dif-
f<>rcneo 'if preasure Kllmiioa does not take place; and, the filter reuainiug
of iho name ualure and in tbe same cotidlliou, tbe amount of filtrate i*
ilcjH'ndfDt III) tbe ainuunl of pressure. May we apeak «f the procav of
treu*iidatii>ii uh a nimple pnicess of diftuHion or a siniple proocat of filtnttun,
tlial i# to my, cuii all the pheuornena of traiiaudalioii he explained a* nrnpir
the results of one or other of ihete phytical pn>oc»8e«? DiSuKion by itveCf
will not account for the reaiilU; for Iho pniteids nf the hloiNl-plwinm arc
iodiflusihlo or very iieitrly no, and yet the lymph coiiutinii a oonifidenible
(juantitr of thwo priitiridi). We have no 8alislui-l<>ry kRowledgc of the cxa«t
c'impoeition of lymph m it exist* in the lyiupli -spaces. In the lymph of tbe
larger lymph-trunks the dilTusible saliDe Eunstances are present in about tbe
tame pruiKirtion, and Ihe iDdiffunble proteids to about or loss than half as
much an in bloud-seruoi ; and we may perhaps atsume that tbe lymph in the
lymph'ipaces contnins relativelr len proteids hut has olberwiae tlie same
ompimitiou ub bloud-plaania. Mere dilfueion would not fci''^ rise to n fluid
of Mich a nntim-. <'an we apeak of IniDsiidalioa tlien as a filtration? TIm
blond is un<louhtcilly (lowing tbrnugh tlie capillanea and other amall rexaeU
unricr a certain pn^eiire; ue have seen (^ llti) that the preMurc 1* rouelily
epcaking al>ant 30 mia. Ilg. ; and it would be ponible to aelectaucb a filtor
or piirou« partition a* would at about this praHure permit the pawngc of a
certain qiinotily of ihi- inorganic and oryatnlline cimittitiienls of hlouii-plmuna
to paw through in company with a relatively nmnllrr fpnintity of the prot«ds
and a large quantity of the watrr, the red and white ourpuiclcw being ex-
cluded. Such a Hltrnlc would Iw morR or let* of thr- nalnrc of lymph ; and
80 far wc might he justilicd in ip<-nking of the Irunmidntion of lympb aa a
prooeeaof KltratJoD. But ibo trani'it through tbe living wall of the blood-
venel ia aflboted by circumstaocrs in n manner w> dJHcrcnt from the manner
in which the sane circumstances nflijcl the transit through aii onlinary lifeloK
filter, that we gain but little, and may be led into error by sfteaking of (he
grocer as a HltralJon. Hulntanees in solution or otherwise, paas through a
Iter nlien the pmeure is sufficient to drive them througn the pa»ia>(e>
funiidhed by tbe nilersticeH existing in the substance of the filler. In tbe
cn»e (if an ordinary filter the substance of the tllter is within liniiu ii&rma-
tieat, and Uie ixiMages correspondingly ooDSlant. The livio;{ walln of a
Mpillary, however, in aot a cmatant unchanging thing. The e]>ilbelioiil
plates mid other elcncnta whieb conatitule it are alive, and being alive are
L<oHliniudly uudtrgoing change and are especially subject to change ; mora-
^^^ 'TtlK NATUBS AMI UOVEUKNTS lU' I.YIiiril. 415
^Mi^l «v httxt H«n (^ 22, 2S>, ibe vascular walls atipeiir lo Ik- raiiiiriuslly
sdioa u|iun nod Iwiufir ac-ieil upon by tbe blouil. Heiift' a elmiigi> in ih'o
blnod t«ndii lo nujc rhatigM Ju l)i«ai ; am) thwt- cliimu;A* may mnii'riiilly
■Act tn oi>e (lireetiiiii or au<>tl»i-r iWir uc-tion lu 61t4-n>. In an unlinnry lilii>r
bu'lBiae of pmnir« n«ccaMrilv I'lilnila incn^-e of lillralion : in it living
fiter it inajr or mar ma, awl inc umc iiicnuLw of [imuiirc may ucfMnliiijt to
cutnRwUaoea product very diflerent rcMilt-i lu rrgardt the traiuudatioii of
Tbu it ttetm rcMonablc !■> sappoM a* we have mggoetod (S 227) thai,
' ihiam being iW Mi»«, an ii>ercnm of blood-preMure 'hould iK-«nariljr
I tlie trsiMuiliiiioi) of lymph. Hvnrc when a tmnll iirlcn- ililalca,
•JBM Uw praaurv in ihii >till emalk-r branclHw nml capillnrica of tliut nrtcry
ia. ■• «v bave toon- tlum onco |>otnlod out. incmuoiJ, inor« lymph appean in
tbe lirnpb-^«4v ; indrod il is oda of the main ptirpoecs of ibe widMiing of
tmrnli artvnw to eupply tbe elements of the tiwne with more lymph, lliai U,
wiUi more food, llut it does doI Ihorefore folluw that under all circum-
Mabm widmtng of the artery should increase th« paaeage of lympb : Mme-
tl^Bg mar oceur to countersct tbe natural eR^ot of the inoreamd prawure ia
Ibe lMoodT«eMlB. An instance of this seems to be ait'urded bv tlie case of
tb* •nbmaxillary eland, witeo tbe chorda nerre is stimulated while tbe gland
b Qftder tbe influeiii.-e of atropine. Aa we have aeen, tbouk[h the arteries
dilate, DO fecivtioti ukis |ilac« : and ne cannut explain ihi.- aWnoe of a flow
bla the alveoli hy HU|i|Hntn)j: tliat the extra arnixmt of lymph which would
ia nonaal drcunulancu fiirru part of the secretion, and in tbe case of a fairly
eopfaaa aecrativo would Ix; cotwiderahlo, now panea away by the lymphatic*
wiUtout reaobioK Ibe cella of tbe nlvei)1i, fur in micIi eiMin* no oxtra flokv in
the lymphmlka leading from tbe gland lia« beat olurrvLiI, and ihrn- ii^ no
accnmulatioti of Iymj4i in the connei-iivc tiMUe of the gland. Apparently,
fur Mime n-nwn or other, in spilo of the increased prca»ir« in the hlood<re»-
H^BOn lympb than umibI Aik-s nul pase into the lymph-spaecs.
tImo again, as we ahall presently have occa^on to point out, an increase
nTprnmiv ia the blooilveneU produced by c^tructioo to the venous ontllnw
ie onicll atore efficient in promolingan increase of IransiidalioD, al all events
an alnormal increase, than is an increase of arterial pre^ure ; and the dif-
fiircs«e between the two vases ap[)e«rs to be too great to be accounted for on
iIm firouud that an obatruction to tlie reaoua outflow raises the preasure
within tbe capillarivtt and small veeseU more readily and to a higher degree
than diw9 tbe widening of the arleriea. itloivorer that olMtrui-tion to venous
oMAow does not produce il« eflects In the way of transudali'in niniply tad
BMnljr by raising the capillary pnMiire in shown by thv fact ihjit the same
•notfnt of ob«trncti<m may nr may not give riM> to exceasive traiuildatioa
■ecording lo ttie ctmdiiion of the blood or other drcumstanoM. For instanee,
tbongh Uia ofaatnioliun prodaocd by ligaturing a vein frequently eausasvx-
caoiir* tntMudaiion, il dooa not always cause it, aixl the femoral vein of a
doc may be licaturvil without anv cx<r<V!<ivo iraneii'lalion taking place; vet
if, after the Tignturc. crrtnin cluuiges be inductil in lh» hl(wd exceeuro
tnuHddalMii occurs in tbe log, the Tain of which hia been ligatured but not
■laiiwlurri I'tHUting toward tbe mme conclitnon ia the fact that exceauve
tmodatton njnrn readily occurs when a roiu is plugged by a thrombus
arising from abnormal conditioos of the vascular system than when a vein is
Mmply ligatured. And in genera) we may my, and this is a point to which
we tball mum, that two things ehieflr determine the amount of transuda-
tion: the preasure of the blood in the bloodvessels, and tiK- oondition of the
vascular walls in relatiofi to Ibe blood, the latter being at least lu important
as the former.
41ll TKK TISSUKS A N' 1> MKCHANISMS or DIOKSTION.
Atiutber a«pect of rW matier morenTer deserves attention. In tfltralioo
the iiioveiueDt take^ pIuL-e lhrt>u|;[h the lilter in uiie diremtion unlv. irbureiu iu
the liviu(; body, the na«sa^ of mnlerinl UimuKh the uapillary wall mkM
]>!nce ill tun uppoHile directions. In ull the tiwue^. [htiti);)i taum iifrhup* in
cerlaiu tissues lliaii iu ciUiera, th« passage IVoai the hloiHlvv9M.4 into lb<?
)yni|ih-spnce h aiXKiuipanied by a (Mwage ttrvm the lymph-Kpuf-e into itir
liliiiid ; while food for the tiiaue pones in one directiim, u*a»Ut pni'lucla p*i«i
in the other. In n secretiug tcl^nd the (greater piirl of thv lymph euming
from the blood vexM'U, the water aud other mutter*, ptuv away into the In men
of the atreoluK iil'ler uiiderj^iiig chanfcc* in the rell ; but eren in nuch a caw:
there in »ouie return from the cella iiil» tJie liloodvrMoIif, nrl>onic a<:(d for
instance if nothing eW in given up by the cellH to the blood : mid in auch
orgiuis ns a mnsck- nr the liver, ibt; backunrd Klreum of mnleriul from ibi-
tJMite ti> tlict hl'jod is cxtriixive and inijioTtiint. Moreover this bookward
strain) workH agnin^t pn^Miirc; iiidetMl. m niny be «cen in a uuscle, it u
when the blumJve^>sel.'> iirc dilat)<d and the prvnnru in the eapillarim and
itiiall vresel* high<*t, u« during and aflcr the cuntrnction of tht> muscle, that
the pBa88g« from the liMue into the blond in most ciier)>etic. Many of the
waele products of tho titane are it la true diffusible, and we mi|^ht be tempted
to say that while the lymph which feeds tbe tissue traverses the vaHcular
wall by liltralion iu tbe direction of prewure, the wnste products return to
tbe blood Hgaiust pressure by diffusion ; but siieh a vieff cftnnot al preoent
be regarded as proved; and if it be true as is mainluned by eome, that
lyiupli, including the proteids, may at times be reabsorbed from tJie tJsaue
into tbe bloodvessels, it is distinctly contradicted. We shall have to return
to tliis QLiestioii when we coiue to deal with tbe secretion of urine; but
meanwhile we may adopt the cuDclusion, which is especially supported by
the phenomena of disease, that while dillbsion and filiraljoo plav their n.-
sfWelivii parts, flitfusibliT iiubntaiiw" pawing in and out of the blood cnoro
readily tnan indid'iiMhle Mibilancm and an incrcaw of prtwurc tendtng to
to pn>motr trunnudntion. the condition of the vaKular wall so profoundly
iitflucnct« tbe tranxit of material h« to render the proccM vcrx* complex.
We may probnblv regard it ns Urn complex In be compiircd even with liura-
lioii through a filter capable of widely clianging in texture from lime to
time, and ns niore nearly rrscmbling the proocn of >^ccretion.
C«uceniing (he passage of the Ivinph from tho eonbnci lyrapb-siMioes into
the open gangrcnys of the lyniph-capillarics wc know very little. If, as
some think, tbe c«vity of tho lymph-canillnry is "hut otT on all sides and
oompletely by a continuous lining of sniiioua epithelioid plates. tbeD the
passage from the lymph-space into it must be rc^rded as a sort of repetition
of the passage from the blood-capillary into the lymph-snioe as a second
transiidulion. liut if, as others think, and as on tbe Wnole seeoiB mote
probable, tbe lymph-«pac«8 open at places directly into the lymph-capillaries,
tlie passage is a simply mechanical atfair determined by the freedoot of these
o|)enioga.
In either esse the flow from tbe lyuipb-spaces will ba ftcilitated l>y all
erenta which promote, and checked by those which hinder the How of lymph
along the lympb-capillurie^ and tbe other lymphatic channels.
^r^ ouiy here remark a* iulluencinjc the ({uantilv of lymph in the lymph-
spaces aiMJ veiwels that tho ijuuiiity of lymph laVeu up from tbe lymph*
qMe€s by the actual elements of me liwie mav vary considerably! We
rentarkeil in g 30 on the puculiar relations of living tissue to water, and
tht^n.- are reasons for thinking that the very substance of a cell or a fibre
(a muscular fibn-, for inxlann.-) may hold in ilKelf n larger nuantily of water
at one lime than at anollxr. Tlie water thus taken up or given out. nod tbe
MOVEUE.VTS OF I.YMPII.
417
»
■otatutn* which may tw rarri«) in ^ituticm hy that water, ooroe fnini and jto
U> thv lymph. The cotiditiou uf tliv tiioue ilHurminM by iUelf ihe unxmiit
of Irnipli IN iho lytti|'h tpaoc*.
I Mh. I'ntlvr ihv intliieitM of all thmb hktbI iK-(ionii tlio lymph in ths
TmnaoM lynph-mm* ul' the t>oi)y varicH in Buioiint fnim lime bi time, but
noder ounpil nrcuinxiancM never cxttwd* certain limit*. Unilur juithu*
kvi^ ooodilioot lbnM> limit* may be vxck-CiIoiI. aiiri thu retult in kouvo a*
■■■inMOr drafwy. ^inilar vxcci»iv<' acoiimtilntioiis of lymph may occur nut
in Uw unlinary l^pb^paca*. btit in tiitvr largiT lympli-i|iHcai. thi; wiroiu
aarilica, any iarK« •xom of fluid in iha pvritoQnd cavity being kuown M
to
Tht paaiblo nuta of oidvma arc. nn lh« one band, an obstnintion to th«
of lymph I'rom the lymph-spnrw, nnd on tbf othrr hand an irxoxwivo
tniMDduioii, thr lympb enthtiriiig ill (lie lymph-spacM fiuU-r than it can bo
(mrrifd away by a mirmal t!ow ; ivilh t\w former tlio lymphatic tyiiUsm itacif,
vith ihr latter chi«(lr the vaiMriilnr vyMcin is «)DC0niei1. A« a luattor of
bet, huwcvcr, imicma is almost iilwnys, if not always, due to abnormal oondl*
tioaa of tbo vavcular srslein, and ie the result not of hindered outflow, but
•f MXcnaive liansudalton.
Owing li ihv niintenius anastomOMs of the lymph-vrw«U nnd the conso>
uaetit HUblishntrni of oollateral Ureanis, obstruction iti tin- lymph-pMNi^^
ihiiiiwlini rmrely if vm give* riae to isdema : and it mav be here rviiinrlceil
that inring to the sane me oollateral communication iM^twccn ibi' lymph-
Ttaela tbe labyrinthine piuwages of the lymphntic-gtnnds do not oiler (he
a«ciuo» obatacio to the onward flow of the general lymph-strenm, aa might at
ifM ai^t bewppoaed. Nor have we at present any knonled)^ which would
lad oa to mppoae that any phy»iolo^teal changes in the walls of tbe lymph-
•lic^TtascIs or of the lymph-capillanea, or in the lymph spaces, by giving
rtee in aomo way to ub»tnch* li> the flow of lymph, ever lead to an accumu-
latioii nf lympb in the latter.
Otw kind uf u-ilema wa haro already loufhvd ujmiu in 6i>eaking of tbe
cspUlarj cireulatiuu ($ 183), viz., the "inllammatorr" a-deiua. In llua
kiod of ndetDB, owing to cluwgoa in the vascnlur null*, a lur^L-r amount of
tTBtnudation puaea ium thu lymph -spacea. and that traniiudBtion Ih richiu^ !n
protcid matter*, and cunlaiiin a lar^r nm'>unl nf the fibrin fnct'ini, or at all
•VWtt* i» much more distiiietir ciinguliihle than onliunry lymph, n* well nt
crawdeil with migrating oofpiiaok*. Atlictl Ui this itifla minatory mlema i*
tbt iocteaae of lymph, abn appurenily changed sontewhat in charnrtrr,
which appeu* ■• "eH'uaion " in the serous cavities when theae are inl1ainc<l,
M in pleuriiy and peritonitis.
One of the moM common forms of redenin is nn itdcnin ■>f ]>riinarily.
ibough not wholly, iDcchaaiml origin— wdenta arising from obelniction to
Uw v«ioui flow : undiT tbtee circuiiMtancM more lymph passe* into the
lympb^paioca than the lyiDph-reasela are able to carry away. If tbe femoral
vasn bo tied tbe 1^ may become oedematoua, and, as we have snid, (edema is
a commca rtwilt of the piuggini* or obeimction of reins ihrongh disease;
tbe ondema which w so comnton an anwro pan intent of heart disease, involv-
Utcobat ruction to the return of vciuhib bluod to the right side of the heart,
au Ibe aacitea which folluus upon hiudmnce to tbe porlal flow, are instanns
of ttdema of this kind. We have already remarked on the relation of
traosodaljan lo btoixI-preBMiio. and in vi-iuhiv ultsi ruction the ri&e of prcMure
witbia the amall lilmxlvessels i» dwintinisht-d from that due to arterial dila-
tion by being accum|ianied with a want of ade<|Uaie renewal of the blood ;
this probably afiects the epiihelinid lining of the bloodveieels in such a way
aa to increase the trantudaiivn. And, tniWvd, as it aetn la caae of In-urt dia-
■It
TBB TlSSUSa AND HBCffAKISMB OP DTOBSTIO?!.
our uiih )ir(i1oDgcd or re|>eKted venom ob^lruclion, tlic uvlema, a* liin« i
on >ti<l ilu' liiwiie* b«ciime impaired, lit mure eiisily i-xdted nml vritli ^
Oifliciilty n^iDi^v).''!, lliuiigh the nctiiitl iiiiiKtiiit nf ulntrii elicit). ili« actiiK)
incrpni''?' of pmioun; in tlin .imiiU vuiucU, ruiuaiiiii the Mnii;, or nt Ituul i» uut
pmpiirtionatrly iiiirrcii!«?'l.
Slill nntitluT kiiiil of (nleuin is oni^ Htic to chaugn t«)ciii)r placn in the
blood, quite apart from variaLioDs of Mood-jirmure. Tliin kind of •nlema
ia »een in eutiie diwMiws of the kidiirr, in *' Bright's iltfcuc" fur iiMiano*.
In Auch cases lliu blood ocinLnin« \v» jirotcid*, and indeed 1cm solida. i« more
watery and of lotrcr specilic cnivity than i« normal. But Uto tcdcnui is not
iu lh«4e ea8e« lo bo cxnliiinciroii tlit? viciv thnt the more wat«ry blood ponn
more readily thruni;h llio capilUry walls, for ll mnv Ih.- shown ozperimvnliUly
lliat tli« mere thinning of the blood, as by the injection of noniinl ealiiw
•olutton into llie blooil vessels, will not at once lead to dedeaw, at least in the
limbs and trunk, and it is these which iu Urifilit's disease especially become
<BdciD&toue. Iu all prohabililr the oedema of Bri;Fhi'8 disease, if it be really
due to the abnorniat character of the bhmd. ih produced by tlie shnorrnat
bluod 10 sotiug on the bloodveseels that these allow a transudation i-reBter
tlion the normal.
But ihow are patholofrical qnestions into <nhich we must not enter here.
We have toudiea u[iou them beoau»e ibey illustrate the important procisoe*
taking place in the lyinph-apocea, and, as we have more than uncv inuvtied.
the lymph in the lyupli-BpaL'ee is the midiUninan of all the tiwuM, ami b4>noe
fad* ilUiitiniting the laws which govern the flow of lymph intn and out nf
the lyniphKpacM are of flindamental phyAiulogical iuifHirtnnce.
S SOS. Lijmjih-hrtiTl*. \n the IVog and nlhcr amptiibiii and in reptiles iIm-
flow of lymph into the venouH iiy»teni \» awiMcd hr rhythmically pulsating
muscular lyiniih-hi.'artK. which ]>r(«i^nC ninny ciinuus UDalugies witli the
htoud-hearl. The frog poasMsa ti>ur lymph -hvarls. Of these two. belnugini;
tu the hind limbs, arc placed one- on each side of the coccyx near iu end,
and, being covered only hy apiint'uriMis and the skin, may without diaseccioo
be seen heating. Two anterior ones arc placed on the transverse puimwn
of the third vertebra, and are covered from view by the shoulder ifirdle.
Kach lyitiph'heart is a more or less oval sac lying in one of those lymph naot
or cavilieii lined with sinuous epithelioid plates, which, as we have taid, are
preeeut iu the frog. It is c<inlinueil at one end, by an oritioe guardbl with
valvea, into a small vein which opens, in the case of the posterior hedit, into
ft crural vein, and iu the case of the anterior hcarta into a jugular vein.
The undl vuuoiats of muscular tibres arTauh-ed in a plexiforni nuiuucr and
snpportcil hy a considerable aniunnt of cunoeolive tissue. Thuae iit>r«« an
Btnated and hrancheii and are intermediate in character between cArdisc
and skeletal muscular librea. Nerve fibre* terminate in tho>e muscular
fibres, anil the muscular wall, unlike that of the blootj-heart, :» viipplied
with Lttpillnry bloodveiwela. The interior U WotA with epithelioid plates of
MKUous outline, and tlib lymphatic lining io continued along a number of
openings or porcH. by which the cavity of the hcan opens into the surrounding
)ymph-«p«ce. Whej) the XtviWl cotitnuits the content* are ilriven into tho
vein, tlie lyniplintic pore* being closed by tli*' np]>ttiximntioti of lh« cod-
tracting muwular fibm ; when the heart dihites the fiuid in the rein b
prwenteil from returning by the valve* at its mouth, while the lymph enters
readily from the iium)un<ling space through the now open porea. In the
flrojlg racular IvmphntSc vcsmh are scanty; Ticncc these lymph-heaitB beeome
of ooostderahlc imporlnnce in promoting the How of lymph. The Irinpb-
hearts of reptilia arc i>iniil>r in siniaure and function. In the fn>g, in
which they nave been ehicHy Mudied, tbo action of the lympb-bearta is iu a
iXTARV CANAL.
419
I
tBMMr* dvptixlMii Ml tbi> »piiiii) r<>rH. T}i« potieriur Ivrnph-hcnrU beUing-
iac b) U>* bind limb* »rv •.'miitMlcd by nieitas of tbe (Iclimic leoib pair of
»fml twnr«i> wtih n R%Hon ofihccartloppcMtc ihf nixili »rwvfiiih vtrrtcbrit
ftflMH^Rwayibnt M-cii»ii atth* oerrc oratHniction of ihr jMiriiditar rvgion
^if llicotnluiiiiMniiliiordiidmvi iWinidivity. Thi? aiilori'ir [Kiirnn- »imihirlT
«nODfcte)l wttb a region <■!' iIm' iipinal fun) ii|>]K»itL' tbc thinl vcriobra. Eacli
yir llwnifew awCTW bi tiBvt>«"ci.*ntTv"inibi' An>iiial cord; but it in probable,
ViMMigb obMrren are niA whittly ngrcil. tlinl thv bi'srti>, nftcr dratriiclian of
^bair spinal cvntre, ultimately rfsame ihrir rliythmic hfMf, «o tlinl tbe
^ItppDilmce of their iictivitr on tltr 8)>in]il cnitra is Dot aii iibmlute one.
Ijkr ibe beart of t)i« blootf-sycteni. tlw lympb-hcnrts may be inhibil«t, and
that ill a Ktlex matiiwr, the iiihihitioD ocntrtr being morvover in the medulla
«l>'i>i)e>l^ If a rro|> be nrcl'iilly ohfervod. the activity of the lymph-heurta
«111 be linind lo vary larj^ely. and thcw vnrinlions a]i{i«ir to be iu purl due
la aerxmm indo«ne«s. w that iu tliia n-tiy tbc iiiovcmeDt of lymph, and hence
iW ptocMw uf abaoq>tH)n, are in this aniuuil directly (le|wii<Ii-ul on tbe
MTTMn syriem.
Absori^iok pbom the AuMEicTARy Canai.
S 306. We may now rttura lo conrider tbe abaoqiliou of th« products of
dtewtioa, tliitt ia lo My, tbe paiwmie of ihaw bodies from tbc interior of the
abmrntary caual. where tbeV are really uutaide tbe iKxty prupfr. into the
body itwlf For siniplicity n itake we ena^ conal<ler difn-stioti in a browl
way aa tJie eouvuntofi of pmcli>'ally non-ditnuihlu pruteids and elttrch into
BOfv dtffiiuble pv^loni^ and highly dlfTuiiblc ^ngar, and ua the eHiuUifyiug,
or dlvirioD into minut« iMrticKv of fats. We have reiuon to believr that
nmt of the sugar may be ebaogml into lactic acid ur even inln butyric or
otbar scidt, that some of tbe protctdB ar« carried beyond tbe pvjitone D>odi-
6ao (oto lendn and other bodies, and timt some of the fat may be HH|>onilied ;
bimI it may be that some of ibc jiroteid material of Ibe food piuucv into the
body mn albunii»e or even an i>anij)efitone. or in some other little changvd
roMition. But we may probably mith safelv. fur jir^wnt puriiimii^ iiwumu
tbat tbe ctvalcr part of the proietd u l%\)*^^T\iv^ as peptone, thut carbohydrates
ar« iiiainly nb*>irlied as sugar, and that the >[ivater Diirt of the fat puaaes
tfltu tbe body u emulsified but otbi-rwise uui'ltanjfcu uentnil fat; and we
nuy iwigkct tbe other conditions i*f drgtsted food as iub«idiary. and as far
•a alaurplloa ia concerned, unimpurtunl.
W'r hare seen that twi> {>a(bi« iifL- iigmn for thne products of digestion, one
by the caiiillaries of llut portal system, the other by the hicteals. It eannui
br a matter of indtArmce which oounw ij< taken. For if the products paw
bv tbe ladenU they Ihl! into the general htiiod -current aAer having under-
gone only siieh ehnngra ns ihey niay experience in the lymphatic ayiteoa ;
while if they piu« int<i th<- [xirt»l vein (hev are fubjcete<l to eerUiin mwerfol
inlluenci-s of the liver (which we tilinll 'tudy in a future chapter:' iR-tuir ihry
find their way to the right fii'le of the licart. We may. tlierefoiv, euiuider
ftM which iif the tno paths is. as a matter of tact, taken by tbe m-veral
products, and suliaetjaeDtly study tbe nbecliaiiism of absorption in tbe two
Tlu Omrte latrn by ihe Sexvral I'rodudt of Digttlion.
I S0T. Prom wliat hii* nlrendy beea said we have l>ceu led to regunl thu
viUi aa tbt moat active nrganr of abaorpltun. and Iho ■tructaiv of a villus
oa fltrther to conclude that the iliffiuible [H'titoiMi and augar laus,
lerwlth lbs water in which tlwy an- (lissolved. into the superfidalty
420
THK TISSi
placed capillary network of ilie viLlna and «i into the porta) anttm, irbik
the merely entulsiti^d tat, unikble lu travi^rae tbe trail of ibe i-npillarir, puKS
on lu the deeD-eeuted liioteal radi(;le, aiiil so finda iu wny iiilo liitr lympbatie
ajMeni. Anu tbe ix-aultK of obBervHiiuii and cxjMtriineni, an far iu> tliey ^,
tupptiri lliia vien.
Fat*. AIYer a lueol containing fiit th« lyniph of ibe laeleftU oantaina fat,
atxl IK niiiv (udled cbyla ; and lh« ricber titi; niuil in fill lli« ntrire oo«*picw>iu
M tlif fuC in llie lymph-vewelx. W« [-aiirml. liowL-vcr, [imvu that all the fat
of a Picuil ubwjrlictt rniin lb« allnieutiiry in pi)uri.i] by tbu ihorudc duct into
lh« vcutuii!* »viit4'ni. If ft iiiutl nmttiiiiui^' h known (jiiunlity vf fnl lio given
to ft d»^ and ihc »niutl <|iiuntity nt' fal [irtwiit in the fecn correepondiiig to
tbo nifaJ be MulUriiL'tcd fnini thnt nnimiiit, wp can dcti^rmino the anxiiint of
flltabaorlicd, for we haw tio ci'idcnccl wliiiU-vcr tbiil any upprtciablc nm-xial
of [at undergocv a ilcstrm-tiv)- it<»ri>mpoftilion in tbo nlimcnlnry i-iinnl. Col-
lecting by meaiit of a rjumla iiiMTli-d into tha thoracic iluct the whole of
the chyle during and nfti:T the meal *u lung ne it rcniainB milky, ebowing
that lilt i« lieiiig nbmrlK-d, we can tircertain the ijuanlilv of aheorbed ftl,
nbich would, bill for the ojicralioii. have pnwed into t(ie Tenoiie ayatom.
When this hiM been done, a very rcmnrkablc dcticil, aniutiiiiinj* it may he to
40 or &0 per cent, hue been obeorveil ; that is lo eav, of every I'lU |>artB of
fat which disappear from tbe aliaieutary canal only about tiO parta Rud ibeir
wav through tue thoracic duct into tbe venous system.
Are we then lo conclude that the mining quantity finds iia way into tbe
portal eystem ? Now the portal blood does, during digcetjon, oontain u cwr-
tnin quantity of fat; indeed, the seruiu ia itaid at tiiuea to appear milky from
the presence of fat. But the whole circulating bhiod durin)c tbe digntion
of a fatty meal coDtaina. for a while, the fat poured into it 1>y iliv thoracic
duct ; and it ban t>een a.icertaiiied in the dog that the blond oi' ibc portal
Vtin during digistion coutatnii not mure but Icea fat ibno the hlood of tbe
carotid nrlery, m> ihnt tlit? fut nhiub nppeum in the portal blood during
digestion if, (or ibc UKWt part at Uywl. not fnt ahM>rl)ed by tbu capillaries <?
tbe aliiuenltiry cniial hiil fiit ahsorbi-il by the WtenK Moreover, when the
chyle of the ihoracic duct i* diverlvd tlirough a canula, and not allowed to
flow into ibe bloifd. the <|iiantily of fat in the portid blood as in the blood at
large is very small iiideeil. Lastly, when a villus of an intcMJae in full
digeition of fat is tn-ateii with uiitiic acid, fat cannot bo reeogDiud by the
nucroMOpe within the capillaries or other bloudveuel*. thimgh it abounds
outside tliem in the substance of the villus niid in the lacteal radicle.
We may probably, therefore, infer with safety ihol all or at lea.-<l vi-rv
nearly all tbe fat absorbed from the intestine takes the path of the lactealiL
Aa to the deficit mentioned above, thai is as yet vilhout eiplnnation. It
may be that in some way, on its course, in tho lymphatic glands, for instance,
tbe fat is taken away from the chyle, bidden so to speak somewhere aw-ny
from both chyle and blood ; but on ibis point we have no exact iofonuation.
% 308. }VaUr mid nJU. If. iu an animal, the rate of flow of lymph or
ebylo through a cnnula place<l in tbe thoracic duct be watched, and water
or, lo avoid the injiiriuUN i-fTect uf simple water on the mucuuit toeinbrnne,
normal (ullne nolulion Ik then injected in not too great iiuantity into tbo
intestine, nt> niarkcil iucrt^aai! iu the fiow of cliyle thrnugo the cunula is
obeervcd. Fnmi thin nu nmy infer that the water of tlie inlcutinal conl«nla
is absorbed nm into ih<- la<-teat» but into tbe |>i)rtat •ymeni. If, however, a
very large quntuliv t>l' l!i<; nunual Mtlitie *i)liitiiin he injectcil >n ai' to distend
the iBttstine, iIkii tiiv tlow of chyle i? increiwd Iu wmie extent. It would
appear, thcn-forc, that while under normal conditiooN the WHtcr jawMS from
AUSORPTtON KKOM TIIK Al.IMKSTAKY CANAt.,
tiiM ntuinlr int<i the purtnl blood, eome of it Difty under circutn-
_ Hi int'> tlii> lacteals.
With fvpird to lh<> coureo laktrn hv ordinary aline mattera we pnaacao no
]|isl inlortnatioo. When special s>lt# eucK as potswium iodide ind
, CBsilr ncogDlxed by appropriate tml«, are iiilrotluoed into the intes-
. (key mir be apcedilv delecleil both in the blood and iu the contents of
' tho thoracic duct : but n liciher. in »iich oaset, these sails find their war into
' tb* ihontcjc duct by tbe lacteal radicle of the villi, or pass into the lyinpb
[aifWim at tonw later pajl of iu counte. ue do not know. Nor can no with
nprd to Mich a salt as aodiuin chloride, state absolutely that it pedaca mainly
with tlie water into tbe ]>Drtal blood, tlioiiKh we may fairly suppose this to
bt the case.
$ 306. Suynr. Both hl<MMl and chyle contain, nonnally, a certain uuall
tmotiat of auj^r: and careful iot^uiries «h(>w that tlie percentage of sutfar
b chyle and tn jj^eneral blond is fairly constant, neither Iwin;; to any marked
tilent increswed by wta amylaceous wcalx: on ihe mhi-r hand, a meal ood*
Itiniiig suftar or starvh does teaipurnrily incrMiw ihi- <|iiiinttty of ougar En
tbe portal blood. From this w« nnir ind^r that nurh porlion^ of thti wugar
_; ff Ine tnttvlinal conteiibn as arc abmrlH'il on ^lugar jituu rxchiKirrly by the
Cal rein. We may, howi-Vrr. hen- call attcndnn to thi? ililliculiitsi attcnd-
an nrgumenl of tbi* kind. In ihv timt plu<i- the tjuaiilitativc dctrrmi-
of a sfnall amount of^iuftiir in to coniplvx a fluid lui blont) i.-i atlcndcd
jfTMt diffieultics' anil uncertainties. In the Miond phiw a vi-ry largo
quantity of bliHfd t> ol anv one iixinK'nt strcnminjf through the capillaries of
Utt> >JinM-ulary <^nal; and wc may |x'rhn|B speuk of the quantiiy which
Baawa ihroagh them during tbe whole pcriuil of dig«»lion lu In-ing cnormiHis.
, I1«M» though each KM) c.e. in pawing thrnugh the capiUarics might take up
(quantity of sugar »c> uniutl m to fall almost within the limits of errors of
ration. yeC tbe whole tjuantitv absorbed during the houra of digestion
_ _ i be oonaiilerable ; or to put it in another way, an error of observation,
-maTOidable with our present meaiu of analrsio, on a sample of blood taken
frtm the porta] veoaels might lead to a wholly unwarranted eonolusion that
majpu WAS or was not l>einK abs»rbed. Making every allowance, howerer,
Ifrr theM diffii'ultiea, thu incrense of sugar which bus been obsen'ed in (he
pnrtal blood during diction seems too great bi iiermit of any otlier exclu-
sion than that sugar it really abM>rl>ed from the Hiimcntury canal by the
blondTcoaala.
When, howerer, a large ((uaiitity of sugar diMoIved in a large i^tiaiitlty of
water b prewnl in the ini«*liae, the sugar (n the cbyle in naid to be tnorciwcd.
In such a case the eioMs of water, as stated above, |xun=i into tbe lacteal*,
I CIkI in wi doing ap|ipiiri> to carry some <if the >tigar wllh it. '
S 310. Ptntriil; The dilficullics atti^nctiu); the ■■x|H!ri mental determination
of tha path lakrn by protoids are greati'r even than in the ca»c nf lugnr: for
tba aiaot iiuantiiativc ostimation of [wptone in bli>oil (and wc an; asnimiog
that protcida arv mainly nbsorlHtl as peptone) is a laxk of the gmlfst dilii-
evUj. cMM vomiiaml with which that of celimnting sugar appears almost
MBjr. Bearing this in mind we msv <tAte that all ouerverv ara agncd that
I peptMte ia absent from ehrle or at (cast that its nresenee cannot m satis&c-
\ lonly proved. On Ihe otiier hand, while «onie oneervcrs have succeeded in
Bnding Peptone in (he portal blood after food, but not during fasting, many
bare tailed to demonstrate the preMOn of peptone in the blood either of the
portiil vein or of the reesels at targe even after a meal containing large
J QUaDtUiea of protdds. Of coune, us we ai^ued in speaking of the absorp-
l-non of ausar, tbe qnantity of peptone pawing into the portal bloud at any
i.nomeat n^ht be small, and yet a oooaoerahle cjuautily might so piusduriitg
■
tlie biiun of digevlion. We may »iipi>iHe, iDorwver, thut ihnl which
pau in iniRii'iliBtel/ oonvorced. {ws^ibly by some ferment Hciioii, into ofiv or
olher of ihtt imiiinil proteide of the lilooi), or otlienviae diepueed of: and,
iiHleed, i>0.|ili>nc iiijcctcil carefully Had elowly into a voiu disappMls from lb*
blood, thntt^h little or even none pastes out* by the kidney. Aim) I tit' view
tb»t iifpcnup is *o cbaD|;;ed. poKioly in the very act of absorption, ia siijv
porli'd not only by the 8Uteni«Dt Itmt peplono may be found in the practi-
cnlly bloodlras wall, that in. inutouit iiiembnioe. of the iutesline removed
from n dcnd miiiual even wbeu it appeitra tn bu abteiit from the bh>nd, but
idsu 4Uid cepocially br the follonint: obserirutiun. If an nrtilicial circulatioD
of blood be kept up in tbe medenterii- urli>ri<w !<iij>plyi»i.' a loop of tnteatine
lemoved from tbe body, the I<H)p may b<? kt-pt nlivc fur kdiiiq ootwiilerahle
time. DiiriuK lliia Hurvivul a conifiderablo cjuimtiiy of {wpiuni; plawd in the
oavity of tbe loop wilt dituitipnir, i. /-., will be- ub«»rlH-d, but cjinnot bi- rvoiv-
ered truai Ihe bluod wbicli i« bdi))^ used for tbe urtiiicinl t-irinilntioa, and
which e3>ai|H-.-> from tbe veins idler IravvrNng the intcstimtl t:ai>illuin>. The
ditupiieitranee is not due to iwiy iiction of ihe bUxjd itself, for iMptonc intro-
duced into tln! blood K-forc it i« driven through the mwcnicrie nrleriiT* in
Lhe exjieriincnt mav be recovered from ibe blood ns ll csL-npea from tlic
iDOOntvric veins. U would ttxm as if the peptone were ebnn^l before it
actually gel> from the iiilcrior of the intestine into the interior of ihe
cniHl liiries.
Bttt llie nrt;<iDient that the abeeuee of peptone from the blood is nii pntuf
tliat the itcploue i§ not absorbed into the ulood may also be applieil to the
cbyle. and thus leaves us unable to draw a eoncluBioD as to tbe path of ths
protdds. The fallowing indirect proof that peptone does not pa« into the
chyle has been olfere'l, but it too ix 0)>eu to objeetiou. We ttliall see here-
after that the absorption of pmteid tuaterlitl lead* to an iiiereaf« in Ihe
eliniinuiion of urea by the kidneya. So marked ia thin increase, thai uulea
thvrv br- cIcHrly some other c»ii«e» at work leadine to lui increase of urea,
■uvh ti:> lever tor inrtniiw. lui incrwc of uren in uie urine following upon
the admini^trnli'in of proleid food nniy be taken iis a proof that tJie orouid
food baa beeil digested and ali»orbc<l. Now if in a dog tho thoracic uuct be
auDoeflsflilly ligatured so thni the chyle cannot paw as usual into the blood.
•nd the dog be fed on proteid foo<I, as free as possible from fnt, «a as not
miDMesanfy to load tlw obslruoied lacleals. an increase in the uroa of tha
UtilM b observed aa usual. Obviously in such a cow the pmteid food t>
abaorbed, and obviously also does not pass into the blood through the
Ihorado duci (tbe success of the lijjature having been proved by post-
mortem examinations But the evDeriment. ihoniib as far as it goes sup-
porting, <low not Tigi>r*iUHly prove llie view that the proteids are ahrarfaed
Dv the i'«]>illanen itf tbe aliiueutary canal : for tbe thoracic duct and lym-
phatics bi'low tbe Itxalure vntn fouiitl laritety distended, and lyionb and
cfaylo npi>Mir to have eicaped front the veueU; hence it b passable that
•onM! at lca«t of the jiroleidH were sbwirbcd by the lacleaU of ihe intCM-
tine, but finding ihvir usual {wtli blocked loaJe tbdr way int4> the bloo<l
aln»m.
We may, therefore, say that the multa of cxpcriinont while they do not
delinilely prove, give some supix^rt to, and at least do not contra^lict, the
view which we a little white up* put forward as probable, nanely, that of
proteids, ttanffornied into ditfusible pc{>ton<^s, {wss into the bloodvewels aiul
not into the Ucteals.
Hut, if ibis view be provisionnllv accepted, it must he on the underttaixl-
iug lltat it is pmbable only; and il may l>e that protdd* do not take tltc
same |i«ths and are not nbeorbod in the same condition in all animals. TIm
ARflOKrTION KBOM TIIK ALIURNTABV CAKAL. 428
opntiDmbi ju*i rvlnk-d wore twrliirnicd on dogs, thiU is ui «iir on carnivoe-
Mu miinal* iih<)m> 'nnlural) toad vonlaim u c(»ii»ilerat>le i^uaniity of fal,
■oJ «h<*« laclcsls Riigbt, itiercfoiv. he coaaidered afl preoccupied iu ibe
*'wiir]4iiiu rif fal. Tlie fond of berbii'orit un the other baud cniitniu* a
'i''alit«lv ■mull atitoutil of fat : and if in ibcac animnU all the proteids nod
nrbotitdrnln are nWirbed hy the blood re«»elii, then* U companuively Ittlle
''ft fir ill)- In('ieii]» to do. V«i in theee aniniaU the liicieab aod lyrapbatio
kT **" ''''^ *■'**!*"'■ '" 'I"* villws of a herbivoftiUB guitien-iiig or rabbit,
"j^^'ijeli ihe reticular tinue ia vtry icanty a« com|jared niih lUiil prfewtil in
™*J *illuji of a dog, ih« liu-ieA) caaiuber ia relatively to the diaiui-ter »f the
"''Ixa, Dot merely aa large as but much larger tliau in ibe dog. It in dilfi>
^'t Iu iu|>|>ose l^ai this wide chamher is ioteoded solely for the abtorpt>>ii)
T^ the relatively small amfiuoi of fnt j>tL'»cnt in vegetable foiK). The >jik'S-
'*Xt nbich ne are ducuMing i^ cleurly at prescut to ba rtganlL-d us by im>
"•*■•!» acUlwL
Thf MerJtaniim of Ahiorption.
9 311. Thr nbtorplioM oJ/uU, \Ve hare now to conuder the manner in
^Vidi ib«e severnl subaiances pam into cither the lacteal radicle or Ibe
^'^ptllarv bloodvessel. It will be cxmvenieut to begin with the aboorption of
Wt- hate iWCD raaon. S 281, lo think that the fats, remaining chiefly a«j
*a«uiral fats, are eniulsiRed in the intestine, br menus of the )>il« and pMH|
^muie juice, ihe small iiuautity of soaji whicli is formed pnilnhly acrving '
vitvply ih« piirjHve of fii<-ilitatitig the emuUifu-ntiun.
TIi« neutral fnln so vmulnilied pant in ilie finrt instance Into tite budit* at
tbv rulumnur tvlU of thi- villi [Fig. 12-1]. It has, it U true, bc«n main-
^shYfl by tvmo that they piuu briieren the cell* and not inli> them; Ikii the
«vifk|ic» i* distiDdlT ■gaiiii't thU view. Thuc.t'lU ma v again and again be
Mvn rri'Milird nith ml, and iho cosci in which the lat haa Ikcii seen birtw^o
the (i')U and hot in iheni iin- due to ihv extniMon of the fat. durin;; the
•hrinktnt; of the villiix in the cotinw of preparation, from the cell' into
•psctx ■•etwecn lh<^' otlU. In llic fmg, in which there are no villi, and in
«bicb ibo folds of niucr'u* nicTiibrane eervin;,' tho purpurea of villi do n'>t so
mulily tbriuk, Ihe presence of fat globules in the celtH after a fatty meal can
alwaya be easily demonfltraied by osmic acid preparaiinna. Since no auch
ooilrcU'oos of (ui globules arc seen iu the cubical cells of the glands of Lte-
brrkuhn we tufi-r that thnte hiive nothing to do with the alaorption of fat.
How the fat eut«r« into the substance of ihe cell we do not know. We
■nay prt^uiue that the stnalcd border plays some part, but what part we do
not know. Thouffb, as nc have atKU, the rodj making up the border ap(iear
able lo more, to cnangv their form, we have uii evidence ibat the fat ia in-
tnidaced into tlM> cell* by tm-an.t of any movemenlfl of theae rods. We may
iimcine ifaat the globulM piun intu ibc o^ll MilKitnnce by help in some way
of tnne rials, through ammbuid niovemenia comparable with tlie ingottve
ffloveniinit* of the IkhIv of nn amrrha ; but we have no jHuitive uvidencv to
aup|<»rt Ihi* view. Wi- Hiiid 4$24i) tlutt bile proRioles the |>awagc of Alt
ibrKucb membraiif*. jawuibly by in luime way pmniotiug n chnvr coatoct
bctirnn the juirticlt-F of fiit nnd tlie xiilititnncu of the membrane : but evan
if bile has this vlTet-l on the stirfaee of tbi- celln. it« iu.'liou iu this reapoct CW
be stibsidarv oulr-
Wilhia llie columnar cell the fat may be won, both in uemic acid prepa-
ntkfns, and in fresb livini: cells, to l>e ilti>|>(«ed in glnbnloi of various sizes,
•utue Inrge and some souill, each globule placMl in a spaoo of tiie proto-
plasmic cell subatanoe. It doea not fotlow that the fat notually ent««d the
A. tncnui or ms Viuu* or a Ktt Kivr.Kii hohitio r*T-xBW«ra<Mi.
9i, tpllkelliuii : ifr, itrliital border 1 e. Iyin|.li relln : f-. lyniph-cclliln OicopMwtMni ;
'. Mulnl litplral lonulnliigilliliitniimlnii l<tni[>li-«>rpiuolMi
B. Hooont UcxBKU'i or FBO»*i iNmnDti t>tiii[!in rAt-Atmotrnttx.
t/r. tfUMUaai : ifr, «iH>Wd bonUr; i^ lyui|il>-air)iuinli>t ; t, UoWkl.]
From l.li« cohniinar mII the fat [latMS iuto ih« anaoea of tlie rcticular
Umig of Uio villuH. U hiiH, it is true, been mntendeu that it paiuui aioog
tlio subAance uf tliir bunt of the [«ti<^ululu ; but in uarcfiiUy i>r«par«cl oaiaic
acid apecSnicnv of a villus in actire iligevtion uf fatly foud, the fat may be
distincll}' recoraixod m InrKcly lillinif up. !ttill tii the form of globiilca of
varioiiit rnux, the >jiac<w in flip m«hw" of the n'tioiiiiim whidh are not uocu-
pieil by the Iciicocyt« i>r iitliral iiumlcring ccti»- Wc have aeoD (S MI J
that the biKo of the columnnr iiill.i, ihniuKh the gap« in the baMixieoi mctn-
tmuie, directly nbiit upon the liitiyrinth »1 Riinom ; imd the fat onee nnt of
tin base of the cell id free in the xpaeo of the labyrinth. How it iasuee
from tjie cell we do doI esaclly know ; pomibly by n proceaa analogous to
t]i« eser«tton of solid matters by an aintcbii.
Froni llie Inbynnth of cpacca of (he rclieuliim of the rilluE the fat pasMa
into ibe cavity of the lacteal radicle: and il is worthy of note thai io the
IMMBge it undergoee a e)ian);e, In the interior of the inteotine, iu the aub-
■tanee of the oolnmnar cell, and apparently in the labyrinth uf the reticu-
lum it is aiiuplv einuUiflt^d fat consisting of globules small nnil large ; williio
the lacteal radicle it eonHistfi jkartly of the aame ea«ly recognized globules
but part3y of the extremely divided "molecular basts" ($300); it is now
DO lonf^ enmUiSed fat but chyle. How and by what means thw extremely
miiiut« divijioii «f the globular fat intu the " molecular basiit " lake* place
ABSuBJ-TION KKOM TIIK ALIMENTARY CANAL.
426
■ •tdoBM know : nttr do «r« ktiun the «Xficl manner iii wliieb the fat jmumx
■ frw tlw Bpmces of ibe reiiculum into ilie interior of tbo radicle. If th«
V 4m cf niiiiouH epidieliiHi) platM wbieh fi)rnu tbe aole wall of ibo cbamlicr
^L fcdBcooliiiu'Mi*. prMenliciK here aod there ini|B Iwtween tbt- pUten. the pM*
^^b|ppitseu[B 11(1 ilitliruUy in il««lf, but itiiCH mine the iliffii'iiltv why there is
^^Bbgnftt • (lilfi-r«iioe iM-lween the chyle iiiiii<le the chnnibcr hd'I iht^ fat oiit-
W Ak On the other hand, if as ohaerTaliona aeem to nlio^' the linitii; in ques-
■ Ml b actually coiiiinutNia, the fill must pan into the hict«al radiole either
I ^<Ki|>b lb« subHtanoe of tlu- pliiles or through tlx' jnnrtion line* of cement.
I A>ch a ptuaa^ prrvenudiHiotiltiiv: hut u< the mmc titnu we can oouceive
I '^t ill the itrusglca of >uch a [iiuutu^- nnniv of thi- hil might be converted
' i***' the tuoleculnr iMuin.
W'c may Imt^ nrrha|M reninrk that the (»Dt«iit» of thv lacteal radicle
^>*iM IMM eiclu«tvplr of 1st. hut of fat accooiiHinied by thi? pntteid and
■'htti nbauon* which jro to make u[> the chyle. Proteid and olhoir sub-
••■^Jow hiwidM fin an.' also pmciit in tde lymph which owunic* in part the
"^yrintb of lh«- lM«ly of the villus, and arc derived, like ihf lymph el*e-
'f'^ow. fmm the bloo.| of ndjacciit capillaries: at Icswl. they are in part si>
•^]»"iv<d. th-High it may be not wholly, for aa we have jiigl aeen iho Jiaiwage
f*" pmirid niutrrial from tbo interline into (he aubstanoe of the villu* paal
"^^ capillarin though not prove), must still be comidered as ponible.
^« haw Boen (§ 'i6'i) that the spaeea of tbo reticulum of the villus are
^^kr» or lev occupied by wandenng cells of which wo spoke under llui
S^MiM>I lertn of leucocytes. These do not atl present the same appenranon
^*kd MoM probably are not all of the same kind. A number of Ihem
''^y br distmi^uisbed by the fact (hat the cell-body is loaded with discrete
C^^nulo which stain readily and deeply with certain aniline dyeB.and which
**tougli Dot of a fatty nature turn black with oamic acid.
Some of theae leuctioytea waiider nut ouly Ihmugh the labyrinth of the
*^ttrtiluni but pa» into the epithelium between ihe oelhi. and may project
IhroeMMM ini'). or even make their way eventually inin the interior of the
KBIaalloe: or followini^ tlie n-ver»e eounte niny wamler fmm lietween the
^ptlhaBum oelk into the ttody of the villus ; iinme of them, moreover, un-
Qoabtadly eontain fat. Hrnon thi^ vit^w hai> W>n ninected tliat thcae leuco-
oyMa arp im[>ortjiiit agent*, indeed the chief agenia in the abtorplion of fat.
Xi bu Itrvu nuppiiMrd that they, rcii-iving the globuW of fat into tliL'tr cell
■nbataace. in fiict eating thv'fnt exa>'tly nfifr the manner nf iin ninteba,
«itlNir while pnjeoting beiwcen the cnlumnitr n-U*, in « hich caiw they c«rry
tbrir bordfn of fat throiieh l.h'> ppithdiiim into the villii*. or whili- w»ud«r-
■ng io the labyrinth of tat- villui', Ix'-iir it awny boililv into the lymphatic
•jitMli. lint thi number of Imicocytos rvally cunt.niniii}; any nnprrHable
quantdy uf fat is too smalt to account for the nmouiil of (at nhfiorlml ; mdco
MM we ju*t nointed out in a certain kin^l of thrsc cells, and thix kind ii ofli^
vrrr abundant, the granules in the cell subslUDoe which stain with uamic
■nd are not fat. Nor is the abundance of leucocytes in Ihe mucoaa mem-
brane during Ihe period of digeetion a sure proof that they are conecmed
Is abaorptioD, but rather an indication only that active chaDgea of aoine
kitKl arc going on, since after the administration of a saline such as magna-
tiuni sulphate, which producee eHiKts the very reveree of absorption, tnSM
leucocytea are preeent in unusual numbers. Moreover, under some drcum*
fltancea, aa in toe villi of a new-born puppy after a meal of milk, thev are
ahaent even when digestion of fat i< noidly gislag on and the Uuiteals are Ailing
wilb fat. In fact, what we stated aoDTe ooitc«ming the preaenoe of fat in
the bodiea of the coliimuar celle shows that 1euci>cyt4« can have little to do
ill timnaferring fat tnm the interior of the iuteatinu iutu the body of villus;
rilK TISSfE
Rml then! urv iio m]ui[iinle rcjifnii' for iitlriliiiting in lliem imjr mA
ill llu- intadfvii-iKri; <if fiit fri>ni ttio Ixiily <>( live viltii* into ihe 1(
c bum til- r.
^ 312. Tlic Inctciil ctiunitior ojHtnK iil ibi.- base uT the vitlii* inui th« r«lre(l
lj-ni|>hii(io viwjcis lying Iwlow. nnil in iht'w: the lliiw nf Ivinpb .oliyl*l is
In-inj: ]ii-i)mol«l by llic vorioim ciiiiMv ^cintlcMl in S :i01. 'flic prewun;, for
iii»tiinco. I'xerteii liy the niTisliillic contrnctions nf ibo inuisiini.- belpt to
vmpty the lyinplmlic vt^Hvl ioti) which « Inclcnl chamber ojhim imd w pro-
mote tho cmplTing of the IkcIit. In mltlition to this the )>lniii riukuIv
fibrM of the viliiia Biipcily ii ^jK'dnl niuKiilur pump for the «ni|>lyine iidiI
filling of the lacicnl cbiiniber. These fibm atid email l>iincll<» of ubrt*.
though niitning in various directions ( § ^ti^i) uod rarring in nunilwr luld
Hrmngcmenl in diflerODi animals, take OD the whnle a lungitiidimil dinvlioD
pantllcl lo (be loiif; axis of the villus. It has bcea supposed thftt in odd-
tmcliog and shnrleuiug the villus they coinprees the laclcul and thus omptjr
it, and that when they relax and the villii» doaj;atea a^ain, the emptiM
cbfliiiber fills ouce more. But a diRl-reTji interpretation of their action bw
been oltered somewhat u# fullowa. When the muHcular fibres contract tk^H
shorten the villus, in tbii& tKH-'Oiuin),' »hcirter the boily of the villus b«codH
proportionately bromler, since nrobably no uixiai ekauKe of bulk in I ho
reticulum lukes place; in tbia Lruailfiiin^; the part to ^ive iiay vrill be the
lacieiil ehiimlier, mhich thim lieconu-* brouder and tarKcr. When the fni»»-
culitr fibn-tf relax, the n-ticulitni, the )i»r< of w hicb have been put on the
streii'h in ii ]iit<-r»l ilirv«ioii, bv daiilie raiclion briiii^ back ihe villus to ila
fiiniier li-njij^th, and tbe lacteal oluinilier ulongiitcs iiinl niirrows. Un ihb
view the niuvcular contraction ex|iiiiidii anil *<> liDtt, nbile tlie reluxatioQ
twrmu'* and w <:niplie» the luclcjil rhanibcr. Which ever view we adopt, we
may at IviiMt conclude that contracti<>ni> and telaxationit of the musculnr
fibrtn in wime wav or other nllornnlclv lill iiiid eniptv ihi' lacteal chamber,
and ill iill probability, utall cvoul:< during dieeaiion.rliytlmiical cxHitimclJon
III' these tibrce are cimtinually ffowa on. \Vben the villus is sliortened bv
tbe contraction of tbe muscular fibres, the ooluninar cells are oompnmo,
becoming longer and narrower; when the muscular fibm relax anil the
villus elongates, the columnar cells return tit their previous form. Tbt
Bltemaiin^ chnngeit of form to which the columnar celU arc thus enbjei-twl,
and the alteruuttiig elian{(es of pressure takiu^' place io tbe reticulum, may
alsf) serve t» promote tbe passage uf nuitcrial through the one and lliroiigli
tbe I Ft her.
$ 313. The abtorfitiiin <if tli^'anbU mh*l'>urM and of tiviler. On the pro-
Vi8ii>nal aaauiuptiuu which we have made thai tbe prateids ore coiivcrtcal
into peptone, we may cuiiiider, for tin- pn.v4'ni at all events, ]ic|>ti>iie, sii):ar,
tad Miuble salts as together forming a cIiim ■liiilinjiiiiabed fVoiu fal» by tlwir
beiuK dilTuaible, some more .-u) than othcn. Atul we have made the furtber
proviatonid aMumpllon that ihcac pim lutu tbe btoodvc*»cU and not into the
toetealM.
The netnork of oapillarr bloodvcmels is spread, as «c hare wen ($ 263),
immenlbitvly liciicatli the bawmviit membrane, and all the nuilerial which
enter* the lactriil vhnnibcr bu« to run the gauntlet of the luusbes of thi«
network. Ihirin^ iligmion tbe capillaries of the intmiiic are filled and
di«tcndcd, to that at ii time when absori)[ion is taking place these imshea
Ijctwcrn the capillaries ate unusually narrow. From iho interior of these
capillaries, here il* cl»cwhcre, transudation ii taking place ; thctc capillaries
supply tlie lymph irhich helps lo till ui> the labyrinth of the reticiiluiu and
the Incieal chamber. Hut to a mitcli jnvalcr extent than elscnhcrc <cf.
$ ^4>.'t) thi» current of transudation from within the capillary to without i«
ABSORl'TlOX rHOV THB Al.lM£NTARr OAKAt.
427
Nc<r<>iiipiuii«d by b reverse rurreol froio without to wiihiu. Tli« diSVisible
suIietflnoM in <)Uwtion poM from the inlesitne throujth the layer of epiiheliiim
oelb, tlirou;;h ilie aii^nuated re(i<:ular lymjilixiHice betneen the Itasemetit
membrane aD() the i-apillary wall, and tbruugli tlie uiitillarv wall into llie
blood current. Their {mwaije cuiiuittUi of two otjt^; tlial l^muKb the epi-
ibelluni cella (Voni the inleatini' to thi> lymph-sjiaoo, and tJiat tVom the lymiib-
s]Miee into the bloudvcsaels. Thenc two Rtagm may beenpeeled to diliW,
seeiuiK tliat the titracttireit mmciTmitl arv dilTuniil ; but »<• nmv nt fint nm-
uder tbeni oa one, bikI upvak of thi; piMutge from tli<- inttMiov inio the bliHiil
aa a dingle ovciit.
In Mpeaking <>f ihnc «ulwliinct« as liift'iinihlf, in an; uting the term in
referenoe I^i thi* well-known piiwngc of such xiilwlnnoes through tbin mran-
braii««or pon)u« partiiloiut. When a strung solution of sugar or of common
mII i* wniinikd by a thin nieiubrame fvfjgelable parchment, dead urinary
bladder, <l<!a<l intestine, etc.) fn?m a weak salution of ftiigar or of sidl, the
iUKiir or mU p«s»es with a certain rafHdicy Irom the stronger to th« weaker
■altllion, and naicr passes tVuni tbe wenker solution to the stronger; if, to
bwin with, simple water be aubfltitule<l for the weaker solution, thi? eHbct is
■1 nrsi still more striking;. I'eplone peaaea in tbe same manner, but. as we
hare seen, much more slonly. The procees is spoken of as a jthysinil one,
since it is not sct-ompnuied necessarily by any chemical chan^ in the dif-
fiising substauce, nor ia iliere any neccaMnr change in the njemhnine or
partition. The rate at which a luostanve A\ffuxii, and the total amount of
(litt'iieion which can take |flaoe, are deleniiincil l>y certain (|i]alitieti of the
Bubetaiice (wliicli ire mav call hhvsical, lliouuh they de)iend on the chemical
nature of the Bijhsiancc^ in relation to certain i|iiiiliiic:i of the membrane;
thus tnn f&hi may >titriiM! thmugh the mun' membrane at dilferenl ratea,
with ditfvreiit rattan in l)i<- iui«iiciat«<I ciirtruc cf wnlfr. thu o«m<itic current,
at) it is catted, I'nim llic weaker lo the :>tn>ti)^-r solution ; and the -ami- nub-
ataiice may {ia.-ii at dilli-rcnt rel*» through d^iHcivut nxtmbmncii. Bv a num*
Iter of obaervationx, in which various subftancw in S'ltuliim nndi acvenU
known ineinhmneo or partitions have bocn employed, a ct-rttiiii number of
" Ibw» of dtlTuHon " have been f«tiihli«lieil.
Now if by t)i<> statement that dilfitsible substances pan by difTusion into
ibe hloodTapillariee of the itiiestine, we are led to expect that the piuwag<.>
takes place exactly ticcortliug to the laws celabllshed by obscrvntions on
ordiiuiry aHMnbrtuMa, wc should be led into ermr ; for the disappearance of
these substanoea from the interior of the intestine doee not take place accord-
log to the laws which regulate their disappearance from one side of an
orainary diffusion eeptum. This can be aacertained by introducing solutions
of tbe substnncee, of various ^trenglb. into a loop of intestine, isolaieil in
tb» living animal by the raelJtod de«cril>ed in i '2oO, and waichiii;; their
dieap|>earaut'e hy anatyflin of the oontenl» uf ilic loop. >'u very lar^-e num-
ber of esperimeiitu hiive l>een maite in ihif way, hut «uch an have been made
all show tliv differenoe on which we ttrv dwelling. For ioKlancc, acMlium
sulphntv )HUBe» ihrtxigb an onJiuary difliiMim M-fitum with n rapidity rather
fCreater than ibat of dextrme, whereas dextniM! dioappcjtni from the intestine
diatinctly more rapidly than Molium »ilphate; |<ept»ne which dilTuititt vi^ry
•liiwly, indecil, through an ontinarv difPiuiion septum diMippcan rapi<lly
(lliiMigb not tH> rapidly hi lii'xtnnvj from the inlc;>tinr ; and when the delaila
of tlitr dii>ap|>ni ranee from the inteeline of weak solutions of two nails which
(lillUw ihriiugli an onliniiry mendiranc at dil)!.Tent rates, whicli bavu, a» it ijt
aniil, liillVn'nt <«molic enuivalentf , are stiidietl. these details arc quite diflcreiit
from those of ttnlinary dill'iision. Tits more tbe matter is studied, the mors
428
TUC TISSUES AND MKCIIANISMS Of IMOESTIOX.
tlwid^ly npparent becnmca the diHerence betuc^ti orduiar^ dilTiJiinn and
tlip nlMorptioii of ditTuHible substaoces from the iiiteslino.
Moreover, in such exj>erimeiiL! od an iaolHled loup of iniettine. the di»-
ap[>eaniiR'e of maleriul rnini the iiiteetine in iii.'ct>m[>«niei) by ilie ii))|>eiiriiaoe
of inatenni in the iiiteeliue. immely. proieid and other fiu)>alsii(.-ee ; these >K
derived from the blood. And the f(uestiim arises. If w allow oiiraelvee to
regftrd the passage of nmteriai from the iut«rior of tb« inteeiioe inm ih«
blooil aa carried out by onlioary [lifTiiAioii, why we should not reicard the
pAsag e of material from the bloixJ iiiiu the interior of Ibe iutntine ss beiog
also carrie«l out by means of dilTusion? Bui such a panoKe w« sneak of
el.Hevrhere us a " secretion ;" and everylhiiijt which we have hitherto learned
baa led un tn the concluitioii that HecruI.Ion is a diRerenl ami mneh more
complex thing IVom mero diH'uMoii. Kvmi adinittini^' that the snc«u> eo>
teri«us in of -lubordinale importuncc in carrying out dlgeMlve chatiKCS, we
cannot doubt that the glands of I.iflM-rkiihn wecrete, and mav with •mue
neoMO auppoM that the uolumnnr im^IU of the villi do m aW Henoe sren
if we ainimo the esixtcnt'c of nn ordiiiiiry ditTunion current from the blood
Into tho int«vtin<', aci^omnnnying and compK^racntary to nn ordinary dilfitrioo
current from thi' intcvltnc into the blood, wo urr comiiclird In admit that
with this llicn> coi^xi«t«, at times at all cvcnl* and in varying int'-n^ily. a
current of u iliffcrenl and more complex nature, n current which !• tbe
result of^M-rt't'iry activity. Atid r»iilt« which at fir»t fight seem cxplicabl*
by the former niny, ntYcr all. be due to the latter. Thus the Sow of water
into the inti^<line. with tho subsequent production of a watery stool, which
follows upon llic introduction into the alimentary canal of a concentrated
solution of magnesium or eodium sulphate, may at first eight seem lo be
simpiv the osmotic current passing from the weaker solution of tlie salt,
namely, the blood, to the stronger solution of the salt, namely, the inteatittal
contents. But the ditlerenoe between these effects of a dose of magnesium
nilphate aiid thiMe of a correaponding dose of sodium chloride are much
greater that) can Im' accounted ror by the dilTuMon phenomena, by the dilTer-
mg immoiic LM|uivali!iit« of the two ■ubscatictft; and the wore the matter is
Mtiidicd ihn more reastm linvc we to believe that the Bow of water produi^
by tlic former is to a large extent the result of nuddenly increased swjvlory
activity. So nliu> the fact thai tho contenU of the small intestine thrunghoul
il« length retain the name nmnunl of wntcr relatively to the mlidii, that in to
way. maintain tho same or iie-arly the mrno fluidity, whcn» in the large in-
tettine the water rctativelv dimiDi»hes until at last the fvcc» become firm
and even dry. cannot be wEioUy explained without calling into our aid varia-
tions in active focretion m distinguished from mere physical diiru>>i'>n. .\Rd
in tho case of a purgative, such as croton oil, producing a watcrv «tool. when
only a minimal, we might almost say an in6niteeimal amount olMta own sub-
Stanco can at aiiy one time he present in the intestinal walls, the result is
obviously due bi active secretion.
If, however, we are thus driven to the concliuion that the passage from
the blood into the intestine is a mflnife^tstion of secretory activity in which
epithelium cells play a imrt, gradually becoming little by little more intel-
ligible to us, why Simula we tiot admit that the passage from the intestine to
the bloiMl, which ae we have seen dues not accord in its phenomena with
known proccMts of onlitiary dilTusiou. is aluu brought about by the activity
of cells; is In hot ■ kind of inverle<l secretion, and hence tike ordinary secre-
tion prewota |miblems which cannot be solved by any off-band references to
known phyricttl pTooCMM? Indeed tbis U the concluition toward which
observation and experiment seem to be ntwidily leading us. Were the
alveolus of a Hilivary gland habiiiinlly tilled with a fluid of mixed and
FTltiN rnoJI THE AhlMBXTAKY CANAL.
429
I
rsrinl lutura like Ihe cootenu of tke aliiueoiarv caoa), mi; dboulil prubxMy
in nur *4U()y of ilt« f^lniid Gmi imnielv« ciiii)|ii-llr<J tu «|i(.-uk »f u ilouble our-
rrtit «■ cxbiiog ID ttie gluud. uf a curreiil fmiii (he oelln to ihe lum«n of the
Klreolua, and of k ciirrmu fmui lite luineu lo tJi« ck\U. And tJI nloiifi: lliv
iltlMCiilt both Uiv columnar niid ciibimi cells, wliicli I'vtiryn'hcre In^ir tlw
Mttka a£ bvlog " active " cell:), m*v |i«rlin)!ii be n>ciirili;d ni> eiiffiiged in n
lika donblo ftioction. Ovrr the villi (hv rts^ptive tunrtioii, in tli<' (.dniiilK of
LWbsHtuhn lh« rj«ctiv« ftinclion is [>rrdonnaivut ; but lu we linve vuggiwtvd,
i W6| in tbs zlmutla rrcvjAion probably ia tint ntinlly idmciic, mid uc may
haagiiM ibal in Ui« villi aomt amount i>f ejvcti»ti (ijiiitc njnirt fmni the
acdon of the gnblat 0«lb) idat t^o placv.
If this view lieaocqtted, ir we admit Ihiil the ciitran«> uf (lighted food
doM BM lalce olaoe by ordinary dilfunoB. the <)u«tion nmy U- niked why are
tbv dqfMlire cbanjm dircctiMJ toward Jnoraiaod difiuaibiliiy, why are protci<la
aoBTCrted into diflii»ible peptones, nnd why is starch converted inta sugar T
BMnaae iboui^h the cell is not nn apparnliis for diflbsion, diffusion is an
iiMlMiiMiiin uf which the cetl make* use. When we say that peptone does
■04 mtei tbe blood by ordinary diiTusion we do not mean that diHiision has
aotfaloK (o do with the matter. The nctivilv of a living cell is an activity
bQili op upon and timkint; u»e of various cbeiuical and physical prooewea ;
ia it thr proceaicii of urdioary difl'uuon play their part as do tlie proceaaes
of ctnlioary chemical decoupoeitiou ; but the cell uacii and inodiHea them for
ila <twn roda. If, aa we have every reason t>> lielievt^. Ihe eell of a villus
puBV the »ujrtr iinchauKed from the intestine into the blu»d (.-iipillary. it
nalm uw uf difl'usiuu l<> elleot that paaaage ; and if it doea chaii;je the prt>-
UM iniii ^'mt-ihin^ eU« before il paaaea it uu, it receiver it into itself in tJiv
8m iiiatniic* bv li«lp of diffueiiiin. When ne auy that subataDees do not
cnirr thv blood "by oriliunn- dillunioo we mean that tbe ditliiMOD which taken
place in m living cdl i« iu>RK'tliiiig m dittt-re^it in ihc results from ordinary
difltaMOB tbrough a dead mcnibntnii that it is undi-sirable to »pcak <>f it by
Uw MUBe nani«. In ordinary diHiision ihc n'sulls depend on tbe rcliUiun of
ibc noleculea of the diffusing substance to the minute por«« or canals or
•pwa in the diffusion Mptnm. Tlicso t-nnals or spaces are constant in an
ordtnarr septum : but a Kim of a living cell may be conceived of asa diUlision
septum the pores of which nn continually varying, and moreover lu closing
up or openin)* out at the touch of this or that substsjice ; hence the passage
01 maierial through tbe pores of a living cell lakes place according to laws
qviie dlflereni from those of ordinarv- tlinuifion.
{ SI4. The whole act of the absorption of subsloncw with which we are
dMling ountisiB. aa we have said, of two parts ; the pa/wage IVoin the interior
of tbe inlctune lhrou|*h tbe ejiilheliuin cell into ihe lynipb'Spnoes or reticulum
af the villus, and the paiBiij-e llience through the capillary wall into the blocKt-
■dsUB, In tbe experimeuis refernd tu above, it has not been possible to
distiMuisb between these two stages of the whole procew ; in each case we
ban nail to ualte use of the terms " from thv interior of the intestine into
tbs blood " and " from the blood into the inleriur of the inuatioe." Never-
llial— tbe remarks which have just been made may be taken a* referrinij
mont sspeciallv to the lini itiaji.'e. They lead us to tbe oonduaion tliui both
fills and dilliuible sulvtancea, tboii^'h in different ways, are carried into tbe
Interior of Ihf villus by tbe activity uf the e])ilbeliuni cells.
In mpect to the second siage of the abcturptiuu <>f dilfuitible substances, ]t
might be expected tJiat purl uT one or other uf ibcM •utMtanevs, part of the
sugar for inslauce, arrived inxide the hiuemeiit uienihnuie shuuld ulip by tbe
npillary bloodveswl nnd iiu.-i*inK tbniu^h the ntenbe* of the capilfary net-
work make its way into i1h- lacteal. And indeol, km we have Mien, S 30!(,
4S0
TUB TISSUES AND MBCHANISMS Of OIOKSTIO.V.
under ^■^^tniIl ciruumilfttioe* attmo Binouiit of NUgkr ii|)iMin> in uk<* ihw vourie.
But, tin w<- liiivG hIii) Mcn, undvr urilinitrr rirc'uiuMtino)^ thi> ciirn-iil. kIiM-
ever \tr it* I'xiu't nntnn', from llii' narrnw l}'ii)[>h-H|H>CN lyin^ iK'Inrvvti lh«
S'thcliiiin itixl the <-ii]iilliirv itil» tlir I)l()i>i1-8lri.-9iiii u strung enough l<> carrv
nr ticnrlv nil thu :<it^r uito Ihn Unnii. In lh« tautblislimuut of itiu car-
rent, in thi» iirc-onil siiigio of iibforplion itittuRiuii alwajs (ilnra a )>(irt. and
probably o Htill more ci)n«{>U'iii>ii» am) dccidnl pHrt than in the first eutge.
seeing that tliv fpicliclfoid plate of t.hi; capilliirv wall i« a far ieae active
structure than the olumnnr cell of » vilhis. ImWd it might be open for
us to i^uDtend that lhi« second ilage wta merely s matter of ditTusHH). what-
ever might bo the nature of the fint stage. But remen)b«rin|^ what was
said nbore, § 'M'-i, in dieciiating ihi? traiisuduiion of lymph, it seems more ia
aci(.s>r4lauce nith what we ali'eady know, U> coni.'hide thai iu thin seooiid
Btaee also diffunoD is the twrrant and nut the master uf tiw liviu]; capillary
wall.
A word may be adde^l concerniiix the special case of the peptoDOi As we
have said, the peptones in being alMorbed apj>ear to UDd«rgo a cbanffo some-
where in the inueuns nienibrane. We do nut know t-xiuitlv irbofW oroow tlie
clianffc talctri place. It seems probable ilmt »> uuirlcnl and difficult a cbaitgf
abouid require the iuterverilion uf Kiime iirtive living tiwiie, ami we may
therefore suppose that it is etrvctvd by the epithi^lium ve\W ; hnl wv have no
oxovt kuowleilge on this point. If the change bi- thn:> cjirritnl out by mHna
of tbe epithelinm wlla, tbeti the latter ittagc nf iIk' abmtrption of prottifls
namely, the passage from the itpiibeliom into tbr inlvrinr of ihr csinllary is
not a paiMugD uf clitluiiible [luptuue, but of minii' other nun dil1i]#iblu kind of
prot«tu. It may bf, howevw, that ihn i:)inngr l«ki» plaec during the very
passage of tbe material tliruiigh tlic cnpillari- wall.
The vi«w tlinl Icuutcyluti are the agcnli> of the absurption of lal, by bodlljr
taking up the fat into their cell-substance, has by some been extended to
nroteidg ; it has been urged that these Lake up proieids either as peptones or
in Bomfi other form and so carry them into the lymphatic system. But the
•rldenoe fur this view is even less c-uijviuciog than iu the case of lat.
AFTER II.
BfSPIItATHJN.
The i^rurrrvBK or tiik Ldn'os atcd IIkoxcuial I'AnAoai.
f SU. 0»E {wrticulir item of (he bodyV income, viz.. oxygen, is ^>«4-uliuly
■Mnciat«d with one paninilnr item of the body's waAte, \u., carbonic acid, in
M mnrh a> Ibe niemm irliiHi nre iip])lied fur the iutrodudiou of the former
■ra mho ut>«<l for the gettin]; rid of the latter, lioth are gases, and the iugnw
of Uw ofw a* well a* the ef-ren of the other is far raare (le|>eudcnt on the
rimplt physical process of ditrusiuii tliaa uii any aciire vital processes carried
on DT aimiu nf ti«ue». Oxyj|[i-i) passes from the air into the blood mainly
hr diSWan, and mainly hy iJiil'u-iioD also from the blood int^i the tismea; lU
IM matp way i-arlninic acid oattcs muiily bv diffunon from ihe tixsuw into
tb« bluotl. anit frum ih<- blood into the air. Wheremi. bh ue hiive wen. iji tba
arcTHion uf tlic digtvtivf juic(« the cpilhelium-ri^ll pluyn an all-ImportADl
part, in rcspinili'in iho vxtntiiM' uf oxyftc-ii fri>m the liingn into the blood.
Mid fnim tho blood into thi- tiiwii>^, and ihc iiawtigi- nf carhnnic airtd in the
ronmry direction, an- atTvclcd, if at all, in n wholly xtibunlinnte manner, by
ibe Miavior of tbi- piilmnniiry, nr of ibi^ cniiillury vjiil helium. What we
famrcbi di-al with in Tvspinilion, then, ill not ku mm-b the vilal activities of
any particular liaiU*, ■• the varion» mc«haiii»(nii by nbich ii riipi<l jnlcrvhaagB
between the air and the bbtod is eKcclcl. the mcnns by wbidi the blood la
enabled to carry oxygen ami carlmnic acid lo and from ihe tissues, and the
maniwr in which the several tiMnce lake oxycen from and give carbonic
acid up to ibe blood. We hare reason:* fur thinking that oxvgen can be
taken into the blood, not only from tho liing« but also to a certain small ex-
tent from the skin, and, as we have eecn, from (be alimeniary caual also ; and
<mrtmiic acid certainly |<aflse» anay from the skin, and ihniugb the variuiu
•fentioaa, as mcII aa by the lungs. -Slill the lungs are so eminently ibe
channel i)f the interchange of gascH between the biAly and (he air, Llmt in
dealing at the preaent niih respiration, we sball conllue imr^lvea entirely to
palmonary r^piratiun, leaving the consideration of the subsidiary rwpirainry
piucwu till He come to fiudy the secreltoDs of which they respectively form
pan. We may turn at once to ihv Htmciure of the lungs and bronchial
paMagta. including in ibe latter the trarhea but leaving the larynx until »o
coma to study ihv voice.
S S16. Tbi: lung takes origin a* a divcrliculuni fmm the alimentary i-aiiaU
and we may cuuiUler it a* a large branclied H[ieciBlly-twHlifie<l gland line<l
with mucous mvinbrane and eonsisliug of a r»nducting portion and a M.-crL-tiiig
portiun; the trachea, the two bronchi inln which thi> ([fvldea, and tho numtr-
ooa brotKhia, or (mailer jMUHgi-* branching out fnmi tbcae, rrnrawnt ducta,
oik) tho ■ecnrtinz alvtoli uf nn onlinary gtund an- reprcsenti-il by what we
shall |>rtaentlY descrilM as aim-lU or imlntoiiary alvooli ; but il must be
bomr in mind that. Mt we Have jurt Mia. active secretion by tltc cpilholium
Iming these pulmonary alveoli b reduced to a minimuin or pombty abient
•llnjRthrr.
Tb» complex strticturo of the mammalian lung will be rendered earier of
ooiiiprehervion if w* nrat Bay ii few words on the structure ■
luiiK. nut-h 8B that of ilie newt i>r the frojf,
1 be lunK of the uewt is a lun^ oval sac opeiiJiiK hj a short mii;:1o lironcbtu
into a very short trai-hea. It mav, by iDllatiiin, Ix.- largely ilMieiiiled, and
when the premure U renidveil collapara and sliriuks to a very stnsll bulk.
Its walb are, therefore, highly elastic, in tbe Rose in irbicli we have lo on«D
need ihnt wnnl. Tbey coiisiBt, like titucuus uienibraiie «bewhere, of au
epithelium resting on a coQiieciive-timiie baBiB, This ui>nDecliv»-tMMie baait.
which is very thin wh«n the lung ia diBtemled, conlniii* n very large number
of eWlit- Ubrcs of vbHoub »m-a but inustly iiniall ; iheae give the wall ibt
elaiilicity JtiKC spoken uf. 'the piilniciniiry arlery, carrying vcnuuii bluod,
«livi<les iiuir the neck of the tac Into br»uche.s which, running in tbe ooo-
Deetive (issne of ihi- wall, brcnk up into itn exceedingly clo»r-»ot nrlwork of
rapillurie» inimciHult'ly nndcrnrath the epltbeliiiui. Tbe cnpillntint are
tbcrnwlvos rvlatively ividt- but tho mwbi-x are verj" nurrow, being in mnnv
casein !<'» thnn the dinmclvr of a eapillnrv. Tlic epithelium over the whoM
of the sae consiMi' of a nngle luycr of relU. which, ('xicpt at iho iii-ck of th«
mc, are modified into thin philes in n »oni<-H'hai peculiar manner. Tbrwor
more cells converge together townnl the middle of each of the tslantls or
iDcafatt of the capillary network. The nucleus of each cell in placed within
the area of Ihe nie»h or iebind near the conveT^gence of the cell wilb ila
neigbbom. but a large part uf tbe call stretches over the capillary surround-
ing the i'land to meet a similar extenaioa of another cell whose oucleiia it
placed in the next island. Tbe part of the cell in which the nucleus is
placed, though thin, has some little depth, but the txirt of ibe cell stretebing
over the capillary ia reduced to the merest titm. lienee each island or luesa
is uceupied by the nuclei, and by tbe thicker parts of two, three or more
converging cells, while the capillary network surroundiuj^ the island i* »c|wi'
rated ftoni the interior of the lung by tbe extremely thin tiat oxjuaiMtou uf
cells heloiigiug to that and to the neighboring islaniis. The blood (MSsiiig
Ibrougb the capillary if m couM:?i|ueiRi-' separated from the air in the lung by
liothinji; luort llinn tbe capilbiry null ilnelf and a GIni, wbieh has not uveo
tbe Ihickni-ito of a Hut eptlbdium cell but is onlv a wing-like extension of a
cell itself Hal. The cttpilliirtii' nie in Ittcc Imbedilcd as it were in tbe epilhe-
lial layer. By tbi» nienn» iIj' [Diriilion between the blood and tJi« air is
reduced to alinoi>t the nnrrniM -i |iii.>.->ible liuiita. Ni-ar ibc neck of tbe (*e
the network bccoruvs more open, und nt the neck ibr pr<culiar epithelium just
deacribed somowbat suddenly chnngi-s into a single layer of rather short but
Otherwise ordinary columnar ciliated cells.
Tlic outer pari of the c<inneclive-tia»tie basis, away from tbe epilbetium.
becoming wmcwhat looser in texture but slill richly provided witb elaatio
libm, contains brHdes Ibo small arteries and veins belonging to the capillary
networks many small bundles of plain muscular fibres, chiefly running in a
circular or transverse direction, ^ninll brandies of the vagUB nerve Mtaa to
the lung, running in company witb tbe pulmonary veins; connected witli
thcM, toward the unper part of tho lung, are numerous small groups »f nerre
edJs. The nerve fibres, which are cbicdy nou-mcduIlal«d, though niedullated
Sbm are also present, end probably in tbe muscular fibre* or in Um btood-
Tenala. Branched pigment cells are also present.
S 317. Tbe lung uf tJie frog repeats, in structure, most of tbe featuras of
Ibe newt's lung Just describecT bm is more eomplicntcd. 'Hie cavity of tlic
Bac. especially id ita upper part, is broken up by a number of partitions or
septa prujectnig into tlie interior. Each septum is a fold of tho wall of ibc
cavity, and con^iaiji of a middle basis of connective tissue, cuvered on «cb
aide with epithelium. From these primary septa start in a similar manner
438
<f^m 7'*^*'^ tfl* of ■ similar -triitnure, projectiDg into tlM open ohnmban or
2ff .■"vUtMoTthe whole sac. formt^d by ihe uriiDary MpU. and dividing; UlSM
^H 'Mo MMller upcu t^liBntbera : uid miiny ol lti«te wcoinliirr ^teptn benr in a
^H ''Mikr nuiiKr ritniUr tentarj segilu. iliviiliiif; thv ttfcnnilnry diiimliur< into
^H '''^''^ chambent, or nlveuli. In lliiH way, i!ii|itfciiilly in iln ujip-r purt, llie
^p 'VritT of tbc lui))i i^ divid«il iou> a buntrymnili nf chnmlK-n or iilveoti, the
I *<iniilrf or tertiary alveuti ojH'iiiu^ iiilo ihv M^'imdnrv chnmlterf, the ncond*
"7 into ibf primary, anil tnc primary iiilo ihi? |cvn«ml <-iirity uf tlic lung,
I •fcitTh in lhrti{i[wr|«rt cif itic lung it rcducnl toawntrnl piM>itg<? surriiundnl
^m V C^M hnfK'yc<>iQt) work uf the <'hflm1>rn!k. In tiuwin^ down lr>>m lh« upjior
^1 t" Llw lower |Hirt of the lung, we find rhc wjrtii bccQiwc fewer, iind the hunejr-
^M **'>t>l> man o|X'n ; the Ivrtinrr aepta toon (at], then the Kecondnry, and at tn«
^B '*S^ boCtom or vnd of the lung mat the primary lepln arc nWnt.
^B^ ^Sach tcptuia conti«ti> of a niiddlp hati» ot'comioetive tiMUe, rich in clutic
^^^I^^Kmta, provtdcid with cI'mmxcI iiftworl<» of cnpillari^ and coverc) on eadl
^^F'^'^h with epttbelium, the elinracter' of (he fpichelinni and it# relati>>n to l)>«
^FJ^C^tllartos D^n}[ okuch the uuitc nit in ihe newt. Uence tn cuch Mjituiu the
~^«^ isfreely ex[>>K'dtoth«niroDeftofa«d«of ibenptum: and the ari«ng«-
**^*iti>f tli« hunL-ycuiiibii'orkof tbealreott iacrvasea laricely the total surnee
*^ l2?^ '" ''*" "''■ *'"' *' 'I'l^reaaeo the expoenre of the blood.
rhv y]sia muacular tibres present in the general wall of the lune peas to
^ Certain ext«ot into tite Mpta. A* in (he u«wl, at the neck of the sac the
— P^vuliar llal " rupinit'irj' cipilhtlium, for now we may ptTbK[iA so call it,
^^T«%aitgn into ciliated epitWIium: trace* of ciliated epithelium are aliio pmeot
H^% iho exIfvnM! endii of the M'pla.
^^L S 318. Kat-h of ttu- lobn (if whitth the mammiiliau Iuiik i* nnule up, maf
^^^« moKt, at limn aomfrwlint indiKtiuetlv, tn br divisible into lubulc. Th«
H'afw«*eAi». nr diriaioDH of th« right an(f Icl\ bronchus rcapectiveljr. dividing
*lirboli>R>oUFly, and running between the lobuIc« as interlobular bronchia,
wccdrupanied by branches of llie piilnit>nitry artcrv and ]>iilmi>iiury veins,
Qlutlly plunpf into and end in lobules as " lobular ' bronchia. Within the
lobule* the I'ibular bronchia diviilo in a mnre or !«« n?otangutnr manner
into snalJer " intralabular" bronchia or brnnrJiiJrr, often apukcn of aten as
tatetKthr piMtayt*. [Fi;;. l'^5.] Bicb such bronehinle ends in an eolaree*
toefit haviii)* more or Imb the form of an inverted cone, called an infundioH-
i^wK. [Kig. IJC.] Each infundibtilum repeals to a oerlaio extent llie strilo-
Uir* nf the whole lung of the frog, or rather \a inlernmliaie between the
laag of the frrtg and tiiat of (be newt. The more or leea conical eliamber of
ibo infundtlnilum narrowiuK into its broDebiote in divitled br a number of
■•pU into iocondary chambers of a (omewhat pulygnnal form, tne Mpta lieing
simple and oat as in the frojt bearing aeeotidnry and tertiary nuptiL Each «M
tblM Moondary cbanilien ix called an almoiuA ; it hiu a haw which i.i jiart of
tbo wall of tbu infundibulum, Hide) which are firmefl by the septa, and a
BMjath which openit into the miernl cavity of the infundibulum and *<i into
lb* brMtchiola. Kimilar but Icm deTdoiMsl iii-iitn arc prMJi^ted into the more
tabular cavity of the bronchiole iUelf. dividini; it, lea* oomptcttily, into
alveoli ; heneu the name alvwlar pn««g« ; iIm^i wholly ditinptiVftr balore tbo
bmnchiolc on its way out from the lobule bevnnics n delinilv tironehbini.
Each infuiidibuliim is surmtimlt^l by connective tissue carrying blood-
1 1 Ml ill and lymphatics. A number of tnfundibula with tiMir raspeetiv*
branoblolus are b-jund together by i-»nnc<^'t)ve tisnis varryinE larger bloo>i-
vmmU Io form a lobule, the bMiK-hiidr joining u* form the loTiular bronchia.
number of lobules are Iwond to^'ether with interlobular brrjuchiii and still
irgcT blomlveHels to form a lobe, and several lobe« join to form the lung.
'bm a lung is inflaied, and vrbcu as after deufa the bloodvessels arc for
I
n
43-t
RKiPIBATIOK.
1
l!i« moot jwrl cinplietl vf Uloud, tlie infgiiiiibiitii niih their ati-«oli form br
br tbs ]{rt'HU>r (iiirt i>f itie bulk of ibe Iudj[. Uenc« a sevtion takeu throufjh
ft hnrdtiMt) uikI iiirpurtil infliiud luiiKBcctm to be made upalmmt m holly of
n nuiuli«r of [N>l^gDDiiI ur fre<|iiirDtly Mxa^nsl spaces, winch are secliuoa of
alvfoli. iiml iimong irbirh iim imi>ii M-ctioiia in variuua plniiie of brunvbia,
Miiiill UH'I lat)M anil of MixiclvtwutlK ; here unil tlit^rc lli« sedioii may duclcMB
titi' ii|>«-utiig of a bmix'hioli- into uii inrMDiiibuhitii. aiiil tbe divtsiou of one of
tlw lobtitiir broiichia iiitu a nuiiiWr of hniiK'tiiolea.
fna. m.)
>^
or A Uiooiiv <Ciiii'rEfiinii>l iiuiiTt]> air* ■kmiki. litataltfil Ui liitm,! a. tvcmimt bron-
(llt>l 1«Ul: bit. lafUmikiLla , ( •-, alTn-Ur n—mii.)
IPm 19. -Tw" iuiii.1 aiMiii iir Aii-i'nu. oH tRTi'miiuut. u •■ : Willi klt<r<tllii. Iih, kihi Uia
klvtvla' [■■«««, *r. wtUi wliirk Uk •lr-«41* poniuunliaM. Prom ■ rmlwrn phlict. (Aler
KdU-KUt. I
$ 319. Tlir itifuiidibdlum irpeats in atnictnre. tu» we have said, llie liitiK af
Uie uewt or ibc frop. A wptuoi or uall bvtwecii two coiitigtioua alvenli oon-
liaU uf a thin mnliaii basis of coDiiective listuc, ciowded with a clow-art
rnpillary network, aixl covered on each side with an epilbeliuni. Tti« oon*
nective tissue is lichly provided with flue elastic fibres, but the ordtnaiy
E^latiiiifi-ruus fibrillft> am ituperl'eclly dcvdnped. lti« blood vetoela h«iiig i» h
rgc t^xteol imbedded iis ii nere in n I)<>ni0]i«iii »ui> maiiix. The acptniu,
Oipvcially loniird iia atiinniil, is ol'len no thin ibat thi- ciif>illury b cxpiiiH-d in
iM air on buth sidr4. TIh- ct^lla of llie rpithvliuin. which an iiiut'Ii boiler
■liOWD Id ttiv Itin^ of n yi>iiti>; iininiiil. and, iiidriKl. t* in tin: adiilt vrry difti>
eiilt to WCi art) for ihc iniut purl t niiiaf'otnti'd intofiiiall tlitl tmnHjiarvnt pliitca
iVxim which llic nurl<-i huvi' diMip|iear)'d ; tiM'ir millim-H may fir dinliiiiAly
slwiwii by lilvrr nilinli- (iv-nlnH-nl but oibi'rwiiMi an- uttcD v«ry itidiitliDcL
BotWKti tbi-M' cl<«r tint plat*-* iht'ri' iiccur •niall gmu)M of itIIh di>liii|iui>ihcil
hj powtMint: ntu-lfi. and by iltcir cell niitmliuicw i<tiini j.'rai)ular aiiil »[NintDg
wild ibe oroioaiy rciitfeiiiH. Tbi-M granuliir w\h, which nn! ihii-kcr ihnD
tlw clear platca. ur« placed in i;rou|» in iImi nie»liu» ol tbc capillary nrtworks,
•u that tbi' mpillnriw tlwiiuctvcs ant oorend only by th« thin ducImi»-I««
[dntce.
Tbo wall of tlic infiindibuluni which foniu! tbe bnM« of the ecveral alreoli
tins B Mtnilnr flnictiirv, aud Is Itoed nith au cpilholiuni of similar characi«r,
Ihe ch'H'f ditlcrcncc botwceu tbe sidea and tlM> base of an alvoliis bcin^' that
while ibc bUiul iit lh« caiMllurieg of (he latter is exp-iwd to the air of the
iilv(H>litiiuii th«aietid« nnly. Umt of lli« JurnH-r is uftcn ox|wm>I on both
guK-w ut' even ilio rnim* cftiMlIury.
() 330. Ill (lii>i'riliiiig the Immchial pntMgei w« hii<l perhnps bottrr Ix^i^in
with the irnt-hcH.
The (nichm consisU or ii cilintml muroUB nien>brtiti«, resliii^; on a cout
of connective lis<tie, st ret i^t lira l-iI with honpH or iiujtorfcci ring's uf cnrliln;^
BQ<i provide'l witli ik cerluiii aiuount uf plain muscular tissue. A vertical
section of tbe mucous luembriute showe nn e|>it}ie>iii(ii cunsislio); of tliree <ir
more layere of cells, thoao in tbe uppertuoet layer beiuji columnar eiliateit
eells ($9^1. nod thuot^ in the lower layers amall rounded cells, the cell Hnl>-
BMitce beinK si'anty in proportion to (he nucleus: it in suppcaed lIuU some
of ihcM email eells may at iiiii» develop into ciliaKtd <x]U in onler to
replace loc. Among ciliated cella are »eeu n trerliiin number of yoblet cells
(i 2lit!). Beiieatb the epilti«liiint nimt a Ikirly ilinliiiRt linwrneot inembraHC,
WkI below tbU in turn U two some lint! retieulnr tiiwuc, like tinti in tlie
unall intCMlinc <$260j, contiiiniup in it« lueNhc* a nrlnin number of
leiiciHryte«. Mtx<^ up with the reltciilur tiiwue. which in iliflcrent animaU
Tiiriifi much in ibe amount prrsent, arc tt*a ii certain but vitiiabic numlwr
of line T'liu'lic iibrra. Then Mruclnrca constitute t<welher tlw mucous mum-
brane. Ih-Iow which is b somflwhat conspiciwui layer of elastic fibres,
nrriin(;ed mor^ or Icra !n n network, but running distinctly longitudinally
Wid funning a longitudinal elastic layer separating the mucous mombrmiM
above from the looee submucous conncctire tissue below. In this submucous
tiwiie are placed a number of small mucous or albuminous glauda, like
those of the a;eopbagus, the ducts of which iMtasing through the elnslic laver,
MicuUr tbsue and tpiibelium, open into the csdaI of iho trachen. The
outer part of ihta submucous tissue forms a sumewhni denser cout of crninec-
live Uasue, ia which are lodged hoops of hyaline cartii»Ke, that is to say,
rinjca vihteli are imperfect beiiind. Siretchint; tmosversely between the
ends of each hoop of cartilajce are Mveral bundles of plain musi^ular fibres,
eomjil(.-iing the ring aa it were bv a lutacular band; a few lon^'itu<li»ally
tUajHixMl mnscnlur bundled may a)tu> be seen outside the trao.irerw bundltM.
Tmw two aelo of muM-ular lib'rM may be taken as being the remains of tint
original cirmplete double muscular cout of the atinieolary canal, almost
ol(liiera(c«l by tbe intrtMliietion of the carlilnginous hoop«.
TIm- main |^Mir|>aM aorvetl by ihrne wveral oCructure* iK (o pmvide a widt^
Bvxible clastic tube, the bore of which ntmiiins liirgi;! ami oiii-n iind tlw
Ibling ainooth during the branding of the tube. Tbi- mueoun (luid Mwnilnl
"by tM f[oblM evils and sniiill gliinds liel|iH to arnail ixdid parlicb* ciirrii.tl in
f>y tlw invpireil air, nhile thtt cilia arc continually driving that mucus, with
tite |>»rlicW cntniiglcd in it, upward to the Inryiix ami mi into iho mouth.
Tb" clastic layer ada|it» the mucous membnne tu the variations in the
length of tlw tube during its bending, and so keeps iCMHOotb. The traiw-
vera: niusoles by conlnicting can wimcwbat narrow tlie bore, when required ;
but llietr ellect in this direction can be slight only.
i 321. In iKUMing from the trachea to the bronchi an<i lari^er bronchia
Iyig. laTJ. tne chief changes to be observed are that tbe earlilagc« are no
DDger io tJi« forin of rei-ulnr booi<e, but are plate» place<l irre(;ularly, be-
^eoioinx smaller and more irregular in di»)iu6iliun the smaller the tube, and
that llie transverse muscular <lbn» beoome more and more prominent,
funning a distinct circular coat of some thickoeas. The carttlaKes, sup'
portt^^d by a fibrous coat of connective tiatue, lie entirely uui^iide the uius-
eular oiai, and the small glaudii have their ducts lengthened hi that the
bodio ol the gluuds instead of lying in tbe Hubmucona tiHiue, lie outatile the
mtiacular layer which is pieixi^l by their ducu. The tube become* now
I
ASm
RKSIMRATION.
i
diatiticlly u niiiMular tube, tboufrli ilie fMeoey of ita b-ire sod it c«rlBln
iin;4Niiit 'or rif^ditj ooinbiaed wilD tlexibility U Mill wcured by tb« «»!•
Ur«d plale* aod futlcw uf oirtibge. After acatb, amag (o tbe coatrftction
of lh« droiilnr niuncular flbraa, the mucoaa ia«nibraii«, like th« internal
i>Ml uf mn anerr in tb« nanw dmunulaDee*, b ihrowo into lon^tudinal
in*, in.
fABT C¥ * TiAlon-nH; Satkin or * Imomiiiul Tt nn rti'S mt l'i'< MnoitiM MOdUJO.
«,Mli«i«lniii«M>U)r«r: ft, nuntiilMr U}*r : r. viiMnal nbtmnlaxw; 4, vHO»'l'l
lajr^r;/, iinauf tlw (iimmlultiiii iltpatl. |
In tbe ntmllor bnMiclitu ihe cMrtiU^es dlaappciir alttmtlMr, nod tbo tub«
then coDsials of nn outt^r i-oiu of connective tiMiie wilh abiindnnt elastto
flbr«B KtA a oonaid«ni)ilc number or ciroaUrly difpuaed muscular fibrw, and
an inner cuni of iniicoiia membrane uith ita pVn elaatio layer ; the supply of
•mall K'nuds etill (xiniiniiee.
As one of tboee bronchia plungiof; into a lobule dividn into brooohioleri,
the culuinuar cells of the muootiit membrane lose their cilia, become shorter
•0 as to be cubical, and are disposed in a single layer or at most in two
Jayttnonl^. At tbe same time the muscular fibres become more scanty,
and lira iliii|Kieed not as a continuous coat but in scattered rings, tlic omnee-
tive'li>«o<> oint becomes ihioner. and tbe glands disapjiear.
In lilt- bmncbioka tbemseira an tbey prepare to opi'n inlu inlundibuin,
the c]ntbeliiim oslla beeoroe Sat Ihougfa sltll rvtaliiiuK urunulnr cellbudiea.
AmotiK thew, however, may novr be aeen iMtehw in which tlie n-Ua are Aat
tran*|>iirpnl plaleit, many of mhieli d<i not i>owe*e a nucleus ; ami toward tbo
JnAiwIihuliiin ibent: jmtchat iiirruuic in number until the Kpithi-lium niMintes
ihd lOinraclcr which w<' prtivinunly dcrn'rilied ax I'hamctrrL-iii? of xhf alveoli,
'llic nniKi-ular fHirra ilL-apimar iir Hprvnd out lungiliidinully, ami the [tre-
VMMialy compact layer cifcWtic lil)r(H now lnvixiini M'nttcred an<l upreail out
over the Blvi»>li of thr iDrniKlibulum and brimchi<il<-. In thin way tbe
rtructurtt of the bronchiole gradually nivritc" into thai of ati iiU'im1u».
Jr3SS. In an iDfuudilMiliiiii and in each of its conotilin-nl nivuoti what we
may oohskUt as llie origiunl wall of a pulmonary piiwsge, namely, a
tuucous memhraiH) iM-)>ara(ol br ■iibiiiiirou* conncciive li^iic from a mu»-
eular ooat. is rwlucod to a thin sheet of connt<«-tive tiwue in which bundles
of fibrilla' are wiutty or even nlwvnt, lUhl which In nitli«r to he considered! as
a Rienibnino of h»magi!n«o«is nature omtainiiii; imbedded in itself a larye
Uuiuber uf clastic libnm and Hbrils with a few connect ive-liiaue cnrpusclea.
STBl'CTDBie OF TOB I.UtfOS.
437
I
I
I
I
aiid tt network (if cftpillaric* ao olotte Ml ilint the iiK'.inbrttiie Metns to tw
DMrtilr elastic luatrriNl filling up thu lucahts of the nutwork. On lh<t uni-
m\c, ttiiii rtt|>illNry nu-nilirniiii, if n\' may M oil it, t^ oatilJnu'Mif iritli lliff
lootNir onliiinry conm-clivc tiMuo, iitil), however, ocmliiiiiini.' nt)iin<lniit oImIio
cIvnwnU. which ciirriw the »nmll nrtiM-int Btiil v«-iii« K"'i"li t-i anil ottiiittig
from tho ciipttlary network, nnH which uiiilm tUv inriinilihiilu ■ii'l hron-
cl)ii>k« inU) lohulra. On the iTtfiHo li«< the nltriiualivl i']>i I helium, all the
cells 'if which arv flat ao'l soiiic of irhich arc mere nucU-UH-lew plau«. Thft
muscular librra hav« either wholly disappeatvil or, iicmrOiiig (o eome
obMrvert, penbt u a few straf^'Ung Itbree s^trMtdiiig over tho infiiDdibulum.
The lerminal poriioo of the pulmonary pamage is a sac, whow walU are
reduced to Blmosi the grealest possible thioness oonsiatent with theii reuuii-
toir verr KTeal elastic power.
The bronchial poasa)^ of roediuni sixe nn eatentially elastic muscular
tubes, capable like the arteriea of varvinf; their calibre, but udIcm their
muscular fibrcH are Ihrown inl» unusually powerflil contractions, remaining
ulwayn fairly i>(k>ii: the iunaller ones, however, those which arc devoid of
oaniia)^, may perha^u cluoe by i-'illajxte. These paamgca are tineil by
miicouji membrane, the edla of whioh aiv well formed and active, iinme
aei'ri'tinjz mucuH, and iithi;ni by their diia driving that mucuH ouward (itwarcl
the tracnvu. The air wbioh pn«Nt» into the hmzM tx frcfpienlly lailen a*ith
im|>untii->>, thrw nrr cnlanglnl in the niu'-ti* of Uie pnfwiy™, especially tho
gmnlh-r on«i, and m aro i,-itli<T carrinl upward in tlie miieua, or lut wc ahall
Me othvrnixe di«p>»ed of.
ThiT latT^r piKKiigi-ii arc open flexible inhw beooroing more rigidly open,
and l<w> »UM«ptil>li' lo change in calibre by niitwular conlrnction llie larger
they art-.
§'3£S. The tungs are well provided with lympliati<*>. The reticular tiasuo
tmderlyiiig the epilhrliiim of ihe mucous mfmbnine i:s here ami there devel-
opei) into masses of true adenoid tixsiie eri.'wded with leucocytes, that is to
say, into more or leas oompletetv diHi^reDtiateil Ivmphalic follicles, and
similar Iblliclea are met with in deeper partA, Among the llat polygonal
epithelioid plates which form the eurfaoe of the pleural niemhrane inventing
the lung are numerous stomala (^291); ami during the rhythmic move-
ments of ihe lungK in breathing the lymph or scroii* fluid of the pleural
cavity is ooniinually being pumi)ed into the lymphatic ve«»eU of the lung<i.
These lymphatic veiaeK arising from lymph-spaoei iu all parta of the lungs
including (he runuective ib»tH! around tbo alveoli, ami running in the ooa-
nerttve liwiiie binding together infnmlibula, hnmchial liib«« ami bioodvraMli
into lobulea, and the lobults into IoIk-j, find their nay at la.-<t. alV^r lraven>-
tag MV«ral lymphatic (l>r»nchial) gianiU to tb<- roots of thtt luugH, wliencO'
Eliey pan) from the lelV lung Id the thoracic duel, and from the riglit lung U>
the nght lymphatic trunk.
The impuritien in thi* in.-<|iin-<l air spriken of above ux arnwled in tho
murns lining (he bronchial |>ii.i«i^'n>, ollon maki* (hvir way (hrough the epi-
theliom into the lymphittii's i>t<l<iw, anil, ''Hrrii'l away in llic lymph elrmm,
are niim ri^t«iu»t !n lh<' brxrichinl lymphatic glaml«. At limcH these glaiKb
boeoriM' in (hin way loaiU^l with jairlii-ti;* of carbon.
Tii4> blooitvnMU of Iho lung* do not call for any special comment mv«
perliape tliattha pulm<mary veins are destitute of valves; and that special
arteries, the bronchial arternv, starting from the aorta, are distributod to
the walls of the bmnchial naamges, (o (be blao<lvemeis, lo Ihe lymphatic
fdantta and (o the sub-pleural lieeuo, llie blood returning from them along
tne bronchial veins into the ngbl vena aivgoa on tlie right side, and into ihe
superior intercostal van on tlie left fli<le.
|324. The oervet to tlM luiif{a cuitie cliieSy frcnn rStrS^aST At, on <
■tde, tlii; vagun iiuTVt wiodl round tlv^ nmt i>f ilie lung. i( Kiv» ofl* in fnmt
bnuidte* t» form the viUrlor piilm'Hiary jilcxux. and l\ti-n. hehia>\. *t<iuret
bnuid>e> to form the prMlvrior ]>iilin»nary pirxux. BdIU llin*i\ but rtfii-
eM\y tbc luLUir, nn; joinnl hv liUroi-nU ff»ni lliu sjm^Uivlic tyslcoi, man
Miwcrnlly from lh« socDni), tliinl, unil fviirLh ihnrjicic guogtia; >di1 it ii
iiiiiinliuiiiM] l>r some (Imt Sbrv* piu» ilirccl from ihc K;>insl (inivrcoMnl)
oun'cf intu these piilnir>nnn- pIcxiiiiM. Thn iipp«?r p:irl of tbv trnohM k
supplied by twigb from the rocnrrenl laryngeal norvo on neb sid«, uid lli«
liiwer jutrt by tiriga (tracheal branches), coming diracl from tho vngni
trunks.
Home of tbe nerve fibres thus renchin;; the lung nlung the vagut nvm
are eRi-rent fibres for the inufculnr tibrce of the broD(?hial iiBSSages and tni-
chea. llut, oa we shall see, the chief and most imporlaot Bbra are aflereni
Kbr» c"noerDO<i hi Ihe regulation of respiration. The function* of the
fibrw <Miniux lV»m the ^ympatlieiic system hare not yet been clearly nccei^
tained; but (here h evidence that aonie of the fibres coming from the
thoracic gauf^Ua ore vaaoinoLor (c )Ustrict»r) fibrei for the |Hilinoniry
TCM^Io.
Thk Mr.'-nANK^ or Pi'luoxakv Rrhpiration.
$ 335. The lungK ure placed, in a ittnte which i« always one of diliteniion,
H<>mcltincii grrutcr, )i(>mi'tiincj> K'ju, in tho nir-tjght thomx, the cHvity ol
which they, logcther with the hwirl, great bloodvcMcU, and olher organ*,
«ompteI«Iy fill. By the coiitraclioii of cerlain niu»cl« tli« cavilv of tlte
Uiorax is enlarged. Thr lungs must tiillow this enlnrgcnie.ut and be ihem-
tclves enlarged, otherwise tho pleural cavities would be enlarged ; hut this U
impossible sd long us the walls are intact. Tho enlargement of the lung
CKUsiaU cbielly in an enlargement or expansion of the pulmonary alvmli, the
air in which beeumes by the expansion rarilied. That is to say. tlie piware
of the air ivithin the lungdi becomes leaa than that of the air outside tne body,
and thi» difference of prei»ure catiaw a ruith »f air ihrougli the tnicliea into
the lung* until an eriuilibrium of i>rei»4ure in entahliubed between the air
Hiiiidu iht! hing!< and that uuiaiile. This conxlituti^s inapiralion. llp<m iJie
Tvlaxalion uf lliv inspiratory museic i the luuNclia whuM iMntractiviu have
abmii Ihe thoracic cxpaosioii), the elaitticity of the lungs and ckeat-wftlk
aidnl ]NTh»ps tn lonio extent by thr coJitractiiui of certain muwiea, eauMa
tli<> clK^t to return to its original tiw: in coQM^ucooe of ihiH the prenura
within Ihe lungs becomes greater than that outudc, nnd thux uir rushw out
of the Iracben until equilibrium is once more established. Thi.t ooiwlitntca
expiration ; the inspiratorT and expiratory net together forming a rotpira-
tioo. The fresh air introduced into the upper part of the pulmonnry |>as-
nges by the inspiratory movement contains morv oxygen and less carlionic
acid than the old air previously present in the lungs. By difiuaion the new
or tidal air, ss it ia frequently called, gives up its oxygen to. and tnkcs ear>
lxM)ie acid fnuu, the oiil or /'talionai-i/ air, as it has been called, and ihoi
when it leav«e the ehetil in expiration has bc«n ihe meaux of both iutro'
ducing oxvgeii into the chest and uf removing carbonic scid from it. In
tills way, uy the ebb and fiow of the tidal air, and by dilfusion betweeo it
and the Malioiuu-y air, the whule air in the luugs ia heiug constantlj
rencw»l thM^i^h llie allrniate cxpnuiieu and eontniotiomi uf tM cbcst.
g 326. I» iinlinary ix^pinilixn ihc exfiannon of the ch«>t never reaebes
lis maximum ; by OMni funibli; munonlar eoulrar'lioii, by what is ealled
laboml inspiration, an additional thuracie vxiuuiBiun c»n tie bruugbt about.
THE M KCIIANICS OK PULMONAHY K£«fIRATIOX.
■ladlog Ut Mt iiiruob of n rertnin additinnil iguanlilv of nir licforc eqiiill-
bMB » fstBhltalteil. Tlii§ atlilUiunnl (lUHiititv ix oncn npulten uf at rom-
ptm*%lal dir. in the mtie way iu ordlnarr res^irntion th« <;»Dtractioii of
Urn ebnl mter rrnrhia ils maxiaiun). By nlliiiff into \ne a^lditioDsl mtnctee
Va ktntrd rKpirmiion an uddmonal uuantityor air, Ibe eo-cnlled reeenv or
'^plemmtal Htr ni«y l>r ilriveu mil. But even after the nnwl fi>rHb)e esfijr-
•tMO, ■ cnimtdKrabft! i|uuility uf air, llie rfilthial air, etil) remains in Ibe
'tngt Tbr nniunil i-riudiiioii of the liiiig* in itie chttt in tii furt oue of |iarliul
^■McBiion. TW i^lnKiii- ]iiiliuonar^' timie ia itlwayii (o a cerutiri extent on
*^ mtnteh; ii i* iilviirN, to lit «j>cHlc,*iriving to |tull aauuder the pulmonary
ftott) ib« [lariMAl pleura ; but thin it mnnm <\i<, Iieeaiife the air can iiave no
*oc«ri« i» tb« pleural caviiy. Wbfo. Iiowcvi-r, tJie clie«t ceoKtt ci lie air-
^Htfkt, abwi by a piinolnrc of lb<^ cbnit-wul! or <tiniihni)tra air \i freirly
^^BB'Bihofd into tlitt piriiriil •'hanihcr, the cliinlicity of tlif luii^ ptillii lU«
P*"! viiKianr a«ny from tlwt mrirtal jilcuru and iln- limgs (wtlBjw-i. driving
?*«*. by tbe Kio'l[iii>e n c»miiiiTiib!tt iiiiiiniity of iho rmiiliinl air. Even llirn.
^**^<«r. the lunn are nol compMfly emptied, mnn' air niill remaining in
^^^in ; Uiis is pn>tiably air tmprimnod i» tht? infiindilitilii liy collamo of the
W* K>atii'%o\n. whicb, as ne bare trva. have flnccid and nut ngid walk If in
R^ %Tiii|[ aninal the premurc of tttc atmoaphcrr continue to have access (o
j^^^Mttideof a Inng, Uw air thus iinprisotit<) if gradually abiorbe<l and Ui«
— ^^ *g beootnes solid. TIk- eanto result may occur fruio tho pressure of fluid
r«fliaUied in the pleural vavity.
^ It need liantly be added that irbeii the pleura is punctured and air can
^B^in Jnr admitlanoe from the exterior in the pleural chamber, ttince llie
I to the entrance of ihe air into llie pleura) chamber is far leas than
radatanoe to the entrance into the luuga. the cd^t if the respiratory
■mnonts u uimply to drive air in and out of that chamber, instead of in
out uf ihr lung. There 'u in cunwciuL-nfi.' no ren<;u'al uf the air within
I Inogs under tboae nrruraiiiiiiii.'i=i. If there Ih^ a suOicicut ubMacle to the
^mtrwocB of nir into the pK-ural tharolier, micb as a, fold of tissue blocking
vp lite opMung. the expannion of the cb<sl may Ktilt lead to a ilineolinn of
the luo]^ ana in this way in iu>nK- cast-* ixinmin- of the chest walU hn» not
•eriously laterfiifcil oilh m>|iirnti-m. Tin- parietal and pulmonary pleura
ar«. Id Domwl rirtuniBtances, separated hy a very thin layer only of Huid,
•0 that we may. perhaps, speak of them as being iu a state of" adhesion,"
taud) as obtains bolneen two wet mcmbranM sii peri in pitted. And it hashe«n
aOQBSted Ihsi lhi» adheeion, having M be OTercome before the two surfaces
out separate, assists in preventing the entrance of air into the pleural cavitv
after puncture of the thorax; but it has nol been clearly shown that this is
I really of iiojiurtance in the matter.
$K7. Before birib the lungs contain no air; they are in tke condition
raJlol atrl/et-ilif. The walla (^ the alveoli, the ^litbolial lining of which is
a] thai time well developed, consisting of distinctly itcticleated cells with
granular rell subMaace, are in eoniact. the caviiy of the ali-i^ilus not having
■a vet ninie into extMence; tho wslln nf the bnmchi'iln are similarly iu a
n>lla|iaMd cumlitiou, with their nalU (nuching ; the ni»re rigid brimchia, like
ibe trMchmi. puMwa* some amonnl of lumen whicb, howi^vrr. in uci^iipicd by
6uid. When the chest cximudit with the first breath taken, the pr^iuure of
the Sn«pi(«d air has li> ovcrottiuc the " adhesion." obtaining bvtwetn the walls
of iba alvcoili tbu< in ooiitacl with eaeb other and aUo thoae of the hroncbiolea.
Tho toKv e|>ent in thus uuaning out and unfolding, so to speak, the alveoli
simI bnmrhioles is conMdurnbTe, and in the expiration sucoreiling ihe first
in»|iiration most of tliv air thus iutroiluce<l remains, tbo force exerted br tlM
CM»t in roturuing to its previous dimen^tnns after the brcathtng in, ana tbo
1
440
R£SI'tRATIOS*.
«)«slir nclioD ol' ihe nlT«oli beioK inBufficient to bring ili« walls of i)ie aktoli
agaJD into coiilacU Succeeding; brcalba unfold lli« lu»},'i more and more,
until all the alveoli ami broiicliioles are opened ii]>, and (h«ii lli« whoki force
Df the expiruiory aci is directeii t»> driving out the i>revk>*i!J_v inspired »ir.
It in tKit. Iiowever. iiDtil douie time after birth that the luu;;* pft*i into that
flirther rlisteuded slate of wliirh we ajHike above. In a iiewly brirti sniuial
tliere ii no negative {ireiaure obtaining in the )ileural caviiits; the lun^
vbeu nt rast ore not na iW airtii.ch. aJid njieninij the ihorax tint* not lead to
coltajxH- of ilie luDgv. Tlie niuie of thiuga obtaiuiDz lait-r on i» «MnblijJ>ed.
not nt once, but grnilually, and in apjnrently brouf^t about br itie tbtms
firnwing more rnpidly, and no bvi'oiuinK relaltvrly moru raniidnuK than tlte
ilD^. The <}ii>l<-uti»ii of the liiiign in iWoiIolt may l>C tamiliiirly di9<!rib«l
M iK'ing diK' t<i the chcTil lit'ing loo larei- for the Inun,
S 328, In mnn the iimunre ext-rud by llm claiticjlr nf iho lung* atoiM
ani'iiint« lo about 5 or 7 mm. uf mimrury. Thi* i» wtimati.'d bv tying •
mnnonictcr into the windpipe of n dnd Mibject and uWrviug ihv Hm of
mercury wbith takes plac* when thr chc»t-vnill» an punctun-d. If wo took
7.6 mm. as the prnmrc thin uonld be just ^ in of tho pressure of the atmo*-
pbere. If the che«C be forcibly di»tcndc<l beforehand, a much larger riM of
the mercury is observed, aniuuntiiig in the case of a distent ion corresponding
to a very forcible iuBpiration to IJU mm. In the liviiif; body lliiii ntecbanical
elastic t'orce of the Iunf>i may be nwtsled by the contraction of ibe plain
iiiUBCulur fibres of the bronchi ; the prej^iire, however, which can be <>xerl0il
by th««e probably doett not exceed 1 or 'i mm,
Wlien a manometer is inlroduced into a lateral opening of tlie windpipe of
an iiniumt. ihe mercury will fall, indicating; a Degative pr(^iin.-.nH it i:' called,
duriac iu»|iiniti'in, and riiu-. indicating a positive prcmuri^.durini: aspiration,
both fall anil Hm? l)i.-ing Blight luid varying ai-curding to th« mnlom with
with which tlic air pnMO in and <iiil of ihn ch<»l. AVhwi a mAnon>M«r i»
filled with uir-tight cliwure into tt»r nvnith, or Wttcr, in order t'l NVoi<l tbo
suction- act ion of the mouth, into one mtilril, the other nostril and thu mouth
being closed, and eR(>r(» of in»pimtion ami cxf>initi<m are made.tbr mereury
falls or undergoes negativt* prowurc with in^fpirntinn. and Hm* or undcrgoea
positive preHure during expirnttun. It Iiab hmn found in this iray that
the negative pressure of a strung iiispinitory effon rasy vary frDOi 30 to
74 iDin., and the positive prcBsure of a strong expiration fW>n) 63 to
100 mm.
The total amount of air wliicb can be given out by the moat fhroibte
expiration folIdwinK upon a most forcible inspiration, that is, the sum of the
eumplomenlal. lidal, and rteerve aire, has been called the " vital capacity;"
"rxlreme ililliTential capacity" is a better phrase. It may )>e measured by
a molilication of a gnA-inetor called a t/iiromrier ; and though it variea
largely, the average may be put down ut :iOUO-400i) cc ('iOU to 2A0 cubic
iucnes).
Of tlic whole n>oiL#ure of vitid rapacily, about 500 c.c- (.10 cuhi* [nchni
may be put down ai* the avemgu nniouni uf tidal air, the reoiaimJcr boinp
nearly C4|unlly divided Iwlwceii thu coniplemcntal and rfsurvi- air*. Tbo
Juaniilv Icll in the lungs aflcr the dccjieal expiration uinoutil* to about
400 or 'iOUO cc.
Slice tbo respiratory movemcnta are •» MMily nflccled bv vRrion* drcumslaoccs,
tkt aimi'le fact "f aurntion \mng directed to the breatnlnfc beiag latHcleat to
eatiM luodificalioDH both uf the rnte and deplli of ihe respiration, it Mcomaa rtty
dilBcult (o llx itie volume of an atersge bmth, Thun varloua authors faav« givM
ligMm nrying from M c.c to 'ff'i c.c. Tho statement made ubove i> the mean of
oStrrration* varying frem 1"7 lo itSW cc.
THR lIRCUAyiCS or PtTLKOyART RKSPITt ATlOy. 411
I
I
I
I
ii 3M. Urnphie monU of rupiratory moivmrji/^. Tlidw iii«y Iw fll)Uin<i)
in uan,v various wajra.
The limplMt, readirat, Kn<) pMbapi th« inoHt generally tHeful meili'ii] U thnl of
recnniinjc ihr movrmonu iiT the (oliimn of air. ThU inajr be effected hy iotro-
dncins a T'l'i<'(-'*^ >"*'* tliv trndira. one cmw-piwo boinx let open and llio otiior
coBuocinl witli ■ Jlur^y'i taiubour or witli n ri'ceivor. which in turn in ci^imi-ctrd
wilb a UuilKiur fsee Fig. 73 and Fig. liSi. Tlie inoveiui-utH of tbe column of nir
in tbe Iracbe* are traoKmiUed to lli« taicboiir. the cnnsequeDt expinHionH and
and eoniriu-iions of wlik-b urn Iraiumitttd to lb* recording drum hy meant of a
tvrtt rmtini! on it.
If. a receiver bc^ng UMcl, tbe ofien end of th« H be cloaed. tbe nnimni lir<-Hib<»
into aiid out of (be receiver, and tlie inoretnentn of tbe tambour are gmllj in-
creNKcd. TIih h»'> (b« diM'Ivnntaga (hat the air in the receiver *onn beoomea
unlit (oi further reapiraiion. \ aimilnr incr*<w« of the mnremeuta of tbe levar
nf Ibe tainbour may be olitained by coancctine a pie.iv of India-rubber tnbitiit to
the open end of tbe h* - I'.v increaiilnK tbe lenglb of thb lube, or sligblly e»n-
•IrictinK it, tlie niovemenia of the lever niav be increased triiboul very seriously
intrtfrrinE vilb the breathing of tbe Btiimal.
In nn'ither method the mnvcmeocii of the cbnt ar« recorded. When ii mnall
Aiilinul. Micb an a rabbit, in iii«l, (ho wbolv animal may be pUtvd in un air-tight
box. breathing bvlng earried on bv tne«n« of a lube imterled into the traobeft and
carried (brongli an air-tiitbt orifice in the wall of the box. Ity anothier orlBco
ai»<( lubo tbe air in tbe bm i* brought into connection with a tambour, which
aeootdingly regiKlm the chnnKcs '>f prewute in the air nf the box produced by
the iiwveneDta of tiic che*l (and body), and thu» indirectly tbe movemenia of
Uu> chtet. In man and larger anim^a tbe changes In the girth of the cbe«l may
be convenlraily recorded by mean* of Maray'a paeunioKTii|!b. Thi« uoneiKt* of a
hollow elantie cylinder, nr a cylinder with clastic coda, the interior of which U
connected with a tnmlxiur. Br lucan* of a «irap attached to each end of tbe
3 Under the iniitruQicnt can be Luekled round the uhesl like a girdle. When lb«
M expand", tbe ends of the cylinder are pulli-it out. and Ihe air within the
chamber rarefied ; in coniic<iiieni'c the Icrcr of the lamlKiur connected with ita
interior ii deprewcl ; ■.imTerHely, when Ibe eheiit oontrnctn, the Icrcr i* elevated.
The pneamatigrui-b of Flrk i* winiewliat alnillar. Or cbangai in one or other
diameter of the cbcst may 1>e recorded by what may be called the " calllpeta "
method, tm io the recording Htethooielor nf Itnrdnn ^Anderson. Thi* conaUl* of
a rrctiinitolar frumcwork MiDilracted of two ri^id ]iiiriillct ban joined nt riKhl
angln to a cnitu pirre. The free eodii of tbe barn, tbn dintanee betwueu which
can be regulated at p1ea*nre. an- armed, the one with ■ lambour, Ihe other aimply
with an tvory button. The tuml-our bwara on the metal plate of lla meonbraa*
(w'. Fig. 73^. a i>maU ivory button in place of the lever. When il i* desired to
record the rbnncM occurring in any dianietvr of the cbnl, r. •;,, an antero-piw-
tertor diameter from a p»int in the aternuni to n pidnt In the back, tbe inatrnnteot
la made to encircle the cheat somewhat after the faahion uf a puir of oHlll^cn. the
ivory button at one free end b^g placed on the «pine of a vertebra behind and
tbe tambour at (he 'tthct on lh« sternum in front in the line of the diameter which
b beine alndicd. The dtitance between Ihe free eniN of the inatrutneut being
car«fulTva)Uuit>-dM>thut the bviilori of the tambour ptetitw lightly 00 the oti-rnuiii.
any variation* in tbe Iciivlb of the diameter io iguottion will, aince the fraiuewoik
of the lanibout in immobile, give fine to variation* of preacure within Ibe tam-
buur. Theao vnrintinnanf tlu: " reociving" tambour, as it !■ called, are cnnreycd
by a lleiible tube containing air to a aecuod or "recording" tambour, the lever
of »bi<.'b record* the Tariatlona on a travelling surface. For Ihe purpoae of meas-
uring the extent of the moveinetitM iIk inHiriiroeot miMt be experimentally grad-
uat*<l. <)tbcr form* of ciillipem may, uf roun«, bo need.
By Htill another method the variati»n» in intra- thoracic preuure. by meaim of
which Ihe movementi of thr cheat wulU proiluce the movement of air in tlie
lungs, may be retorileil. Thl« mav be eTeried by introducing carefully. Io the
tolal excJtmion of air, inioa pleural cavity or into the pcricHrilial cavity, a canulu
connected by a rigid luW with a manometer. Wilb each inapiration a nqratlre
preasnre, or mihrr an increaae of the exiating negative ixcHure, in produced, ibe
meroiry, or nuiil, In the manometer reluming at Mcfa expiration. An easier
THK VICCHA^*tCfi OF IT I.MO.VART KESn R AT1I}:C . 448
I
nielbod o( neotAiof ibia iiitrH-iliorHcic tir««iaui« is t<> inlroituc« Into Ibe i»oi>Iuj;u«
aDclM«i««»iiiitl (lUmtlai-Ui Ihc cnrduc tound, Fig. 73) connei:t«d withaunbour.
TIlC <BIOpbBf;iit within llin tbutnx, tikr Ilitr liriitt .inci gtr-M vn>vU, a* w(i nhall tier,
UafleciM iu«'i.-ll on till' luiigB tiy the THtUlii>n* of intra -tlKinidi- pmuure bmughl
alxiut by tlit^ reKpirutory [novMrenu.
In yet itnolUor nieibnd the morfineata of the dUphrnj;!!) whicli. M we shall
■M, Mrrre >« iho prime affrnt in bringinj;; hWiI iHp wlnrgcinpnl of lh» thomcic
cnvity att recorded. Thi* rony be lione by imwrtiiij!. lliri>U)cb an incisiuii iii the
nlidocalusl wall, « flnt claatio b«( tie(i>e«ii (be diapbragin and abilnminal organ*.
Wbea )a Inaplniion ibe diaphragm dcaceoda, itvxarln oa the bag a preaenre
which, bv meaiM of a lube, may be comrauoicated to a umbour. Or a tiwdl*
mar bo (IirD«l through th« clitt wall «o an Ui rvsit upon or imntfix the diA]>hragiii.
anil tbe hMil of lhi> nt^edlc iiulaide ibe b'ldy connected b;r a tbri-ud or othern'iM
with a lerer; e*cb u|iwitid Kiid downward ti'itvement of the bead r>f the ne«dle,
GorrespondlDg to the downward and ujiwRrd DioTemenli vf the diaphragm. U
r«Kiat«red by the leTcr.
VarioM inodificationii of llinie Mivcrul metboda have been adopted by varioiu
obtierv«r». They hII. however, leave much to be devirad. A very ingenlout
Dielhod of register! nff tbe conlruLliuaB of the dUpbra(;in ban recently been iii-
tmdiKNs]. In the rabbit two allmofmLixcular Abree forming |>nri of the <liii|ibrjiK"i.
one on each »idc of the eniiiibnn cartilajtc, are »o diapiMcd and p'la'CH* nuch
attaclimenU that one or both of them may be imlated without injutr to either
nerves or bloodraaebi, and arranged no tlul while one end of the iilfp u wcurely
fixed to (lie cheat wall aa a fixed poinl. the other end can by a thread be brought
10 bear oo a lever. Tlie «!ip, even when thus arraoged. apiwin to coiiirucl
rhyllimEcally in complete uaitan witb the contrnctiottn of the whole reil ot the
diaphragai ; it aefvee, eo to apeak, lu u mniplo of the diaphriigm : itiid heoci^ itfl
contraclToua, like tboM of tti« whole diaphraijra, muy be taken ai a record of
rmpiratorr ninvemvnU. The record lias to be correlated for rarialions in the
pmiitloii of the lixcd point.
Tautm ae TituRjtcii) linrtiiiniBi UovcKim oiir*iiiu> nv nutMor Mtaiv't
■■HieHuulitrn.
A atalii raaplnilMjr t'lu* ■• n><ii|>rlvl beiaotn a knil n. IntplnilMi. ditiluf nUrli ilie lover
it'oviMb. exwwdlng l>uni« lob. uid eiiilnUuii trum Ho». Tliv iiiidutalkai al eafeMiuol Iijr Ibn
hiMii'* b»t
^ 330. In these various wap curvce are obtained, wbicb, nhile dtlTering
to deUil. exhibit the mmo g«iwnl fcMlnree, and more or \vm rescmbte lh«
curve ahoii'ii in Vig. Vi^i.
Mv«adf4ecec4 lllMt>(^, liwi'irJ <•< obxii i9uu< cither elCMd or twnUlly otmnpcnl at flouiue
\yf Meant or the tfwii ^fninii ^ From lh<i Jat fintri^eilt a thlnl |ilv<^ *'f tutitiijc if, rnfuiivE<il wilh a
Mfnr'* laatioi*! m Im 1^ T3I, (bo leivr ul wblcb I wrlM on the nonMliiR fiiifaiY^ M hen Ihe
inlB li |» iiptB Ills uiIiibI bcvatlM* ftMly IbKiiiali Ibf*, and ttia louvoinunti In Itir *U nf 0 aiHl
mn>»|ucn<>T In ilie uiaIbui an >nil>t. i>n<balD| Ih* tikmpi: lt>c nolniftl timilm ouli ilio atr
omuinttl In Uh it. ■iM ihe luovnseiil* of Ibe lovor ut Uio Mataar booMno aHWotneiillT nnieh
mi-ni niarkM.
B DcloH llio lever te vcn * tuMll Umemtrkfr ■ CMtneeWd iillh an clMtmmacnol. Ihv eunenl
H UinniRli aliicb. niminx nntn* lallory It* I he iHpb i ■lul |r. It madoaml tirokco by ■ clock -nntt o«
^B lUMaiuiuc,
144
KKSHlBATItlX,
Ab itie Rgttte ahon;), jiiRpifiiiion begiuti sumewhac suddtmly aud itdviiiioc*
rapidlr, bc'iiig follnweit iinmeOtatel]^ by fK|iirnltuit, which i.i carrk-il iiutu
flnc nk[iiilly, but iincrvrnnl nmna aixl iiion- ilimly. ivueh tiaiuet lu are teta
Ufluidly t'LTiir Iwlwetin llif cm! of cxiiinitiiiii Atid Hut bi^niiing uf Entpin^
UoD. In tiorinnl bmithiiig liunlly iiny •iicti {)uii»c rsi*U, Dut in cum nhero
tJie rvupirutinn twuoni*^ iiifruu(ii-nl. pauHci' of confide rah 1« Ivagth may bo
obiervuil. An wi; fh»i\ wo id (ictail noreafter, ihc (rvural jmrW nf ihc wlwlo
•Ct vary iniioh. iinilcT varioii* circiitu stance*, in rrUcion lo each othct.
SomotiniM oxpirntion. soiui?[imce inapimtion in prolongs] ; aai) dthor id-
•ptralinn tir pxpiration may be slow iir rapid in iu development. At titno
thv dic»t may remain for a wliile at thf- height of inspiration, thus Risking
a pan»c bctwenri ioBpiratinn and expiration.
In what may be ocin»dered aa normal breathiu;; the rwnir«t(>rr act i» re-
pealed nboiii seventeen iiui« a miante. the duration of ttie iuapirntion u
oomparvd with that i>f the expiration tand *uch pause aa may exist) being
about as (en tu twelve; but the rate variet very largely, und in this, aa in
the volume of each bi-ealh. it is very difBcuU to lix a HaLiAfaclory avera^.
the fii;ura given vurying fruin lwi.-iily tii thirtt^n a niiuule. It vorica
aooordlag to at^e and »<>x. It i>> iuflnrnciHl by ilxt piwitiiui of the body,
bMDg (|uicker in ulaiidiiig than in lying, anil in lying than in titling.
Huscular exertiim and emolti.inal onditiimi' atlcct it dwplr. In lact, almost
•rerv event wbirh ncciini in the body mav inlliK-ncc it. Wc ibnil liavo (o
consider in detail licrcaticr the nmuiicr in which thc«c inHuimmv an.'^ broiiglit
to bear.
Wliru the ordinary resniratory movemente prove insnlRcifint to ofTuct tbe
nec<THary chiin|,'e«' in the blood, their rhythm and character l>c«ime cliiuigwi.
NiinunI K^pi'nition gives place to labored re«[iiraliou, and thia in turn to
dmjmiai, nhich, unless Bonit- resiorutive event oecur», urminatce in OfphtfTM,
Tnme abnornml enuditiuii» vre Ahnll tUiAy more fully hereafter.
Thr Rftjiiraiory Mavement*.
S 331. When the movenienls of the chest during noriiud breathing are
watched, or when a graphiL- reconi U taken by one or other of the methods
just dcAcribed, it in seen that during inspiration an enlargement takes place
in the antcro-piMricrior diameter, the Hternum being thrown fltrward. and at
tlw nuDo time muvinj/ ujivrnnl. The lateral width of the cheat is also in-
oreaMd. The rertical incrcaw of the cavity is nut so obvious fruni the
oub<idc. though when tbemOTumentitof th«^diai>hnigm are watched by meanf
of an inserted needle or otherwise. It is okar thut the u(>j>er iturface of that
organ deeoeods at each inspiration, thr nnU-Ti»r wulU of the abdomen bulging
out at the Ktme time. In the female human imbjrct, the movement of the
upper {Hirt <jf the cbe«t is viTV conxpicuouK, tliv breiuit rising and falling with
even' respiration ; in the male, however, the movemenlit an- n1mi»l eniiixlv
ODO&tcd to the lower part of the chest. In labored rt^piration all iiar(j> of
the obeat are alternaiolv expanded and contracted, ihi^ brca«t riiing and
falling ID well in the male na in the female. We have n<iw to c-ini*idur thoM
several movements in );reater detail, and to study tlie means by which they
■re carried out.
$ 339. Intpimtion. TItere are two chief moans by which the chest if
enlarged in Donnnl liisinration, vix., the d«neeiit of the diaphragm and the
filevaUim uf the ribn. The former causes that movement in the lower part nf
the chest aw) alxlomrn ki clmracieriiilie of male breathing, which is hence
called diaphnigniiitic ; tlw latter oauMH the movemeu of the upper diert
TBE UKCUAKICli OV PfLUOV AHY RKaPl It ATION .
446
rlinrAclerittttc of t'vmiiln bn^Uliing. nhich is tMillfd cimuI. Thmc Inro innin
Jmclun are umuOocI by \v>% in){H>rUnl untl aulwitliitrr <ivi.Mir«.
Etl-ii iu the female human »tibjccl, tho nhan.* Uik«n in rcspiniti<m li,v the
Hi«j>bnifm is no imporliiDt oik-, in the miilo the (lisi>hrngm must tw rcgmrtletl
as ihe chief rrapiraloTy ai;eni, and in gome iiiiimnls its n«e, for this piiq>08e,
is 90 profuinent that the movements uf the ribs may in norma) hrealhin^ be
aJuKMt neglecled. In the rahbiL for inMimce. in normal breathing, almost nil
ibe Kepiratory work U done by the conlr ael ious of the diajihra^'m.
The dcfloent of tbe diaphra^fm is etit-L'tt^d by menus of the (.-on traction of
its m«»-ular tibrea. When at reel the dijiphragin preaeuis a oiinvex surisM
U> lli« thorax : nht^ii cuntrscted it beeomen much nailer, Hiid ill euusequeitoe
the level of the <^hi«t floor h lowerod, the verttcle diAm«ter of the cliest Heing
pr»)M>n innately ^nlarct^d. In d«ecendiii|{, the diapl>ri^;m prtesea on th«
alxloininAl viaceru, niiS bo canm a pn>j«ctii>n of tb« flaocid nbdotniital whIIr.
Fmiu its altachmNitx to th« M«n)um nod the fulve rilu, the diaphracm, while
contracting, nattirully tttudit to piilt the Ktenium and the ujiper fiiiao riba
downward ami innnnl, and i)m! tower falae rib* upward awl inward, toward
the lumliar »{>iiH!. In normal hreathine, this tt^ndency prodiii^ea liltl<! elTecl,
betu^ cnuDtvmclvd by th« nrcom puny inn general c-ohuI ctevatiou, and by
cenain sfiwial muscles (o ha nicnt!4)niil prwcutly. In foroe'l iDspirmtion,
hi>w<^vt'r. and muocially whvre Iherv w any olMtniction to the entrance of air
into ihc iLinn, In* lower ribs may bo f> much dniwii in by iIm: nmlractirm
of Ihi- dinpnmgia, tlial the girth of the trunk at thi« point t« ohvimisly
(limiiijflhed.
S 333. The «t«vatioo of ihe ribs it a murh mort- complex m:tlier than the
cloMent of the diaphragm. If we examine any one rib. Htich lu ihr fillh.we
find ihat while it n>ovea freely on it vertebral aitioulati<m. it iucliiMi wlien
In tbe iMisilion of rxvt in nn obUr|iie dinn-tion from (he tipine to the sternum ;
hence il is obvious thai whi-ti the rib in miiuid, itx xlornal ntlachmeut must
uot only be earricil upward but aimi thrown forward. [Fig. PtO.J The rib
uiay in bet be regardeil a« a radiuy, moving on the vertebral articulation at
n centre, and causing tbe sternal attachment to descnbe an arc of a circle in
tbo vertical plane of the body ; as the rili is rvrnvtl upward from an oblique
to a more horizontal pucilMn, the steninl attachment must of iMceMity tie
atrriml <\irther away in front of the uplnc, Since all tW rib« have a down-
ward slflutiu); direction, they must all tend, when raised toward the hori-
xontol )>0!dti<m, lo thrust the slcnium forward, some more than otliCffl
Uti
BESI'IBATIOS.
according; to their slope nnU length. Tlie claslidty of lli« ttwniiini and
cu^isl cikrtilages, swivteti hy ihe nriit^ulsiitiii ill' lh« Uenium to tite clxvk'le
above, pt-rniiUi the front Aiirtiioe of the cImuI !•> )>e thu> (hriwi I^Twmnl a*
well a* upniird, nheti tlie rttia are niJMH]. By ihu actiou, llic niilcni-|H)»'
leriur dmmeler nf tin- cheat is i^iitnived.
^iiicc Uii! ribit fiinii nrches which uietvaMi iu th«tr iweep u ouc prtkceeili
fniDi the lii>l ih)wnu'iiril nn tkr at Iciist a* the seventh, it !» (<vi4luiit that when
a lower rih »iiih iw ihu fiftli is uicvaicii ai as t«) occupy «r lo imiiroach tnwanl
the {Mfitigo »f the one above it, tli« cheat at that level will l>ec<tnir widttr
froDi aiilo t» ridv, in projiortiuii n» tlip liith arch i« wider than tliv fourtfa.
Thus (he elevation of the Hh iricreii)>c8 not only the an toro- poster ivr hiit alao
thir tratiBver«e dinmeler uf tho chest. Further, on account of the rrai»l«nc«
of th« sleniuni. the nagles betwecti the HbH and ihoir cartilages are, in the
elevation of tho rihs, somewhat opened out, and thus al*o tho trai>BVcnM> na
well as the an tero* posterior diameter, somewhat increased. Id more than
one war. then, the elevalioa of the ribs enlarges the ilimenHions of the chat.
$ 334. The ribs are raiwd by the contraction of certain muscles. OrUma
the external interci'v^talB are perhaps the most important. Kven in the esae
where two rihs, tiuch as the tifth and .'>ixth, are isulated from the real uf the
thoracic caee, by sectiun of the stnictiifi^ occupying the iuiercoetal kp*obb
above and below, the eontrui-'lion of the external iniercuolal iduaclfl of the
iiitertening space raises the two rilis, thus bringing thciu toward ttie posilion
in which the librcs of the ntuscle have the sborie^^t lenuth, viz., the buriiontal
one. Thiii elevating action w. in the entire vhon, furtner bvored by the fact
that die firat rih is le»s mciveuMe tbuu the decund, and sn aflonU a oompan-
lively fixed bate for the action of the niUHele> between the two. the wcond in
turn supporting the third, ami so on, while the Mcali-iii aiuscles i» (ulditiou
«rvc to render fixed, or lu raiw, tli« firel two rihs. So thai in normal re^|i(ri>'
tion, the act may probably hi- de»cril*ed as beginning by a contrac-ti'in uf the
ccaleiii. The HrM two rilis being thus raised or at lca«t Sxi-d, the contnietioD
of the leric:' of exlenud intercostal muscles acts at n great disadvantage.
While the elcvatin^f. i. e., inspiratiiry action of tho extcmid intcro(«lals is
admitted by nearly all aiithon, the function of the inlcrnnl iiiterciMt«ls hie
heel) much disputed. Home ret,-ard their action as wholly inspiratory ; ollieiv
mainlaiu. what is perhaps the more commonly adopted view, thai while thon
pariB of them which lie between the sternal cartilages act like the external
inlerc<talala as etevatora, i.e., aa inspiratory in fuuciion, those parL^ which
lie betnecu the oasenus riba act as depretuors, >'. e., as expiratorv in function.
In llie well-known model conststin); of two rifpd bars [Fif;. 1^1] repre-
H-iiliu)!; the ribH, moving vertically by nieanu of their articulations with an
upri)|>lit TL' p risen tiuj; the tipine. and connected at their free endd by a piece
tvprcM'ntinjf the Meniuni, it ii undoubtedly true thai sirtlched ela^^lc bauda
Bltai'Jitfl to ihe ban in such a way as to represent resjieclively tlieesternal and
internal iut^-rciMlaK viir., itlopin^ iu the one cii^ dowuwani and tiirwan), an<l
in the other downwanl and Imckwatil. do, on being left free lo coutract, in
ihe former oa>e (-levate and in ttie latter ileprcM the rib«. t^eh n model,
bowevor, dotv not finirly rvprwent the nattinU onu<iitioiiit of the rilio, which
■re not ■Iniight and rigid, but [leeuliarly curved and of varying elasticity,
enjiahle moreover of rotation <>n their nnn nxes, and luiving thi-ir mowmenta
dt'lcrmiii<-<I bv the cluiriicleri of their vertebral artiriilation*. The nivchiui-
icul ei>ndilions in (art "f these niuwlw are »o complex, that a dctluclton of
tlteir actions frcni simple mcehanicnl principle*, or from the direction of the
fibr4«, must be exccolingly <lilliciilt iiud dangerous. Actual cxpcrimcjits on
tl>e eat and dog teixl Ut show that in ibcse niiimalx tlie contntctioo of the
internal iuterouauUs, tlong their whole length, take* ptaoe, in pwut of tinw,
TUK HKCIIAMce OV 1'Vl.UOSAKY KtSl'I RATION. 447
vIltnMvlTiftilithKtortbedinpliraeRi. ami thuRafTord nn irgiiaMiiit in fiivor
^ ihrw niuEolcfi lieiii); expimlury in fiiriclinn,
Sm ill inipurlancc lu the cxtvniiil intrn-ofltalH omin ihr liivnlon*
'■Qauram, wbicli, th»ii);li »mall iiiiiH.'le8, arc able, Iroin thv tiviirnraa ul' iWir
^■aacal bu*rtti>tut i<i the fulcrtiin, lo produc« aiii«<]pnit>lt> tiiovcnx-nt of ihp
tsraial coda uf the ribd. Tbe exlernitl iiiier<.'iw tali and llie levatorc* tiistanini
*> t h Uw acaluii iun,v fatrlv be ta'id lo bu i he elt-valora of the ribs, i. c, tbft
■"-hiwrf BaKlea of covul iuipiraiioii in Durmiil brc-«iliiug.
Iff f
I
I »>rt«i.»'> >W lan.nnli^UUiiK ihi tlw iKfllo*! terr; i |i ui4 vi art two rutiUir baiiilt,
tviamttit mpmittvlr <>>r viumoJ mni UiMful iMdwaul MuMte*. Hacnni a r«pn>c»ia
HflM la > Msi* of rm ir. »'iw, (htisixl «aa4r*iu ftcc la Kuitnel. tl it •ibriuiK ilui In
■oMMIn III itirvvai Iriitih. I1 uoiild ■hiiid** i<ariilnn MladlNcno*-' If. ■>"•• IIh I>i»I w
■ «■* n<« ii> WOTP. Ilir (MiiDiw niniM laVv pUr* iif iTlHit «■ lecn *l>m r ud ■> bniudv
I Tiniv. In ilUcmn Ik xtve ftuo^lAfi afv ^l«]|*i^M>ii-
TWMnaif awl ■ RT|MMmi ib* mIbiwI ikiomoiai iiiiui<l«. iiiiidi. tir •»»ini^ilaa.*I':TMii Ike
fn*. T)>*liui4> a kifl I fii«BcM tlw iiiuniBl lii<iin.-ii«al*, wblrli. Irj TniilmvUnit. iktirrw ihi> rlln,]
It niuxt be ai]<led, bow«v«r, that swue oWnera deny that eitber act of
iii|rr«nataJ hiiikcIo) take any iiuporliint |Nirt ia raUin^ ibo ribe^ Tboy hold
■hat tlM> cbiff if uiii lb« only une uf th«M.' iiiiik-I<9 i* by tlM>ir contnu:u<in to
rrwlrr llic intetxislat «|>ncea 6na ami tbv urbulv lbi>rii<:if oiigc rigid, so tbat
Ibo Iboras i* luuved lu n irbi>le< by ibe other mtiicb-e mvntiontd, and tbe
iattrcaMul (pact* do m>l give way during tb<! renfiinilnry RiiivenicolB.
Addhioiial npuff in Ibe Immvent* diameter i> alfonlvd probably by ibe
ntalifMl of ibe rib* oii an uiili-n> -piiatcrior nxiti; but ibis awv«m«nt is quite
Mibeidlary and iitiiiiipiirtaMl. Wtion iIh- ebest if nl nwt, the ritia are eonie-
■ bat ioclinnl witb itirir lowvr bonier* •lirvetti] iiiirnrd as irell sa downward.
Wbru ibey an- diiinn up br ihc nclion of the iiilercoalal inuH.'Iefl, their
lu«i>t )i«nlcn aro evrrlctl. Tliu* their lliil f<i<li-i> iirc prctcnled tu the iHoracie
tavilr, nbieh t* lbcr<>by »ligbily iiRn-iiMril in uiiltb.
i $39. lioiaml iutpimti'iu. Wb«u rwpiiatioD beOMOcs labored, olher
■Duarlt-* ar« bruuglit into pbiy. The evalciii are Btronf;ly oonlmi.ted, m fta
diatiDL-ily to mtae or at leiiM give a very fixeil HipiwK to ifie first and aecmd
rilw. In tbe «anie way the aerraiuD {kwIicos sopenor, which deaceud* IVam
Ibr fiKiil npiiie ill tbe lower cervical aud upper dofanl regiona to the •eroDd,
ihini, fiiiinb.aml llftb riba. by ite oontractioin ruiaea (hoae ribo. In iaborc*!
breailiiog a funt-lion of tbe lower lal«e rilis. not very iMXicenblp in riuy
bn-aibii>([, v»OHa into (>lay. Tbey are deprew^i), retraclol. and fixed, thereby
giviu); in(-r*«Md MipfH>rt l» the diaphra^iu, and direclitig the irbolc vnat^M
of ibat UKiMjle lo the verlieul enlarpioieQt of ibe cbc!)!. Iti ibijt way the
torralu* praticti* inlrnor. wbich pnww upward from ibe iunilioir aponouroaia
bi tb«- laA four ril«, by dt-preaaliig wd lixiii); tboM rib* l>ecoiii«« an adjuvant
J
iuipiratorv miiecle. The r|Uiiflratus liinilioriint nnd [OW^^iamont of til*
MDro- 111 tubal i ft iniiy have n Biniilnr liiiietion.
All these [tiutcles mar come iuto ovtioD oven in brculhiiic which, tliOUgh
deeper Ihaii ueual, can hardly piirhape be call^ labored. WheD, however,
thv nei'd for greater inspiratory cITurle bccome» urgent, all the muscles vrhiob
c«ii, froru any fixed point, act in eidargiii^ the cheei. come into ]>lay. Thus
the unua and shoulder being fixed, the terratuB ma^nu- paaung fr^im the
acapnia to the middle of the lirtl eight or niae riba. the pectontlia rniDor
pnanns from the coracoid tii the front nans of the third, fourth, and fiftb
rib», the pecioralis major uans^ng from trie hunieru* to the costal cartilsj^w,
from the MCRund to the sixth, and that j)urtion of tlie latiatiroiia dom whidi
pOMn from the humcruii tii the lii»t thrc« ribrf, all iiervc to elevalv llic riba
and thii:* to tulargc llie choit. The ^terno luBNtoid and other muiiclcs pnMing
from the nt^k ui the xtcrniini, nr<! alno railed into acliou. In fact, every
muscle which by iw ciintrnrlion can filhcr cli^vnlv the ril» or contrihuiv tu
the Hxetl siipnorl of muscle-* which <l» elevate the rilw. nuch an the Impczins,
levator iingiili »ctipiilic. and rbomboidci by lixinc the ecnpiila. may. in the
iuapiniloi'V cfliirl» which accompany dyepmca, be oroiight into ptav-
tl 33S. Kipimliiii. In normiil eti»y breathing, expiration it in llie main a
simide effect of ctnslic reaction. By the inspiratory eHori the elastic tiaaue
of tlie hiuj^s is put on the stretch; so long as llie inspiratory muscles coalinae
ojotractinj*. the tissue r«muin8 stretched ; but directly ibcae muscles relax,
the olsBliciiy of the lungs coiiim into play and drives out a portion of the
air coiitaini.-d in them. Similarly the ebiMic sti-rnum aud ciiatal curlilaga
Mv by the elevatiuu of the ribs put ou the mreich ; tbey are drivcji into a
position which in unnatural to them. When the iuiercxfiul aud other pI»-
THtor luuscien cease to cootriu-l. the elasticitv of ibe sternum aud eostal car-
tilagcs causes them to returu to their previous [)«»iti»n. thus dcpreasing tbe
ribs, and diminishing the dimeiiiions of the ohest. Whcdi the diafihraem
de»cciidft, in imxhing don n the iibdoininni vitwora, it pule the abdomioal iralb
on the strrtch ; n]id hence, when at the end of inspinttion the diaphnifrm
relaxes, the abdoniinul wnlls return to their plate, and by pressiug od the
abdominul viMvrii, puth ihe dinphrngm u]> again into its position uf rat.
Kxpiratiun then during ciisy brcnthing is, in the umin, simple elastic reac-
tiau 1 but there is probubly some, though pcKsibly iu mi«l cases, u very alight,
expenditure of muscular energy to bring the chest more rapidly to its former
condition. This is, as we have scen.supjHised by manv to l>e nSurded liy the
iulerniil intcrcostnls acting as deprewors of the ribtt. If these do not lua in
tfab way, we may suppow that the elastic return of ibi- abdominal walls ii
accoiupauied and assisted by a ci>Dtmc(iuti of the abdnmirinl muA>clt9. The
tiiantfularis slemi. the etlect of whose cimtrnclion is to pull down the n<Mnl
«artiGigc», may alM be r^^arded as au expirutury mn-wlf.
Wheu expiration becomes labored, the abdominal muncW Ix-coine im{>ort-
ant expiratort- agents. Br prcfsing on tho conti-nt> of the abdomen, they
thrust lliem and therefore the ilia|>bragm iiImj up townr<) the chctt, the vertical
diameter of which is therebv lenteiied, while l>y pulling down the sternum
and the mi^ldle and loner rilis they lemcn also the cavity of the chest in its
anicro [HUteriiir and trantvcrw diameter*. They are, in fact, the chief PXpira-
tv^ muscles, though ihey are dmibllem aMisted by the scrratus posticus
inferior and |iortions of the tuicro-iumbalis, since when the diaphragm is not
oontrscling, tiM deprimitm of the lower ribs which the eontrsclinu of these
miucles causcD, serves only to narrow the chest. As expimiion tiecon
more and more fiirced, every musclar in the body which can either br et
tractiug depress the ribs or press <M the abdominal vi^i-era, or utfonf fixed
support to muscles having thwe actiaits, is calle-l iuto play.
CITAK0B3 or THE AtB I?: RESriRATIO:?.
449
S 387. /new/ and hryngeat rwpiration. Tin; (honieio respiratory mtvt-
taenia .ire accorapuiwd bv s»ociat«d respiruon- itKivemenu ul' otber |MirU
of the body, mora mrticularl;^' of lh« tkoe and of ilw );lutiiit.
In Qomial heallhy respiration tlie current of air whioli pnium in and out
uf ihe lungs, irareli^, not Ibniugh the moutb but through thn nose, c)ii«f)y
lhr(Wf(h tbc lu«-er naxil mealuii. Th« iiivoinj; air, by exjxwiira to tliv va>-
cular RincouM membrane of tbe narmn ntiiF winding; naxiil pmn^M, i* mora
effldeuily wnrmiHl ihnii tt would be if it pMHtd through thr mcnith ; and at
the sattH' tiiiK- tb<- inoiitli i* tliurcby protected from the dmiccatlng cilf<.-ct nf
tbe continual inmnd of omipanttively dry nir.
DuriDE encli iiupiralory effort (be noelrilx nr« expanded, probably by the
artlmi of tbedilalores naris, and thuD the eniranoe of air fiicilitnted. Tbe
return to Ibeir previoua condition during espiratioo is etl^ct«d by the elas-
ticity of the niiMil cartila|;e8. assisted perhaps by tbe coniprewone narie.
Tbie movenietit of the noetrils, perceptible in many people even during tr»tt-
quil breathiDg, beromea very obvious in labored respiration.
When ihe mouth L> cloaed, tbe mil palate which 'a held somewhat tense,
ia swayed br the respiratory current, bitt entirely in a passive manner, and
it is not uuti) llie larynx is reached by ibe inw^n^ air that any active move*
menta are met with. When the laryiix (the details of wbidi we sliall have
to deal with at a later jnrt of thi« work) Is examined with the lurynvosoope,
it is frei)uently Men that, while during inipinilion the glottis i« widely open,
with each exnirataoD tbe arytenoid curtilaKcH apiircMcli each other ho m to
ttarniw iIk* gli>ttis, th« cartilflMS of 8antorioi projecting iiiwnrd at the aaine
liine^ Tbuit, ityiichronouR with t)u- nvpiralory oxpanawn and ouilniutioQ of
tbe cben, aiid the mpiratory t^lrvaiion and aeprcwmt of llie alw niui, tbtrt
it n riiyiltmic wMleniug iinil unrmwing of Ibe glottic. TJkv tbe movement*
of tbe novtril, this rmpiraion' action of the glotti* i* much more evident in
labored than in trvniinil brr-jithing. Indiwd, in the latter com- it i« frequently
abMDt. The manner in which this rhythmic opening and narrowing is
«fleet«il will be <loaorib«d when we oome to study the production of the voiO&
Whether there exiala • rhythmic contraction aixl expansion of the trachea
and bronchial posMgea, eapecially the smaller and more exclusively niuDcular
ooee, eAected by m«aDfl of the ]>lain muscular tissue of those orj^ns und
ayuchroitous with the respiratory moTemenla of the chest, is uncertain.
Chamoeb op the Air ik HrapiRATioK.
^ 398. Dunns: its stay in tbe IimgM. or rather durini{ its slay in the bron-
chial nanage^. tlte tidal air (by me-ans of diffusion chiefly) elTects ex^angea
with toe stationary air : in ooiueipienoe the expired air dilfera from inspired
air in iieveral important paritculHn>.
The temperature of expired air in variahlo, but under ordinary cin-um*
atanow Ik higher than that of thn intpirtrd air. At an avera;^ tempi:r.)ttire
of the alni'Mphete. for iniilanee at about 20° C, the tetD|){traturi< of expin-d air
ia, in iIh' mnuih 'X\.9'. ia tbe nim; !U>.S°. When the external li-mpi-rAtiire i»
low, thai of the expired air xink* tometvhat. but mH lo auy );rent (-■xlenl.
thus at — K,.'!" r. it i* ■iit.H" C. Whcii the external tempnrnl'ure in higli. the
expired air may Iwcomo cooler than tbe ini*pirv«l. lhu» at II.IP it hai been
roun<l l<i t>e M.]", The expired air taka< ilji tem;>orJi)iin.i from that of tlia
bixly, that is, of tbe blo7<l, and thi« as we sbnll sec later on while generally
hiffher rosy, at timm. be lower than thai of tho atni >t|>hi>rc. TTic exact
temiierature of the expired air la fact depends on the relative temp^-ratumt
of tlie blood and in4pife<l air, niid on tl>e depth and rate of breathing. Thv
480
RKHPIRATIUN.
(i«]re*n.
IlltrnvTII.
ntrUiiilc arlil
80.91
m\.s
U.(M
l(i.O»»
7BIM7
4.86
cliiinK« in (emper8tur« lakta place Dot iu ibc luiit;a but fiMhenr
and (hiellv iu the nuae «ni] pnarynx.
^ 338. 'rbe expired air isluuded with a<|itctiu« vapor. Tlic point of lain-
ration of aiiy gaa, that in, Itio utmost <|iiHDiii}' of u'u(«r vhirh nny givra
volume of (tan am inkr uji a* Htjui-oiin va]i-ir, vnntw with il> ti;ni[)cniturT,
beiiiK higher witli (hi- higlior ttrmjicrulinv. Fur ila ohii (rnipvnitiirc oxpiml
air it, iicv>rdiiig tu tm>M obvcrvon, siiLuralfd with iiijiirous vu|K>r. The
inoii«tiiri-, liki' the wArmlh, i» impuited n<>l in the depths of th« iung but iu
th(> iip|K:r piuHogiv. The inspircn nir ae it pluses into Ihv bronchia is already
wilurnlri.l with moi^turu.
§ 340. The rxpimi nir conimn* uboiit 4 ur 5 |>er cent. I«n oxygen, sud
nbout 4 per cent, more cnrbontc iicid ihnn the inspired sir, the quautilv of
nitrogen siifivring but little chungr. Thus
Innnlrril Rir contains .
Espirvd ■• ■■
The <|iiantily of nitrogen in the expired air is aonietiioes found to bt
sliebtly greater than, as in tho above table, but aoroetiines e<|ual to, and
and MDieiimt^ less than, l)mt of the inspired air.
In a aiogle bivath llie air is richer in carbunie acid (and poorer in oxygen)
at the end than at the beginniug of the breath. Hence, tli« loneer the
breath ia betd, the greater the (artificial} paui^e bi-iween ianptration and
expiration, the higher the percentage of carbonic acid in the expiretl air.
Thus, by increatiing the interval between two oxpiraiionH to 100 Mojuds, the
percentage may lie raised to 7.5. When the rale of breathing reoiaiiu the
•ante, by incrvasing the depth of the breathing the ]>ercciitage of carbonio
acid in cneli brentli la lowered, but the total (|tiaiitity orcarbooic acid
expired in a given time is increased. Similarly, when the depth of breath
r«n>ain>> the Enmo. by iiiickcning the Tuto the percvntage of carbonic acid
in vnch breath is lowered, but the (luantity expired in a given time is
in created.
Taking, ns we have done, the amount of tidal air pawing in and out ot
the chest of un aveni^'e' man at SOO c.c, lucb a pereuu will eX)>ire ahdut
2*2 c.c. of carbonic acid at each breath ; this, reckoning the rate of breath-
ing at 17 a minute, would give over oOU litres of carbonic aeid for tlie dav'a
production. Actual determiiiatiouf. however, give a mtlier smaller total
tban this ; ihua, in a eeriee of experiments of which we shall bave to «{M>nk
hereafler. the total daily excretion of carbonic ncid in an average man «u
found to be SOO grms,, >'. r.. rather more ihnn 4U0 Htree (40(}), containing
218.1 grmft. carbon and .581.^ grme. oxygen, the oxygen which actually dis-
appeared from the inspired air at the aame twing about "00 gniu. Thia
amount, it *huuld tie said, represents, owing to the manner in which ilit
experiment wm coDdiicted, the gaaa given out and tukeu in, not by the
lungs onlv, hut by tlie whole body ; but the amount of carlionic acid given
out ny otWr channels than thf lungs is, as we shall vce, very slight UO grait.
or even les*), so that 800 grmn. may be taken iia the average jimcliiction of
carbonic acid by an avertige ninn. The <)uuutitr, however, both of oxygen
consumed and of carbonic actil given out, is subject to very wide variatiooa;
thus, in tho obtiervations of which we aio sjKtiking. the daily ijiianlily of
carbonic acid varied from 686 to 1265 gmii., ami llial of the oxygen from
894 to 1072 grms. ThtM variBtions nnd their cniitcs will be discufeed when
wo eome to den! with the problems of utitriiicn.
g 341. When the tolnl <iuanlity of tidal uir given out m any expiration is
comjiarcf) with that taken in at the onrrvsponding inHpiraiion, it is found
timt, Ixrtb being dried nnci nienMinid at thv Mine trmjKiriiture mxl preaaure,
the ux|nftt<I Bir m l«ai in vulumc tliiui thi; ii)«pir«i) nir.iho dilTpmicc nmoiint-
in^ to nhinit ^tJi or ^th of tbc volume of the iiilter. flencc, when un
nnimiil u miid^^ to hrcalhc in it ounlirmt s|ifl(-c. the iiir n ub«uliitety diminiahed
in voluiDi'. ThvBppruxiiDKtc cquiviili-nci- in vol u me hoiw«CD inspired and
expinvl iiir iirUc fmni |]i« fact llwt tlie vohimo of any given quantity of
cnrlwnic neid i» c<jijn] to the volume of the oxygen eonsuined to produce it ;
tbo tlight tnlliug short of the expired Air \* due to the drcuiuslancc that all
tbsoxygeii inspired does iiot reappear in the carbonic acid expired, some
liBviiie formed irilhiii the body other combinations.
4 342. Besides carbonic acid, expired air contains various mibetanccs
which may be epolien of as im])uriti<s, many of aii unknown Dature, and all
in small amounts. Tnca of ammonia have been delected iu expired air,
oven in thai latten directly from the trachea, iu which cose its prvecuce oould
not be due to deeomposiii); food lin^^ering in the mouth. Wheii the expired
air is condeosed by beitif; conveyed into a eooled receiver, the lujueoti*
product is found to cuniatu urj^iic matter, vrbidi, from the pmcnce of
micr&urguiisins introduced in tlie inspire*! air, is very apt rapidlv to putrciy.
The organic suliHance* Ibua shown to he oreueDt i» ihv expired air are the
cause iu part of the odor of brc«tb. It is probable that some of them are
of a poisoiioiis iiatiire. either ]>oiiioaouH in tlieniaelves as comiug direct from
and produced in some way orothcriu the pulmoaary apparatus, or poisonous '
Bs beiug the products of {Kitrefactive decomposition; for various animal
•ubatWKM ana fluids ^vo ris« by dvcampoaition to distinct poisonous pro-
ducts known as ptomautfJi, and it is possible that some of the constituenu
of expire! air arc of an iillied nature. In anj case the mbatanoes preeent
have a dclclenouti action, for an uliiKWiilicrt- containing simply 1 per oont. of
carbonic acid (<rith a corrc»pon<lin^' iliniinulion of oxygeo) has very little
rtRvt on tbo animal M;oi)oniy, whereas an atmoephere in which the carbonic
iiciil has been raised lo I per cent, by breathing is higbly injurious. In fact,
air rendered so far impure by brealaiiig that the carbonic acid amounts to
0.08 per cent, is distinctly unwholesome, not bo much on account of the car-
bonic acid as of the accompanying Impuriliee. Since these impuriti(« are
of unknown nature and cannot be estimated, the easily detenulued carbooio
acid is usually taken as an indirect measure of tbeJr pramnoe. We have
seen that the average man loads at each breath 50<) c.c. of air with carbotiio
acid to the extent of 4 |)er cent. Ue will accordingly nl each breath load
If litres to the extent of I per cent. ; and in one hour, if he breatlie IT timca
a minute, will load rather more ibun 'JOOO litres to the »aiue extent. At (he
venr teas*, then, a man ought to he Hupplied with this quantity of air hourly,
and if the air i» U> be k«-pt fairly wholesuin*.-, lliul ia witli the carbonic acid
n-doeed below O.t ytr cent., Iiv slioutd have even mure tliau ten times as
much.
Tub RKeriRATOKY Chanoeh in the Bi.onD.
$ 343. While the air in iMwung in and out of the \an^ iit tlius rubbvd of
a portion of its oxygen ttml loatk-d with a certain quantity of carlionic acid,
iho blood as it streams along the pulmonary capillnrit^i iinilirrgiiiw ini|K>rtaiit
correlntive chan(;e». As it leaves tbc right ventricle it ii> venoiix blucid of a
dark purple or maroon color ; when it falls into lli« led auricle it is arterial
blood of a bright scarlet hue. In passing through tbc cnpillarin of the bodj
from the left to the right side of the heart it is again '-hanged from the
arterial to the venous cunditioo. We havo to inquire. What are the esun<
tial diflisrences between arterial and venous blood, by what means is the
452
RESPIRATION".
veoouB blooJ chaneetl into arterial id the Iuiiks, and the arterial into venous
in the^rest of the nody, and what relations do these changes in the blood
bear to the changes in the air which we have already studied?
Vi'i, laj.
rilAliKAIIHATIl' 1 1.1,1*1 mm IN OV T.l UlVI'i - MEIHTBUr, OA--ITJII',
A »nd h on.- Iwu gluw plobes ciimei'dil hy sirimg liijia.mlilwrtiil"*: i unci ). wild tnnfirailar
glon ^lotiu*<, A'unil It'. A K Air1hi.T coiini<<'Ieit \ij m^Liis of ihi- i^iiiiHri'rk r wlrh t^c rect'Lver i'
POOtainintf the tfliHHl tr>r oilier HiilcJ) lo be nnaty;ft,<(l, unil b \iy menh^ i>f Iho siiijh-ivi'K vi and Ibe (utv
mltb Ibe i¥<'virer Ii lV>r rvrdrliiR tbt-KHW!. A hiiiI H hivhIni (iinrn'OIiil uilb tmb i>ihvrliy means
of llip BlofHVH'lts/BdJ i;. ttiL- ifliiCT bt'liii! «n umiiiin.-U lUiii 11 Bl-a) niiiimiiiilr(ii>-s «iib lllj; llii.' imssagB
g", A'Hii'l B'beiiiB ("H "f mereiirj- auj lUt c.iok- I: f, ii, and rl Wiii: fiii'ii. Iiiil r iimi u' tliiseil. on
ralAlnif A' by iui^hiis uf tho jriilley ;j tlio iDprrury irr A' itll' At drlrhiL: iiliI \he iiiim-oiiEkiiuiJ iiHi Uitu
fl, and ttiiont lbroui;h e. Whun Iho morcury hii,-' rli^-n iibovu <|, /i* flusiiT iniil v' )it'iii^'Hp|ieni?d, H' Is
In (uni rBlseil nnlil li 1^ i-unii'kii'ly Hlk"! wllb nitrcnry. all lln'nir imvinnily in ii UiiiL.- ilrivi-n uui
tSroiliiHe. I'lion i-liwtnK ■? ami IubxtIhb B' Uio uliiilu of llii; infrrTirj- In It fill I- in if. iitid n v«i iiiiiii
coiuWiuenlLj- 1' pttablisliLiL In Ii. I'n rlii>.inKf;', bul iil-etiMin r/. f. "Il'l *, "nd lnucrin:; .V. ii lai-iium
iitfilmlliirly o^Inbllnli^l In A iind In tin: Junvllnn IrumH-n A and II. Lf tin- ri>cL r ]h- ni-\\ iifiiHiud,
IhCBSsen of llic IiIoinI in C i~caii' '"I" I hi' vwtimn r.r A an<I ll. Ity rai-inv- A' ;ifn.^r I In- Lln-Lin. uf "■.
and ii]ifiiiQg of ll. the i;u-*» "i xl frvc aiv i|rlyi>n fmm A InIO H, anil liy [hi' r,ii-ini! <'( If Inim 11,
through f Inii' rhi' ruoeiVLT [>, -^EHndlin; 'iv>t nion'nry.
The fncla that venous blood nt nnoe liecomes nrtoriid in aii|".'iirniii-L' <'n
being exposed tourahuken iij) with nir or oxygen, and llmt arliiinl lil I
THE BS9MHAT0RT COAN'OES IN THE BLOOII. 4o3
"mnca vtnoiu in a)ifmimice trhen kept fur twine little time in a olow<l
^Mil. V ohm •uttiniltvil tu u mrreat f>{ »uqi<> iiiditlcrciit gas such ns iiilru-
^or bTdrogVD, prvpitre ua for the Blatentent tlial th« futi<lainenlal diflrr-
*vec tictwtrn venous ami arterial blood l§ in the relative proportion of tlvc
"VK^n aod Mrlxmic aciil ^i^eieH cuntaiDed in each. From both n oeriniD
'lUMititf »f gucuii be e.\tni(.'lt^i bjr means which do not othervriw materiallir
■jtcr tlw oocutiliitiou <>r thu blood ; am) thin khs ulieu obtained from arterial
UdmI ia Ibtind to (MnlatD more oxvgen and K-m farbunic acid than that
■MUtititl fnini veuouA blnoi). Tliiii i» th« real difTcreiitial character in the
**> ViIoihIs ; oil olticr difTi-rences arc tilher. iis wi- shall .-we to be the caiw
*w tlie ooW. dcprmb'nt i>ii Hm or ure iiTiirup'>riant and fluctiinliuf;.
J^ tbe quantity <if gn* which can Iw extnu'tcd by the mercurial air-pump
100 fok. or blood be moaturc-d at 0' C. iin<l n i>raeure of 760 mm., it
lad to ainouDi Id round numbore to i30 vol».
, ^/^bv tacuuai produced by tlie ordinary luechanicul air-pump la inaafllcient to
^^Uaet all iba ga* from blooil. Hence it becomes necesrary tn uiw n mercury
^^ttp capable or prodndnj: a Urge Torricellian vacuam. In the form of mercu-
rial paain which iwan Ludwlff'* nnroc(Fiji. VAi! twu large globee of Kl>a*,oae
~^ua BBU the other movable, ere i;uuDFCt«il tiy a 6exible lube ; tbe SlM glob* b
*tole to oxamunicate by oieaaa of air-tight •ioiicocIch altornatciv with a roceivcr
^notaliiing Iha blood, and 'nilfa a rccetTer to collect the gu. When thr movable
filled >ilh mefcuiy ia roiawl abnva ibe fixed one. the mercury from tbe
t nuia Into Btid carapletdy fltU the Intter. tliv nir prevoualj preaent being
OBI. After adJuMlsf the cocki. the uiun-Kkle globe b then depreaaed
thirty incbea baloir tbe 6xM one. In which the MtiKe<iucttt fall of the mrrcary
raducea an almort complvte vacuum. Uy tiirninK the proper cook this rocuom
pot laid 004)aection with I he ret' river cpntainiiiK the bltHxI. which thereupon
beeoiea propcrtiuiiaiely eiliauxied. By again HdjiialliiK the oclt* ant] onoe
Mra eJavatiag llie noveableelulw, the giia Ibun extroclea ia driven out of the
^fti (lobe into a reeeiver. Tiie t-actium it then unce mure eatabliahed and the
•fanUrao noeated aa loog aa gaa coutinuet to be given olT from the blond.
A laodfliea fbrBi of pump working on the nainu principli* aa that of Ludwlg,
but Invnlvins the tuc of oaly one globe to he mado vacuuuji and one morable
naervnlr for mercury, hiw teen oouatraclcd bv I'lliiiref. It preaeota aeveral
adeaotagaa over tbe one {uhi deeoribed, the chiei being thai (I) non-deflbri anted
Uaod auy be aacl for the eitracUoa of gaaee, (S) tlu) vacuum into which the
paea an evolved i« large, (3) ihia racnam ia hem drv by being connected lalci-
allj irith a vacuooa dianbftr ooniainiaK mlpbunc acid. Tfaedeiaila of ila eon-
Mractlon are. bowever. oompllcHtcd. and the greateet care ia required in ita uae
ti> avoid breakage. Of Inter yrars a almplllled form of pump Iim been tntro-
diM*d for laboratory work. It na* flrat aaMl by (tri^haot and i'aul Bert, and la
now frMHfotly callfol an Alverpiiat'a pump, froni the name uf ita preaent maker.
Ptff. ISa give adiaKraramatic repreaentatiun of Itn con»tructioii.
A M a gloM bulb auioe five Incbm in diameter, blown oa toaglaia luben below
and on lo a vertical tube b above. The lower end of i it oonaected by ■ thlck-
wallcd India-TubbcT lube witli a icucrroir fir meacnry B, whiuh can be raiaad
and lowi-rrd hy mean* of a ■irinj; ihuwok over a DUlley r. The vertical tubo'^
Ib Ihickenrd ■! one place, and into tbia thickened iiortion a Ihrco-way tap 'I b
lirouod. The upper end of A b prolonged lahuve ihe ihice-way lap) inlu a line
paint. Th-* (KMot pawH by a tight joint throaith the botloin of a vcMel r, which
ran b* |>aTl!r HUrd with inrnuty, and over vhuh a receiver/. Illled with ini-r*
nirj for the L-ullcclioii uf tlie i:aw>. can be inverted. A tube y fuaed on laterally
to due tiMolng of the tbre«-way tap 'J plaeei the lailer in connection with a
iblck-aralled Woulff'a bottle t'cuiiiaintng a layer nf mrong *ulphuric add. Tbo
«H;oniI lulMilure of tlib bottb t* •imlturly connected bv an claatic lube with the
ir«el i>. iiilo which blood or othrr llui<l tuny be iutiuluced by inean^ of tbe lap
A. All the movabtu joints nf ibi: a|i)ianiius arc jiiotected by India-mbber tubM
loin wb'ch water can b« poure<l. and a nieial caalng round the lap if, which
nmj alio be illled with water, limilarly prevoota the powiibility of any leakago
bM*.
464
BB8PIRATI0N.
The pnmp is lued u foUom. By placinK the Up d in the portion ahowD ia
the figure and rutiog B, the balb A may be filled with mercurj up to the top,
the coQtaiaed air bein^ expelled through the upper end of b. Bf a elight tnni
of the t^ all connect'on between A and either the tube g or the npper MTt of b
may be cat off*, and on lowering B a Tacnum U eatabliahed in the bnlb A and
Eart of the tube a. A may now be connected by the tap d with the tube o, and
ence with (7 and D, and, h being; closed, a partial Tacnum ii eatabliahed in C
and B. By meani of the tap d the air In A may be cnt off from g, and on raia-
lug B and placing the plnr of if aa shown in the figure this air may be expelled
throngh the npper end of^. By elightly turaing d and lowering B a Tacuam is
Main eatabliahed in A, and aa before a further portion n{ aXi in (7 and D nuty be
wlowed to pas4 over into A and the racaum in D and C increased. In this way
Fio. lis.
Q\
^^::^
DUOKAII or ALVERaM.tT'B I'i'sir,
all the air in D can be extracted, the final stngea being facilitated by the admis-
doQ of a little water into D, the last truces of air being ilriven over into A by
the rush of vapor from the water. A known volume of blond having been col-
leoted over mercury in a small tube is now allowed to enter D through the tap A
ud yields up its gases to the vacuum. A repetition of the processes by which
thA wr in J) was originally extracted will now remove the goi^es which have been
Sven off from the Known volume of blood, the only difl'erence being that now
e tube/filled with mercury is inverted in the trough e over the iip]>er end of
the tube b. In this way the gasee originally in D nrc not allowed to escape into
the air, as was the case when the apparatus was being originally made vacuoDH.
but are collected in / for anhseqaent analysis. During the extraction of the
gaaea from the blood the bulb l> is immersed in avesaelorirarm ivater. to facilitate
TOK HESPIRATORy CHAXOES IN THE BLOOD.
4S5
iIm) pxit of th« giwc* >ti<I. t>v raufin^ tlio r»nn»tion of Urj^ quAnlitin of aqueous
vnt<or, U>MrMpiltc|C>uc*r*pi(lljr urcr ioto A. Tb««ulpliuriciK:i<l cbsmbcr CdrJM
tli« vnouum befote ibe admlMion of the blixij into O. aad hence makea it more
|i«tf<>ct and tnuBM Uie mott oo(Di)l«t« and rapid evolution of gun ttnta the blood .
The nverage rompoAition of tlie g&§ thus obuiitied from «nch of the two
kinds of blood (the arloHsl blood being taken from a Iiir;>e arlery, and tbe
venous blood from the right side of the hearti U, slated lu round numhera,
aa foUowB :
From too vola. may b« obtained—
or oxyvtn. U( (vrbMilo aald. Ot nlln««a.
Of arterial blood, SOrols. «> toIs. I (o S vols,
or Tcuinu blood. $ti>12vok -k! vols. 1 to S vols.
•II ateuured at '<X) mm. aud l>' <:.
That is to Bay, venous blood, as compared with arterial hlood, eootaios 8
to 12 per cent, lew oxygen and l> ]>er «eiil. more oarbonic avid. It mnat be
i-eiiiem)>ere4), however, tliat while arterial hlood from whatever arlery takeii
h>N alwayti nearly the itame proportion of gasMw, or at all cvunl« the xame
•mount of oxygen, the amouut of uxvEen in venouN bboci, cvui when iiikea
from tbc »me vein, may Thty a good deal, utill mort< w> when it is taken
from diHercnt veins. The rea«on of ihi* we Mhall fix hereafter.
It will be I'onvnulenl to cj]D«l(lcr the relntiona of endi of these gases sepa-
rately,
TAe Rdaliont of Oiffytn in Uw Blaoit.
$ 344 When a liquid such as water is exposed to an atmosphere contain-
ing a gaa such as oxygen, some of the oxygen will ba disaolred iti tlie water,
thai b to aay, will be absorbed from the atmosphere. The ouantity wliich
IB to abeorbed will depend on the premure of the oxjgen in tlie atmmtphero
' above ; tbe greater the pressure of the oxygen, tbe larser the amount which
will be abeorbed. If the pressure of the whole atmosphen' remnin thi- sante,
at 700 mm. of mercury for instance (tbe ordinarv atm<isph«rii: t>n:N>ure), the
priTtiurt! iif tbe oxygen may bi- incn-ased or iliminished by increasing or
diminishing the proportinu of uxygni in thu iitmimphvre. tio that with an
Bimonphere renutinmg at any givi-n pmwurQ the 4uanlity of oxygen
•bwirbed will depend on the quantity jirexwt in that umosphore. If, ou the
J^thcr hand, wat4:r, already conliiiiiing a good di-nl of oxygen dimolved In
it, be exposed Ui an Rtmosphen- coDtainine little or no oxyge^, the oxygen
will escape fK>m the tralrr into th<- ntmiHplicre. The oxygen, in fact, wbioh
is dinolved in the water, like the oxygon in the Htmospnere above, stands
iti a crrlaiD prmsure. the amount of pressure depending on the quantity
dissolved ; and when water containing oxygen dissolved in it is etpoeed to
any ntniMphere. the result, thai i^, nbolfier the oxygen escapee from the
water into the atmosphere, or pusses from the iitniosphere into the water,
depends on whether tbe pressure of the oxygen in the water is greater or
Jeasthan ibe pressure of the oxygea in the atnoephere. Ilenee. when water
v% oxposed to oxygen, the oxyc«4i uthor escapes or is absorbed uutii equili-
I briiim ia esubliMied between tne pressure of the oxygeu in tbe almcsphera
f-above and tbe preastire of the oxygen in the water below. This result i^ as
(hr as mere absorption and escape are concerned, quite in<lependaut of wliai
oilier gases are present in the water or in the atmiisj)bere. Suppose a half-
litre of water was lyinj; at the hoUom of a two-litre tlask, and that tbe
atjncapbere in the flask above the water was otte-Lbird oxygen ; it would
make no diS'ereiice, as far as the aluorptitm of oxygen by the wat«r was
cnQeemed, whether tlte remaining two-thirds of the almoupbere vai car-
boni*' bpkI, or nilroMB, or hvdrogtrn, nr wluttlivr tb^MM nftoTC
mi It vncuiim tilled lo ODe-lliml Kith pure oxyeon. Hcdco. ii ii> said tliat
the nbicrplioii of any gns (Io])«iidi' on Ihi- jifiriiaf pfiuurt of thnt gn* iii tb«
atiiKM'phGrc to which Inc lifjuid i» exp«scil. This ie Inir iioC unly of oxjr^n
Biid WDtcr. but of 111! gusce and h<]iii()ii which do not rtitvr iiiUi rhctoic*!
c-onihiiinciotfuith ench other. Diticrent litjuide will, of rotirto, stwirb dif-
ferpDl gflecs wilh diftcring rendiiicMi bin, nilh th« anie g** itiid (bvMiW
li(|(iid. ihc aniouiu aheurbed uill depend directly i>n the pnrlini prrsflirv of
the gfls iu the oTerlyiOj; space. It should be added that lite proc<«# i» Riucb
jnBuenced by temperature. Hence, to stale the nialter Renerally. tlw ab-
soqrtioii of aDv gas by any linuid will depend ou the nature of the g». itie
nature of the liquid, tfae pieuure i>f l\ie gua, and the t«inpenUure at wbkb
biith stand.
Notr it uii^bt be Buppote<I, and Indeed was once supposed, that the oxygMi
iu Ibe blood wtu Dimply diwolved l>r ibe blood, if this were so, then the
amount of oxygen preteut in iiiiy given quantity of bloud exposed to any
^ven attnoaphore, ought to Hue and fall Ntendily und regularly as lite par-
linl preitiiure of oxygvii Ju that ntmo«pherv in iiicrruwd or diniiuiabed ; the
nliMirption (or tscaptO of oxygen ought lo fijllow wlml i» kuoivn a* the
Heurt-Dalton law iif pn-miin-s. But tJiis is found not to be the ca*?. If
wi; expose bloud conUiiiiiiig little or no oxygen to a siiccreiiion of »tmo«pheie«
contninin^ inurcusiiig <jii»iititips of oxygen, we find that nt tina ihi-rciia
very rupid abeorntiun of llic nvniluble oxyjjcn, and tht-n this Mtmewhul »ud-
donly censes or Dccomes very aninll ; nnd, if on the Other h»n<l vt: xubnil
nrlerial blood toeiicoesaively dimioishing preieures, we fin<l that for a long
time rerv little oxygen is given off. and then suddenly the etcape beoomM
Tery rapid, The abeorplion of oxygen by blood does not lolloiv the general
law of ohearpliori according to pressure. The pheDomena on the other hanii
BUggeBt« the i<leu that tbe oxygen in the blood is in some particuhir combi-
nniion with a substance or some aubstances present in the olood, llie combi-
nation being of such a kind that it b<il(U good during a lowering of pt^oaim
down to a eertuin limit, and that then diNfociation riaidily occur*; we nay
add that ihiit limit in vory cloudy dcp-tident on temperature. It u, bow-
evi*r, not to l>c NuppniH.'d thnt ii* the pri.fwure is luwMvd, no oxygen what-
ever it given off from the ■ubntitiicc until a certain point U reached, and
that at that point the whole store is iu an iustant disKciatcd. no more r«-
tnuintne to bi^ given ntf. The citao is rather that while pressure U beinz
lowereiTdown to a ccrtnin point, no appreciable dissociation takes place, and
tlutt then h.-iviiig begun it increases rapidly ivith euch further lowering of
pT7«suie until the whole of the oxygen is given otf. During this narrow
range, between the lirsl beginning to give oH' oxygen and the coiuplMion of
ihi- giving oH', the compound of the oxygen with the sulwlance or subetancca
may be *])oken of as partly, that Is more or leaa, dioaociaied. What is lb*
eulvtance ur what are the substances nitli vrbieb the oxygen is tiius pecu-
liarly conibioed*
If serum free from red corpUBcle* be u»cd in Nucb ahitorption experimeou,
it is found (hat. as roi:i|iared with the entire bkHid, very little oxygen is
absorbed, about as mud) ax would )>e nlworbcil by the MUne iiuantity of
water; and such an in abnorhcl ilix-s follon' the liiw of prratum. Iu natural
arterial blood the ipuinlity of oxygrn which cnn l>e obtained from serum is
excetslinglr amall: it <lot« not amount to half a volume in one hundred
volumM (if the entire blood to which the serum bclongetl. It ii> evideat
ihnt the oxygen which is priscnt in hlooil is in some way or other jK-culiarly
c»nnccl(.-d with the red corpuscle*. Now, the distinguished feature of tlie
nd ciirpusclefi ix the presence of hwmoglobiu, We have already seen 'J 24)
^
THK ItesPlRATORT CRAIfOBS \S TKB BLOOD.
457
I
I
tliftl thi« cotiMiluics !Kl per cent, of the drie*! ml i"or|HUi<;le«. TJi«re cnu be
A priori little ^l<r^lh^ tl)ii( this niUHt bo tb« subatntit'O wilh vrhiuh the oxygen
w (Dwiriatcd, itnd lo lli« propcrliM of this bmljr w« muA tlicrefuK <lirecl
our ntleiitiou.
S 345. Ifirmoqtobiii. When »«|>3rii(eil from the other conGlitiivnlii of the
sertini, hwii)oi>Iol>iD apjiears hs a fnibeiauce, eitlier nmorphous or erj'ntJilUac,
radily solume io wiitcr (eapcciully in Marrn wHi«r) and in wrum.
Kincr bntnoglobtB i* noluMc in serum, luiil Miioo tliv idrnlity of the cryMaU
»l>srr*r4l occuKiaally irjlbio the corpiHolcit irilb Ibnae obtained in olber ways
ahuna tliNt th« ha>inuelubiii a* it cxinu in (he corpittvie it tbe aaine thing n« that
wliicb is arliflrially (irepared from blouil. it la evident that Bom« peeuliar rrlutioii*
ahip b«lwern the Vtrcnia aod the bwnioglobin muM. in natural blooil, keep the
latirr froai l>einK iliuglviKl hr ihr lemtn. Hesc^, in prrpurinf; lucmnglobiii it ia
atemmty Am at all tn brrnk'up Ib'u cnnnTclino and I'l wi tbo biemnglubtn free
from Uie corpuaclvH. Thin mny be done bv the aililitloa of WHier, of ethrr. t>f
chloroform, or of bilp *«li«, or by r«'|>eAtealy fret/.iiig and ibawiriB; blood »»
treated becoraoi " laky " ((/. { 24].' U Id a)»n'of advniitaee prcvicnmly lo remove
the alkaline ■erum lu tiiu<;h a* piHwiblc. hi iw to oj^rnlc only on the red coriinitcJe*.
Tbeatronta and ba^mugiubin btiiif; tfaiM tvpHraied. a Butiitlun of bu'niotflobin in
lb« rMult. The allcalinily oT the BDluti'-'ii, when pmeni, being rcdiiceii by the
catitinut a'ldiiiim of dilute aeetic aoi<i. and the Mlv«n( power "f the wiiieoii*
m<diuin beini; diminialml by th<^ iiddilion of onft-l'ourth tl* bulk of ulc<illul, llie
mixture, lei ajtide in a lemperalua- of U* C. in order atill further to reduce the
aolubililjr of tbe hieiuoclobln, readily cryatalliiea, when tbe bloo>l ii^e-i i' that of
the dog. eat, hor»e, rat. eiiinea-jii^, etc. In (be eaae of the dnf, indon), it la
aimptr atilBcietit to add dbrr mrefully to the blood iinlil it jual becomr« " laky."
and then lo let it viand In a coal place; tbe mixture toon beoomeaa miMof
crratala. The cryitaU may be (eparated by lillralion, rediaaolred In water, aod
re-cT)Ualliie)l.
Hemoglobin from the blooil of ibe rat, guinea-pit;, X]>iirr«I. bMlt^bng,
bDrre, cat, dog, gixve, and some other animah, crvHlalliites readily, thii crvn-
tab being generally blender, lonr-eided iiri^mu ttcloukriuj; (o llie rhonioic
qratem imd oAeo npfesrini; (|uile
adcular. yFigs. \M. 1^5, IM.] the
fiiTTBtds from the blood of the giiineia-
pig ar« octahedral, but also Moog to
the rhombic aystein : those of the
SQuirrcl are six-eided t'l^'*^ I'he
blood (if the ox, ^heep, ruhljii, utj!;,
and miiu cryMflllizi(« with difliriilty.
Why the*« ililTereuns exist ii not
known; but the c«in{iii#ilii'u niid the
unountof waier«fcry»lKlIi/alii>ii vary
soineuhnl ID ihtt cryntHlii ol)iaiur<l fr-itu
difttrreiil anii»ul.<. In the do){ the
[>crceiit»;ff (•omimaition of the cry*-
IhU ban Ixt'ii detcnninrd im C. ri!l.'<>5,
H. 7.32, N. 10.17. (>. m.M. S. O.XU,
Fe. 0.43. with -i to4 percent. of naler
of cryttnlli/HlioD. It will thus tte
wen that hnmoglnbin cunlninK, iti
nddition to the other cleiuenlR
tn-.. iM,
■ *-^.
Tn««iiMiN4L raDH Iiiuoe> o» tiu rio.i
to the other cleiuenlR usually present in proteid Biibelancc». a
certain nmotint of iron ; that is to My. the element iron is a distinct part o{
the hwmoglobin molectde.n fact nhich of itself renders biemo|jIobiii remark-
able amoiii; the chemical subttances prcwnl in the animal body.
i 346. The crystals when fcen in a sufficiently thick layer uuder the micro-
act^ have the tome bright scarlet color as arterial blood has to the naked
4r>s
IPIKATION.
«)% : when aeen in a maas tbev natuniUy n|>peAr <lnrk@r. An aqueous Bolution
of tmnuiglobiii, obtained b7<)iaatlviu|( [luHlied or^tnls iu ilLslilleil water, lia«
atoo the HUne bn){bi arteriitl mior. A. tolerablf dilaU Mlution phoed before
ibe apectriMcope U fouud t<i alHorb eertaia rays of lis;bl in a peculiar and
characteruuic manner. A jiuiiinit of Ibe red eaa of the apectruia is absorbed,
(no. Hi.
[ni>.iM.
ti-
l^*"
<
llRIAiMMill.CBVvrtMl'IIOli tlUKiDnr KgltlDRll.
KT<i»i<*il InstWIlBM ig*na«r. noi iinfn^iitnllr
llmttTiL' r«uM llinttx RijMB.]
M i» also n luucb larger portion of the bluo end ; but what is aiml "trikin^
ti the nreaeccc of two stron];;ly marked abfl'>rptioii bands, lyin)^ between tbe
Rolar linex /> and A'. (Se« Vig. 137.) Of these the one toward the red
side, fiooielimea spoken of as tbe band (a), in tbe thiDDeM, but the inoat
intense, and in extremelir dilute soliitioni (Fig. 1^7. I) is tbe only one
vittble; its middle lies at some liule distance lo the blue side of I). Iu
position may be more exa«t]y deRned by eipressios it in w ave-lenutbs. Aa
la well-known, tbe ravit of tight which make u|> tlie Hpeclrura dilTer in tJte
length of their wav«M, dimioiHhing from tbe red end. where the wuvei* are
longMl, lu the blue end, where they are ahorte&t. Thus. Frauenhotbr'a line
D correspODdii to rayit bnvlujc u wave-leiwth of 589.4 millinntbH of u railli*
metric U»in^ tbu name unit, the centre or thix abttorplioii, bund », of hiemo-
glohin corrvMpondii to tbe wavc-leuj^lh 578; us may bo seen in Fiu;. 1^7,
where, ho«ev<ir. the nitmbcn of thi; div-tuon* of the dcalc indicate nnly
100.000 of a millimetre Thi? other, iii>riietime« called ff, much broader, lie*
II little to the red tide of E, ila bluewanl odgc, even in modcmtely dilute
■oUilion^ (Fig. 1S7, 'i), coming close up to that lino; it» centre corresponds
to about wavelenglh o')0. Each band is thickest in tbe middle and gradu-
ally thins away al the edges, These two absorption bands are extnmely
characteri)itic of a siilution of haMnogtobin. Kven in verr dilute solulioDs
both bands arc visible (they may be seen in a thickness of 1 em. in a solu-
tion containinj; t Kroi- of liiemoKlobin in 10 litre* of water), and that when
scarcely any of the extreme red end and very little of the blue end is cut
off. They then appear not only faint but narrow. As the strength of the
solution i* increaieJ the band* hmiden and bsome mire inlen*e: at the
same time bith llie red end, and still mire the blue end, of the wlfito spec*
Inim are encroached upon (Fig. 137. 3). This may go on until the two
alMorptioD banda beoame fused to-^ther into one bnia 1 bml (Fig. 137, 4).
1
BRSIMRATI
The only ntTfi <>r liglii. nhicli ihen pom tlirnD^h thi- bwi
are those in Ihe |>rccii between the bluennnl eitge iif ibo united Iuidi1> buiI
the geil«ral abeorplinn, which is now rstiidly sdviinciii); Cmm thf blue toA,
aod Umm ill the red betuceii the united mind^ mid the geiierni alK»r)itioa ml
ihe red end. Jf the aoliiiioii be elill fiirther iucreoaed in &lrciigih, Inv inter-
Val uu the hhie xide of thi^ united bands beconiefl absorbed atai, bo tb»i ilia
only rmya which i>asii ihrtiugh are the red rays \yios lo the red sido nf 0;
th«M arc the but in disa|){ieHr. and lieiice tlit.- iialunu red color of iIk- wlu-
lion ii» unea bj trnuiiniittLiI litcht. Hxacilv tW sanae appearances mn ttta
wbdi crvMHlit af hiemiif^hihin are ixumiueil with a inicnvspeolroscope. Tker
nre itN" m-cii n'h<-ji arterial bluod il^i'lf (dilutvil with ^ine Mitiitiotts, to that
tlw corpuscles remain in n.-i natural » condition lu puwiihie) u exaniiued with
(he sprctroscopc at well a* when u ilrop of btiind, which froin the necemirv
exposure to nir is nlways nrterinl. ii" cxaniimid nilh lh« micro-sitectrowofL
In fact, the Hpectruiu of tiiciuoglribiii is ihc «|nviriini "f nurinal arterial
blood.
5 347. When crystals of hacmiiglohin, prcpttrcd in the nay dcuciibcd above,
are Biihiectcd to tlie vacuum of the mernirinl Hir-|nimji, they give uff a certain
quantity uf oxygen, and at the same iJnie they chniige in color. Tltc rjuautitr
of oxygen given olf is deSuile, t giiu- of the cryttalf giving off 1.5i* ce.
of oxygen nieaeured at 7l3U mm. U^'. ami U'^ C. In other word». the cri-»taU
of bKnioglobin, over and above the oxygen which enters iiilimatciv iut<> iha
oompoeitJQU of the molecule (and wliich alune u f;ivca in the eleiucu
coiDpontion nri'viuuHly stiitw)). cnniaiti another ijutuitity of oxygen, whit
in looee conifiinnlion only, and which may be dissociated from them by •
jectinj; them to a sultimnily low pnjwure. The chau^ of color which
ensued when thi> Imwcly cuinhined uxygen ia r>:miive<l, iti churacieristic ; tbt
crystals bci-umv darker and more uf a purple hue, and at the aaiDfr tim*
dicJiruic, so that while the thicker ridg«« aru purple, lli« thin edges apfwar
gT«eni»1i.
An ordinary solution of bicmoglobin, like the cryrtttis from which it is
fornted, contains a definite <|Uantity of oxygen in a similarly pcctiliar loots
combinntion : ihia oxygen it also gives up when tubjocled in the air-ptimp U>
Butliciently low prcMurc, becoming at the same time ot'a piir^ilish hue Tbii
loosely combined oxygen may also be reniovet) bv passing a strenm of
hydrogen or other iudiHereul gas through the solution ; the stream of
hydrogen acta tike an oxygen- vacuum to ihe hienioglobiu and thus di»a»o-
cialion iseSecIed. Carbonic acid >;as is uusuitablel'or this purpose, since, at
ue shall see, huiiis; an acid it acta in another way on the hemoglobin. The
oxygen may alto be rt^movcil tVum ihi- hitmoglohin not only by phyucal bul
also bj chcinienl means, as by the iiaie <•( reducing ageNts. Thus if a few
drops of ammonium sulphide ur of uii alkaline .■Hilntiim uf ferrous sulphate,
kept from ]ireciiiiiHtiun by tho pn'M-nce of tartaric acid, he added lo a solu-
tion (if hiemugIol)ln. or even to an un|>uTilicd Solution of blood eor|>uscIea
•ueb II* i> airunhd by the woshiiigv from a hluod clot, the oxygen in looee
combination wilh thy hiem«);Iobin is immcdinlclym.'ij!ed ujwn hy the reducing
agent. This may he rrcitgnized at once hy the chBraclcriMic change of color;
from a bright scarlet the solution becomes of a purplish claret c<dor, when
teen in any thickness, but greenisli when sufficiently thiu ; the Color of tbo
reiluccd solution is exactly like that of the crystals from which Ibe looM
oxvgcn has bri?n removed by the air-pump.
Kxamined by the B^iectroscope, this reilueed solution, or solution of nWucvd
hanaoyliAiti, as we may now call it, otters a spectrum (Fig. 1 37, fi) very dif-
fereot from thai of the unreduced soluliou. The two ab'oriiliun Wild* Imve
diBaiip«re<l, and in their place tboie b seen a single, muca broader, but at
THK nK9FIRAT<
461
I
I
I tU mtat tvat niucb fainter Imixl. wliimc miiMlo <>i-('ti{m« ii iiiuiilian ntmut
mUnr betvecn Uk (wu nbiiirpliuti ImikI* nf iIil' iiiiix'ilucnl solutinu, tlxnigh
tUftivKld edge of tbe bnnd ^IiikIi-h nntn n»tlii-r I'lirtlK^r townni (lie ntl
tin don the other edge townni ihc liltir ; iu ci^nln! cormnioiiil* lo iiliuut
tmlMglli &S5. At thv Mime liniir tbr ^mi'ra] iilwnrption i)f the »pceiriiiti
I lidiArnl fttMD that of tli« unrcilucvil mliitioD ; IriM of \lw blur vutl i»
tituhld. Eren wlien the wlulinno bccnine tolcrubly cowrentrnlcfl. maiiir of
lit UaUi-grteo rajv to tbe blu« rkIc of (ho siDgl<i bitiid still y>n» through.
Bnmha dHhnae» in color Wwoen Iwnioelubin which relniriB ihi^ InoMly
owiUnMl oxrgrti,' nnd hinDoglobin whicli hiu Icwi ii« oxrci^n ntid liccome
'•diKtil. ]n tolerabljr ooncenlnitec) solution*, or tolerably thick Invcre, th«
^natr Ivl^ Ihrnugli lti>- ml and the omti^-Tcllow rays, the )»lter tbe red
'Ofl ill* hluich-^'nvii rave. Accordingly, the ono iipi)cnr8 ecnrlct, the other
Ijurplc la dilute •ulutioDe, or in n thin liiycr, the reduced biemo)>!obin leta
'^>^gh H> nmch of Ihe green ravn that they jirepoudernCe over the red. atut
'**« t^ltins imprewoD ia one of t;reei). In lite unreduced huemiit-ldbiii or
^Ky-hviDo^lubin. tbe jiotem yellow vhich is blocked out in ilic reduced
'^^■ofllabiD make* Hmf fell, so that a very tbiii Inyer uf oxy-hieinoglobia,
** b a ainjcle eorpaade seen under the mteroscope, uppeara ycUovr rather
*ha« red.
It tniiat be reroerubeml that wlwu we fl)ieak of reduced hKiDoglohiii (or
tare briefly iueaio{[lohin), with s puri>l6 color aix) a characteriuu! one-bunded
iftetnta. «e roean h»tnoglob!i) wliidi ha* lu»{ all iia lonely Msitciated
mygea. If a <|uaDtity of osy-hRnidglohin t>e expowed to an iniiulTicifiitly
iam pveMurc. or to the adioti of im in.tuflicieiit ijunutitr of tliv retliiciiii[
leUon. it give* up a part only of Ui' oxyfren; it in only jxirtly rciluceil.
ShH) a IwrtlT n.'duccii (ulution still shnw* thv Iwo iMmtsofoxy-hienioglubiu.
S 348. Wlwn the hatmoglolun wlutioii (or cryslul) which hiu K>et its
iajgm by the action either of the nir pump or of a reducinj; ngent or hy the
pMMgtOT an indilli^rvnt gns, is expoeed to air containing oxygen, an nbeurp'
doa of oxygen at oiioe takea place. If suflicient oxvcen ne present, the
fc— inglnhin aeiiee upon aufflcieiit oxygen to obtain its full compleioent. each
■laaHM taking op in comhinaiion 1.59 c.c. of oxygen ; if lhen> be an
MarfMeiit qnaotily of oxygen the bn.>iuogl<>bin still remains partly reduce<l:
or pertia|» we mar »ay thai a juart only of ibe hemoglobin gets iia allowance
■ bila ibe remaiudor contiuuea leiluoed. [f the amount of oxygen be auffl*
cieM. Um tolutiou (or crvxal I, a* it takca up the oxygen, regaina its bright
timiUi ooloraitd iia characieristic abaorption spectrum, ihe single band heing
iitilioul by the iwo. Thus if u totulton of oxy-birinnglobin in u tnl-iutie,
■ner being rnlucnl by tlie action of a drup or two of ainnicmiuiii nulphtde
mlkatiim and thui> itbowing the purple nilor and the nngle baml, Iw «hake»
m with air, tbe bright acnrlcl color at onc« returns, aixl when thv Huid is
plaeid btlhr* ibo epodroacopc, it is anvn that the sinslv laiut broad hand of
ihe ndnrrd birmaglohtn has wholly disapitearcd, and that in Ita place are
the two sbarp tbinm-r linnds of tlw oxy-hirmoelobin. If left to stand in the
toC-tubr the <|(inntilr of tvducing agfut still pn«.<nt is gem^mlly niRicienl
aMatn t<i nd> iIh- hii-Mi»glol>in of the oxyg^^'U thus newly acipiired, and noon
toe acKriel hue fade* bacJt again into the ptirpic, the two Itands giving idaov
Lo ibe dUe. AiHither stmke and cxiHuure In air will, liowi'vcr, again bring
Iwek tbe scarlet hue and the two biin>t> ; and once more ib(«u niiy dbappMr.
In &c<,afeKdro|Mtif tbe n-dut-ing Hniil will allow this ^anM of hMmoKlobin
uktBg oxyceii (r»ta the air and giving it uj> lo tlw reducer to be plnyM over
• fs l«vtUr*i ■•>• HBOMTcall Uir li'ii»«l»bln MilaUiIna iiiiBii Ui k<iiH<iiinlilBallv*,AnpAr-
m^bb* M>l Ih* lu>*<f IuUd rViiia ahiili titii huKl; cvniMbfil iiDXn'i li>< bBra rriiH>TT>L (vlocnl
kaaBaUtn •> d)i|<| liBin>i«hriiin
462
RKSriRATIOK.
ud over BOiQ : at meb tiim of the frnme t)i« ciilor thifi* from Nmrlf t lo
{Hirplc, aixlfrutii i>iir)i)i,>tu*c«rl«it, wliifv ihc two biintU «xc-biingv for the uni,
mid the niiu lor llic Ini*.
S 349. (.iflor of vrnoH* mid itrltriat klowl. KviilcDllj vc luive itt thcM
propcrlien of hicmogluliiti nii i-x|>laniition of at Inut oucOuilf i>f the gTMl
rwi>ir«tun>' |>n)(.Ti«, hihI tboy ttwL-h u* ilx' iiinming of tlw cbuigo of coW
which take* I'laoc u*h«n vviixiis MikuI Ix^-imit* iirk-rinl or an«rbil veDous.
In venoiis blooil, a« it ifnn-n tVum iW righl ventricle, the oxygen pr«wa(
IB iiuufficwnt to imti»fy wliully ihc hirmoglotiin of the rm! corptiicl^e ; the
hftnioglobin a, to a. largo «xt«ul, rediifvd, htnic the purple color of veiiaas
bhxtd. Wben ordioary venous blood, (liliiicil wiihuut acceu of osTgen, ia
brou^t before the Mpectroecope, the ino bonds ot oxy-bieinoglobin are wen.
Tbis is explained by tbe tnct that iu partly reduced bwmoglobin, which we
may conveniently regard as a mixture of ozy-haemoglobin and (reduced)
bKtuoglobin, the two gliarj) baDda of the former are always much more
radily aeen than the iiiui-h fainter baud uf ibe latter. Now in ordinary
venoua blood there is always nuoie louae oxygen, removable by diminiahed
prtwure or othern ise ; tiie hwmoglobin is only partlr reduced, there is alwava
•ODi«, indeed a considerable (quantity, of uxy-bsmoglobin as well as (reducvcl)
bttmuglobia. Ii is oiilv under special cin-umMancea, as for instance after
dMttb oy what we mIihII presently !<i>eak of us asphyxia, that all Lbe Ioom
oxyno of tlie blood di-iappcuirw ; atui then the tiro bands of oxy-hwrnoclobin
vanub too. If even only a Miialt (jUHnlilv of oxygi'n he prMent, so dtaiinct
are thi) two buuda that a wiliiliou »l'c«mpMely nnluced ha'inoglabiu may be
utwd HM a t<»t for the presence of oxrici-n ; if oxvgi^i be prrsrnt in anv fluid
to which thv rt«luced hucmogluhiii m iiddcd, iliv «i»f;Ic Iwnd immediately
give* way to the two blinds of oxy-h.-i-nmglobin.
As the venous blood puwrw thmueh ihr capillaries of tbe lungs, this
reduced ha-uioglobin takes from thi'ptiTnionnry air its complement of oxygen,
all or Dcnrly ul Ibe btemogloliin of the m\ ci>rpiiH-lni becomes oxy-hmmo-
globin, and the purple color forthwith shifts into K-urlct. For carefbl
<^i«ervatioiu> shon- thnt the h:iMHogtubin of arterial blood is saturated or
nearly nturalcd with oxygen : it probnhlv falls short of complete saturation
by aliout 1 vol. of oxvgcn in lOU volt, of blood. Ry increasing tbe pressure
of tbe oxygen, un additional (luautiiy may be driven into tbe blood, but this,
after tbe nwmaglobin hos become completely anturated. is efleoted by Bitnf^
absorption. Tbe qunniity so added is extremely small compared witb lbe
total quantity combined with the hwmoglobin.
Paaing from tiio lel^ veolriclo to the capillaries of the tiasuee the oxy-ha>-
noglobin gives up some of its oxygtn to llie tissues, becominf;. in part,
reduced lui-u)oj|;lobin, and tbe blood in cunseijueiiee beeomea once more
veoous, with a purple hue. Thus the red curpusclee by virtue of thnr
htnnoglobin are eoipbalically oxyKeu-carriers. Undergoing no iutriusic
chniigfT in iisclf. the hieinoglobin (M>ml>ines in the lung* with oxvgcn, which
it i-arri<^ tu the tinbues; these, mure greetlv of oxygen thau itseltTrc^ it of
its cliargf!, and the reduced luemoglohin liiirriea hack to llie lunpi in tho
%'enoUN blood for unother porlitHi. I'he change from venous to arterial blood
is then in part (for as we shall tiee there are other events as well) a peculiar
combination of hiemoglobin wtth oxygen, while the change from arterial to
venotiK is, in part also, a re<lunioii of Dxytiieninglobin ; and the diflV-rence of
color between venous ami uricnal blood de])eudM aImo»i entirelv on the fad
that tbe reduced hiemoglobin of the former '» of ]>urjik color, while tbe
OXyJlRmoglobln of the lutter u* of n scarlet color.
Thero may be oilier caiiffn of the change of color, but tliesc are wholly
subsidiary and unim|>ortant. When a corpuM^le tiweJIs, it refrmclivc power
TUB RKSl'lRATOltV CIIAN'OE!} IN TUE BLOOIt.
4dS
I
I
I
I
I
u diniininubcd, imd id cwiiiequMice tbi.' numlwr vf rap which piMt iiitu tsui
are abeorbed by it nrc invrcnsctl at tb« PX|«nM of ihuw ruflccli^l rnxii iu
•UfAeo ; anyliiing iht^rcfurc vrhich iiui-Dk the corpiji>cJn, »wh tu iW ii<ii)itiMii
of nntor, ten<l» tu durkeo tilood, and uiythiag, «ucK •» n c^iccutniCed siiliiie
tuliitiuii. wtiicli ntitcs th« corpUBcInf tu shrink, \em}* to brighten blootl.
Ckrbonic iicid hii« npfmrcDtly i>»m« inllucuce in swdling tUe ocirpusclee, mid
tberalbrv muy nid in durkcniiig tho veiiotM blood.
^ 360. W« biiv« epokcn of (ho cxmibinution of hiMtiQKilobiR with oxjgea as
beioK « peculiftr oae. Thi> pcciiliiu-ily conaats in the facta i^nt tb« vxyiiea
tDBT M Mtociated niid difsocintcd, without tiny (fcncrnl disturbauie ol Uie
tDoWOole of hieiDoglobiH, and that disHidBtion niny be brought nbniit very
readily. Hnmoglobin combinw iu a wholly similar QiaDn«r with otker
Kases. If carbouic oxide (monoxide) be passed through a sxluliDU iif
flseotoglobiu. a chsnge of color lakes place, a peculiar bluisJi tinf,'« making
ita appearance. At the bame time the s^ieclrum is altered ; iitu bands are
Mill visible, but ou accurate measurement it U seen thai tbev are placed more
toward die bliie end than are the uthcrwue similar bands ui uxy-nieinoglubin
(see Fi);. 137, ti>; tlieir centres oorrw|>aiidiug respectively to about wave-
len^ctlis oi'i and r>3:t, while thuei^roxy-hffiiDuglubin, as ire have weii, corrc-
.ijMnd tu 578 and 539. When a known quantity of carbonic oxide ens is
•ent tlirough a httaioglobin nulution. it wilt bn found on examiuHtion triat n
certain amount of the fiai> him brcn n^tninrd, nn c<|iial volume of oxygen ap-
pvariog in iti plait- in llic^nii wtiirh imum from the solution. If llie solution
su treated ht cry»Iiiili/x^l. i\\v ■rplnlj' will have the mmc cJiaracteriatic color,
and give the eamu ahmtrption Kpi.-clnini us the solution ; when subjeeted Iu
tha action of tlw mercurinl pump, tht-y will eivi- off a delinite quantity of
cnrbonic oxide, 1 grnminc of the eryiflnU yirlditig l.'>9 c.c- of the gns. In
fact, hwmoglobin uoDibincs loo«ely with carbonic oxidr just ns it aoea with
oxygen ; but its affinity with iho former is greater than with the latter.
SVnilfl carbonic oxido nwdily turns out oxygen, oxy^o cannot so readily
turn out carbonic oxide. Indeed, nirbonic oxide has been used as a means
of driviiyt out and measuring the -iiiiflntity of oxygen present in any given
blood. This property of carbonic oxide explains its poisooous nature.
When the gas is breathed, the reduced and the unreduced unmoglohiu of the
venoua blood unite with the carbonic oxide, and hence the peculiar, bright
cherry-red color observable in the blood and tissues in oases of polsoningliy
this gas. The carbonic oxide hterooglobin, however, is of no use in respira-
tion ; it is nut an oxygen -earlier, nay more, it will not readily, though it
dooBso sluwljr and eventually, give up its carbonic oxide for oxygen, wlx'u
tbe jioiiiODoiia fpe ceasea to enter the oDen and b replaced by pure air. Thi;
orgabisro is killed by suSbcalion, by want of oxvgen, in unite of the blood
not aaaumlBg any darlc venous color ; to adopt a ]>t)nuR which has been used,
the corpuscles arv pnrul^ »cd.
HiCDid^loliin Miuiiarty fortiiit a com[H>inH', having a characteriiitic spec-
trum, with nitric oxidr. more stable cvi-n than that with carbonic oxide.
It has lieen suppowKl liy H>me thai the oxvgen thus iuw>cialcd with hnnio-
globiu Is in the condition known n» unouv; f>ut thv arguntents urged in »ui>-
port of this view art iucoiiclusivc
J^rvdttcts of the DtfQmpontimt of Hamtujlobin,
g 961. Although a crystalline body, haunoglobin difllises with great ditti-
ctilty. This arises from the tact tJiat it is in part a proteid budv ; it con-
sists of a colorlew proteid. aMocialeit uiih a colored KulMianre, w^ich may
separated out from the fawmoglobtn, though not in the exact con<iitio» in
RKSPIBATION.
I\' exisl« in lh« corapoiin'l : tliu subtfanM wh«n HpnnilMl
BfeDvnt'h-re)! hoily known tin turm-tliii. All rite iivin b«-
pnglobiii i# ill ivttliiv nitnclietl to th« liicninliu. A eolutioo
tvh«D heatod. coagtiliitc«. tbe rxaci decree at wbiclt tli«
place (le^ndini; on ttie amount of dilution; at tli« bame
rti from the seKiiig free of the hwinaiin. If a Btronc •olu-
tin be Ireaioil with acetic (or oiherj aciO. the tame nrown
ppenrance uf hseinatiu, a observed. Tbe proleid codmUu-
lot coagulated, bui by the action of the acid i)a&«t Jntu
kibuiuin. Un itdditi}; ether to the mixture, ami tbeikins.
3J9Molv<!il ill tbe supernatant acid utbcr, irhicfa it colon a
ich, lixnmined nllli tlit «[iecti>uroi)e, iit f»nnd to p(M»e** ■
truin, ilie Hiieciruiu of the it»-culle(i acid liicmniin i>t Stokts
'he proU'id 111 thv water Im-Iow the ether appeani in a ci>a|^-
to the action of thi' eihcr. In n .■"uncwhnt nimilnr maniter
.in^mo^dubin into a prot«id cnnrtiturnt and b»nMtin.
ire of ihn jirotfid coiistiliiwit nf bipmoglobin ha« nol a* yet
rtiiinol, 11 wiw "ujipoiii'il to be globidin (hcnco lb* itaiiH-
conlractcd int^ biemoglobin), but ihoiieb belonging lo the
has ch.iractcre of il« own ; it is poHibfy a miiture uf iwu
proieidB. It ha« been proviiiionally named fftobut and i»
)iu ash.
n wbcn Bcpnrnted from it* proteid fellow, and purified,
rk-bi'owu amorphous powder, or aa a sualv mav with a
iving the probable ooniposilion of C„, H„. N,. Fe. O^.' It
n dilute acid or alkaline solutioas, and then givee ctiarac-
i-ig. 13M,2.ft).
fvatiirc ill hwniatin it that Its afhtlwe solullim ia ca[Mible
by rcdiK'itifT agent*, tlit' Mpealruni changing ai lh<- Mnte
), and that the r«.-diicc'l solution will, lik« iho bieitiogldbin,
Igain on being brought into contact with air or oxrnu.
to indicate that the oxygen-holding power of hamogiooiD
uivoly with ile haiiiiniin con'titucnt.
)f strong eulphurie add btNoatin may be robbod of all its
liDB the feature of poowwng color, the solution of iron-free
dark rich browni^h-rcd : but is no longer capable of com-
tli oxyeen. This indicates thai tho iron it in tome war
be pei'uliar respiratory functions of hntuo^lohin ; tl)oti;>n
error to suppose, as was once suppooed, that tbe change
ierial blood consista easentialty in a change from a ferroua
MalUnblB itaelf, biero&tin furnis with hydrochloric acid a
ring In ntlnote rhombic cn'stals. known as hixmin cry^lala.
I left until it decomposes, the haerooglohin is very apl to
Bto a ptTuliur boily linowu as meilnrmoglobin, in the apec-
vcrr coiixpicnou-i band i.i Keen in the red between rnnd D
"the same change may be bri>U);ht about by the action
ch as carbonic acid, by ozono, an<l iiy other agetili «uch n*
pKBi pcrmaneaiialc. When a strmin of rMTlMnic add is
pm or through a solution of hiemoglobin tb«t baad In the
of meth:cmogk>bin soon mnkc* it« appearance. Mvtb«f
rat little, if at all, in wlcmpiilary composition from biemo-
Dtained that it coDlains the Mime <|uantity uf oxygeii aa
Mil 4UW "f llap|<«.7vylat ^ Ui>tKi*««>" )Ui.U Hid niMe (Moat auAfnteUr
4m
REitrlH^TlaN.
oxj-hwcnoKlobiii but in a more stable ooodition, nore iatinuudy asBocimH
with the molecule.
Ill ctitid union . the c-oiiililion of oxr^eu in Lite blixxl b lu fuUtfirs: Of ilw
whole ((iiaiitily of oxy^u In the bluud, only a minute fractirm En siaplj
ilbitorlwd ur ili.iniilved ai-i.-ur'litiK to the Ixw of |ireitiure« (tlie Uoory-DnlloD
law}. The gtcal iiiiLw in in astute of combination with the hnmoglobiD,
the cunn«cti«ii being of wieh » kind that while the licnKif[lobiu rcadilr
combine* witli the oxygen of the air ta which it it expoacd, diMariatioii
rcni]ilv octrurH at lnw pn^KiUKu, or in the [ireM^nce uf indilrercnl gium, or by
the iiclion iif ^iilMliinrist having u lerenler afBnily fur AXyj^n than has hirmo-
globin it^t'l)', Tlic (lIir<'n'n(N- ln-lwucn veiiou* and arterial Muih), tu far m
oxyeoQ \t I'lmt'ernvd, in that while in arterial blood iho hn-niogloMti hoUi
nearly its lull oomnlemuot of oxTgrii and may lie >|>i>ken ufiks nearly
wholly oxy-hirmoglobin, in Tonoiiii blood the hn'tunglobtii \a to ■ lar^- bat
variable extent, nxluccd ; and thu cbumoterixtic uniors ol' rcnoui iin'l arte-
rial blood are in the niiiin due l<i the fact that ihp color of reduced hsnw
globin is purple, while that of osy-hiem»gM>in is scarlet.
T^e Jteiailoni of Ihe Carbonic Atul iit tic Blood.
g 363. Tlie umence of earbonic acid iu the blond ^)>ear» to he di.'Ur- —
mimed by conditlonit more oomj>lex iu tlieir nature and at proient not eo well
underatoiMl a.i thu«e irbidi deierdiinc the prcaencc of nxygi'ii. Tha ntrbonic =
acid is iK't »imply dixtiilvf.fl in the blood ; itit alMiirplinn by blood diira noL^
follow the law af prci«iirc-'>. It. t^xi^ta in iwwii'ialii m with Hi>mc »iib«tanoc tr-
eubetonenf in the blnoii, and its wcapcs fmni ihc lihiiid io a pnicivs of disM—
ciatiun. We cannot, howerer, iipeiik ft ii iw beini: KK«K-iHte<l, to the MniB
extent aa is the oxygen, with llic hienioglobin of the rrrl oorpiwclea. &»
far fruui the red corpuscles cootaiDing the grvat mass of ili« carbonic add,
the ({uantily of this gas which is present in a volume of serum is aceurdiug
tu Home ubtervers actually greater than that which is pre«nt in an *.i)ual
volume of blood, i.e., au ecjual volume of mixed corpuscles and Merum:
thai is to »ay, the carbonic acid is much more largely aaiooiatcd with
theverum (or, in the living blood, with the plasma) than with the red ntr-
pusdea.
Wheu Rcmni is siibjecled to the action of the mercurial pump, by far tlx
greater part of the carbonic noid is given off; but a small additional ijuan-
llty (2 to ^ vok |)erc<tnl.) nmy be extracted bv the lubsetiuent iiddiiion of
an acid, Thii- latter portion may be spukt-ji of* a^ " fixed " carbonic acid in
distinction lo thr laigrr " looM "" portion which b- given off lo thv vacuum.
When, however, iho whole bloiHl vt suhjwted to the vacuum until the car-
bonic acid ccasM to ho given "If, ihe Mibfeqitunl addition of arid i» said not
to eet free any ftirtlicr quantity : so (Jiai when K^riim in mixed with cirnusclea
idl tbe carbonic arid miiv Im- spoken of as " kmav" : and it i* stat^id that the
excess of carbonic add in ■ quantity of serum over that pretenl in tlie aatue
bulk of entire blood, corresponds to tho fixi.'-l [loriiuu in snrura which has to
be driven olT by an acid. Moroovcr. even ihooe who maintain tlint the
quantity of carbonic acid iu entire blood >* Imh than that in an equal volume
of nrum, admit that the carbonic acid vxistii in N'>mc way or other at a
higher preeuire in, and is more readily given olf from eatire blood than frum
serum.
If tli«M BtatemenU be aceepted it seem! iirobablo that the cartMuic acid
exiua aMOciated with some suMtauoe or sulMlanci^ in the eerum, or rather
l>IasiDa, but that thecondillooaof itaasmoiation (an<l thereof iudiMociaiiao}
are dcUrmined by tbe action »f some Bubaiauce or subitauces present in ih«
RBI
467
«orptiecl«£. It has been suggMted tliat ibe aMociaii»i) uf tbe «Hrl>uniQ m-id
ID Oie p|[u>ma b> with one or other of tfa« proteida of ilie plaaiua^ but it ba«
alao been eu;-i;t«i«Hl that the turaciatioD ia one with eodium a.i »oiIiut)i biunr-
bonate, and further ibat tU« hnmoj^lobin of the corpuDelen [itay!i a iiart iu
promoting tJie diasodalion of the Htdium bioarbonate or ewn the caruoiiaic,
and tbu» keeping up di« carbonic acid of tlic wtirc blooii. Oilier obwcrver*.
bowever, mamtatn that the plaania doe* not holii Ihin ex<!tuNiv<! po««won of
the carbonic aoi<l, but that a ci)UNi<lcrab]« miaiirity uf iliUi ^aa \t iii mmv way
•taociated with the ml corpii*cti.>«. Iiii]<«<i, ftirilicr iiivwugatioiu arc n«coa-
Han,* hcri>re tho matter van \k mill to have l>ecn plimd on a satisfactocy
foot I tig.
7'A« IMatianM nf the JffUrogfn in (he Jtlotxl.
i 304. The email quantity of tbia j^as whi4:li a present in both arK-rtal and
veiUMia blood aeemi to exist in a state of simple eolutioii.
The Rr.4i*titATi)RY ('iianoi» is tiie Liinim.
The KttlniMfe of Oxytffn.
i 353, \V«> hnvp already twn ihal the blood in paxtin); through the lunga
lakee up a certain variable oiiaiititir {from H to 1*2 vols, per cent.) of o.ty^^.
We have further wen that the quantity hi taken up. putting aside the insijf-
niflcant fraction dimply absorbed, cnterB into direct out loooe tiombiualion
with the b»m (■(;'** bin. In drawtng a distinction between the oxy;:;en simply
alMiirheil and ibui enterinc into coiubinatioa with l)ie lueaioglAbln, it must
itot be understood that the latter is wholly independent of pressure. On the
contrary, all cJientical compounds are in various den^reas subject to diieooia-
U(in at certain pressures and lenii)eralur«4; and the exutenoe of the aome-
vfbat loose Gorapouod of oiygen and buuo^Iobin is dependent on the partial
prtaiure of oxygen in the atmosphere to which the hKnioglobiii i» ex]>0Md.
ifol only will a suliition of hiemc^lobin or a auantilv of blood citlier ahnrb
oxygru, uud thus undergo aa»cialion or unaergodtmociulion and give off
ox'ygL'ii According t» the [lartiul preaaure of oxygen in tlie a tin imp}! ore to
vrbicli it t» expiiabd ia high or Inn', but alto th« amount taken up or eiven off
will df'piMid i>D the d^rvM <>f the jiarlial iinMurs; the hatmoglobiu as «•
have tixn may be partiallv an well an wholly reduond. Tho law, how«r«r,
acojrdiii); )•• which abwirpli'm or nicape thus tiik<« place it '{uitc diflerent
froni thai olNUTved in the simple alMorptimi of oxygen by liquiii*. The aara-
ciation or dimiiciatioo b fiirtbormpoeialty dv[H-ndent on tMnpcrnlure, a hi^
temperature favoring diiwociatifln, ao (hat ul a high tomperature laea oxygen
is taken up than would be taken up (or, a* the ciuc may Ixi, niore gtraii off
titau would be given off) at a loner temperature, the partial preaaure of tbe
oxygen in the Btin<a|ilMTc remaining iho oamo.
Moreover in the liloud ne bare to deal not with hicmoglobin in simple
wolution. in which tho moJecuIca are dispersed uniformly through the solvent,
but with the humoglobiu aegregated into minute isolated maaees, battled up
iin it were in the individual oorpuHcles. The hwrnof^obin of each corpuscle
is aeparatef) from ile fellows by a laver, ihin it may be but still a distinct
layer, of colnrleas, bwninglobinles^ ptasma. As the corpuscle makes its way
through live narrow eapillary paths of a pulraonar^,' alveolus, it \a s^karated
IVoHi l)ic air of Uie alvtvolua by a thin layer of plasma a^ well as by tbe film
of dm ooDJoiued capillaiy and alveobtr walU ; and a like layer of pi&tmia
aopantiot it from its Iclluws us it journeya iu company with them through the
466
RKSflRATION.
wider pawy of the arUrin aud veins. Through tlib layer
iibidi conMuttng no luemofclobtn ma holil oxyxeii in siiiipln iuhiUun nnlj,
Ibe oxygeu has to jtam on lU wfty t<> ■»•! froui the corpuade ; ud vverv cor-
|himIc may be ci>iiaidered as ^veniiuK. ns Car as oxy^n b oonvemed, n xone
uf i>lHSuia imiii^ilialely HurroundiBg iudf. Tbe corpusck uk** iu oxyg^ii
directly iVuin this taao hikI gire* u|> il« oxy^u ilireotly ki thi« xono; nod
the pKwiire at which at any momisit the oxygea cxhU in this lone will
dapend mi the prenumif osyKttni>uUu)i;the x(>lH^, in th airof thepulmonarj
nWeolun for iiistaiK-c. him) i>n lh<^ Kniallar or grvnter nmoui>l of oxygMi
«M()ciat«(l with the hn'miiglohin of the curj>u«flc
The film of tho c<u)j<)inc<l cnpillnn,' nnd alveolar wall u a thiD membrane
soaked with lym^h and wet ; tre caunot speak of it a* ac4uallr secretins a
li(]uid wrretiuii into th« alreoluit, for the cariiy of the alveolus is filled with
air which, though saturated with OHUsture, is air, not a liquid : still enoucfa
paw«s through the film to ke«p it oonUoually moist. Throui^h this film t&e
oxygen has to make its way in order to g,iua accem to the plasma and »o to
the corpiiselo; it makes lis war dissolved in the fluid, that is the lymph,
which keeps the film moisL This film, moreover, is comjxieed uf liviiiK
matter, and the con^derations which a little while twick li 3i:{) w« ur;^
concerning the difllbaion throufh a Uvint; membrane uf wjUiI »ubstancea in
solution, hold good also for the ditltuion of Rases in solutiim.
We have now to consider tlie queelion, Arc the conditions iu wbiob ha»DO<
globin and oxygen exist in ordinary venous bliMvl b.i it fluwn to tho lungs, of
such a kind that (he venous blond in pairing ihrongli tliv [iiilmonary captl-
Uit\(* will find the partial prwsure of the oxyg<-n in the pulmonary alveoli
.lufiicient to bring about the niMciciation of llic additionnl ijunntity of oxygm
whereby the venous is (x>nvertwl into arterial hlooil?
We mar say at i»nce that we haip-c, at prwent at all events, no witufnetory
evideoee Uiat (he film spoken of above exerts any influcnec, w a living film,
on tho entrance of oxygen from the alvcoliu into ih« hlood. Nor have we
any evidtmcc that u# a mere membrane or septum it exerts any such influ-
ence : (he oxygen appears to pass into the blood in the same way that it
would, if Ihe blood were freely exposed without any intervening partition (o
the alveolar air. Further, the eridenu'e. so fur as it g<Mw. seems to show that
blood absorbs oxy^n in the same way as an aqueous solution of hjeotoglobin
of Uie same coooentration ; the zone of plasma spoken of above as surround-
ing each corpuscle behares as far as regards the passage of oxygen to and
ftooB the corpuscles in no caaentially different respect from the way the mole-
cules of water, belonging to a molecule of dissolved hemoglobin, behave in
regard to the absorpliou or the giving-ulf of oxygen by an a<(ueau8 solution
of hiemoglobin,
li 394. In man, aa we have seen, expired air contains about 16 per cent, of
oxygen. The air in the pulmonary alveoli must contain lets than this, ainoo
lite expire<l air connisls of ti<bil air mixed by dilTuiuon with the stationary
air. How much lew it ei>o(nin« we do not exMitly know, hut prohuhlr (he
dilferenre is not very great. At the untinary atmunphenc premure qI 760
mm. Iti fter cent, is e<(uivalent to a purtinl prcMiure of 122 mm. The ques-
tion, (htri-fore, stands thus. Will venous hiooit. expiisecl at the teni[)erature
of the h»dy to n partial premnrc of liM than 122 mm. (less than Id per
cent.) of oxygen tiikc up sullicit^nt oxygen i fn>m 8 t« 12 vols, per cent.) to
convert it into arterial blood? Numerous cxpvnmvnis have )>«>pn mode
(chielly hut not exclnsively on the dog) to detonuinc on the one han<l the
oxygen- pressure of both arterial and venous blood (t. c, thi> parlinl prcssnre
of oxygen in an atmosphere exposed to which the nrtcrini blood neither
gives up nor takes in oxygen, and the same for venoti* hlcxKl), and on the
i
TUK ItESflRATORY CIIANQKS IN THE Ll'XGft.
I
■ not
other bwul tbi* beliiivlor nl tlio t«ni|>eralure nf itie budy or ni onliimry t«n)-
pomturai of UIoihI or of toltitioiu of hicptoelolrin (for the (wo o» we hiivo
jiiM snid, bdiiivv iti this respect very mtiuh nliko) towttn) on fiti)i<i«ph«rc ill
which the pitrtitil praBura of oxygea u made o viirv. Without going into
di-tnit, wv may state that these eixperimenta shmr thnt the nnrtial prt«ura of
oxygen in the Iiingv is amply sutHcienl tn bring about, at the tctnpcratura of
(ho body, lh« awociatioii of that nddilioiial amount of uxyceu hr which
WIWU8 blood beoomM arterial. When a solution of btemogiobin or when
blood is suoDearively expoced to increasing; oxyg«n pr«wura, as the partial
grcmiirv of oxygen is gradually increaned, the curve of absorption nsos at
rat very rapidly but aAeruard more slowly ; that is to say. the latter adili-
tione of oxyjten at the hijtber ])rewiirea are proportionately leas than the
rlier ones at the loner prewunw. And this is consonant with wliat appears
be Uie Imcl that tlie hwroo^loliin of arterial blood (bough nearly Batumted
itb oxygen, ■'. c, aajuciated with alinori. its full complement of oxygen, is
not quite caturated. When arterial blood is tlioroughly exposed to air it
lakea up rather more (ban 1 vol. |ier <«nt. of oxy^^en ; and that appean to
represent ihc. difference be1«eeii exjMving bloml to pure air, nuch an e:itenor
ouKht to enter ibe mouth in inspiration, and expiwin); Uloixl to llie air an It
exislHin the pulmonary alv<»ili. The greater rclnlive ulixorpliiin at the lower
preaturen linn a bvnelicial cflcct iniiNnmch uk it !>tin rx^riultx a (!iin»iil«nible
quantity of oxygv-n to I>e nlwrlMHl evm ulicn lliu [Hirliiil pr)rfi>iire ot'oxygco
in the air iu the luiigx is lureely reduced, as in aw^ndin^ to gmat hcighCi*.
Obwrvations made both witb dog's Mood and «xV blood worn to sbon that
arterial blood i-va^n to take up oxygen and begins to giro oil' oxygen ; in
otJM!rwor>U.lhnt dixnMocintion begins to take place ^rhen the partial prmHi re
of ihv oxygen in the atmosphere to which it is exposed sinks to n1>t>iit IHI mm.
of mercury- ; that is to sav, when the whole atmospheric prcMUre i» rwlticed
fnmi 764) mm. to about SUO ram., or when the percentage of oxygen in the
ntnt<«pherc is reduced by decidedly more than half And this accords with
the observation tliat in man, when the oxygen of inspired air is gradually
diminished without any other change in the air, symptoms of <lyBpncea do not
make their ap[iearance until the oxygen sinks to 10 per cent, in tlie inspired
air, and must therefore be l«a than this in the pulmonary alveoli. Wa
may remark that at ordinary altJtudee, eTeo taking mto auoouiit (be diminu-
tion the oxygen undergoeH befure it reacbea the pulmonarv alveoli, the partial
prenure of the oxygen in Ihe ntmosphere leaves a wide tnaruin of safety.
But at an attitude of 5o0i) metric (17,00ii feet), at which the preesure of
tlte wltole aLmcephere .itaiKln at almut the limit given above of :iOO mm., the
|,>lMrtud prewure of the oxygen will be Mich that tlie veuoUH blood mnnut
nke up the <|uantilviif oxygcJi pniper to convert it into arterial blood, Miice
at thiH limit iirlerial blixid Tit^iun to give off* oxvgen. \V<- may mid tlinl it
is at tbio altitude that breathing l>eoomce especially ditHcuit, but lo this we
idiall retuni.
§ 357. Tbo statements nuulc so far refer to ordinary breathing, but the
(|a<aiiou may be aj>kcd. What hap|)ei» when the renewal of ilie air in the
pnlmoimry alveoli ceases, as when the trachea is obstructed? In such a
caw tlH- oxygen in the alveoli is found to diminish mpidly, so that the
partial pressure of oxygen in them soon fnlls below the oxygen-preestire of
onlinary venous blood. Bui in such a case (he blood is no longer ordinary
vanoua blood ; instead of being roodentlely, it i;> largely and increasingly
leduced ; instead of containing a compsrativelv small amount, it cuntains A
llL^^ and gradually iDcrauing amount of rei^uced hwmoglohin. vVnd M
the redut^tiou eontinuce to iDcreoM. tlte oxTgen-pre^ure of the venous blood
also continues to decrease ; it thus keeps below (bat of the air in the lungs.
470
Hi-tipc. apparcnily. even ilie Ia*t trnt-ee of oxygea ia ttie Iudj^ ni«jr be takra
nj' by the bldotl and varrkil away tu ihe li«u«ii.
The Erll o/ Carbame Add.
4
th^.
obyj
USOCf
!f 356. It m-env imtiiriil to KUpmi^c ihni the carbonic acid would otoapr hv
(lifTiiftinii Irom the bloud of the Mveolar rapillari^ into llio oirol'lhc alveoli.
But ill order (hat diRbsion should thus lake place, the carlMMiic acid prcwuiv
of the air in the milmonarr alveoli niual always l>e leaa tliaii tltat of llio
venoiia blood of tnc piilnionary arlcry, and ouglit Dol to exceed tbnt of tbe
blood of tbe piilmonury vein. There are, however, many practical diffi-
culiies in tbe way of an exact detenniimiioD of iLe carbuuto acUi preamire
of the paltnooary alveoli (J'ot, though it muKt be greater than thai of tbe
expired air. it Is difficult to ^ny how much greater), and of the carbonic
nod jircftiiiro ••( the blond at the .tame time, ao an lo be in u p<i«iiiou to euoi-
iiatf the 'iiie with the other. In the ciu« of oxygeu thtrv u alwaya preaeut i
III the Uiii)^ii a iiiirjiUiH of the ffii*. n imrtion oiilv beinji altwrbed ai each |
brcaili : in the niife of carbonic ucid iIk* whole t|iiautily comei direct from J
the bliHid, and any mad ill cations in brcnthiiig ^crinui'ly allcci the amouBU^
given out. Tbtif, when the hri:«th i» held f-ir tuime lime the porociita^ oT^
carbonic ai-id in the einired air roach(« T or K |ier cent., but we Cftnnot t«k»^
ibi* H* a im-aiiure of tnc noniuil percentage of carbonic Bcid in th« |njl- j
monarv alveoli, *ince by the men; holding of the brentli ihc carlxtnio acid. \
in iW ^ilood. and kciicv in llic tiiilmonar; alveoli, is inrrca«ci) Iwyond th<
normal.
The difficulties of ihc problem seem, however, to have been overcome
an ingeniou« cx{>crimcnt in which there is iutrodnccd Into the broncbu
Ihe lun^ of a do^; n calheler, round irhich is arrantced a email bag ; by the
inflation of ihia baj; the bronchus, whenever dt»ired, oaii be completely
blocked up. TliUH, without any luarkcd dbturbauice of tlie^neral breathing,
and thcTcfuTe without any uiark«(l change in the normal profmnions of |]>«
gaM-K of the bhiod. the exMiiinebtcr is able to ulnp the inf;r«M i>f frmh air
ml» a limited puriiiin of tlic lung. At the >ame time ho ii enabled by nirant
of the catheter to withdraw a inniple of thi; air of the Mmc llmitwl jiortton
and by iiiialyKiii Ui dclcrmim* the nmouiit of cnrbonic acid which it i-ontaiiM,
or ill other wi^rd*. the partiiil prcwure of the carbonic aeid. Tbe blood
pMiine through the alveolar capillaricx nf thi» liiniied portion of the lung
naturally pomcisca the same carliouic acid prcwiirv as the rest of ibe vcnoui
blood riowing through the pnlmonar)- arierv— n prcnure which, though
rarying slightly from moment to moment, will maintain n normal avenisv.
On the supp'iKJiion that cArbonic acid pnsMS t»imply by dilTusion frooi the
pulmonary blood into the air of the alvivvli, because the carbonic acid
prcMtire of the latter Is nonnally lower than that of the former, one would
exjiect to find tluit the nir in the occluded portion 'of tlw luni; would oon-
linue to take up carbonic acid uuiil an er|uilibritim was establl-ned between
it anil Ibe cnroonic acid pressure of the venoua blood. Conseiiuently. if
after an occlusion, say of some miuuii-ii iby which time the djuilibrium
might fairly be axiumcd to have been i-atabliahed), the carbonic add presaute
of the air of the tKx-luded portion were deterniine<l, il on^hl to be found to
be eiiual to. and not nioro than eijutU to, the carlmnic acid prenure of tbo
venous bl(HHl of Ihe pulmonary artery. And this is the result which bm
been arrived at .- it liog been found ifjul the prrwun,-!! of tbe carbonic acid
of the occluded air and of the veuoui blixHl of the right side of the heart
arc junt about eijunl. Ilenco tbe evidence, bo far m It got*, it dinlim-tly in
ftvor of the view that the ettcape of carltouic acid tVom the bli>o«l into
TUK RESl-IRATOKY CI]A!IGE« IK TH£ TUSt'KS.
■171
pulnioDsrr alvool! ui (imply dut* l(i •liflUsIoti, ami tliut llivrc b do aCfid to
itvk fur atiy riirtlii,-r fxtilniiiiliiin. Tlit-re b, ii» far ua we cnn *ec ftt preaent
at tM rnnn. on iifcvmity, iiny more ihrni in the com of oxyjutn, U» MippnM
■tuU lb* wall of lh« piiimonitry iilvci>li liii> »ny niecilic micntory power of
itiarUrLJnt; catboiiir a('i<l Iniiii llir blimil indvpmnently of, nr iii iinlii^oDl'ini
to. iIm tnflgrno' of prcf»ur<i'. or ihat il rxcrts any uprcial influent M iill ait
■ €]iinn>oD H-ptiini.
*n>n arv midl- faL-te tihicli trtm to stiggnt that the exit of carbonic acid
'V^cam tba blood n usiMed by (hi- simiildini-ouB entrance of oxvgin. biit Uii*
W nrt il<-6niieK' proved. If Midi nn iii') is given, it is profmbly brought
^IxKit lij the c-liaoce in the hainioglnlnn in )«)Rio indirect nay rniciiig tbs
S*V'uKiiv of tbe caroooic neid in the bluad.
Jb far iheu aa can be seen at preiwtit, both the enlranoc of oxygen and
*~*^eeiil of earlxinic acid by which venous blood is converUd into artcriul
-*T llir simple physical results of ihe expoeure of the blood in ihe pol-
■ry capillary to tbe air of the pulmonary alveoli.
P
Tar. KEdPIRATORY ChaKOBB IK TBR Tli«DE8.
eSSS. In pnwing through (be several tiwues the arterial blood bvcomea
WKT DK>r» venon*. Tbe uxyha-nioKb'bin lieromes coniiderably reduced, a»d
■ qiMatiir of wrboiiie iiciil iwwm from tbe Umihs Into the blo»d. The
■aiennt nr chanice variea in the various- tii^uei, and in tbe WLHie timue may
TW^ at dilPeretit litnoii. Thua, in a gland at rvst, ii» we have sceii, tho v«ioits
blood is dark, (hewing that ihe lia-inoglohin is to a large extent in the
ndoetd ooadili^i ; when the glnml is active, the venoiia bluod in its color,
■sd in tbe extent to which the hienrnglobin is in the coodition of oxy-bicmo-
■lobia. fwcmblee closely arterial bloud. The blood therefore which iiBaca
Bnoi a gland at rest is more " veiions" thiin that from au aetive gUutd ;
ikoagb owinj; to the more rapid flow of blood which, as we saw in ou earlier
action, aeeompaniofl tbe activity of tbe );land, tlie total i{uantily of oxygen
taken up from and of carbonic acid di^charf^ed into the blood fmoi the glhud
in m given time roav be h-realor limn the latter The blmnl. on the other
luuid. whii-b cimea Aom on active, i. «., a controetiiig muscle, is, iu »pitc of
Ibe rooTv r«[4d flow, not only richer in carbonic ncid. but also, ifaoDgh noi
U> m corrofmnding amount, poorer in oxygen ilian the blood which flowa
fnini a miiacle at rest.
In alt these raira the great nuetition which conien np for mir conKtdc ration
ia ihi* : Doci tHo oxygen (mm troiu tlie UIoim] into thn liMnuii, anil d(K« tlic
uxi'lalion take jiliicr iii ibe tiwucs, giving rlM la carbonic acid, which passta
in turn away from ihr I'txatt* Into the blood T or do certain oxidiiable reduc-
ing ■utMlancen piiik> fmm llif li»iie< into the bliifHl,and there become oxidized
iniii rartHiiiic neiil and other pnHluct«,u> that the chief oxidation takes pluce
in iIm- bluiHi iwellT
Tbcre arn, it i* inie, nnlnnng oxidixable tiibstsnce* in tbe blood, but Iheae
■n sanUI in aroiHint, and the •pianlily of carbonic acid to which iboy g;ive
rin when the bkind conmining them is agitated witb air or oxygeu, b w
MBoll as scarcely to execeil (he errors of obaervatiun.
On tlie ntlter hand, it will be remenibeml that in Bpeakiog of nmacte. we
drew ailcntioo (§ I>1 1 to the fact that a fn)g's □iiiscle reinorea ttom the ixidy
(and the same is triieof the muscles of other animal-i) contains no frve
oxygen whatever; none can be obtained from it by tbe mercurial air-pump.
Yet MiHi a muM'te will not only when ut rest go on producing and ditcfaarging
a cennin >juaniity, but also when it contradt evolve n. very oondderublo
472
RESPIItATION.
quantJ^, or citrlMiiic uciil. Al<>n^)vcr, ilii» ilitdiiirge
go OD for II I'erluin tiini' in muH'lci' iiikUt rirniniftanon in urlucli il is iin-
poKfiblc for ihfin lo ohisin oxyKrii from wilboat. Oxrgcii, it is ini«, ia
nccwuMiry for iho life of llw mii^Rfc ; when venous iiieU-Aii of artwiid bKiwl
n Milt tbrmigh ilio blooilvceM;!* of a, itiufcIp, the irriuibility upcctlily <li«-
npiicnra, and unlcn fresh itxygi-n bo admin isicrt'd the muscle aoou ilia. T1h>
muscle nmv. however, liiiring the inicrvtil in which irrititbility i* still r>
tnineJ after the supply of oxygen has been cut olT, eontinuc ta contract
TigoroU!<lr. The supply of oxymo, tliough neecssary for the iiuii»i'-niiN« |
of irrilaliiHty, is Dot ncceemry for the man^e^ation of llial irriubility, is
Dot Dotweary for ihnt «sploBive 'lecomjKMition wliieh develonH n contrnc-
tion. A frog's tnuade will continue to coiitraet and to prouuce carbonic
acid ill an atinoepfaere of Iiydriigeii or oitro^eD, that is. in tb« total abaenvc
of fhse oxygen, both from iu«lf and frum th« medium in which it b
placed.
Thus, on tlie one hand, the muscle seems to have the property of takinirj
u|> and fixing in some way or other the oxygen to which it in exposed, <rfn
BtioriuK it u^ in iu own substance in suc-h a CDuditiuu thai it cannot be re-
moved by simple dimiuiflhcd prewure (so that the pre^ure of oxyKcn iu ih^k-
miMcular subttance may Ik- cuuiii'Iered as always nii), and yet )uu nou
entered into any diatinet conibiunlion which wu can speak of ta an oxida-
tion, but ia still available for such a purpose. The idea has beeii put fur—
~ ird that the oxygen iu Uiis comlilinu js physiealiy atladied lo and 1'
between llie muleouTes »f the muscular siitHiiunoe without tieing clirmicsillr
ownbined with tlieni, and henee haii bt-cn siiolcen of lut "inlra-molecular *
oxygen : but we linvc no exnet knowli;idgc lu to what its oondili'in rcnllv la
at this uliigc. On the other hand, the muMrnlar xiibslanoc is alwitvH under-
going n decomjitmilion of such a kind that cjirbunic acid is sut free, some-
timo, ns when llie mnsele ii nl rest, in small, oometimee, as during a con-
tractioo, in Inrge (piantitics. The oxygen preeeut in ihis earhnnie acid, ai
an oxiiiatioD product, enmee from the previously existing <^ti>re of wbicb w«
have just spukeu. The oxygen taken in by the muscle, whatever be ib
exact condition immediately upon \V entrance into the muscular subataooc,
in the phase which lias Ik-i'h cAJled "intra- molecular," sooner or later enten
into a cum bi nation. i>r perimps we should rather say, enters into a »«ri» of
combinatiouH. We have previously urged (§ :tO) tiiat all living aubatau<«
uiav be rc^ardfid a> iDoewaiitly uudergoiug changes of a dmible kind, changa
of building up and cluuigee of breaking down. In the en'I-pruduct« of liitt
breaking down, in the carbouio acid given out by mnsele, for insumce, we
can recognin; an oxidation product ; but we do not knuw exactly nt what
•cage or exactiv in wliat way the oxygen is ouiuhined with the oarbon. We
may imagine ibat the oxygen, as it coinai from tlie bloid, i* caught up so lo
atHUk by, and disap{»cnnt in, the building up proceMes (forming. poMibly at
toe very Ix^nning, witli noiuc convlitucnl of the mutteulnr siibstAuoe a
combination like to but firmer imd niiire Ktablc than its combination with
haiDloglobin^, and that through ihosi.t procenM^ it is made part of irouplex
dteonjKiubie subslanco nhwe di'^;ompo»ition ultimately gircs riw to the
carbonic nd<I : but, as far as aetutil knowledge goea. we caonol as vet trace
out the steps taken by the oxygen from the nioment it slips ftom llie blood
into the muscular suhst-tnee to the moment when it issues united with earboa
as carbonic acid. The whole mystery oi' lite li'« hidden ill the story of that
Iirogrci^, and for the present ve must be content with simply knowing the
iCKiimtng and the end.
Itut if the oxygen -pressure of the muscular Useue be thus alwaya nit,
oxygen will be alwaya paHiiig over from the blood corpusoles, in which it it
i
1
TlIK R88P1RATORY CDAKOES IX THE TlSSl'BS. 478
« I cnopvstiv«ljr high prcmre, ibraugh llie nliumn, through the <-npi1lar,r
valh, lbs lynph'>|MC«t, aixl the sHrcalcniDa, into the luufciiliir nutwtauco,
lalaitoap W il nrrivcH lbi>r« will hv iti ngim- mgiiiiirr or other bid<I«n nvray,
laiiag iho oxyifrn-pmgiiin; uf llii^ mutiulnr niitwiiirivc onoc more nU. Coti-
Kifiljr, the c»rliiiiiii' mciil produced \>y ihr liivwmpojiilioii of llio miin-iiliir
•Muee will u-inl Ii> riii»e llw eHrlxmic nci<l im^iiri' of the mmclr until it
mmit that of (Ih- hloml; whereupon carbuiiic ucii) will ]ia» frtini tlw
ill Into the bluot], iu i^nev in iIh- miiBcuiiir eub»tjince being siipplieil l>r
i]f gtncntcd luppliea. There will ttliin)^! in fnri bo a nlrcfttn ofoxygm
I llw blooH to the diumIo uid of carbuiiic ncid from the muscle to lh«
UonL The r<«|iimlM)0 of the muscle then doca not consiM in tlirowini' into
iW Uoo<) oxiiiixable eubetincee, there to be oxidized into cnrbonic nciJ and
^ilT nuiu-n; but it does consisl in thonHumption and sloriiif; iipaf uxTgMi
^OMhim or other in its anbetatice. in the btiildini; up by help of thai '>xTge»
^*xploMve deoomCMMRbie substances, and in llie cnrrvin^; out nt' deeompjei-
**'*oaa whereby enrbontc acid and other matters itre cliscliarKed Hrat iuto the M
"'"'iMlce of tite miiKle and subaettuentiy iuto the b!ii>>d. f
^^ 360. Our knowlei))-e of the n-xpitaiory e)iau)>ea iu muscle is more com*
*!**'<• than in the oase ofany other tissue ; but we have no reason to suppoeo
''** the pbeoomeDK of niuwle are exc-eplioual. On (lie coolrarv. all the
■ ^'nilable evidenee gctea l<i »h>iu ihut iu all linun the oxidation talcee pUee
l'' the tissue, and n»i in the n<ljoiuiii;c blood. It is a remarkable fact Ibat
^■»l>h. serous Huitb. bile, urine, ami milk cootaiu a mere truce of free or
*****ly eonibineii oxyiteD, but a very cousiilurable ijuantity of carbonic acid.
, ^Url vre may prvbably asMit with Mifety nilli rejmrd to all the liaaues that
." ibc tiviKi ihemaelvea, in the Ivniph which bauies their lymph^pacee, and ■
*» iho **ofcUoni whicli Mime of lliem |M>ur forth free oxygL-n in eitlwr wbollv ^
*bicnt or M tnuity that their oxygen -prcMu re nwy be ngnnlti) m mU, whife
^Krbonio aidd is to abundant that the prenura of carbonic acid in them may ■
(■■ rvgsnlad as exorciling that of venous blood. An excciilion seems to b« fl
piiimiiliil by the case of the lymph floving alon^j the lar^r lymphatic
Vviaala. for in Ibi* the amnuni of carbonic acid, while usually bicher than
Uutt of arterial blwd, if lower than that of the general venous blood ; but
tbia tirobably la doe to tlic fact iliat ihe lympn in ita pannce onirard is
lAfl^erly CI |viec<] to arterial blood in the conneciivetisimci and in the lymphatic
^laui'i^, where llie production of carbonic acid is slight as compared to that
KoioK on in muscles. All llie facts poini to the conclusion that It is the
CliatMa. awl not the blood, which become primarily loaded wilh carlmnic
■»H<1, the latter simplv receivinj; the saa IVon the former by diffiision, except
^^^hp ([imbnbly) small ouantity which results from tlie metabolism of the
^^^Kod ooqtusolei; and that the oxy;-en which paassa from the blood into the
^^Hbmm i* at aaee taken up and placed under such conditions that it is no
^^l0M>f remorable by diminished preniiie. ■
^^ Id Atrther Mipporl of thL< view may be urged the fact that if, in a tng. '
the whote blood of the IhhIv tx- replai.-«d by normal saline sohilion. the total
metabuliaiD of the body i.i lor >omt- lime unchau^^. The saline metlium is
able, owing to tlic low rale of wrtaholi»m and large (cutaneous) renpiraior^-
wrfioe of the nuimiil, to Mijiply the linmet with all the oxygen tliev need.
and lo remove all thr carbonir acid tliey pn>iluc«. It i* difficult to believe
that, in such an exixrimi-nt, the oxidation took place in the saline solution
ittcif while circulating in th<' bliMHlm-vIs and tiM(ue>«paoea of tlie animal
We mav add that ihc oxidative ponvr which the btoml itself removed
bom the &ody is able to exert »u ■u)>c<tnnco' whii'h arc uiidoublciUy oxidixed
h Ihe body la M small tliat il may be neglcclcl in the pnvt-nl omiidera-
I llgwai If grapxcgiu' lie addol lo blood or lo a solution of haemogloblu, the ^
1
474
KBSl'IKATION.
tnixtitre taty bo k«Di for s \oajt time ni the umperaiure of tli« body with-
oui uiMktgoing axtclnlioD. F.v«d within i)i« body a xli^ht excea* nf sujiar
in Uie Uowl over a eeriiiiii (wrcvnta^ wholly vnc«|)m i>xi{tnltiiD. nud is d»-
ohorged un«h«ngwi. Miinr ciutty oxiiliiutl xilwrtunccs «uch w pyrDjpdlw
Mid, )MM InraoJy througb the blood i>r n living biMly nnd mn dbcMrgM.in
the iiriiii^ witliotit being oxKliwd ; thniiEli |>rri)niHi in ■nnir of Umm CMH^
wluU ii[>iicnr8 lo he iin nbMOM of oxidaUion is rr-olly nn niidiilHUi fbUowwl
by m mibwijueDt ixjuivulcnl mluctioD Uiking |>i«oi- in thi? urin« or elMwbsre.
Inc oreanic Mciil«, such ttt ritric. ev«n in comhinntiiti) witb HlkaHtut bakw.
■n odIV fxinially oxidixed; when lulininHWnNl m Mcids, ud tuA of mlts,
Ui«y iirt- h«r<lly oxidixod U hII. It w, of course, quite poasiblo thnt the
chnn^ra wliirh tb^ blood undergom when shed might interfere with its
oxidative si-tion. and bonce the fact that shed blood has liltle or no oxidizing
pow«r, is not a satisfactorr proof that th« unchanf^ blood within [ho living
TCOeJs may not hnre eiicb a power. Itut did oiidatiuD lake place largely
in the blood itaelf, one would ex|)e<ct even highly diflbsible siilwtanccs to tit
oxidized in tbear trauHt; whereas if we suppose the oxidat>i>n to tak« pbuM
iu ih« tissues, it keeonies int«tligible why such divisible siibsiano» as tboae
which the tiwuw in gei>eral refuse to take up largely should readily \»ea
unchaii)^ from ihe blood ihruuf;b the exeretinK orpuia.
We have seen that in niustile tl>e ]>roduc(i<in if?carlMnic acid ia nol directly
deiiendent on the co4isuuiptiuu of oxygen. The uiii«-le [jnMluces carbuntc
acid in as atmosphere of bydrugvn. What is true of muHcle is tnie aUo of
otlter ti»u«s and of the body at huge. It wan »hown lung ago that aoiniata
might coniiuuc to hnMthe out carbonic acid in an atmosphere of nilrogCD
ur bydriijccn ; and thin liiui more recently been illuNlmlM) by thi- n'mHrkal)l«
exjicrinK-Ml thai ii fr<i^ kept at a low lemjirratiiro will live for wcvcral hourn,
and i'unlihu« li> produce carlKuiic iK'id, in an iitmii*pbor« alBiilulely I'rm
from oxyg«R. Tim nirlx>nic lu'itl pro<liii-i-«l ilnring ihtit ixritxl was made hy
bi'lp of tbo oxygen io«pircil in tiK- Wiir* anterior to the cmmmciMomenl of
the pxin-rtmeiil. T\i« oxygen then iilwirKni was Ntowed away frtMU tb«
liii'iuuglohin into Uk- tiwuics, it was made umsof to build up the explosive
oomjMiundii, wltow cxphwions later on gare riw to the carbonic arid. Or, to
adopt a simila which ha:> be«n euggmted, the oxygen helps to wind up the
rilnt cluck ; but once wound up the clock will go on for a period without
t^irther winding. The frog will continue to live, to move, to produce car-
txmifr acid for a while without any fresh oxygen, as we know of old it will
witlMMit nny (rvfih f<iod; it will continue to do i>o till the explosive oom-
pounds which lliv oxygen built upari- exhausted ; it will goon until the vital
clock hii* run donn.
!i 361. To Mini up, then, the results of reapiration in its elieaitca) aspects.
As the blood passes through the lungs, the low oxygen-pni^sure of the venoua
blood permits Ibeeotraiiceof oxygen finm the air of the oulmonarv alveolus,
through the thin alveolar walls, through th^ ihiu i-npillary sheath, tlirough
the ibin layer of blood-plasma Co iho red cor]>UM-le, and tlw rei)uci-d hietno*
globin of the venoua liloud becomes wholly, or all hui wholly, oxy-hsDino-
^l<d>in. Hurried to the iLnsuee, tlie oxygen, at comparaliiv/y high prvwurt
to the arterial hiMHl, pustm lurgely into tbi^m. In the tHsucii the oxyg«ti-
preanure u always kept at an cxce«<linglv low pitch by ilic fact that thvy, tu
aome way at prateni unknown tu uh, |i«c1c away at every monK-nl into some
Mible combination each moh>i'ule of oxygen which they receive (rum the
blood. With its oxvhienirwlobin largely out not wholly rvducrd, the blood
SiMtm on tw vcnotit blond. To what extent the hicmoglohin is ntluccd will
lepeiid on the activity of tl>e twsue i(*clf. The ij^iiantity of hicmoglobin in
tba blood ia the in«asun) of limit of the uxidixing power of the body at
T«K SKRVOCil MKCHASISU Of KEdP[ RATIO^f .
475
larfTO ; 1>iil within tbal Umic the amotiDt of oxidation is ddcnitinod tty the
ttMU«, aim] bf th« liaeue ulooe.
Wo vAiinot irtoe the oxyBcn thraucU iia sojourn io the tisMio. Wo only
know ihM sooner or Ut«r il comee nack oomhined in carbonic x^id v«>id
other matten not now under consideration). Owinn to iho continmil prii-
duction of curbonic acid, ihf pn-vsure of that jim in the extrii-Tiwoular
eleincnU of the liaeue it nlwaya hi;;ht^r than that in the blood: the |^
■coordini^lir patse* from ilie tiwne inl« thti hUmd. and the vctiou* hli^id
piuwa OD not only nilh it« hictuuiilobin num or tnw reduce*], ('. e., with it«
oxy(,'eu-pr«eauredecraa«<t, but ai.iii witli itH carbonic acid prenure increiisccl.
Arrired at the lungs, th« hloixl liiidtt ihi: pulmonary air at a lower carbonic
Mid prenure than iUelf. Tlic gM acwrdiiiKily ittream* throuEh the thin
va«cul8r aiid aiveoUr wallii until ihn pruMure wiihmit the hloodveseel is
equal to tlie )>r«aMnre wilhiti. At the wme lime the biixid liiidi in the air of
the jiulmonarj Blv(n>li a ttupply of cixvgc^n, more ihaii adc(iuali> to convert.
i>ot entirvlr but nitarly wi, the rediiiTil Wmuglohin hnck again to oxy-hieino-
ghibin. IhuH the air of ihti piilniotiary atvc«li. havinz given up oxyKen to
the i>looi) ainl taken up enrlKtnio acid iVum the blood, having id conwqueuue
a hi^licr carbonic wid prewitre and a lower oxygen prcMurc than the liilal
air in the bronchia) piuettccs, mixes rapidly with this by ililfu»ion. The
mixtiin.- in further iu»i)>ted by ascending and desoaiding currvnta; and the
tidal air inaues from the ebott at the breathing out poorer in oxygen and
richer in carbonic acid than the tidal air which entered at the breathing in.
Tuc Ni^ivovH Mkchahisu op KlSPIItATIOrr.
i 36S. Brenihiog is an iovolunlary act. Though the diaphragm and all
the otiier inu«oIe<i employed in respiratiivn are vofuutary muscles, i. «., mus-
cles which can l>e called into action by a direct eflurt of the will, and tJiough
respiration may be modiiled within very wiile limits by the will, yei wo
habitually broithe without the inierventiun »f the will ; ttie normal breath-
ing may ooiiiinue, not only in the aWnoe of oiHuciiuimmt. but even at\er
the removal of all lite psru of the brain above th« medulla 'ihlongaUL
We have nlieady seeii bow oompliealcd is even a ximple renpiralory aoL
A very Urge number of muHcIes are called into pluy. Many of tlutMi are
very far aimrt Ihitii Mch other, such as the diaphragni ami the iia.'otl oiuHcIea ;
y*t they act in hartDonious sequence in point of time. If Uie lower iiiter-
coMal muscles contniet«d bcforu the scaleni, or if the dia|>liragiii contracted
alternately with the other <■batt-mu.■^:]eK, tli<: ntiffiictory entrance and exit of
air A'ould let imptMiiblc. T)i««i muscle, moreover, are CMirdiniiled iiIho in
iwpeet of tbo amount of tlu-tr Mronl contractions ; a gentle and 'irdinary
ooBlractiun of the diaphragm is accompanied by gentJe and onlinarr otui-
Iractiotia of the intercostals, and them are preceded by gentle and ordinary
Cnnlnictions »f thi- scalcui. A forcible contraction of the scaleni, followed
by simply a gentle contraction of the iutercoetals, would jwrhape hindor
rather than awist inspiration, and at all events would be waste of jNiwer.
Further, the wlwlc complex inspiratory ellbrt is often followed by a Ins
marked hut still complex expiratory action. It is impowible thai all tlieea
so carefully coordinated muscular cmniractiona should be brought about in any
Other way. than by coordinate nervous impuUcs deiicending along elfervut
narvca from a coordinating nervous centre. I)y experiment we find ibis to
Ijc tlie case.
When in a rabbit the trunk of a phrenic tierre ia cut. the diaphragm oa
(hat side remains motionlea<, and respiraliun goes on without it. When both
m
RBSPIRATIOS.
cut, tlie whole diaphr
quiewuni, though th« coctal
nervre
rcApinitioii becomes «xcemivcly Itkbored.
Wlicn Ml iDier(xi<lal nerve is out, do active rwpirnlory nioviMiM>Dt« an
«Mn in the iutercoetnl muscles of the correBpoixliiiK epnoo. nnd when the
spinal cord is dirided below the ori^'in of the seveath i-ervical spinnl nerve,
lltat is. below the exits of the roots of the plireoio uerves. coetnl respinttion
ceases, thou>:h the diaphrasm eootinueB to a«t, and thai witli inc-reased vij^or.
When lite cord is divided juet below the inedulla. all thoracic movemeDtB
cease, but the respiratory actions of the ncwlriU hikI );loiiis Mill couttooe.
These, however, disappear wheo the fadal aiid recurreut laryii^ceal uerres
are divided. We have already slated lliat afU-r rviuoval of tbe Itraiu nhox'*
the m^ulla, respiration still cootinuea vtry much as usual, the iU'>iiifii-A liana
which 4'j)9iie I'roni the km of the brain being tin essential. Ileiice. |tiiiiing
all these facts together, it is clear that thu nnpiraturr iiinvcnitrDts nrt', iu> w*
suggested, brought alwui by oocirdinated impnlsnt which, devdopitl iu tha
oenual nervous systein and ittarliu)i in tbi- Rnl inHlanuu in tlu! nmlullii, find
their way along the several efllsruot nerve*. The proof is coinpletMl by ttio
fact that the reniovnl of, or exUmiiivi' injury t^i, iIm^ nicdiillu aloiio is, Mvi- in
excejtiioiinl cast* which we will discuw presently, at oiic« followed by the
CMHlion of all reepimtof)' moveoHuit*, even though the rest of liie iiorvou»
nrstem including «very muscle nnd ercr^' nerve cuncemed be Utt intact.
Nay more, if only a smalt portion of the medulla, a tract wlii»e limitu have
not been clearly defined, but which may be described as Iviug below the
vasomotor centre in the imtncdiate Deij^iKiriiood of the nuclei of the vagtis
nerves, be removed or injured, rcdpiralioii tvAHcs, nnd di-iith »t once ensoee,
Honoe thia portion of the nervous system km called by Fluurens the vital
knot, or )^t;Iion of life, na-iid vital. We ehalt »jieah of it as the rc4p*mtory
Wttlre,
<} 363. The uaiure of this centra roust he e-^ceedini^ly complex ; for while
even in ordinary tespiraiion it given rise lo a whole group of cioordinale ner-
reus impulses of inspinitiou foiloncd iu duese<|ueace by a smaller but »till
coSrdinale ^roup of expiratory imnulses of an antagonistic nature, iu Labored
re^iratiun fresh aud laiyer iinpiilsea are generated, tliougli still in eoordina-
tion with the normal ones, the expiratory evenU beinn eapedally aui^mented ;
and in the caaw of more extreme dyspnea and asphyxia impulses overflow,
mi to Hpeuk, from it in alt diredioiu, thoush only gradually looung their
coiinii nation, until almuttt every muscle in theltody is thrown into contratitioiu.
We must not, Itowever, ci>iievive ul' thin oentrc a* one of sui-ti a kind that
the impulses leave it fully r(M)rdinaU^l bimI (vjuippul no ihal n^ithiog rcmnins
for tlieiu hut to travel, unchanged, alontt thir m'vitiiI cftiveiil nerve-fibrc:' to
thfir Mvcral niuwculnr destinations. On ibv (xuiliary, we have reason to
think ihat the nttpiratnriF- motor nerves, like other niirtor nv-rvw, are con-
ni;cU-(l juKt US they an? about to tnoe from the spinnl eord, uilh a nitrvous
machinrrA'. in which mtrve eelU play a pnrt — a point which we shall consider
mori.' fully in trcuting of the spinal ctinj : we have reason Ui think that the
r^T-pinil->ry impulsw stttrting from the respiratory' vcntro paw into and are
iiioililit^il by Mecon<larv spinal nervous mi.'<chanisms beforo they issue along the
riiNi.rr nerve rootA. imteod, observatioiM show that under particular oondi-
lioDH, and espc-ialjy in young animals, respiratory movemenis may be carried
out in the entire aliseiieo of the medulla oblongatA. Thus if in n kitlen or
puppy, or young rabbit, after divitiion of the spiaal cord below the medulla,
artindal respiration he kept Up, and then pauses be made in the ariilicial
respiration, during these jiauses uol only may what appeer to be respiratory
movements be intniced in a reflex manner, by pinching or by blowinj; on the
skin, but, esjwdalty if tlie excitability of the spinal cord be hdghtened by
TRX XKRVOUS MKCHANI&U or RKXI- 1 RATIo:f .
477
I
I
I
I
mII iattt ot Mryehnine. own »|K>iitiin<H>iis c(l<>rW of brcutliiii^ tiiat occa>
itmiflT bftobMrrod. Thnoare tlicxcvjiiional imtmioe^ nii-ii Honed nbovo.
Km a ncfa nun the rhvlhmicnilv rvpenlc<l movomciite of llio respinitory
■mtIm arv •Oiiivtinx-e ncconiptiniitf by rtiytltniic movcmcnle of the fore soil
inl tiaiba not rMpimton- in natiiro, it mttv be doubled wbetber tbeae
'VptrilMnts rwlly prove tW exb>tcn<« of dUtincC reepirator^ centres ia the
ifml rOrd: ind at moel lh«y merely show that the respiratory nervous
■Ktiabm tt DuC enlirvty contined, as was oace thoki{;hl, to the c«>»tre in tbe
•Hblla, but also eoibracea other Huhiidiarv mechanisniB, which may perhnpa
b ipakcD of aa centre*, in the spinal coni beloir. It has. iudeed. bt«u miiiu>
liiaad by eom« that thme lower aptuul centra are the chief ueotres and tliat
licinedtiUary c«mre acta merely in ibe way of ref[ulatiu); th«iie; but U ditK-
rait to leoMicile ibta view with llie eapericaoe thai inli^rlert^d with the
""lulla, limited eulirely i<> the medulla, ao ofton IcxU ti> the entire alxjli-
"m oftbe i«e|)iratorir move men m. This motier is not al prcacnt thoroiifthly
*vrkc>ii otit, but we shall probably not greatly err iii regarduig the rrnpira-
^^fy iwrroiua system as in many waytt analogoua U> tlie Toaomottr m-r^'oua
*?M«n, wiili its bead centre iu the incilulla, and secondary centres eluowlifrc,
*)H in c<naiin(iin|{ Iu «peak of tbe ci'uire in the nu-dulla n» iM-ing " the nwpira-
^rycwnirc" nhilL- nilmiltiiig tluit it works lbri>u^h otbiT nervniix machinery
l*Wcwi lower down in tlie «piiud curd, ami that thin iiuliorliuaU' ninchinory
*^r in Dxci'ptional ranw, carrj' iwt, though iuadcc|iialely, the work of the
*iim omtn.
t 964. Adinilling then the existence of tliis medullarv rmpiratory ccatr»
till iititatioo uainrally arises, Are we t« reeard its rhytnmic action as dua
l^mitially to changw Inking place in itMlf, or as due to alTerent nervous
itaiMilBn ur othvrtlimuli which nHb<a it in a rhythmic tnaniior from without T
in otber words. Is the action of the centre automatic or purely rcficx? We
know tbai ll>e centre may be inHuenced by impulses proceeding from without,
■ad that ll>e hnAlhin;; mav be ellecled by ilie action of the will, or by aa
amotioii, iir by u d&*h of cold water on tbe skin, or in a hundred otiter wa^;
hilt tbe &ct that tbe acliuD of the centre may be thiiH modified fn>m with-
out, is uo proof that the continuance of its aL-tivity is dependent on exirinaio
In atiemptinf; to decide tbb question we uattitvUy turn to tbe {Mieumo-
t— *'if' <" °^°g t^^ nerve most likely to serve as the channel of aJTereut
laapmlaes setting in action the respiratory centre. If b»tli vaKiis uervea be
diviiln). ra[iinttiuii >till oontinuea, thou^lt ia a modified furm. Thin provca
(UrtiiicUy that alfereut impulses ascending those narve^ are not the ■■tBcient
CMHtof tli« renurator}- movementa. Wo have ween that irhea the spinal
«acd is divided b«low the medulla, the facial and laryngeal movement* Mill
continue. This provts thai the re»|iiratory oenire h »till in action, though
its activity is tmable to manifot itielf in ikny thorado movement. Rut wlHin
the c«rd is thus dividcal, the respiratory centre in cut otT from all sensory
impalsci, wre th<»« which may put* inli^ it from the cranial imfvcs of tua-
tarf AiMtioa : and that these tenixiry cranial nervea are nut speciallv oon-
ccmed in drvdojMng the activity of the respiratory centre w shown {»y Ibe
foct that the division of tlMSC cranial nerves by thsoiMlTas, when thentGdiiUa
and spinal cord are lell intact, <lne( not do away with tlM: ccintinuniic« of
respiration. One erauiid nerve, lis we nhall Mm, iit winxially iimk^rrn-d in
rtapimtiun, vti.. the vagus iwf%'e: hut if alli^r r^-ni'ival of the brnio above
tbe medulla both vagus nerve* he dividiil. rat|ufation :>till goes on ; iudo«l,
tbe Kspiratory impulses proceeding from tbe ceulre are, though in a ])eculiar
way, exaggerated. Hence, though we cnanol put tbo matter to an cxperi-
ntal IMl by dividing every sensory nerve in the body, while leaving the
478
nifsfinATtoN.
iDDlor Derv«e of rM pi ration iniacl, eucli nn op^initioa beinj; praetimlly iinM^
dble, ire may infer tlint the respiitilun' impulnee proce«liii)» from the reeFHra-
lory centre are not sininly atlereat impuUvd reaching the centre aluiiK a*rerent
uorree aud trans formen! by letlex action in tlml ceuire. They evidently ttlan
dflnovoAvm tJM centre it^ir, liaweTermuch tiieir chuaoMrs maybe aflbcted
by adfercnt impulseit. reai-'tiing tlial centre al tli« time of their heing geaeT*
•ted. llie action of llie centre is automatic, not simply reflex.
S 369. We find do tu(|uiry that tlie activity of the centre is priifoundly
intUifiH-«^ by two clawee of evmtA. Tb<9e, as we mijftil exnent, are, on the
one liuiid, cventA |>rtM]tidn>; ohan|i:(« in the 'lualily of the blood distributed
t<i the mi-<lulla bv the lull rtnlricle, uipecially ua re^anU its frasek, U>at ia to
Mv, evGiiu niixliA-tMx the lutcnrhant:!' inking place in the liinc:*: and an the
Mner hniid, nervouK Jiiiptjiim »tarii.-<l in viiriciuM wny* und n-iu'liing th<' cctntre
Along varl<Hi» uervoi ur nervous tract*. U will bo convenient to cnusidcr the
AA'crent iwrvnus ImnulHCM may alftct t1i« centre in many varioua wmy*.
Tbi^ It hole net of bn-mliiii); '>r of taking n breath iii n double act, oonaistutg
of an iiiipiratiou and an cxpinition. and tier\'oun impu!*** may MpeciAlly
aff«-ct the <hh: or the nthcr. One mode of brvalhing inaydiHTerfnim another
in ihv depth nl' the individual breath, in the volume of nir taken id and
givfin ont; and norvoiii> imfmlMa mav incrtMae or may diminish the depth
of a breath, the voliinic of air tctpired. One mode of breathing again dtfmv
fVom another in Ihi: rapi<litvnith which one breath suoceeda another, that la,
in the ntc of rhythm ; and nervous impulses may slow or may quickt^n tb«
rate of rhythm. Then, aj^in, combinalioD of edects mi numerous and varied
as almost to baffle description may result from the inffuonoe of various ner-
vous intpulses. Emotions inav aflfect a sin^e breath or a long senes of breaths,
may (juicken the rbvthm while making each breath more shallow, or tttMj
at the same time ma^e each breath deeper, or may sloiv the rhythm in eJth«r
Ok one or the other manner, and may liear chiefly on Inspiration or oo
expiration. Moreover, there is not an nflereni nerve in tJie nody nhicti by
means of a fle real impulses pacing alone it may not be the iuatrumout of
influencing the re§pit«tiiry centre. Of all the automatic ceiitm in the body
tlie reaptratory centre is the one wh«!»e Independence is moHt olucurcd by the
the repeated alTcctti of aflerent mirvous impubee.
ru, tm
KmNT on HK^NAtiiMt OF Swmeat or nm Vmvt.
Tlw nKU* WH dlvldoi) dI Ibc folDI fntrkpil i, Thr van* hm iitiUliinl tij imnii* of ■ Uin)»(tf~
toiuwM lAilt « noflrar IdIo vIMli tlw ■nlmkl (nliblli ortBliial. « aboun in FIk 133, Ihs IcTSi
lalUlwtnlnipl'*!''"'** «t> tt aurlMl qM lit iha tainliour uul rtilii|| IB «i|>lmli>ti*(lhui>lr faliini*
IfwpiBtlMi IxcUuBi ■ unlen^al k tCittnUon Iwclni M b uidaidiat o Tha Isiwgniiliiklly
klb tntintu c N&d < owIbc in Ibo «*Mp« of ■!> (Mai Ito ■wwlw.
TIIK NKHrorS IIECIIANISU Or REsrtKATIO.V.
479
ftnain Rfllcniit iwrv**, however, npix-nr w be more clwely cmiinwlwl willi
il ibn Mbm : ud<1 of l)ii»C titc Ri'«t ivmHinciinitH iiinl im|H>rUiut iirc tlic two
ni|u* nrrvCB. irhu'h nv hitvi- iilrcnilv nu'iilimii'il in llii* niiiiK-ctioii. Tli^ir
bapMlUMO M Wi-ll illunlralwl brthi' fnllnwiitlg c\|HTini<'tlI*: (f onO VIl^llH Iw
dividtd is ui oMtDarv WDV, wiihoui nriv <|>i'i:'inl {inrmiilion!'. ibi; ri»|)initii>ii
(■ dlW not inBltrri«llv cDODgrd, or it jll.i mI lutMinw nlowt-r >Fig. 139).
If hn\t be ilivid«l i.Fig. IJOj it lici.nin- viry >!•>«,■. tin- pHUHm lictwi-on
expindon Hml iofpinilioii bt-iog mnrkrdiv pmlongiiL T)ic charni-irr i^f the
'«>DtrEUir]r riov«i»«iii, inn, it nmrkvill}' cbatigtsi ; vnvU n-Kpirniicin i» fuller
*ih) drcptr, so much m. iiKlwd. Hint, iicrordiiig to Mint- olMTvrn.wItnt Ia lott
KtTtrt on ltBaiiiiTii» "f liutios -f Burn Vir.n NiiMTn.
Tk« con* »M (MiliMl lallitwiiuii>v>*rie. I». TlwwcaiultiiinwtKrtoHvdlTMcdMK.
*■> rate is fitted iu extent, llicamoiint of carbonic acid produced and oxvgt-n
OaoMiRiMl in A given [icri<id n-niiiining afitr divisiou or the nerve* iibou't the
*mn>n as wbeii titwe were iiitacl ; bul it is undesimble to iiiaiu too mnch nn
Mm •Hctnen of thu cumpmmion.
FtO. ML
■* Usnunw »T OBnu »nin'UTiMi up t>s CarraiL Ink or nil V»m-» Ta«wi.
««• aMaliiM la Uk mow war m Flvk IK l«. SUmNlMlM tt tlw tMU* ta«M at k
•III] dufad at J .
WlicD Kftcr divulpD of liuth vogu* ncrree id tbe itcck, llie iDfldiilln b«iog
^intact, tbe contnd flump, thiit ooun«cted with the cenirnl iktvoos fvslem, of
■ of them is slimulntwl witli K gMit)« interniutel i.-iirreni, the I'lWta «re
■IwMfB tbe Mtne; oDtt of two rfiMilte may fnllow. mid tbat whichever of
BtSPlRATlOS-;
the Inn n^rres be used. In a ceniiiii nuuiber of com;*, and theae inntr. pet'
Imiw, lie TWftnrdMl as the more topical one*, the rL-a(itrutt(ni. vrhich from the
i)ivi»itiii if ilie nervca hud become slow, U ((Utckfued Uffaio < Ft|;. 141), aod
with i,^are, by u projier upplioatiou of the *tinmlu«, the nunnul reiiipintor]r~<
rhj-thin ruuv for tt time be retlored. Upon ihe oesRatioti of the »timulu.-> th
slower rhytnm returnii. If the current lie iiicretued in «tr<!n^h, ibc rliyth
may in smaiK cime* bn (o ■'.'twlerutt^l ibnt inn pi ml ton buginH Itcl'uri; thv espini'
tiuii of thi; prcciiliiig brentb ia completed (Fig. H2), «nd tbin lanj go
rio. Hi
^
7L
/
eimiLlTKll Ol' V*lll» iJADIN-i T.) 1 Vll'IIUlllBV IHiH«*«,
Thlicntvr, nnllko Uiv luncedlof. Hwobtaltitd Iif InwllUE a nmlleUiKni(li tha bodj «all. «■
UltBLan (he itUi'hnMrn. >iiil nlliu'liiiiE ■ lei«r lo Ibe iiL-cille. i9cc)ISi i Tlia l«nr tliei wItliMCk
ConirullQD of ihc dlitpUracni. •<> 'liu' liKlilrslltiii bv^iu nl ■ kiid and* >l b, ciplnaoB bqiJna il b
•nd null ai o, Uio Interval bclman c &ud ■ eormpnndltiii to th« |i4uc. SUmulaUnn of tba nm
bcslniat X. It will b«w(Dttvu upon tilmu1iiUoiiIhDliifpli«iiiiTrtK*ortbp|«r«rteKtBloacli«IH<g
ilMpraealluit aplimtlotUBnconipleie.
iinlil at last the diaphmgiii is brought into a coDdition of prolonged tciAUot,
■nd a standstill of respiratioii io nii exlreme iii«pirnlory pb&ae is tbe rcniU.
Oh the other bund, in a ccrlnin number of ciml's the result it of an opuo*it«
character. Even though the rcepinilion l>e already slowed by divi«[oa of
the nerves, etimulatiou produccA a still further slowiiij,', tlie paUM<« beCwevo
cacb cxpinitiou and the succeeding inspiration are prolonged i ef. Pig. 14-^).
and in a certain number of cases actual standstill is brought about, but a
Btaadslill of a kind the opposita of the one just desoribM, tluce tho dU-
phragni which in that case was in prolonged l«tanus is in this aue conipletaly
relaxed and remains for some time in the condition in which it is at the don
of an ordinary- breath. In a cerlain number of cases, and ihoM arv not
uncommon, tti'c nwuli is intermeiliate between the two above extremes : the
dinphracm stands ntill in a prolonged contraction in a position which is
intermeilialc bewecn the height of inapiralion and expiration.
T)m« ranltA nuggtvt the conclusion that the thbus norro (we iir« dealinu
now with the main trunk nf the nerve) contains n^rent fibres of two kintb
oonnectcd with (he nvpiraKir)' centre; one kind augmenting the action of
the ccntrv somewhat iu tbt; .<aniu way as tlie augincnlor cardiac fibres au)^-
racni. the beat of ihti heart, and ihe other kind hnving nn inhibitory c&ect.
Apparently sometimc-i the one and sometime* thi- other kind is. according to
circumstances, moe>t provokol by the stimulation, much as in the same way
as stimnlstion of (he vagus in the frog, which, as we have seen, § 158, is
the channel for both inhibitory and augmenCor cardiac impulses, produces,
sometimes inhibition, •oitietimos augmentation of (be heart beat. To affect
TOE NKRVOta UKCUAXISM OF KBSPIBaTIOX.
481
ikthMTt of course the Mlniitstion of the vakils raunt be centrilbffal.difected
ImHlbe |ien]>liery, wImt^ab to ailed the res pi rat ion it nmat be oeotri-
fial, tpslml to the part of tbe nerve eotiuci-iixl with the bntin ; ami while
tbtHral «ifbct on the bean of onllnnry mimulHii'iii nf tlie vagiu u iiihibi-
tba WMnieDUtiOD unir oix-urriiix In ngwi'iiil cil-hm, llie uhiot cftmmOD efibet
(■nqmiMit b au^entali'iii, tWi^b iiiliiliilion i* not unfrMiiientI}r aval.
V^ the PX)KTinM>ni m f<)n(liii-(«il no iiii nomiiil iiiidcir ihr full inQiMinoo of
(UonI, *(iiiiiiliiti«ii 1)1 tlie vngii* goii«'rally [inHlticcM iiibibitii>n of reapirft-
(IM, pnihal>lv bocniiiw' ihn clil'irnl ri,iii)?n> the rrspinitory centre more sua-
TJble t'> iiinibilory iiifliii-tKiii.
9M. \Vf wiid just DOW "tiie acliuD of tbr criilro"; but the respirntoiy
Mtn ii n il'iiible ooe ; it givra n«c to innpiratory and to expiratory eflerent
ibmiIms, BtiH thoc arc aiilngonuiii; th? one to the other. If hispinilorjr
w ripinilory impuJMw itMimJ fmm the centre at the same time and in
"|uk] potency, there o(hi[(^ bo no breatliing at nil, they >irould noiitrattta
■^ch i>iher'« eficcts : am), indeed, any nmiiuni of inspiratory inipulBO is
"■laiCoiiiBlic to n siiuullaneow expiratory inipiiise. and rice vcreri. Henoe,
"'' tbe adeouate aervioea of the rcapinttory centre we niigbi extwct to find
^^•t eaeb kind of afferent impulse asoeudinf* tbe vaKUH aSeetea the centre
'(> a double and opjui^ie uiiy, iuliibiliu); expiruliuit while uugineulinj; iiupi-
'Vtitio, or iuhibiliu); iuapiratinu nhiie au;;iueiiiiu>; exptralinn. If ne allow
*^iuwlvea to g|)ealc of the whok n^Hpirainry centre a^ couaintii^ uf two parta,
*^ tim inspiratory (wrt, or in^ipinitory i^^ntre •xiucerned in the irtaue of iu-
4|iraJnrr impulaea, and Ibe other tbe expiratury )>arl. or expiratory centre
fwcemt.'d in tbe iMue of expiratory iuipuUt-a, we may auppoae that tlieae
(•tm are mt related to each other that afferent inipuUn. reiiebiu); tbe
Mdulla, which augment or inhibit the one, necHMrJly inhibit or au>;iuciil
ika other. We mxil {vrhajin hardly add thiit of tbete two centres we .diould
•Ip«ct lo find the iufpiratory ecnire the dominant and tbe most rcsponiivo
OM; in twrmal brcalhinf; it comes alnioit alone into obviou* uw, aince na
*e have «evn the expiratory muaelea have then a very flight Inflc only, tbo
(icat beinjt enipliea chiefly by elaatie reaction : and, e|M?aking generally.
IiKathtDi; in is tbe firtt conaidersiion, w« breathe out nuiatly bwauw we
kmre already brtalbed IL
There are many facta which support this view of the double antapmistie
action of aflerenl respiratory impulaea. If tbe central end of the siipeiior
larrnicefil bnutch of die vaKua be Hliuiuluied the elTecta are much more coD-
•unt than tbui»e <>f atimuhUinK (he main vagus trunk. Wbelher the main
^Arunk of tbe nerve be nrevtoualy aevered or not. (he reault of centripetal
BKimulation of the iuperior lap'ngeal branch in always in the direction of a
^buwiaKof tbe rmpiration (Fig- 143); and thia may by projier aiianilatiou
Fba earned ao ftr thai a coaipfele »tand»till of r«»piratii>ii tu the pliaw of
r«sl i* brought almut. While the main trunk of the vagua ooutaina Rbrea
iif twu kind*, both aujimi-nlor and inhihiUirv ui inHiiiration, the au|>erior
laryomal breoch npp<'ani to (-''Ulain nnc kitui unly, tlniw.- which inhitiit in-
Stratiua. If now while ihia )'X|H-nmi.-nl u being conducted on a rabbit the
idonieii be wnlclx-d, it will be aei'u that the inhibition of inKpirnliun la
acconifianii-d hy a cmtrarlion of tbe abdominal iou>clti>. that ia by an elfort
at expiration -. iIm- atimulnlion of the Dcm whilu inhibiting Tvnpimli'm pro*
Ivukea, ti> a certain extent, expiration.
i 367. That the trunk of the vnf:iia is the channel of these two kinda of
ImpulM-a, of a niutaally anlag4>iiii^tie character, ia further ahiiwn bv apply-
Inj; what may be considered aa natural Mimiili lo the ending* of the nerve
in the lunjt* ; and the reaulls ao obtained have an eapecia) valui> ainoo the
artificial Mimulation of a nerve fibre at a [tart of ila courac by menna of an
91
482
electrii' forrent U at Imt n r'llgli prm-esiii, by wliich w« caonot how to*
tnore tliau appruximntc to tlii: rwult* uctutiUy taking pUc« in ln« livJi
Swmso a* tuviaAtioa vt finiicLinoei at Suruio* Laimoaiu. Knni
TM«ti)n*wMnliia1iit>Iln(hiimneita)'**>^P' DIMM 141 ■■><) lk« bM«tt h*M Um tam
mMOIng u In tbaM Igure*. SilBulatlao liOftiu at 2. aad mdl at p
body wbcn the nerve is stifniilat«d nt its eadiDx* by nattiral utiniiili ; u
the Approximation ia perbaiH len in ihe cise of tb« exiiuiaiuly Moiili*
respiratory centre than in many olher cases.
Flo. IM.
^ I i_
_i I I j_
-I.
B
mrwv or DnmtMN uiB Couurw •» Ij^mi. illub.1
BKb cniTM an ilewtlbfil hyo let*? MIWIibI, ki ilalcl l>|»M, to ■ illp at llw dltptantaM
rabUl. A ^wUMlloli «f llie i)l«|*ni(ai lliu|iln>d(iii> nlut Uiu Inret-. during raluUlon oTU
itt*ph»t<n-*>>"l>**' AiUt- la A. (ho tecimUclofnl u j. Ihr hplEhi 0/ tiniintloii: ■ t<««iil
lomdinliic whiehiha lenr cwIimI])' iliiki until an laoiilnitloa (i ■tvts pDRuirul mii'i M* Bk J
a. Uisitactwatoclowilfliiiiooiiilor«irl**U(n.': UM»Ci)lo«r luaornil iiuiiini^iau.
4
THE NERVOUS KKCnAyiSU 07 RKSPIBATION.
483
If in KB animal m wliich « i-areful L!ra|ihie record of the rwpiralnry
>m<>V«nM;nU U being lal(i:ti, th« truchcu be sudilenly 4!liwe>d iit lh« «iiiiiniil at
■ui in«|>intion, thu rcJiiU ii » piiiiMr Ix-forft the ouoCMding iii4 pi ration folluwH,
llinl is U> my, ■ juutinl or U-m|>i>rary inhibition of in*piriilioii ; nml if iliirii));
Moti iin i'X|>criincnt on n mhith m curvo he bikvn ]>y iu«nnit of the iw)Ititvii
slip of (Ik- iliaphniifni, ^ '^2V, ii will be «vcii { Fig. 144, A) that tlic «Ii|i vlon-
giilc« Homcwhiit ; ihat i* to my. previously in n t!tat« of slight tonic eoBtmc*
tioi). il cluuigro ill the <lirvctio:i of cxpirution. If, ou ih*: other hun<l, the
trnchcu l>c ouddcnly closmt M th« i>ii(l of »n expirniion ( Pig. 14tt. It), when
the lungM have returned to thwr emptied condition, thu n«iilt i» an increaea
of the E«4|itent inapi rations, that ifl to say. an siignK-ntniion of iiwpiralory
imiMibeB. If the che^i or if the lun); only be geatljr intlatod a temporary
ceasali'-n of nil iHi]>iratiuii may he produced, accompanied sometimes by an
Attempt at expiration. If, on the other hand, air be sucked out of tiM
-L.
.*-
_kU~
I t^
Ktnxn or Sipumi tsruTiuvs, riiuriti Vixtuaiiot' (llut>,>
TtM limn curv* la dMtiltrod. u In Pij, tu. by • limr stMoiMrf lo s illp iif ih« ^Ikfkngm. Tbe
iinntMimitKmitbv InniUoiu from i ta ;. wlilcli w*n Inwiii wlllioul aur ■Uaiii|< lo diaw lliv
klroaliiiMeh intiaiiun : tmeh Hhihi llili aarrD <loiial«> an tnlUilnti, it wID M otMned ihai n>
Uio loMloM an oinUiiuvJ Uw nfplMorj manauoH uT the dlitfilirktin arv xndtnJIr ■■ knocKol
down."
chest, or if oiM lung bi- nisdv lo collapse by puncture of one pleural cham-
ber, a prolonged inapirnliou is the frequent rejult, tlie diaphragm being
thnmo into a prolonged inspiratory tetanus. If the liini^ are repealcxlly
inllate*!, without any meaiu bciog taken to draw i>iit tbe air all«r eadi infla-
tion <FitC- H'^).* procedure which we may s{>Mtk of as positivtr veiililatioa,
the retiilt in iliat Uie iusptralory etforl^ are diminished, and if the ventila-
tion ta continued mar cease alto|^ther. If, on the other h«nd, air ii repeat-
edly sucked out of the lunga. without any c(irresnondi»;t inllatioiio, negative
vtrntilaliun, the unpiratory eni)rt9are increased (Fi|;. t4ti|, and thi> iiicrsMa
may be nucU u» to briiiic tlie diaphragm to a state of leCaiiun. And in
Snemt, though i^'vurul coiiiplicatioas occur which we cannot <liKcii)« burv,
I- ri'atilt.H of inllaliou of lliv lungn on the one band and of rinction or »>!-
la|><H' 'if tbr liiiig:> on lilt- oib<,ir hand. *how tbut tlic mi<re inflation or por-
X rather the tmrn- dUtifii«i»n of (he lung ten<U to inhihjt inspiratory and
in rxpiruttiry itn[iiil»s, while colla|WO of (hv lung tenus to inhibit
•xpinUorjr ami tn dt'vrl.tfi iti>|>iralory impulwM, the ulbct on ibo inspiratory
iiapaltM, M might he <'X|H-c[c'i from the domioanoo of the iiupiratory por-
tion of the centre Ix-iug niurc nmrkivl thao the dffiict oil thv expiratory
impuWs. That the iii«truiuetil by which tboM eflecla are nroduoed is the
vague nerve is shown by tbe fad that they arc no longer distinctly rvcot;-
niuible when both vagus nerrea are divid»l, And (hat the results are due
to the mere mechanical expansion and coltansc of the tung in insulBalioa
and collapse, and not to any chemical influence} exerted by tbe larger
484
KESPIRATIOS,
amount or amftller Rmount of air preseDt in the lung in the two cases in-
creasing or dimiDishing the absorption of oxy^n and escape of c&rbonic
acid, is shown by the fact that the results remain in their main features tbe
same when some indifferent gas such as hydrogen is used for inflation in-
stead of air or oxygen. We infer, therefore, that the expansion of the
F»i. mi
-^-vWWW\A
ElKIi.-!^ ur KEIT-ITEII SlCnilNr. tlV THE I.INi.S. NlIllAtlVE VKSTlLATillN. (ilEAD.)
Till.' pun 0 corrtsjioiulEt i.'XHCI1>' to t'lg- H'l, uxcn^lit Thut iJii! Uuigs lire niiUJ^-rliHl 10 repeated lUctiona
H'llboiit «)rtvi>iinUIiiK iiillHliim.''. Ttiu n.-.iiill Ik Iliat lliu liis|ilm[!niii nn: n.'K'ilt.il Id uicb a way u
to leoil iiliiio't to«n liisi-lnit'iry lelmiiisof llii; 'llni>liniEm.
puliuonary alveoli in some way or other an stimulates the endings in the
lung of the pulmonary bmnchea of the vngiii'. that impulses are generated
which ascending the vagus trunk inhibit the inspiratory processes in the
respiratory centre ; and that conversely eolla|>^e of the lung similarly gen-
erates impulses which are augmentative of iusjiiraCory impulses. And,
assuming on the strength of analogy the existence in the vngus of two sets
of fibres we may buy that expansion stimulates the endings of the fibres
which inhibit imtpiration and concurrently tend to augment expiration,
while collapse stimulates the iibrcs which inhibit expiration and augment
inspiration. The respiratory pump mav ihiis be looked upon as a self-regu-
latmg mechanism ; the exjiausiun of the lungi' which is the result of the
efferent inspiratory impulses tends to check the issue uf these impulses and
to inaugurate the sequent expiration ; and the return of the lungs in expi-
ration tends to set going the succeeding inSi>inition.
The regulative influence exerted by impulses normally ascending the vagus
nerves is further shown by the following striking experiment : As we have
already seen, the brain above the medulla may be rcni'ivcil without any
extraordinary change in the respiration taking place. Wv have also seen
that when both vagus nerve* arc divided the respiraliim is slower iind deeper,
but is otherwise regular. If, however, after reiuoval of the brain al>ove the
medulla both vagus nerves are divided, if the resjiiraturv centre be cut off at
one and the same time from impulses pat^in;r down frutn ihe hi<.dier ports of
the brain and from impulses asceudiiig ihe vagus nervi-:^. the result is that
the respirations take on the form of a series of lon;;-<'oiittiiiicd inspiratory
spaems. It would seem as if there were a tendency in llic ret^piralory centre
to go off into tetanic inspiratory explosions, that this [endein y is held in
check by impulses from the brain when the vagus nerves are divided, and
TUB SEttVOL'S MECUA^ISM OF KKSPl It ATION.
485
' h^ Mftubra aliKii; i)h> vb|£ii> iH-r^-ea when ihe lintiii i> reniuveil, lull iiin-U
vubno MJequntr rluvks wIh-ii im)HiUf» from liuth Miurccn ur« cut off iit itio
matlitae.
t368. HyiwrtboM* have bevn pui fomiinl to «zplidn the cbatifntt in t)i«
rMftatorr ivntrp which Iciul to the rhylhrniir iliat'hiirKi* nf in^iiirntury nnil
•XpiiWorr ini|iulwii and tbc fuHhirr I'hniip^ which ronult I'min ihi- nilrcut
of ujtnti'otiiig iiRil iohihitorv iiii|>ulM»; hiil thmc m yvt rt'riiiiin inrrr hrji-j-
^hww. mil it wiHiliI Hi>t 1m! pmAinbk- todiscuw ihcni liirM-. Wi: niny mid
ti^Mi Ihiiugh tin' »nal<>!;v of the cnnJmr nfrvoun nn.-chttnwm. in whioh wr cttii
■Ostarakall^ dttlJRgtiuh Iwtwccn itneint^nl'^r and inhibitory fibm.jii!>tilii!:t
5" to ifwakiRg uf Kugmentfrmnd inhibitorr »nd rapinitorr Fihrcn tie cxislinc
ia Uw T^us aem, wentv not tu yot nhl« to difttinKuish Ihem bv nniitumical
"■Nlvidft. We may further ndd that. «o «xqnint>>Ty senaitive ie the mpira-
***';3r «ott« to th»e Bli«r«ni impulMse tliftt stimuli loo elight lo prndiira any
*f'pniciablp «l1#cl when Applied to aflcrpnt nerves connected with an i>rdiuary
***ttn; such u u spinni retlei centre, mnr produce marked etlet-td on the
'*"|Mmlory centre. For inslaDce. the Tceble electric current which is de-
'^Iftpr't whew the out eiid of a divided vagus is replaced in tbe wound, (be
*^^^.^|il belwM-n the cut end and ibe lougitudiunl Mirface of the nerve beinjf
^Voml ilir'>ii|;h the hl<Mkl or lymph of the wound, ia ofleo lufHricnt to de-
**1b|> inhihii-jry iiupulMSL A^in, when tbe eouncelion of thu rt^pirntory
^«Btre with the lun;pt through the vagus nerves is aboli.ihed, nol hy tH'ctinu
of llie nerv<>«, hut by [ntft'inn huth nerves ai aome pitn of Lhe courw nf i-ach
Bcrrv (an operation wbtch, while cuniidrivly hhickiiitc the iiMsageof impiilM!*
liomtl ^* nerre fibrat, doui not it^if net u a Htiniuliii), the eHfitct on the
tvpintarr RioTeincntii it niucb more in tlie dirv<rlton of iiu'riikiing iind pro-
loD^Og the in*}iimtory acl than «f Klowinj; the rhythm. Hence it wotild
ippCT that what we have prvviousir duscribed n» tliv result of dividiui
both vague nerves i* partly due to the blockin); of natural impulBes and
pwtljr lu the sertiftn of the oervfe. and posnibly t« electric curreul* <le-
veloped as 8ii]ci;estcd abore, acting as «tiinult s»d thus giviD][ rise to artificial
inpulwa.
( 380. The double or allerinile n»i>iratury action of tbe rajius nerves on
whkb we have dwelt tibove muy Ito iiikeii as in a j^ieral way illutilrative of
tba nanaer in which other atliirent nervee and variuiiit \nnti of the cerebrum
aiC tnabled to influenee reapinitiun. Am we have ulrt-udy onid. and. indeed,
know from daily ext)erience, <>f all iht^ n|nrchicul nervous ouilres the lespirtr
toTf omtre i» Inc one w bich i* m»cl fr<'<(ueiitly and most deep); aflbcted bf
B«rmus inifiuljMa iVom varioue iiuurleo'. Bi^iiiloi the chaniNS brought about
hf tbe will (and when we brualhe voluntnrilr we probably make uiw to aome
•aWsl of the noniiiil nervou* miii'hinvrv of n'^ipiriition. working tltrouvh
tbi*, rallier ihaii siMiding in'li?|><-»d<-ut volitional irti|>uliiM dirtTt to tlie dia>
pbr«gni and olhvr n-«pirRlory muM-lct), we find that emotionK and juiiiiful
BMWfttittn* alter profinindiy the cliaractcr of the Rvpiratorr nioveiix-nt*.
Ami lhtMit:h ibnN.' c ITccIm may he partly indirect (the emtXion modifying ihe
rt-l)«at or the timun arterive. and s» inlluencin); the flow of blood thn>ugb
riBpiratnry centre), th«r are chicHv duo to the direct action of nervous
[miIhs, reaching that centre fronu Wher psrU of the brain. So also
impiiltai fnini almost every eentieot surmee or pnaaing along alinosl every
MBSory nerve may ntudily respiration in one direclwHi or nnother. Tliic
lafloence iu this way of Htiniitli applied to tbe akin is well known to all ;
bttt, perhaje. itext to the vaguii ibe nerve moot clutely oounecled with tliC
resplraiory wnire is lite flfih nerve, branches of which giianl the
nvplmiory rhiinnels: the -ili^thteiit mi mu In I ton of tbe iimtriU »l once
thtb£Mthinj; an<lmii»t lh!«|ueutly iirnats it. TbeellccUof -aiinuli of rariwis
ih tiK m
uatal I
affects ■
^ariwis M
486 BESPIRATIOS.
Strengths brought u> bear on various nerves are very varied. Sometimes
the result is an increase of inspiration, and that either by a quickening of
the rhythm or by an increase of the individual breathe or by a combination
of the two. Sometimes the result is inhibition of inspiration accompanied
or not by an increase of expiration, and sometimes, as when the stimulattoa
causes a cough, the expiratory rexulls may be out of all proportion to the
nioditicalions of inspiration. While in the case of some nerves, tor instance,
as we have seen, the superior laryngeal, and it is said the splaDchnic Dervefl.
the effects are exclusively or at least chiefly inhibitory of insniration and
augmentative of expiration, that is expiratory, and in otters, perhnpe, chiefiy
augmentative of inspiration or inspiratory, in the case of most uerreB the
effect may be according to circumstances either in the one direction or the
other. Perhaps, as a rule, weak stimuli tend to augment and strong to
inhibit inspiration ; but the effects of artificially stimulating sensory nerves
are complicated and often confused, because powerful afferent impulses by
fiving rise to pain may, through impulses generated by the pain itself anc)
escending to the medulla from the brain, act in an indirect as well as in a
direct manner; and tiie prominence of the indirect painful impulses will in
any experiment depend on the senesthetic used. We may say, however, that in
all cases the effect is very largely determined by the condition at the time being
of the respiratory centre ilselt ; ami that is in turn determined not only by
things which affect ila nutrition, such as the character of the blood circu-
lating in it, but also by the nature and amount of the other afferent impulses
which are playing upon it at the same time. Thu.i, as we shall presently see,
the etTect of a stimulus applied to tlie vagus, when the respiratory centre is
inndefjuately supplied wilh arterial blood, is nut the i-anic when the centre
has its normiil supply of normal blood. So also a sliniulus which, applied
to ihe vagus or I<) another nerve in iin intact niiimal, simply quickens
inspiration, iijiplied in an animal wbifc ct-rcbnii hemispheres have been
removed will cull forth a pnilon^icd tetanic inspiratory jiiisp. The respira-
tory crntre reFpon<ls, iti fad, in ihc nuist iutricnlr' iiud varied manner to
nervous impulsi-s proceeding from all parts of the body, and thus deli-
cately ndjuslM the working of the respiratory pump to the needs of the
economy.
s;370. The complicated nature «i' ilie respiriilory centre i.-' further shown
bv the fact that it iipjiears to consist of tmi inlenil halves wliieh normally
work in unison mid yet niav l>e made to work iude|)ciideiitlv. If the medulla
oblongata be carefully divl<k'd in the middle line re:'pit'u[ii)n may continue
to go on in quite a itorumi fashion, if, however, oue vagus he then divided,
tlie re!'}iiratory movomente, both costal and diii]>lir:igninli<*. on the side of the
body on which division of the viigiis Ims liiken place. becTiie slower than
those on the other side, so that iho two ."ides ;ire no lou^'er syuchronus ; and
a stimulus (Hinfincd to one viigus utliTls tlie respiratory movements of that
side of the body only. So also a seeliou ef a lateral hull' the cord below the
medulla stops the respiratory miivenicnis on that siile alone.
§371. Besides these nervous inHuencfs, however, there is another circum-
stance which, perhaps, al>uve all iitliers, atf'eels the res]ur!itory centre, and
that is the condition of the blood in respect to its respiratory changes ; the
more venous I leps arterial) the blood, the greater is the activity <i(' the respira-
tory centre. When by reason either of ntiy hiiidrjuiee In the entririee of air
into the chest or other interfereuce with the due inlercliunge between the
blood and the pulmonary air or of a greater respiratory activity of the tissues,
as during muscular exertion, the bhxid becomes less arlirial. more venous,
I. c, with a smaller charge of oxygen and more heavily laden with carbonic
acid, the respiration from being normal become.-^ laliore<l. \Vc loay speak of
Ttre XERVOUH MECnAMSU or RBSPIBAT10X.
■187
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tbiag as eitfiH'ta, nm) sav iliat ihiM, wh«n (lie bK>o(i is insufficiently
mierBtlhed; jMrnx ini» ilyronaa, nn iiiterni«diale aiujff. in which the rtsfiira-
tAiry moveineniii Hr« .■•iniply vxiiggerHict), beiti); known us hj/jtetjiHoa, Tbc
nKHlifinalionjt uf lirt-Blhiiig i)iu» chumkI by d«riHi.iit urterialixalioii of lilourl
lire viipcci«)l}' chaniclrriM^t hr an incrt'-iwc in the lotat boer^v uf \)m napira-
Iiin>' iniptil»n Ki-iicrntcit, ami in thia n-opvcl ilitlV^r iViiin the nuidiliciiiiotii*
reuniting fniiu iiili-rfrrrnci; with Ihc ncrvnii* nrnuignnciiti) nuch ait thiwe f<il-
luwin^i! ii|Hin section nf tbo rttgii* iwrvif, in which ciiwv n» wc have WH-ii, iho
rhvlhm i» much more proloiirHllv atlcflrd ihnn llic Jinnniiit. In ilynpnceK
lln- hreatliinp ts frequpRttr tjuickcr an well ns ilccper, then.' n en iiictx-iiiw in
1\m- Hum of cflrrent trspirntary impiilm^. and the uxpiralorr iropxiWi!, w hich
ill uornial mpiralion are very slifiht. acquire a pronounced importance. As
thv blcHxl boeonwx in ca»eN of obslrucliou Ie8< and Icm arterial, more and
more venous, the diechiu;ge from the reepiralory centre becumce more and
more vehement, CImI instead of confining it^lf to the Uiiial tracts ajid paee-
tag (Idwu to the ordinary r^^ipiratorr mUBcles, ovorflowe into otiter traela and
puts intu action other muHclea. until there is, perhaps, hardly a muscle in the
body Vfhich is not made to feel its effects. The muscles wiiieh are thus more
and more ihmnn inf> action aie especially thoae teiidinK to carry out or Ut
snisl expiration; and at last, if uu relief is alForded. the riolent but still
definite resniratory movemetiis give way to ueneral convulsions of llie wluile
Ixxly, w'hicD. howi-vcr, have to a certain extent an expiralon,' character.
With (lie oniN-t of these ccmvuluons dyapnoea is said to have pui«ed iiiKi
iitp/tyrin. By th« violence of these ooavuMons the whole nt^rvouH xyatent
ItrconMv exhausted, tho onvulniona cesM, and death is usherol in through a
lew inl'rcipii-nt and Inng'ilrawn brcatlis : but to thin maid-'r we iihall return,
TImi effect of vcnoii« blond, then, ix to augment all thoik* natural cxt>l(i#ivc
dee<>nip<i*ili<iii<ol' (he fulwtancc of the centrul ncrvHiii Hvntcm which give riw
to respiratory inipulM^; it incrunMv their amount and aW ()uick«Di> their
rhythm. TIio latter chiuige, howcA'cr, it much Icm marked than the fciriaer,
tbo rtvpiralion being much more dec|>cmvl than hurriol, and the several
n»|>irtitorT acts are n«rer ff much hiislcned as to catch each other up, and
6o (o )>roduce an iuspiraiory tetanus like that mnlting from stimulation of
the vagu>. On the contrary, especially as exhanstiuu begins to set in, (be
rhythm bec<omea slower out of proportion to the weakeiiin); of the individual
tuoreraents.
S 878. Tii« question naturally ariwe, Does this condition of the blood
affect the substance of the central nervous system, that is to say, the respira-
tory centre iu thd medulla (and the subsidiary spinal uervons mecbauisms)
directly, or does it produce its effect bv stimulating the jieripherul emU of
oiftfent i>en,'es in various jMirla of the ^dy, and, by the generation there of
aflbrent irojmUes, indirectiv modify the action of the central nervoiiB ^vsteni 7
Without denrtng the potPihility iMt the latter nio<le of action may lielp in
thv matter, m- r(-gurilr> not only the vajiUR, hut all alferent nervts, ttie follow-
ing faclji utrf.m to shim' that the niaiu vfftvt 'n pniducMl by the direct action
of the bliHid on the e<.'nlral nervottn svittcm, and, imteH), on the mcdullaiy
rapirulury <H'ntre it^tf. If thi^ n)>inHl oinl be divided l>eIow the miiluliii
obloogHta, and l><>lh vagi be cut, want of pn>|H-r acrulion of tbf hlood ■till
nrodiiceM an incn-a»-«l activity of the rw|>imtory centre, lut shown by the
incn-aM^l vigor of th«t facial rc^'pinitory movcmcnti". If (he nupply of IiIixmI
be cnt olT (mm the im-ibilla by ligature nf the carotid and iutervrrlebral
arteries dysima-a is j>riHluced, though the (iiwrwtion prodmv at first no
cliange in tlie hh>od generally, but limplv nnecu the n^>i^l^•rv condition
of the nie<lulla itself l>y cutting off its hi uu<l- supply, the immediiile ri-sult
of which is an accumulation of carbonic acid and a |>auci(y of availnbl«
tof that r<4^inD. IT l)iO Mum) in the rnnrti'i
nliDVc lliir iiormal. n <W*\taiK* x* |ir»<lu(Y<l
oxrgtMi III inp n«
•rti-ry in uii nnir
whicn, though iipimrentlr not niiilv ii i<-i)iic-iil with lltu ilTAinnrii i!«iiw>l hy
ini|Hirfrct iirlt-riali/alioii of ihi- lilooil, "hunrs thai lltc Cuii high tci»|H.'nitiit
of the lihxiil iliriTllv iilfiM-l* l\n- iii-tivity ol" lliv rcjipinitorv wiitre. Wo ma*— ^
oUR'hiilf. ihcn'r^'ri', ihnl ihi.- <.vii<)itiiin of the hliioJ ntlocta ntfpiretioti l><-^
acting iJirwrlly oti ihu rt»i>initc>r_v c«nt«i, Moroovtr. it fa tho nwiliilian— ^
centre which, at till vccule iu uduU aniraula, is atTcctotl by th« too vi-rimi ~
hload, eitice iiflcr divisioD of tho »piiml oird bp low tho meilulln. il)-#pDiri
thoracic rpepirntory move men Is nn<I convulsious do Dot follow upon exclu
of air. They are, however, dated tu occur in newborn oniniali), indicalini
that Uie subdidiary niechaDHms in ibe u[>|)er epinal cord, of which we enok
iu ^ ^Gi, may be aiai) affected by the too venous bloud : but ihe doubu wliicl
w« previously urged hold good in these cases also.
While the rtepiratory centre Is ihui being atTected by th« urn veoom blood ,
it ii>, until exIiHii'tion begiuv^ Iu aet in, more irritable, mure eiuily iiud lar|[el>r
aflt-cled by ulfL'ri'tii impulses llmu iti iu nurnial i-»iiililii>H. Diirinu; dyspniB^
a KtiinuliiB which applied l<) the va(r|iM or in mime nlher »uii<<,>ry iiervt; tiuder
nurmnl i^oiiililiiiiiH would produce littl« or no edlict, may stitrl very ffuvitrfaj
^373. bi-lici<rut lu-rntioii produces tivn effects in blood : it dimiui«he4 iht
oxvg^in and ini-r<.iu>L-ii the carlnmic add. Do both of lht«e changed aSwt
the rr:<[>iruli>ry ct*4ilr6, or oulv one, Bud if si>, which '! When nn animal is
made to brcntho an atmiMplierc contiiiiiing uilmgeu only, the ozil of aw
bonic acid by diflugioii w tint alTcctv<], and the blood, ns is pn)v«l by actnal
Bna1yai», contiiiiiTi no cxctv* of carbonic arid. Yet all the phenomeDa of
dyspncea an: prcwut, and if tho cxporimcnt be cuntiDiied, conTuUiooa cnroe
and the animnl dies in asphyxia. In this case the result can only be
attributed to Ihe deficiency of osyKcn. Ou the other hand, if an animal be
made to breathe un atmosphere rich in carbonic acid, but at the same time
containing abundance of oxygeu, though the hreatbiii^ becaraw markedly
deofier and aUii mimenhai umre fre'|ueul, there is no culmination iu a eon>
Tulsive asphyxia, eveii when the quantity iif carbonic acid in the htood. n
ahuwn by ilirocl analyvia, is very largely iiicruased. Ou the ciMilrary. t)>e
incmuM; iii the rvspiratory moverrii-iiU may alV'r a while pum off, the iinimal
becoming uncoiiKcioiis. and appt^ariiig to he vulfcriug rather from n narcotic
poison than from Mmpic dynpniiii ; llie ■■xci.^iw of carbonic acid in the blooil
appears to afltict other parls iif the wnlnil nervous i>ysteiti, and ojH-cially
forlions of the i>rain, more pro^iundly than it does the reypiralory cenirc,
t has been muintatncd by K>nie that while a delidcncy of oxygen pr()mot«>
inspiratory movements, an cioon of carbonic acid »timulHtr« Ihe expiratory
movements, ihe nervous madianismg being so arranged that a lack of oxygen
leaits to an eflbrt to get mora of it, and a too great load of carbonic acid to
au elfurt to get rid of it ; but the facts are opjiuseil to the eiisUmoo of any
■ucb teleolo^cal aduptntiun. It is obvious, however, that a lack of nxygeo
and an excess of carbonic acid uSeci the iespirat»r>' centre in rery different
ways, and that in ordinary cases of interference with tlie inlerclianKO in the
lungs, as in deficient ai-ration, it is the lack of oxygen which jnaya the
prim-ijiiil part in developing the abnormal nupirutury uuventeuta. We
inay infer ihat it too is chielly voncerueil in n-gulaiing the more normal
nepirution, hut cannot as yet say what !• the exact .iharc to be attrihutis) to
the earbonic acid.
AVe may here (xiint out that it is n»l (o bo suppomd that each brcatit it
detennincl by the condili'in of tho blood flowing through th(< cnpillnric^ of
tite medulla at the moment |)rocedilig that lirenth, il i« not to be imagiusd
Tns NKRTOt'S MBCnAXISM 0>' RESPIRATION*. 4S9
titat eacli breath u tbe result of t)i« lack uf nxy^n felt i in mediately before.
Od th« iMtilrary, ta we have |ir«rio(iHl_v ur^j, the rwpirntoTT,' rnitie, like
tUtb rjirdiao Milwlnim. U nii auloinatio i;e»tre, i)i« respiratorr i(ii|>i]UeA Uaue
fVoni it in rbylhnii<r wnM sjt n rftiiill of tlui nxilcoular cliiinK;«a of (he
inetaboliani going mi iii il« niiKhIikkv ; an^ whiitrvi-r atlH'tK lliat rli.vihiii,
L w)i«ther feir or miwy b«*t« be iiifliii'iioHl, proiluora iu tiMull liy mi>rlitViiig
I that meUboluiin. A lack of oxyi^'n in itir blood, or n iKrrvuiin imixiliti
along un nt)(in'iit fibrv, Ixilli nAVct ibv (^'nlri.' by nimlifyiiif; its ludabofiMn :
Imt I'luh probably aflVcts it in a liiffcrrnt way. It i^ bfyond our preMnt
kiiowWgv lo explain how either the one ur the other mrt*. Wc may imagine
that a lacl: of oxygen on the other hand hat a morv prolbiind utli-ct in modi-
fying the wholi? oom]>lvx eerie* of metabolic cbuigcs. tb« whole chain of
building tip and breaking doirn prooesws, thus in Bonie war or otbi-r rrn-
dering the whole edifice, m tu spmk, more unstable : mid llial an attcrent
InugniCDting impulse (and poa&iuly an excess of carbonic acid) acts nilh«'r
nnvr the niehion of what we are accustomed to call a slimutiie, and tires otf
n lormr amount of the already stored up explosive ixmipounds. And wo
may further imagine that the spectat feature of llie substance of the respira-
tory centre is that its metabolidin is eo wranged na to be thus, unlike that of
other livioK suhslanoes, rendered unstable and more explosive, not siniplv
diininisheil or deadened by a laek of oxygen. But tbeee as yet are matters
of specutiition.
»We iDKyf lierhape, add that, under i,-arious nutritive eoudilions. tbe seu#i-
tireoen of toe metabolism of tlie reapiratory centre to lack uf nxygeu ma^
vary niilely. Thus, while undouhledly under tbu normal niitrilire condi-
tions aflorded by the ordinary su]>|ilr of ixirnial blood l<> the medulla, laok
L of oxygen in that liliHtd at onoe pnivoktv inrreiued rMpiniuirv niovenienta,
f it oee^ not do s^i nmkr ntlwr nutritive condilionsof ihe medulla. By trans-
ftision a large pr>i|Hirtion of ihi- liaHnoglobin holding blood may in an animal
l>e gradually rt'plaenl by liiemogiobinTcMs iiorniid #al{aeaolutton. hi ^urli a
oaae the amounl of oxvRn brought to tbe meiliilla by the diluteil blood
muM be greatly diminished, a»<t yet if ihv eliange Iw made suiriciintly
slowly, no 00fiMHrtiou.-> dvHpno-a in jircHlticetl ; tinder llie new vtrange nuuitive
Icofuirtioiu of the diltiUiil bknid the niedulU is not iifll'clcd in the same way
U Iwfinv br lack of oxygen.
i 374. I'herc are reasons for thinking that conditions of the blood, other
than variations in ihc amount of oxygcii aitd carbonic ncid. niuy niao nMtt»>
rtally iitfect the vorking of the respiratory centre. It is a matter of common
exjirrivooe that mtiwrular exertion, especially if at all exoes*ive, incrvosH
tho rcMHnitory movements ; violent exercise soon puis a man " out of
breath.' This incrrasod activity of the reepiratory r<cntr« is in large inoiisnr«
at all events cauted bv the character of the blood, which during;; and for
some liule time atU>r tlie movements is curried to tbe medulla, and not by
any iierviius impulH>eti «ent up to tbe medulla from (be contracting muscles.
This i« j^iown by [be fact ttial if iu an animal tbe spinal <-nrd be divided in
tbe dorsal »r lumbnr region and the bind limbs be powerfully tetanized, tbe
resniratory niuveinents are increased ; the animal pants as it would do if it
hail l>cen runuin;;. In such a case the only connection lielwet^n the bind
linibe and the rwpiratory centre ia through tbe blu«<l: it muHt be some
diauRe in the blood caused by the muscular couiractiitns which affects tbe
medulla when the blood noaaea from the hind limbs to Ite dbitributei by Ihe
beort to tbe medulla. Vow when a mibwle ooulrocts ito on.tumplion of
oxygen and produeiion of carbonic atiid, especially tbe latter i^ t>:t}, are in-
creased ; the bhnid Iniviiig tbe muscle is more vtmous than usual. Hence,
when ntany niiisdca are contracting powerfully, the liloml carrier! lo the
m wiivu ouiny
^
riglit "ill"" onn^niSin in more vrmiun tlini) iiftiiiil : nivil wr nit^t ^xpM^lfiiiT
it w this iiniisiinllf vrirnus blmMl fuiliiig v> bo ncliijiiJiU-ly ariprUIi/j-H in th»
lungH. amt Iipik-c rFsching llif nuilnlla fmni ihr left side of ihti ln-arl in
more vrtiniie, Utn oniDplctply iirtcrinllicl oimilition thnii luiinl, tvliich «lir»
Mp tlie reapirntorv oenm- lo incrcnwi nclivity.
On exnniiiiatiati. however, it is f'oiiii'l ihnt the blood Icariof; tli» left Md(»
of ihe lienrt in •■iicli «ih:« ii> nol lc«s itrtcriniized. but, if Hnytbing. niof^b
arterinliwd ihan usual. The iticrrastsl respiratory movemenla indiicvd bi^
(he chani^d blood soon prove sufficient, or even nior« ihnu sufficient, to givo
the blood ttie extra qiinntity of oxvf^n and to remove tlie extra qiuniiiy of
cnrbonie acid. Obviously the blood coming from the (etaniKe<) muscles
tifects the rear^iraiury centre by virtue of some (luality which, unlike tbu
flue to the deficteufry of oxyti^n or exeeas of carbonic acid, is not initne-
dialely nflected by the jtatHafce through llie lun)(s, Wliellier the ijualitr ii
(|uciiti»ii be dep^<t<^Dt on tiu exceea of enrolactic and. or on borm other
ptnduut or product! of nitiacular luetabolism, we do tuA u» y«l know. But
the fai-t that nubtUwce* in ibe blood may An alFoRt the m>|iim1ory centre 'u
Inlen^tirig, nincr it fihow* by how inauy wufegiiardit the working of the re-
•piralory centre iii carefully adnpUs) l<> ihc nredit of the economy.
ThuK a elian^F in llu^ ciriHinintaneei' mirrounding an animal InHly.nra
cIiHnj.'r in tliv bodv itmrlf, iniiy in one or nion- of wivprni wavK. bv nrting m
n ti^imuUio to ininiit artcrint nerve* and no K-ndin^ up alTen-iit m-rvoua fm-
pnlRcs to the respiratory centre, or by interfering with the interchnng* of
Knt» in the liinp', or by otberwinc altrrinR the- proportion of the givc:> prewnt
in the blood reaching the re»iiinit'>n,- ecnirc. or by generntitip or incrntwing
in that blood some snbstnnec or substances tending lo aflcct the nutrition of
the respiratory centre, affect the working of the all important breathing
tncchaDism. And the alfcction »<> wrought ha>> generally an adaptalivp char-
acter; it generally teuds to protect the organism against the evil ededaof tlie
change.
$ 376. Apwra. When ire attempt to hold our breath, we find that ire can
do thin for a limited time iiiily ; tooner or later a brcHth miiu come; but, as
it well known, the linif iliirin^ whleh we can remain without breathing may
on occasion t>e niticb prolonged, if we finil of all take a iN-rie« of c eep breaib«L
It in probable, though perhuju not ilintinctly proved, that when we breathe
Voliintarily, or irlicn by nti act of the will wo. hold the mtpiratorv apparatni
in any one respiratorv phm»e, the nervoii* impiilmii, gt-nersted by the will,
do tioi pass down by a direct and independent roiiwe to the re^niratorv mua-
cles, but that tlie will makes n»e or niodilies the nctiviiy of the medullary
and fpiiial nervous respiratory mech«iii*ms. The brcnth tmoner or later
inevitably follows because at la»t the natiinil impulse* proceeding from the
respiratorv centre become too imperious to be any longer held in check by
tlie imputsei^ of volition pai^sing down to the centre from tli« brain. The
fact that a series of deep breaths, n thorough ventilation of tlw luncs pnat-
ponea the victory of the unconscious centre, shows that inch a ventilation in
•one way delavs the development of the natural respiraton,- impulses'. A
rimilar but sli^l more marked delay may oReu be seen in an animal under
artificial respiration. If in a rabbit artificial respiration is carried on very
vigonHisly for a while, and then suddenly stopped, the animal does not imme-
diately lii-jfin to breathe. For a variable period no respiratory movemenla
nt all lake plnce. and breathing when it does begin occurs gently and Dor-
nially, only |<ni<t>int; into dyxpnu-a if the nninial u unable to breathe of itself;
and even then the iranniiton in (piiti? gmdual. Evidently during thin period
the re»pirato(v lentre is in a ataic of complete rest, no exnlo»iuna are taking
place, ltd rexpiratory imputiN-di are being generated, and tliC quiet tianvittun
THE NERVOUS WECnANtSU OP RESPI nATIUN*.
-191
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from tbu condition Ui thut iit' Dornml rrvpinili'in »1k>w>i ihiit iliv Mulivranvnt
generation »f inti>iilA>ii U nttciitlcil t>y nn gmit (li»liirl>iin<N>. Noc au\y ■■'• the
centru Hi KM, Itut it ix Inw irntnhlv thim iW Tionniil ; iininilsoii nloiig the
vugUK or (itlicr ucrntu wliich olbvrviiw wmilil r>n)iliico mtpiratnry Axplixiioiis
arp now im-fltTtiiiil. Thi« «atf of thingn ii^ ktmwn ii;" ihnt i>f npn<ra. the
ooDverecof tlymitKi-ft ; iiml thr longer pniU'C in hmitliing incniionol aHorc
U poMittk- iifiVr ii»ii6iiiil vcntilntioii of llic liing« may Ih- rrgnnlvil ii» n brief
npnivii.
Xtm- it Mvnied nntiini) to tunpow that »tich a vUilo »i r«M of th« rf«[>irii-
ti>rv centre wiu> bTt>ii);hl nbrnit W the more thnn necwsiirily »nipl(^ Hi(i|>ly of
oxvgrn atKirdci) by llie ))revioii» increased in«]>imtory n><>venirnt« ; niKl,
indeed, it Kas Diaitilniiied l)i«l Bpn<»n wna th« resnit of loo great. )iiBt ns
dy>)>uiTa is the result of too liltif arterial iznl ion of the blood reaohing the
re><]>imt"ry mitre. It was argued that owing to the increased vigor <if the
nrlilirial refi|)irati>ry niorentenlii llie haemoglobin of llie arterial bliKxl, which
in Dornial lirealtiiug is not ijuite saturated nith oxvgen. heeame almmt eoni-
pletety BO, and thai at the same lime the ((iiantity of oxygen dimply dissolved
in the blood became laigdv incrvased and its tension largely augmented.
Unt tliere ar« reasons which rt-wler such n view unienahle. In the first
Claoe there ia no direct and sntixfactory proof that in apnwa the arterial
lood ia overloaded with oxyg«n no fiini nosed ; indeed, during the course of
apncea before it has come lo iin end the bl'md t)econ>eit distinctly leas arterial,
inofe venoiit than usual. In ihf second place ii[ino>« if not entirely impossi-
ble, is much nion? dtHietilt lo bring nlMHit irben both viigus ncrv«s are divide<l.
aixl if it d<ieH ort-ur after Hcclioii of the vagus ncrvcK hiu> not the same char-
acien a* onlimiry H|>ii(i'n. Now, when nrlificial revpinition is Wtn^ carried
on aection of the vagus m-rvi* can have no cffi-rt '<n ihc i]iiantily of oiygun
taken i>|> by tlie blood in the lung*. Out ihr vagu* ucrvt* are the diaiinvl
of impulse* aftwtiiig (be renpiriiiory centre, iin>l thi« relation «if the apunOR
ta tJM) vngus nerves »iigg(«t' another and diflcn-nt i»ter|ireliition of apn<r«.
lAs *ni bare •«», eximnKion of the lung by acting in »>me way or other on
It* pnlmMMrr terminations of the vagus nerve sends up along that nervi?
impulse* which inhibit inspiration. And it is nrgiitd that re|iciiled forcibk
inflations of (he lungs pi«duce apDoea bv generating poteni inhibitory im-
pulses, which by a kind of summation of tlieir elfecis in the medulla slop for
a whila the generation of respiratory impulses in the respiratory crntra.
This conclusion, moreover, ia strongly supported by the fact that uu apnom
may be {Hwluced. so long im the vagus nerves are intact, by forcible arlifictsl
respiration with hydrogen instead of almoeplieric air; in other words, the
inl^ibiiory impuldea generated in the vagus nen'«« by the inflation are sufti-
cieni wholly to neulrnliKe tbe development of reapirator^' impulses which the
deticieni arterial iut ion of the blood would otherwise have produced. Tbe
exact nature and development of such a summation of inhibitory impuUes,
eapecialty in tbe pre»enoc uf correlative atigmentaiivi? impulses called forth
by tbe currtviiunding aueceMiiTe <-olla|]eii=i of ihf lutigs, ia loo complex a
matter to Ifc itwcit <in hen-. Moreover, an a]>nie<a may b« produced inuugh,
as we have ratd, with ditfimllT Bl>er section of both vagus nerv'<«; but in
thi» case air nnd not hydrogen niu>t be nseil fur inltation, the nae of the
latter, in c<inlm»t to ihc n,-*ull when lh<' iiervi-H are inlai.-t. leading to dya-
pnuea. The subjeet eauiiot il< vet he eoRsiderc<l lu fully cleared up. That
apnu-n as ordinarily pnxtuced n in »ome way the rmult of inhibitory impulsea
genenii'sl by the iuHalioiw enn how-ever hnnlly l>« doubti-<t.
5 378. Srfouilitiy Tf'ii'ralory rtiylUm — Vhrynr-Sf'tkr* rr*pitnH»H. A re-
markable abnormal rhythm of respiration, liret ohservei) bv Cbeyne but after-
ward moTV ttlllv studied bv tStoKee, nn<l hence calkt) (iv their combined
482
DKuw*. cic^-iirx ii> cvrtniit |i«lho)o|;ical ca'^8. Th« T^pir^tory monraMtll
^uluall/ ilwresM both in extent and ra|ii(liiy until thev oeaae alUigMlHr,
tod » conililioii or apiMoa. laatiof; it mar be for Mveral loconda, enmMi
Thb it fulltiiTcd by a leeble respiration, succeeded in turn bv a sometrbNl
nirnii^-r one, and thus tbe raepiratiou returu^ ^'raduHlly to the iiortiial. or
may iivro rise to hyperj>u<ea or slijtlit dys|imea. after which it agnia ditK-lim
ill It itintiliir niaiin«r. A secondary rhyltiin of respiration i^ thus ileveloned.
[wriods of iionual or stisbl dytt]tii<i:i« reepirulioa alternalinK l>y gradual
Iraneitioiu witb perioiU of apuoea. The cause of the phenomena is not tJior-
oiighly undertoiid. Whellier llie waninji and waxiu^ of tiie respiratory
movements be due lu cwrnrspiimlii)); rhylbmio chanj^ iu the nuirition of the
respiratory centre it«elf, or in a rhythmic iticrawe and decrease of inhibitory
impulses playing upou that c«nir«! from otiier )iiarU of the body, for instance
from hi)(er i-egiona of brain, hii-i not yet bi^fii iwtllcd. it fre()U{^ntIy a|>pe«T9
iu oonuection nilh n fatty cimdilinn of the Ix-art, hut has b^u met uilh in
rarious mnliulln*. Clowly Mniilnr |>hciiom<-n« have been ubaerrcd durioK
aleep, un<lerj>erfe<-i!y nortnal coii<Iiiion« ; and thL-> fairt » rather in &v»r cif
Uie latter of the !«<> rxphuiiitioiis jiMt Kiven. Tbc phenomena prem^nl a
Mriking analogy with the " groups " of hearl-lxmtM wi fn^uently weru !ti tin
ftox'a veutncle placed under abnormal circnmnlMiooa.
Tub KpFBore or CiiAM(it» ik tiik CoMroHrriox axd Fbbhubi
OF TIIK AlH ItltKATllKD.
S 3T7. The preceding aections have shown ns tliat the renpirntory mechan-
km is arrunged to work satisfactorily when lh<r King* ar« advquaUiIy NUppliad
uilb air of the ordinary comjiwilioii of, and at the ordinary praasure of, tin
atmosphere. We have further seen that tiw mecliaTiiKni can adapt itself
within certain limits to chaugOH iu the oompoaition and prrmuro oflhe air
supplied. Wo marnow consider briefly what takes place when ihoM limiu
are overstepjXMl. The most atrikin;; elfects are seen, wheJi, on account of
oocluuon ot the ireacliea. or by breslhin^ in a confined space, or for other
nwoas, a due fiu|>ply of air not betng obtained, normal respimtion givea
place. thn>u)-h an intennedialA phase of dyspmea. lo the condition known u
as[thyxiii ; ihi», unleaa remedial measurea be taken, rapidly proves fatal.
Atphifria. Aa soon as the blood beoomea leas arterial, more venoua than
norma). tlM reapiratory moreinenta hbcome' deeper and at ibe same lime more
fmiucnt ; both the iii^pinilory and espimtory phasea are exaggeraled, the
supplementary miiMle? »|>oke» of, $ .t-^. are brought Enio play, and the rale
of tbe rbyihm b biirriol. Th4v« effecu, as m liarc aecdi, are chiefly to be
Bseribeil to tbe deficiency of osygttii in the blood.
As llie blood contiimcii to become more and moiv venous tbe rcspinilury
movemoils continue to incrtawe both it) fonw and fnyjuenn-. a larger number
of miisclAi iH'ing cullixt int4) airlioit and that to an inorviuiug extent. Vcrj
siKid, however, it may be obserwd ihat llie cxpinttorv mnvi'mcntM are Iwcom*
tng m<iri; niarkcxi tluin the inipiratorv. Every mUHcle nhich can in any way
aatiitt in oxpirtilion >* in turn brought into play ; and at taal almont alt tlie mufr
ol(H of the Wiy lire involviil in the Mrngglr. Tbe orderly expiratory- mov*-
ntcnU culminutc in expiratory roiivuUioiii', the order and in-|ueiice of which
are obsctirol by their violence and cKlenl. That ihcMe coiu-uUionn, through
vhich dy!piia<a merges into asphyxia, are due to a slimuhitioD (by tbe venous
blood ) of t he medulla oblongata, is proved by the fact that they fail to inakt
their Bi)|)earanee nhen the spinal con) hio' Ix-en pfwiously (lividid Mow the
medulla, though tbey still occur after those jxirtiona of ine bruin which li«
iibuvc the ni«Itillit htiTc Im-ch rcni'ivdl. It '» Uftiml to «|>rHk of n "convul-
Bive centre" in th(> mcdulln. the stiniiilntiuii of which givee riec to these
CORviilsioDB ; hut if wo acoep; the existvnw uf euch » centre wo niuflt at lh«
MUDe time admit that it is connected br the cloNWt 1if« with the nnrmn)
expiratorT diviaiun of the reepiratory centre, tiaix every intervening step
may l>e ooserved belwecu a simple sHf^ht expiratory moveiuent of normal
respiration and the most violent codvuIhou of asphyxia. Ao addiiiDiial
p^Klf that these convuUions are carried out by the uj^ottcy of ibe medulla is
■tfurded by the fact that convulsions of a vrhoUy similar cliaracler are wit-
ueti8«d when the supply of blood to the medulla is suddenly cut itfl* by lij^a-
turing the bloudves^ls of the bead. In this case the nen'ous ceulre«, keiue
DO lnnK*^r furiiishe<l uiih tn^h blood, become rapidly asphyxiated through
lack ul iixy^eu, and cxpiratorv coo vulsioos quite niniiar to tliose nf ontinai^
lUphyxia, and precedea like iKem by a pawing pbaae of drapnixa, make their
Hppearauce. Similar "aiitenaic" convulBtana are seen aher a sudden and
iiuffi low of blood ttom the body ut large, the meduitu hriii}; ximilarly stimu-
lated by the Uwk of arterial blntHl. In ordinary fuintiti);. whidi ia Iam of
COD»oiouiDen> due to an inHuffimnl supiily of blouil to tliv brain, the dlminii-
Uou of blood-nupply in not ^rr^t eDougli to priHlucn tlM»« ■■i>nvnIaionK.
t^uch violent «tiiirt> Hgiecdily exliaunt th« nrrvouN *v#k-ni ; ami thir convul-
■iuna after being maiiitainttl f(ir a lirinf jwriod Dudncnly t-eaK* and aiv fol-
lowed by u [wriiid of cJilni. The oalm i* ope of exhaunlinn ; tlic pupilit,
dilated to the utmost, arv unafltnitet) by light; touching the cornea cuIIh
forth DO movemvut of the cvclido, and, indeed, n« rvflex actions can any
whcTp be produced by tbc Mimulalion of i>ciiticnl ituriaw*. All expiratory
active rouvciuMita bare ouued ; the miwcUv of ihc body are flaccid and
quiM : and though from time to timv tbe respinUury centre galhcn enHii-teiit
energy to develop n-Hpiratorr niovementa, these raemble tboM of ijuiel
aonBal brcalhiu);, in l>eing. as far as muscular nctioni! are concerned, almost
Mitirdy itiKpintiory. They occur at long interval*, like thoff«- nlUr ecction
of the vagi : and like them are deep and slow. The exhniistcd r<i>piratory
MOtre tnkee som« time to develop an iiupirntory explmion ; but thi- im-
' « when it is generated is proportionately strong. It teems as if the
itance which hod in each case to be overcome was coosiderable, and the
effoit in oon«equcnee. when »ucce»efut, productive of a large oHbd.
Very suod, thwe inspimlory etTorts become tees frequent ; tbeir rhythm
beoomee irnsular ; long pauses, each on« of which seema a final one, are
oucoeeded by several somewhat rapidly repeated inspirations. The pauses
beoone longer, and tJie inspiratory movenenta eballower. Kach inspiration
la aeeomiwuied bv the oonuacliou of aceeosory muscles, eapecialty of tbe
face, so tnat each breath beoomea more and mure a prolonged gauk The in-
spiratory gaspe spread into a convuliiivc stretching of the whole oody : and
with extended limbs, and a straiKhlened trunk, with the head thrown back,
tbe nwKilh widely open, the face drawn, and tbe nostrils dilated, the last
breath t6 taken in.
Thus we are able to dixlinguiiih ttiree stages in tlie phenomena which
rtMill firom u continued detirieiicv of air : 1. A stage of dyspniva, churac-
t«riud by an incrMwe of the re»|>iraiory movemeala botJi of iuspiraliou nuil
expiration. '2. A convulnive Kaj^, chaiacterixed by the dumiuanct- of the
E~" iratory effurl.i, iind culminating in general convulsions. 3. .\ >tage of
aui<ti<in, in which lingering and loug-dnivu iibpi rat ions graduatty die
out. Whrn bp>ught alMxil by sudden occlusion of the irat-bca tiuae evNila
run through Ihvir raurse in about Rmt or five miniitm in the dog, and in
about three or four minuua in the rabbit. Tlie tin-t stage pannui grmlually
into iba scoond, convulsions appearing at the end of (be lint minute. The
404
•
iriuuitioD tmm tlw Ncond tinge to the tliiril is eomcwluit kbnipt, the mn
viiliioDfl tiuddfinly oeasiDK early iu the see»u(] iniouie. The roaMOiiii; tini«
is uccu|>Kil in ihv iliinl stage.
The iluruiiiiu uf ud|)tiyxia varifi) not ouly in <lifler«iit auiinals but in t
mmo animul umter ililfvretil oimumMaiieeB. Newly b>jrn mid iroung itninui'
□OmI much longur immcniuii in WKt«r l>efi)rc deatli by aapliyxia occurs thi
tla adulU. Tliu.t wliil* in n rull-grnirn dii^ recovery from drowning h u
lUual nlW OIK.' und n lialf niiiiulMi, m iiewbitni pufipy has l>eea knowu
boar an inini«n<i(>u nf tt* muHi m 61\y miiiulm. Tbe cause of tl>e dilfereD<
lita in tjic tuvt that in iliv i]iiiu^ yuung or r«clier jiut bora aoinial ihe rwjN"
nUiry clmngt* of tlw tiwiiw am muclk luw active. These oonaume Ie«
oxyeni, and tlic fnosral Rturv of oxygen in tln^ bloml haa a leas rapid demand
oiaiu: u|H>ii iu 'uiiB raapimtorr activity of tho ttnuen may aim be Ictnened
by a <kfici«ncy in the circulation ; hvnm bmliw in a Aat« of ayuoujie at the
time wken ibc dvprivalion of oxygen begins can endure ihir liaa fur a much
kxigcr period than can bodies in which tlie circulaliiHi is in full Mwiiig.
There being the aame store of oxygen in the blood in each caMc, tho quicker
oirculalion must of neeeaaty bring about the apoolirr exhauiitiiin uf tti«
otore. 80 alao aiuBsthetias may diminish the etTooU and ilvlar thi- final m-
aults ; Urge doaea of anaalbietics may iirvvcni tho exag:^rniv'l and (ronvul-
nve movements In many caaes of dronning, d««h is biirtcnei) by tiie
entrance of water into the lungs.
By traioiaji;, tbe reepiratory centre may be accustomed to bear n scant;
aupply of oxygeu for a much longer time than UMial before dyspnoea Mts J.
a» ia Been in ibe case of div«r«.
Th« phenomena of alow naphyxia, where the supply of air is gradual!
diiutni»ned, are fuodaraentally the Mwe u» those resulting from a 6udd<
and total deprivation. The annie stagei are Mien, but their developine
taken plaoea more slowly.
§370. DeRdency of air ratultA not oiilr i» a diminution of the oxy^
but also in lui increase of the carb»iii<- acid of the hlood. We haro seei
however (§.17.1), that ibo phenomena of aMpliyxia are in tbe main due
tbe former, and that the accumulation of uaVbonic acid io the blood has
subsidary efli.>cu only.
If the pvrocnuge of oxygen in the inspired air ha increased instead of
diminislifio, tho total prasure of tbe almosplM'rv reniamitig the same, the
partial prtasure of the oxygen alone beiug chiiMgi.fl, »•> marked results
follow. We have alrrady seen (S 85S) that the pcrovniage of oxyjien in llii
ordinary atmospbrrc Iciivph a widi- nmrL'in of oafely. and that ({) HT.*)) thi
phenomena of apnoui are in the main at leasl to Ite explaincti as thr rwu'
not of an increase in the oxygen of the blood but of nervous intpiilw^
aioendiiig the vagus nerves. We have no satisfactory evidence tluit, pro*.
vided the respiratory mechanism is in good working order, an increaM of
oxygen in ih« inspired air even to a whole atmiiapbere seriously modifies tbe
respiratory act ; and it may >» doubted whether any etl'ts;t is produced eren
when the mechanism i» impaired.
$ 379. The conij>u«iiii>ii uf the atmoepbere, tiie pressure remaining the
oame, may be modi6ed by the intnxluciion of fureign itaaee. To some
those tbe respiratory mechanism is indifferent ; for ioaUuee, hydnjgeu raaj
be substituted for nitrogen without any change in the n^piration, pmvidfN
of eouree that the oxygen ia not diininixhi^l. Other gases may produce
poisonous eSecls, either by iaierleriiig with some of the respiratory proceuaca
or in other way^. Thuit carhun tu<iiii>xide, by eombtniug with the hiemo-
glubin of the rcsl cirpii^rlfs. and in preventing the corpOAcliv from acting
a* i>xygeu carrient, |>ru(ldC4-i naphyxtn through deliciency of uxygeii. SuU
lIU
ro.^
RrritCTS t.tr CIIAXOES in ATltUSPRKRtC l-RKSSrnB. 499
phurelted hydrogen inlcrr«rca wtih iii« oxrgenBtiuii uf iht' bliKii) by acting
IIS « reduciug ngvot. .Soiii« f^tMctt uiiilc nlloniiig the onJiiiary rw|>iriiU)ry
chaiigcB of Ihe hlotid ui go on w iiiiiiiil jimdiieo toxic vfle<;U> t>v itk-tiii); uu <>ne
or otner of tii« tis»ui«. Thiw, lu »«? have »wii, iin exctw oi citrbonic iici<l
in the blood taama U> hnve n Mirf-iiil rffccl on the ccotrnl ucrvoua system und
so acl« a» a oarcotlc pi>is<>ii. Tbo pvculiar utlocU of inlrous oxide (Uugliiiig
Ras) BK kiiuiliiHv due 1» the din-cl acltim of the gas in th<; blood on the
i-eutral Dervous syvtcin. Sonw gases an irrwpirnblc and iita)' inlorforv witii
r«s|>initioii, ev«R cstisiii^ sufiocatioD. on acoonnt of ibcir ciuisiiig e]>a«ni of
the glottis, and itiin w snid to be, to a cenaiu extent, the ciuc with an atitios-
ph«r« whirb i* nitollj or largtly compoeed of carbooic acid.
§ 380, The. effetit oJthange» in almMpkerie /)m«>iirf. DiminulioM of prt»-
turf. The piirtial prenura of the oxygen in the iaepiivd air may be
obanf^st, Dot only by altering the coinptMition of the air entering at the
onjiiiary alruoeplieric pre8>ure, but also by altering the total preasure of tlie
atRMwpliere nithout dtangiiig its oomposilion. The rwulta of the latter are,
»howev«r, complicated : we liave then to doal not merely witli the efiecU on
the intercbaoge of gases in the luugsbut with the elfects on the whole organ-
urn. All (lie coinpiioated niacliiuery of ili« tiudy is adajited aiid arranged
to Work UDd«r what we tiiay call onliuary atimirplieric pretnure, that lit to
my. within th« limiUof 7liU mm. mercury at the him level aiid aliout SOU
■niu., cori«8|iondin{[ tn an altitude of <<000 feel, this being lh« range of i>rdi-
nary huniaii dwellings. Any great iiienawc »r deoroaso of preauir« beyond
these liiuiiH will uHi-cl n»l only the vxit of ciirbonic acid from and tlie en-
tnoce of oxygen into the blijin), but, in varying d^roe, »1! tlie phyntoU
and dietnieal proctww of tliR iHKly. A grom instance of tiii" U M-eii wlien
an animiil is MniMiiiIy Dul>j«%t(Ml to ii gritit diniinntinn of pnuwinrv, lu when
it ii" plaocd in the nwvitr of iin iiir-pump mid the reot^iver nipidiy cx-
hiiuKtnd. The aiiimal i:i soon thrown into final €imviil«ii>ii», which are in
part, ttnlflniy in part, due to tlie libornlioD of ga« from ihc blowl wilhin
the biooilvrwels : ihc gas so set free mechanicnily interferes with the circula-
tion, ns by obstructing the pla? of the cardiac valves, or by plugging the
9>niiillt'r bbwdvessels, and thus hdpe to brin^ the machine to a uandstiU,
Tliv free gas found in the vceaels upon examination atler death is add to be
ciomposed chiefly of nitrogen, the carbonic acid and ihe oxygen, which pro-
b«bly were also set free, having been reabsorbed before the examiualion was
made.
But, quite a|>ari from gross efiects of thia kind, it is very obvious that the
organism muM in many oaya suiter from a diminution of prewure. The
ootnpl<-x and delicately balanced vascular svstem id constnicted to work at
the ordinary atinoapheric pressure. U'he toroe of liie heart-beat and t^e
ttmie r >ntriu-tion of the small arleriee are, so to apeak, pitched to meet the
influcnrv exerted on the outside of the bloodreaseU by the ordinary precaure
of th« aimiMpbei^ ; and any great diminution of that pressure must produoe
a Krenler nr law diwir range nirut of ihe vaacular mechanism until it u ooun-
terbalancod by «»niv compensating changes. And a liltle reflection will
s«M>ly many other iiuUnces.
We liiive alri'iidy i-allcd aileiition ($,155) to lh<- fuel thai, the total pK9-
surv of llwr ainiiHiphen.^ n-iiiaining the mine, the iwrtlal pn^wuiviif the oxygen
ill the iuHpircd tur muv he roduced as low aji about ,6 mm. flO per cent.)
without eerk>u»ty nvxlifying tlifl respiration. In tmU-r to attain thin diini-
DHtion of the partiid nrcssiire of the oxygen without diunging the ooinpiuition
of the atmosphere, the total prewure of the ntm'wphcre niti'C be rvdud^l
to the limit of :i(M mm., corresponding to an altitude of 17.000 foct. Now
it is a matter of common cxpcrionoe that in iMoendiitg a mountain "dislraw"
496
is f<'It Inng l>i:furc Hiich nn nltitiid« is re»ched. Tlw ilistreee fell on bucV
Kccnginriii it probably due nut so much, if indeed al all dirvclly, (o ihq
diminution of axyi^eD lu b> a c^etioriil iliaarraageiDont uf ihe orRaoisni ana:
perliiipe more (iDTticiilnrlr of tnc vnscular system. The iiae&-bl€>eiling whicli
u so fre'iiiutiL sn occurrence under the oircumslnDOOs showa that tlie minute
bJoodvcwelg more directly expoaed Ut the diiuiuiitioD of ureasure are pro-
foundly ndected by it; and what is true of ihetu is, protmbly, in vuiout
ways Kud to dilfereDt degrees true of (he wljole vaecular aysIetD. Tbr
breath iea&uess which i» so mnrk«d a feature on Ibese occasions aeenuduB
not BO amcb to llie faot that the blood which reaebca the respiratorj
nervous ceiitreo is deficient iu oxyp;en, as ui the fiii-t that the tmubkfi %-u-
cular Hvslein fails to deliver to those iviiItc!* their bluod in au adetjuato
(Wiioii.
It h a featnri! of lliv vnitculiir system, and indewl uf the other raechantuu
of the body iu wbtdi ticrvoun factom iiitcrVMic, ibitt tbi^v pmaoM the powtr
of tida[>tiij^ themtolves to chnnged couditioDit ; and il> it t» ircll known, ibe
huranu urgiinium somewhat nipidly become* accuMlomtyl to thvr*^ mtxlrrale
altitiidf*. Priiciici^ and custom have liir IcMcffcd, thi>ii"h ibcv havo some,
on the more funilamcntal nnicrKsn depending nn the net mil supuly uf oxvntD:
and it in al the extremis nllitndcs, where in tiddilion to the olDcr trmiDwe a
deiicicney of i<xyg<-ii delinitcly innki>» itficif felt, that the body teetat to &U
in adapting iti^lf lo thi- new circiinwtjiiices!
The addition uf th<'>c trotihU-e not directly rcspiralory in nature, when Ifae
«upj>ly of oxygvn i* dittiinished by n diminution of the total pressure, per-
haps explniiis wliy thoui^h an adequate lowering of pressure will pruauce
n.-^liyxin. that nsphyxii) is somewhat ditlcrent Irom the ordinary asphyxia
duo to (leprivnliun of nir or oxvgeo. Convulsions which are easenlial to
ordinnry asphyxia are at times wholly absent : the nervous svsteiu uD«ler llio
peculiar conditions does not respmd to the alimutus of the lack uf uxygen :
and other nervuus symptoms, suoh ns a rapid onset of (eebleneas aaiuuuting
almost I" paralysis, are apt tu make tlieir appearauoe.
^ 38t. TliK efferJn of imrm*! n/ atmo*phmo jirvjvure. Theae ara Id many
ways remarkable. I'p tu a jirewure uf several utiuoapberea of air, the only
tymptoma which pr»eiit thfmM>tv<st are thm somewhat reaembliii;; imrcoUc
poisoning. The animal become* aleepy mid ntupiil, the result pirdtahly not
so much uf rwpiraloiy chauen, at of the elfi^^to of the incrctUMil prcK>iro
on the whole orgtmiam to which wr hnv<- juHt alludtsl. Al n prrwiiirv. how-
ever, of IS atmospheres of air, or what amouutii to the snnie thing, of 3
atini«pht?rvv of nxy^n, and upward, a wry reniarkahio phenomenon pre-
tenlN Itself The animals die of asphyxia aud convulxions. exactly in tbe
same way as when oxygen i^ (kficient. Corresponding with tliis it is found
thui the production of carbonic acid is iliroinisbed. That is to sny, wbea
tJiciircmnre of the oxygen is increased beyond a ccrtcin limit, the axidattons
of tile body are dimmished. and with a ttill liirther incressc of the oxygen
are arrested nllogcther. The oxidation of phosphonw is jwrhajM analogous ;
al a high preesure of oxygen phospborui^ will nut bum. Not only animals,
but plants, bacteria, and orgaui;r«d fermi^nts, are similarly killed by a too
great prcseure of oxygen. J^m
The Rblatio!« op tbb Resviratoby System to the Vasodi^r ajto-
Otheb Stbtemb.
£ 388. Slany events iii the body show the inllaence which the respiratory
moreineiit* exert on the cirrulnliun. When tlte brain of a living mamnuu
ia exponent by the removal of the >kull, a rhythmic rise and fall of the i
RKtiPlHATORT VSTtVLXTlOSt.
m
i
P
bral luHse. n pubntion of lh« brain. iiuitcdistiDct frocn the niuvvmi^nts cKuiisl
hy the pulse in Ihe arleriM of ih« brain, is ob««rved : and ii{>iin vxAmitialion
it will M Ibund tliai thcsr iiioveiii«>te niv synchronoiia with tbe roepiniUiry
jnovem«oto, tlie brain Tis\n^; np diiriii;! exptrntioii and BJiikiiig during impiri-
tiou. Thty disnp|>ear wbeii the arlerioa >;'>■■>!; ^ the brain arc li^Muraa, or
wlien Ha Tenoua sinusea of the dura ranter nre litid opcii ho ai lo admit of *
ft-ce escape of ihe venous blood. Tbey ei-idontly arise fraiu tbe exptratorr
■novements in some way hind«ri»K aud lb« iDBptratnrr iiiovemeDU aansting
lh« retuni of blood from the brain. We have already ($ 116) atat«d that
(IuHmk iniif'iraiion t)i« preasunt of blood in tbe great veins may become
iic^tive, i. r, luayHiiik bolow llie pressure ot thealmoapliere; and apitneture
of one of thftv veins may cau»e dealb by air beinft aciuatly drawn inio ilie
v«in aud lliu* into the bearl duriiijr an inspiratory imiv<'meiiL Wli«ii tbe
veins of iin animni are laid Ixira in tbu d<acIc and watched, the M-i-allcd
puUuf ■<c(ir>riM may be obiq-rveil in them, that is, tbey swell up diirine cxptra-
tion and diminifh again during insm ration. And indtiil a little vonsiduralion
will i>bow that the i>xiMn»i<>n nnii c-'intrarlti)n of tin; ch(^Kt must have n da-
cided effecl on the flow «f blood (hroiijrb tint tbornctc pfirlion of, and Uiub
tndirevtty od ihnt through tbe whole of. the vi^culnr syslvm.
This is well illustrated by the cffbctf of re:tpinili'in on artcria] blood-
E rewire. We hare teta, while Ircnntig »f the cireulalion, that the arterial
lood-nreaaure ctirvee arc marked by undidntionv, wbicb, since their rhythm
IB synehronous wilh ihal of ibe respiratory movrments. arc evidently in some
way connected wilh refiplration. Similar uudnlatiuus may be obsarred in
ibe pulse tracin;^ taken from man.
na. iiT.
''H^
/
OuaraaiMS ur HuxM-ramL'ss Ctsvk yum Cvim ot Iitsa'Twism-k: raiwc-KK. iDcn.|
|b ilio bi-wd-pif ur» tvnv lak*n tv mutiuvl » monun cutuoiDclar : ll tliaw* Uw mplniUin'
sUlluio. iho tUfmr totit ini ihv 4««rmi( Miis tkt owikcd. A I* iho r.itciv vl lnln-lliiin«Ii'
I wum DliUIiwl by nitinoi-Uiiii one tlmb of ■ m* oometcT wllli llic ^Iciinl «(li)'- Ii»|iLr>Uon
liaslatsl '■ *i|''n>ll4ni>t (. Wllb Ibv Ufliwliig <>r liM|il>aliiiii I'l IIicviiaiUiliMut Itit rhnt niiUH
■ ■■ubnl fitllnr iliomcrmiry iDiliflntn-tlionclcBMiiomalct. bui tlistdkctseoiidlinliilibak ■Hue
ilMtMHOiHS ■■riiiir»iliiimiilc|«aBim4cm nol boMon Ike luinomeicrtlMiKlMuai Ihaluntsabn;
•ikl 4* Um ioiigt •vpAiKl moTvaadmanilwiklUiiilifaianriiry b«am«ahnt mkI Um until tovard
llinfnitt iif Iwttfiitliiii llii> <«rre tcoonicsirBp' lunrlykitnlglitllnt. Cvnveatlr, Uta ivluni or ilir
ulwtl M tbi> l*i;li>n|ii|| of aittmtian frt |>p»liinB nl nnt ■ mitrlud riic of (bu incnurY In Un
Wlliininliir tml Ibli «nn mw« u Uw ulr Iimvm ttio ah*(l 11114 lb* Iiiiih* (brliik, iiti«nn>pan Its
curr (kill ilonTf.
Wlieu these uiidnlalioiia of the blood-pnwMiro curve arc compared care-
fully with the rospirntory movements ur with ihi- variations of inira-lliuracio
jireasure. what is most commonly olwcrved i» Ihnt while the blood.preaaure,
uu the whole, rises during inspiration and falls during expiration ueitlier the
nae miir tbe fall ise.taclly syncbrunous nith either iiupiration or expiration
Fig. 147 fiGSwCTwRiiiciiig* from n ilog lalii.-ii nt the MinMiniT^w^ir
■be ordiDiiry blooll■]'n'i•^klrl,■ cunv Inm ila- ciiixitid, iiml llic olhrr. h. «[ ^
fCnling tho <-oiu)iltt'n tij' itic iiitrii-tbunioic prni#urc ■« obUinnl by carvlml^
bringine n lunnomtlcr iiilD oniini-i-tKiii with lli« i>kuni] citvitjr. On rom-
pnriiig lilt? IV.XI curvet it ie evidvot ihui ucithtrr the rue nor the ftitl of nrlcrial
prmiirc cniiicitli's exactly dUier nilli ii)«pirnl)on or witb cxpiroiion. At
the beginning <ir iti^pinitiun (i) the nrteml pirsiuro ia seen to be falling ; k
euoti, lionevcr, tiegiH^ to rise, but dura not reach ibc iDaximiiiu until son
lime alVr cxpirotion (f) hui begun ; the Inll eontinues during the rvniaiDder
of expiration, ond psffcs on into the eiieceedtng inepiralion. Tbis mgRetU
the idea ibal, while inepirntiun Und* to iner«aee and expiralioD to diminith
the bloud-prceeure, there ore causes at work which in each oaae delay ibe
effect.
Extended obscrTatioDB. however, thaw that such a reJation oj that ilioirn
in ibe dgure, though fre(|ueDi, is not coosiant. in fad. Ihe eOecm of ilic
re«iiiralory movciuents on blood-preasure are found to vary %-eiy widely
according as ibc reapimlion ia <juick or slow, easy and »hulluw, or Inborn
and deep, uud eapednlly as (lie air euiera into the chat readily <ir with diffi-
culty. Mcreuver, rt^piratonF' undulalioua of blood-prewure are teen not i»nly
with nntuml but ulso with artificial redpiratiuo ; tn tli<- Intlcr the ntccbnnini'l
cundilion;! are to a large exicnt the reverse of iho»e of ihc fonaer. and might
fairlv be csjicctei) to aftbci the circulation in u dilTerent way. The (-iui'>.iti>)<i
of IIKVC rcvpiralory uudulationd iw, in fact, complex. Tbr rcspiriitorT act
allccUt the vuNeuliir »yiitt>ui iu nevrnd dlfierent way>. and the eenernl elTect
vanes according a* one or nlhcr indui-nce is prrd»minunl. The«? several
nction« are suffi<'i<-ntly intertvlin;; nml inigiorliinl tu doterve discuMion.
§ 383. TIic heart iind gn'nt hloodvi-Melii arc, lilce ihe lungs, phiccd in the
air-tight thoracic cavity, and are subject like the luiigK to tho pumping acttoa
of tho rotpiralorj- movement*. Were there nu longs present in the chest,
the whole Ibrcc of the cxpitneiuQ of the thorax in inspiration would be
directed to drawing blood from the extra- thoracic veBselti toward the brart.
and convereely in espiraltou the eHect of the return of the thorax to lU
previous dimensions would be to drive the blood thuB drawn in back again
iWini the heart toward the extra llioraeic veasela. And, even in the iireweo
of the lungs, some of tbis clTeet is still felt. The main purpose and tW main
result of tne expansJon of the chest in inspiration is. of cuune, to draw nir
into the lungs; oy that expansion the air in the puluionnry alveoli is rnrilied
and brouuht to a lower j)n?S£ure than that of llie alin(«]ilicrc outside (be
cheat; and the diflerence of pressure thus set up lead^t to an inni<h of
inspired air until an ei|uilihrium of prcasure is esiabliohed l>ciwccn the air
in (he lungA und thai outside the cbetit. Ilefore, however, ihe inspired air
can Gil a pulmonary alveolus the eloMic wallti of the alvci^lus have to be dil-
tcndeil, and that disteunion i« ellecleil by means of (be prtmure which caWM
Ihe iuspircd air to ruter. Part of the atiuMpheric i^imvitre. in fad. whieb
cauMa the entrance of (he air into the lung is fpi'm ni overcomiug the elas-
iicitr of (he pulmonary panagt* nud <*lls. So tlmt while by the inrush of
inspired nir the diflerencc of |)re9>t>ur« between the nir inside the puInioDatT
alveoli and ibnl outside (he ('h(x(, brought about by (he thoracic expanaioa,
is completely neulmli7<ed, ilie difference between the pressure to which the
parte lying within the thorax, but uulsidc the lung«, are exposed and (hat
ou(eide the chest is not to completely neutralized. The pressure on these
patU always falls short of the prcMiire of the atmosphere by the amount of
pnature ncccstary to counlerbalnnco the elasticity of the pulmonary nnsBa|[W
and alveoli. Conse(|uenlly, any structure lying within the thorax, nut out-
side llui luug6, is never, even at the conclusion of an inspirativii, when lbs
I
»
MTo lilli'd nrilh nir, eubJMt to a prcMiira m great m that of ibe iiltnM-
|>lM>rt<. Ad(I, Mnc« Ihe rraotion of lli« nlnxteph^ric preMura which is ihut
e|>«-tit in distcmting tlw lun)^ inPTvasps »» ihe lungs boconie mi>r« imii tnoro
sIreteWI. it follows tbal ihc t'ullpr the inspiriitlon tile i^renlvr i» th« (lilloivnM
brlneen the prentiro on strucltired within the thorax, but nulside the luna»
and the ordinary prenure of the stmoephere. Now, w« hnvo seen that ibe
prvaiire nwenarjr (o cauiiterhalcnce Ibe elasticity of ihe lun)^. when lliey
are completely at rest (in the pause between expiration nnd inspinilioD), is
in niaD about 5 to 7 mm. of mercury, and that uhen the lun^ are fully dis-
tended, as at the end uf a forcible iuapiralinn. the praeeure rma to as much
aa ;10 mm. of mercurv. Hence, at (he hei^'hc of a forcible inspiration the
trei^iire e.terled ^^u llie lit«rl and ^reat veeaela within the thorax U 30 mm.
»i ifaau tlic ordinary' atnioapherii^ preraure of 760 mm., and even wlien Uie
chnt ia comjdetel)' at reitt, nt Ihu end of aa expiraiion, tlie preaiuro on the
bean and f;reat VMm-lit i* «li;|chtlr (by iihnut 0 mm. of mercury) below tl»l of
the atmoatibere. We may a<ld tlial any ultntitcle tu thu free ingrMt of tlie
iuipired air, any diflicully in the full cx|>itii»ii>n of the pulmoiinry ulveoli, of
course inrrai»*=i tlw negative pn-wiire to which the thoracic iitnictiiri!!( out-
side the luugTt iirr Mibjrcted by the ex|uin»ion of the cbwi. Hence, when
tlie (rueh«a it clawed a vi-ry lar|2c jmrt of the thorado expanHiun in directwl
to inereiuing the nc^tirc prcMure around tbe heart and grmt blood veMM'l:*.
During an inspiration, then, the preniirenrnuiid the hejirt un<l gn-at blood-
Twwla Itecomn conttdembly lf«e than that of the atmovpherc on the ve^M^la
OUtsUlo (ho tborax. During expiration thin prcuure returns tonard that of
the atmoxpherc, but in ordinary breathing never quitfi reaches it. It is only
ill forcible exjiiralion that the preMnr« on the thoracic vascular organs
iH-acltw or cxece<l» that of the atmosphere. But if durinc inspiration th«
pmwun.- iK'arini; on the ri>;ht auricle and the venie cav-v become Iceh than
the jmtwire which » bearing on the ju((tdar, subclavian, and olh«r vmita
outMde the thurax, this must result in an increased tlow from the latter into
iIk- former. Ilenc*. duriuK each iitut pi ration a larger (juantity of blood enters
ibc right side of the rtKht, Thia pruliably leada to a Htronger stroke of the
htarl. Biwl at all eventii eausea a larger <|uantity to be ejected by the right
ventricle ; th» causea a larger quantity to escape from the left ventricle, and
thus more blood is thrtiwn tnto the aorta, and the arterial pressure propor-
tionalely inoreawd. During expiration the converge takes place. The
ErtSRire on the intra-thoracic bloodvnuieU ivturns to the normal, the How of
lood from the vein* outWKle the thorax into the venie cavat ami n^ht auricle
is no longer asaisied, and in r<>iiMt{ucncc Icm blood passefl through the heart
into tlie aorta, ami srti^rial prvssuro fall* again. During forced ex pjrat ion
tite intra-tiiofocic prt-wuri.' may tw so great as tn afford a diatiiict obstacle to
the Bow fhun the vein* into the boarL
The e&evt of the r<vpir»t<>rr movcnienls on lh« art«ri«« ia naturally ditfer-
ent fium that on the vcinv During itupiralioii the diminution of ]ira«uro
ill (be tlM>nix around the aortic airb tends to oxnaml the aortic arrh and
thu« to che<rk the oiinai^l tlow of b!«<>d and to dimmish the preasun- «if bliiod
within the aoitii. During expiration the increase of nreMUrc outnide the
aortic arch uf courra tends to increase also the blooil pniwure \vithin lh«
aorta, acting in fact jusi in the same iray as if the coats of the noria thcm-
Kwlvm contracted. Thus, as far as tulcrial blood-presaiiro it conccmerl. the
eflectif of the respiratory movements on the great veins and grent arteric*
re«[>cc1)vuly are aiilognnistic to each other ; the effect on the vein* being to
increase arterial iir(«»ureduring inspiration and to diminish it during cxjiini-
tioii. while the Hiect on llie arteriw is to diminish urterial pn^««urc during
inspiration and to iuerense il during expiration. But we should oaUirally
«ESI'11tATI0K.
ez[>cci [he effect on ilie lli in -wallet) vcJti.t to be greaicr Ituui (hat »n ibi^
etuul, Uiiclc-Kallvil arivrieit. su luticli »o thnt tlw direct tftvcX ud i1i« arteri»
may be ueKlect«<l. Tliul is u> Har, we sbould expcd Utc bltwcl-prcMun' Ut
nM during iiiiiii>i ration ami to lull liuring ex{Hration. Tht*, n» wo havr k<ci).
i« frcc|iH-iil1r tati cane, um), iixli-eil, n)i«ii llie breathing i» decii nitd laborcK),
wul M|>c<rinllr doriiig violent and iindili^n rexpirittary inorciiie»t«, llir inHu-
encn in tbi» (lirvi'lion on the blood ■]in-».'>iiri: mrve of tlte piimpitig ncttoa of
ll)D chmt in nninislaknldc.
In iill<ini|itin^'. hnwi^vcr, to Mitniate the tffeci itf the roipimlnrf' ninrsoMnib
oil blwHl-pn'-wurr tvr mini li<-iir in mind wbiit in Inking pince in tlie sbilooMfi.
In iiispirniioii the dtwi-nt of the dinjihrogm <-»ni|ircHiv tUc nbdominti) viscera,
and M, wbik' iil tlir vcrj- liml it drivw ■ nuiintity of blood onward niong tbe
infei-ior vonn I'avn, i>iili»c<iu<MilU' binder* the upward How from the abduiueo
and lowrr limh«: nt llic nimu- time, by (^miprvMiog the abdominal aorta,
it tend« to niisc the prfwaurv in ibr thoracic aorta and its brnncbea, while
lowering thai of tliv abdominal iiorin and its branches. Tlie elli.>ct of bmt
expirntion would be the I'onversc of lhii>; but in forced expiration tlw
prewure of the contracling abdominal mu»clc» would, as tu ioapimtion,
ant tend to drive the blood onward along the vena cava, but aiihsequentlv
to hinder the flow both along the vena cava and the aorta- The ef^-t of the
abdominal movements therefore U mixed and variable, and tbcir indiKUc*
on the blood- pressure in the femoral artery muBl be dilferent from thai on
tbe radial arien* or other branch of the tborado aorta. It w difficult |a
pre<liel what in nil cases the eflect would be ; and the matter cannot b« mi-
llod bv el i initiating' the movetnents of tbe diaplirai^m throiiifb iu»ction of tba
phn^nic nervL-o, HJuoe iu such a case the nlii>k- working of the rej|»irauirf
puiup i.4 tnaliirially all^cie<l.
^ 3M. Ill nddiii'm hi the tnAiu-ncv thui> exerted by thiMhonidc tnoTenenU
on thi- great vein» hiding to and the grvat arteries leading from the heart,
we have to consider iho bohavtor of tho pulnionarvrci>scl» iheniselvea under
the varving thoracic prnaurc. ThcM. like the venai caw and aortn. tend
to expand un<ler the influence of the inspiratory cx|KiDsion of the che»l. and
thuH to becotnc fuller of blood, very much as they iraiild if the whole btiif
were placed under a large cupping-gla». The lirst elfect of tbit incmwoa
filling of the pulmonary vewels would bo to retain tor a while a ivrtaio
(|uaQtilj of blood in the lungs and thus to lea»eu tbe amount Inlliiig into thr
left auricle. But this would be temporary only, and tlie widening of the
pulmonary veseelsnotild speedily produce an exacllv contrary elTerl. naiuvly.
an increattcd flow through the liinKS due lo the diniiuiohid rufi»taiico oBerod
by tJie widened pasmgn. Convenicly. the first etlect of ex|>inttii>n would be
an increased flow into the leH auricle clue to tbe additional iiuiiniity of blood'
driven Mivrard by the partial colla{K)L- of thr pulmonary veMrl>, foliowot by
a moT« ugnifieiml ilimintalieil floiv cauoiii Iiy the greater nvixliince niw
olTervd by the namin'or vascular channels. Thii5 lh« d&ct of iiinpiration in
ii»i way wouM Im' tir>l tti diiniiiUh the flaw into the left auricle and «u itilo
the It'ft vi-nlricle, but nflcrwnrd, for the rent of thn inn pi rat ion until ihe
beginning of expimtion, to incrtiuc the flow into the vcntricio : while con-
veriely the cH'rct of exiiiratioD would be linrt, for a brief period, lo increase
and aitcrward, during tlie rret of the movement, to diminish the flow of blond
into tl>c left ventricle. Further, while tlii« may be coniidered as tbe efleet
on tli« pulmonary vtMola, large and small taken altu^ther. the inHueuoe
both of tbe thoracic negative pressure durinc inipiration, and the return iu
a positive direction during expiration, will near more o« (be Ibiu-walled <
iHilmoniiry vrins than on tbe stouter pulmonary artery; that is ta mj,
■DBpinitHHi betxNtiea eelnbli»lied, there will be » dimiuutiou <>f pressure in l]i«
BBSPln^TnUY I'SDCt ATIOXH.
601
pulraooary vdns crntlcr tliaii tbnt in the pulmonary »nm-,aiii) l)iia nill Iw
SD ■italilionnl iiilliionre fnvonnt; the Row iiilo Llie l«fl veniriric; during
vxpimtioci a nmilar tlitlerence nf offecl will be fell iu the contrary direction.
Daribfi the incmse of tinw into ttie ventricle the- 'lunnlily of blood ejected
tt «*cflflroike itill incrcaMt. and each stroke will (^ 1<>2) be incrensed in vi^or,
io eoaM()a«-nce of ubich tlie arlertal pnMure nill ride. C'onverwlr, during
tb* decraue of lluw into tb« ventricle ibu iirteritil jirewure Mtll iiill. IIcDce
U>e mienil eHrct of tbc n>iivvmeiil« of ibi? cheut un the piilmonHrr veesels
will be duriu): the )>f(tinniti){ i>f impiiiilinn tn i-cmiiniiu (he luwcHn;- ot' arterial
IWiwarr. whtL-h wu taking* (iIiicl- ihirint; t^xpimli'in but [iubiie(|iieuily to raiM
llwarlprni prvMiun-; nnilcunri-Rii'lyiit the lieginnini^iifexnirntiuu tOL-oDtinufl
Umrinof arlvrialpnamtn' uhicli wiu (akine place; ilnrioe iii.ipinition bulsub-
K^urntlr til Inuer urlrrinl prcfwire- In i>ri1innrv brcuthing, hs ir« hnve tteen,
vbal tnay be conwdcreH n» the normal ivlntiniunf blood-prettture lo llie ntpirn-
kry tniiTpninibi ar« tmciwly of this kind.
i 3M. KflW'ti* of lb* rw|>imtnry morements, however, are •ren n<ii unlv to
aataral but abu in artiticial re»prmtioii. When, for instance, in an animal
■uxtvr urnri. anilicin) is substituted for ualural reepimtion. undulntionn of
thn bIt'Ni-pnwure curve, syni.-hroD<ius vith the reflpimtory ninvcnicnbi. iirv
anill olwem-d (Fi);. 14?*), tbough generally leas in extent than th(w« Ken
timlrr nnlural conditions.
NnH in uriiticiat reapiration, tlie mei'lianioal conditions under which iho
tirw'ic viscera are placed a* re^urdj^ proii^ure ai>e tbi- exact opposite of ihoM
during natural respiration, for nbcu air is blown into the trachea lo
Uw lunga, the prea»ure >viibiii tbc clieH ia increased instead of
^linitntshed. Under theae circumstances, nppJving the conaideratioiu Ittul
tittnn in the procedint; paragraph with regard to natural respiration, we
■Inuld ex|)eet to find that while the firU eAict of an artificial inspiration
Kid be to drive an additional ijnnniitr of blood out of the litrign into the
rtotiicle. nnd (hue to raimt arterial pressure. thi» wotdd he in turn fol-
>d by a fall of arterial pnvsuro due to the incrmsed rvsintaiice olfered
both to (he pMMge of blood thn>ugh tho lungs and to the entrance of blutxl
ihmugb the vtam C9r» into tbc right auricle. C-onverscly, tlio eflcrl of the
Mcvecoing expiralitm would be an initiiil continuance of the lalt of arloriiU
proanre tuenxded by a rise. In other words, we should expect to lind In
artificial reapiration effects exactly the reverse of thuse which we lind in
aortnal reapiralion : and inde«<l in many curvee of blood' pressure taken
4uring artihcial reftpiniiiun tbi« is die am.:
Bota in natural and in ani^cial respiration, however, the featurw of tbc
Uood-pmmre curve vary iiccordinfc as the breathing is burrie<l or flow,
■halhiw or deep, aixl according to the facility irith which uir enters the
cbf»l, so much ao that at [iin«) ibc bloo<)-prewiur« curvea of natural and
arttticial reBpiruttiui may cluiely memble each other. And a liitJe nio-
•ideralicn wutdd leud uv to cxjiect thii.
We ha<re M«n that the nw in arterial pri-»iiri- w hich miirk.i the respiratory
undulalion is in tl>e niatii due tn a leni|Hirnry greiitt-r amuiint uf blcxxl thrown
into the aorta by tbo left ventricle, nnd thai r<irr(v|>(mdingly the fall nf pres-
sure runiplrting tbc undulation iit in tbc nuiin due to a tcmjiorary lessening
of the amount hi thrown. Though the cnuN-a diwuKwd in ^ Sl^:! undoubtedly
make iheniiwlvw promim'nt in liibowi nnd vioK-nt renpirntory movements,
«t may mficlude that in onlinary n-xpiratinn. Ixxb natural and artificial, the
main rvent* prodnring the reiipiratory undulations are (bcwe di»cti»MHl iu
f asi. Wb may restate ihe conclusions of that dismasi'rti by *aying that the
ropiraturv inoraments alli'ct iIh- aniouiii uf flow of blood into tbc left vidi-
t Ml the disebnrge of blomt fDiti the loll ventricle into the a>irta, in
two inniti nnrs. In tliclimplHcc. ihnnii^'h [li>' widening; w narmwini; of tK«
pulmoDnry vcMt-ls tlioy nller llic (Mi|Micil y of tlie vc^ttels to liolil liluinl fur (lie
lime being. Id the spcoml place, in ci>iKei|ueiice of ihe diUvrenru of n«i«t-
anoe, flccntttoneil by the uidening or narrowing, ihey alter tb» mic vf Haw
ihrnogh tbe pulnuiuary vewel§. The tint fuclor id m bnof nml |wiMtiug titM;
the extra room due to widening ia souu tilled np, tbe namiw«d v<r(McU mmb
dbcIiiu-Ke the (luantitr which tlieir can no lun^rpr bold. Bui lb** awotij
factor u> It more lajiiiiijt one; ao louc tm la tlw rtwpinilory iiinremtot
ibe vumcIh remain wideneil or narrowed hu loog u the iiite of tUm
incTCAK'd or diminished. Theiie two fuotdry pnidun- "ppiMie eflrclB, ami
hence tb« the lolnl result of any imrtJculsr kind nf rapinition ivill (l^wo<l
on tboir rulntivi* pro mint- nee. Wilb 'jijicklv m|(e«i«l re«pinilory more-
Dieni» tbf lirat foclor t-oni.w to tbe fnuit ; ivhcn the n^pirol'Ty nioveiMSti
are more slimly n-.iii-iittd and niorr «lowIy cnrricd ant tbp eecond faelor ii
the nior>- polctJl. Hciin* it cnnK-M ttl>iiut thai io 'juickly lepciiled ani&cial
respimtioii whvre the fint factor is pmloininanl. and the prominent eSM of
each intlalion i» Io diminii'h tin- mpiicity of, iind eo to empty (lie pulioooaiy
veiidet" and to iiici'riv»c thti H<iv intu ilie v-niriclc whereby (he pmsura riici
in inllitiion, tbnt ii in in»pimii<in, (he liloo'l-prtMiire curve Miniubi(«8 that of
a slovly repented naliinil rc»piralion, where ibo pressure aUn riaes iu iiu|)ira-
tiou, Imt where. llu' second fiiclor liciiii; predominaui, tbe rise of preimire
bn>iit{bt about by each iniipiraliun u due muinlv to the more rajud How
throuKli tbe widened pulmonary veaBol& Aad oilier illuatmtiitiu uf a like
kind I'ouhl be pvcn.
S 386. Ilesldw tbe mechanical effccU of tbe respiratory raovements tbe
vucular system ia influenced by refipiralion throuKb the changes iu thv gaxa
of tbe blood.
rhanf[«s in tbe blood may aSect on the one liand tbe vaMtootor syatcm
and on ibe other hand tb^ heart. They may further afl^t the bean ritlier
directly by acting "n the cardiac lissuen or iiidins;tly by means of iW
inhibitory and augmrntur cardiac nerves. They may almi probably affect
tbe inTipbcnil v<»cls, nol only tliroiigb vnwmotor ncrvec hut by acting
directly on ibc watU of tbp smaller vcmoIi!. Wc hiire indications of an
action of mpirali<m on tbe cnnlio inhiUil»ry tnlvm, even io normal iniei
respiration. 0nc8lrikincf«aluro of the r<9pi1-.1^>^y undulation io the blood-
preoMire curve of the dog* is the fact thitt Ibn piilw-mie n i)uick«D«<l duriuu
tlie rise of the iiiidulatioii and b<-i'<imcJt ^ilovter during Ibe fall ; see l'i){. 147.
A similar influence mny be *ven in piilse-traciugt taken from ntau. Tbe
quickening of (he beat might be coni^iitcml a» itself partly accounting fur
Uie rise of prcaaiire.or on the other hand it might be urged tlial the incretued
floir of blood which craiuee the rise of preraure. at tlie Miuie time leudti to ilie
r|uickening of the beat, were it not for one laol. vis , that the dtHerL'uce is ai
once dune away with, without any other essential change iu the undulatMnw,
by iieclion of both vague nerre?. Evidently the slomer i>ulse diirin;; llie fidi
is ottued bv a coincident stimulation of the canlio-luhiliitury centre iti the
tnrdulb oblongata, lh« (|uieker puliw daring the rii»e being duu to (he fact
thai, duHng that interval, (he centre U cuinparalively at rRC Wu have
her« iodioatious that, while the respiratory centre in ibe nicilulla obluogMa
is at work, sending out rhytbmli- inipiiUea of iuspiration antl cxpiraiioo, tb*
oeigtdioHii^' card iii-inhibi lory i-eutie is. lU it w«re l>y :'yiu|>athy, (brown Eoto
nn activity <d' .luch a kind ibal its intluirnce over the hciirt wa\i?i with vadi
cxpimtiou aiiil wanes wilh each iunpirution. We cannot a^ yrt explain
eiacllv the manner in which (he a«.'livitv of the <>ni- eculrv iiiflitiiices thai of
> tnUienMU.tlMrMiiimioij'uiKluluIlonsiliniish wallnurkisd. p(«wM>rtrrtinnndlKcHnM*t
fuk»-nlo la tb« rbt knit Ikll.
BKSI'IRATOHY UKDUIATIONS.
903
it may Ite tlmt <)iirint; iIk; uxpiratnry phtutc ibe blood Kockln^
In H aai i]iiit£ «> vrell urUiriiklitwl, tupttcially m fur tu the vsciipc of
vnrli'inic add it oiiccrnt^il, lu during thv inipirnlitry nbiuc, kckI tlmt the
4:nr(lii>-inlittiitc>ry criiln; in Kuffidi-ni))- xen^tu'e In ii{»]>n-c)ste lh« ilight differ-
vac«; Imt nf thii' wc ojuinol be miro.
j 387. Who4i tiiroiigh iiili>rt«rcncp >r!tli tli« ptilraonnrv iiilcrchKiig« the
lilmxl cnit out from tbo l«ft rvntricte bi>coinm nnd continiiM to be toM
MrU'Hnlixeil ibnn nounl. the oITl-cU oil both the Iicart and the vjuoiaotor
Hvalvm bwome conipicuoiii. The rhythm of llie hc^arl-brats is moat distinctly
sluncil. This, under ordinnry circunutanotf when the vn^jun nervei are
intflcl, hi probably in pnrl the r«siilt of vaRUa inhibitiot), the veiioiia blood,
a? sUK)jeet«d above, atiniulntiii;^ llio cardio'iiihibitory centra in the nieditlla.
But toe frlowio); in not vhoily caused in this way, for it is Still moHpicuous in
■u animal placed under nrnri nnd with both v:i>;ii^ uervea divided, dunpare
currea 3 and 4 with I and 2 in Fij{. 148. How thia slowiii); 1b broiidu nb»ut
b not very clear. When veuoiu blM>d is sent thiuugh an excised lieurl, the
beat is, it b true, slowed, but it is aUa and still more oondpicutmsly weakened.
N'ow vrben the blood becomes too v«uou!i, a.-i is shown in Fi^. 14J*, «ven ntler
the action of the vaxuH nervett has het^n eliminated by suction and als>i by
urari, the alonini^ is out of pri)|M>rli<m to tb« weakotiin/. siuoe, as we shall
presently see, the blood -preMtirf rif^ea ; and though that rii«e i.-> ohi<-tly due to
vaaomoitor coiiMrieiion, still it could nut (uko plac^ if tbc cardia<- slnilcc v-hte
T«ri- notably WL-aki^ned. It may W thst the vvnous blood miniulaCes the
cvuxliu! augmcnfir uK-chnuium in such a wav lu to brint: ab:iiit an augmenla-
tioii of iIm' canliar iilrokp rather llinti a quickening of the rliythm ; but thi#
luu not l>cen ddiiiitdly provcil. In anv caiw a slow biiil, with such a inain-
tMMUoe of til'- Kirctiglh »t' the cardiac strokm as permit« Uie conliuuauce for
•ome COBslderahlc lime of a high b1ood>prM8urc, i« met wh«D the urlerinliut-
ti'M> of tbc blood is inu'rfcn.-<l with. S>oDer or tat4.T, howov«r. the delicioiKy
uf oxyiieit in tbo blood diminishes the store of cxploeiv« compoands in tbft
canliiiv rauoculor Bubttanoc. ihc ImuUs lawn ID force, oAen showing a 1«4U-
porarr increase io fre<]uency, and won become Irrogular.
{i 3s8. The etfects of deficient arlcrialixation on ihc viuomolor »yst«m are
well shown when in a» animal placeil under a moderate doee of urari so aa
to eliminate the complications due to contractions of the skeletal muaclea,
with both vagi divided so as to insure the elimination of inhibitory impulsm
from th« medulla, artilicial respiration b BUS|>endod. 8oon after the raspita-
tion b sIop{ied, a very Urge but steady rise of pressure U observed. 8ee Fig.
144. The rise so witnessed b very nniitar to that brought about by power-
(Ullv stiniulslin;; a number of raso-coDstnolor nervca : and there can be no
doubt that it is due to the venous blood stimulating the vasomotor oeatie in
iIm medulla, and thus cauniug comtriolion uf the small arteries of the biKly.
eepecially thune of the splanchnic area, Mftce. as we shall see, in speaking of
the skin, a too ven^Mis blood leads to a widening of the eulaoeous arteries.
We say "stimulating the modulbiry vasomotiir oeutre," because, thou'^h we
Lniausl admit that, sinoe n rise of preaaure follows upmi dyH^uxea wheu Dw
'^ionl cord hai been proviouKly divided below the nietlulla, the; vuioiia blood
may stimulate Otlter viuomotor oenlns in the vpinal cord and punilily even
nrt ilirvi'lly un local ]>vriph«ral nicchunlims, yet the fact that the rise of
pm^un- io much Icm uudiT iImw,; rircumslant^^ showv (hat the nic<)ultary
orntre plays tliv chivf jiart. As wc have just said, the oflccl of this vnso-
eansiriditMi in raising the pr«Nure, if not as«iste<l hy an inerc«M, ut all
vveiite. is iMt neutrabxcd by an ndeqiinto ijetircaMi of the cnnliao slnrke.
U|wj the ceaestioo of tW- artilicial r'^«|iimtion, the respiratory undulations of
Courae cease also, so that the Iduod-pruMure curve rises at lirst stMidily la
C04
RKSPIRATtOX,
olniott A sirKiulii line bn k«ii cnljr hy ibe Lcart-beaU; Tel nflcr a wEiilc nrw
undulRliom, iliv no culled Truub<^ vr Trmtibe-Horing curvfc, make ilieir ap>
peanmcf (l''iK- ''''i -• ^)i very Himilnr to (he previous oii«e, except that (heir
curves ara larger and nfa more Htrcepinc character. These new umtulaliooa,
siDce they npiicar in ihc ab»ciicc ol' allthonicii.' or pulinooaiy muveinents.
pn»ive or nclivc, and are witncraed even when both va^i ar« ciit, miuil be of
VBSonioiorial origJD ; the rhythmic rise must be due to a rbvtbmic- ctin-
•triclion of the tmnll arlcrif^s, and ibii) probably is cautMl by u rbyibuiic
di»cbar{;c from vasomotor cenires, and («peciHlly from the medullnry vaao-
tootur ceolre. The undulations are maiulained u» long us the blood- prciau re
roDtinuea to rise. With the iucrea»ing venotiity of the blm^l, boirever, Ixwh
the vaoomolor centres and the heart become eufei'btcti ; the undiilKtioiif di«-
iippcnr, and ibe blood -pre^Mi re ra)>tdly *inl(».
Pic itt.
mmmfh
A
-o^
A -
. y \ 'JiAA/ V \i ^ y \i A A A' s
Oumi-nmn'ai Ci'mvib niniiini ^ ei>««n»OK at Bkrathiko. iTajn-Rf-llntni. Cram i
nenmot 1, ^>, 4^ y %m \t^r%l»iHitt^lixi<A ftranctnc Irinr rotitlauou* Inolur faiinli>c ilipnniM
<f > prakmttd iitovmlltin. m tlml %h* •vvcml nittvt it|iTM«ii1 mrmtlvr U*gr* «r (|i« •*>■« exiMt-
mml. t^tb curve 1i plarcd In 111 |in>pn rnlllun rIiUvt Ih llif Ian tlsv. iitiifh, l»!«r« ■pMn.b
onliuil; *iiil llI•L'^vluu•l^Bl, lUrUiiH tnai ite liLaiic ii| invDifd Vj I.iIie bloodpmnnwnNi la
tUlem :!, iLiiiiil I. tint MInnKiklii In Mufii.V nimi 1 liulion tMiiia (wflol wbvu ■rUndiU i«»|4iUhMi
■attxJDg ke|il op. snil ibe uiiiluliitliiru vMhlc siv Ibow I lie niiliitpof iiblrb hair hwti (Uicunadl
ilw t*|iu ncritt* buTlnit bmii cm ibv pulMilana oil ihc nii?cnl und ilnc«ii cf ibc Rndalatknit do
notdllltv. n'b«n lb* uIIIIfI&I miiinhtm «iu antiwiiilttl thcw undulklluii* iltanxiiml. M>d Um
bliic4-p(*aiite irm Mnidltjr vbllp ibe bnn-lvad bvcome ■lou'iFr ^oon.wtbniitn lnrarvi>X.M«
niKliiInUuiu »fpaut4. A Ulllc Ealcr Ibe Lluol^|JRHUfL■ n» Mill rlilns.Ulv han^bnt* tliil ilaiinr.
tnil Ibe Dii'liiUtlniit illll tnurr uIivIdu- 'i-uii« ni. I^Utl iiit*[ Iciirtu I; Uic (rawin* vu ttlil U^bo.
bullhu brnnibtali «rK quicker ■nil lhFUiid<il«tloii>niitli-r. TIi» pmaun Ihpi) br^faii toMt npldlf
tttayti^. iDd nimliiiicd lu f*tl umliiuiiii- time Uier nnini'lkl m|4miian nu mumtil
We may here incidentally remark thai the occurrence of long, alow iioda-
Itlioni i« not depciideni on ihe rewatinn of the reapiratnry movcmenlt, and
on nn abnormally vcnoiin condition of the blood. They are poinetiiiieH
(I-'ig. M9) tccu in nn animal whuM hreatbiug is fairly Dorinal. We need
not discitM them any further now, and haTS introduced them chiefly to
illiMlrate ibc fact that the v»i"im<itnr nervuuB ayalem i» apt to fall into a
ciiOdiiion of rhythmic activity. It hn» been atiggeatol inat the Dormal
RESPIBATORY ITXDl'LATIOXS.
506
K'rnKiry uiKliiliitioDB may b« due lo a rbylhtnic I'lMf und Tnll nl' ilw
ity of the ntsomotor centre, syuclminouii, lilco ilmi nf the cnnlio-
inliibilorr ceiitr«, with lli« reeiiiratonr movemt^iiu. Thrm aiii. hovrvver. li«
iioiloubl Ibm the r«&|iinit(irv variuliuu* in lilcmil-piVKViitT iiro iliir to th«
mci-tianiciil comlitioiiii ducuftKd abuve, «ii<l ibRt vnwiiiii>t>jr intlueiic«« ihIit-
v«i« but Utile if ftt alt.
no. iM,
«-iiB CnTK OP i IUriiit. lin(<ni>iiD ayi a ?ii>h|:V Movmii Si-iimcn. ia wow
fnioamr* wu •tctcnicii not b; niMntof ■ inircniT muiumdn. but lif as Iruinimciit dnilir to
Ihm w* IdanUcitl hIIIi Fli-k'i (prliiK Vrioivnt^.i latacli bMTt-bcal (bo nimiird ■nd iloKDaiiil
Mmt* arg rttf cine Inrf'bei, biil inn; bv Twll)' •llilliisal^bin) br Ih* brip ot a IvnK Tilt nndalR-
Uont oT lb* ncM ofd«i kic that or nvjilnilon. Tbc wider luMi* ■(« (h« Tnutic-Ilctfni furia. ut
wUcb Iwo coniploie cunn nud |»ttlni» <•( i«o <nb»n ate ilionn, Kicli Tnubf-HnliiK cnna
(OMpAlH ftbool tiln* [v>Filnl<irT currm. anil rarh n^iJuliuy ciiiia tUw ihu Mittf nomber of
ItMrt-bnu.
$ 388. The f\inb«r geticral etle^rtn, ^iniilnr to the above, on the vnacular
eyslem of tletldeni art«riiili;[alii>ii of [lii^ hlmn] iimy be *lu<li«d by tftkJu^ k
blood* preastt re trnciiiK fruni the oiiroiid or utht^r Krlvry of uu kiiiniAl while
the iiilvrferviiw with rc?-|iinilioii in [Hi!>hi-<l uti lo n fntiil a>phyxiu. Diiriug
the fint ami Mvond ntiigc-* of thit nvfihyxiu iht; blood- prtvKutL- rixcii rapidly,
allX)iiiii)C a iK-iKht far above tlie iioniml. During the third Mtagu it falla
even more rapidly, rrpiiMiii^ the normal aod becoming nil ns death eiitiiie*.
If the Hniii.al, no uran having been ^ivc-n, w breathing of ilwif, and if, aw
uiually in thv cute, the a«phyxm is brought about by occlution of the trachea,
t>ii that the mochanicnl enecto of the n^apirntorv movements arc exag^rated
by the air Iwiitg unabl<t to enter the ehcKl, tlio renpiratorr undulntionR of
tfiu prnoure-curve due to the n>echniiical eaiiH-i> diwuiwed above are. Fs{ie-
cially during ihi' lirsl stage, eslennve, abnipl, and irregular, the inspiratory
movemenis being aecompanted by a coti^pieiioii» fall of pmsure. When
ibi; niiimal has liceu previoiibly plac«d under iirari, m liiat ibe tcqiiratory
impulses cannot nianitest themselvc!) by anv muscular uiovemcots, tbe riw
of the pmeure-curve.aswehnve already aaid, i« at first st«ady and unbroken,
but after a variable [leriod Traube's curves make their appearance. As
during the ihint tila);e the pressure Mnks, ihe^e undulations poM away.
The henrt-beau are at Ural somen hat (|uiekeued. but sjieedily become alow,
at the san>e time, b« we have seen. Dot notably losing force, so that the pnl^e-
curves ou the tracing are exceedingly bold and sirikiog. Bui (he bolduetB
uf the curve of the OMreury manutueier is, it mtat be reineniberM), partly
the mere resiill of the »lo«Deu of the rliyihai ; the mercury ha* time |o fall
largely between eaeb Ino heats. * Fig. 1 48, 8 niHl 4.) Kveii while tlie blood*
[>retwire it Mnking, and when the eardiae Mroke is now certainly livM-ning
tl yifffT, the «loinncM of the cardiac rhylhm is still sufficient to maintain
aomewhal time characters of the curve. Tlie strolH-s at la»t, however, rapidly
506 U£S1'1HATI0X.
fkit in strength and became irregular, though the heart continues to beat fur
some seconds after the reepiratory movements have ceased.
If the chest of an animal be opened under artificial respiration, and
asphyxia brought on by cessation of the respiration, it will be seen that the
heart during the second and third st^es becomes completely eorged with
Tenous blood, all the cavities as well as the large veins being distended to
the utmost. If the heart be watched to the close of the events, it will be
seen that the feebler struk&i which come on toward the end of the third stage
are quite unable to emjpty its cavities; and when the last beat haa passed
awBV its parts are still choked with blood. The veins spurt out when
pricked ; and it may frequently be observed that the beats recommence
when the over-distension of the heart's cavities is relieved by puncture of
the great vessels. When rigor nwHU sets in atler death by asphyxia, the
lefl aide of the heart is more or less emptied of its contents ; but not so the
right side. Hence, in an ordinary post-moitem examination in cases of
death by asphyxia, while the left side is found comparatively empty, the
right appears gorged.
These various phenomena of asphyxia are probably brought about in the
follonisg way :
The increasingly venous character of the blood augments the action of the
vasomotor centres, both the medullary centre and the subsidiary centres in
the spinal cord, and thus leads to a constriction of the small arteries, espe-
cially of the splanchnic area. This is the chief cause of the markedly m-
creased Wood pressure ; though the venous blood ma^r possibly also aet
directly on peripheral vasomotor mechanisms, or, what is more likely, may
increase the peripheral resistance in the capillaritj themselves, since there
are reasons for thinking i,S ISa) that venous bloixl rich in carbonic add
meets with more friction, and pas.%$ lets entity through the capillaries than
does blood less venous in character.
This increased ptriphcral resistance and the high blood-pressure to wbidi
it gives rise, while tending to increase the distension of the left ventricle and
so indirectly helping to augment the force of the heart's beat, soon becomes
a direct obstacle to the heart emptying iUelf of its contents. On the other
hand, the labored respiratory movements favor the How of venous blood
toward the heart, which in consi'qnence becomes more and more full. This
repletion is moreover assisted by the marked Infrequeucy of the beats which
is soon developed. This in turn de(>enils in part on the cardio inhibitory
centre in the medulla being stimulated by the venous blood; but, as we
have previously seen, cannot be wholly accounted for in thi* way. The
increased resistance in front, the augmented supply from behind, and the
lung pauses between the strokes, all concur in disteuiilng the heart more
and more.
When the large veins have become full of blood, the inspiratory move-
ments can no longer have their u^ual eS'ect in faiililating the venous flow
into the right auricle. The chief effect of liie chest movement, as far as the
circulation is concerned, is to widen and so to increase tlie capacity of the
pulmonary vessels, and at the same time to diminish the pressure around the
lai^ arteries; hence the marked sinking of the blond-pix-ssure duruig eacb
inspiratory movement.
The distension of the cardiac cavities, at first favorable to the heart-beat,
as it increases becomes injurious; and the cardiac tiMucs at\i'r a while be-
come enfeebled by the action of the venous blotnl, so thiit the strokes of the
heart become weaker and irregular.
On account of this increasing feebleness of the heart's bent, ace iiupanied
by more or less irregularity, the blood -pressure, in spite of tlie continued
RESPIRATORY CSDULAT10N8.
B07
>
I
I
trtemi eonairielion, l>egins to fall, stnc« leu* srid lew blond U pumped iutn
aneria) system ; the iKilduaw of t)i« ihiW curves kI tlib «ag« b cliiefiy
due to the iiifrc<)u«iic_v of th« AtmkeiL Aa the (itmntily which pmMi IW>in
tite liviiri iiilo ihu arlcrieA bcciiim Iaw m^iihI Iiv neci^ni), the pRMiire geu
lower niiit luiror, th« d«c«iit Mag nwij<lvil hj lU- iixhutintjon of (he vma-
lOMitfr centre, until idina«t before the ImI benln it bm siiDk Ui zero. Thiii nt
. _. oli»i' of M|)hvxiit, while the h«iirl and venouit B,vstotn are ditticndi-il wUh
blntxl, the nncniil oyeUHD ia Ion llinii iinrmnlly lull.
S 390. Whilo chiutgM occurring primnnly in the mpintory emtem tht»
*H)(Cl iht YMCular systeto, eoiiv«ra«ly i'hnng<m oocurritig primarily in the
vncular tyttem nffoct the rcepimtory sT0t«tu. Tiro kinds of change in (ho
viix-iiliir »yst«m bearing on two parts of the respiratory syMem dcwrve
Mpwial altoDlion.
lo (h« firet place the rMpiratory meohaDism may be olfecied by changes
in the blood-supplv to the respiratory centre in the medulla. We have
iilrctidy Been (| 373) that ibe sudden cuttinft off of the supjdy of blood to
the ntedulia gives rise to dvspnivio respiratory movemenla sn<l may lead to
expiratory ranvulaiona. 'fhat b an extreme ooM; but. short of that, the
nctivitv «f the re:ipiratory centre, the exlonl and obaraoter of the respimtorv
esplijsioui wbicli take pla«c in it, may 1>e varied aoRonlinK as the eonstricled
or dilated omdili'Mi of tlitt dmali iirlerim branching otT from the basilar
artery or of the iMwilnr artery itM-lf allows a scatily or a full flow of blood
throui'h the mo'Iulla. .Vnd it in [HMiibU- that w>me forma of dya]nu!ea may
be brought Altoiit in tbii> war-
Mnch mure ■■nmmon and iinporlant. however, iii llu- trctnd kind of obangie,
that nlTeiTliiig the oir^'uliuion Ibrough lh« hin^d. In liic normnl organism an
•derjnnte nupjily ol' arlcrial hloixl to the ti»«iic« is sccurcil by an adequate
renewal of the air in the pulmonary alveoli, and an udoiuiitcly rapid flow of
blood thMxifih the pnlmonnry capilliiricK. When, as by ohstnietion in the
ptilmonary nrlerics, or by failure of the cardiac valves, or, and perhap CMpo-
cially, by an inxutticivnc canliac stroke, the stream of blooil from the lungs
into the led ventricle is lessened either in amount or in rapidity. Ims oxy^n
is carried to the tisBues, includint; the nervous tissue of the medulla, and
dyvpuoea or" want of breath" follows. When the circulation through the
luDgs b in full healthy swinn, the lucmo^lobin "f the red corpuscles i« ait we
have seen Mtiiiraicl or nearly italurated with oxytj^ou. [f owing to a slower
•tream the red corpuscles tarry longer iu their ))«»a;{e alon)- the walls of the
pulmonary alveuli they cannot thereby take up a oonipensatin); addition of
oxygen, indrul. It l< doubtful if they can take up any additional oxygen nt
all. The bl<MHl fiillintr under these cireuinstanoes into the led ventricle and
sent tlienoc over the Ixidy b not more arterial than tinml ; at the i>niiie time
the amount of bhxHl tent out at each henri .nroke is Itm, oflen mnch Iws,
than the mirrnal : mid tlto mednlla a* n-ell a-> the Oliver timHia milTer iu cim-
sat^uenre fmm a dctiHeocT of oxyg^'n. Thi< deficient supply to the medulla
manitWiw ilvlf in dv^pncrtc or at least in lali'ircd tirentliing, which sometimes
throng)) the nifchaiiical inrtiirnoi* disciiwed abuvr, hn»t the happy result €)f
imtirovini; tho pulmonary dreuiatioii and m prodnns conipeitMting erllbctai.
Wnen the pulmonary artery if xuddenly plugged with a clot the primary
and urgent symptom is " nant of breath," though air ■■4)i«rr« frocty into the
rliest : and " canlia^' dyspmea " is a common symptom of canliac disoaee.
li 381. Other systems of the bodv are also re!»(e<l to the rrapiralory srs>
t»m, tl)oui;h by tiai lees slriltin;^ than thow which bind to it the vascular
svstem. We have seen that delicient nrterUlizstion of the bloo'l stir* tip the
niu.tcltf of the alimentary canal to increased activity, and we shall pivsently
608
RESriRATIOfr,
Ki' 1 lint till- runic<-oii<lit ion Iirntn DOtftMeHlect in )>roaiiiliiig the perspiratitw;
it prvbulily bus t> similBr iiifliiciKw<tverotJi«r wcretioiko. Uo ibeoth«r htatA,
ss we hnvv tvcn. i :174, (here are r«afone for thinkinf; ilial ibe ac-tivilv of ib«
mpirnlory Miilre iiiid *o ibe energy of ih« whole teapiraturj act is inffncnnd
by chcmicnl chmigcs, oth^r (hmi tlw <l«crew« of oxygcD am) invmiM.- ur
carbunic ucid, bnitigbi about iu the blood by the activity of th« alcrltul
miuclee.
The cloeeiiecs and the intrican' of the ties which tlius connect thv mtnin-
tory tysiem niih nhiio«l all perls of the- Iniily inuv t)« illuslmiol bv ouiuitlrr-
io); the edects of muscular work on the body, ami the «oiiditionH nhich. a|<nrt
fn>m tl»e capa<-ily of l)ie rouaclea theinaelvn aiu] of the tnolur ni-rvi>iiB u|i|ia-
ralus which jiuis them to Hi>rk. detcrniiiH- ihr ])oitfr of the body tu do worh.
Duritt^ work. CHficdally urdiiout work, ihc muMUlar ooiiinictiuns rob lb*
bluixl III' niiich oxygen and luiul it with much carbuiiicucid. Thie rhunp
ill Ibf blmid nixild itM-'lf liirrciMtt the nciivily of ibe rmpiratorr centre bdiI
lb** (iicrgy of the rtvpirutor)' niovcnicni*. and might he KiiflitiViit to securv
euch iin incrcOM- of ihrwr nrnvt'iurnttt, ihiit ihc dcticitncy of osy^en anil
infTcnx' i>f nirhitiic iii-iil i>hould never ovrrctep certain limits. Dul, att «e
have raid, aj'pnmitly olhcr proditcis of rauectibr nietaboliim act «< jKitentJy
in Mimiihilini; the nvpiraUiry centre, that th« reepiralory moveinenitt are
more than sufficient lo couipciuale tbo changes id the gaeee of the MiwmI.
The efficacy of the angnicntcd lespiraiory niovenienie is much iucrenM^I bj
B cciicomilant increase iu cardiac activity and a swifter or full«r strrani of
blood ihroiigh tbe lungs : indeed, unless hacked up by the cerdtnc incrmve,
the mere increase of the puhnonary veuiilaiiuD might prove inad<^|unir.
Hence ihc capacity for arduous uiasculnr labor \s determined mil by the
rtspiraKiry inechanUm alone, nor by the ra!>culaf system alon^, bitt by
both, snd «^pcciatly by both working together in harmooy and concert
Tbe incrcamd venlillaiioii uuuld he idle unless it neri- accompanieil by a
(|tii<'ker circulation, and tUv i]uickcr circulation would i>iniilarly Im- of com-
turativt-ty liltle iih' unkry aciiimjiaiiici) by IncrraM'd vcnlilalion. To a
vttandrr ihc working of llic tci-i>iraUiry puinp in much nKirv obvious than
thai of Ihc vascular »yKl*ni. sad iijilnd ihc xuhjtci himwlf is inucb nxm-
directly ccnfcious of cliangi'i> in ihv t(>rni<-r thiin of cbnngi-s in ibc latter.
Hriice «hen the orgsnimi <di«i'x to he able lo ricW tbe demands nhicb ibe
labor is making upon it, the subjfct is said lo he " out of breath." tht>U|;h ID
a large number of cases (he lailurc lifs much more at the door of ilie v»-
cutar than of the respiratory system. And, as a rule, it may perba|H be
said that Hbtn two men diflcr in their capacity for strenuous nork, such as
tunniiijf a race, the ditlereiice, ihuugh it is oOeii familiarly spoken of aa one
of" wind " or power of hreaihing. i» in reality not a dillerence in venlilaling
capacily but a ilitfertnce in the power of ilie heart lo keep up lo and wora
in haimtiiy wiib the tncreated letpiiatoiy niuvtmeiiiii.
Thus there ere two main factors in re«piraiion. ihe mtpiralorr mecbaobtn
proper, and llie circulation, the <<iie hnnjiini: the air to ihit blood, and ibt
other the bidod lo (he air. W« may remind ihe rc-udcr ihat llK're ■ n\*n a
third factor, nnd (hat me of urenl moment, the amount of hirmogloMn, that
ia, (he nnnibcr of rtd tdrpniiclis, in ihc bkod. The anioiint «f oxygen taken
Up frcni the Inneti depends iiot inlv on ihc stroke* ol'thc n«pirat«ry and the
varculnr {'.uiii)s hut aho en ihc riclintss of the htrod in reti corpuscles. A
b<<ly wbiih licm Irss of bleed cr fmm disease is ansrmic is thrown out of
breaib by very slight exeiticn, not 'o much bccauH the respiratory or t)ie
vascular ptimi) ia week, hut because, through lack of oxygen carriers, with
their be»l cdona ihe combined pumps can only deliver to Ihe tisaucs,
RKSPIKATORV MuVCMKNTfl.
fi09
Hluiinp th« iti«tlull«,KD inftdequatenippl)' of oxj-|{eii. Aod ftii psrsoiu,
■Kit tt'itr of bictawlobiii io proportion to tbeir body wi-t);hi i» aln-a^r^
^« jmr. sK proverbiall]- " scant of breath."
MoblFtED RrariBATORY MoVEUEKTS.
i 998. Tbv roHpiratory Rii^nfliantMii with its ndjiincl*. in nddiiioa to lla
'Hptntorr fuortion, bccomitt uf aen'icv, ivipccinllr in tlio nue of iiuui, hh a
ikratBa iif rs|>mwing rmotioii*. The rnpinttiiry ciiliimii uf nir, moreover, ia
■I* uit from (Iw cbmt, t> rmjin-nlly iiinilc um: of ia a meduinicul veay to
J^pvl bodka from tlir upfMrr nir-piiMMif^w. Hcncn ariM a number of iiccu-
Q*ri* Diixlificd anti mure or Ivw coniplicntt^l r(W|>initon- niovenieoia. tighiiu;,
Bpu^iDg. laugktrr, rlc, adaptiKl to Mciirespvcia) ends wbicli aro not dw-
'Xifilr rcKpiratory. Thoy am nil cneDtiallr ntfltrx in ohamcl«r, the alimulitt
'^vtrmiiRioK Mcb movcmeDt, snnKtimm affecting a pcripberal uflVrvtil nerve
*^ ia tW i-aae of cou^'hing. fnnirtiinee workiog thmut;b the hijcbur piirta of
'•M> braiu At ia laughter and trying, KDnictimm powiblv, lu in yuvrning nnd
*^li)Dg, acting oD the respiratory ce«ilre ileclf. Like llie simpk mpirntory
^ct, tlMT inaT with more or loss Buooess be carried out br u direct elf ^rt of
M<;Ain<7 U a deep and lonx-drairn insplniiion, chiefly through tbe ncwe,
follownl hy a somewhat slioner, but correspondingly large eipiralion.
VaicNiiiy iJt (iinilarly a d«ep iii»pinillon.deo|>erAn(l long«rCi>ntiniied than
^ alKh. drawn ibruugb tbe widely ojieu tuuiith. and acornnpanled by a
fxculiar deprtmlon of tbe lower jaw and fre<|uent[y Ity an elevation of the
•bouMrrv.
Bteemiyk coiicisI« in a Nudden in.->pinitor^' contrnotion or tlto diaphragiu, in
VW nmne of wbidi the glottic soddi'idy eloeot, so thai the further enlraDoe
I *>( air into the chest is preveiiliil. while the impulse of the column of air
joat votrrinit, as it strikM upon tbe closed glottis, sir as rise to a woll-knnwn
MMMapanyinK sound. The aflerent impanm of Ine reflax act arc conreyod
b<r tlw gaMric branches of the vagin. The closun of the glottis ia carried
oat hr nsana of the inferior laryni^al nerve. See Voitv.
Id soAl^ia^ a series of «imilnr convulsive insniralioua follow each other
rinwlr. the glottia being closed earlier ihitn in the ciue of hiccough, so that
L Kula'iv DO air enter* into tl>« chest.
f Qniflting consists in the first place of a deejt and l-iugdrawi) iii»{>lniliou
br which ine lungv are welMilled with air. This i^ fidlowinl by n complete
eliMurv of the gtiiiti». nud llit-n comes a ludd^ii nod forcible cxptriition, in
lie mirlst of which the glottis suddenly ii{i«iiii, and thu» a hliiM of air is
drfna through the upjier reapinUory |ia»rax<*t. The afl«n-nl impulses of
Uiia rvfWx act are. in most caaai, as when a foKiiri IrMily it lodged io the
luyax or by the side of the epiglottis, cotireyed l>y the ftuperior laryngeal
nerve: but iIm- morenient may arise from rilmiili applied to other aShrent
branches of the vagas, such as thuw xupplyiug the broachial pas!ag«s and
•bNnacb and tbe auricular branch di.itributcti to the fnfilu* Ktlemtt*. Sliinu-
latloo of oiher nurvce also, sucli a< tliosv nf the skin by a ilraught of cold
^r. mar develop a cough.
t- ■■■--■■■iti the general mnrvnipnl is r>ventia11r the some, except that llie
[ •>; m the pharyiix into tbe mouth is cloMd by the conlraciion of the
anc-M' r j>illars of the faucw and the <Uc'ccnt of tlie soft [Mlate. so that the
force of the blnst is driren entirely through tbi' now. The afferetit impulsea
hen mually come fium iIh- iwmI hrancbM »f the fifth. When meexing,
510 RESFI&ATIOX.
however, ia produced by a bright light, the optic nerve would seen
the afferent nerve.
Ijaughing consists essentiully in an inapiratioQ succeeded, not by <
br a whole series, oflen long continued, of short spasmodic espiratii
glottis being freely 0}>en during the whole time, and (he vocal cord
thrown into characteristic vibratione.
Iq crying, the respiratory movements are modilied in the same wa
laughing; the rhytnm and the accompanj^ing facial expressions ai
ever, different, though laughing and crying frequently become ii
guishable.
CHAPTER III.
THE KI.IMIXATJON OF WASTE PRODUCTS.
,S 393. Wk have Uaewl the foot! from ih« alim^Dtarv c«niil into the blood,
antl. ilitl the 8l*t« of our knowle<ige (>erniit. the italurAl couree of our slmiy
nould be lo iraoe the food from t)i« blood iuto the tissues, ami then to follow
the products of th« actiTity of the tissues back into the blood and so out
of iDe body. This, however, we eanoot as vet salii-faciorilv du ; and it uill
be more ronvenieut to Mwly ^ri>t the final produclji nf ifie metabolism of
Ihe body, ai>d the manner in whii>h tbey are eliiuiiiHled, and afterward to
return to the discussion of tbe inli-rveniiiu; iilC[ni.
Our food on^isls of certain fiKHl-atnfhi, vix„ ]1^>leidll, fam, and rarlioliy-
dndea, of various salts, and of ttntcr. Id their iiiimagu through the blnod
and tinuea of the body, the |in>iei<K faCn and i-arbuhTdratt-ai un- c-oiiv4-rt(T(]
ioto urm (or sonie cluwly alhed Ixnty), cnrlxinic ncid aod water, tlx: nitrovrn
of tbe urai btiiij; rtiniii.hi.-d by the iintteklt alon«. Muny of the ]>r«l«idM
OOOMin Hllphiir, mid uli«> huv« jdioKphoniK iitucht<l to thvm in »oni« oiimhi-
DAtlon or other, ami Mime of l!i<! (ul*. talcfii n» food contain pbospborus;
ibtae tWn»atr iiUimniily undergo oxidation into phoophatn nnil snlphatM,
nod leave Ibc body in ibnt form in company with tbti other »ii!t».
Broadly i^[M-ulcing ihi-ii, the waMc proiluctc «f tbe luiininl cronomy are
iirm, mrtwniG Mcid, null:* and wnter. Thr«u It-avv the body by one or other
of (liroo oiniu cbiiniicK the Inngv, the skin, and the kidney. Sinie piart. it
is Inn'. leiivei" tlw bwly by the bowel*, for. an wc have peon, the feces eon-
tiin, Itroiileii nodigntrd porlions of food, substances which have t>ei-R
BMrcftd into the bowel, and are. iherefore, waKlo prudiiet« ; hut the amoutit
of lhi«c ia so small that thcv Rwy bo neglected.
The luncB Dcrve ii# the cfuinnci for the d>ichu;ge of the greater part of
th« carbonic acid, and a cmmderable ijuantitr of water ; this discharge we
buve JHst sludifd. Though tli« akin there lenvc llic body a oumpnralively
small <|uantity of mIis, a liltle carhouic add, and a variable but on tbe
whole large luantiiy of water.
Tbe kidneys discharge nil or nearly all the urea and allied bodies, the
tcrealer portion of the salts, and a Ur^e amount of water, with an iiu-igniti-
cant quantity of carbonic acid. They are nefiecially important since by
them practically all the nitrogenous waste leaves tbe body, and to tiient we
will turn tirst.
Tbe Btrvctobe op tbe Kidxby.
^ 384. Th« kidney ia a secreting gland constructed upon the geoerd pka
of a <'<>ni|Hiund M-erelJng glaud, but ixMseasin^ special features. Tbe eecret-
ing [M>rlion«, in nbicb the divisimui of the main duct or untfr end, are not
relatively »Iiot1 tnben with branchings or lateml bul|<ings, that is to say, are
not alveoli, bnl are cxlreinely long narrow ttibtdes, will) no branchings or
lateral bulging*. The whole Ixiily ef the kidney is made up of tlieae oon-
lililuent tiibultn, uriuiferou* lubles, tubuli ttrinifiri, closely packed together
'the Ki,IMIXATH
[no. ISC
with iuftt Its much C')niii?clit-e-tiBsiif> as is sufficient lo oirr; s l>rg« <
blDoaveeaeU, a certaiu Dumber of iTmptiatiire, and ucrves.
Kach uriniferoua tubule, consisting; nf a single layer of epithelium rnttiD);
OB B basement membrane which over the ^ren[ jiart uf the leu^h of tW
tubule is couapicuous and diniuct. bej^-ins in a peculiar i^iruclunr called a
Malpigliian i-Hiiaule, and fjr tlie fim part of its course puiitues a path ithi^^b
U, uii the vrbole, very tn luted and dt^cioua, diirlDK winch ic may, for ifae
prawnt, be spoken of tu a ini:4tL-d tubule, correspoiiditijc t-i the tubuliu mm-
b>rtui of old writers. U Mul>»e()uenlly tukeri n inun; Mrai^ht courM, aud u
then called a Atruight tubule, fu&u/iu ir^fiui. At tu l>eginning and duriii|f
it« tvr luted cotirM!, thv tubulo li<si, f>>r ibe laoBt,
neur llu> Furfacv nf the kidney, farmiuff tlie
nuiiu part of the enrfrir of the kidner {Fis.
ir>()]. During Ita vtnttght iiiunK it rum
lovrnrd tlw dcrpw parlK, ponvi-rgiug lavanl
Ihc Conciive border or bilut of the kidocr
where iho ninin duel or ureter tuttm : the
couverciiiertraiuhi tuhiilc* forming together
tin- mrJidii of tlic kidney. While ptiniitn^
Ihr- lirsl twisted and devious part of tbrii-
coiirac. during the gnater put of whichj
as vm nhnll see they [mmsosb marked t«cre —
tor)' chAniclers, the iubulee do not join «afh
other. Diiritig the latter siraight part ct^::^^
ihdrcourae, when lu ue sliall k« tlteir char— —^i^
Rclere are thr«c of conducling rather thaiEr.^ to
of secreting tubulcx, ihev repeatedly joia
A fler each junction the tuWle, though wide
thau,^h of the two tubulee which JMOed
form it^occupies 1cm spiice than ifae ti
toother; Bimce the inedullary suhtiAnc* be^'
conui l«M af it convergai lonard l)i<- hihi»<
The medulla is, morvoTer, divided inio r k >
number (varvlng in dilfi-rcnt nniniaU, brin)t^ ^*f
(>ne in the rabbit ami the ral, ntwl about lur ^ ^
or twelve in man) of uiaives. rnch of which '' •^
»iiiei> it diminiuhe* in hulk toward the hiluk ^5*"^
hiw lh<' form iif a jiynunid. i<jri^Mi</ of J/a/— '^'"
}Hghi, with its Kpcn direci^id nidinlly toward the hilus ami its linso nstin^^^?
on and becoming cimfuscd w'nh thf cortex.
The iirrttr or nuiin iliicl of the kidney when traced to llie kidney te found
to expand al tlw hihis into n fllunel-^hnpn) irtivity, the prlri*. which divide
ur branches somewhat irregularly into a number (equal to thiti «f the pyra-
mids) of ehort, broad luW, niAjeff. somvirhnt in the way that the htiuil uf a.
{(love divides into the tingore, but more irregularly. Into cadi cnlvx ih^
summit of a corresponding pyramid projects fur some little way in ihe form
of a nipple, or ;>i;>iV/a, the epithelium lining (he calyx bein^ thus cmitiuuoua
with and, OH it were, relleeted to form the eiiiihelium ciivering the projecting
nipple of the pvrainid. llie straight tuhuW forining. aa we have seen, the
pyramid, thougli nunicrous at its base, become by Te]>eated juQctlun^ fewer
and larger, and tlually form a nutulier (in mau about a score) uf relatively
wide tu1iul«a which a|ieu into the calyx at or near the vety tunimil of the
nipple : here tlie epithelium lining the tubule* hccoino coniiauoiu with ihe
epithelium covering tlic papilla.
Heuce, in a mdical »ectioa of hunuui kidney (one taken io the long txU
Sacao* Til Rill an tik Kuocet.
SIMVnM Tir( MEI'l't-LUTY j>u Ojk-
TIKtU ttoKCW*. A.1U TUB BoilWIISi
ornit Vnnni lUuiixj
n. unl«r : b, ■■rlviioruiauieMrif.
(•pUIiE lutnunile'l t/j oillm "I tho
•avivlatT lube; <l. [lylnmlilikl pat-
tlDnt -, t. nrilMl ]«nl(iii of i>w kld-
u«j.j
THE STRfCTCBE OF THE KIDXKV,
I
I
I
I
I
I
htina iirffrtaMt}, lli(> wlioli- imter |H>riiitu of t)i(^ orgiin. ii)l iVMini] vxi-cpt at
the hilu.*. Hill Iw ««n t" be uctupiffi liy tin- fairly uniform cortex, which,
being wmpodwl, ii» w« h«vc Miiil, tnninly (if iuIms l«ii>iiii^ in (it) dirMtious.
EreMOU f»i »ecliop lo lh<: iiiikrtl eve n gniniilar iinpect. From this cortex will
It **en oonrerfriiig tuwiinl the h^ihi* fl rerluin number of pyramids, each of
which) Muc« it i« niuiiily oomDufcil of ndinting itraieht lubules. imd since
iho minulo bl<x>dvriwel« minifjring ia it hitrc a Mtuilnr radiiuiiig Blrai^ht
cour«o, will pmetit (u I he niikf<l vyo n more or Imh marked radiating t*rain
or striatiun. Tli«np6X of riirh pyrnmid whcrctbc scclion has paased tliroutt'i
lh« apox will bo SMti projecling into its approprialc cnlrx. the calyc«a will
bo t«ea uoitiDg lo form the p<^lvis. and provided that the plane of seel ton
hu jMiwed ifarough the mouth of the ureter, the pelvis »iU be seen narmw-
ing into the ureter. ITw section may, of course, have miaud tlie urotcr : it
ia also very likely to have cut one or other of the pyramids higher up than
the HtlKchiueut of tl>e calyx, in which case of course the prctjectJon of lh«
pMpilla of the pyramid into its calyx ia uut seen.
Jhfi pyramidi are teparnled from each other Uiterally ulx^ve the nttacb-
mi-nt of their reHpeetive caljcea partly by a small ijimntily of cortical "uh-
sKiimr which creep* down their udes toward the pelvis (coliimnit c.f Bcrtiiii),
anil rIko by the larger branches of the bloodvesacls which, lying uut.iiile the
GIvis and' dividing as it divides, plunge intu the milMiancc of Ihn kidney
twe«u lh« aily<^^ and •» between the pvraiuidK. and then run otitwunl
loirard iho junction of the cortex and meduflii to be di»lril>iitod it) a mnntier
which we shall d<*cribe oremnlly. Tlie kidney it really, m »ecn in tho
embryo kidney of man and indicatoil by the adult kidney of some nntmnls,
Cfvnipuaed of lobes, each lobe consiitting of a more central metluila in the
form of a pyramid. covcfkI i-«[>vcially at ito huse, hut also to n oerlwin extent
nt the sides by cortex, and opening at iln ap-x into an appropriate divi»on
of the uMer. As in other elandif, the larger hmnches of the bloodvenels
run in the connective ti«>u« joining the lohe* together, and paas tlienoe into
the lobce. In tli« adult kidney the lobes have become more or lets fused
MMlher. In the cortex the fu-ion in complete, but the pynunids Blill uiain-
tiiiD the medulla in a iobcd condition, wparatrd. however, laterally by noth-
ing more than by bloodvc6Mle, with the connective tissue carrying them,
nnd n n'mnant of cortical Kubelnnce. The surfncc of the kidney, save iti
abnormal cases, ibows no indicniione of division into lobes; the uniform
level i)f ihc cortex is bounded by a capsule of connective tiaiue, which may
b« easily tttrippcd off from the cortical substauce below, and which at tlio
hilus is contmuoufl with the connective tissue suntiunding and binding
together the ureter, renal vessels, and mial nen'es. A quantity of adi|>oM)
liss(i£ not infn^nently turrounds ihe kidney, being especially nbumbtnc at
the hilus.
i 386. I-'ach tubule begins, as we have ssid, in a Malpighian capmlo
tFig. ir>l] somewhere in the cortex, either near the cnpxule or nor tbe
ase of a pyramid, or at Home intermediate level. From thenoc it runs, we
have also said, first aHatwtMed tuhuleandsubeequeatlyaa a Mtrnight tnbule;
but in the Bret part of iio coui«e its path is so peculiar that the word twisted
does Dot Bccnnilelv <le»cril>c it, Sloreover, t!ie cbamcu-nt of the tubule
dianite BO markoUy nt varioiiii parin of its> course, and thotc chnnges are
pnihably of Bu<-h great itii)»ir(Hnce that a desi-ri]>tIon of tiie liibulc at stio-
iH.>wive steps of its jirogn*!' along its whole length bet'oncs advixabte, though
we at present do not nndemtaml the meaning of the varioun changw. Afl
we Kball see, some of thcMt complt-x ptruliarilicn of the mammalian kidoejr
uv partly explained by the stTuetnrw of the kidney of one of the lower
33
bU
TUB KLIMIXATIOX Of WASTG rRODUCTS.
BmniuK tudi lu a frog. Il will be conv«nieul lo dnoribe flntaome dcMili
of th« |£uiii-nil cciune iiii<l U) study tbo changes in (.■hftra>-|<-'r »iilMM|UuiitU.
Lenvmg tlic ai{<iiul<} tbu tubule fornm id the iiciglibunug ntrlC'S •evvnil
Hlmrp but roiinilcti luriix. kucI in ihia jMrt of it» coune is vcrr ili>tiuclly i
tirislAd, coDturted, oonvoluteii lubule. It ih^ii, ccuiiig to bo ifUlinetly wa
[Pio. lu
inuiiiAK vr TiiK Uui'ut or Thu tTniMrcituiv Tiui'Mki
A. tonux : B. tiMiivtio due : V, iai^Uut omm <^ <'>'> looliiUi : a. n', •ui«rtkf»l uid 4iv|i l»TMt
tifrarMi ftM rtniD rloiuorull,)
voliilcil, tiikcs on a waryurgMillvNpirnl ur Komotinua almocl tflrai^flit cuurtt,
iKJng diroctol mdiiillv tonnrd tlic tuodulla. In Uiu j»trl of ita couree It u
spoken of as then>irn) tubule. Still couliDuiag its nidial course, tbe lubulo,
Buddeiilv diininisniiig ven- mudi iu widtb, passes on f<>r some dUUiDce right
doirn into ihp p.vraitiid b(>tcin- uniil, ut n level which differ'* wilb the difRireal
tubiilc«, b>]l is alwuye iit ^^nie di«tanoc from ibe n|>ex of the ]>Tramii), th
tubule betide Bbarply round nnd pur«iM» a bncknard, nearly siraighl eour
TIIR aTRUCTUBE OF Tl
610
»
I
I
I
I
lU former oiic, until it fimlH itself huc-k ngiiiii in lh<- cnn^x nt
nee fmm tlio iiKfliilln ; iho lulmir, in fnct, in contiuimtion ul' tlic
spiral MgniMil, miikn u I'lop, Ihr /fiji of Iltn/r, dipping down iiilo tlio niedulla
for n Kfinii) distaticc. nnit (-oniiMin^ i>l' n ilricrniliay and un a*r*n>i'iHg timb,
bolh of tlMrm niiiiiing ■ riutial itnirsc, which is Mraighl or nviirly t«. The
dMcendin); limit i», n« w )mv« wtid. very narrow, but either before it inakea
the iMrud, or jiixt nt the bend, or at some liltlc distanoe beyoitd the bend,
«h«) it has nlmdy bpcom« thv n«cending limb, it ealjirgtH Romeirbat and
cliHnt;i?» in chiirador, thongh not renching the diameter of the H{tiral or oon-
voluli-'d tiibtile. Having rosched eome part or other of the cortex in a more
or lens strHight mdini lioe, the aacending limb of the loop of Henle ehanites
again in characler, becuraes atill nider, nud runa in the cortex a once more
diatiuclly twisted fourse; the twista. however, are not roiliKl but aiiKulnr,
jtivioff the tubule a sigiag apiwaranee : hence this portion of the tubule is
called the zifraiK or sotnetinies the i\regular tubule. Very soon, however,
the turns of the lubole become rounded, atid the tubule, still running iu the
Ctftex, aaaumes chamclera almoit identical with thoav of the initial convo-
luted ]>orUou : it now receives the name of tlie »t«<Md murol'ilfd tubule.
After several turns of thij« kin<j. all confined to the cortex, the tubule once
mure chang^e^ in dninicttT. and, running u «eci>nd lime in a *tTaighl radial
courw toward the Ma-ilulhi, bui^iimcit a m//Minf/ tnhulv piinuing a straight
rndiai eunrte diieclC'l toward the apex of tlir pyramid. The collecting
luhulc, joining other collecting tubule* and clinnginL' olighlly in charai'tcr,
while by ro[Ma(4!d junclioni beoomiDg larger, '» continued lu a durhaiy/lny
tubule, which, joining other tubule* kk it jMiwn toward the npcx of the pyni-
mida, oponti at InMt into a calyx at or near the summit of the papilla 'if the
pyramid.
Thus each tulHiU- starting from a Matpighinn capsule becomes in buccks-
eion a first convohttml tubule, a i>jiiml tuuule. a descending and iiaoending
limb of a lix>]t of Henle, a xigIl^; or irregular tubule, a second convoluted
tubule, a collecting tubule, and tSnallv a di»char)^ng tubule. The diKhnrg-
ing portion, the lower part of the <'o]lcetiDg portioo, and some part or other
of tne loop of Henle he in the medulla and forni part of one or other of the
pyramids. In all the rest of its course the luhule lies in the cortex ; but
from what has been snid it is obvious that the itart of the tubule contin>-d to
the cortex can not be called, alon;; the whole len^'th of iti course, a twisted
or ooAtorted titbule. The upiier |)art of the collecting tubule, though Btill
lying in the cortex, runs ncarlv Htmight ; the beginning of the descending
liinh and tli« end of the ascending limb of the loop of lleule, though lying
in the cortex, are nearly straight ; and even the spiral tubute is not fltr
renvoveil fVom being straight. ^ that the cortex does not consist of gouto-
lut«d tubules only, but in |iart of tuhules more or tea* straight. These, linir-
crer, are nut dia[ter«e(l uniformly nmunj; the convoluted tubulea. hut are
gathered into bundlen which run in a ladial direction from the bases of the
pyramids thnHigh the cortex toward the ca[Kiule. The bundles, of which
tAMV arc several to each pyramid, are calleil mtdittlarii riiyt or pvramiiU of
Rrreit* (the large pvraniid.i of the medulla being iheu distinguisncd a« the
pyramid* of Malplghi).
Bctwvon and i>urr«unding thew^vcral mtslullary rayi* are mnww of cortex,
eeen in riflinl si-rtionx n» ctiliimiu bi-twfcn two adjarrnt rays. coniiiitin|( of
oonvohitiil lubuUi!. Iwith lint and xi.s-ond. of i!iu:iiitg tubulin, and, an w« liiall
aev, of Malpighian capnuhv; all the luhiil<n> in the cohimn an? ntont dinttuctly
twisted and cuittoriol. ninw the c->lum« contains only the very lii"^'inniugs
of the spiral tubule and the collecting iiiliule. Th>' spiral tubule beginning
iu the column of cortex iNrtwcen the mcdultarv ravs miikes nt umxi for n
old
THE KLIWI.VATIOX Of WASTE moUt'CTS.
ntodiillitry ntr, ilown which it ninit lo broome n dnccndii
of Henle ; the ■scpinling linili oming tip froiu thv mi;iliil
lury ray im<l 'inly 1crvc« it to becumo ii zigng lubuK-; lun) t-itcli cnllcrtiDg
tubiile ruii? sirniglil into n nmlullnn- ny Uid iIiodco aiiiiv into tlic nH->]iiilH.
Ueacecuch my I'ousifte of spiral liibulW, deeceoditig anil aiKTCivdiiig liiuUi
(enieoiRUy the latter), of the loops of Hmlc, and colTediug tiihutn.
Since each medutlnry ray remvea »]>iRil tnbuW and collectioi; lubulcf,
and gives off zigzAa tubules at difler«nt leveU above the baaw of the pyn-
inids, it must be thicker below, where it Itotde all the tubules which it Iim
ret-eived or is about to (-ive ofl'. than hiRlier up, where it hm^ already (pven
offiome tubules and han not yet received all the tiibuUe which it will receive.
It ditninialie.1, in t'ai-t, pyramid fiulji"" <henct.- ibenune mraiuid of Ferrein),
toward the surface of the kidney ; n»d, ludee<l, jimt belon the capsule then
in It Ikver uf miinc Utllo thiokueM L-unHimin^ entirely of cortical dub^tanee,
that 1:1, of convoluted tubule*, the nieiliiUarv ray» not liuviu){ tu yet becun.
It is obvious that the upper iwrt of eacli m-raniid of the medulla diffen
from the loner part, in lo far s« tinit while the latter con taiiu siniitcht lubalM
only, nnd thiMt nxwtly dim'hnrging tubulin, the former Fontaiiu. iMaudM ool*
lectin^ mid diKchnrging tubule*, the ondit of the loops nf Hcnic. which arc
mlly pnrEs of tin- tnbuin' in what no have ndli-d gvncrnlly their twiaUd or
deTioiM course. Hence the upi>er nnrt of the utcdulln contiguous to the
cortex is sometimes spoken of a* the boumhry iont or intmiwlintr mite.
$ 396. Hiiviiig thus traced out the doviwis >ind complex [Mth Uikcn by »
tubule wo must study in more detail the special characters of the •cvAral
sections of its cours«.
Thf .\f<i{jii^hinn mp»ulf, Kach tubule begins iis wo Iwvc said in a globular
expansion, having in iiinn a diameter of about ^IJO <•. the Malpighiaii capeulv
or eii<tcapsule. [Fig. lit'i-] The several capsules are dispoaKl Ibr the mtML.
[FMLUl.
ru« of ins v»«iiti < ii]i<<iiTtt> wmi tut tcvtlu iBcnrxAXi
t. a.lubDls: th «0iMni>taU(iTt <■. intrr-i-jTHinhliil antrr: d, ■ninviii )<mnrb •. ■loiuerulnf.J
«llkl«nl Wlhi ft lilMUa nf niallaitM aniund Ibv Inbnle. K Rullcln of ibe nlM
ondaa; 4. aStctM, soil /, tfltn-ni twtl of thr rIuiusriIulI
pan in serin of circles around the medullary rays along their length, so thai
in ndial sections of a kidney they are seen in double rndiating rows in the
eoluinna of cortical subetancc between the medullnrv ruy». Kacb capsule is
ewenlially a terminal globular expanaiuu uf a tubule, and conrisls, like tlw
tubule, of a distinct and con^picuiiua baseiuenl meiubmiH-. having the ordi-
nary characters of a buaement nieuihrane ($ 211). lincil by an cjnthelium.
At one pole of the sphere the caj^ule a cuniiDued on into ifae tubule, i\i
A
TUB aTRt'OTURg OF TOK KIDNKV.
617
I
ll ineiukraiie bihI ito «jiil)i«Uuin bi>ing vontiiiiioUM with i\»p bufmciil
membrane ami the t!|iiiWliiiiii nt' thn mbnlc ; iti Ihi' jiinctinn nf th« two thm-
ijt a marked <^oi)i<lrictiiiti nr ni-<<k. At llx- ii|>jiii«ili- |>olt' ii «hitrl xtrnight i^muU
mriery (of vhirtf rclBtiniii) wi< will .'>|>i,!nk |>r<wntly). riu a^<-r«n«, ruii» intri
Ihe cAptule, Hrivinjr U-ron- it nml iiivrrtin;^ iitio th« cnvitir of tbp oa{isiilo
Ihe b«j»rtK-iil nicmtininr itnci qiithHium wjiiii-whni in the way that om- niijrhl
ihrtlM i>ii<.-'i> \t.< int<! hikI r>» invert at nnv pnrt the wall of a large (ligtvut'Teil
cliwciv Imll. Imra<vliuiirl.r upon it« rntnuici; into the cnp«ulc the iiHbrGnt
artery Hividn into ii itiimlHT of hraiiehes. Each branch fiirlh«r splits up
ii>to It niimbor of ciipillnry loops, the reUintiug limbs of the wv«rnl loops
juiaing, without lateral aDawtomoM*. to form n sinj;lfl vein-like vraed, txM
^tren». The whole lobiiluted buDcb of brauctiinff nnd looped vefnels has
more or Ima the appearance of a knot, and is i-nlled ihe ghnurulnt. The
exact mode of diviiioii. however. difter» in ditl«ren[ aniiuals nad appArently
ID different ca|«tde8 iu the tami kidney; thus in the ratixulee nearer tlie
roeiliiUa iIm; ;;lomeruli are larjcer and more subdivided ilmii in tlinee nearer
ibemirfoce. The vas eflerens slariiuR from alioul llie middle of the hunch
laniee from the t^upMile Hide hy i^ide with the va-t alTerena, thu urilii'e formed
liv the inversion of the rniMale bei»K not wide hut narrow w) aa jiiat to admit
the entering and inuiu^ veneb. Heuoe the glomerulus hann ao il were
into the ravity i>f tli« cnjiaule !<u*pend«d )ir a narmw neck eonwi-iling of the
alftrent and the elTercnt veM«l. wirroundt-d bv ihi- commencement of the
inven4.>d porlii'ii of the wall »f the caiwiili'. WEii-ii tho bloodvmtcia are fully
d»lendod with lln- hWA the glumrruliif filU the greater part of the Oivity
of the CHjwute ; when they arc conxtricLed and cootuin liltle blood, a apace
uf Mome file i* 4lcvfl'>jM'i) l>i'twtx-n the etirfaw of ihi: glonwrulua and the
<^poHile wall of the iii[mii1<.-.
The onilhclium lining lh« wall i)f the eapeuhi <y>tuiKla of flat polvgoiinl
nucleated celU which have atm'ut an epithelioid character. Indwcl, they
are se<Mi wiih dilRcuity and are best brought into view by the silver nitrate
metlMxl. These ct^lU mt on a baseinent mcmbraDO which as we have said a
(lirttnct, and in optical or other aection preeuniJ^ a sharp ouiline.
The basement membrane over the glomertilue cannot be so readily dis-
tinguMied. It appears to be completly fuwd with the mil of the capillary
leofMi which like other capillariei> consist of a liomogeoeoue membrane of
nucleated epilhelii>id plates cementeol loji^ther The epithelium cnveri&g the
f^looeruha, whieh follows the ine<|iin lilies of the surface, forming a covering
or and dippinj; down between the Krou|M of the capillary loops, and hence
is in cloae omiact with Ihe bloodvewels, is said to differ from tne epithelium
lining the wall of tlie capsule inasmuch as the cells do no closely resemble
epitbelioi<t plates, but are Battened cells, olten irregular in form, each with
n imiiHpnrent or faintly granular c-ell'HulROauoe and roumled niicleu!i ; they
are diminctly cnhioil in the newborn animal hut l>ec»nie tlatter iu thcr adult.
Thnit r«eh of the capillary liHips [|f the glomLTulus appwini t<j |)ri)jeft. into
the oavily of the <-ji|wuh: in such a way that the blixHl in thu viatwl i" j^cjia-
rntol from the cavity of llie chimuIc, and mi from tlii: himen of the tubule,
first by n thin film ci>mp<nwl of the cuiiillary wall (with which tl»: Iwscmenl
niembraiH' of the inverlcil purtion of the ca|«ulc hon liccome fuK-d), and
next by an epitltvlium cell of Komewhal |Kvuliar nature. As wv xhall proa-
ently see some of tbo prohlcmn coiirerniug thi- Hfrction of urine turn on the
nature of the ptnoeMi* cnrned out hy thia iitm covered with this epithelium.
Kach cnpeuto is surrounded hy a uninll ([unntity of eounective tiMae which,
vety scarce in the kidney genertUy, i* more obviously here than elsewhere.
Asuall amount of cohneciive limue aUo surrounds the atferent and the
effisreDt veeeel of the t(lon»erulus, hut a minimum of this timue is carried into
ihe capsule with the gitMnrriiluB. Indeed, the prcmtL-e or ouiinceti^
to tbrni a middle lo or n support of a lutip or wen iu the deplhfl of (he
glomerulue. cannot be definitely demoOBlrateil. Hence, lbou);n vre hnve
reaBOn to think that lymphatics exist ia the tissue arauod the capsule M elae-
wbwein the kidney, tlfhoa been luainlnined thai lymphaltt'a are abaeot io
th«|tloDieriilua between the bloodveaseU In ai all evcuta the penpberal
portion of each capillary loop, eovered as it ^eems lo be elowely by epithelium,
the odIv exit of material thnui^-h thu cajitUury wall Ituidii direct thmujch ihe
epithelium iuti> tho cavity of ilie uijiatile.
Thi; cajxiulv i» c-outiiiued uti iuio a tubule by a shoH constricted nnrtinii or
Deck ; and hero the epithuliiim Auddenty chaDMa i» characler and put* on
the fcsturcs which we are >iow about tn de«cribe.
In the/rol nnroluleil tuhufe the boaenient membninc ii' dii>tinct an<) coii--
vpiciioux ; indeed, wr may *Hy at onco that thin dinttnotiu-a* and rharpocM oS
ouitiiie of th<? htLtcintnt mfiiibraDO hold* fcotid for the whole li-iigth nf tb«
tuhilii' urhiiffrtiut until wc reach the di^'har^ing itibulc* io the inedulhi
and hiTt the buR-mctil tiK-iubrane it l»*t to view, wuiply beniUH it beooii
fiiK'd with ihc cnncctive-tiwuc gniinidwork or #tnimu. which ia eapeq«ll; y
well derclopi'd in the lower part of a pyramid. KlMtwbere the '
oicmbranc niny eni>ily bo recognized as an independent inembraiie.
[ fui. IM.
I
r
Tl'miEll ISijU a ^U-Kn.i ui nil, INu'* KlIiHIT.
0. <«|>«l« uhoIuIdi ilu> (luMiriuliu: ri.DivIc ol iliriapulp, c.r. nnmbiiol lubulat:ttl
lutaloi: d, c(itlw[ln«iuto^ r. (.iplnll tiibni/ parlor IbaMnndlasIlmtior U«lilt'>ISDf^lie><c
Um w4«IUrT i*ti Htm*-]
The epilbelium of the first convnlute<l tubule [Fig. 153] has the followl:
chamders. The outlines of the cells are very indistinct. a« that not unfi
<|qeiit]y the tubule eeenx tu be lined by a layer of eell-^ub»tauec iu wbi
rounded nuclei are imbeddeil iit intervals, wbeti the oulliniii are maflu o
]( u *««n that eai-'h cell, which hns u rounded nucleun plnitvl nt nUiut i
middle. i» avm or le^ cubical. Hometime^ of iiuch a betglit an tu leave a niw
nw, noiuelinies bo Iuw as to leave a fairly wide luinan. Tlio outer portion
THE STRrCTirJtE OP TUT. KlIiKRV.
521
I
I
I
I
I
tiou is ill tuni «iu-(?eedeil by a (UriHon of inucli lurger diaineler, in nhkli
tho ocIIb are moet dutiiiclly strialed aiit) otberwrise itAcmble the w]h of lli«
coDVblut^d tuhulce of tlw maniniHlinii ki<liiey, ihiis ooii^tiliiliiiif (he »eoi»i<l
Mcntinf; portbn. Tlie suceeediii}; iiurtiona of tlie lubiilt' bi»v« tbo chnrBfler
of oouductio^ iitbiile»<, and Join rbeii- felli>wa [o fall ullinuttely iuU> Uin
ureier. Obvioutily the svciind secrelio;; |M>rliini is repre^'ntt^l in tiur iniiniiniil
by (be ae^oiii) ci»iv»IiUh1 lubule, itit- iU)ir'-'i)iE tubule, «ii>l tho nw-«iiiliii^ limb
of Hcnk'fl lui)|}, wbile tbe ilcMMMxIiD);
^
ri jj
*.i
\
>J/^
limb of HetilcH l'i"|> oirr^iiands lo tno. iM.
tli« Hrcond ciinducling dliaU'il [>i>r(ii>ii
of ibc ninphibian tubule; tlir cilia,
Iwursrcr, bBVc fDtin<ly ilixnptMarKl,
«nd the liktDcw \k cnntincil (<> tW nar-
rownoK of ibi- whulf tuhtili- and (he
•bttliceof se«retiugcharmc(ei> iii tliv
oelln. Why, however, th« kidney of
tbe lowiT nninwl poaweHS (bit redit>
|>lieaUi>i) of aecreliiiji; aild coiiducliiig
porlioiM, and why reinaioa niid re-
mains unly of llie redu|)linitioiisb<Hild
tbii^ be |ire»erved iu the mwniuaiiait
kid(M-y Iin8 iM>t vel been Miii^fuclorily
«Xi>lnined.
$ 388. 'I1i« vn»vidar nrrnu)n,'infiilH
nf lb« kidney <li9ervi! i|N-ciul altt-n-
tion. The renni nrli-ry aiijirimcbiiift;
tbe kidney at ibc; hiliiN divide* iiiiu
bnincluit, ubicb, ii]i|i{mi^ round ibc
Iielvis, vMi±r into tlic Kiibtilanirof iIk-
iiiliK^v Ht lli^ iinKlt-e fonneil liv llii'
bnttii.'liii)|i; of ihr jH^lvis into c:ilvcc«,
and tberelorc iKHwi-en ibt- pvntniids.
Ituiming riKliiilly JH-twert) tfte )>yrn-
niiiU the bninchcs. reacbi»;r the
buuitdary Wtwei'D cortex and nie-
duDu, divide and, 8pFmdiU];> laterally,
form at (lie bnsea of the pyrniiiidn
areb<« mure ur kes coofienlric with
tbe hilus. Prom Uww uvfaw [Fiif.
1^4 j, vrhich anaxtonioM- lo a rerluiii
extent with other, rei<a«lii jinic^vl »ii
the one hand (n the oiitex and oti
lb« titber lo ibd nKHlulIii.
To ibft oorUx are givoii otl' rvla-
lively lyge arterin which run in a
radial direeiiun towiml the nurfnce
in tite inaatc* of orU-x Ixawii-ii the
medullary raya. From each of these
inta--loliniiiir or nuiiatftrli'riet.aalhey
are oalM, abort relatively thick
braucben are given off at intervaU
on nil tides; these takin|> a course aomowluit curved, with the convexity
dirx-cta-d toward ilie surface of the kidney, end, witbixit branching, in
Maljiigbian cMjiaule^: they are the airerenl vmrnU N|iok4^n nf tircriouxly.
Other branehcs of the sanie radiate arteri^v break up into cnpillniiiw Wr-
y:
.A
VAfei'UR SL-ri-i-v nr Kicmct. (tuui-t?.'
1. laHof nilcrlBlAieb: b. intei^WKilatBnBfT:
r, k1'^uu--'<>''u<; d, cObrtat v«anl punliii; lo lu*-
>liilu 1^ Mtr nrlartir rwlir: f, ni'lllarliaor OM-
!<■» . /, M[illt»rt« of mttlullii ; tf, nii'iiu nrh : *.
uni<(Iii rtlnnof iixaliillii;'/, •nut •lalhila: (. In-
t«<^loblUu>vln.)
Xl'2 llIK Kl.IMIXATKIX UF WAriTB l'IH>Ul-CT3.
r.'im, litis; tin- iiiliuK-!<. this beiiij; especially the ca^ near the furfsice of
ilif kitlnev. Tlie vtlereii! ve*iels from the Malgiighinii capsules also break
iiji into a i-a|iillury iii^twork which, eTnbraciiiij; the tiihtiles, becomes con-
liiiDoiis with till' iiilior network, the meshes being ronndetl or polyj^nal in
till' I'oriiial sill IK I a me. hut nmre elongnterl radially in the nieiiullary rays.
The bIoi>ilsii]i|»ly here repeats mi ii small wale the |)ortal system of the
livi'i", siiiit' a vt'ssel tbnneil by (he tinimi of capilliirie!< breaks up in capil-
larii-s "iiiH' iihire.
Knmi ihe siiiiie arleriiil arches at the boiimiary of the ciiriex and medulla
branchesi aiv also j:iven utf to the luedulla, that is to say, to the pyrami<i«.
ThiiM' niiinin^ in a sirai>;ht or rather radial direction down the pyramids
H» ui'l-riir iri'lii; but sihiu breaking up into bundles of smaller vessels also
ruuiiin}; radially, supply all the mediillurv substance of the pvnimidtf with
lilooil. foruiin^ eapillarv networks with meshes ehintraled rndiallr.
From ihe I'lipilluries of tbi' pyramids veins are gathered up, and these
runniuj: mdially iiiiwurti full into venous arches, which, like to and even
belter doveloix-il than the arterial arches, are jdaced at the boundary be-
twii'ii ibe cortex and medulla. Kolluwin^f reversely the course of the
ni'terit-s these venous an'bes, forming more luimerous aniistonioses than do
the arteries, tall into veins which, lunninj,' radially tietween the pyramids,
Join together over the jielvis of tlie kidney and Ibrm eventually the renal
vein; this, running in compiiuy with the renal artery, falls inUi the vena
cava interior.
From the enpillarii'i' of the eorti-x, includiii;r tht; medullary ray, the
bliHid, wmie of which, as we have seen, hiis (liis-^ed through the glomeruli of
the Malpighian euiwule-*, but some of whieh has not, is gathered up iiitu
radiate veins, which, riimiiuii radiiilly inward to the boinnlary zone, fiill into
the venous arches sjioken of above. .Vt the surfuee of the cortex the small
veins are apt to be arranged in u somewhat star shapeil fa:^hiou, and are
«piiken of lis rriiif fh/hl'i:
Relatively to ihe bulk of the kidney the renal arlery hiis large dinieusioiH.
Coming oft' "direetlv t'nmi the aorta, wher.' the hlood-|iressur<' in very high,
and being com i in ri lively shon, il allords favorable eomlitiims for an ample
• ■' i" ■i.T.lv'of the pvraiuida, consisting eliielly of eondiiviins,' tubules, is to
"" "^ ir'* extent distinct fmrn that of the .■..rie\. wline the tubules are
^^'^™rM«*t"lUt for its sine the kidney is most iUnmdantly supplied
* - In wclioDS of hardened and iirej.ared kidneys, the nrU-rii-s,
TO_ i'i-»- , i,_g extent the veins are emptied of their bl.Mid. and
»r.L.uf» M» ...lUns^ Hence, judging by such speeiiLi.>iis alone, the
-^ ^r,ili«r«.J-l»P^ J^,^, of tubules i.lo,,,.; but i. must be
ski:.- «"'*'' '/^.^.rinT life everv tubule U netted ronnd with fairly
.^TW z. mill.; ina- ^.''""^, ' „„'niore or les< lille-l with blood, and at
„^ «...:an^-m.h, a ^^^.^_ _^^ ^^.^ „„^ „,„ ,. ,„^^. ,,_^.
'™*^ "^4,.' -■ iTi-*"** <ii~ ^f ^'**"'' ""*■'"»■' 'I"""":-''' " '"■'>' ^■»'">' '■'^^y
■»«rl> Z *''"■ _^^ CMU. whieh binds together the lul.ul.s and blood-
m 1i^ -*'=**^!1"^;,™, ^T,a!l amount ente.v will, the bh.o.l-
-«!S!**C3r ■«»»' .■ . .J, i^rpif branelie^. but in the cortex
- -wTTOo- ^'|r^^l,anlhe basement memliranes of the
TUK STBirCTUBK OF THK KIliNKV.
533
I
I
I
I
I
tubulen, nilh u few ccinn«ctiv«-ll>«iie corputiclci' imhcditcd In m »cant_v lionin-
flliyWJW ikX iihrillntcil luutrix Iviiig )wtwC4-n them ; nroiiiii) the c«)wiil(w t)ii»
^itiVn* \» FHtlii-r mor<! al>iiii(liiiit thnti ? Ik when', und hcrr i» mnictimi'n
filiril luted, In the pyntmUl*. Mpo<.-mllir nt thrir timer piirtfi, « liirgt'r
niiiKiiiit iif u ciiiiilnr liomogcncoiH matrix, cooliiiiiiiig wniiDi-tivv-tiiwic cur*
E>ii]ucl<'«, In found Wtwi-co the tiilxiles; nnd *iii<M! lit-rc thfl bnwmcut ment
iniiHi vf t))v tulmlo i« luwd nilh il« Blroron, tli« tiibul* appMra u a
UibiilBr cavity hollowed out of tli« niHtrix or Rtrunin nnd iine<l with
vpitholtum.
Tbc whole kidney is eurroun<led hy n capsule, <xiii»iMing nf ordinary
coDlWctire liMue, and ('onlinuous at the hiliu viih iW connective tissue
forming tbe outer vrttlls of the pelvis and ureter. This cnp^ido may uller
<leath be peeled olT, Mid slender proceeacs of connective tisiiie, with sodw
bloodvMselfl pfUBtng from (h» eapaule into the c<nt»x. are iheu disclMed.
In (be scanty eiTonia are nuinerouH lyni|>h-ftpactB, the lymph from tbeat-
being colle(.-leil into lyinpliatic vrswU which in part lenve tlie kidney hy the
hiliifi t»f^lli«r with the blood ve^eeU, aiid in |)art run in the ca]«ule and
leave the kidnev on its convex surface. The capsule i» dewribn) n.i iH-p-
arable inW two Imytn, nud the lyn)]>lial)C vnsels run chiefly livtweeu thnw
lay en.
§ 400. An the rennl nrlcry pnsKv to the kidney it l» inveatitd hv n niinibiT
ar(tw<'niy or lei's, in thu dog u iloxcii ^r nion-) ncrvi*. arrnnEcl in n niL-xiiK,
the rnirtl filrjut, Tbe ncTvi^nrc compi^rd niirtly of nivdii) lali.it libm of
very ditlV'n-nt fitee and partly of non.nHtlullaitiJ filtm; niitneroui' snmll
ganglia, difl«ring, however, very much in rixe, arc scultered over thu
piciiu.
The mrrvn thus forming the renal plexus come cbieHv from the great
polar plcxu«, and appear to bo more immediately connected with (he part of
tfant plexus which is cwlled the sciniliinar ganglion. Tbe plexus is, there-
fore, indinx-lly conoccte<l with the nerves entering into tbe solar plexus,
Mieb as the right vaguti and the abdominal splanchnic nerree, great and
Hnall. UeMdes this llie ^lanchnic nerves appear to send tilamente directly
to the renal plexus; tilnments have also been traced to the left kidney from
tbe lell VBKus (which does not join the stjiar plexus), ami it is coutendeci
that filaiueutd from the right va^u^ also make their way direct to the riglit
kidney without distinctly et>mmiiuicaiin>: with tlie aolar plexus.
As ne flbull see there in exi><'riuii.-uliil evidence that. In tite doff, nerve
flbrea from the anterior rontn ot the eleventh, twelfth, and thirtnnith dorwtl
aninal nerve* and even u few lihreii fniiii Mill liiwt^r nerves liml their u-uy to
toe reual plexti* and *o to tin- ktiliiey. Tbtw make thi-ir way fn>m the
aympuliieiic chain, inlonhirh thi-y limt \tn*ii either hy joining llie vplanchnie
nervi^ low down, or by a niortt direct course, to tlie solar plexus, and thence
to tlie renal plcxtin.
Nothing verv definite is known of the termination i>f the renal nerves
within the kitfney. ^nie of them, and considering how vascular is the
kwliMiv. probably a large number, end in the bloodvesM^ls ; but some of tliem
must have other ending*. We have, however, no evidence that any of theoi
are coitnecled with the epithelium of tbe tubules. Since under abnonnul
rircumsiances alTereiit impulses sufficient to give rise to very great pnin may
(MBS op to llie central nen-ous system from ibe kidney, at least from the
pelvb uf ibe kidney, some of the fibres of the renal nerves are atlerent
flbrca; nnd soiue of tlie medullatni libres are probably of this nature.
Thk CtmixwiTioK Axu Characters op Uhike.
it 401. These are so flilly dwelt upon in special mirk» titat we amy cuuftiw
ounwivea here to uilietii poiiitn. Tlie healthr urine of man U a clear y(>ll»«uh
sli^litly tli>ure«cri)t tlu'u). of a (le^-uliar ixlnr, Mtine taste, and a<d<l ntictwa,
liaviiiKo Qxuin sjicvific gravity ol' lO'Jii. aud pii<-ntlly holding in i>u*{k>dmiiii
a liltle itiuciii*. Ttie miuouh, witcii prrii^iit, coiaw Iruin llw uriiinrv |>msni^.
as do al4(> tin: oircnitiinm! <![iitlH-Itnl ii'll*. All tlw rwt of tlit iiniiv innr t^
I'onHiitiired a* llie xwivtioii of ilx' kidni-y.
Tlii> iiriiii' n» vie huv« Mtid !■ the chief chaiiDol 1>r wliieli ■uUd mnitcn
leave iliv IhmIv. h ■mall <iuanlitv only jiaainnK hy tlii' xkiii and practimllv
nuiM liv the luiigd. Hviicir. tiegWting for lli« imwiil the -kin, we mar mt
Uial all till' iiiln'taiK-o takrti into \hv oody wouorr or later leavr ihe body hy
tbo uriiii;. Nivf iho fow «iiheliiiic<-s nhicli may bo reUiiHi) permatwntly within
llu- iHMly and the eutwiaiicos nliich iiiako iii> tbv budy al thi- nion)«i>( of itt
di.iilh. Wc acL-onlini;ly Hnd that the iiriiifi ouiilaiiis a large iiiiniher of sub-
•UUecv, th« cxiu-l ■iiiouiil of eaob sulMlauoe prceent id a nven quaulily of
uriM vtr>-iiig, in the rase of every atibatanoe «om«vrhal, and ii> (be cmm» uf
nuuiy «iil»t«iife8 very laruely. from lime to time. Tbe oonipoeitioa of urine
t> IM»* odIv complex but extremely variable.
Moreover a liille consideraiioD will ^low that tbe Mwnil sttlxHanMi
preeenl in urine niuM bave very diflereut hialoriea. ^lue of the con-
Miluenta of urine appear in it in the exact form in wliii-li ibey wrv iiiiiw
dticed into (be mouth ; they have been aimply absorbed from llie alirutroiary
<.-aual iuto the hloud and exvreted by tlie kidney wiibout undergoing chaogv;
tliev are dtTivvd din-cily and without cliang« fmm tbe food.
Olherti uguin an- thu prodiii-ta of obaiigeo uhicb ibe fof^l liaa iindergunv
tbe bixly : and lli<v« chaiigiv may bo vlight or may be extenvive, Nn<l ini
take ]ilao<: on tlx^ one blind in llin allnH-niary canal, or diinti>; a brit-f into
of the Nuhatance in tbo blood -nI ream, or wen in ihi- urine itiielf, nmy an
sjieak be ituper6dsl ; or on lbi> otbcr bund may tiikv pla<* in the very dupi
of llw tiwue* and Im- clwly aMOciatwl Wiith lbi> wry life of the Iwuva. V
ahall, hourvrr. have to return tj iheiN; mntien' liiii>r on. and m*y heri> hrtel
oinHKlcr wliiil NubetaiK'c* are, normnlly and alxiormally, (irMunt in urine, aoi
the Mvf features of ilie fluid it«elf.
MOS. Besides wilfr, the const itiient« of urine arc:
yitif/'j^nous erynt'i/liiie lio'iUji. Neglnrtiuc tiw Biiiull |>nifKirtion of lb
InmIics ivhiub, e«nccialty in (he caao of lleeh entvn. are incnMluced in(<i I
oCtiiioniy with the food, as kreatin and the like, and so pnss into the urii
with no or with oomimralively lillle chanf;o. wc mav on (he whole leKard
■ubnancea of tbia olaaa as tbe ptoduote of (he cfian^m which the prot
mattera (an<I allied sulistanoes such as gelatin and the like) present in f<
bare undergone either while tbe food wa« nmnly food, still in the aliinenta.,
oaual fur inatance, or after tbe food bad been built u]> into tlie tiwuea of the
bodv.
(Sf thcac bv far tlie most important, in tlie urine of man and niammalla,
la l)i« IxkIv urm (N H,(X>). It in the chivf form in which, in thm^ animab,
nitrogen leaves (Ik^ imdy. We ithnll have to diwcuM the n-latioii* aud forma*
tion of urea lnt<'r on, but nieaiiirhilc we will i^impty (tate that it han rcioark-
sbU double eoun^wti'itii' with two gn-at grounii. On the one hand it ia
ralated to tba ammonia ei''>up. and by hydration >• rmdily conv«rted into
annnoninm cnrbonau^ i NJ[,C^> t 'JH,0 - i NH r,CO,h On theotlter baiKl
it is related to tbo gmt cyaiiogon grouj>. aininonium eyinate and urM boing
A
Ubdln, «ilb K (til ci)niieel!ve-liH)ii<^ orpusrlett imbedded in h H^auly lioitm-
po^MM nut fShrillaUsI matrix Whig Iwtweeii Uiem ; aruiiii<l die t-u|tsi)lc8 Ihiti
Know ii mtlior nion.- nliutnlsnt iliiiii elMwherc, and liere it soRi«tinw«
B5rilla(«>d, In tW pyrnmiiU. e«pedttlk at li>cir Inner jiarts, a lai^ger
—T'llt* nt n oimilnr liiiiiiu)frnci>u« niiiirix, cotiluiuiriK coiinL-clive-liMtut.' vor-
Marlca, i* loiiiid liclitmi ilif lulxiirs: niid nince hi-ra the bawmt'iit lucni
•TKn* of thf lulxilr i* fuwril willi it* ntniiua, the tiilnilc nppcnn ni ii
■obtiUr cavilv hidliiwctl out of the miilrii ur stri>mn nnd lined irilli
^>JlJNJiuin.
Ilw wbule kHln«f ia PiirrouiKled hy n capsule, oomliting of ordinnrv
'"OiauiMtivv tiMuo. and continuous al the hiliit with the nmnrctivv timiA
fram mtlag the outer wnlls iir the p«ki« nod iireUT. Tiiie cn|wii]e niav after
'*^^Mti be peeled off, nnd slender pmcewcs nf <'i>nn<^'tive tiiwDe, with tonit*
IwinItonu pMHDft ffom iha capeule into the cortex, are tbeii discloeod.
In tiM aouitf Ptnima are iiuiuerDus Ivniph-spueee, the lymph from th««o
colleclef) into lymphatic vessela which in pari leuve the kidney hy tlw
tnjretlier wilh th« bloodvoKU. and in port run in the capsule and
ve the kidoev on ila oouvex surface. The cnpule is described as rep-
Tahle intu twu layers, and the lymphatic ccwek run chiefly betireen tlteae
ijer».
i 400. Ah the reuul aner\- (hums to the kidney it is iovetted bv a number
if <,lweniT or leu, in the d'>^ a doieu or mnre) nervea, arratived in a plexti»,
be rffiaf pifsnu. The nervi's tire compciwH nartJv of me<iu1lated fihnit of
~^err diin^rcnt nut and purity iif n<>n-me<lullateil fibrex; numerous nnall
^nnglin, difStriag. however, very much in nijse. are ttcatlcred over the
plecua.
The otrvt9 ihtit forming the renal plexii» come chicflv from lh« gift
N'lar plexuN. and ap|)car !i> he more imme<liatoly connected with the part of
that plexus which is called iho semilunnr gnnglion. The plexus is, (hcre-
fera, indirectly connected iriib the nerves enlcrinK into ibe solar plcxu»,
Mch aa the riglil vtigxts and the abdominal splanchnic nerrea, great and
wnall. Uoidea this the splaDchniv nerves appear to send filaroeiits directly
1(1 (he renal plexus; lilanientA have aUo Iteen traoed to lite left kidney from
iiie U-A v»)cus < which does not join the solar plexua), and it is c»ntendeil
tlwt filnmentB fr»m the ri^lit vngu* alio make iheir war direct to the rigbl
fcidoAf without distinctly cumminiicniuii; with the :>iilnr pivsue.
A« »e nhall we there i* cxmrimeutftl evidence that, !n the dog, nerve
SbrM fhmi the anterior loou or Ibo eleventli, twelfth, iind ihirieenlb doraal
•pinnl nervtM and rvcn a few fihn-s from still lower w'tvv* tind their way lu
Xnm rvnol pU-xiis ami wi to the kidney. Thiwe nmke ihi-ir way fnmi the
■5'ni|wllietir ehnin. into which they Ural pnxs either br jninin): the splanchnic
iMrrea low down, or by n nx^re direct courw.-, to ihe solar plexus, nnil thcnec
t*» Ihe rcDiil plexus.
N'tXhing very delinite is known of tho termination of the renal uervra
within the kidney. Some of them, and considering how vascular ia the
Vtdivev, probably a large number, end in llic bloud vessels ; but some of them
■BUit nave lUher endings. We have, however, no evidence that any of them
ate udinecletl with the e|rithelium of the lubulea. Since under abnormal
cirwuRisUncea atfereui inipulseit sufficient to give rise to very great pain may
fmm up to the central nervotis «yi*tem nt>m the kidney, ai least from tba
l>rlvi» of the kidney, wtue of the librtu i>f the renal nerves are alfertnt
flbm : and aome of the niedullaied hbre* are proluhly of this nature.
*
i
i
The Co«positii>s axd Characters op Urine.
§ 401. T)i«*e niv Hi> fully di*«lt ii]iou in ftjii-cinl work* tbnt i>v mnr coofis
ounelvca here til «a]iL-iil p(iint)>. TWlipnlthTurincuf Duinwa Hmr yellow t
allg^htly fluDKuccnt fliiu), of n [H'c-uliar <iil<ir, mViiw lnrt«, nii'l H(.'i>l rnwtMi
liavinga mcjin i>iii: !.■!/)(■ fjritvily nf I02il, iind gV'iK-mlty hiililin^: in MMpcoiia
II litllo niiiciio. Tliv tiiucii*. wlifii ]trr»«nt, romra from the iirimm-
iM il<i iilv< (hv (KTcaKiotiiil <^]ijtlicHiil cell*. All tho raU of thft nniH- nuy
cnnnidcn-d n* iheewn-lioii of the kidnrr.
Till.- iiritio as we hwvc (mid i« lh« I'liicf clianoel by trhich wIkI mat
lonvo Iho body, a rdkII <|UaDtilT only piiMing by thi? rkiii nod practicall
none bv tho lunge. Hence, iiefi;lecUiie for llic pre»ut the 'kin. ve lunv
that all itifi siibdtaiicee taken into the body soooer or later leave the boily
the iiritie, Hive the few fiiibelaDces which may be relained i>erRuiuently irilhi
the bcKly and the subalanees which make up the body at the niuiDcnt of i
desiJi. We aciyirdtti^ly litid that lh« urine cimiainii a lar^ number of sul
atanco. llie exn<-l aiii<>utii uf each Bubaiaiioe present in a civen quantity «>/
urine VArytuii. in the uate of every euhetance eomewbal. and in the caaa o/"
mzaj subslauL-eH very largely, fmtii lime t» time. The voinjMMilion of urin*
is not only comiilex but extremely variatde.
Mxreiiver a little cinixidcmtion will »hi>w that the aereral anbatanctt
]>re(tei)t in urine iniist have very ilitlerent hiatoricfl. Some of the caa-
stitiieiil.i uf nrinc appear in it iu the exai-l form Iii which they irere itrtn-
(iunil iiil'i the mouth ; they have been »imply alwirhed from the alintentaij
canal into th<; bloml and excreti'd by the kidner without uudergning clian)^;
thev nn: di'Hveil direcllv and withuul change Iram the fni)d.
Other* again are thcprtxluct* of chanp* whirh the )!Mid lia» iindergoiw ii
the boily ; and th(!«o changes may W flight or may be eklciMvu, an<l mar
lake place od the one hand ia the ulinientary oaiial. or during a brief Innat
of the Hubfttnnt.'e in the blood -stream, or evon in the urine itself, may io to
Apeak be Huper6cial : ur on the otlter hand luay lake place in the very depths
of iJie tiitwieB and be ciiisely awociated with the very life of the tituuca. We
shall, however, have tii return tj these matters Inter on, and may here hrieflr
ooiiBider what HulHianeeii are, noriually and abnormally, prewnt in urine, anS
the chief fcaturcn of the fluid itaelf.
$40S. Bc*idi?9 uiili-r, the consliiueala of urine are:
Xilr<,iirnoii» crytinlliuf liorlieii. Ne(rleetinj( the ftinall proportion of lhee»
bodiea nhich, twiurdnllv in the ciu^ 'if fla>h eateta. are tntr<)duced into the
economy with itic iuwi, ii« knuttiii aud the like, and a\.< pav tulo the urine
wilh no or with commralivdv Htile change, we niav on the whole regard the
■ubtUUKM of Uiti clasa a* tlie jiroduvtit «f the vbaagea which lh« praietil
matter* (and alli<'<} *u1)i>lun<'i* nurh a> gelatin and the like) pr<-^at in fund
bara undergoito eilhrr while the food wax «iniiily fund, rtill iu thr aliitu.iiiary
canal lor iiMiaiice, or after lli« food had bi-en t>uilt up into the tiwuc* of tho
bodr.
df theee br far the most important, in the urine of man and nuunmalia.
is the body urea (N,H,CX)). U is the chief form in which, in tiKse aiiintala,
aitragea hbtm the body. Weshall have to discun the relatiouH and forma-
tion of urea later on. but meanirhile we will nmply alate tluit it has n*inark-
able double conuectionB with two gmi groum. On the one hand it it
relatnl to the ammonia ifroup, and by hydration is readily converte<l into
nmmuniuni carbnnale lN,II,00 + 2H,6^-; tNH,',(-'0,i, On the other hand
it ia r<-taied to tho great cyanogen group, aimuouium eyaniile aud urea being
fADACTKR!} Of L'RtNK. ^25
I
iiKiiiuTic. fl»i] lliv former 1>r «iRiple Iifutmj; \mng converted itilo the Iniltr
(NH.-CXO-N.H.CO). '
TliMigh a hnw, fonniii^c ntlu with Hi-ida, such as Ditrat«a, osalalm, Hi'.,
(irctt oociini in Mrine in n iVw tuid iiidcjiemlvnt (."omlilion.
CIomIj aUit'd tn uo-n. occurriiiu ai>|i«rpiitly tw a bv« urwluet of the wiino
lin« of inetAbolitiii, in xWc acid (C,H,N,0,s whidi m found atnajr* iu ihe
urine of man, oocurrinji; in itnuiil but variable (|iinntity. la tlu* iirino of
BODW iniuMlo such as binle and rcplileH il ucnirs in iitiiindancc, and inde«d
in Ulfln replaces urea m the chief nitixiuiiKHiN i:xrrctii>n. Uric acid ia a
inore complex IkkIt thiin nr>ni. one molecule of nric acid :<pli[ting up, under
tite influenee of cerinin reagents, into two inolcculat of tirea and a com-
(loimd of oialie acid, ha [tecom posit ton product*, however, under diSbrent
reag^nlfi, are very uumeroii» and coinplox. though urea occurs amoDf; them
flre>|uently and eharacleriHlically. Uric acid may be synthctictilly produced
out of nrea aitd Klvcin fglvcocolj.
It is a weak dibaaic acid, and occurs in nortuat human urin?. not aa a fVee
acid but as an ucid salt, being combined ivith potassium nnd sodiuui. and t»
■ Um extenl with calcium and ammonium. In ouitc Dormai urine tlicM
mIu are aolubl« in the urine, even aller the fluid ha* cooled down to the
ordinary temperature of (he air: but not inffe<|uently the urates, soluble in
th« urine at the teoiperaiure at wliich it leaves the iKtdy, are precipitnt<«d
when tite tluid cuoU, fumiing the well-known "depiwit of urate*.' On
furtliLT ttandinf; Ihe salta are apt to be decomposed and thiu to give riae to
eryvtaU of urte acid.
Btaidea urea and uric add the urine contains small but variable (]UBntitic«
of inonvr leM nearly allied iMxlies such as kreatinin, xanlhin. Uv{H)xniiihin,
aiKl pianin. (Vmcoriiing ibeno w« will at present only say that kreatinin is
a bydraled (orni of the l)o<ly kreatiii which we spoke nf (i Ii2) a* a eonetiueul
of mu«cl«f. Kreiiiin hy hydration ia readily ronvLTttnl into ki\-«tinin. and
kreatinin by dehydraliKU inli) kreatin; krutlin intrndiierd into I lie alinitrutary
canal or into tin- blood appttir< iji the uriti<^ u.-> kreatinin ; and iu lle:°b enl«r«
some at t«!ii*l of the krvaliiiiu •>f tin- itriiii:' i.i di-rivinl dirvclly from the krealin
C-eaetit in the meat eat4.-n lu fund ; hut we sludl diwuM the subject of krealin
teroii.
BeeMea the abvve, twh bodi«f aw leucin. taiirin, cystin. allantoin and am-
mnnium oxalurain are occaeioiiaJly found iu urine, hut cannot be regarded as
cofutitueiits of normal urine,
In ihe urine of man hippuric arid oppeani to be always ]>r«flent in small
i]uanlilia, and in the urine of herbivura occurs in large iiuanlitics. [n theae
lailer it ia derived more or l<n» dirvctty, by eluinges ol which we shall have
to speak in a succeeding chapter, fruiii conMiluonu of the food containing
bodies belonitiog to (he aruuiatie group (benxoric acid series) ; but the small
iiuaniitv preaeut in man and other earnivora appears to eome from the
motaboiiani of j>rotei<l matter which, as uo have already seen, contain* an
aronatJe wittUlueni. Another member of the aromalic );roup, lyroMin, is
nocnuonally preeent in urine: and as more regular eunstiUienta uf normal
urine may be mentioi»ed certain phenol compounds, such as pheDysulphurk
acid, (be phenol coniitiiuents of which are derived fVom tlie action of niicru-
organisina in the alimentary eanal. see {283 ; these subetanoea though they
no longer ctwiaJu uitro|{en, lake uriKiu from bodtee of the aromatic series.
Similar ehauaea are al»» the source uf indigo compounds (iiidiean'i in the
urine, derivwi fr^im indol. !>ee § 'J4!f.
§ 403. lHori)»n\r Mitt. Thette for the most part exist iu urin« in natural
lution, tin- <y>m{)<Mitiiin of ilte Hdh almost exactly corre* ponding with the
- ilu a( the dinct au»ly>iH <<f ihe fluid ; iu this respect uriue cnnlraata
626
TlIK BLIMtVATtOX OF WA3TK ritOlJDOTa.
forcibly wiiU Wood. l\u> ntU i>f irlilch U Inrgfly oompawd of Inr.rii :li.
fltniicvs, wliit-h jirrviouK l» ihw iiicincriilitm fxixtocl iii |MictilUr . n
wilh prok-iil iiiii] <aImt corijiIok limliiii. In ihc ii»h uf uritii' ttiiT<- it ralbr;
more sulphur than <\>rmp<intl.< tu llu- Miljihiinc aci<! ilin-i-lly <lrrtvrmii>f«l ;
thie iuiJictiU« the «xi»ICDw in iiriix- of Miiiit! itul{>bur-hiil<litig cotuplvx bodf-
And there mrv IrsMs of iron, {xituliiig In »<itiH- ^iinihtr iroii- holding sub-
Malice. Hut qihcrwisc, all the suWativtw found in tbfi iwlt trxtst as wlb m
iIk imiura) duid.
The chief bftMa are aodium, potaniiini, cnlcikim luid magneeium in ilr
fonn of dlloridcs, ^osphntea nud aulphtilM. Th« exact way in nbich ikt
Bevenil haaes oud acids itre combinecl is to some extent a mailer of uii»r
tainty; but sodium chloride u ooriitinly |>resent and in cnnsiderabli; (luan-
tity; it it the inosl abundant and important inor^auic coneliluent. A Itttgi
Itortion of the phoaplioric add seeata to exist as acid Boditiai phuaphiUe, the
rest aa soluble calcium aud magDOuum phcaphatee. The reuainiag chMi
mlU, occurring. Iiiiw«v«r. In smaller quantity, are ]M)taa&ium aikd aodiiiin tnl-
phale, Bud calcium chloride.
Ammonia occure in Hmnll quantity, alkaline carbcHialea an frtHjuaUlr
fiiuml, tract'* of iiiiriili'H uru at ull eventii oci'ani'iiially i^reaeiit, aa ttloo ion-
cations pf ulicatcs and of i>ul|ihii-i-yaiiHte».
The |>lttwphiUe« are iltTiviil partly frxmi the )th<Mipha(M taken na such is
food. |iiirtly from th* ph.Hphoriw or phtuphaKv (ictuliarly awociatrd inlli
the protciiU. aixl {uirtly ft'ito llie phiupbnrui' of cerlaio oumplox fats eudi
as lecithin. When urlni' bvcoinca iilkulhic (and. lu we nhall preaeutly set.
it may da so by changes taking place in iu<i-lf ) the calcic and nia|;iMeic
phosphates are converted into du«c Mllif which, being i»»>liihle. are pte-
cipitated, the sodium phosphate retiminiiig in solution, When tl>e alka-
linity, as is ffvquently the case, is due to iimmooia. nmmonio-magneaiuin
phosphate is formed and is apt to ap|>e«r in crystals. The sulpbaten are
derived partly from the sulphates taken ns such iu food and partly from the
sulphur of the proteidE. The carboualee. when Dcciirrini; in large quantity,
generally liave their origin in the oxidaliou of such sails as citrates, tartraUs,
etc. The baiw« prttteut uejMind largely on the nature of the food taken. Thai
witli a vegetable diet, the exceca of the alkalies in the fou<t reapiMim in the
urine: with ananinial diet, the earthy bases in a similar way come li> the frooi.
§404. Xon-nUrot/enoiui boitU^. Ihese exist in very small quantities, aad
many of thmi are probably of uncertain oL-ourniiice. i^Hno of tlieM arc
organic avid*, the most constant |ierha|i^ being oxalic aold ; to thia may b«
added glrccriu-pli>mphDr!c. lactic, formic, aoblic, butyric anil possibly suc-
cinic acids. Intuit hax alio been mid to occur normally. It has boto maili-
tainc<l llint miniili' (piaiititicji of ^ugar ('(k-xlriBu.*) an- invariably pnjscnl in
even hi^allliy urine; tUix. howcVi^r, haii not iu> yet been placml Wiyuitd all
doubl. Tbe nattin- of iht; siibttanees uhich give to urine its charadcriMia
<Mlor bus not liecn mnile out ; ]iro)>ably thrn- arc more sucfa bodic« than one.
§405. Pi'jmcnU. Urine i» always colonil. Ihc tint varying from a light
to n ilark ycllon with an admixture of briiwn. In the course of twentv-
four hours, a not inconsiderable quantity of pigment must leave the body
by Ibc urine; but the nature of the normal pigment or pigments of urine
is at present obsciini and the subject of much controversy. The matter
is apparently further complicated by the presence in urtue of what bat-e
bi-eu called "chromogenf^, * that is to say, bodice which are not colored
tbcntEelvra but which reiwlily give rise to pigments upon oiidnlion; and it
is probable that soom of tbD»e " chrumogeus " of the iiriiie are rednctloa
pnMlucts of the respective pigments, the reduction lakiux place in the
urine atler aecretii>n, or during or cren before secretion. There is fretpiently
TUK coMi'osirios Xwl^irHfACTKKO upTk^^. 627
|irt«eiil ID uriii«, i-»|ici-iiilly in cmra of ffvtrr, » pigment nhiob has Ixvii
)ii<>lHl«() ami ilrlcrmii)).-)], which him ii thantrirritlic *i>o>-trutii, anii which
Wing iiininUincd br nnDio lu bo » drriviUiv* »{ biliriiutu, has be«i) oillcii
tiroifilin. It 18 not ihis iitttbilin. howi^vcr, which f(iv<e* to urine iu ontiunry
color. Some obeorven, on (he oth«r haml, ninii)tain ibat normal uHiio <Ion
coDlaiii and, id pnrt nt Icnst, owes iU Domial color to a soiuewhal Hiinilar
but dilterenl body, which In ocMi(ier)U«noe tbey have nllt-d " iionnal "
urobiltu. It is in fact Dot pomble, at th« present niomenl, tu make dclinite
and ealbTactory itotenienia aa to whether urine coutAiua one or n>i>rc than
uue normal piffinetit, as to its or llieir nature, a» to wlietkvr they arc (lerived
from bile iM^ment or directly from the hwmatin or htemosloMn or in otl»er
wajrs, or as to the several stepe by which tbev are pniiliuxil. Tliere are
iilao abnonual ovioring: matters prcoeut on ocnwion, auch for imtnnce as ilie
pvculiar reil coloniiK nmttt'r <iceurri»)t >oniet!nie« in tbo tirine of actite
rhvumali^m, wiiieb haa l>eeii caDed uroerytlirtn; htit our knowWgc coit-
ivriitnK lliftw nre very ininerfpct.
!i4W. f'rrm''nU and othrr htKlif*. Kven normal uriiw bn« frcqiientlv'lieeii
found t» contain a vniall <|nantity, hardly amoMntin^' to more than a trncc,
of prot«i<l material, a|>pari'ntly an albiiniiu ; but the nornml presence of
vv«D ibit amall <|uaiit!iy lin« t)«cn diHpulcd. Urine, however, certainly cvn-
lKin« funuiMit Ix-liiv.
Wlwn urine i« trcatMl with many \\a\vr it> volume of alcohol, a Knuiiilar
or docculonl pm-i|>itNtv is thrown down, cunnistinf; chielly of phii«|>hnt<«,
tAfCther with Rome other fubtttanotw or probably «i'vcral other snlminnces, in
very Moall (|Uantitie». An ai|Uvo(i« solution of tho precipitate, which maj
be freed from the pliiiephatc«, i» b»tb awvlolytic and proteolytic. . Porments
may alao and more rendily Iw esiractrif from urine by allowing Nbrcds of
librin to soak in the urine for a few hours, and then removinji and washin);
Jthem. Tbe ferment* become entangled io the fibrin in iiui?i a oav m not
lo be aaeily removed by wasliinft. The waabad shredi* nil! mnvi.'^ri «urck
into sugar; and when treated with dilute hydrochloric ncid di;^«t them-
•elvea, abuvriuK ih« presence of pepsin. By thit method it hii.-> b<vn asoer-
taineit that an aniylulytic ferment and pepein are pn-sent in <|unniitii-fl which
vary in the twentyfuur hours acoordtux to i\\k me:*!^. K-^nuin has aba
l>c(^n found, and at tiatea at least, trypsin. From tbi* it apiwars that some
of the fcrmi'iits uf the alimentary canal «c«)ie from the bwlv bv tbe unue,
Ix-ing proltably re-absorbcil directly from the reajwctivtt ((land ; tike (juantity
ntoreoTcr nhich ibuaeseapu ih intii^cuilicant.
A Riuall <piimtily of ga«, about \h vnU. jwr vcjit.. oiin hv extracted by the
mercurial pump from urina revdved direct from the body without exposure
to air. Tfie ga* no obtained coo^sls cliK-lly of carbonic acM. nitrogen being
ivary ■'*^i>tTi and oxvf^-n oocurrin|t in very umall <iuanlttiM or beiuff witolly
absent. The meaning of thift we nave alniady touehoil upon in ^leaking i>f
respirntton, see $S*K>-
$407. The quantilws in which iheae mullifariotw Wlies, all of which, as
we have seen, we may perlmps regard as coiMilueats of normal uriiM, are
pruMDl in ditlerenl apociroeiis of urine, vary within very wide limits, bi-ini;
» dependent on the nature of tbe fond taken, and on the coitditionn of the
iMtly. Tlw amount not of water only, but of many of tlw othcj- M-rcml
ooiiatiuients, varies widely and indeed rapidly, so that the pcrceutiifte o»m-
niKUtlon of urine will vary from hour to botir if not from minute to minute.
Tbo causes which determine these variations in tlie nature and amiviinl of
urine we shall study later on. Meanwhile what may be railed the averajie
coiuptMithHi of human urine is shown iu tbe followin); table in which the nci^i
gd \m*n aro put down aeparalely.
628
AuouNTs OP TB£ Sbvcral UniN.tRv CoNaTl-n'B^m• Pawbo
IX TwE.tTT-pouR Hctcid. (Afcr Paiikis.)
Bj an ■«»■»
mui or IK klloa.
IV. I Ul» ~
of b(dy>iir*«^t.
Wnter
1.1(M.0ft0gnioiiRr>.
£I.UU(>0 f;raniin«.
TirtuI ii»liiJ«
I.UmMi
Ure»
83.)>>0
O.AOUU
tJrIo Ai!(I
U.S3A
a.oo»i
Ilippiiric ndfl
(i.«0
0.0060
Krratioia
0.UO
0.D1W
I'igairnt, uiid
oiUer Hulutuiicea
10.000
0.1. ',m
Buljiburlc atiil
3.013
O.OHOS
Ph<««|j|nirii' acid
S.I64
0.04S0
riiiiiriiio
7.000
(8.21)
0.19(10
' Aminoubi
l).770
PoMariom
3.5D0
Sodium
11.W0
Calnum
0.26n
Ua){nfriiuiii
OSiir
T3.11IXJ
I; 408. Ttie iieiiiily of urine. The lienltliy iiriue or niim u ncM, owing t«>
thu pri.-M'iit-t- <<t' aui<l fioiliiini }>hi>s|iliate, the aliaeRce uf free iu-i<l lx.>iug ilto^wt)
|>j ilii^ fuel thiit UHliiiiii liyjioaulpiiiie ^'ivei> no nreripitaic. Tl>c nmotint of
eiility i.4 xlriml e<|ijivaleiil i<i '1 ;,'raiiiiu«i of oxulic iid<l in tw^niv-four hoLa rv,
. Jt tlii^ ili-yrw of adililv nl aiiv oue time varies mu<-b during: if'C "luy. tx^ing
in ■& invoret- ratin to llic atuoniit of add oecrMMl by tli« itomnch : tlits ^ il
cl«cmis»> nfler loin) ii> tiiki.ii, utid iiK'reiL<nt »)ptiii iw g&slric digmion cokkio*
to ui end. It vnri<.-» nitli the iinlnre of (lie IWI ; wiUi k %-<cet«ble diM cfar
exc»B uf alkniitf in t)ic fixid, Imng ^ci-n-Ktl by the iinii«, l«ads to allcKlin i ty.
_or at leaiit lo diiiiini»ticd nridilv, wti<'nMU title cfl^ct a wintiog with au
~ limai diet, in trhtrh tho n1knli«'« am l(W abiiiidnnt, nrtby bsMv nrc-
_ oiidemtinK. Flcnco iho urine of nirnivora is |.>?iieT«lly very acid. wl»itp
tliHt of Iierbivora St nlknliiK'. Thr Intirr, uhcn fasting, are for the Lain*
being carnivorous, living Milircly on their ovrn bodies, and hence ihdr uri"*
becomes uiidi;r tUoe circtimstaiiev« acid.
Tlic iialunil aciditv increaem for some time afler the urine has twen «3I»-
cfaarged, uwing to tfie formation of fi-esh acid. a[>pnrenily by somo kind oX
t^roientaliou. This increase of acid frequently cauMB a urectpitnliun of
urato. irbieh tbe previons aci<iity, even after tKe cooling; *» the urine, h<^
becu iiwullident to throw down. After a while, however, the acid n^ctt**"
give* way to alkalinity. Tbi^ ia cau»eil bv a couveraiou of the urm io ^^
ammonium carbonate ihrouj-b (he agency oi a ^teciBc " organi/e<t " f^rroei*- ^-
Thii> <ermi;nl ti» a gciit.'ral rule doea not make its appearance exiiept in uHk^^**
exjMiMfl lo till' air : it U only iu unheallby cMiditiona that the rermcntali<=^^^*'^
lakc»|>lnix- within tbe bladder, and in sueh ca*ea \* due either t» miW^"**"
organumi> iiitniilucrd into the bladder IVoin without, during ibi- into of inHlf
mentr for iiu'nncc, I'r to the action of an unnrganixcd iVrmrnt, tccreted a
parerillv by ihe wall* of the bladder.
■; 406. Ahnfrmnl foM*litnfnlt of urine. The ttructunil clemcola found
the urine undrr various drciinixtam-i-Tt tin- blood, pu* and muciiR corjiuwil'
epithelium from the bladder and kidney, and tipeninitoxun. To thewi
be added the n>-c«IIc<I " rnet« " which are eithrn* " e|)itliolinI casts," thai Is
■ly cyUixlere of ntore or \c» ultvml upitlieltnl celle »lied frura tlie tutiules.
(
I
THE SKOBETTOIT Or tTItlKK.
■MB
»lnicturrli^i). "tibniiDua" caola. wliich an cyliiiden of ]>cciiluir nmti-r'uil
nxfukle^l in th« liiiuiim uf ibe lubulea; ibe exact nature of thb mati-rial ia
nt {ire»ent a iiwiler of doubt ; il b uut alwitya lh« ttime, but apjKwn mA to
f tibriu.
The moat ctiintuou aix) iin[Kirtaiit abnurmal coiMtitueiitA of urinv urv
umni, giving ri*c to HltHiniinuri*, ami twjar, giving rise (o glycvuria or
<liitb«Kv». The Mliiblu pmlciili! giMicnilly )i{Kik<;n of m " albumin " id the
iirliii- differ in liifftront cnnv*. The exact tletvnnination of iheir nature i* a
niitttcr of wimi; (llfliciiUy, olnci-, m wc have «cen, we have in (litfcreiilJnUug
thi- viiri4)ti» proteitU to trust laigely to their buhuvior >■■ reganln precipiUtion
iipun the H(l<liii(iii of c-erloin MHoe boilim ; ami the ))m>enco of mliiM bodiM
in llir natural tirliii- introduce complication)). It wotili) appear, honever.
that ibc proleitk umially prment are scrum-nlbumtn and glomiliii ; the«e are
not however as a nile, if ever, preecnt in tbo Mine relative pruporliona M
in blood-plasma ; and eitlier the one or the other may be pment by ilM)lf.
A form of albumoee (i 20$) called Itemi-alh'imot, is Komeiimcs found, and
and indeed probably verr many distinet kinds of proteids are from time to
time pretem. If egg-albumin be injected into the blood it appeat^ in the
urine as egg-albumin, and peptone similarly injected apjMara ae peptone.
The sugar which is found in the urine of diabetee U undisliugiii«hal>l>>
from ordinar7>' dextrose; but whether it is absolutely identical with that
Ikody, or wlteiber the sugar in all eases of diabetic urine is exactly the same,
caiiiio* perhafie as yet be n^anled as definitely S6ltl«l.
When blfiod is mingled with urintt in tbe kidney and in the urinary |)as-
sages the citnutiiuents of the former an.\ of course, added to tliowe uf th«
latter; and whim, bh Mimi-tliiiej* hHp[H-ii», chyle fn>ni the lavteals ntakra il«
way into the kidni-yv, the urine contains the fats and other ccnstituentit of
chyle. Fal^ h»wcver, may be present without the urine being distinctly
" cbrlous."
Cliolosteriu, bilc-acidH, biIi.--]iigincnU, luid one or other of a large niioiber
of bodica arising from a dinonhn^l mrtaholinni of lite body, such as leudn,
lyrneln, acetone i in case* of dialxTies), oxitlic acid, taurin, cysiiu, and many
oihen ant alio found more or kn rreqiienlly : some of these. inde«-d, luive
bein regarded as iitmnal eonsUtiient*. Itcsiidea these the nrinu «erves as the
chk-f ebanuel of elimination for vurioii* bodira. not proper constituents of
f(MM], wlm-h may hapinn to have been taken into the system. Thus various
miiieraiis alkaloids, mits. pigmentary and udoriferouR mattera. may be passed
unchanged. Many «(ibetances thus occasionally taken undergo, however,
changes in ^uasing through the body : the most important of thew. since tl»c
changes wbich th«v undergo throw light on the meubolic procesece of the
body, will be considered in a eueceeding chapter.
Tbe SscRenoN op UBnre.
§410. Tlie facts nhich we have Immi in a preceding section concerning
the structure uf thf kidufy have Hhowii iia that that organ, unlike the oUier
sti'R^ling organs which we have hitherto studied, oonslsts of tnu parts, su
distinct in structure that it seema iiii]i(«ible to mist lite conclusKm that
ibtir Aioctions aru different, and thai tlie nu'chaniMU bv irfaiuh ibr tiriiw is
BKVMtd b of H doiililc; kind. On the one hand the tiibnli uriuiferi, with their
eharvderistie epillielium. seem nbviouidr to be actively Hording Mructtircs
oompnrublc to tbe Hireling alvenli of thi^ Milivary and other glands. On
the otln-r hanil iIh- Malpijiliinn <-a)i*iil<'s. with thi^r gluiiieriili, are organ* of
« pMuliar nature, with an nlmwt iu«igniliciiiittpitlielinm,and their structure
41
630
TllK KLIUlNATtON* OT WASTS PII0DU0T8.
irrrttnibly flUfffCCBtolbatthov acl rnlher its whiil idav bocnilet) in ii jjeuprnl wmj
a lllierinf; lliaii tu a Irulj' secreting; niei'lianisRi. ilenoe liiu nri.'^ii rJie ri««
which frtijuentlr bears Ihe ounic of SitwiDao. aiiim he whs lh« tint Ci ]tul ii
fi>rwur>l, timi reruin coiwtitueniB only of (be urine iir« eetn-itvl slVr Uir
liwbioti i'( >ilh«r settrccing glands by ibe Mbuli iiriniferi. and that the rort iiT
(h« i.i)Di<litu«utA, iocluding a great deal of the wuterwiih such highly •oluU*
mmI (liBiiaible wlu as |>r«'xisl iu adequate quantity in the hliKxl, iiiT, n» k
wera, filtered off kv lii« glornoriili of the Mal|UK'>iAU ca|wile«. Wo ahall M
later oil rciunn tn clouht wliiititcr we are justified in applying the t«nD " filta-
tioD," whicli )iiw a tli-(ini(i> phyMcal lueauinfc. M the proeeM 1>y which water
and oth^r tiiilR'tiuKX'" |>iu« fh>Ri the bloodv^M-ln of llie glonivruliw into tk*
liitncii of tlic- tiibiilc : fur thiit ]>rt>ceM in, ait we ahall find, pcctiliar and com-
plex. But mich a d<Miht n«Ml liol prevent u* frnm rvrcwnizini: that the wbole
art of Bccretion of urinr ooimkUi of two purl*, one of wliii-h is much more
cloeely depeodeot on the Sow of hlooil thnnigh the kiihii'y than is the urdi-
nary proccae of aecrotion auch a» han bitlMirto vome hcfiro us. and aaMber
part which seenii to bear ihe anrac relation to the flow of blood as does
ordinary secretion.
That the work of tti« kidney is to an unusual degtM dependent on tbt
lion of blood ihrou^b it, secnii FUgpsted by tbe vascular arrangenK-nlfl ; for
tbcao are extremely favorable to a full anil rapid streoiui of blooil iJirouicli
die organ. The abort and relatively broad renal artery conwa off direct
from the abdominal aorta, where tbe blood-preasure is extrenielr high ; the
ntnal vein opam directly into the vena cava, where the blooa-pnwtirr if
extreuidy low. Between tbe niouth of the renal artery and the mouth of the
nmal vein tlie differeuioe of preaaure ts very great, indeed ; and, na we have
Been in treatin^iof the vmu-ular system, it is thedll^reneeof preMtir« botwwa
two [Kiints of tltcvajiciitnr tract which l» the actual cause of the flow of blood
Irom tbe one point t<i the other. The difference of |if«isure, indcfHl, which
drivf* the hliiod ibmuf^h t)i« linittetl area of the kidney is tlw Minic difference
of praeun^ which driv^w tlio hliMxl along the abdominal aorta down both
lea back again to the vc-na cava.
This Itc<' and abundant supply of blood is regulated, is either increased or
diminishcii, according to tli« noudsof the moment, by (he vasomotor system;
this is shown by experimental niid other results, which it will \w pn>f!t«hle
to study in some detail. Before entering into these details, however, it will
be well to call attention to the fact thai when vasomotor events modify (bt
flow of blood through an orf^'ao, they produce their efleets in one diriKlioo
or another by workin;; on arterial blood-preMUre. Thus, as we ahall me.
when stimulation or section of a nerve inoreasea the blood through tho kidney,
it docA so by inereadiuK the premure in the small vcmcIs of the kidnev, iot'lud-
ing the eapillary loojis of the glomeruli. In »uch a caw the wnlls of tbo
glomerular loops, tlinnigh which the pawage of materinlii t4> form >!|Hirt of)
the uriite takea place, are subjected to two iufluenc«s — on the one hand to ■
AiUer, more rapid flow of blood peat them, and ou the other to an increwe
nf the prewuTv which that hlood as it posees along exerts on them. We
ahall )ia\-e snihstHpieiitly t» iliscusa tbe share taken by (hew two influences in
determining luid niinlifyin^ Uie passage of material throu^jh the walls of the
glouu-rular liio])i< ; and this will bear on the quwtion of fillnition to which
wo have above alluded; but fir the pnv^nt it will lie convenient to deal
with the eflfeet« of variation in blcml -pix-Mure apart fo>ni this Eccondary
qucMiou.
ji 111. Thf vatamator merhattunu of lh« titlMy. It may he shown experl-
nionlally that tlie kitlney is supplier! with n vasomotor mcchanifun as well-
developod, IwhajH. ii:> tlwt of any vtlter jmrt of the bi>dy By means of a
'hKTION or UBI!fB.
681
mmliiicntiiii) of die |^elh}-«m<>gmj>li (Pi^ Io.i, ISti), we cma rwdUy ohwrre
tbc vnriaii,nis wlikb uko J»l«c« in tlie vuluiue of tli« kidn«j.
The inslrninent coiubla of two iisrtii. one of which (Fie. 155), wllod thi> oaro-
in«I«r.' U >pplip<) to tbe orgao about to be «ludl«d. while lh« other I fit;. l-'SlI),
c«lle.J Ihr oii.xiicrapb, i* Ibe ncordtur iwrt of the appitjatiu. Anjr (limlnution in
tb« volume uf lli« orjpin (Fij(, IM. £). kidavy. iip\rea. tic., h the caee m*y be.
diiniais)ieslli«pNi)*uru AnlbcDiiiilin th«<clllllIib(^^<I;llOlIl«oftbe0ui(]in thecl»m-
b«r ^(Fig. 130) Mcoidlnglj jmumm ihroueb lh« cube A'lFig. 1S6) &Dd liie lube T
<Flg. 15.^) to the diatnbera: the pbton j^kccordlnglv faWa aod iritb it the lerer
/r. Simttuly ma iocroiwe in A* volume of the orgau cauiee tbe lever lu riie.
a
^
I
Rkhal QncMum. (Seen Id nMUon. Monl-ilUcnunnMllc. i
f\ r, tomI* aiiil nama Imtwlilod ui Ui. otc. raloriiiR tilla* atoratn: ft, C< auil r. (^
'<»ulw *»i lancr niaUil mimiln nnwml UfvUwrbj'Ui* tfw AaiiJ bnlillnf Mmm lti(ai)lui(dE«
aribauioiiiboiMo jr.wMchappUM ltMiriollwiiirIlu«arih(Udii«ir and fimuvtih tbcmeUlnp
Mile two dMiotoixa uul B, una ol wbtcib^Jti Iteloaol b; a pliur dllliii; iho aptiiJDit It, hLIIu ib«
oUier (n) eamniiatnUM lir* lube T* wlib ibo nKmliiig liiMiuuMnt, The otbrr op«nlna. CimbM)
bi «taa*4 br a omU tap). U IMiihe pori"^ «< illUni Uw ohaiuber a wKb mrm all alto tbo KUiwr
haatMM itewd la UmIok, Ih* oUwr i-hainba JI baTlaii been |<n>Tioial7 laitlj UM. tbaquasUl)'
InlmdDcaA lalaUitaiiianiUniiitua ti«UMor tbaUdner.
The volume of tb« kidnoy tunj bt incrvased l>jr a swelling of itif ooti8titu<'iit
cells sod otli«r structural ckmindt, br an aucumuUtion of lymph in iu
1vinph-«wow, and by a i)ixt<:iitioii of it» ltlti<Klv»iM)l». Compared with
lite tliira, ihe two former ciiuecs are in lu-jillh »o liiMgniHcunt M>d iiroh-
lematical (hat thoy may be disrvganltxl. Purlhcr, the diatDDtion of ihe
binodveaaeld will in graorul de|>end on the constriction or dilation of the
renal arterim and tbdr raaiilicatioaB, for duWntioii duo to venous obalruction
Kms Mmi^ bulk.
es2
THE ELIMINATION OF WA8TK PB0DCGT8.
win only occur in 8pecial cases. Hence variations in the volume of the kid-
ney may be taken ai; a meaaiire of variations in its vascular supply — incroK
of volume indicating dilated renal veseels, and decrease of volume iiidicatiii(
conBtrictiou of the renal vessels.
Fio. IM.
6khi-[<iuirahii*tiij SKCT]O^A;■ Vrtw or ONi im>rai-i<. [Half nHluml bIz«.>
A', tiibt connecting limninipm wlih unciiiiictiT -, D. iilsi"ii Uniting on oLI contained In (tae niftr
U; Itiv oil 1h prvvuiitcd fixiDi evcapliij: by 1h« wlv of flu- pin^u by ihc delicate Heilble menibt«tw£
whirh iloen not liitcrriTC uilli (he iiinvticictit.^ of I lie plilon : Jl. ri'OjrUiiig lerer coDUccled with ilie
pitluu bf a necllc fi innslnG tbroiigh Itie gulilee F. f. fin- i^tieu c l>. for ihe purpoHe of clamping
the edKeiiKbe membmne beineeii the two rlntt-sbuiicil -iiiniivi'i bi ,V, wlillu ihc itlde tubt L li rortlie
pnrpo«B of lillm^ Ib^j in^lrunivnt.
When by means of the instrument jtiat Jeserilied » tracing is taken of the
volume of a kidney in what may be considered a normal condition, some
such result as that shown in Fig. li>7 is obtaiueil.
Fir., l.-J.
BL 0 u 0
P n L ^ z u p, £
KfDNcr CUBVE
Bl.Oui>-PRE(t<ri(E TraI'INu, ami Ci Hve rtioM Henai. ij><.'<»it:TEU. i^ntnml size.)
The Lilood-pretture abwliwiL line hut been mlKil 'l.i:' cm. ilbe lU'riinL mciilnni blood. preoEiu^
bavtng been 115 mm, Ilg.). Tbe ilme-cuTve |(lviw lriiL-TrU['llon^ rvcurrln^ t-very ibTri- efooads.
The volume of ihe kidney is seen to be so delicately responsive to chaagea
in the mean arterial pressure that the curve reproduces almost exactly ■
blood-presaure curve, showing not only the respiratory undulations, but even
TUB tiKCRETION OF t'fllKK.
fi83
I
I
I
I
I
I
tlic rite nii<) full due ii> Uie imitvidiial li<'arl-)>f!nU. With each ri*e of tneaii
iirtrriat prcmiirf nv)re blood iit driven into ihn roiiiil v<u»i:U and the IcidnHy
HWclU; with I'licU I'all of prx^triiirL' Ikm l>I(i';d i-iiti'r;< and llir kidney shrink*.
On other trectngw tukci) in thi> >uini« wnv may (dlRii he ■cim (not (hown in
Fig. l'>7) the nidi;r viiriAti<.)nit oiminnoDiling to the Tfiiiihr-H<iring cnrvc«;
but it will hv iiWrviil thiii in ihit* the kidiioy shrink* with the ri»; of iirmc
Dure Mid BWvlU with 1\k fall, F»r lut w« hiive socn (^ 3^:i} tht^ rise in thu
Tmoho-Hrriiig nndiiliitii>n it due to an AUEmetXation ofporipherid rosi«timce
c»iw»l hy th« ran>>triclii>n of miniito nrlents; and this constriction occiir« in
the kidney lu clMwlutre; the reniil arlcriolM Uikc ihoir »hnre in nrodiieing
tb« reeiiU, mid iu oonseniieuue of iheir tonjtriclion the kidney shrinW. Simi-
larly tiK relaxation of tne renal veig&eU contribules to bring about the sequent
All.
i 412. In the course of a discitiHian in an earlier part of thia work (S 171)
on the local and ((eneral elfccls of arterial constriction and dilalii>u. we saw
that the local blao<i-i>rwdure in and flow of blood thriMiijb the capillaries and
other minute vea»eb of thin or that vaaoular area may be inemaieil —
1. By an increase of the ^ueral blood -preaaurA bruuKht abmit — {a) by on
incrcHMnl fonw. frtniuency. etc., of the heart'* heat, (b) by thr i.-onstrititiou of
Ike Mnall arteries siijiplyinK ari.-tk> other tlian the aiVA !» <|ii<»<tii>n.
2. Br II rwlnxatinii of the nrlisry (ur arteries) Aupplving the ar<.ti ttMdf,
whirh, whilo diminiHhtn^ the |ir«w«ure in thf artery itwcif, incn-iww the prwt-
sure in the cnpilhirii* iin<l »mntl vmou wliirh the artery ifupplicii. It nucil
hardly W add<->l llmt thii local n^Uxnlion muihI uoi lie uccomjuuiiod by a loo
great dilMion elsewhere.
The tnnK' lo^al Idoud-pretMiire and flow of blood may similarly bo
dimiuitheit—'
1 . By a constriction of the artery' of tJie nr«a itself fand it« branches),
which, while incn^aini; the prmsiire on the canlinc side of the artery, dimin-
islws the iiri«sure in the capillaries and vi.'ins which are supplied by the
artery. This ajfain must not be accompanied by a too Rreat cunsiriction
elsewhere.
2. Bv ft lowering of the tjewral blood-preasure, brou;>ht about — (n) by
diminished force, etc., of the heart's beat, (6) by a jteneral dilation of tba
nmnll arteries of the botly at lurjre, or by a dilation of vascular arena other
than the area in question.
Applyiti;i; tfawe considerations Vt the bloodvesseU of the kidney, we should
enpoct tu fiiul the follouiiij; :
A rise in g«i«ral bloud-presanre, and that mean* a rite vf preMure in tlie
abdnmioal aorta at the mouth of tbi- renal iirlrrr. will cnusK a greater flow
of blood ihfoueh, and au an expansion of the kidney, provided tliat the renal
nrteriiw ihenivelvm are n'>t umliily c'lnstricted at the same time. This is well
sh«>wn, IU we have settn, in the curve tfiven nliorc, where the incriMM »!' prm-
Hurvdaetowicb bi»rtdMrat,M well il* ihnt due to ench n-«pirali>rr inovcmcnl,
l>uing of onnlral origin and n<it due to nrUTial ci>n*iriction and being unnc-
oompunieil by any oompen-ialin^ oonstriclion of the renal artery, leada to
cxnanaion of the kidney, tlial is, to u greater flaw of bluod through the
kidni-v,
If, liowever. tlio rise of gtriieral blood -preMti re be due to events which at
the same linxt canse a eon«triction of the renal arteries, tJie flow through thu
IcidiH'v may not only not be increased but even be diminished ; the kidiwy
may shrink instead of expanding. Thus tf dyspnoea bo brought about, M
by stopping artilicial rcepiraiion during an experiment, the kidney at once
shrinks ; the too venous blood xtimnlaieri the vasomotor centre, and probably
aba by direct action on the bloodreMeb leaib to a KCneral arterml constrio-
634 THE ELIIIIXATION' OF WASTE PRODUCTS.
don and so to a riee of blood-previire; but the renal vcskIb are involved is
this constriction, so much so that their constricted coudition more than coun-
terbalances the general rise of blood -pressure, and less blood flows throng
the renal vessels, ^o also when the medulla or spinal cord is directly stinii-
lated hj induction shocks (the animal being under urari so as to eliminate
the com pi i cut ions due to contractioas of the skeletal muscles) the renal vei-
sels share so fully in the arterial constriction which results that, in apite of
the great rise of mean pressure which is induced, leas blood than normal
passes through the renal vessels, and the kidney shrinks. Or if tbe abd<HDb
nni splanchnic nerves be stimulated, since as we shall see these carry vaso-
cuuBlrictor Hbre.'^ for the kidney, in »pite of the rise of blood- pressure which
folluns, the kidney shrinks on account of the great constriction of the r«iial
vessels.
On the other hand, if a rise of blood-preasure be for any reason not accom-
panied by n compensating constriction of the renal arteries, that rise, whether
It be brought about by general constriction of arteries other than the renal
or by an increase of the canliac delivery, causes the kidney to swell, showing
a greater flow of bloixi. Such a conaition of things may be induced by
section of the nerves of the renal plexus, whereby the paths of all vsso-^»&-
strictor impulses to the kidney are blocked. After this has been done a riee
of general pressure whetiier by dyspntea, or by direct stimulation of the
spinal cord, or by stimulution of the abdominal splanchnic ner\-e8. leads to ■
greater flow through tbe renal vessels and an increased expansion of tbe
kidney.
A rise of generiil bloo<l- pressure then may be accompanied by either s
shrinking or ii sivelling of the kidney, by either ti greater or a lesser flow of
blood through the kidney, according to the con comi Cant condition of the renal
vessels; or, indeed, may under certain ti re u instances be accompanied by no
change at all in the renal circulation, the local etlects exactly counter-
balancing the general ones.
Conversely, in a similar way, a fall of blood-pressure leads to a lesser flow
through the renal vessels nud a shrinking of the kidney unless it be accom-
panietl by a dilation of the renal vessels out of proportion to the general
fall. Thus when the spinal cord is divided below (he medulla the fall of
general hlood-pre-wurc is, as we have seen (§ ll'A) very marked, being due to
an iibolitiim for the time l>eing of wonted constrictor impulses. The pres-
sure in the aorta falU rapidly, and nt the f^ume time, owing to the more open
pathway through ilic region of (leriphcral rcwstjince iu the body generally,
the pressure in the vena cava is increasi-'d : the dilicrence of pressure between
the mouth <jf Itio renal artery in the aorta and the mouth of the renal vein
in the venu cava is so largely reduced that in spite of the concomitant relaxed
condition of the renal vessels themselves the flow of blond through the kiiiney
is largely diminished.
It will of course tie understood that, the general blood-pressure remaining
the same, the flow through the kidney will at once be im the one hand in-
creased by dilation and on the other decreawd by constrictiim of the renal
vessels themselves. The constricted or dilated condition of the renal vessels
can by themselves produce hut little vHect on the pressui-e either in the aorta
or in the vena cava ; and the dillerence between the jircpsore at the mouth
of the renal artery and that at the mouih of the renal vein remaining the
same, the more o|>en passages of the dilated renal vessels must lead to a
liiller, and the narrower passages of the constricted renal ves3i.>ls to a scantier
flow, through the kidney.
§ 413. By means of the oncometer, watching the shririkinc and swelling
of the kidney and thus jutiging of the flow of blow! through it, the results
TRE SKCItETION OP tTRIXG.
nan
I
I
t
being nlwiiys interpreted with rellerenoe to th« gejieral blood-preaaurv on the
llttee of tbe nbove discuMiioo, ihe |iatlia of vaeoiuotur imptileeB to the biilitey
baT« b«en a))|)roxininlely ninde out. Vaso-constrirlor 61>ns fur the kidiioy
are Mpi>lied from nbat we have iirevioualy (^ UVJ and eUewher«) spoken of
OB ibe Taao-ioDHtriclor regiim of the spinal oord. They ii-»ue fn>m Uie vpiniii
mrd bv the aolerior nx>ta of a large number of lfaS~3}>iDat iiervea tiikLug
ori;[ii) troni thia region, and may be tncml (in the dog) iu> bigb np an the
MXtb diintal. u few [lerbujM even to tbe fourth <)oniul, and iw Kiw <l<>wn tu the
aeooud lumbur (lourth lumbar ir»iily lhirt<t;n nerves be (^xinusl iw ilorMiI) ;
but im»t necm to imm hy tbe elevcJuli, tweinii, and thirleejith dunml iicrvea.
;h tne eorm poll dill);; )2;nii);lin nf llin upliuicliiiii' (Dvinpatlietic)
cbniii, ihi-M; tilin-x reuch the »[iliir {ilesiifi mid thiix tlie nrual plexu* )>v the
I
I
abdoiiiiiiHl «{iliiTK'liiiii- Rurv«; ih'iu', howuvrr, ei>iniHg from »iiiiu of ihv lower
nerves ■p|>nn'ntly<lo not cDiiinhuio lo the KpImK'bnic nerve, but takeaMt[>a-
rale rounte. Onlrifugal Miniulatii>n of ih'rse anterior roola produces shrink-
ing of the kidney, ah the more marked and diHtincl in the ca«o of the
elevMitli. twelfth, and thirteenth dorwd ruot« because the oHecl on the kidney
b tlMfl not M> much masked by vasomotor effects on other organs, t^timiiia-
Uon of the higher roots also produces shrinking of the kidney but leas
toarked, »inee In these caaes the stimulation bears at the same time largely
on vaso'constriclor fibres for other abdominal iirgvns. and so by miging the
l^eral blood -preHauiti tends l» neutralise the local elieel on the kidney.
And even tbe very decided shrinking of the kidney which reaulu from the
etimnUtion of tiie mlauchnio tjunk itaelf is lean than would take place if the
Himulfltion atli;cled the veK«e]s of llifl kidney only.
ji 414. We stated in j 16^ that liy tbe mediod of slowly repeatetl rhytli-
uical aiiniuiation the prmenee of vaao-dilator fibres in tbe sdatie nerve
niiKht he detecteti, though tlicM are largely nixed with vasoK^onstrictor
fibres; and sliiw rhylhiuienl vttniutatiou of tfiv luiterior rooU of the ahove-
roeuiiiiued lower dorml iierveM l<wls. not. an don> ordinary rapidly inter*
rupted stimulation, to abrinking, but lo awelling of the kiitn«r, showing thnt
iheae mota euoinin vaaodilaior tihrw a« well lu viwi>-consirietor lit>re«. The
higher (anurior l rot)!* iilim appear to contain some renal vaeoililalor Hbrm;
but i)k- cSiKl of etimulatiii); them bv tiie slow rhythmic mctlioil i* more
m«*ke<l by a concomiliuit dilation o^ the veaada of the other abdominal
ofgiuM, the roots in <{uest4on eonlainiiig vaao-dllator as well as vascMWDalriotur
fibres for th<«e organs; this leads to a fiill of genoraJ blood-preaauiwirhonby
the tendency of the kidney lo swell is oounteracted. As far as can M
■scertainei) at preaeni tbe paths of tbe renal vaso-ditator libnis are similar
lo iboee of tbe renal vaso-coustrictor libroa.
Tbe kidney then is well aupplied. espedallv tiirougb the anterior roola of
lite elevenili, twtilflh, and ihirleenlh dorsal nervea, with vaso-coustrictor
fibrw, anil is alui xupplied with va^ki-dilator flbree. Borne reeulls have
aeemed to .ihow that the libn-s passin;; along the roots of one side of the
•ninal eortl guvi-m ibe veieeU not only of the kidney of the same fade, but
UM lo a irvriaiu extent of the kidney »f tn« otber sule ; it seenia doubtful,
bowtver, whether ihi* i* really thi- civn:
There is no Mtisfai-tory evideiicr that the vagu:> nerve of either side oon-
Uline any vaaomotor film-x renchiiig the kidney (sec S 400).
Ji 418. It is i>bTioiin (hen that by niejiu* of this vasomotor mechanism the
flow of blood through the kidney is governed by the oeutral nen'ous system
in sucb a way that nlTerent impulses, started in this or that region or lur-
face, and (laSEine up lo tbe central nervous system, may lead cither to con-
atriotion or lo dilation of tlte renal vessels; and to sueb actions of this kind
we ahall preaeotly rMuni. Meanwhile we wish to udl attention lo tbe fact
J
58fi THE ELIMIN'ATIOX OF WASTE PRODUCTS.
timt tbc volume of the kiilnev is remarkably sensitire to chemical chuga
taking place in the blood. 1?be injection into the blood of even a small
quantity of water causes a transient shrinking of the kidney followed br a
more lasting exjtansion. The injection of urea and some other diuretics
produces the same cfTect to a more marked de^'ree, leading especially to ■
swelling which lasts fur some considerable time, while the injection of normal
saline solution, and especially of such diuretics as sodium acetate, causes an
expansion from the very lint, the primary shrinking being abisCDt. It it,
miireover, worthy of note that these ettects of diuretics and of chemical
changea in the lilood are observed even after all the renal nerves have
apparently been completely severed. Hence the changes in volume canaed
by the ])re8ence of these substances in the blood must be due to the sub-
stences actiug either upon some jieriphoral vasomotor mechanism, or, evea
more directly, on the bloodvessels themselves. It may he added that they
will produce considerable effects in the kidney itself without appreciably
modiiVing the general blood- pressure
S 416, If, while the kidnev is iu the oncometer, and the various experi-
ments on section and stimulatiou of nerves and the like are being carried
on, a cauula be tied in the ureter, the secretion of urine may be n-atched at
the same time. It will then be seen that the How of urine through the end
of the canuln is not equable, and does not either increase or decrease in an
even manner. On the contrary, it will freijuently be found that a sort of
gush of urine takes place, several drops following each other in rapid suc-
cession, followed by a cessation of flow; and if the ureter he watched it
will be seen that the gushes of urine are synchronous with waves of
peristaltic contraction sweeping down the ureter. Obviously the urine
collects, to a certain extent, in tin' [iclvis of the kidney, and is driven
thence by nmscular action from time to time; to this point we shall return
later on.
Making every allowance, however, for these irroj^ularities of flow, we may
take the rate of flow from the end of the canuhi ns a measure of the rate
of secretion; and it i.'' found that as a general rule increased How of urine
is coincident with swelling uf the kidney, that is, with a greater flow of
bl<MMl through it, and diminished or arrested flow of urine is coincident
with shnnking of the kidney, that iri, with a diminished flow of blood
through it.
A striking instance of this is afl<>rdod by the ex]>crimeut of dividing in
the dog the spinal cord I)elow the lui'dolla. The blood -pressure then, as we
know, falls rapidly, owing to the removal of constrictor impulses from the
small arteries and tlie general diminution of peripheral resistance which
foUows upon so many small arteries becoming dilated ; and though the renal
arteries probably share in the general ivlaxation yet, owinj; to the fall of
pressure in the aorta conjoined as this is by a corresponding rise of pressure
in the vena cava, the flow of blood through the kidney is largely <limin-
ishcd. We lind that after the oj^ration the a'cretion of urine is greatlv
dimiutsbed; indeed, in nio^t rases, the flow from the end of a canula is
almost arrested. In fact, we may almost make the general as.-icrtlou that,
when in the dog the l)IiKid-|)rcssure falls to abr)ut SO mm. Hg. or leas, the
secretion of urine Is for the lime stop|K?d. These and other results support
the view stated above that the secretion of urine is in quite a special way
de[>endent on the flow of bhn)d through the kidney ; and we may further
conclude that the secretion which is so particularly influenced by the flow
of blood is that ."pcclal kind of secretion, allied to flltration, which takes
place through the glomeruli, and not the more ordinary kind of secretion
by means of the epithelium of the tubuli uriniferi. But before we proceed
THE 8KCRKTI0N OP rBIKE.
S7
itSt
I
C^dklillMllSott tW incn>(u>F(( flow of blixKl iiicraue« lh« glomeruliir flow of
'arint. m must turn (o voiMitlvr Iho fiinvtiong of the ppilh«Iiiim of the
tiibtilt.
SetrttioH Ay fA« /tf»al Epilhriinm,
S 417, The glomerular ineclmiiUin ia. nflcr »\\, a, small portion only of the
whole kidiMV, and the epithelium over a large part of the course of the
tubiili urinifrri bf»n mo»[ dislinclly the rharact«rB of an active iecreting
epilhelium. These ihctA would lead a* ri priori t<» BuppiMe that ilie tlow of
urine ia iii part the raull of an aaive »eerelJo« coruparahli^ to that of ihe
salivary or other ^.dan^U whicli nc have already ntudiud. And we have ex-
petimeatal and oiIkt evi<lenoe that Muoh in lh« cnf».
In tJte (init pliici', ii ihiw of uriue niny !>a uriilioinilv oxt-itcil oven when
the DBlura) tlow liiu hwu iirrtnU'^l by diminiiltoo of btoott-proMtin:. Thus
if, wlieo tli« tirint^ hag cwuwcl t<> Hon' in riiriKiiciiiencc of R werlioii of the
medulla oblotl^ta, orrlniii i<uht>titu<V)<, Ruch lu ur<». urate*, sodium ai-ctnte,
and iW like, he injtMwl into the hlotid. a mori; or hM <»|>iou<i wwn-tiou is at
once Mvt up. Thw aecr«tioii is, or nt h-iuti may be, unnccompuDiud by any
riM of f^envnl blood -f>remu re suffiduni to iiocouiit fur the incniued secretion
M the mero mnit of an incrcnsod flow of blouil. It is true (lu wv have
Mtn, S 4]>>) tliiU the injection of those pitlMttinrM leads to an expnoBion of
tht kidn«y. an oxpnoeion which is probably due to a local dilation of the
snull notl arteries; but the flow of uriuo nhioh is olMarvivl in these ciMOS
■0 too mat to he accounted for by an<r increase of How of blood which ths
local dilfllwu mar bring about ; and lienc« we concludi' (hat the iucivast; of
fiecr«tion is of a diBereat kind from that which folloir? upon merL- inoreaM
of blood-flov. It seems much more reasonable lo sui>pose that the presence
of the abovs sulMtaucea in tlie blood excile^ the renal eiiitheliiiin cells to an
uiiwonied adiviiy. caadoff them to pour into Ihe interior of the tubules a
copious secreiiou. juHt as the pn«eiice uf piloear^nne in tlie blood will cause
the salivary cclU to |>our forih thuir Accretion inio the lumen of their ducts ;
and that this aclivitv of the epithelium i-elU in Hccmnpniiied, at*o as in the
case of the lubmaxillnry and other jflaiiilit, hv a vn-Hculnr dilation, which,
limugh adjuvant and hon^Bdal, i* nut thu dutinci i-atue of the activity.
This view i« further Hup)H>rtiHl by ibc; follontni; rvmarkable experimeilt,
wbioh got» far u> xhow that of Ihe variouo no luioiKfLi nhich, having fouix)
ihctr way into ibe blood, are thrown out by llir kidn*-y. ^ome paeut into the
urine IhnHigh ihc glomcinili, while olhen aiv ili.iiinctly Hccjvtod hy the tuhuli
uriiiilvri, the diichiirt^' nf the latk'r bMiiy; nccomjutriieii by a general activily
of the iH.i:r«*liu^ cclli. ax iihown by ihc flow of water taking place at the
Name time.
Ill Ihe nmi^ibia the kidney ha« a ilouble vascubir sumdv; it receivee
Brtvrial hliiml fi im tho renal nrtdry, but ihers !■ «||ii) poured into it venoui
hhn"! from another source. Tlw- fcmoml vv\u divldcn at the lop of the thigh
int'i Ino bniiH- lies, one of which runs along the front of tlto abdomen to
mee* )i» fellow in the middle line and form the anterior abdominni vein,
white iIm) other |>as»ee to ihe outer border of lUe kidney and brsinchc* in the
substance of that organ, torniiog the w-called renal portal sy*t«m. Non-
tbe glomeruli, in same spciciea At least of these animals, are supplied oxclii-
sirely by ihe bmncbes of the renal artery, the renal vena fuTUe only serving
to fonu the capillary plexus around the tiibuli urinifcri. which is also sup-
plied by the eflereol vessels of the glomeruli. From thii it is obvious that
if the reoat artery be tieil, Ihe blood is abut olf entirely from the glomeruli ;
and actual observation of the kidney has, in tl»e aniiuala in (jufiotiou, sliotvn
llml ilio volume of the kidni^v U remarkably senaitirc to ebeniical rlt:
takiiiK place in the Mooil. "the injmitioii into tUe hlood of evtu a iina
({uaiUity of nater causea a traiuii«iit sbriiiktiig «f t)i« kidiieir rulbwed br
tnotv latling ex]iaii»ii>n. Tlie iDJecii"!) »!' urea iiml »ume otluT (UurMi<
produoG* lilt) name cfTect tn n ni»ro innrked d«jcrce, ItailiDK wpcciaJly to
Kvrelling wbicb ImU for Home coiinidcrablv time, wbilc tlic iiij«ctioti of norni:
Millie •olutioD, iiii<l cMpCcially of micb (liur^^litrii ur WHliiim ar4!tate, Miuea
«xpan«ioii fmm the rcry ftnit, thf primiiry «hriiiLiiiig l)vtti(! abwiit. It
moreover, worlhv of nolo ihiit ihcsto i-irccW of ditirctin aii<l of clicinii
chaiigvs in tlic blood are obwrved cna nfU^r nil tlie reitiil nerven ha
appantotly been coraplvtvly Kvunvl. Hriicc thf cbungofl iti vulunie ema
by tii« proaeDCO of tne«c eubBlimccs ■□ the blood must be due to the m
sUtacee acdag either upon some periphi-ml vasomotor tnechnuii'iii, or, er^MV
more directly, on the bloodveaoole themselves. It may be added that the^
irill produce considerable effects in the kidney itself without a)ipreciablf
□ludilyinj;' the ^neral blood- premu re
S 416. If, while the kidner is in the oncometer, and the various expoi-
ments on seclion and stimulatiou »f nerves and the like are beinj; carried
oo, a canula l>e tied in the ureter, the ^cretiou of urine may l»e waccbe'l it
tlie aaine liuie. It will then be ceen tlmt the flow of urine through (lie eo4
of the cauula is not etpiahle, and doeit nut either iiR-reaM or de>:ren£e in u
even maniier. On the coiilrary, it will fnxuiently be found that a «vn of
gush of urine tiikes place, wreral drujw followiu); vnch other in rapwl uc>
«eaBion. followed by a c«itfuiiioii of tlow; and if ihi; un^tcr \n- waichol it
will be ?een that tlie )[Uiih<=t uf urine ar« michronouii with wavcu of
perislHltic cciutnii-tioii n«XTpi»]^ down the urtttitr. Ohviouxly the urine
collecui, to II cvrtniu extent, in the pelviii of the kidnvy, and ■• driven
thence by niuncular ncliou from time to time ; to this point we shall relnrn
later OD.
Haking every [i1]oi\imce, however, fur these irrcgiilaritice of flow, vn naj
take the rnle of How from the end of the canuin as a measure of the rate
of secretion ; and it in found thot as a general rule increased flow of urine
is coincident with swelliu),' of the kidney, that is, with a (■^"'c H'^*' '*f
bloitd through it. and diminished or arrested fliiw of urine U coincident
with shrinking «f the kidney, that is, with a diminished Dow of blood
throu)(;h it.
A Mrikiug inaiaueo of this is alTordeil by the exi>eritueut of dividiujr in
the dog the xpinal cord 1>elow the mohilln. The hlocl-preuiire thi^ii, a* wo
It now, Ml* rnpidlv, owing to the I'cmoval of constrictor impuUesi from the
small artcriot luiii the gcnernl iliminution of peripheral rMiitance which
Ibllow* upon to many >iunll arterice becoming dilalL-il ; and though the n-iial
arteries probably nharc in the general relaxation yet, owing to ihe fall of
pressure in the aortn conioinnl in> this n by a corrtaimnding rise of prcmirr
in the vena cava, Ihe tlow of blood through the kidney is largely dimtP-
isbed. We lind thiil sl^cr the opcnili'>n the wcrction of urine is grmtlr
diminiehed; indeed, in most cases, the flow from the end of a caaula it
almijet arretted. lu fact, we may almwt make the general aasertiou that,
irben in the dog the blood- pressure fulls to about SO mm. Hg. or leas, tbe
secretion of urine is for the time stopped. These and oilier r«<!ults Bun|Mri
tlie view Mated above that Ihe secretion of urine is in quite a special way
dependent on the How of blood through the kiduey ; and we may further
ooncludc that the secretion which la so pnrlicularly inllueuoed by the flow
of blood ix thai .-jiecial kind of secretion, allied to tiltralion, which takes
Clace through the glomeruli, and not the more ordiuiiry kiml of ^ecrolion
y nwnni of the cj>ilhelium of the ttibuli uriuilert. But before we prucvwd
TDK 9KCBET10N Of I'BIKK.
587
I
I
to (lisciMj bow the increiued flow nf Mmx] iiicreaaea tbe glomerular flow of
uriiip, w« tuuBt turn to ooosiiler ttie flinctauit* of tli« i']>Illi«liiitii of tlic
tubiiti.
S<*irfiioH A.v 'Ac /friin/ EpUhfiitim,
§ 417. Tko gUHneniliir iDechiiiiiiim i«, Hftor nil, ii KinttU pDrtioii oitly of tb«
w)iol« kitiiK'r. mill thu viittln'tiinti ovtrr it largo piirt o\ thr i^itirw of the
tubuii vrini/i'vi Infir* numt dlMiiiolly llic 4^hiiniclcr« of iin ucliro RccrtltiDg
opitholiimi. TIktm JacU wtiulil l«ui ii» 'V priori to nipjicwe tbiit ihe flow of
uriiif in ill pnrt ibo nsiult of tin iK-tivv scrrction ctfnipnriibte to that »f th«
K«li\-iir_v "r other inlands nbidi ve htivc already studied. And w<? bave «x*
p<^rinH-uuil iind other evideDCe thai ouch is tbe nue.
in th« first place, a flow of tirilie mar be artiliciallr excited even wlicn
tltc Diittiral Hon hiu been arrated by dimiiiution of blood-praseure. Thus
if. when the urine has ee«»ed lo flow id consequeuoe of a section of iImj
niodtilla oblor%«ta, cerlaiti sobitanoee, euob as urea, urates, sodium avetatv.
Uld llie like, be iiijevted into the hluod, a more or Itsa copious aeoretion is at
onoe set up. This secrelion is, or at least niair be. iinaonorapnuied hj any
rise of fteiieral blmtd-preK^tire sulficiem to aoci>utit tiir tlit! increased aeoretion
OS the mere reeult 'if an im-reased tlow of blood. It is Inie (as we havo
Men. ^ 416) that the iiij^L-tinti of ih«4>^ rulistanees leads to au exj)ati«oD of
Ihe kidney, ati i3X|>uiL'>i(>ii which Is pmhahly il(i<- ti) u locnl dilation of Ute
small renal arteriiii; hut thi- ttow of urim- which i« ohstervexl i» these cases
is too ureat In be iirroiint'il for by »nv tmrrciM- of How of lilond n'hidi the
local dilation may briri^' ahiiui ; luid hciKV w« ci>ii<^luik llinC the itinn-iuw of
secretion it of a diffcrnit kind tV»iii thiit whii-h follown u|x»i mere iooreaae
of blaod-fli>w. It tu'i'mn much nmn- n-iin>innbUt to suppoAC that tbe preaonoe
of th<- al>^iv« KuWiiiici^ in the litnod cxcitt^'* th«' roiiiil ttpithcliuui ceili' to iin
niinciiitnd iiciivily. ciiuoin^ ihem to pour into tb'! intcnor of the tubulwt u
cofHoui SMfttiioii, jiint m» tne preMnoc of pilociirpine in the blood will ntuw
llie salirary cclU to [mur forth their iwrotion into the lum«n of thoir dutft»;
and thill ihii iictivilr of the epithelium <m:IIs is ncoonipanied, al«'> ik in the
aiM.' of tbi.^ nnbmaxilhiry atid other glands, by n vascular dilution, which,
thotigh ailjurnnt and IwoHicia], la not Ihe distinct cause of the activity,
Thix view ii> further viipporled by the fitlowiug remarkable experiment,
whirh goes tar to show that of the various substances which, having found
ibfir way into tbe blood, are thrown out by the kidney, some pass into the
urine through the glonienili, while others are di^liootly secreted by tbe tubuli
urinileri, tbe dischar;^ of tbe latter being accompanied hy a general activity
of the secreting cells, a« shown by the flow uf water taking place at the
aame time.
In the amphibia tlie kidney line a double vascular sapulv; it leoeivee
arterial blood fr^m tbe renal artery, but there is also ])aurei] \nt6 it venous
blood from another source. The femoral vein divide at the top of Ihe tlii|tli
into two branches, one of which runs along the front of the abdomen to
nte^t ita fellow in tlie middle line and form thw anterior abdominal vein,
while the other paaiwi In tite outer border of tbe kidney and hrnni'litM in tbe
nulmtanre «f that organ, forming the uncalled renal portal nvHiem. Now
tl>« glomeruli, in Ni>ine xpeciisi at least of tlMM> animal*, are itupplicd cxelu-
Nivcly by tlie branches of the tunul artery, the renal vena porti« only serving
tJi form the caniltnry plexu* around the tubuli uriiiif^ri, which is also sup-
plied by the efferent vtwu-ls of tbe glomeruli. From this it is obvious that
if the renal an<!ry \w tied, the blood is *hnt off witircly from the glomeruli;
and uetuAl oWrvalion of tlie kidney lius, in tbe animals in <tuestion, shown
TllK CLIMINAl
thai umlcr lh<wi drcunMADCM Umhw U au it>|Iiis fruni tin' cnpiMnrjr ttetwn
cdrnniiiilitiu: iW tiilxilo bnck to th« glmni-nili; Itio^ kUv kidavy by ti
•iinpla owmtii'ii ix tniii>f<>rnKil ioto tut <iolii>urir M-cr^iing gland devniil •'(
imy sfMcial tillrriit); mrcliiiiiUiii. Such n kiiliiry niny )« iiattl la aan^iuia
tthiit MibrttUK'M nru oxcn'ted hy the gtonM-riiM. nml nhut In- the tubules io
some otiicr ^mn »f thHr wxirw. It is riniiii) ilmt iireii iDJccted inu tk
hlond giv« riM to n fountioii of uHiitt when the renal arleries aro tte<l : tbit
•ubdniice, therefore, is secreted by Uw epilheliuni uf tho lul>ule8, unit in
being so secreted gives riM nl the Mine tlDie to a flow of waler thrmi^fa th'
cells into tlie interior of the tubules. 8u|nr and peploiiea, on the oltiPt
baud, which injected inta the McxmI reatltlj {xtw tliruugli tliu unUMirW
kidney and apiiear in the urine, do a<A pass tlirough b kidney the mial
arteries of which have been tied, m-ea when a diuretic aucji aa unv >■ tfivta
nt the same time iu order to secure a lluvr of urine. Thcae Mibalaaina, U)tfn>
tbrv. are uxtreted by the glomemli.
The valiiliiy <i( tliia ex peri men t. whii'h may be lUTcepted u» in . a
lunrkcd difl'pr(!Oce Iwiweuu gl'imi-riiliir Ktrcretiun uu Uie one hand i .--
liul iir tulmlnr^rtTi-tioii »u I In; iiih(.-r,di.-|ii'ud.i on the absence of any mllalrnl
circiiltilioii wlnn-hy tin- glunn'riili may In- nuiiplled with blotxl iillier ligature
of the renai artery. In these animals anaMonKiMw occur between the rooal
arteries aiw) thi- nrterin of tlw generative orgaun; and uulctt llwiwial artery
be »o tied as to avoid thme t.'olliuerul coiumunicatioDs tlie rteulta of tJie ex-
periment are diflereiit,
Additional evidence i» fiivor of the Kcrvtor%' activity of tba ■■]>ilholiuR)
cells is aflorded by the folluvring obofrvatiuii. ^nto the voJaa of nniinals in
wlueh tbe urinary flow bad been urraeled by section «f the siunul cord l>elo»
tbc medulla a i|Uantity of (h<< blue oloriug material known ns sodiuta
i>iil{>)iimli|;otale' w injected. Tbii' snbrtmice is rapidly excreted on ibeooc
hand by tlii; liver in the bile, and on the other hand by the LidBey. By
Tarying tlw tjuautiiy injected, killing the auiraals at appnipnate LiiaeB aA*r
the inji-trtion of the material, and examining tbe kidiMya mioroecopicolly uil
othcmii^, it may be ascertained that the pigment so injected pasMS from tke
blood into tbe rcnni vpiihelium. and frum llieuoe into tbe ebanneh uf tbe
tuhulw. There beiti|^' im niniiiu uf lluid through the tubules, owing to the
arrest of urinary l|i>n by nti'uuit uf tht- pn^liminary oininilton, l)ii< pignK'Bl
travels vcrj' lilllc way dun n tbu inttiHor uf tbi: tnbuica, lunl rental iw veiy
much wWro il was ca»t nui by the epitbelium cellii. There urn no mots
whatever of ihd pigrmml having iwaed by the ([loaieruli: and the cells
which appear most diatindly tti take up and eject it, are thim.- lining sunk
portions of the tubules (vii., the linil anil •ccoud otmvoliitrd lubuliii, ligxag
tubules and aaoendiug limlm of tht^ l()o|« uf H«nlci im fmm their niictw
Mopic features have been «upp<jv<il tu he thcnctindy HTrrling |><>rt><tRBOf
the entire tubules. The folhiwing oWrvnliou which hux l)een miulo i» of m
peculiarly interesting character. Alter injecting a certain <|UHnttty of iiig-
inent. imd allowing Aucb n time tu cln[iBe as mi^ht be judged front previoill
es|iL'riiutiilii notihiHultitH' for the passage of the material through the epilbv
lium to Iw pnMty well iMmpleted, a second quantity was injected, it was
fbund that the cx<!ix'tii>n ol this seuHod c|uaDtity wad iui»( incomplete anil
iraperfci'i. It xeenied as if the cells were exhausted by their previous ellitTU,
just as a miiM'lc which has l>eei> severely lelsuiixed will not rsepond to a
renewed Mimulation,
The above obwrvation may be objected to on the grouuil that this oolorloc
matter doesnot occur ns a constituent of tbe blood aiher in In-alth or diMOM,
I lomrUuiM dillM IndlisiMunnlnf. thnusli tin* mm* b mgn PKqsrl) aiiplMit U a crwto; tmim
UlllNE.
rinlly ilml tlie HWnce of nuy oonoomiUui (liB(-)iaii;e of fliiM fVoni
Hi l^x^iI<1l ■u*|>i(.'H>u thai the procMa ob»er^'«ll was not rmllv mie of
Hitm ; fur the iojeclion of such nuIiMnnws u urea ur uratei iiiio th*: Woo.)
don raiiH a i-ri|>i<iua Buw uf tluii), am) imltnil lhtj« preveut» iht* niicroficipk
ttadciog imt nf tbi^lr pmange, which in th« cue of unua miKlu othi-rwiw Iw
AttM much in the Mtae vrmy a» with ilx; wodiuin sulphiDdiguuii-. Mi)T«ov«r
othvr nfatenren hmvt nuuiirBtiicI ihni rliv MHliuni oulphindliKoruic doeti like
nrrlfnary cnrmin* Mfs thmtifch lh4> gh)mcruli. Kiit ihvir niiull* iimy proh-
vhtv \-r rxpIaiiH'H by lh<- )iloiii<'nili having licrn dnnitijiicil hy n Ino rn[iid or
loo abuiiilaiit ii)J4-(Ht<Hi: iind in thf c«iw i^f th*- itnijihihiaii kidti«y, wlifn
Hum *td|ihiiiiligo(at<' «• injcctrfi after ligntiirc of reiinl iirlcriw, no iiriiio n
>im1 io the bladder, (ml iho pigtiicnl win he trtiocd thmiigh the <'piilii>tiuni
t^ werelinK portions of the tutMiUw. AVilhouC inoitting to<> much »n the
I of the aodiuiii Nilphindigotate experiments, ihcy mny bo takvn tut fairly
■pporting Ihe view which we are cunFiilenni;. We mny ndd that ii> binU,
Lbo uriDe of which coDliiiiis lilllc wnier. iirnlcs mny be delecte<l in llw' epilhe-
lium of the lubiilea but not in tho cnpeiilcs.
Tbou);!) much remaim to be cleared up, we may, for (he present, conclude
that the secretion of urine doc* consist of two neparale and distinct acts;
MTtvUon by the i^looteruli. which we may for brevity 's sake »peak of u
gfemerular («cn?<iuii. and seci-etion by the epilht<liiim of the ttibiiti, which
■t msy sfieak of siniilariya« tubular secretion. But these forms of sccrelion.
specially the former but Io a certain eiteiil the latter al»». differ from the
•Kreliion of »uch a iiland na the aolivary, oiid both deserve moie special cou-
•idemtion.
■ S 418. The nature i>f glomfnlar trerelion. We linvo seen that lh« expnn-
■Won <tf the kidney which hait for its accompaniment no increaeed flow of
tarinc bi one bmn^il about hy the renal nrUTV and il* varinUH hranclMB
txcinniog dilated, under mch circnmetoucn thni the dilfcrrncr between the
blnaMl- litem re in Ihe anila at the mouth of the renal artery and the Mood*
KiTMUiv at the vena cava al the mouth of the renal rein i» at the Mime time
*ncreiw«l, or at all mcni!" is not diminished. We say ihc renal arleri- mid
it^ varioti* bnancho' aince nnr present knowledge will not enable ii> l>i make
» more eiad siatemtmi. It i* of oourse pOMible that nervous impulse* [mu*-
int; al'ing pnrtJcaliir nervo fibres should confine iheir ellorls to relaxing the
mats of tne vna aHircniia of the ^doiiienili and not juuib to ihc other hmnehrf
tif the renal artery, in which viw the circulation of the iflomemli would W
exclusively lor ne«irly so) affected : but of this at pres«iil we kitow nolbing,
aw) the ^neral nf;;iimeHt remains good if we sitealt simply of the brancbca
of ibe renal artery as n whole.
In dealiiiK with the vaacular sy^ieiu we saw that relaxation of a small
I artery, lakin^ place without any marked changv in tite t;ei>entl blood-pres-
Hirc and in iieir^h boring arteries, leads t<> a fuller mid more rapid Mream uf
htdod ihruugh the capillari«!i supplied by Ihe iiriery, and that at the same
tine the prCMure in the capiUari>-« thcniM-lvi!* u> increoml; owing li> the
ilrtmue of peripheral roiitunee throiigb ihe widening of the artery, the
fmi fall of urmtnre (tte $ 111)) »t charncteri*tic of the [leripheral region U
(■hiflrd from IM arterial nideof thecupillarHii, toward the vcnoui' si<le oiu) to
Ihr cBpillnrice ihemoelvi-n.
Hence, M we have alrendy tuiid, when the renal arlery dilate* two thinga
ha|ip<'ii in the loop* of the glomeruli : a fuller, mon.' rapid slmiu of liloud
|aa»n ihrriugh tlwm. and that bliuxl as it Howi ihnmgh Ihem ic exentng a
l^reaier pn-wtiru than bi-((.>r« on iheir wallx. How duM each of the oventa
ataml Ifwani the secretion of nrinc?
Wv liuvc tMt at pr«4«nt Ihe meoiia of iitducing a Ailter and more ra[
640 THE El.IMIXATIOS OF WASTE PBODL'CTS.
flow without increasiii); the pre^^ure; but we ninv easilr obtain iacreaae uf
prei«ure nithnut the fuller ami more rapid How. If we hinder or obstruct
the outflow through the reoal vein we at once itierea^e the nreaaure Id the
gtomenilar l<H>|nj ns itt the other capillaries of the kidney. Xow, when tht
hlood-pr&<8Lire in the glomeruli is thus raised bv partial obstruction to the
venous outflow, the flow of urine so far from l)etng increased is dimiDished.
Obviously, then, the passage of water and material through the walls of the
glomerular lon)t)<, to go to form the urine, if not the result of mere pressure,
and cnnnot therefore be spoken of properly us a jiroeesa of filtratioD.
(t'f. S 303.) And we may here draw a cDnipariaou between the passage of
water and material through the wall of a capillary in an ordinary situation
to form lymph and the pat'i'age through the wall of the glomerular loop to
form urine or part of urine. The former, as we have seen (§ :}03), appean
to be directlv dependent on pressure, though influenced as we have also sees
in a very material way by the condition of the vascular wall ; and hindrance
to venous outflow, gii incfbcient in promoting a flow of urine, is as we have
seen es|>ecially favorable to the transudation of lymph. In the former cue
the sulMtances which paas through the capillary wall mav be described a* the
constituents of the blood t.'enerally, proteids as well aa saW and other soluble
and diH'osiblc lualters. Through the wall of the glomerular loop there pa«,
so long as that wall is sound and intact, neither albumin nor globulin dot
filirin tactor, but only water acrampanicd by some, and apjmrently a selection
of .some, of the soluble ilifiiisible constituents of llie blood ; for, as we have
said, the presence of proteiiU in normal urine u contested, and, at moel, there
is prciM^nl a very small quancilv only (which moreover may come from the
tubular epithelium). This difterence in the material which passes through
may be referreil to the diHerences in the nature of the partition. The trans-
udation of lymph takes place through the capillary wall ; between the blood
on one side and the lymph in the lymph-sjiace on the other is only the thin
Aim of conjoined epithelioid plates. Kul the corres;Hinding wall of the
glomerular loop is covered over and wrapped around so lo speak by an
adherent layer of ceils, which though reduced and thin are etill epithelial
ccIIh : the materials which go lo form urine have to pass through these cells
as well OB through the film of epithelioid plates. It seems to be this layer of
cells which determines what shall pass and what shall not.
Obviously the passage through this epithelium is of a peculiar nature.
The neces^nrv conilitiim for the due accomplishment nf the passage is as we
hnve seen a full und rapid stream of i arti'rial) bliKid : the high pressure
which accompanies lliHt full and rapid strciini, though probably under normal
eiri-u mstanci-M an adjuvant, is by itspU' hclpie*!. Thu^ when the pressure is
raised by venous olh»tniction, in which cai* the high pressure is accompanied
by a hI'iw stream or by actual arrest of the llow, even tlie passage of mere
water is retarded. Seeing that many of the Constituents of urine are dif-
fusible suhstanci^ ccrtainlv preexisting in the blood, inorganic salines for
instance, and »t*ing that, if we may trust ibe exjKiriments on the amphibian
kidney .*jM)kcn of aliovo, diH'iisible ahnorinal iMnstilucnta of blood, such as
|)epli)ne ajid .*ugar, pa.-s into the urine not by the tnhular ejiithelium but by
the glomeruli, wf might expect that didanion. in contrast to tiltration (.see
•: :ii;>), phiyeil an im|>ortunl part in the |>a3s:igi' ; and a full rapid stream
would und<iid)tedlv tiivnr diOii^iion. Birt diffusion l>y itscif will not explain
mntters. Efnr-nlbuniin dillcrs very slightly as regard-" dilfusibility from
KTuni-nlbuniin, au<l yet while at the most a minute ijuanlity only of the
laitci' posscri into the urine in normal circumstances, the former when injected
into the blood at once maken its wuv into the urine, presumably bv the
gbmieruli. On the <ahcr hand urea is an etuinently diffusible body, and yet
i)io (■x{>cHtueiiui on the iiiuiiliibinu kidney, the Dinin maMSt
■II rvTtil* lit thr iinii of titv uriue |iiiM«s by llie ejiitbi-liiiiu t>f (be UiIhiIm.
TlivitBporlaul |iiirl [ilnytif by tlii-epitbeliuni urhuKii ohrii th^e[iitlif]iiim
!• (lena^l. Jl' the n-mil artitry bi- temporarily li>:utiirt.il nr othfTwiw
obMnicit*), to ihnl tlkC glomenjli nn nhiit ntl' IV(>in tbcir btiHulstipjily for
•eOM liltir tini«, Xhti iwrciioa ol' titinc !» !>t(ii>]ir(l ; on nt^ablUhmrut of' th«
^fcalslion tKr frort-tioH nftirine vlonly rctnrnK. •in) Ibc urine i* thin (oanA
«0 bt albuminoti*. rrniniping m fnr wmc liltic time. Tho :>criim-tilbiin:iin
«Dd globulin nliicli c-ouM not mm through the intact rjiitbcliitin. ciid pnw
Vbrougfa wlH'ti ih« t-|iilhHhiin is (Uitia^cn b}* intcrfervncr' with its nutrition.
The iip|)fanow of «II>umiti in lh« uiia« (Klbuminuriii) i* a not infrm|iMrnt
■jnptom of kifioer diMiuc, and it« prcecDce in other than minute qusniitits
itHiimee iiuperf<y;lton« in the eloin«ruliir i^pilhclium. Uut even under
UBbmllhr condiiiona (hui epilhelium Ftill L'ovenis lo a cerlun extent llw
Cmape of material ; fur thcproieideof ihi>blo<Kl-pln£nia do not psse through
<dil\ ur in a proportion whicb correepoude cither to the retutivc projHiriion
hi «hieh iliey esiti in the plasma or to the relative eaw (or difflfuliy) vfiih
which ihey pais ihrfiuj{h membrsncs. Though the "albumin" of ulhu*
iBinou» urine Iroqucnily comiMaof both serum -album in nnd (;lol>uliu, tkrae
■k nut Dvce«Mrily (kvqt in the aaine proportion as in bluoit ; they vary la
■rina much more than they do in bluod; and indeeil the one or the other
mmj be aWnt ; moreover librin factors are very rarely found.
nwinoKlobinuria, or ihi- presence of hmDi^lohin in urine, loav be broUfrht
about liy injii-tiufc into the bloTKlvt-^i'dii laky bltHfl, or some •nlMtani'O Hic-h
•* pyrogallir acid, nhicb will "brrnk up"thr corpui>e]ea oftbo blood. Now,
in aucb cam there i* evidenec that the hnrmoglobin panes through the
^lom^ruli; minute difo-likv ratatt* of h»moglobin, lhL> fo-nlled "nieninci,"
mre, by appropriate mcthixls of pre|HtralioD, found in rilu in the cajwultv.
6urb ■ p«wag;e t* very far removed from being n proccn of diHiisioD.
We niny conclude. ibcD. ihni the |)(i<saee of malerial ihrouKh the );loni-
*ruli, like ihe tranmdation of lymph, ana even lo a more marked extent,
is • complex affair in nhicb the ordinary physical proceasea of dllfliaion and
filtr«iiou may plav their part, but are not masters of tlie situation.
S419. 7"Af wori 0} Ihe rjiithflitmt n/ the tubule*. As we have aaUi, the
alrunurvl features of the epithelium cells of the tubules seem l<> jUKlify the
eoDclu»iun that they exerote a secretory actJvitv comparable witli that of iv
Mlivnry ••r a gimtrii' gland. But their itork !m in maov nay* [wciiltar. lu
ibe c-n>L- of I hi- -iili^an'.gHMri?, and pancrcttlic glaudu llirrc can be no duiibl
that I hi- vfiecific Cunflilueni* of tbe tcvoral Kcrelions, mucin, jiegnin, lirprin,
and ihe like, arc inaDufaotun-d in the alveolar cells out of aiilt-C4-deuti< of
fome nature or other. The evident-, *n we have teen, it all agninct the
view that tbne glan<l)' mi.-iply wilhdntw, Mcrcle in the old sense of the word.
from tbe blood lbe«i- lulwtnnccs preexisting in the blood. When the Bulivary
^nds are extirpated, or the pancreas or the stomach removed, (here is no
iccumutatioD in tho blood of the fpecilie constituents uf the correeponding
secretions, tro also nhen the liver is extirpated there is no accamuialton in
(be blood of either bile acids or bile pigment. With regard to the kidney
■D<1 Ibe iDOet important eonrtiluent of urine, namely, urea, tbe eaee is differ-
enl. If tbe kidnersin a maiumal be extirpale<l, or if the kidneys by disease
or by liitalure of tte ureters be so damaged as lo be unable to carry on their
vork, an acrumulalinn takes place in blood, nut an wat once thought, of some
Mnlrei-iioiit of un'A, »ueb at creatin, but of urea itiwlf. In the caM.- of birds
■ nd r«[>tilr> which exircte not ureu, but chiefly uric and, the iK-eumiihilion
]• line of uric acid. Obviously in >ccrctlng uron the work of the epithelium
of ihr tubules is largely, if not exclutively, conliutd to >imply [Peking the
542 THK E1.1MISATI0N- OP WASTB PE0UCCT8.
ur«n out of l)ie blocx] aui) [uiBbini^ it, »i to »m«k. iuio Uw Iiimlnn of the
luhulfv. Wt< lut^hl. |>erluips. Bay exclutivdr, lur there U nu vviilcncv itiai
«tiy rirfn lU ull \» actually iiiHnurA(Ttiin.-d in ihr kiilooy.
Uut e\'rn thiit iiii-re tiickiti); uji ilii.- una u afWr all not n nintplo pixiccM;
tbe epithclitil cell »f the tubule U iiui n tiieri: jiiunivi- «i<-ri- of jioculinr Mruc-
ture, oepcciall/ tMUjttcid to siraiu dIT the urea from tlic l)l<x>d. As nr have
aln-ady nen, when iireu or uric aui<l Ih injecloil into the blood (tie nmilt u
not a mere iocreaac in the proporliuus uf iirvit (Vtr uric acid) present iu th«
tiriiw which is being nL-orctn). The injccliou Iviuti tu ua increaseil jlof 0/
MriW, tl>e whole oelivily uf thv cell ii* fllirrrd up. Kiid other ooaslitueiiU, pot
at th« momeul like tliv urcn cxisEiiig in exocM in lh« bluod, arc dtBckarfEtd
iuto the luinina of the CtibuUv toother with the urva.
How the titea, which u in thu peculiar manner token out of the blood,
conwi to make iu appearance in the blood is a problem !□ which tbe kidney
ia DOl conoerned aod with whicli we shall deal in treatiof; of tbe tueUib>ilic
events of tl)t- body generally'
!i420. In tlt« caoeofeome other coostjtuentsof tbe urineweluiTeevidwce
that tbo colU do something mure tlian iiinijtiy pi«k the couitilueut uut of the
blood. Iiii>i>uric acid, as we have ^««u. occurs Id small ijuEuititv in the unnc
of mao, and in Inr^-er amount in tbe unue ofbetbivoro. Now, liippuri<: add
may be foruKx) by the combiDatiou, with dvbvilraiion, of betuoiv Kcid and
alyciu (0,11,0, ■i-C,H.N0,— H,O = t;H,N0,>: and benwic acid inlro-
duoed into the alimentary caual ur iuieciAiI iuto th« bUiod rvap[>«art in largo
measure in the urine a* liippuric aciJ. HouHjufaere in tlte body tfa* beiuoK
acid [ueet« with and combine with kIvvid- And w« hara oicp«riinantal
{roof that the oombiiiaiiun may lutd j>robsbly doc* take |)lac« in tbe
Idney,
Ifn circulation of blood b« kept up through the bloodvcBwIs of the kidiwr
tVeshly removed from a living luiimal, and benEoic acid and Klycio be added
to the blood as it ia about to enter into tbe kidnev. bippuric acid will br
found in the blood isAuing from the kidney, eapeciaily if the Mune blood be
iwned through tli« kidney several timee ; the blood used must be bluod cud-
taiuingoxT-luuiDO{i;lubiu,c«rlM>iuc-oxide-hiG[aoKlobin not producing thu rHvcc
Tbe mere mixing witli the blood iteelf is insufficient; and if tbe blim«l bs
Mut not through a kidney just reruoved from the living boily, but tbrougb
one taken lironi a dca<I body or one which bos been left to itwlf for some
lime af^er removal from a living bmly. the sjDtbeaia will not bv cfi^ctvd. To
carrv out tbe i-ombinatiou by mettus uf the kidney which lia* been removed
friiui tbe body, the kidney mui>t retain for a while it« own lite, it must be a
"(tirvivinK " kidney. Nor is it absolutely uocuMory to bring the benioic
acid Olid glyein to tli<- kidney by meona of a blood-ttreiua. If n"»urvit-uiii"
kidurv be divided ra|>idly into small piecca and the bcnxoic acid rapidly
misc^f witli the pieces, hippuric itcid u forKfed. Nor t* it nccewary to fur-
nish the glyciu. if bcniuic acid hIoiic Itc \uvd, bippuric acid b formed all
th« Mme. ' <«lyctn, as we have previously said, cannot b« rwogniied as a
normal oitstitiient of any of the tissuea ; nerortbeUm, as we have aeeo in
speaking of glycocholio acid in the bile, and as wc shall kv« later on, glvcu>
must make a momentary ap[>eaTnDce in various nietaliolic proce^es or the
body. Wing immvaliately on il« appeaiBOoe converted into somethiiiii ehc, f>
that it iMver remains as ^lyoiiL It apparently is formed in (be kidney, and
is tbua momentarily available for tbe converaion of benzoic into bippuric
acid.
It seems prolNible, therefore, that, with regard to thb poriicuUr god-
Milueiit of urine, bippuric acid, tbe cells of the tubntea hare iJic potr«r
elleirtiug a oombiualion bulwofn the beoioic add brought to ttMin by *
THK SECBItTtOy OP tTBtSlt.
548
I
I
hUnnX utul ibe xlyciii wliidi they Airiiisli bv iii«am of (heir own inMKbotim,
bikI in iliiit WHy produce hipjiiirio acid.
Not (Nily beiiEttic ae'iA. tint timuy <i(iier bodice tnk«D into the lystem,
rcnp[)«Ar m ihe uriii« ronibioed with ;;lvciD. oml in their csbm also the
comliitiKCion |>n>bab1y ttkkts place tbnugf) the activity uf the cells of the
tubtilM of tbo kidney. MoriMver, other cban^ than the naauniptinn of
glyvin, ihft various clumgut which many chcmicst aubsunces taken into the
sy»l4rin HiidiTgo Wfont rwipjiwiring in the iirine, [ir'thahly also take place to
a lurcc •'xlMit in ihn kidney, aiid nn.: aUn oarned i-iui by i:i«aiia uf the
epithclliiin of thv tiibiiW
What other coit^tiliicnu of iiorinul urinv an' produced in tliia or a similar
loanncr wc do not iw yet. definitely knim,-. Tlic pigment tirobiltn. which, an
we have ewm. i# «ipw»M tn bo a derivative friim bilirubin, may he brought
ready farrac<l from liic liver or may hare Ihi- lini«hi»g ti>iich(M> ^iveii to it iu
ibe kidney itself: »nd the other normal or abii-trmal iiriuary pigincnig pos-
sibly ftriM either directly from hannoglobin or iiidirt;cllv from ibal bmiy
(hroueh the biliary pi^^ment by a Initi^formalioii taking; p\aix in the cells of
the tubules. There is also evidence in Tioga th&t acid aodiura pboephst* h
(Urnished by the cella of the tubule*.
In oonchiaioD, then, we may say llial the activity of the epithelium of the
kMi>ey appears especially nioditi«d, as comiksred with other secretiug glands,
to meet the special object which the kidney has to secure. The purpone of
the kidney » not to provide a fluid, urine, which oan be made uae of fur lh«
mttd* of ihe body, but to cast out wast« matters fVom the body. Heuoe its
M-crctory activity in limited largely to the mere dlschar^ »f mattera which
rcKch it pre<>xifttcnt iu the blood, ihnugU in several cases it give* the ItnnI
8ha[>c to the excreted substance b^ore it puMws idIo the ureter.
it 421. We may illiislnte the preceding discuiMon* by brieflv puMiing Ed
review »ome of the mot« usual ways in which the wcreiton ol urine » En
ordinary lif« modified.
In the precedinfc Motion the <vim|HiKiiion nf uHne wa» illuMnited by the
daily output of ibe WTenil oniHtituiMila. ruther than by a percentage account
of auy specimen of urine, for tbo rviwon lliat the composition of urine varies
within cxtmnely wide Umilm. This is c«[>w)ally the case as regards the pro-
[mrlion of water to F>ilid». Ouo urinr may be of biyh gpecilie gravity with
n small amount of water nilntively to the solid;, while another may have bo
Utile color and inch a low spi-eitic gravity as to appear hardly more than
water. The reason of these extreme diH^nom lies in the fact that the
kidney U uot only the channel by which waste solids leave the body, but
also an important outlet for the discharge of the stream of water which, iu
order that the various procowa of the body may be duly carried on. is cou-
tinuully passing through the system. It is frequently of adrantoge to the
body to discharge through the kidney a large amount of water, more or leai
imipeclive of llie solid matters which are, so to speak. wa.Hhed away with
it; nd henc« the advantage of the glomerular mechaui^n no s[iecially
adaptfil for the special dtsoharge of water.
As wp shall see prenfiitly, l» the iikin also falls tlie duty of disdiarging
large quaiililieH of water. The n«|)irat<iry organs also, as we have seen,
8pr\'e for the discliarve of water; but the amovnit which the latltir put out
csui only be varied by the inconvenient mHboil of incr«uhig or diminiiihing
the wholi! act of breathing. Hence we tind KpiTtal relations Ixrtween tlie
skin luid the kidney*, eurrvIaltDg the work of ibo one to tlial of the other M
regard* this gnuticular work of the iliscbargi' of witlor.
When the Ewdy is cxponwl to cold the ilinohargtt of water from the "kin in
ch^:)ic<l, and iho cutaurouit vessels are coustriuteti. At
J
544
TtIK BLlMlIfATlOS or VTASTK ritODUOTS.
the Mrao timr llic blmHlvpiwU •>!' i)ii- tiliil'iniiniil vUcrni. incliidibg the kU-
neys. ■»% <)ilii(tyl. but not out of piMjiortion to iW oitfttrntion of ib« nik-
oeoas vcMets, fur ilx' general lil'>od-pr«N§iir« <Ior« n»t fal}. )>ut if aDjlkin|
rlMfl BomewKnt. Tliu* ibcr? iit trslubiished juet the »tal« of tliiD;;^ w^iob m
&Tornble lo n full ntii) rajiKt stream of blood through the reiial fclomenli.
and an iucKiu^ How of iiriiw neiilts. It is poteible, w« nwy. jwrban, m
(Mobahle. Ibat tlie nen'oiift Bjeiem nlTords a special tie beint^n tbe bxia uii
the ki^lnej, to ibat under tbe circufitttaiicea in ciuealion iJie reDal artcriat tn
diUtei) even more iban thoe« of tbe otiter alxlottiiial Tisoera ; but ihla Im
not been proved ex peri mentally. It U also poeeible that br anotbcrnflec
mcolianisai of tJie central nervous svstam the skin may work tipon iba kid-
ney, pot by tbe vsKimiitor nerves aloue, but also by nerves piveminiL' tW
fMretoiy activity of tlie lubulM : but we have no MUiafaoiory iixticnitoiij nf
any such mecbaninmi, and it m-t-mn iiiL<r>- jirnbablu that theconnitction miouU
ba with tbe gldmt-nilar niei'liHui^ni. ^lll^■- ihii chief object at all evi-iilc i» M
got rid of KUtcr
Convi-n«:ly, whtn ihe body In exposed to warmifa the ikin perapirvs (nth
and llw aitam-ous vtwel* are widely dilnle<t: and conversi-ly al*! th« renal
aitd other ulMloniioiil v<-n»eU an^ vunxirictvd. m that a slow ami iininll slnaa
of blood trickles tlinAigh iIk- ginnioruli, and tlw urine whicJi Ja secreted k
scanty.
^ tiS. Even m"Tv imporlnnt (hau iu relations to the iikin are tho relatiooi
of the kidney to Ihc water abwrlictl bv tbe alimtsitarr cainil ; this is eajwci-
ally seen wben large iuanlili«« of fluid are dnink. The whole of the water
thus introduced into the alimciitarv- canal pOMce iubt tbe klo<Ml, for in a
healthy orji^nism no aiDouHt of Huid drunk, unlcw it throws the ectMomj
out of order, can eflect the amount of water present in the feces. But tht
addition to the blood of ev«n a very larf^ quantity of fluid do€s not, as m
have seen, by ita mere quantity (§ 1S(J>, increase the K^neral bloo<i-pn-Murf.
and therefore cannot in thin way produce nbat it umiimbtedly doett pnubiiv,
an iocnased flow of urine.
The fluid so absorbed may act on Ibe kidney iu two ways. Oti tbt <um
hand, as we have seen (§ 41o), tbe injection of water into the blood prtiduM*
a local dilHtioii of tl»e renal ven«l<i, as indicated by iJie sucllin^ of Ux
kidiMty. Thu» the al»orptJon of mere water fmni tbe alimentary caual may
stir up to (greater activity the glonit^rulur iticchaiiiiiiu, and tii so doina; nay
b« aswtcd by the i>r«seaice of various subataaeoH alHorlied from the nlimen'
tary canal with the water, for some of these nlm may similarly lead to
dilation of the renal viiwel*.
On ihc other hand, M>me or other of tlie chemical Iwdin ibuapasrinft oM
the blood with ihv water drunk may excite the eccntory activity of tU
tubuIcE, aiMl that ciiIkt by acting din-ctly on the epilbelium as ihev are car-
ried through the kidney in the blood of the mtal artcrin, or indirectly
tbrxiugh some inlervenlioti of tbe ceiilml nervous sysleni.
Our knowledge is at present too scanty to enable us to dactde which of
these two meiliud« is the one usually employed by the organism ; but tlie
inordinate flow of urine, so poor iu soli<ls as to be little more than wat«r.
■rhich may be directed through the kidney by means of an adequate "drink
log bout," would lead us to conclude thai in such eawe ihe organism, ulriv*
ins, ibougli too often in vain, lo fV«e itMlflhini the evils to which it w Mag
cubjecte<l, has reeourse rather to tbe simpler flonenilar tneduinism than to
Iha more exjieohive [iBMie-nastinf; activity of tbe tubules: and the tirine in
nich cases pri>biiblv dioihurged chiefly br the method of dilating tbe renal
vewelx and thus throning the poisoned blood inio the glonirtuli.
When, houi^virr, llui<l i» taken simply as a projier acoom|iauim«nt of mjIM
THE SKC'RKTION OF tfBIXE.
MO
toad, the incnaae of una« wliich ksuIis trnn pmlirilily iiiiDthvr origiti. As
we have already »aid, nnd tti wi; »hiill jH>iiii out iimix! fiiliy liili^r on, ihe
abM>rptioa of proteiit luutxTml, which in ii cxiii.->ti|iiciit umI generally ii oon*
»|)icuou« i-onstiiiibiit »f tivvrv mviil, IviuU u< n lormHtioti of iinni ; iind urm.
oa we have iweii n>»»oii ti> tfliovo. diix-clly "limulalw tho <-t)ithi:liuni of the
tubulcn to HCCrtitor/ notivUy, Apd whnl «ociiw prominently tnic of urm U
probubty tniv of niitDV othiT procltirU of iligcslion ; to Ihitt thv ii)crciu!nl
How of uriiM! which foriow* an •>n)iiinry inn) iKvompflnicd by not morv thiin
tb^ ordinary amount of Htiid, is tho result of the liibora of the epithi'litim of
th4> tiibiiln' a« well iis of tbe fuller stream of blood through the glomcruti.
Ii 423 Whiit hits just beeo Mud concerning the inRuenee on the kidney of
food itnd wilier niny t>c np[)lie<I nleo to the action of subatance*: which being
•Specially ctlicjicious in promoting' n tlow of urioe when taken into the body
■re c»lle<I "diuretics." The several actions of various djorelice are veiy
Taried, and it would be out of place to discuss them ftdly. We mav, bow-
ev6T, MV thitt wfai)e the action of some appears simple that of others is
complex.
Such aicenls M sodium acetate nnd pola^um nitrate proluibly produce
iheir efl'eec ohieflr by actinf; directly uti the kidney, indiieiiifc. as we hare
seen, ^ ilfi, locml vascular dilation and ho workiii); ou the glomeruli, but
|>ri)bai)ly at the same time also xtirring U]>, ai\er the fanhiuu of uren, the
epithelium of tJie luhulee to weeretorv activity, the aecomiia Dying fuller
Mreuni of blood tbrotigh Ihe tvholn {lidncy oiAog, iw iu tlH: cbmi of Uic
mlivary and other ijlaiiiiH, a UM-fiil ndjtiviint.
The diuretic eflW't <if such an ugi'iit im digitalis U probably more complex.
Sy incrMuing the cardiac Mroku, and at the Mime time oouHlrictitig many
SnuiU vcmmIs, digitalin rsi«c^ the g<>ni-rn1 i)lo<iil-iimMUre: but tho tendency of
tbe increased blood -proxiiri' to incrcii>i.- tin- flow of urine: may be counter-
bnJanoeH by thr conMrii-iion of the rcinil vi?«els iheniulvcti. And while it
U n matter of common i'X{K'rieticr that iligitali« u Tory dfectivo na a dinretto
in cardiac dwerotc, thin- is great douKl whi-llicr il rrafly artu as a diiirMic in
liealih : in <'ar<linc ili^-iwe it prolwibly miw* iIm; b!i»od-pr*n«ure by improving
the cardiac Mroko and n<>l by cmiMriction of the blootlvvMcK Ditt even in
the abeence of cardiac dineiin.-. dii;it»]i« hiu* been found in ivrtnin uiRce to act
<as a jkowerfol •liiircli<-, nnd in th»e caMe« either il miiKC act directly on the
tubular epithelium or il* iHcets in eonglriclini;; ihe renal arteries must be (eSB
than its effecia on other small arteries or mma iituo olf before tbe inHuence
of tiie heighleneid blond -prenti re has dimpiteared.
§ 424. Quite removed from the intervention of chemicnl substances in tbe
tihvxl and yet m<et striking in the iutlueneo on the kidney of the ceniral
uervoiis system. Tbe potent influence of emotions in promoting the socre-
tion of unoe is proverbial, and the general features of " nervous " urine, the
water increased out of proportion to the solid constituents, eapeeially seen iu
tbe "iiriua byaterica," which is hardly more tban umple water, often dis-
ebarget) ill c]K>nnous ipinntity, at once suf^eais Ihe vievr that iropulMS oHgi-
tinting In tlie btain and paaaing down to llie kidney along the vaao-dilator
ihrtt, of whuM existence evidence was given in ji 414, lead to dilated blood-
VMMband gie*t play of glomerular activity, witbuut perhaps produeiugajiy
otbtr dirf«t effect on the economy : (bough pudsiblv the same emotions by
oonairieting tbe cutaneous and, it may be, other veteels may raise the genenti
bhxxl'prrMiure and *•• help the dilated renal veMels. In the coMof tbe uriiM
of bytu-ria we are li-uipii-il, more perhaps tlian in any other iiulancQ, to
accept tbe hint prcvi<iu«ly thnmu out that il b giowible fur the rata afibr-
eutia of the glomtnin to' be alone dilated, so that tbe gr««ter part of tlio
S6
546 THE ELIMIXATIOX OF WASTE PRODCCTS.
renal bl<><id is directed to the glomeruli and the epithelium of the tubula
left in its iistmt quiet. Itut thi^ ii as vet pure B|>eculation.
Thk Discharge of Urine.
•t 42fi. .'ilrurlure of l/ir iirrter. The ureter, like the large ducts of other
glands, CDiiifist of an epithelium resting ud a connect ive-tisaue basl^ Btrength-
eiii'd with plain muiM-ular fibres. The epithelium is iu its characters iDter-
mediate between that lining the ces»phugus, which as we have seen (§ 221)
rcsemlilcs the epidermis of the skin and that lining the ducts of the glandi
of the ulinienturv caiinl. It couxiBts not of a single layer but of three or
four layers of celU. The luwernuKdt cells, next to the baeement membnue
which limits the cod ne<:tive-t issue basis, are oval cells placed vertical I v, in
one or two layers. The cells of the next layer are irregular in form aod
ofieu |iear-i«hapod, with a narrowing process dipping down between the celU
below. Above these, forming the i^nrface of the epithelium, is a layer of flat
or of flattened cubical celU. All the cells are nucleated, and there are do
special features in their cell-stiljstnnce.
The connective ti^ue is, av in a mueons niemhrane, delicate iramediatelv
IkIiiw the enithelium, but l>cconics coarser and more fihniue in ii£ out«r parti
The muscular fibrfs are arranged in three liiyera, an inner loagitudinal, i
thicker middle circular, and a thinner less n-gular outer longitudinal layer
lietter develo])e<l in the lower juirt of the tube than elsewhere.
Nerves jiatw into the ureter at the ui)]M.-r end from the renal plexus and at
the lower end from the s{>ermatic and hyjiogai^tric plexuses, and at the two
ends nerve-cells are scattered anion>! the nerve- lib res.
The jielvis of the kidney is an expansion of the upper end of the ureter,
anil is lineil by an epiihclium like that of the ureter, which is continued ioio
the calyces and over the projecting |>a|)tlUe of the pyramids. The circular
muscular Hhrcs of the ureter are continued over the jielvis but form here a
relatively thinner layer, while both longitudinal layers are very scantr and
graduulty become lost.
At its hiwer end <'uch ureter ojiens hy an oblique opening, serving aa a
valve, into the cavity of the bladder.
S 488. aii-Hchirc of (he blaililer. The e]>ithel]um of the bladder reaemblea
in its characters that of the ureter, but the ap|H-iirauce9 presented by the celh
in sections of prcpareil bladders will imturully vary a good deal aeeordiD);
as the bladder was hardened in a coutruuteil or in a distended state. Thii
ejiithclium with the underlying fine (.-onncctivc tissue forms a mucous mem-
brane, separated by submucous itinnective tissue from a well -developed mus-
cular coat, which in turn is investe^l with an outer coat of connective tiasue
cuverul over the greater |«rt of the organ with |H;ritoneum.
The wcll'<leveloped, i>laiu muscular libre-ccltH which constitute this mua-
rnlar coal are gatlicrwl into rounded bundles or flattened bands, which in
turn are arranged in a plcxiform manner, being btmnd together by con-
nective tissue carrying bloodvessels an<t nerves. The direction of these
buiidh« is not very regular, hut they may lie regardcil as forming on the
inner side below the niucouK membrane ii circularlr disposed coat, better
drreloped at the lower part of the bladder amuml the oj>ening of the urethra
tku elsewhere, and outside this a longiludimilly disposed luat. the loogi-
ndinal direction of the bniiilles lioiiig belter seen at the front and back than
M tW aides. Many of the bundk's and nelwnrksuflninclles. however, in both
<m» nin > c<>urst> which is neither exactly lun^ritudinal nor circular. The
■Mr Vwgitudinal coat of the ureter apjieurs to be rejiresented bv a very
617
thiu und inmuHiKtiotis \t»ytr. Thi- thicker nnil better developed portion of
chf drculitrlr tlis[KiM<I cuat is eotDotiiiice epokeu or m ih« mhiiuiter rftictt,
anil llie longiliulinnlly <li8prei.il cmt t» similarly aoinetira«s calfe'l the defriuor
urinir : Init, ae ire Bhall pee, |Iim« names are uudeairable. In the ttog th(i
lonptudinal butiillcs are mucli belter developed tlinn the circular; but tlic
relative projxmion of the two eets of bunttlea eeeius to varjr in different
aniniabi.
The bladder b sujiplied with iwrvea (torn the liTposasiric ])lexti3, tli« fibrw
beiuff both mediillaied and non-meiluUated. They ap{>Mir, tu, in the cue of
Ibe rectum (§ 277 ), ii> have a double ori^n. dimiiiL' (lO (lie oite linnd from
llie lower <I')r«aIaud upper lumbar spinal curd ibri>uKli (lieiynipaibeti<'«y«tcti],
and on ihe other hand, in a more direct manner, from tlivwifrsl spinal nervee.
More abundaut nmniid th« neck uf ilie bladder than biglu-r up. they run at
iir«t in th« outt^r oonnective-tiioiic iiut beneath the pt-ritoucuni and, funning;
pIcxiiMf. iillitnalcly end piirtly In the hloodvocicU and partly in the muii-
cuUr tibrcs. tli'iiigh iH>m« fibrus are wld to have been innvd to (be epithe-
lium, f iri>u|M "f nerve-cell* occur on the pkxneM, espi-ciallv ni-ar the neck.
■j 427, Tbc oriuf. likr ihc hile. i« Kicrcied <>ontinumMly ; tfjc flow may rbt
aix) fall, hul in health neror atwotutely cciuc for any length of time. Th«
<-i-*Hilii>n of renal activity, the km^IIm Mipprewion of unoe, entalb apeeily
death. The minuie »treHiiii> pouung continuously — now aion> rapidly, now
more slowly — at'ing the collect inc. ind discharging tubule*, arc i^thorvd into
the repal pelvi«. irTicm-e the tluid is cxrriitt aloug the ureters into (ht; blad>
<h»r partly l>r prcMtire and gravity, and from time to itmv partly, as »« have
alrendy said (§ 416), by th« peristaltic cantractiona of the muioulnr walls
of the tireter.
If in a liWDR animal a ureter be laid bare and nimutated, mechanically
orulherwiM, at a part of its course, ware^ of periotaltic contraction may be
aeen U> pass in both direcltitis from the snot nimulated upward toward the
kidney and downward the bladder. In toe abaeoee of artificial stimulation
i<{ir»ntaDeou« waves of contraction make theirappearauce.aometimes repeated
with tolerable re^laritr (abnut every twenty aecModa in the rabbit), aome-
tln>M occurring in jrroujid with longer pauM« between. Those MKHilaneoui
contractions invariably i>aM in one direi^iiou, from the kidney to the bladder,
and their rmpieuiTy and vij^or seem to be determined by the activity of Uie
Mcrt'tion of urine. But they ar«r not directly called forth by ibe urine,
either mwhaiiically diMendin); the till"- 'ir chemically stimulatms the inner
surface, for regularly reoirriuK <v>n tract ions may be obwrved in a kidney
iind unrtcr removc<l from the body, or even in an isolated excised piece of
tthe ureter.
The rhythmically repealed contractions arise spontaneously in the miia>
oular coat of the ureter much in the nmu way as tbc similar cartliac coo-
tradiotia arise in the muacular mibstancc of the lifwrt ; and it may hen be
raentioned in support of what was iirg«d in ^ i't-'i ivith re^nl to the hc«rt-
beals not beinf; M4irled by nerve-cells, that rhylhraically re|)eated sjioHliincous
neristallic cnntmetions have been ohwrved in isolated pieces of ureter taken
mm the middle of its course, in which uo nerve-cells and indeeil no dij>tinct
nerve-fibres could be obaen'ed.
In the living- body Ihtse spontaneous movemeols, beats they might Im
enlird, are subordinated to the flun- of urine into the pelvis; the more active
the iMtcretton of urine, the more fre<|uenl and vi>;orous are the bcwis of the
pelvis and ureter; but the exact mechanism by which tlte Mcretiou and the
jDorsnunts are autintaincil in liarmuny lina not yet been cleareil up.
I
648
THK ELIMINATION
PRODUCTS.
MirtHrilion.
$428. In the urinary bUd<ler ibe uriii« a coIIucim), iu raturn iob) ik
tir«ivra beio)^ ]>revetileH by Um oblique entrsriM iiilo llif? Iitiukler niii) vain-
lar iiaiure of ilit oriticea of Utoee tilbe», and il* di-chitrge (rvnt llieocr io
CMKftderable uuantily ia ertcoiwl fr.mi time to tinio bv a ^lucH-bai cumpifi
muMular inecbaniain, of the uiUure and workiof; of whit-h i1m> followtuij isi
brief aucouut :
The iuvoluDlnrr luiiacular tibrea forming iJk ^realer part of the vatctl
mill* are arrnii|j:vd, a* we Itare nid, {MUtly iu a ii>»re or loa toriuiiuitiiul
direction, and |Minly in a circular manner. Afler it Itaa been eiitpti<>*l th)
b)ad<lvr ix oxilnu-UKl aod ihruwn into foliU ; lu tbe tiriiie fcradunlly oul1i>i.i>,
llio blBd<l«r b«wuiDea mure aat\ uihk t)iUeu<ted. Tli« e«i-a)i« of tb*- (luid i> ia
]iart iirevcDU^d by tbe reaiatance olKired by tbe eJaiiii; ^brua in tb<> wulla of ibc
urvlhra, nhidi lielp to keep the urethral ehaiidel clo^^d. But tbia ia iiui all,
for ohwtrvitlion »bi>«« that fluid 'n retained vritbin thi> Itlatblrr up tr> A pn»-
ture of twuiily incliva of mater, to long an llie bladder ia )|[»v(-riioil by aa
inlnct Dpiuiil c»rd, but give* way to a nreMure of aix inehc* only whvn Un
lumbar spinal coni i« <b5'trovetl or the viyucal m-rv)* are *t.-vurcd. TU
aRorili> vm~ ttrong rvidciuf ifiiil tho oWtnioti-ni al tbe ncok of Uw hla*iil(*
to the «xit uf urino dopendx on fonic Kiiiit- niudcular i-ouiractioD mnjatsitwd
by a reflex or nuloiDnttv action of thv lumbar K|)inal (M>rd. And it ba^baoi
maintained that it i» the circularly dispuacd fibrot apocially d4'v^I(>|K■'J nr<itiad
the neck of the bladder, which an; the aubjects of this tunic con trae lion aal
thus the chief cause of the intention ; h<'ncr the name «^bincter vesioK. IV
voniinuity of tlieae fibres, however, with the reet of the circular fibres t>f tht
bladder sugiteeis that ihcy probably do not net n< ii sphitu-ter. but ihat tbur
une lies in their coDtraciine after tlie rest of the veaicul fibres and tku
fiiiifhiiiK the evacuation of the bimldcr. Tbe resistance in i{ue8iioii b sun-
I died by a tonic contraction not of th«ee circular fibre* of the bladder itM«,
Hit of the muscular fibres— partly plain. {>artly striated— surrounding (br
prostatic piirtiou of the urethra, and cousliKlliDg ihe ^fhinrtenvjiint Fxl'-mut
or pi-iul<ttinii', or ajihiucter of Henle. It ia stated that artiticiallv exetted
<.'»n trail ion » of ihi-ne fibres will rei^ist a pressure of fluid in the l>la<l(W.
When the hind'ler baa become lull, we feel tli« need of making water. Ok
eaoaation being hcitilitem-d if not eaused by the tricklinii of a lew drop* of
uriiM) from the fiiH blsddi-r iuiu the urethra. We are then consctuits uf an
eflbrt: during ihiv dfort the liladder ia thrown into a lonirLonlinuisI con-
triiclion of HU olim-un-'ly {icriidaltie nature, tlie force of whicii in more ibna
8ulfid<.iii 1^1 oventinir ilio n-ii.->lunce oflbred br the urethra, wwl fht' atim
imues in a stn'ani. lli<^ •phincli'r vcMneexwmuaueiugul iheHunc tiniii I'ltbif
relaxes! iiflcr ih^^ tiuhiou of the M>hincter ani, or ailemt overconx'. lu iu
paMiige along Ibc urethra, the exit of tbe urine, at nil evenlf of thr last pM-
tiom, is lorwardwl by irregidarlv rhvtliniic omlractioni' of tlw bullju-cavcr
DOHis or ejni'ulnlur uriuic niii>clc. ifie conlroetioiu »f which ciiiipn-es ihu
urethra ; and the wh'dc net i* further muiKled by precnre on t}ti- bladdsr
exerted by means uf tbe nbdotniiial muscle*, very much the snine a* is
ddiMMitiain.
In the esse of the rwtum we were able (^ 277) to distinKuish between tk(
MOtioas of the longitudinal and of the cin'ular cimU. luxl we Mid that tbt
two coats liad di*linet nervous supplies (Fig. l'Jt>). The bladder has. as wt
have said, a Mmilar nerve supply, and it is very prubnble, hut iii>t yet dis-
lluctly proven, thni this, like double supply, bas a Itke <l<>ub]c action. Stimtt-
Inii')U iif the branchr* coming from tlw sneral nerves, at lh<^ same tinifl tint
" Uiium thr Km^rilniliiml tiiai i.f ilic roc-lurii iniii rontnictiMU, brings aUuil
'B ci>* iUifi. iu whirl] the loRKiiiKliiiiil lilin'« of iho hiNdtUir mre niii<.-li mora
flMfMiiK-nl thu tbo cin-iiUr, powcrrnl vc«iral miilriutiunH. )Ior<<over,
tlataliUHdi <>r lh» Mcrnl n«ivv« on otw rvU- producvs uiiil«t«ntl mjotrai-tioa
oC thr ttUilik-r. Fn>i» thi» we mar inf«r thnl (he mcrnl nerves govern the
*OQ|titi)<lituil c)«l. KlimulBtion ol th<- hr|>nsguttric norvcs cnrryLoe Bbrae
A'i'ni tlir ilonail luii) upper lumbar oml (mv Pi);. ]'H)), irhili> thfotruiK lh«
Ctrculmr cjhI of the rct'tiim into elrong coninietion^, gives rieo to mieal con-
tliMtticHn, but the«>e lira )ir no meuiis fo nmrked at tlioee wlticli sp|icar wlieii
Mm axmil ucrvm ara Hliiuiiinied- We may probably Ly>nelu<le that ibe Piore
■niportaiit tibrea in the fnntlus of the bladder, whicb ara for ibe nioM put
loDptuiiinal, ara to be rejuanled aa goTeraed for the moat pari by ihe Mcral
■MTve-iibn^. nhile the circular rouMiular fibfes around Ui« neck of tbe blad-
der, wfaoAC cnntracli^n cufupletat. lu it were, the emptying of tbe bladder,
Ar« tIttMe ou which (he byputraeiirii- nerve-fibree have tbe chief influeiMje.
^ 429. We Mid juM now " when the blndder iiaa becwne fuli," hut tbb
iniii4 Dot l<r underehiud to mean " when the blailder liaa received a certain
niutntiiy «<' Ouiil." Ou the i-imtrurr. it is n matter of coraiium ex [wrivnctf
tluU w« Utl the di3iire to luukc wuier iuiuieLime* when a lur){e i]uautity and
■oimwiiw when a iimall iiuantitv of urine has accumulated in the bln4tdcr.
AVe have cvidenec that Ine hlailder poaacaaai lo a very high decree Uiat ob-
•curv eoatinuoua coDtraottMii which we tpeak of as " tone " atui, funber, that
th» amuuiil of its time is cxcreitinifly variable, the organ, quite inde])end'
«otlT of dintinct rfliirtit at micturiiioii, bciiifr at otic tim« contracted anil at
mnoltier flaccii) an<l iliMCmled. When it ii> in a coiitractMl Kale, a smaJl
ILianlity of Suitl mnr exert the muic clfi^ on (he vtMcnl walls as a larger
<|uaatiiy when the bladder io fluedd. Hi-um.-, while ihv drtvrmiiting cauw
of tbe deoire to make water is tin; prcsture of the urine upon the vesical
walls, the quauliiy needed lo produce the necessary fulneae K dependent ou
the amuunt of tonic contraction of the muscular libr4M enstiof at the lime.
Awt we have evidence that this lone is regulated by the nervous aysteiu.
(430. Micturition as sketcheil above wema at firat aght, and capecially
■tuna we appeal to our own con^^ciouHDese, a puivly voluntary act. A volun-
tary rfliirt thiuws tbe muscular fibrm of the bladder into conlraolioiu, an
hruoipanvini; vuluninrr efli>rl leeaena llie bme uf the sphincter extcniut,
)ut>lMlilT i>y inhibiting >t'^ oenlrv in the spinal curd, while other voluntary
fdna tlirow the ■■iacnliilor and abdominal luuiicles into oonlractioiu, and
I&e ra^istanne of the urethra iM-iug thereby overcome, iheexit of tlw urine
(Utlurally foHowa.
Tbrrv are facta, buwcver, which prevent the acce]>tanoe of Mtimplu ariev.
In tKe 6ni plnce. in caan of urethral ohslniction, where tlie bladder cannot
lie •niitiM-<) when it rcechtv it« acciMl'imed ftilonw. ilw incrvasing (listenUoo
ijaU up fniitlMs ImU powerfnl contntcti<m]i of tbe ve«ica] wall*, conttactioas
vrbtch are dearly involuntary- in nalura. which wane or dixappcar, and return
AK^in and again in n rhythmic innnDcr. and which may Iw m strung and
|H>w-erful u ID cause graai •ullVriDg. It spi-iii« that tbe lihres ot the bladder,
like all other museulnr libres. have tltnr •■•mlrnctionii augmenitvi in pnipor-
tion M tiiey are 'ubjecled [n lcii»ii>n. JuM as a previously iiuie^eenl ventricle
«r a frofc's lieurt may be excilnl to a rhythmic beat by dixicndint: its cavity
with blmid, *> ihe •juiesceni bladder may, i|uite indepcndeni of the will, l>e
*scilr<l, liv tlie disii-iiniun tf its cavity, to a peristaltic action which in normal
(■act i* never carried beyond a first cITorl. liiioe with that the bladder is
MBUied and tlw rtimulus is renmvcd. btil which in casea uf ohatniction is
(Babied clnrly lo raanifesu iti rhythmic naiure.
In tbe Meoad place it bus iJeeu tliowu that <|uite normal micturition may
MO THK BLIMINATION' Ot WA8TB PROPtJCTS.
lake tilitce iu « 4lng iu whk-li tlw htnilmr r«^on of (He apiiial curd bw bwo
coiiiplpi^lv kikI |<«riiiancnilir weiwnim) by Mvlioa Aroix tho Mi>|M>r dunl
re^rioo. I" xiicU a oiiw iben- can t)« an txvmm nt volition, miil th« wlxJe
Erocew appeani am b n-flci: iictiiii), Wkni tmiWr thcM cinmniKtancaib
liulder D«e<in>i.-» TiiM (an<l othurwiMi ii|i)iur<.-iitly tl>v ad liiilf) unv mUrM
vlimulii*. viicli an opnnging lli« utiitt or >li^ht |>nwurt> on iho ubtlniiilml
unllo, mi>*i' a romptrto act of micuiritMii : tlve liladdvr i» •.-ntircly emptJnl,
HD(1 iW »trrnni of urine tottnnl ibe vaA of llw net und«rgo«« rhyttimnl
augmcntntioiu due to oontnctiotifl of the ejaculator tinoie. These fwu nn
ODly b« interpnied on tbe vi«w that there exi«ls in the lower spinal cnni (uf
the ilog) what ne may speak of u a micturition nntte capable nf beinn
tbrovn into action by appnipriate aBerent impulses, tlie action of ib« cmlrt
beiDic sucb flfl tu cause a coniraoliou of the iralls of ibe bladder and of the
ejaculator urtnw, ami at the raiuu liro« to auspend the tone of iho apblocMr
Teaicie extemtu. CHuical vxperience aUo ^o» to show the exutoaoe of ■
■itoilar micturition centre in nian, plBcc<l liigher up id the cord (ban lb
corretpoudinK " Ktuiital " centre goveniinK the jtetiital oiYao*.
Moreover, w<- lini'e, in the ciwe both nt man and of ocher anininlii, eipwi-
menial nnd o[h«r <;vid«ncn that emit ract tun i)f the hlailder t.t fn<i{ueMlT
brought itliciiii liy rcllcx action. Thii> \\\« prtrwurv iriibin ihit bladder wbni
oWrvcd for iinv length of tinir in found to Xtv irnhjed to ninxidcraldv anl
nianifold vfiriiitinns. Over and nlmvc |mk<>vi.' i-haiigt« in prwnuiv <Jiie tii ll«
mpirntor)- movi'mcnttt. ihnmgh whii-li iW lilaild«r i* prr"H*il ii[»i« at oadi
dNCrnt of the iliiiplimgtn, nr<ivt< i-<>iilrartit>it«. nf a drrngth iiiadcqunle Iu
bring about micturition, nre from lini*' to tini« oWrrct). Tht^n in eoiM
imtBDCw a]>pear to be sgwiitanoitii, or to be ihi- r««ult of emotions, but tber
may be readily induced in a rcRex manner, by stimiiliiling vnrious seutieiil
fUrfaces <>r »eiwory nervei. And cotiimon exjwrieuce afliinU many iitataixts
where veucal coutraciiouH thus brought about in a reflex mauner aoqain
airenglfa adeijuate to empty the bladder.
Obcervntion of vMical prcMure may be tnont onnvenictilly carried out \>y latn>-
tlucioi; into ihr Iilailder a mtbetrr coonrcted with a water manomoCcr and ■
rrt!>!ili'riiiK ■ppHmtiti. nnd to luranged lu to alluw fluid to tie driven into wr
received Irom the tdadder ui |iteu«ure.
$ 431. Involuntary raicturilion obvioui>ly of reflex nature ha« frequentlf
been (d>!«rved in cam's oC pamlysia from diseawof or injury to the apinu
cord ; and the involuntary micturition which is common iu children, aa the
reoiilt iif irritation uf the )>eiiiii and gtitiiial organs, and which sometimei
ovcuni iu tl>e adult aa tbe result of emotions, or at least senaory imprewioBi,
np|M«ni to l>e the result of rf^flex action. In tbeee several caaee we mar
fairly auppoee that the centnt iu thi; »i>inal cord ia aflecied by afl^reat
impulaee renchini; it along variou* twiivury nervM or deM<tndiug tmra the
bntin, Hencr wo are W tu the conception that when wr make water by ■
cousctoue effort of (he will, what occure in not a diret^t atriton of the will im
the mUHcnlnr walls of the bluddt'r, but that iuipuUm Planed by llw will
dfec«ud from the brain after the taHhioit of iiHcnnc impul*e« and thita in %
rellex manner throw into action thi' micturition centre in Ihvxpinal cord. W«
may draw an analogy between the micturition ajiparatus and the respirulorr
mechanism. We sair maona in the latter ea»e to ihiuk that wlivn the will
interfered with the reapiralorr movements, it did so by acting up«)U the ner-
vous meclmnisin in the eeniral nervous system and not by acting directly oa
tbe muscular Abrn of the diaphragm and other respiratory muKles. And
ibe ewe of the plain muscular tibres of the blwlder erams even stronger than
that of reapiraiory musclea so largely skeletal iu nature. We might alao
RUCTUBE OF TOB SKIN.^
I
I
logy wllh thu iM-iirt. We nrei nul able to throir Into ncUon, \>y
Mr dinet eflurl of tbo will, the i.-mnii«i;! uiigmcmlnr mcchnnism. Were we
*wt lo do H povrerfullj miiI Mi(](li<iily, ito miirht llirow in(o viulent action &
■takljr boathie htArt mucli in lli« Mime nay ihftt vrc empty nn ohwuroly
toQtnMrting bliuMer. Nor i« thii> view ncgntivod by the fnct that
|i&n)y*i« uf Ibc blaiUler, or nitlivr iiinbility to mak« water either voluntarily
pr ia a rvflex m»nner, i* n common eymptom ol' cerebral or «pinnl disesMc or
uijurr. Putting aside the cases iu which the rellcx act is not eallod torlh
b«^c«u8r ttte appropriate stimulus baa not been applied, the future in mictu-
■itioti under tbese cireurastancea may be oiplainvd by BUppooiDg that the
AlM>ck of the sfitnat injury or some exi^naion of the Hiseaae lias rvnilored the
•piluU ceDir« unable to act.
TIm ao-called iiieontineni'e of urine in children is simply an easily excited
and firequenlly repeated reflex micturition. In caaoi at cerebral or spinal
<!!■«*■« a form of incontinence is frefjueiUly met with which seenia to be of a
dtfl^r«nt nature. The bladder becomiiif; full. but. owiu^ to a (kilure in lira
meohaiiijiu of vulunlafy or reflex mietiiritiun, being unable to empty itadf
by K cinuplele cuniniction, a coniinuiil dribbling of urine laltes place through
t.lao uretlin, the lulnen of the bladder being auflicieol to orervoue the reHat-
MBOe M the ueck of the urethra. It in pn)tiable. however, that even in these
«aaai the How i* partly caused by obM'iire, tiufelt, inlrinnit: controvticitu of
«1m bladder.
i 4S8. \\lM)ther, under normal eunditioiu, the unnc undeigoea any notable
^MBga during ita nUty in tlie bladder baa been much debated. Experiments
Aowlbat pi>i»<>ni>ii» Mibclances intccied into the bladder with all due care 10
avoid aiir ahroNon <if the I'pitheliiim arv almirlM-d and nnuluoc thdr usual
iBklS. It ha* ulwi U-eu Milled that if a Koluiion of nrca W injocted into the
bladder after ligature of both uit^tera, and allowol to stay for xonu- houn,
fUt of the urea disappears. But ai pri9>etil thero is no very decided proof
Uiat under ordinary cowlitiom either the water or other cooetituents of uriuo
ar* tu auy appreciable extent ahdurbed by the bladder.
Uttdcr abnormal conditions, as in inflammation or irritation of tlie bladder,
the urine may have iiuderKone nnirked ohaogee during ila fttay in the blad-
^tr. (]i>e of toe mudt eomimm being a change of some »f the urea into auimo-
Oiuni ovbooate, by which the urine also l>eeome8 alkaline. Under Atmornml
ctMidttkNU alao, Ute mncu« of the urine, which in a healthy man h iiiHt},iiili-
<lMit. ibuugli in some animal*, fur iii*tniicc the liorbi', it occura in con»dL-rable
Quantity, Is largely inrreiuol during the utay in the hlailder. Since there
■Ue to man no goblet ndU in the vi^iiail upilbcltum lin the frog they are
ftrewnt) «r mucou* glan<l> in the wall* <>( the hlwlder. tbi» mucus must be
tuppticd by an aboormal metabolism of the ordinary epithelial cells.
TitK 8tuih.t[jke of tiik Skix.
§ 433. The skin, like a muooua membrane, cnnsist* of an epithelium nat-
iag UlMjn a conaective-ti»ue baais; the epithelium, which is compoeeil of
taanr1ayeraofeelU,b called epiiUrmu [F'\k. lot^J, the eon ncct ire- ties ue basis
bralleil ilermu, or eon'iim, or eutu vera. The surlace of the dermis is thrown
Up tuiu a number of elevations, pipilUe, which ditfer in Hie, form, complexity
and arrangement in dilfereni regions of the bo<ly. Souie are small, more or
has oonical elevation*, limpU papiUit. In olliers, n broader primary eleva-
Uon la divideil at its summit into a numtier of secondary elevalioni; Uteae
ara evM^iuid papUliv, In mnny regions of the iikin. oii fur examide io tJte
of the oonds. tltc |>apilhe are arnuigcil in ridgea separated bj ■hollow
&52
THE KLIMlSATtOIf OP WA0TI! I'RODOOTS.
furrows. Tlw iur&oo uf tlw Ekiii, thai is. ilie couUitir uf l>it> cfiiil«rmb, Jtm
ixA Mlow tbe pujiilUry contuiir of tlie dermb: Uic iwjiiltBry tmcor4[ngtl
nppL-nr to pltiiigp luto am) be cuveml u]> by the wore eveti p[>i(](>nnii. tl
6url»oci of wliicli, huwi^ver, U niarkeil by Uie ridgee nnij rurruWM «|>»k(& i
above u nel) u by bolder creosee and folds.
ina. iM
8
•w
-■'^X
II. lieait lajw, oeaMlnt uf i*-Mi-l*I hrnoy *»)<■ ; m, tHnlkti'CM be««y nil* ; • (, il*
luol-lain ; U. raH taiKHoiMi or Uiti-lvhuit lu;vr. (»n<iliiitiic or j>. pncklv-odlt. irienj lowtttmiri
elcmcitoil Mill A>niilnc ■ aliiiil* tlraluin ilmf Itie nfiTlnlu : uvl •jr. airUuncnnnbuinn of tj
haiw..KutIal<n) llMMMimluvldiita. ■, |iarl oT ■ |>l*iiit or MrrwHIbralii lh« •iiivtll'i«t lafw I
tlw sum ifn. Kniiii ibit plEiua Kiw tulaoM aeTFO^Urlla nHTlwli>«<d fmmtat up •wmni i
•iMtullum-osIb of (ba Sliiliil(ttt*u Ufc*.)
Th« Aurface of tli« dermis b not 4le%'eloj>ed iuta • diMluot au'l MijHinlil
biiAi-iueta iiiembnuM, as is .10 itften llic vase iu u iiiucou* moinlinini.^ : bm
tbe DtiHtt Hiiiwrficial porlioDs of ttu: deniiU th<: t'i)»ii«ctin) tiwif nlntwi' litl
ornti tibriltiuiimaiiilconaiswofaboiiiogenemnDiHtrix, innbioh iir<! ini)»-iltlc
<:0DikH?tivir-iiiuiie<^r|>iuK!le«aDd cxtremtljr fineoloKtivfibriw. T)>i;> KupiTticii
jM>rtioii of the (b'niiU, whieli in m]iQCtallj welM^vrlojioil in tlio |iu)iiltit>. servt
Dwonlinjily titu jiurgHHiM of n biuwawut iiicmbntiic. and KliBr|>ly ilclinf^ ll
derini* foiiii the nvrrlyin^ epidcnnia. At a vi-ry littU- di«UUMM fmni ili«
qiirli-nuin, lihrillutioii tiwki^ iiM a|tpoanincc tbe biiridlce of fibrilln ititorladiijj
in a iielwurk wliicli, vcrjr doac tet id tbt outer, inof« suiwrtidal layer
lii-iMiiiM riMiiv aod ntiire open id llie iuiutr. deeper parts. The cunueciii
ii<fui> of the di'rmi* thus paaove iDMmibly ioto ih« eubcutaoeuiis euiiiieL-tii
ttwu<', ID wbivli iliiok iiilt^rwuven bundles of 61>riliie, bearing in trausver
iM.-oiii>» a reriaiii nwiiibUiicx' to Kclioiit of loiidoD-buiidlea. form a ton
open iwlwork, tbe Inrger h|nioc« of which nr« fre^ueaily occupied by 101
TUE BTHl'OTl'RE OF TIIK SEtN. A53
of fnt w\\* of ihc «ul)cutnn<Mtiti niltpon tiMiie. Elusiic libm are very abiiD-
dnni in tlio ()cmii)< {»r»pcr, Iwiii); very 5ti« iiiiniediaU'ly iMDenth llie e|>i<tenniB
nnil Ws-'oniiii^ cu«rw.-r in the (leu)>cr [■t\r[«: titey xre nreaeni also, tliou^b to a
1m8 cxu-iii, in the eubcutaueous coDn«cuve linue. The skin aa a whole is a
Tcrv elofelic structure.
JiloodvMwU ar« very abuudant, formiD;; cl<«e set ca|>illRry nelirorks and
loop* immediaiely tiuder ihe epiilenuia. mpei-ially iu lliif p«pill», and mure
open networks el»enhere; )>ul uo IiIuikI vessel paasea inio llie e{>iderniu.
Lyniplialic vwaeU and Iym)>)ialti.- ca|iillaries are abumlniit in llie dernita,
beuiK cuunecied liere ba in otlier renpous of tbe body with itnialkr " lyii>|»b*
epaAX»."
The eonatderaliMn of tbe uervea of tbe skin it will lie lulvBntt^c(9»ll• lu
defer until we come lo dm) <*Uh tb« «kin w nti orgNii i)f iwuw ; f'>r thuiigh
■ome of lite niCanowiA ui^n'efibn;* ar»Cl)l'rvHt librai diittnbuu-d lo the bloMl*
Towk, aixl prolwiblr lo tin; ^wenl-glamlH and (iibcr etr>li.'turni nol diiv«lly
OOnnectMl wilb iIh' M-ane uf lotK-h, tiy I'ur the j^n-iik-r niiniWr luv alTurvDt
fibnn beginning in dutiuct Uotilu orgaiw, or oUwrwiM acrvliig as Mwwry
xlnictiirw.
J 434. TIm' cpidn-nii* mnnuu of two |>«rU, neparaloi) bv a fairly vbnrp
line of ditiuirciiliim : an innor wirt Inyi'r. the Malpi);hiiin liiyor. or *h\iluui
MiUpighii. aud an outer harder homy Inycr, or rirtilmn rorHrum. TIh; >'kin
u is mil known varies in thicknen in difiiToni n^inno of l\w Imdy. niid ihe
dilfcTroces are due nlnioet exclunvely to varimion^ in iliv thtcknoM <>f the
horny layer nhicb. tvt over tlie lipe. in»y be extremely thin, or m on the
hvcl, excesMVely thick; compared wilb the vnriittiunH in thicknct« tit' the
horny layer, the varifllions io thickue» of tbe Malpi^bian layer or of ihc
dermis may be disregarded.
llic line of demaratioo between (he jMalpi^hian and horny layers follows
tbe contour of the surface of the skin. ni>t that of the dermis, the |w»illM
of which apjtenr in sec^ou» as if imbedded in the )lalpi>;hian layer. When
the «kin after death is macerated, the homy layer is apt to |)eet '>ff from tbe
Malpi;{hian layer below, which, oriitiually «oft and rcjidere^l still mfter by
the macemiion. tbeu nppeara as a layer »f slimy li^ue Mpread out between
the sidfA of and covering the »nnimii« of tlte papillte of the (termio, aome-
what after the laxhion of a network ; hence this layer was in old tiinee
•pfiken <tf an the rrlr muroinim.
Tlte lowennuM. innernwMi [Hirtion of the Slalpighian layer reBlinjc iipoti
tlie dermu. coOHUt^ of a niu{;Ie Inv'^r of e)oMK»ted. «r almiwt cohiuinitr cells
placed viTtii'ally. that is n'ith tht'ir lung axi.i p^TjieDilicular to the plain- of
the ile.rmi;-. Thin layer, which pmcrvct tlvc original ffuturtu of the rpihlaHt
of llx' iMiihryo. an<l which niny lie followed over the jiapillw lU welt aa alons
the intorvCDil^ valleyif. proi^nts n charael<rri«tio apMUnuioc in vcrttcttl
M'cliims of ibe skin. Knch cell, which it ahmit a« large m h tuiioDcyie,
about r2/> by 6 m. consisif uf a fdlativcly large oval nudeus lying lu tho
midot of a voarsely granular cetl-sulwtanci^, which slain* rewtily with tho
ordinary staining reagents. The baf« of the veil abutting on the ilermia
ofiea shows fine jiruocsses interlocking nith oorresiiondiDg prooeates from
tJie dermis ; the mU* of the colls arc in doM contact, but merely i» oontaci,
tlo cement substance existing between thorn.
The nest of tbe cells of the Mulpighinn layer, much like oK'h other, are
polvKi>nnl or irregularly cubical cells, resembling the vertical ci-lli! juat
sjiokeD of iu 8o far thai each consists of a coarsely granular ccll-»iili>l»nc«
iu whidi is imbedded a relatively large nucleus: this, however, is ^phoricil
not ovaL The surface of each cell is thrown up into short ridge*. ra<liating
somewhat irregularly from the centre of the cell and prujediiig at lh« Nc
su
TBS ELllirirATIOy or ir^STB VH0IirCT3.
face aD<) et\g^, so «a to give the cell som«w)i«t lli« appenntice of ha'tae
aruied wiili a ntiaiber of prickles. Ueacc: these cells are oft«n cmlM
" prickle cells." The prickles of a cell do not iiilerluck witb tboee of iu
ndghbord but touch at their poiuls, su that the contact of ivro adjaoent c»ll*
i> uot complete but carried out by the pointd uf tlie prioklea ouly. miiiiitv
ipacea beiii^ Ii!ll )>elwceii. Hence tbe whnk- Malpifchian layer b traveraed
by n liiiiyntilh of minute paiwi^, along wliioh lluill cau pasi beiwceo the
tmicliiiig iirirklnt.
In iIiirK pkins, ns that of the negro, pifimrut nnrtit^lcBi alMuntl in the
lower Malpi^hiiui cdla, esj>ecially in the vertical layer. In Mich caaei
bruiichc<l pigiiieril' cells, connect! ve-tiHue ci>rpUHi.-li» loafM with pigmeat
griiim)rfl,ar«to bo eocn in the dermis aIko; and occnsioiiiilly !iniitar branched
cclU may be seeo in the epidermis betneen the Malpighian cells. Leuco-
cyte* also not infrequently pan out of the dermis and irander among lb*
celUof the Mslpighinn layer.
The nuclei not only of ibe vertical but also of the other polygonal cdli
may, not unrrftiueatly, be observed in various etaeea of karynniituMi
Throughout life the cells of this Malpighian layer of the skin appear tn bi
undergoing in u 111 plication bv division ; the increase of population (bus arir
ing is Kept down by the cells [wssing upward and outward, and bocomiog
tnosforitied iuto ibe cells of the huniy layer.
S 43fi. The line of deinarcutinn between the Malpighian layer and ibi
horny layer is, oa we bavc waid, »hnrp iin<l <listinct. It is furnished by tm
peculiar «lr»ia of e«Il*, more conKpicuuuii iu sume regions uf the skiu tboo
10 otbeni. The lotverraoat, inncrmimi utratum cuiwiHta of a iin^le layer or uf
two or three hiys of ocll« which arn not unlike Malptghian tells, but aie
diR^tentiated by their form, being extended horizontally to a* froiuenily to
■p^ir fusiform in vertical seclioas, by the absence of ptickUv. by Ihtir
■taming very deeply with certain reagents, such ns oaintc acid, and esp*-
ctnlly by their cell siibstance being crowded with large discrete granule* uf
a peculiar nature. Hence this stratum is called the ilnttitm gmnniwrun.
The stratum above this cimM^is of one or two or even more layent uf cells,
elongated and tiaiiened horii'>ntallv, the cell subataoce of which is homo-
geneuua and iranapareut, free from granules and not staining very readily.
In the middle of a cell may freijtienily be seen a rod shaped nucleus placed
boriKimtally. Thenc dear tniuspareiit celU form a tran«|wreai team, the
tlr>itum luel'linn, between tbe stratum grauulo^um and Mal|>tf;hiau layer
below aud the horny layer above.
S 436. The horny layer, wbicb is at we have saiil of variable Iwt nearly
nlwavH of considerable' thicknfev, in formed of a number of layers of cells
whicn, differentiated already in the lowtsi luyent, have that diffnrentialioa
ooniplctcit us theco pnsa unwiinl. The upper, outer portion of this bomj'
layer is Goniinuallv being shed or rubbed on in the form of Hake* of vari-
able si w. Each Duke upon examination, as for instance after diaMciaiion
by maceration or with the help of aikalics. \* found to be composed of tie*
meuts wliich can no longer be recogiiij^l n* cells, and which nmy be spoken
uf as »ca1«M. Kucli scale is a tlnttcned mii«« or plate in which no nucleus
can l>e seen, aud which oonsiata not of the proleidn and other couslituents of
ortlinary cell subiiiance (§29), but almost exclusively of a ninterial called
trnitin' TbiH ii a body, the exact nature of which has not vet been clearly
mmie out, hut which hits the general percentage composition of proteids.
from which it i' a derivnie. with the exception that it contains a considerable
ounnlily of sulphur t,the kertitin of hair contains as much as -^ per cenL);
this fulphiir npp<-an> to bt; soini^whnt lonselv attached to the other eleraeai*
of the ki.-nitin *incc it may be removed by boiling uiih alkaliea.
THE 3TKCCTt*BB OT TEZ 81CIIT.
Thr liiw«rin<i*t |>'>rlii>iH of the liaroy Inyer are composed of elerupnia
wkirb may uilt l>e recuKiiiied a> cells, iotuoiiicb na each contuina & nucleus,
Ibnuch ilii* ii obvioiiiiy uDdersoiag ohkqge Kud on the way U> diaappesr.
(la^h odl U, hnwever, flaiwoea and plate-like.
aoil iu •utwiwiM- nlrcudy oodubU largely of [Fn-H*.
lwr«Ua. lu jMuwiii^ iifiwttrd fntm the lowtr to
tbs man BUiwrficial luirU of thu horny luvrr
•ueb ui iBiiM-rfrct <x-li loan Ju niicleiiii. hihI "htt-
MBMB thr whulty kvnUiiiMi* plnt« Juitt deacnlxrd.
The wboU horny layer eounivta of strnta of cl»-
n«iit«, homy to b«rin vith, but lH.>cDaiinc mon
nnpktely to in the iipiier |iart». Bekiw. in
(untttct nith the inowt Malpighinn Ittyer, thft
komy layer u moisl but the HUiwriicinl pnrUi b«-
^MM dry hy evaponition ; aiut hptv the strata
HHlBmitaMtC Vrom each other, the otitcr oiiet, 08
HKhave Hid. being t^hed in the form of flakt*,
which Been in the dry condition under the micro-
•c*>pe have often the appearauco of irregular
Sbrei.
The karroniitosii seen in the cells of the Mai-
pighiao layer, not only iu iJiose of the vertical
larer but in tlte otbcre at irell. »how, as we have
wkM, that lli<!W miiltttily by diviHion ; ne have
BO triilanc* of loulliplicaliim tekiii|f plac^ elw-
wbcn in the epidcnniii. Thti more .inpi'rficia]
CtUa of the Malpifjhian layer, thru»t upnurit by
the DCW corners, are trniKfurmcl into the cclle of
lie atnUum gniDuloaum ; and although we do
tot ■■ yet filUy UDderetand the exact nature of
ihe trvunformalion we may conclude that the
pvcultar (granules of theec cclU are concerned io
the manufacture of kenilin. Chtinf;cd by the
cuosumplioii of their (-ranulea in thi» mauufao-
ture, the cells of the stratum j^anulosum become
AraS tlic celU of the Biralum luetcum, and ilien
Ike raUi of Ibe diMinvtIv boniy layer, pushed
ti|>«ranl through wliieh bv the new fonnaiiona
^continually suooeediiig below them, they \mim to
Xbv varfaoc and are eventuallr nbed.
$4S7. TlUfuvif-y/""'/'. A tineat-KlamI, like
'«cb«r ehmdi, oooMUof Btecretinz iHiniun and
« coBd««tliw portion [Pig. 1591. The secreting
Ennion i' a KHtg lubulnr alveofn;! cuilet) up in a
nut and placed in (he >>ntK-ntatieoiii< cnnncclive
tiMue at suaie diatance fr»m the epiitermi*. Gen-
erally Ifat gland i» forme<) of onf mch tubule
otUy, but soiDctiinefl two inbuU^ nniic into a
duct. The duct beginnine in the knot,
eonvobilions of trhich it shara, runs a
irhat navy but oihermise straiftht courve vertically toirard the aorfooe
of [he skin on to which its lumen opens.
Through the epidermis the duct ia nothing more than a tubular pasaage
txtmraled out of the epidermis with a remarkable <'»rksorew course, the
turn* of the screw becoming more open and tlie canal wider in the upper
tn:tinMirtMiN.«<ii.un> ramt ras
fxLM or tiu Baud. lUagnUWd
Wdlamoien.)
1. 1, mnloncd iaM», etrnipotnt
llw (Und, and odIUiv la Ivo
axpmlonr diMi ; 1. It, irlileli UOM*
Loio oils finl Mill itet pwa^-
r*(is ili« «|ililDrtul« Hi f. andoptoi
lu iiirDtc* at i : Ih* ■laiiit la an.
■willed In Ikt-Tciricka, utilcli an
THB ELIHINATIOK Of WASTC PKUnrffTfl.
pnR lu ii it])pro«chM the aurfiice. lu llic Mulpi^fhUii layer Ui« cell- k.nUr
ing i>ii thi.' pttMigv are lI»tt«De<t and iiu-liiinJ il»wtivtnnl »>t lu to ntl'ini i
more or lc*i dcfiiiUc lining ; tli«rv i« it fiiniliir arrniijriini-nt but ooi »' "r >
well in lli« c<fri)L-uue layer. K«iu;hing tbo il«rmi». iii m valley b«i«na
ERjtiliii', llifl paiia«f[e beoMiiw a ngulnr duct, witli oti independent epitW
mil "f iUuwii, » dtslinct bsaetBent nMnitintDu cHilinuous wiih the upfar
Hirfiiw uf tJi« <k-rRii«, and au outer cont uf ooiioet-tire tissue elreiiErlliracit,
in Our osuw of mtti^ of tbe Isrirer f;land«. «ich as thoae of tite njciTU, «ilb
jilain muHiilnr tibrea. Tbe «{>iilHrliiiiti cunaista of iwo or ttirvta Inygn <(
Mnmli roiind«(l cellB. ench irttli a relntirdy large but aheoluldy anmll oiiclni^
gonendly ettiiuiiti; <let!jily. Tlie celU leave a uarrow tubular tbivad-lBw
TuiDMi vrbich k lined wilh n vt-ry I'liaroeierislii-- diMiuit ciiliole.
TIh^ ilui^t cuDiitiuM u< |HMWH> iliau' diarai-i(.'r§ after it haa entered tlM knwt
niid \w(^n t« |>iir!>iM! a tvriMwl <:»ur«i!. but soiiu cbaiii;<^ tiuddculv iuui Uri^d
Mcretiui; lubule. ThiK tiiiiy i>f di«tiii|{tiiHh<.^l fruoi tbi; duct by Ixmi); wiiWtjH
Wld by beiiifi; liuL--i by s xiuglu layer <>f cubical nr oiiiiiuuHr chIIa lai^
tluin tlKise uf till' durt, IxTiiriDg far^ger nucl«i. and twhavin^ <liflbre&Uf
tovaixl variouN otaining reagsat*. The himttn though fairly distinct U Dot
liued by nor trutirle im in thu duct. Lying botirven tlnj b««eR)«nt taeni-
biane and iIh^ i.i|>i(hcii«l mll». nr ratbcr iinbodud in the baM'ment tu«iDl>raut
ore seen a nundx-r of jitaiu muscular fibres disposed longitudinal ly or io id
elou){atMl spiral, and o)l(?n lorming a diatinct cont bonnUh t)i« epilbelium.
Ail in the case of other ^jlaods, we are uimble to malc« niiy stat«iu«tiC
to the nork carried on by ihe eirilheliuDt lining the duct, but we may p
ably iissiitne that the sweat is niainly secreted by tlw larger cells of
terminal roiled part of the lubule. Tlieee celU, therefore, like other
cells, are probably " loaded" and discharged ;" but as yet uo marked si
lural changes in ibe cells cormponding to these phnsea hare lioeu mi
torilr ascertained, though nf^cr tbe admin islralJoii of pilocarpine,
causes sireating, tbi- im-IIh of glands hardened in alcohol ^laiii uii>ri;>
Ihan iifiual with <-arniiiii:. It miiiit he remembered, boirever, that the r
couIaioK tiKriiially in'ilhi-r i]iut-ii:> nor pnilcid fiubdiance^, nud u'o sh>»i
llierefiirc. not csiit'i-t t'l oW-rvi' " granulcfi " in the evils.
The ix'culiarly |>laced munciilsr lilirw> liavc been stipuoaed. b^ Ihcir
traction, to assiH in the flow of sweat along the tubule. In certain cutAneai
gluiida of Ihe fi\>g, uf a relatively simple nature, there is cvidt-ncc that
secn'lioR is i-jix-led from the compamlively larm lumen by Uie contnictii
of plain mii#cular fibres in the wall of the jjlaiid. or by a oouinirlioii of
wall ils>.']f. which is contractile without being dislinctlv diflrrt-n tinted
mu-cular tissue. \ad this rather supports the above vU« ; but the tnat(
is at prmeni by no menus clear.
The coil of a sweal-glnnd la well supplied with bloodvewete in the form i
capillary naworka, and n«Tv«« have Iwen traced [o the tub«a ; but tbe exa
manner iu which lliete end U nut ns vei known.
Though pruHmi in all regioDs of tfie skiu (uf man), tbe sweut-qlands
unequally di>iribiiied. Iicinji; PMra abundaul in some rcgiMis, such as
palinsof the hiind. than in ulhetK In the axilla are )• lands of very In
sixe, and in tlicse the ducu piMM« distincllv musoular ciiMts.
i 438. Seh/wMta gieiHiU. [Fig. 160.] Tbtsw are appvudaKeii of tlio ball
A Iinir io H devcliipnicnt. in the form of a eylindt-r, of a cap of come
epidermii' xiinnouoting a papilla of the dermis nmk to the liottum of a (u
lar pit, or involution of the skin, called a hair-folliole. In llie unp«r '
of (he iiair- follicle the walls cunsist uf oi\linary :>kin wilh all it« i
dermie, Malpighian layer and corneous layer, the latter il> usual <>[
sidemhle llii<-kuees. At some little distance from the ouHilh of the foUu
\
tl»e c»ra«(Kis layer siiildeiilr c««m», nw) in the follic)<? bclovt tlm iiv I'pU
dermis is repivseniei) ))r the M)i)i>igliuiii layer, uow called ibv oiiior root-
sheaiti. and two tayen of |>e«uliar wWa,
fariDiujf Ihe inii«r r<H)i-sl)»ith. of wbich tlie IPto, tio.
otiier Ih cnllei) Ucutes aud tin- iii(i«r Hux-
ley'a layer; lliewe mav. |ierhii|#, he r»n>
iiidiiri-d ua cornwpi'iiiiin^ !■> ibc xinitiim
SruniiliMumaDd luddiim rti<;)ootivelr. The
■.•rmi" of lti<! vail of Ihv fnlliclc ix at tlic
Mine tiin« <kvi;l"i>«l into aii imtcr layer
with txiadlw of conni-dive liwue diijKiwd
chiefly loiigitailinMlly, »i)d nn inner layer of
pec-iiliar Dature, thp arr«n)i:cnient iif which
IS tmnsvcrw-. iind nhtoh at lenst etimulnio!,
if it fMilly ittr not. ti nniseiilar transverw
coat, Betvr«;n thi* dermis of the follicle
and the outer rool-«hcnlh or ,^la]I)if;hiH^
layer is a very contpieuous deJiniie hyaline
baseineni ineinbmne, so tliick that it pre-
M-nu a very easily recogniiied double con-
tour.
At the bottom of the follicle the dennU
of the wall of the folliele is c»utinuou>i with
the Hibbtance of the (dermic) |iatiilla, while
the outer roo4-sheath or Mnl|ti};hiaii laver, •^mmnf.
which here be««meH extremely thin iintf re- ^WKj" Vi
duool lo one or (wo layers, i« n-Hceleil over
the ]in|iilln, and there cxjMiidii strain into ti
niHiH of mils whieli like the cclU of the
Mal)Mghiaii hiyi-r in iJio rent of the «lcin
multiply, and by their nudtijilicntion give
ri«- I'l the ciirneriuii tioilv of the hitir. It it
Hii<I ihnt ill llioHe l)iiir» uhieh ji'wiii* u
tiiedulU the v>Tticiilly •liKjiutrcd t'lnirnio"!
C«1U of tiMi Mitlpighian layer are at the
actual •nmmii of the pupilln roiiliniiei)
upward ill the axis of the hair, us the
raediilla.
The layer of Henle, following the >ltt1-
linn layer or outer Toot-she«th on which
it mts, is similarly rellected and forms
over (he hair a sio);le layer of Hot trans-
jiareni imbricated scaler known as the
cuticle of the hair. Huxley's layer, simi-
larly rellerled, forma a similar layer of
nmilar w-nhti, but thin \» connidercil as be- / ^'
longing t<> the mot-Hhenth, and i» called the
ciiltele i>f llw riMit-itht-fltb.
JiiM where the o<ii:iiv>iis laver ahruptlr
leavesoff in thr uugicr part of ihtr hair fol-
licle, a Bcbnccoui gland o}H'Ms into the cavity
of the follicle nn enrh side of the hair.
Each k,'l"i"i coiiM^t" of n iiboTt rather wide
duct which ilividfls into a clii«tcr of some-
what Hask-sliaped nlveuli. The tMsontent
mrmbmne, Ixiili to llie alvmli niM) in llie
\
pighi
llttN-rOUICI-C IM ljU)HllirNKjtL
smiiBi
<^ nuNith of liilUvIc ; 6, nod ; r. diilb -
il.r.dcrnuo coal:/, nutrr ront-ilintli :
9, Innrrml-iliailk : A, blr: t. Ri m»-
diiUa; (, tuilr-lin»t>: *i.ailt|iMeliHln;
a. twlrmawlr, v, |ii;iillK •<! *Uui f,
Itttfltt ufUalr: t, ruiv ii><i<>i>"ui, tMi-
Unoniw tttib auloi Fwri'fbotlb : 1^
bomrlitlvt: I, "itBdjinwBliiBiJ.]
TOE KLIUINATIO.V or WASTE I'BOUt'CTS.
duct. i« liiMil will) n Injrr »r rnilier aniall ctibit-iil n\U r<iMtimi>>iiB wiiL <V
layer vf |><-rji<'ii<licuUrty <lL>piw<l oelbwhk-h I'onit titr iniUTiniBi Invvr i
tho oiilor r>jut-ahciUh M of thv AtnliML'ltiiui lavvr uf the akin geoenll;.
This layrr of colU IfAvm n wide liinM;n Ituth in tlie Blvt^iti iimt in llw duel;
this lumeti. h'lircvcr, is occiijiicd ixit ns in other glands with fluid, but nitli
cells. Botli alri-nii mid duct, iu fact, nn' fillvd with n>imde<j ur pidypuul
OtIIr which ninv be regarded as mixlilii-d cvIIg of l\w Mnlpi^binii Intir,
The whole gland, indeed^ is a sidid diverticulum of the M»liii);liiaD layvr.
In the alveoli the cells next U> the layer of cells ii»nieJiiii«ly Wainf, iht
baaenieni menibnine. though larger thau theee, reaemble iheiu iu du far thai
each oonsisls of ordinary cell eubslaDce siirrMiuding a nucleus nt aribmn
character. Tlie more central cells are ditler«oi; their cell inilMbin^ •■
undemiiDg chan^, uumerous granules or droplets, nn>e of ihem <ih^
of a (attv nature, make tbdr appearauce iii them, and llie niicln mv '^r-
cuming «lirunk and altered. Th« cells are manufiictunng fatty and nthrr
bodien and depositinK the products in tlwir own Mbstance, which, howni-r,
Ik hot heiii;; rt-ui-wed, but la dyiiix- Thee« chantceA are still awn- oliirioU' is
ihi' cell* Iving wilhiu thi- duct; the cells as indicated by the breaking up "I
t)ie uui-Ivi un- di.'«d. and the whole of the cell subataucl^ ha* b^WD tran*-
lom>e<) intii the niuierial I'lin^iiituliuK the secrMitm of the eland, called
M^m, which in <liiicharg«.i] on to the surface uf the skin tliri>ugli the nioatk
of the hair follii^li'.
Iu theiw M-biicnous glaiidi secretions, if wo may continue (o tue the ^nui,
taktt place al1cr ii fashion ditliMreiit IVcim tjiat which we hnvv bitlierto
Studied. In an nnlitinry ghuxl llie c«IU lining tliir walls of the nirmli
maiiu&clurc niiitrrial which they discharge from ihvmsolves into the lumes
to form the secretion, iheir own milMlance being at Um sante time reiwotd.
BO tJiat the same cell may conliiiue Co ninuuliKlDre and diachar;^ the aeciv
ttoti for a very prulong«d period without beinir itself destroyed. In •
eebaceouB gland tlie work of the cells immediately lining the nail i>f aa
alveolus appears limited to the task of increasing by multipttcatiim. Ofthe
new cells inus fanned, while some remain to continue the lininfc and (o carry
on the work of iheir predecessors, the rest thrust toward the centre of ttw
alveolus are bodily transformed into ibe material of the secrelKm, and duriaj
llie transformation are pushed out through the duct by the geoerntian «
new cells behind them. The secretion of sehuni, iu fact, ia a nuKlificallu
of the iinrtit-ular kind uf secretion taking place all over the skin, am] siMtkta
>if iu> nhtNlilbg of the skin. It ia ehielly the chemical tranafiirmatioii wbich
u diDVnnt in ihe two cases. Iu the skin ;;enerally tJie prolopUsitiic cell
KuMaiiie of Ihe Matpi^hian eellB is transformed into keraiiii : in lh« s^w-
CI.MUS glands it is tnuMformed iiito llie fatty and other MiuHtituentt of tke
sebum. 8<>m«, |>erhaps, may hesitate to apply Ibe word spcreiion to aucb a
proceas aa thia ; but, as we shall see later un. tiie fonuaiion of milk, whidi
(vriainty deaervca to be called a seerction, is a procAw intcrmcdjale betweea
the svcrelioii of saliva and gi»trie juicn and tlw funnation of sebum.
The sn-callc<l " ccruminous " glands of the cxtcnuil menUis of the e*r an
ewentially sweat-gtaniln. They are wrongly namoil, since the fatty material
eiM>keii "t" ii" ■■ wax " of the ear in «ecn^t«d not by thcni but by the aebaneooi
filiiud' III rijiiig U> the hain> of tlic meatus, or by the seneral epidi^rniic
tuing. 1 Ik- c>.Tuniinous glands appear at moat to supply Que pigment whioli
colon the "wax."
The Meibomian glands of the eyelids, oil the other hand, are eawniialW
the sebnceont glaiKis of the eyelashc«, the giflude of Muhl being In turu
sweat-glands.
ST OP l-KBSrtHATIoS.
5.VU
TiiK Naturr aki> Auount or pRUfvittAiiox.
t 438. Th« i|UiiDlitT of iii«it«r which l«nvM Uie human b<Klf br war of
iW akia la vmty cuiuid«nibli.-. Thus it hus been ofllimaloi thai while OjO
(nMBie |MBaM iiwkv throueh the lungs per minute. a§ much as 0.8 Krnmme
faHr*throuKhth««Kiii. llwainoimt, hnwei-er, vari«»oxireniely : h\>m beeii
<slcfil«l«(l. Ironi ilnin ^incd by enclueiii^ the arm iu a ciXHiichoiio ba;;, thai
ik UAml uiinuni of pen ignition fmm i\w vrbol« bwly ii) tweuiy-f'Xir h'ttir*
■i|^t nuigv fruni 'i to '20 kiloa; I>ui micIi ■ mode uTcalculaiioD it obvjauily
9fM lo niaoT flwircM of error.
Of ilie whole amount tbiu discharged part paaMe awar at onc« lu vnury
Ta(Mtr mix(iJ with volatile mattara, while jnn niay ivinaio fur a tiuto a<a
fluid oa (be nkiu ; the former is IVequeiilly >|H)keu of ta i-uteiuiihlf. the latter
•■ m^nri&lf, jienpiratioD or &weal. The pr<>t><irti(iu of the iiiMiiMhli.' to ihd
•riu>ibt« |>«nijiiralioa will depend ou Ihe ra))idtty<if theMcn'tion in roleieiicc
to the dryocaa, tfiroperaiure, and aiuuunt uf iiiuviMueiii of the xurr>fut)dini;
mttxttmfiben. Thua, tapptmae the rale of sccreiiou to remaia consUmu the
•lri«r ami hMur the air. and the more rapidly the nratn of air in contact
■with th« biMir are rvnencd, the ttreater it the amount of tCiuible perapiraiion
-wliirh i» 1>T ovB(H<rnliou ocHivertei] iiiIj) the iowiuibk oondilion ; and con-
TTviwIy wh«o till- nir iii cool, moint, aud rtaguant, a larga amwint of the tulal
Mtvniratioii may roniaiu on the Mn tu aemable awoaL ^finec, a» the name
impUM. «a are ourtelvo awarv »f the «rn.-<ilile pcnpiration onlv. tc mar and
fteqoMHlj do«a happen that wc xeciu m otitvclvra to bo penfnrinc largely,
wbaa in rMdity it is nut »o inu<:h the l'>l«l perspiration which b bein)[ in-
cTcaatd aa the relative propnrtion of the sensible pcnpiration. The rate of
aecrelioa may, bowerer, be *o much increased that no amount of drynei* or
bntt. or rooveoMBt of the atniqephvrc. is sulBcient to carry out the neoeuarr
«T«piiniii»ii, aod thus the aoiiaibto ]>cr«piratioa may becooie abiindaiit In a
but. dry air. .\nd praciically this is the usual occurrence, since c«rtAi»ty
a hub tero|ieratiin' cotvluces. as we ^lall point out preifrnily, to an incrvaM
of tbm accnitiou, and it u puwible that mere drvueas of tbe air hn« a similar
tdbec
Thr anioanl of jwrtpiratioii given ulT u iilRtetcd not only bv tbe coivtiliou
af thr Biuinnphprv. but nb" by tbe cireiim>taue«a of lhe'lHx)y. Thua it is
'ii6uenvcl l>v the nature and (juantity of food eaten, bv the amount of fluid
^ruiik, by tin- chunicler of exen-ite talcvn. by the relative activity of the
uiber «xi-ratiog ur^^nN, more jinrlinilarly of the kidney, by mental condi-
am) the like. VariiilitM)* ninv al.->i> Im- induced by drugs and br diaenaed
How ihcw varMHis iiiduvntm produci* tbdr clfecU we aball study
The fluid perspiration, or 'wi-al, when eollectcd, l< found to be a clear.
^aloHans lluid of a distinctly »nll lasle, with a xlrong and diMinctire oilor
^rmryinic according to the part of the body fr>3m which it is taken. Besides
aocidootal e|>idermic scales, it contains no sinietural elements.
Sweat, as a whole, is fumiubcd wrtly by the sweat-glands and lurtly by
tbe aebaceotts glands, for, ns we sfinll mc. tlte small nmonnl which simply
UMiaudca through the epidermis, a{Hirt from the glands, may he oeglei-tvi).
linw, tbe secretion* from lh«c two kinds of glsii^ls dider niilely iu nature,
and tbi! characters of the sweat, as a whole, will vary according to tbe rela-
tire ptoportioa of the two kinds of secretion. The secretion of the sebeoooiH
riuM* appears to be fairly constant, the lart;er varialiuiu of the total sweat
depmdinit chiefly »u tbe vaM,-tns activity of tlie sweat -Klandn. Hence, wlten
•Met is scanty, tli<! ouniititucutd of tbe sebum iiiflueim- lar^^Iy the cluirac-
560 THK ELIMIXATIOX OF WASTE PBODUCTS.
•
ten) of the sweat ; when, on the coDtrary, the sweat is verv abundant, tbnt
may be disregarded, and the sweat may be considered as the product ot tk
gweat-glande.
We are not able at present to make a complete statemeot as to irhtt
bodies occur exclusively in the sebum and what in the secretion of tbennat
glands. The former consists very lar^ly of fats and fatty acids, and appcan
to contain some form or forma of proteids ; but we have reason to tkinV t^
the sweat-glands secrete in small quantity some forms of ^t, and espedtlly
volatile fatty acids.
When sweat is scanty, the reaction b generally add, but when abundaa^
is alkaline; and when a portion of the skin is well washed the sweat whick
u collected immediately afterward is usually alkaline. From this we vmj
infer that the secretion of the sweat-glands is naturally alkaline, but thtf
when mixed sweat is acid ; the acidity due to &tty (or other) acida of da
sebum. In the horse, which is ungular among hair-covered animalB for iti
frequent profuse sweating, the sweat is said to be always acid and to omitaia
a considerable <|uantity uf some form of proteid. These features are proba-
bly due to the large admixture of sebum from the numerous sebaceoua glandi
connected with the hairs.
Taking ordinary sweat, such ns may be obtained by encloeing the arm ii
a bag, we may sav that in man the average amount of solids ia from 1 to 3
per cent., of which about two-thirds consist of organic substances. Tbe diid
normal constituents are: (1) Sodium chloride, with small quantities of-othcr
inorganic salts. (2) Various acids of the fatty series, such as formic, acetic;
butyric, with probably propionic, caproic, and caprylic. The presence rf
these latter is inferred from the odor ; it is probable that many vaTiOM
volatile acids are present in small quantities. Lactic acid, which has ben
reckoned as a iinrmal constituent, is stated not to be present in heahh
(3) Neutral i&ls and cholesterin ; these have been detectM even in plaeci,
such as the piilni.s of the hand, where ^baceou^^ glands are present. (4) ^m
evidence goes to hIuiw that neither urea nor any ammonia compound exvtt
ID the normal secretion to any extent, though some ol>servers nave fbunda
considerable quantity of urea (calculatetl at 10 grms. in the twenty-four
hours for the whole bmly). Apparently some small amount of nitn^en
leaves the body the skin, as a whole, but this is probably supplied fay tbs
sebuiu or by the epiderinis.
In various forms of (iiBefl(>e tbe Hwcat has been found to contain, sometima
in consiilerHblc quantities, blood, albumin, urea ({wrticularly in cholera),
uric acid, caldum oxalate, sugar < in diabetic patients), lactic acid, indujo
{or indigo-yielding bodies giviufr rise to " blue " sweat), bile, and othra: pa-
ments. Iodine and pito^iiiiin iudi<le, succinic, tartaric, and benzoic (part^
as hippuric) iiciits have l>een fouii<l in tbe sweat when taken internally M
medicines.
Culaiievti" Jitsjiiivtloii.
$440, A frog whose lungs have been removed will continue to live for
some time; and <luriiiir thut |>eriod will coutinuu not only to produce car-
bonio acid, but also to cdiisiiux^ oxygen. In other wurd:^, the frog is ablets
bicatbe without lungs. rea])irutiiiii bein^ carrieil on etlicicntly by means of
tlMskin. In mammals and iu man this cutaneous rospirntiou iia, bv reason
of tbe thickness of the epidermii', lest ricted to within very narrow limits ; and,
Mtead, it has been (juestionrd whether it can be s|X)keu of at all as a titie
nntatioo. When llie body n^miiins liir smne time iu a closed chamber to
* '^ the air i>assing in and out of the lungs bais no access i&s when tbe
TBB IVATl'RB AVP AKOPNT Of PKR9PIRATI&S.
I
I
IhkIv ia «DcI()eed in n 1arg« airliiitit b«g fSltiiii: liglilly mund the ii<v\t, »r
wli«n.> « tube in the trachen mrrW nir tn nml from [b« lung> of kii uiiininl
placed ill un nir-ti);bt box), tt i:* found thut th« nir tn (bo vhunber loMH
oxygen and guii» mrbdiiic acid. The amount »ri;urb()iii(- acid wbirh i« lliUH
thrown olf by ihtt iikiti of an ava-ragc nwii in twMity-foiir hoiini anuMint* t<>
about ID grni*., or accnrdiitf; to work- otiMTvrre to (iio more thnu) altout 4
grm*., iHcreaniie "ith « Hiw of lfiu|)cnilun; ■iid btiiig very innrkrdly mig-
mralvd by bodily excrciw. ll m staird thai ihc amount of oxygi.ii con>
Humed t« about ^qual iti voiumt' to thut of ihv carbonic acid given nlf, but
•onw- obMrvGn makr it rather )«w. It mav bu doubted, howctcr, whether
the i-arboiiic acid comes direct from the blood; il may come from dccom-
po«itiqii» tnkinc [dncr in the Hwcat — of carbonnten, for imlaiice. f^mihirlf
the oxygen which di»appear» may be »imi)ly uwd to oxidixiDg aoiiie of tlio
const ttu en ti^ of iho «w«it. It is evident inat the lofs wbicb lb« body eufl^n
through the fekin coDsislti, beeidc a ;mall <(uanlity of sodium chloride, chiefly
of water.
When an animal, a rabbit for imiance. is covered over with an iniper-
meabte vamt*h, such aa ^latin, ui that nil exit or entrance of goeeB or
liquids by the ekin is priM'enied, dealh ^lorlly eosueB. This reeult cannot
be due, as once thoui;hI, to arrest of culancoua re«]iiralioii, seeing how inei^
niflcant and doubtful is the gaseous imerchatige bv ihe skin as eomparm
with (hat by the Iudkb. Nor are the tvmti(4iins at all those of asphyxia, hut
rather of Mme kind of poinoning, marked by a very gi^at fail of tempera*
ture. which, however, seems to be the result not of dimiuiiihed piuduction of
beat, but of an iiicreaMe of th<; discharge of heat from the surfaoe. The ani-
mal may bu rulored, or at alt ereiil* iU life may be prolonged, witJi the
abatement of the itymjitomK. if tlx- gix>at lout of bttit which in evidently
taking place bi- pievniittti by (■overinglhe hmly thickly with <N)tton-n'0(d, or
keeping it in a warm atmoaphera, llie RynipKiroM have not n* vet IxtB
clearly nnalyn.-d. but they »ccin to be due in part to a pyrexiii or fever poa>
sihly cautN-d by the n'lention within or rmhiwrption into the blond of ttome
of tin- ontmiiiuentif i>f ihc ■went, or by the products of onie ■hnormal nvota-
btdtim, and in part to a dilation of itie cutaneous vcs^U caused by the apli-
cation of varnish ; owinj; to the dilated oondition of the ciilitneouK veaseld
the l(w» of heat ihriHigh (he skin i» ahuonnally large, even though the
vnmish mar not be a good conductor.
§441. Abtorption (tv ihv. nl'm. Althimith under normal circiimKlAntifa the
skin serves only as a channel of loss to the body.il liiu> been niaiuiaincd that it
may, under particular circumslancra, be a means of Kain, and the little which
we hai'e to my on this matter may perhaps be said bere. 1_'a«es are on record
where bodies are said to have gained in weight by immersion in a bath, or
by exposure to a moist almoephere during a given period, in which no food
or drink was taken, or to have gained more than the weight of the food or
drink taken ; the gain in such cases must have been due to the absorjitiou
i>f water by the skin. Direct extierimniti', however, throw doubt on theae
•Utemenis, for they show that under ordinary circumstances such a gain by
the skin is slight, being aptian-ntly due to mere imbibition of water hy Ihe
upper layers of the eiudermiH.
Abanrptioii of vnriomi Hulistances takes place very readily by abraded
■nrAoM where the denoi* U laid ham at covered only by the lowest layers
of midermb, hut it ha* hnn ik-ltateil whether suhitaneeM m atpieous solutioti
ewi M aL«iiri>rd by the skin when the epidemiit is intaet, the evidence un this
point being (Hinlrudiclory. In the ease of tJiO skin of the fmg an aliwirplioii
of water and of variniiif lUilulilc milu'tiinct's certainly luke> place. 1» llie ctUM
iouud human skin ihciv arc no rt priori rcawm-- why water cmrrying
Sfl
662
THK EMMIS'ATIOX MF WASTE I'RODL'CTS.
•ubMHiiew <liAi>iU-ect in it dIiouIiI not pasi iiiwiin) thmuKh the oorowui m
well M tlif other layer* of ihe epiileriiiis, the minxitii fi |iiis<iii;; (l«ptii()h>)t.
aniORg ntbor tliiu]^. npun tht? (.■miilitiou uf the xkii); noil cixiininn i-u i
«D00 MM]n]i> to >1ii<w ihnt it iloiat. Nevt^rlhfleiv, the rr^ulu nt' ■ctunl n. - -
iiMtnt are con Hi cling. Si>me dIimtvi'i^ maiulAiii that soluble non-viluil*
RubsUnce* aix- iiul ahiuirbocl, mul lliiil vulntilc itiilwtaiicw, such u iixltnC'
which maybe Hi-t^'cted in ihp NyHietii sfter u Wh eonliiioinj; ihrni.arr
alMi>rlie<) not by llie «kiii. hut by the miicous membrnnc of ihp rmjiirBtor^
organs, the nuliKlaiu-e nmkiug 'Hf wiiy to the latter by viiUtilizatiAD rmaa ik»*
suriaoe of- the btitli. Olhvrv. ngiiiu. hitve fouHil «riileiiec of ahaurpti«*^'
esoeciiLlly uitli volatile Riilwlancoe, «veii when care has been tuk«D la aro m-i
all «nnni ; and the gmater weight may jierhape ht> givcti to these, sincv ib^^
aeooid with Gommnti experieoce. The eonfllcl of experitnental r«iull«, hu^
ever, al iMist ehows that we Ho not fully undersltind tbe oonditions ud ~
whieh Ruch abMrption tnke« place.
There is, mnreorer. evidence that even solid particles can pass through :
iiilaei i^kio. The lyniphatica in the skin of a newborn iufaiit have W\ ■ .11
fiiuiiit crowded with the partielee of tbc peculiar fatty aeeretiun which ein-e— n
the skin at birth ; and mlid particles rubbed into even lh« sound ?kin ui&_
eei>eciatly when applied in a falty vehicle, as, *■. <?.. in tbe well known
cury ointinoni, find tbeir way into the underlying lymphatics. Tbe wande
ing leiii''>cyte« which are at times found amun); ibe e]Hderinic oelb
IwrhaiKt take part in this Ifaii^mrt.
The Mbciiamsu of the B»-Reriox of Sweat.
1& 442. In dealing with ibe manner in which vurioiu drcum3laneu> afi:~^
tbe amount of Hwenc ivcruted w« may, as we have already said, consider -^cJ^
sweat as a whole to hn supplied by tlie »wc«t-gland» alone. For thouft^ it
aeema cvldenl that Mime umouni of fluid muM poM by simple traiiiudac:.^iM
through the iMiliniiry epidermic of ih" portions of »kin interveaios beHr^ewi
tite mouths of tbe gland*, yot on the whole it is probable that the pnrC-M
which M> payees is a i>ma1l fmction nnly of the total quantity ^creieil br tbr
akin ; and direct cxjHTiment shows that even the sinipte eTU|M)ralion of nK«,tfr
in much i^reater from thi.«o tHirte of the skin in which tbe glunds lire al> ain-
daut, than from thwe in which ihcy are eiiiniy. We have us yet no rvidt^Wv
tlial tbe sebaccouB glands vary in activity ; tlieir very jieeuliar form nffc^^f*.
tion, if we may spcfsk of it &» a accretion, is not adaplfl lo HUilden ehiiia ^n,
•nd at all events wc have as vet no evidence tliat cireuriiHiani'e* rapjdlv auiil
largely modify ibe amount of sebum ilirchar)>ed by In-uItJiy scbunsiug gla.Biik
The secreting activity of the skin, like that of ihe other glaiidi*. is umb anil*
•ocompanied and aided by vascular dilation. lo one of the early exK*«i>
menta on divisiim of the cervical sympathetic. 11 wa^i olMervcd that in dit
ome »f the horse, the vascular dilation of tbe fiiee on the side operated <a
ws* iiccofn{)aniecl by increased pe»piratiu». Indceil, the eonneciiun Iwtwvera
tbe slate of the eutaueoiu bloodveaseU and the aniouui of porspiratJ'Hi i>*
malter of daily ohiiervutiou. When the v&iseb of the »kin are ci>nsin<7l'<I.
tbe secretion of the «kin i« diminished; when they are dilated, ii bee* xim*
abundant. In thU way. lu we *hiill Inler 011 point i>ul. tbe tem|ienitiii-A uf
the l>ody is largely regulated. When the -urrDiimling almosphen- in wnroL
Ibe cutaneous vissa-b arc dilate<l. lb<- nnioiinl of #wpnt seervle>l in tncresMif,
and tiK- conse<]umtly nugmcuieil I'vaporiuion tends to oiol dunn the Irodr.
On Ike other hand, when tbe aimwphere U cold, the eulaoeuua vmhIa in
!. p0ra{iinitiaa i* vouity,
ItMMintion.
Llaou)«lo);ir nhlilhcolht'rwvretiutjorgnno ivliich we hnvr nlrfiulr «iudi«d
Ml nflk however. i<> iiiTor tlmt there ttn >|Hvinl nrrv» ilircvily governing
be anivitr of tiw 8iid'>ripur<>iis gliind*, indcfH-ntlcnt uf vniinlions in ihv \m-
illmr fiipply. And nut oiilv t* tlii» \kv MiggriFlH liy miiny fncte. Fiich as
•f (irufitw perapimtion of tfic death Kgony, »f vnriodi cr'itcf of diwam, and
certain mental enioiioii*. mid the mid Hwuntj occurring in phtliisis and
liar nmiadNe, in sU of whicli dte ftkiii ii unwmic riiiher than hypenuwmic.
It we have direct exjierimeutAl cvidedct- of n uervoua meohanbiii of
'rsjiinUiOD u c(>in|)lete ud Itie ru^tmotor ineohnnisni.
Jl iu the cal' ilic |>eripheriit slump of the divided sciatic nerve be siiniu-
l«l with the interrupied cunviil, drop^ of aweul may readily he obaerved
gmcher on lli« luurleaa s»le of lh« fout of thnt side. The sweatia^ is not
W lo any iDcrease of blood-aupply, for it may he oh«erved when the uuta-
oiw reanb are thrown into a «iale of coiiatriction by ilie ttimulus. or even
^•o tbe aorta or crurml ark-ry h claiiijK^d pr^TJoiiB to the Minulation, nnd,
AvmA, may Ih^ ohminud tiy tiimulutinj; the aciatio nerve of a recently ampu*
lad leg. Murvovcr. when nlrupiue hns been injected, the stimulnlW pro-
icos no sveat. lliout;h viLiumohtr elTecU follow tu umuhI. The i>timl<i);y
tlTMO iha cwm-iflamU of ili« foot and »uch a xlimd us the mihtnitxillury
in fkct very doM. and Wf are jimlified in »|ienkiHg of the seuitic nerve a^i
nUiUiiBg Menilory fihrm diMrihuU-d to the eudoriparou* glandi uf (li*r foot,
niilar rvMilli may Ui ohtuiiied «ilh the nervtu of the fore liiuh. And io
inwivr* a copioun itcerciioii of «w<Mt niuy he induced by teUiDizing ttiroiit(h
i<* akin tW n^-rvfls of the limb* or the face.
If a cat in uhich the K-iatio nrn.-e htut l>eGn divided on one vide bo exposed
t B liiuh Icmpemtiirc in a heatrd chamtwr, tbe limb th« nerre of which has
eeo divideil remains dry, while the feet of the other limbi sweat freelv.
tib mull shows that the swejiting which L> caused by exiKwure of the body
t high temperaluroa is brought about by the agency of the central nervous
fwtteai. and not by a local acli^i on ihe sweat-|[lands ; for tbe fiMit of tbe
Inb NhiH« nerve hu been divided is e'{UaUy exposed to the hif;h lenipera-
um. A hi^-h ieiu|>eraluro il is true up to a certain limit increased the Jrriia-
illity of i)ic e|Hthelium of the swont-^tand! and predisitosM it to secrete, just
• it proDHXtis uc'tiou in ilie ca^eiif a muscle or oerve or other f»ru)i of living
nbttflucei Thus sliumlaiiou uf the sciatic in the cat pnxluoei a mueh more
ibuDdont secretion iu a limb ex|>oied to a lemjieraiure »f ■'15° or •omewhat
lbov«, Uuui in one which has been exposed to n ilialiuetly lower tenipenlure.
lad in a limb nhich luu bei-u pUceil iu ioe cold water lianlly any secretion
It all can be ^ined ; but apjiareutly mere ri«e of tvaifieraturv without nerve-
MiMuIatMHi will uui i;ive rise lo a secretorv activity of lh« ({land*. Tlu>
rmating caiuwil by a dy»j>n<eic condition of blood, and lucli nupvan to be
A« •wgat of the oeoih agutiy, is similarly bniujfht about bv die agency of
IJ10 MUtnd nervous nstem. when an auioia) mth the acintio nerve divukd
Ml ouB fide is made dyspnaic, no sweat apiwan in the hind lioib of ilial aide,
tlwugb abuodanoe i* wec-n iu tliv utlicr fevt.
Hwralin^ may )h- lirrnigfal about as n rullea act. Thu» when the central
iluiKp of llie divideil M'ialic is Mimiilntnl xwealing b tudu(x-<l in the olh«r
limbs, and in oiinttlrai clte iulmdueti»u of pungent snbttancc* into the moUtJl
irlll fre(|tu-nlly glre rise to a ciii>ioui |ienipiraiion over the side of the tkot.
Iht baitta* wita nf Um ftM bnt Dot (m sdj imn uf ttw twlr •■ni^d
tUMlu aii.l otuet
K • Ttm <M ■■•«* fMalr In
Hjk kail*. Ttwikfalma**
i ansa s*—iH. Urn Jatu brti.^ .^ , . _ _ .
MsaSSStHar boi bi navl ai slL Tb* tasui oTiha Mg ut>MM tmlj : suil IM nnen ttutiua •whi- j
554
IMINAilUN Uf U ASIE I'RODl
fiioc itod wlgcs. ao as lo give tho coll ■omcirhat the appcanutea ofbthii
anned with n nnmbcr of |>rick)i9U Henofl thwa oell« are oflan odid
" [iricklc cells." The |>ricki(9 of n c«ll do not interlock with Uiom ■( lb
neighbon but (oiicli at tlip'ir poial«. eo itint the contact of tiro a^jaomteeik
i» not complt'tc btil carried out by tfao poinu of th« prickW only, niDMt
spaces beiag lefl between, llenco the whole MaipighinD layer a tranml
by n labyrinth of minute pusfaj^, along which fluid can paw betirHiiibe
touchiui; uricklei.
Id dnrb skins, aa tliut of the ne^ro, pigment particles abooDd in ilit
lower Malpi^hian ci-lls, ihipiidally in t]i« I'ertical layer. In Mch caw
breached piKiiit^nt-cclltf. cimni^clive-tiMue orpusclee loa>lt<d with pigmeal
grauules, are to be lemi in tho dormiM hIso ; and ocnuiuHitlly similar brancliM
cells inay lie xeea in the cpidcrmi* brtwccii the MalpiKliiaii cclU. Leutv
cytea also iml infrciiurntly puu otil of the d«mii» aud wM»dar ainoag th*
celb of the Mnljii^hmii litror.
The nuclei not only of the vertical but also of the oth«r nolygonal c«^}»
may, uot unfrvqucntly, be oluorvci) in varioiix rtag«« of kuryutnitci^
Til rou fill I lilt life tlic cell" of (his Malpigliiiui liiyi-mf th« »kiii appear l«> ^
untliT^<ii]i|r null lip] initioii br division; the iiicrciuw uf poptiUtion ibws^
iiig is kept down by the cells pfming upward and outward, and beooit*^ i"*
tnuiffornu'il into tlic cells of the homy layer.
$436. The line of deraarcntion between the Malpighian layer nad ""
horny layer is, an we have said, sliarji and distinct, h is furnished by ^1
!>C4.'i]iiikr strata of cells, more conspicuous in some ngjons of the skin t--^'"^
n others. The lowermost, innertuost stratum coneista of a single layer o - "*
two or three lays of cells which are uot unlike Malpighian cells, but ^
diflbrentiated by their form, beiny; extended hortuiuuilly so a» frequent Iji^^.^ ^^
apjwnr fusiforni iu vertical secliuna, by the ab«enco of pricklw. by lt^K_j|^
Mtaiuing very tleenly with certnin reagents, such as oamic acid, and '^■^^flr
clally bv their oell tulwlancv l»eiiig crowded with large dincrele granulo^^^
a peculiar nature. Hence this stratum is called thi: *(ru/uin jfranutmtum. — .
TlMHlmtum above this ciin»i«ts of one or two or even more la^en ufu—' ™
elongated and Hatloncd hnrixonuilly. the coll sulutance of which i« ho^ ^n^
feneuuK and tratisparcnt, tree from gratiule* ami not «4jiiiiing very read ^ '"'
Q the middle of a cell may fmiuciitly bo seim it rod ihaped nucleus pla-
horiEontiilly. TheM clenr tninspnrent cclU form a lmn>iittrent >enm.
Mfrviliiiii fiieiihiin, between the slratum grnnulosuai and .Mnlpighiun ti
below and the horny layer above.
$ 4S6. Tbe h»rny layer, which is as we have said of variable but nc
always of couaiderable thickness, is formed of a number of layeni of o
which. dilTerentiated already in the lowest layers, have tbnt differential
completed an these pass upward- Tbe upjier, outer portion of this ho
layer is ooulinimllr beiujc shed i>r rubbed off in the form of flakes nf v
able aire. Kach flake upon examination, nt fur iiutance aAer diaaociaui
by niiiceratiou or with the help of alkalies, is found t^ be composed of W'
roent* which can no longer be re«uguixe<l as celb, a»<l which maj be spaki
of IU )ichU9. Eaeh sciue la a flattened maw or plate in wtiioh no nu '
can Iw Kva. and which omuisu not nf tbe proteid* and otJier contttiiuent* a —
ortlinary cell subatanoo (S>-9), but itlmo^t excluiiively of a mnterial calltjc^'^
fteniin. This is a body, the exact nature nf whicli bii* not vet licen cltarly^it-y
made out, but which has the general percentage coin]Kii>iiion of prottidi, ^^^
from which it is a deriviilc, with the exception that it contains a consMlerabU ^^
quantity of sulphur (,lhe kcriitin of hair contain* as much as A per cant.): ^ '
this sulphur appears to l>e somcivhnt looselv attached to the other clemeoti
of the keratiu since it may be removed by boiling with alkalies.
r
THE 9TRPCTTJa« OP THK fUtlS.
ventiofit }>-.>rlii>n« of the horuy layer ara compoapiJ of eletiK^ntg
|v Mill Ik reoJVTiized lu celU. iiiiuoiucb na each contaiDs a iiucleu*,
Itia it ubviously undeiyoing chaii^ and od the vaj to dLsappear.
U 11, bowerer, Baiieueil nntl plate-like.
■ubitB»c« already coDsisU largely of I"*- ""-
[In pawiD^ upward fW>in the lower to
I tuptrfipial riarts of the horny laver
[tnitK-rfn-t evil 1iim-« ilti tiiicleuH. andW-
B wnolly kvniliiiixiii plntc juiit di-Hcrilml.
K horny Inrtr cimiiitU or ulnitii of t-lt!-
■rny to becin with, but becoming mom
\j to in llw uprxT part*. BiToir, in
rilh the mwm Slalpij^binn liivvr. tbv
iinuaat but thcvutwHicial pnrlfi bi-
by eTSporaliun : ami briv tlu- Htriiln
rrom t-nch other, the "titer <iti(«. tig
Ciid, being theil in the form of ttnhes,
in the ary coixtition uiwlcr the micro-
hre often the appearance of irregular
knoniitosis seen in the cells of the Mal-
Inyer, not only in iboae of the venioal
i in the ulhera as n'ell. iliow, a^ ne have
[ thwe iiiuliiiily by diviaiou: we have
IM of iDulti plication taking ptai-e elxe-
tbc epklermis. The tixire !tu|ierfiriul
Malpigliiaii layer, thruM upward by
icr>, arc imn«fi>nnrd into the ci-ll« nf
im KntttuloBum : and although wo do
fuHy understiind tbc exact nature i>f
ifornintioR we may conclude that the
;nnulm of iheeo cells ar« concerned in
tacture of keratin. Chnnt;cl by (he
lk>n of their );ninul(« in lliU mnuiifaL--
celb of the ('tmlum grauulosum become
celU of the stratum lucicuiu.aud then
I of the diatinctir homy layer, putilied
Uirou^h which by the new fiiriuatiou*
Uy Bucveediiift below them, they pass to
toe and are eventually xbeil.
\The nueat-gliiniit. A sweat-jtland, like
pb, oooHtU of a *eeretiuK ]K>r(iou niiil
pSK portion [1^;. \fAi'\. The lefreiing
: a long tubular alviulu* coileil up in a
pUcea in thv nubcutnuemiH cotinrclivB
mine dintancc fmm the cpiilerniix. Gen-
t gland \« formed of on<- «uch tnbule
: >nnM>lim(!fl two tubuk^ unite into n
lucL The duct beginning in the knot.
iNivolutionR of which it *hnr««, ninit a
\ wavy but otherwise Ktmighl course vertically ton-ard the surface
on to which its lumen ofH-ne.
the epidermis the duct w nothiDK more than n tubular pnsanee
out of the epidermis with a remarkable c»rk^rew coiinte, tlie
e screw becoiuini; more open and tlie oniml n't<ler in the ujiper
V'
8t'i<oKirAai)Mc;u»i>nKiii twa
Paui or Tin Banii lUagnUM
411 dlamrun.)
1. 1. ■xiDlurU'l lolMi. nnninlni
lln ctend. *dA doIUIic In two
•leiMotydilcla^^Z. Which unit*
(DM mw (plni <Bui UiM p«rt^
OKM Ika ipldtnnii ■■ «, and opoa
itaiurbmat^: Um iiuid to «»-
bf Med tn bt-*(tfel<a. nUtk an
Hrli &I 6. S.I
^part u it niifiriMH-hts the aurfnce. In iIk- Malpiiriruiii Inror iho «eUi wfH
ling on tho |juK«»gi- mre flutteaed mid iiK-liiK-<l •l«>tMin iinl *n utoftllix'
' more or ItUt^'U-finilc lining; tliotv i« n gimiUr itrrun^fiiK-iit hul tioi «
H«ii in Lho vuri)c:oii!> lK}or. R«iu'tiiti^' iIk- iIitiuu. in it vittlvj b*t
1m|)iliii-, lb<r pnsMge beconiM a I'Ogtilnr duel, with so iDdvpttodttiit t|>ii V
iuni itf il« own, ft disliDc-l bnsenicnl nipinltmiiv ODtinudua with the u|>^^,,
eurlnoe of tlie ilprni'is, iind iiu outer cout ur i.-oniit>c[irc tissue slMilxlb«CL m,\
in the ciuo of mnw of the lar{^r glands, eiicli lis thoec of iho RxiTlft. v- j;^
I [ijiun niuHcular <ibrt«. The epithelium cunAieU of two or threv larvtv [^
Umtnll rwunded (.'ells, each with a. relatively large but absoluWly (iiuill t)ui-i^«ju^
hjiQerally ataiuint; de«{>Iv. The cells leave a narroir tubtihtr thivad-Lc|(
r IVDion which is lined wil)i a v«ry characierieiic distinct cuticle.
The duel coniinuu to |»aMeit( these i-haraL-i«r< al'i«r il hiu entered lh« knot
and t>eguD l« pumite a twiMed ociiinH*, hul «h>u cliiuiKes uiddeiily into tb«
Kcretiu}( tubule. Tlii.i may be di»tin){ui^hed fnnu the <luol by beinj; viitr,
and by being tiiird by a ninglv layvr of ciibieal ur oolumuar oelu larger
than thine of the ducri, heurin); utrger ntiolei. aiid behaving diflereulj
towani variouii ntnining rva^nbf. The lumMi though fairly dialinel k M
Jiued by auv eiilirlv tu in the diicU Lyi"S l><it«rc«u the bunenwut men-
braoe and the c|i!lliclial rells, or rath<<r iinbcdiil in the bibunieat uienihnuit,
an seen a nuniWr of plain nmecular 6brc« dixpoM-d lungitudi Daily or iii aa
elongated spiral, and often forrniiig a distinct coat beneath tb« epilhi'liuin.
A* in the case of other j^laiid^. we are unable to aiakv any fiatrtia-ni u
to tlia work carried on by the epithelium lining the duct, hul «e may prob-
ably asume that the sweat is iiminlv ^ereiod by tho larger mlU of th*
torniiiial coiled partoflbe tubule. TEiese cells. iherefoni.lik«othi-r»ccrvting
cfll*. are probably " loaded " and <lisch»rged ;" but as yet no marked itruc-
tiiral chanfEca in the i^ells ciirres[>ondinK to theee pbaeca have beea MUk&c
torily luceriaioed, thoujtb after the udokiiiistratioii of pilocarpine, which
cwiscA iweatiiiK:. the cell* of glamU hardened in alcohol staia more deeply
than usual with rarinine. It miml ix rvnicmliered. however, that the HVeat
GOUtaiiiH norninllv neither niueun nor proteid Hulvilaucef, aud we ijliuuld.
lherefi>rti. not expect to obnorvo "gr«nul«" in the cell*.
The pHTuliarly placed muscular fibres have been *upi>oiei>l, bv their oon-
Iraciiou.toasHst in thfl flowofswoat along the tubule. lu c«rtaii) cutaiiiMxu
ghiiidK of ibe fi'i>g. of a relatively eimple natiia-, llicre w erideiiee thai the
w.-erettoL is ejected from ibe comparatively large lumen by the coninieti'Xi
of iiliiiu niuoriilar fibref in the wall of the }.'hiiid, ur by a <viuiracii<>ii nf tite
wr:ill iKrIf, which i.t e»iiu-actile without being dtstiiietiv diBbrenliated into
mi»cular tissue. And this rather supports the above vien ; but the iitaiMr
is at present by no roeaiis clear.
The coil of a sweat-gland is well supplied with bloodvceaels in the form of
capillary netwtirlis. and nerves hare been traced to the tube* ; but tbe exact
manner in which ibtsw end in uol as yet knowii.
Thoiigb preac^nt in all regions of tlie akiu (of roan), the sweut-glanda are
une)]ually dtstrihuUK]. being more abundant in some regions, such a« the
palms of the hand, than in othera. In the axilla are glands of Terr large
Hie, and in tluve tlie iliii-t» pivtieas dtMiiictlv muacular coatd.
^ 438. S'i"te">'u ijlamh. | l'i)c- ItiO.] 't'htwe are appendages of the liain.
A hair i* a devt^lopmcnt. in the form of a cylinder, of a cap of cinieuuA
et>idermi« iiurmounliiig a piipillu of the denuin >iuik to the bottom of a tubu-
lar pit, »r involutiriu nf the Hkin, called a bair- follicle. In the upjter (>art
t>f the hnirlutlicic the wall* eoncifl of onlinary !>kin with all it» parl4.
dermifi. Malpighiaii laver and conioou* layer, the latter at unual ol cou-
nideiable ihickuuH- At wuue little diMancv fnim the muutb ul' Llia ItfUicla
->
1
*
m
n
'•' Uyfr :>u>l')ffilv (^-iiH-^, an'l in tbi' fullioK- in-low lhi« llio C|m-
[«lif , , ;>m»iniii><l liv ihc Miiljii^'hiiin luyi-r, imw onllci) il«' uuter rrwl-
' ■hi'kili. uixl twii Inirors nf jH-c'iiluir o'lls,
forming itii- innrr r<Hil->hr«tli, nt which (he fPi". i<u,
■Hitrr i* •-nllvtl Hoiil«'« Mlij lhi> inm-r lliix-
.l«v'» Uj>'r; ihi,-*) nwr. iwrimja. he cna-
^~^^tn'i ft» comoponrling In ihe stratum
Duliitum Knd lunitium r»peetively. The
Dia of lli« wall of llic rolltcio U nt lli«
kBw linif <kvrl<>j»l into no outer layer rfi
ith iHiti'lh* of oiiiiicclive tiwue <)is|M>!)e'l
"liHlv If'Dgituiltiiiilly. niiii an inner layer of
jluir natiirp, the arranKeiDent of shicb
I Usoavenc. bikI wliicti at lenat titirDuliiiGS,
' it nallr be oo4. a iDiismilur ininsverw
Bctweeu tliis dermis of the fnllifle
the oaicr root-«JiMith or Malnighiun
b a very- eonapicuoun il(>finitr hynline
t nieniliraDe, *u thick that It pre-
t* « very cuily rec>jgnin,>iJ double oou-
'«eur.
I At Iho ImiUoiu of thi- follicle the ilrrmiK
B of tbr wall uf |h« follii'le i:> rontiniinUB with
■AeMilutance ot One (<krnii<-) pnnllln. while
FVe tiiitrr roM'thcnth or Malpiyhinn Ittror.
obtch b«r« bcvomni esircmelj thin nmf re-
i^c^l to tmv or two laym, ii rtllcclwl over
l^ pniiilla. and llv-rc expands a^in inlu n
<baw> <'f nrlU. whirh like (lie celU of ibe
^liil|>i>;hian layer in the resi of the skin
biulti[>lv, and hy iheir nntltiplieniioii atve
i-n' !•> tl>e corneous body of the hair. It iit
■ajfl iIhii in those hair^ nblch {HiM^n* a
ullu th* vertioillv di^puned lowtrni'iol
]m of llie Malpiitbiiin Inrer lire ni iho
«(u«l nimniit of t)i>- jmpillii eoniiiiued
ipwanl iu the axia of the hiiir, ni> the
-tilla.
Ttw layer of Henle, f-illonin^ |hi> Mill-
V>t)ebian layer or nuirr rfHitflivatli ou which
St mta, is Mmilnrly reflected and fomi«
^ivrr tl»e linir a single layer of flat trani*-
{«rent inihrimeid M-alea known as tin-
ctitiele of the hair. lluxlerV layer, ami-
larly rrflwte<l, Ibrms a iimilar luyer uf
atnilar atsl««. but thi» is eom-idered as be-
lancing lo tbe root^icnih, and is called tiw
cuticlv of the rool-sbcalh.
Jiut where tbe eomeoua lover abruptlv
Icarasofl' in the upper pari of the hiiir fol-
Ucle, a aebaoeoiis nloikl opens into (be cavity
of the follicle on each side of tite hair.
Karh irlttnd iiinaistA uf n abort rather wide
dorl niiich ditidai inlu a clusler of some-
■hai fluk-ibaMtl alveoli. The baMfltfot
oirnihtnor, both in the alveoli and in the
lliia-rMurK w LowanvMiH«l>
a. ^siuliuffoltklo:^ nivk;'. bulli^
d. r. Avnnlc «iM : /. oaUtr roD«-d>cal>i ;
#, innci mnt^thMlli : K. Iiatr : t. It4 tiie-
milla: '. Iialrkni'li: •^•^l|«Uw«:
n, tMlr-muKlf^ a. jmvttltt^ tUn; p,
■■pllta of liBlr^ •. >««• Batmiiin. one-
UnuMi widi iMiu* noMbnih -. tp,
iKmjr layw : t, •vtapewia clkiid.)
568
TUB KLIUINATION OK M'ASTE ['BOUfCTS.
duct, is tiaed witli ii laytr of rntliftr sinill cubioil cvlla coiiiiououa wiib i j
layer uf i)crpctii]iculnriy d»p<w<nl cells wlik'h Ibrm (lie innermost Injtr
the outer rout-»licftth u of the IklulpiehikU layer of the skin eenml J
ThU Inycr of cells leaves a wide lumen both ui liie alveoti mid in iht ilu^^
this lumen, however, is occu]>ied not as in other inlands with 6uid. but « 5
cells. Holh alvcnli and duel. In fact, are tilled i^ilh r«>uiided or ImItl'u^*
eells which may be regarded as mixlitied cells of iJ»e Malpighiaii la>^^
The whole gland, indeed, is a s>tlid diverlienluin of ihe Muliii};hiaD Uvi-f_ 1
In the alveoli the cells next to ttie layer i>f cells iiiiiiititiat<rly Huiuij « k
haa«!iiieui nieiubrane, though larger Ihmi tlieae, reaeiiibk- iheii) in m far iVu|
each consists of ordinary* eell i«uluiatiee uurMundin^ a nucleus of onlioaM
character. The loore ceiitrul celli« are •iillvTciit ; their i.-«ll iiuWiam-v ^
undergoing change, utiuierouH j^Taiiutes or dropk-ts. nmie i>f them obviixa « r
of a futv nature, inake their niijiciiraat^ in them, nod lh« nudi-i ar« l>^
coming shrunk and altered. The itilU aiv miinufai-turinu: fntly am) ullK-r
bodies and (ivp'isitiu); the tiroducts in thriir own nubsUinoi-. which, hoitrvvr,
b not bein>; renewed, but in i]yiii)(. Thciv chanjrM are *lill murt* obviom it
the cells Iving wilhiu the diivi ; the celU as indicatvtl by the bmtking ii|* i^ |
tho nuclei an- dead, and the whole of the cell KuhstaDoc )iiv> Ih-cu trao*- <
formevl into the material constituting llic wcrvtion of the gland, called
iwAurri, whidi is diicbarge<l on to tlic nurrHoe of the Hkin tJirough the nt^iui^
of the hair Ibllicle.
In tliMC nebaectiUH gland* ncorctitms, if wo may continue to use the irori,
tak<4 placr after n fashion dilRtrent from that which we have hithcne
atadicd. In an ordiniiry gliind th«i cells lining the walls of the sln^ I
manufacture iniilcrinl which they discliarge from thcmselros into the luim
to form the nccrctioii, their own aub«tance being at the same time renewed,
so that the lame cell may continue to manufacture and discharge ihe sect»'
tioo lor a very prolonged period willioul beiu^ iuelf ilestroyed. Is a
•ebaceous gland the work of ihe cells immediately Hninj; the wall of am
alveolus np(>eai'8 liuiiteit to the task of iucreaniuK by miiltjplicjilion. Of the
now oell» tliiiB fiiruied, while some reiimin to continue the lining and to carry
oil the Work of tla-ir predeire.'uors, tbe re^t thrust fiwanl the ocntre of tlve
«lve»lu« are l>i>dily t ran 14 formed into the material of the i>ecretion, and during
tho trmnnfonnution htv punlied uui tliroUKb the duct by the generation of
new cells behind them. The ;>ecretinH of sehiini, in fact, it a ntudificatioD
of the iMrticuhir kind of iieereliun taking place all over the skin, and spoken
of lu Sledding of the skin. It is cliiefly tbe cliemicid transformaliun which
is difi^rvnt in the two cum.-*. In the skin gciierally the protoplasmic cell
eubetancc of the Malpighiim ivlls in traniduruicd iuto keratin ; in the seba>
ceoia glands it if tranKfornied ii\U> ihc fatty and other cfiustituenta of tbe
•ebom. Some, [M.-r1iii|n>, may ht^itikle to apply the word secretion to fluch a
ptOOMB U tbis : but. a* wc vhiill ore Inter on, ibo f-irniation of milk, which
eertainly deserves to be ci)I]e>] a secretion, is ■ proecw inicrmediate between
thenCKtioD of saliva and gastric juice and the fl>rmation of sebum.
The w-callcd " ceruminous" glands of the external meatus of the ear an
easentially sweat-glands. They are wrongly named, since the fatty muerUJ
spoken of as " wax " of the ear is secreted not by them but by the Mbacaoui
fjands t>elunf(in){ to the bain of the nteatua, or by ibe general epiderniir
ining. The ceruminous glands appear at most to supply the pigromt which
onion the "wax."
Tbe Meihumiaii glanda of the eyelids, on the other hand, are GweotiaUy
the sebaceous glands of tbe eyelu^lies, the glands of Mohl being in turn
vwent-glniids.
Tbk Natvbb and Aiioiryr or PeitaviKATiox.
t 438. Tlie ((uanlitv of matter which leavce the buiuau bwly hy way of
tlw akin to rtrjr ooD«l<li^rati1c. Thus ii lins lioi'ii e»liinaiv<l thitl vrhik O.Q
graniDa paan Rwrnv thniujch thv liiiitc" [x^-r miiiuio. h.h much ti* 0.8 gramnw
MMSlhrough th«*Kin. Tltviin)iiiiut,ii<twevcr, varies uxtn'mvly ; it hiu bi-^ii
oBtcatUftl. nvm dain gniiMM] by (■ucluiiinK ihv arm in a uaixiic^huiio Imjc, that
ibt IflUl KDKMint of pi'n>|>inttioii fntm ihr ii'bult' kiMly in tirvnty-f.mr boiira
Bi^it nuige fruiii 2 to iO k\U»: but iiK-b a mmU nf <3ilcuUtioii is obi,'iou»ly
a(>m lu mHDV nMTvet of vrror.
Uf tht; wtviilr antmiDl ihiii diiurhargiKl |mrt |]ii«^s away at once ■« wnteiy
nfmr iniitd with vrihiiiti; iiiatlors, wbilu part may rfmaiit for a time n* a
fluid (Ml the skin ; tbi' Itu-incr is rreiiufntly 8pi>ki.-n of n» i»mriifi/tlf. Ibc hilier
a* tniMt, penjiiratioii or nwml. The [ir<>|>nrlioti of tho inieiisibk lu ihe
teatihle penpintion will ilcpond on the nipidity of the Hcrclioii in rer^renat
bt the drynne. tcttiprntiinr. and amount of movement uf the Hurruiiiidin};
stacijibere. Thus, #upp(»ng th« rate of Kcrction to roniRiD conelaut, the
dnr and hotter the air. ana the more rapidly the Kirni« uf air in cnutai-t
wkblbe body are renewed, the );reRter i* the amount of iKnuible [lerapirat iuii
whitti b bv evajKiraiiun converted iuui (lie insensible condiitoa ; and vm-
vvnrtv whcu tbe air ia cool, moi«l, and Klagnant, a larj^ amount of the total
f'^'niiimicu may remain on the skin an sensible sweat. Since, aa the name
''npbai, «« ue ounelves aware of tbe seiL^ible penpiration onlv. ii may ami
"HiflMlitljr does happen ihat we eeeni to uur»dv(S to be perapiriiii; largely,
^^Vn In mlity it u not su muob the total penipiratjou whii^h n being in-
^*^m$td M tbe relative proportion of the seiinilile |)eraj)irati<in. The rale of
^~ ■vtion may, however, be m> muob increased that no amuuni of dri-iii.w or
t, or oMtvemeDt of the atmosphere, U autfitieiit to carr}- out the neecsMr)'
k - ^poratioo. and lhu» tbe teuime penuiratioo may beoone abumlaut in it
^*^t, dry air. Ami praciically thia is tli« ii#ual occurroace, since certainly
^ ^>i{h tero{)enitiin- conduces, as wk xhull point out pnwODtly, to an inonue
_^ the aecretion, and it ii poaoible that mere drynew of th« «ir has a aimUu'
Thu amount of pers{>iration given off is aflected not only bv the couditiou
T^^ tlir atmoapbere, but abo by the circumatancea of the bt>jy. Thus it is
^flueaoed by tbe nature and quantity of food eaten, bv the amount of fluid
^*»UDk, by tUe character of exercise taken, by the relative activity of the
*^>tH«y excretiojE orjnuiB, more particularly of the kidney, by mental condl-
^■4a(is and the like. Variations mav al>^o be ludueed by druf^ and bv dUeued
^a<an«)itioDa. How these various in^ueutvs produce iheir effects we sliall study
Kunnedtatelir.
liiv duid pentpimii'iu. or sweat, when collected, is found to be a clear,
^solarloM <lui<l of a diitiiiotlj salt taMe, with a strong and distiiici!%'e wior
^mrring ncr^irding to the part of tbe body fn>ni which it is taken. Bi»iil«s
Accidental •■piilcrraic scalea, it containti no structural elements.
^weat, m* a wtiole. u furnlibed i>artly by tbe sweotglaiids and imrtly by
the sebaiovouN glands, fur. as we »hull !»ee, the small amount nhicn siroplv
tTUtsudea through the epidi^rmb. aiwrt frum the vlandii, may lie neglected.
If ow, the sevTMions from these two kinds of jflands difler niilcly in nature,
■ud tbe characten of the sweat, as a whole, will vary acomling to the rvla-
tjra proportion of tbe two kinds of •wrvtiun. The aecreliod of tbi- seliMOOUua
gtaails appears to be Curly constant, the larver variatioiu of the total sn^t
depending chiefly on the varyinit activity of die sweat^btmls. Hence, wlm
sweat is scanty, the wnstituctits of the ■cbuni influence laiifely the charac-
580
TUF, ELIMIKATtON OP WASTE PROIll'CT8.
ten "f till' »«i-iil ; wliou. i>u the couirurc, the aWMii U verr nltuuftanl, ilie-
rnnr Ih- ■li»r<,>^rili-<i, uiii] ihe aweai tuny be cuuuJerett m» ibe pniliK't of il
BrnfHtglitiiiU,
We sm not iiMu ui rir<«(Uit to miilce a coiu|)l«t« •tatvraent »» m wh
bodies oL'ciir rxclufiivvly iii t\m wtiiiiii iiml uliat Jii tbe Mcreiiuii nl' die #Wi
glandf . Tilt.' furmcr coii^ivlii very \aiw\v vf fata und fiitty iioids, nnd «(>[••«-
ta contuiii bohip form or fiirinv ft pruteiiii ; but 1K luivc iTJUHjn la thiiiK th-^^^j
the ewent-^'lRnd! #t^.-rctv in >auUI quiinUty hmdc foniu i>f t'lit, itix) c«p«ci*l J3t
volfltile fattT nodd.
\\'hei] mem it fcuiily. llic mtdion is mn«nillj acid, but n>i«ii nhumliix-:*^
ia alkaline; nod witen ii jiuriioii uf the »Kin i« w<'II woebi'd the sw<«l whi ^:^
b colleitc'i inniicdintcly nfU-rwKrd is usimlly tilkidiiu'. From this viv nia«r
tafer thai the secretion uf iho Kweai-glnnds » inuiinilly Rlhiline, but tl^«r
nben mixed swcjit is soid ; the additv duo to fiilty (or othor) iicids of thr
llebun). In (he horse, which is Nngular among hnir-cqvrreil anitnale fur itt
IVe<iuent prufii^ sweiilin^'. the sweat is said to be always arid nnd to contac*
a ciin^iderahlu ijuanliiv uf fome farm of proleid. Tbese teaitired are profn-
l>ly due lu the large admixture of Bebmu IVt>m the nunierauB sebaceous Ktea4t
connected nith the liaire.
Taking urdiuary sweat, such na may be obtained by eticln»ing the arm in
n bug, we may aav that in man the Hverage amouni ol' solids is I'rum 1 in ~
per c«Dt., of whicb about twit-thinU eoiuiist af organie subMaucat. Tl>e cbt
normal coiiMitueuts are : (1) Sodium chloride, wilh «<man (juautitiei ofotlii
inorgiinic naltn. 1 2) Variouit acids of the fatty series, such as formic, acetic,
butyric, with probably propionic, caproic, and caprrlic The pru>enii> <^
thivc latter is inferred rrnm the oilor ; it in probable that miuiy nrious
V'dnlilc aciiU are prcMiit in small qiiantilies. Luetic acid, nhichha* been
rcckimcd lis a minnal ri^Dsiiiiicnt. is stated not to !»• prtscnt in Itcallfa.
(It) Neutral Ints and choleelerin ; ihvt/n have been dt4ccl<Hl won in placea,
such as the palm* of the hniid, where whac-eoiis glaixls are preaent. (4) 'Die
evidence goes To show thnt neither urva nor any ammonia compound exuU
in the normal secretion (o any extent, though some oWrvera liaTe found a
coui'idcrablc i]unniity of urea (calculaled at 10 grms. in tbe Iwenlv-four
hdura I'or the whole body). Appntvntly some S'MhII amount of niLmgrn
leaves tiie body the skin, as a vhole, but this is proliably supplied by (lis
lebuia or by the epidermis.
In various forms of diiensc the sweat bus been found Id contain, «omrtiinm
in coiuiderable qunnliiies, blood, albumin, urea (jmrlieulurly in chokra),
uric aciil. cakiuin oxalate, sugar (in diabetic jMtientsj. lactic arid, indigo
(or indij.'o-yiehiinjf bodies ffiving rise to " blue" sweat), bile, and other [«jf.
ntenis. loiline and potassium iodide, sumnic, tartaric, and lienznic ([lanly
as hipjNiric) acida have been found in the sweat when taken ium-njilly na
met)ivin«4.
CUtMSOiM ifoyitrartow.
{440. A frog whose lungs have l>cvn removed will continue to live for
•omfl Ume ; and during thnt jwriod will continue not only to produce car-
bouio acid, but also to c»nstim<; oxygen. In other words, tl>e frog is able lu
hrenihe withotit lungs, rcspiralioii being carrinl on ctKcieoily by inonns of
■he ^kin. In mammals and in man ihis cutaneous rcspiralinn ii^. by reasoa
of the ihickueas of tbe epidermis, icslricted to wiihin very narrow liuiu; and,
indeed, il has been ijncetioned whether it can be spoken of at all as a tni«
rtniiiration. When ihe U^dy remains for some time in a closed chum)H-r to
which tbe air jMs^ing in and out of the luU);8 hoa uu aoueas i,a> whin iba
TUX .XATVRK AND AUOLNT or PERSf IRATIO.V. fitil
fpm
l-ntri]
■4 encltwed id k Inrge air-tight biif; 6uiag tightlj* round tli« oeok, or
m liib« ill lli« tracheM <'nrri<'ti sir to and from lli« liings of an auiiiml
I in an iiir-ti(;lkt box), it is found that tlie air in tlw dmmber loMS
and jtaioti nrbonic acid. I1i« amouni of cnrbooio add w)ii<^)i is tbua
wo oH'by tb« alio of an areraKv man in twciiiy-tour tiour^ amounls to
I 10 gm*., or BCConliDft to lonw otMerveru to (no more llinii) aboiil -I i
iurrvuine with a riM of temperature snd being very luarkedly nut;-
by budiTy exerctfe. It b Btaled that the amount of DXVf^eo cuu-
tao} ia aboul eijuai in volume tu thai i>f the c-Hrbuiiic acid giviMi uff, but
itDP obMFYen make it mtl>er It^. It tunr \-v dotibted, Imuevr-r, whether
rliduir acid i^otnn direct from the hlouil ; ii may cotDe from <lecoin- 1
ni (akiiiic jilnci- ill thu awent — iiT ciirbunitln'. fur iDitanc«. Similarly
osjgen «hi«b <ttfs]>[H-ani may Ih- yiiiiiily ii-iril in oxidtdn^ wiitin of the
fiMMtltitcDia of tli« Hnciit. h i* rvid«Dl tiuil ihc l«** nhich tlic body KtiHbrw
ibruuKh llic *kin ruiuiNlii, Iwviilr ii uninll (juimlity nf widium cbtoridi^ chiefly
ofwUr.
WIm'd an animnl. a mbbit for ioMnnn-. i« onvvrvd oTcr with an imper-
MaUu vamUh, «irb a* griaiin, so thitt all oxit or volmnce of gaMc* or
lk|iiiib by (hv *kin is prfvetitcd. drnih abortly enaun. Tbis mult ennnot
W(la#. ae vaw itMNi^iht, to nrn»l of cii1ni>coii« rrapinilion, Mcing hov inigg-
ailtcaiil and doiilrtful it the gSficou» interchange bv ibo «kiu w romjMrcI
>ilj) that by thu l«og«. Nor art tite svnipti>ni8 al nli tbwr of n»)ihyxin. but
ntJivr of ftome kind of putK<ning, ninrke'l by a very grmt fall nf tcnipora-
lurr, which, howrrer, scent* to l>e ibe result not of diminished production of
amt. hot iif an increnm? of (lie diicbar|i(e of heal from the surface. The tini-
BmI luay lie restored, or at all events its life may be proloDged, with tho
ahftCrniciii of the »yniptonis, if the great lut* of bent nhich b evidently
Uktii); place lie prevented hv iiiver'tiig the body thickly with collon-wool, or
kw|nnK it in a wiirni ainiiMpht-re. Tl»c tymptomN have not an vrt l>e«n
slenrly BiutlyM^I. but iIk'v m^m to lie due in juirl to a pyrexia nr lovvt po*-
ix anaed by the n-iention within or rentworpliun intu tho hIo<Hl of agnic
UM CMMtitiienlf of the «weat, or by the pr(>i.luc<«i <d' wome ntniomtal mMN-
liw, and in jMit to a dilulion of the ciilAncous vcswls oniHed by tbo aplU
c&tti« of vnmish : owing to (he dilntc<l condilioii of the ciilaiietnn ve«cl«
tt»« la* (if bcMt thruugli the ekin is abnormnlljr Ini^, «v«n tJiougb tba
naniih mar Dot be a gwxl cnnduclur.
SMI. Aotorpli&n by the »Jtin. Although under normal rirninuliincca th«
*k>a MTveaonly as a coannel of loaa tn tli« oody. it ha« benn nisinininnl that it
"■^r.WMfer |iarti<'iilar circumstances, hen means of gain, atxt the little which
** aavc to say uii this matter may pcrha]« be fnid here, (.'tuee are on record
*'o»H) bddiea are satd to have gained in weight by immersion in n hath, or
"y npomin lo a moist atmccpbere during a given period, in irhich no food
"*' drink waa taken, or to have gained more llinn the weight of the food or
'•^ink taken ; the gain in i^uch coMs nmitt hiive been due lo the »b«iorption
**' Water by the skin. Direct ex['eriuie«l>. however, throw doubt itn these
■**lcmenU. for ihey >liow (lull under ordinary circuiuHlances such a gain by
**k« akin U flight, bt'ing iipparently due to mere imbibition of water by the
^I>Mr layen of tlie epidi-mii^
AbMrpiioB of variouii »uhilun<^es taken place very readily l>y abraded
^XirfaoM wbere the dermin ia laid ban- or mverMl only by the lovreM layers
^1 ppJdvrmi*, but it has bevn drlmtc<l wh4.-lla-r substancea in atiueous solution
I tw absorbeil by the *kin wlieu ilw epidi-ini* in Intact, the e^'idence on this
il baing cnntrwdictury. In the mm.- of the okiii of the IVog an nbinqrfion
water and of various ouloble sulwtnnccs rertainir ljike> placv. In the ciue
ikr *i>uud htiuiiin akin thi're iin' ii<i <i privri r^'iitons nhy water currying
W
j}62 THK KLIUINATIOX OK WAKTR I'BUDITCTS,
MbataticFH ilifsnlvvfl in jl ibiiuld iml piL>4 inuanl t)inini;li tlir cunM-oiMw
well as (he oiImt Uycre iif ibc epiiltrrnit^, the iiiiiniiiil m imta'tnn ile)ieD>lin;
tunoDgotlier thinip, upon tho ooniltlioit of tlw i^kin; niiii coinmnu i'i;i>tv
tace Menis to ■hoir that it doM. NcrcKhelcM, lh« rMiilts of u^'iunl i-ifirn-
mciit are ooiiflicling. Some obwrver' niuhiMin that soluble tion-volnii't
ruImuuik* are nol ab«nrbed, and that volatile Bubstonoes. aiirh n* tivltai.
wlii4-h may be dere<cted in the Byrtem after a balh oonlaiitint; thrw, ur
Blaorbed i)<>l by tlie Hkin. but by the muouus ruembrane of ilic nsiiiral/irT
nrKana, tli<! sub^tanoe luaking iU wajr la the latter by vnUttilization fpim ih
•urfiwe of the balh. (>iher«. again, have found evidence of abwirpiloo,
(■mclnlly uitb volatile subatanoea, e\'en when can baa been taken lo aviid
all erront ; «a<l tbe };raater weiji^ht may perbapa be g^ven to tlic«a. sincv ihtj
ftoooid with common ex|>erieiii-e. The oonSid of experin>cniu] roMilta. hot-
evvr, nt Imwi uhovt that ve do not fully uuderBtaud the ounditioDS osdr
whioh nucb alnorption lakea place.
Th«rv t*, luonxirur, cviitctiof that even soUil ]iartlcle(t nti [ia«:a throuj^h m
inlnct rkin. Tbr tymfihatit» in the »kin of u newboru infani have bwa
fbumi cr»wilrd wilb ihi- jiitrti(-li« nf tbe peculiar fatty M^-n-tion wbich coren
the skit) .11 liirtli ; aiirl »»lii] jinrticlis mnbed into nvi-n the hoiiii'I akiu niaf.
especially whcii apptii-d in a lully vphii-ti-, tu. r. y.. in ihc well known nwr-
entry oiiiimml. lirid tlii'ir way iiiln thi- utidi-rlyiii); lyniphalit-it The wamlrr-
tag le»eiK-vi<fi whit-h um' nl timoH found antuiig iIm epidermic uells nay
jierbaps taki- pari in tlii* iranapoit.
TiiK MacirAxiMit or nit: Si:i-RKTtow or Swb&t.
\ 44S. In dealing with the manner in which mrioiis rircunwtaooM nlicei
the amount of § neat McreCed we mnv. as n-C have almdr said, ronvidirr llir
aweat ha a whole to be siipplinl by the sweat-glands alone. For though il
Bcnns uvidcnl that some amount of fluid must paw by simple ir>ini>mlaliuD
through th« ordinary epiilermt' of the portions of skin iniervenini; lietHrrn
tlfc motithi of the glands, yet on tlie whole il is probabte that Uw (■•ninn
which m |us»ea is a small fraclJon only of the loial i|iiantity mM^rt-i^fl l>r ilir
skin : and direi-t ex|>eriment shows that even ihe Himpte evap>>raiioii of nmrr
is miirU gn-nti-r frum ibuse parts of the Bkiu in which the glniiiU at« ahiio-
Oant, than from llxve in whieh they are scanty. We bare as yet no evideiii*
that lh(^ «nl>aiviiUH glands vary in activity ; their very pevuliar form <if Mere-
tion. if wf muy s)it«k of it as a aecretion, is not adapletl ii> ouddcn (■)ia»g'&
and nl all cventa we have na yet no evidence that oircunii'l ami's nipidk and
largely modify tlie amount uf M'biini diicharged by h<-atthy Hehacciaia i;I"U<l*'
Tb(? secreting atiivjty of the akin, like that of the other glands, is usuallv
aecompanivd and aidcil liy va>«'ular dilation. In one of the early expnl'
nients on clivJHion of Ihv cervical Kynipalhetic, il was observed tUnt in lb*
case of the horse, the vaaculnr dilation of the fiux^ on the side optmieil m
WM MCumnaoiod by iilcrauied perspiration. In^leixl, the eonneetion twtWMB
tlie stale of the aitaneoua bloodv«wels and [be amount of penpiraiton m s
matter of daily oWrvntion. When the vuewls of the skin an- ennaUicUd,
Ihe secretion of tbu skin is diminisheil : when they are dilalMl, It beenati
ahundant. In this way, as we shall tnUT on point out. tbe lem|>r<mturv of
tbe hiidy is largely regiilalcil. When the surrounding atmosplirre is warni,
the eulanea^us vcsele are ililaleil. the aiuonni of sweat seeretvl ia inereiued,
and the eonsei|uently augm<'nic<) evaporaliun lemts to ciml down lliie body.
On tbe other hand, when lh« nimospliere it cold, the eniaDouua vewela an
TUK UKCHANISM or TIIK SECRETION OP SWEAT. f>63
cotiMricud, perajiifatlon U scanty, aud lees l»eat u loM to the body by
vvnporalloii.
Ihfiiiinlogr with tIi«otli<^rw-creiingorgaDsu'hirh wc have Already tnudied
l«n<l» iiH, liont^vrr, to infer thai ibrrc are f|>ecml iiervu (tireu-tly ipit-eruing
th« m-tivity •>( tbi- HiKlorijtapiuii j^lamU, imleiieinleiit of variation* in (lie viu-
ciilnr itiipply. Anil not only is lliia view nuggttittsl l>y many facut, kucIi hh
thv ]>r(ifuiw [K'ntjiiriiiion of um dcttlh agoiiy, of \'nH'>ti» vriiuw of iIimimr, ami
uf itrtnii) Rivnliil <tni'>li(tn*, and tht ciiUl «wediU (iccurriug in plithinit atid
other maladitv, in uU of wliicb the «kiu iit unaumic ral)i«r than hypernntviQic,
but we have direct cxtH-riinMilal «vi<iwirt' of a nurvntiK mechunisin of
perspirati^in as complete lu tliv vasmuotor mrt.-haiii^fn.
If in the cat' the peripheral ttiump of thu divided sciatic nerve be ■llmu-
lated witb the int«rnipted eurrcnt, anp* of sweat may readily be otwerved
to gather on the litiirlf« tole of the foot of that tiide. The sweating t« not
tlue to any incream of blood-aupply. for it tnay l>u oh«ervvd when ihe cutji-
iieuus reaeels are thrown into a state of c»iW'trictii)n by the stimulus, or even
when the aorta or crurni artery is clamjH'd previous to the stimulation, aud,
indeed, may be obtained by slimuisiiDg th* sciatic nerve of a recently ampa-
inted ]«g. Mon.'i>ver. uhen atropine has been injected, the stimulation pn>-
duoea no sueat. ihnuKh vasoniolor etlc^ts fi>llow ai usual. The analogy
between the tiwval -glands of the foot and such a gland as the auhmaxillary
i> in fact very close, and we are justified iu speaking of the sciatic nerve as
containing secretory fibres diatril>ut«d to the sudoriparous glands of the foot.
■Similar results may be obuuned witli the nerves of the f»re limb. Ami in
uumelvea a copious secretion of sweat may be induoed by letauisiug through
the flkin tlw nerves of the limbs or the faee.
If a cat in which the setatie ner\'o has been divided on one side be exnosed
lu n high lempc^ralure in a healed chamber, the limb tite nerve of which Iium
itrta divided remains dry, while the feel of the other limbs sweat fn-elv.
Thi* result showit ilinc the sweating which is caused by exposure of the body
Vt high teiD[irratun9t is brought about by the agency of the central nervous
«TMt«m, and ni>t by a local action on the sneat-glandH; fur the foot of the
limb whom' nervp ban Imm dividtHl 'mi ei|UaUy exfiuted Ui tlu- high li-m|ii'ra-
turv. A high lt;m|HTature it in true up to a certain limit tncn-iuc^t the irrila-
hility of tlw cpithi'likim of tbesweat-glamU an'I prnliipoMu it to secrotc, just
as it proinolvK action in the enseof a muwde or imrvc or other forms of living
wibfluncct. Thus itimulalion of the sciatic iu the cat produce* a much more
abtindani secrelion in a limb exposed to a tvm^ivratiire of '.i'>^ or KHnowlwt
iibove, (ban in one which has been exposed to a distinctly lower leropeniture.
and in u limb which has liocn placcfl in ice cold water hardly any secretion
at all i-au be gained ; htit apjtarently men- riav oi' temperature without nerve*
stimulation will not give rise to a secrvtorv activiiy of the glan<U. The
sweating e«use<l by a dyspna'tc condition of )>loo<l, and such nupeairs to be
the sweat of the death agony, is similarly brought about by ike agencv of
the central nervous system. When an animal with the scijitic nerve divided
III) one side is made dyspnuiic, no sncHl apjicars in Ihe hind limb uf that side,
though abundance is seen in the other feel.
t^wcaling may l>e brought about as a roflex act. Thus when tli« central
stump of the divided sciatic is Ktimiilalcd sweating is induced in the other
limbs, and in oursdves the introtluction of [xingeut substances into the luoutb
wilt frequently give rise to n c'>pioiiE |>ers])i ration over the side of the face.
1V> oil •woMi mtti kn ilio bklrlcB wla <■( the (Ml bill nirt •m unr iwri uf iu» IkiIj' ii.ivHwd
Alth iMln. t>~
U oflan •Man '
mdviut DMo >
IHM '•! thalioi-
(■I ine ROi eui nui •m unr iwri ur
I '-'"I. lU«UII|t
unit MiMC
run •iniai-
We ari^ thus leml to 8[ieak I'f e«e»l c«nira>, ualoouus tu llie viUKirnoiar
In*, as exutiiig iu iIk wuirat ii«rvoug stbImd ; and lu in (1m? ouk n( %
ntotor ceDir<«. a dispute Ittu arisen as to whether there in a tlomiDaiu ■tmi
ti) th« mecliilln obloaenta or whether »uch centra are nM>r« gfiicrnlli^ ili*-
tribtlted OV«r the ytUiAv nt the »pii)ul oiM.
It does not ai |in-9eiit H|nwttr cvrLtiin whelher the «n'vHltng ciiiSMi br lia:
id onrri^l oul li_v (lin-<'t m'tioii of ihc )ii-HlF<t hli>o<l on thi; »wc»t t.'entres, v
br the hi(,'hi-r leinpcnilun- rtimiilnliiig the i>kiii nni) so wmiling up aflenBi
[mpalses whk'h I'milLttrv ih« efH-el in n rvflrx manner: but in tb« case of
dyMUKM at Iviutt wi- may fairly mippoH- ihnt the artiun of th« veuous bi'mi
H cbwfljr if not exHii«i\'^ly on the tivrvc i-rmlrcs. Some dnii^s, such as t«i>
cnrptnr, which atum sweating, npjienr to produce (heir effect chieHy tiy i
Uk-iiI nclioD on thv ylaDdn. vinco the action oontiiiue« after the divimoiiof tiw
nerves (thongli p4lo<.itrpine ap|]nniilly hag as well some ali;>hl aoiiim an tba
nerve rentm), and the aDtagonistic action of atropine is Hiinilarlv locil.
Pi4>rotoxine and Mrirbnine appear tu prvKluce their sweating action cnialljr if
iml exclusively by aotio); on the oentral nervous syst«m, while nivutlne SMSi
to act birth centrally and ])erii>herally.
§ 449. The aweailitirea for the hind foot (in the eat) appear to louve tht
spinal curd by ihe rouia uf the lower dorsal and up[>er hiinhar itvrva, (mh
along the mmi mmmunityinlrf to lh« ahiloniinal ^yn) put hell c, ami ihtis reacb
(be cciatie nerve Thvv ihu.i follow \'ery much the coutM! of thi^ vaeo-cnc-
strictor fihrM uf the lower liiuli; but the jmrticiilur .ipinal nerves ttr wUcli
the ?weat-fihrrv iMue from the coril havi- iioi yet been d<rliiiit<dy rattled. asJ
Iioa^ibly they are in the dog mid cat the Uu't Ino nr tlic In-it three dorml aai
Irst t<To or four lumbar iierviLV. .Siniilarly the *wca I -nerves for the f(irf!-fi»t
leave the spinal cord by the mots of it'tov of the upjior (chteHy the foiirtli.
but jioisiblv al«o thi- Hfth and sixth) dorsal nerves, dom into tbu tbiinu-io
»ym)Mitbetie, thence into the ganglion stdllatuni, antt so. join the brnctilal
plexub by the line hrnncbes nnasiiig from the gaoglioii lo the spinal uervn.
Tile course to the lore-loot t* fiiiaTly along tna median and uliiur ne:
r<w]<e<.'tively. In the horse the s went -tib res lor the side of th« lace and tn
nig tliose for the snout appear to run iu brauchn of the fifth Ber%-e and i
in the fiirial ; in the latter animal at least some of these fibre* reach the li
nervit from ihe cervioil sympathetic, but appturentlv not all.
^444. Thti fact iueuliott»l above that in the nurse, alter secti<»n of iho
cervical nympatbelir nerve on one Hd« of the neck, profuse xneatinK i* b|iI
lo break out on that nide of the fiiee, has SUKgeoted the idea that lliix nrrvs
conveys inhibitory immilKS to the sweat-ponds of the head and fm^r. ui>l
thai when it ii> dividml ihc; Hweal-libra runuiug in the Hflh nerve, having
iKvihing lo coiintcntet them, «el op sweating. Uul it is prt>bably Miflicirai b
this r»H> to eupiKM' that thr glands predUpoced to activity by the highf
lemixTiilura brought about by the rteetion of the sympathetic diluting tli<>
bl-roilvessels. are mure easily exrited hv anv vtiniiiluii working njion tboiu
through the liftb nerve. And though itc i<fca of a donhle nervous nicohiui-
ism, augmenting and inhibitory, guvcnng the activity uf the sweut-glonds, is
a tempting one, there aro at prssent no satisliictory rcMons for adopting it.
rvw.
i
CHAPTER I
THE MtrTAUOLIO PR<X.'ESSI& OF THE BODY.
I
I
I
i 445. We have fnllowe*! ihe foml through m obaQgea in Ui« alimentary
ean>), nnd have seen it enter into the Mood, either direclly or hy the iuier-
tnedinu cbNniwl nf the lactenlH, in the fonn of |)ei>to»e (or oihemi^e
RKMlifitsl olhuniin I, nu^r, luetic neiti, auO fatf. accompauieil liv rariou* salu
uiul water. We have further wen that tlie n'anie }>rciain'ta wfiivh leave the
iKwly nrc urm, cmrlxinic ai-iO, Nih.-i and water. We )iitve now to Mtenipt to
ouBnect lo^her the fixKl ami the wiwUt proiliictn; Id tmvv »ul a> far a* we
■re able tlie ^iij-iou« »t(!p hy which tlie one r» tramffornwil inln the other.
Tlicn remuiiii ihe fnriher tn^k 1<> ini|iiire iiilo the manner in which the
eneivj set fri-e in ihi" trannfornuilioii t» itititri)iiit<'i| and madp dm of.
Tne master lixeiin of tbo hudv are the inuiicntar an<l nervous ti<«uaf ; all
the irther timies may (m ngarucd a* th<: •^rvmils of these. An<) wc may
fairly pmiime ihnt. ^)«sidt» the digestive luiil cxcretorj tinurn which ue
have alreiuty Mudied, many mris of the hodv arc engaged either in furil^er
elaborating the coinparntivcly raw fimd vrhieli enlers the blood, io order ihat
it may l>e ueimilated with the Icnet |)oe«ible labor by the rnaater (iseiieB, or
in BO modifyiu); the wn^te ))roducl8 which arise from the activity of the
master litwueti that they may he removed from the body as aiieedily a» |his-
sible. There can be no doubt that manifold intermediate enanges of itiis
kind do take place in the body; but our knowleil^ of the matter is at
£r««ent very imperfect. In a few inaianoee only can we localise these mela-
olic actioua and speak of (tl^tiiict metuhotic tissues. In the majority of
CMM we can only tnice out or infer chemical chanKes, without beinj; able to
ny more than tluil they do lake place Homewhere ; and in consequence, {wr-
hapo itoinewlutt luovily, kjteak of them as taking place id the hloutl.
How litllT we know ronceniing the metaboUiini of the master titwiies
tbeinselve* wiw :>hown when we wt'rt- dealing with thme tiMiiKit in an earlier
part of lhii> wnrk : but »ucc(iH >n the ntndy of thew- ^-nn hanllv W ex]ie4'ti>il
until otir knowledge '» iiiemiacd a* regard* llie chaiigts winch the hliHHl
und<in:o*3' before it rraehn> ami after it leave* tiie min'clc or tin- iiTrve. The
fact that a large jwrt I'fthi-alworlHnl food is carried through tlm liver before
it is thrown on the gcniral circulation leads uk to itupiHiM^ that in thin largo
»r)(an imporltint mctab>'lie proceaiKa are carried on i and obwrvation with
exjwrimeDt confinne this view. Important nii the secnHion of bile may be
t)ie olher metabolic functions of the liver are of still greater im|K>riitnce;
and preparatory to the study of these we may now take up the structure of
tlib orpa.
TitB STttlCTrBB OF Tn« LtVKR.
S 446. The liver i* a glan<l. tlM conducting gxirtion of wbicli. the bi)o-diict
or galUdnct. after repeated diviaioD end* in piussnge* lined by svcreticiE
simctures: but the (vnn|inratirely rinipU- arrangement seen in other glantu
in which the terminal ducts or diietuli-« end in blind, tubular or flask-shaped
alveoli is, in the liter, mudilH-i) and to a certain extent obscured. These
6tfR
TBS MKTAHOLIC PKOtlESSKS or TUB BOOT.
nKMlilicKtionR may be nseribed on (be oae banti lo t)io Wt Ihiil tho rvlla mlicii
provide tho ^eiTt-lina, beiDK aim enf^ged an ul< biivo jtiM niiil iti tiii|(tirt>nl
ntcCalxilic iltititrs, nrc ilevclojK^l init uf |)rii[iorli4>H t>i thi.* billiirr [iifea^,
nml on ibe other IihriI !•■ ihe fiirt that ihi.- i-nlinnry viiM-ular Kiiftjily <>l' u
artery (he|<ati(- arloni' i en<liiiK ibruu^'h (-n|>il)iiri» in h vriii i hcjiulit: veiii), u
overHha<iimc<l by ihi- tin-iit fHirliil iiyxlriii ; the f.'r<iil ikud wi<lu vc-rin |iurti>
(liviiJi'iH iiii<j vciiiiuri nijiiUnrii-x, nmi tlnw iirit ^ihorcd «p ngtun ini'i lfc»
befMic mi), nbirli ihitu >lrnii-« itx timin Mijiply of blood fram it rstber ikas
ftnm ihf much sDialhT lii^fiittic artvry.
Tb« «b"ile iiver, iiivivli-d nilli ii caimilv of wnmvtivc tiwue aad markol
DU! inti) ilM wTi-ntl h)lH». i» <livitk->l by mi{iU of conucclive tiawm- ■(•.'■■
niiriitK-r i>f vmall |inmiir.T tioils of loinonhat p»lyKi.'iinl t'unii, ciillnl .
each hv'mg in nin»e iibout lb« tixe of a |>iu'> heml. Th« diftUt)vUrea> •>( ■
IT*^ W.
/
^:i
- U-'
I'MiRH tii'firii K or Tax Ln'Kn.
irtflii. unH II, li>n li.bc; •'.qiiaitmtc ItilH': I>. *i4(«li*n. »iv\ K. ivn^iu l»b« : 7.
' t: <-•■ itnlI-Ma'lilFr; n. nun nvk : ft. toim (arlir: f. niunO IlKiimeiil ; rf. utiUlefaM
ttMmkj cniniinna Itciailr <hiit ; /, e]r(U« duct : p. ooouMin bllodiiMi *. licfuilo anory.l
lobule fW>ni ila nea^bon depends >m the relative abundAnce nf th« oM-
aeciive ll»ue wbi«h aeparateit them ; iitid ibU in much more i-»ns|)icuiMU in
aone luiiauila (audi aa Itie )>i^] ihuii in othera such aa the rnhMl or man).
Tlie larice ponnl vein [Fin. l''l]* ''"^ rava-h smaller bile-duct, ii»d thvatlll
•miilli-r hi-{iattc artery, e.ntcrin); tlif liver togi-ther on itit tinder flurfiK^' nt Uw
porta lifjHilien, or |ini«' "f ibi- liver, arc inve*te<l wilb a etin.iidemhlr' <|ttnnIttT
of (NXinwlivn timiii', ntrryiitu aUn lyin)>balif!< and nervM. nbii^h is (.tie'
tinuotiK with ibe ninnrrtive'lin'iie covering of the whi>W liver and is '■»Il>iT
OliMikn'» mjwnht. RM[>idlv dividing, the divinioni' conliniiinji to rgn l...
•Idu hy aide in ibc Iwdi of' (vmnecttve liniii' into which tiliaum'* (-a|».i.<: ,>
continued, the three vnvelB iilliniairly rmrh (li« oulMdtt of the serenl
luhulee, Ibe ^cpla of connoc-live liK»xii' delinin^ ill" l»lHilca from twh otWr
bcin); the torminalioDH of Oli»eotiV cajmiile carrring ihe tbrM iOta of veSMk
The "luall branches of the |>orlal voin thus rvaebintc tbo mrfuiv <>f lh»
tohiile». and iiinnin;; nnd niia$Ion)<«intf freelr belneen the lobuleH, an
s|ir>ken of as Inlrrlijiiulnr veiiiM, (FiK> Il>^.] 'Hius each lobule is prorkM,
at ilillerent jxurts of its <rircti info ranee, with two. three, or more int«rlnbular
veins, ai^iiitiiianied in a manner uhieh ne xhall devcrilie by ilivisi4>ii.i of tbr
bileijiiet Hi»l hepatie artery, all beini; enibeddtil in a i variable) ipiantiiy uf
eonni'ciivo liiaiie.
TUK STRVCTVRE OK TICK LIVSR.
5/67
I
Eiu.')i l<ib(il« Rt one |Mrl of ite drcuinlernnc« rwis directly, with thcr iutor-
vciitiim of hitnlly nny omiMM.-tiv^ lueiie at nil, tipoii n imiiaII v«iil which It
Uol jMrt of ill'' poTtn) vein, l>iit which when mic«4 otil ia tutiDiI lo past into
Kiid r»rfti tli« Iiirpatio win : it is called a uMiAalar veil). A lobule in fact,
thodgh gvnerally |iii!ylieilrnl bk *«eii in sections at the liver, may be con-
cidsnM lu anmenhat of llie furiii ul' n bniail invrrled llaak. lh« neck of which
rtMe directly •>» n »tilili)hiilnr hraiioh of \\\v hvfiaiic vein, niitl U[hiii the
]>u)y<;iinal Ik"!) ul' whit-h, >iirn>uiiiJ«i by innre or Ick* c»niiivtiv« tiMtuc, abut
at variuus |Hiiiiiii iiiinrlobiiliir l>nuichw< of lh« jxirial vein.
ino. vca.
^^^P lHw>K*iiH>ii' tLirtatwumrva or Two HtrATK lAm'ua.
^^^^1* ttft-hnDit lohulf M npNWiiKK) ullh<lirt»ti»li'1''>lurrtliivutiwlcnilli therigkt'blDiluno Uia
ncitani ■■*« tlio (vwm (rf IIw liiinJ'<balar vtUi. (•. iDwrUibulu brui«h«a of the ^rlNl i*ln. A.
lnlMli>*aiUr hnnrtx* of iha bapaitc miM: •. lublolialiK iMii : f. otWllarlM of Iti* kobali*. lb*
uimr* lii4lniU Ihr (ll>e«tte*i«< Ihcivumof IhcUood, Tlw llTwr-rtUi «r* <inl]F t«pr*itnliri In oD«
pan«f aKfelob<ii*.{
§ 447. The network of inlerlot>ulsr veine stirrotindinif (he cir^tumftirence
of n lolnile givce orifcin to a uuudter of rather nide capillary vetaeln which
run in a radial direction toward the middle of the lobule; llie«e arv omnitKUiiI
by cmm capillnHtv, which, however, are shorter and leaa abtiiidaut than the
radial L-apillnrint, m that the tno!ih«s are elongated, more or lew re<!tanKiilnr
spaces ci>iivrrt:lnu radially (owunl thi- centre (if the ]i>liul«. Tnwnrd lliv
mi<tdle of iIk* lobolt! the cHiiillarii-H mer^tt into a Nin;;le v«iii. calle<t an intra-
lobnlnr VL-in, which, running ilnwn the iii-ck <il' the lIiLak. and receiving the
mpitlariefl of the ucck uh it guc», lalU into the sublobutar vein kpokeii of
above.
Tlie elongated menhc* nf this capillary network oinvprKirij!: rapidiv toward
Uie intralobular vein nt it* lic^'innin^ in ihi^ Imdy i>f (lii> llni>k-like lofiulo and
lU It b cvnliniied alon^; the ni-ck of the tliuk. are (nviipiiil hy n'lniiToly lur^
ivdripnal nui-lealed cells, which wc ithall prtw^iitly di-^rrilK- in diiaii us hryilie
tftlf: The wiillh of a mesti is ^fciicrally such m V" admit unc or Ino ccIIh
ubrvunl.hut its leuuth adniiiH tv\-ernl celb: henne thu cvlU are urranj^cd in
narmw. radiating;, broken columns converging toward the middle of the
lubiile.
The ndtimn* of t-ellR and the meebwork of capillaries practically con-
BtilUle iIm' whole of the lobule, fur batidea a mininiuni of connective timiie
forming an advNititia to the i-apillariee, certain lymphatic |Mu«iig«« atl^irded
by thid advontitia. and exlreniely minute paang«8 which form the bejfinniiiga
M8
TnE VKTAUOMC PtlOCttS%t» Of THE SOI>r.
of th« bile-ducU, anil of wliich wl> shall Hpeak lawr on, uothiiig elw e
nrnenU The lahuU in Ikct consists Urtt of a vaacular tmaev/ttrV. of c*pil'
Urtee, which, (akinj; origin at the surfnoo of ihe lobiile from the intertobukr
Smrlal veiiU«U, are dtspooed in a network with mwbw elon^tnl in a radial
linctioD, and ooiiTef^e at the oeotra of the lobule to form the intralobiilu
v«liiliH falltuit iuio ihe sublobular (hepatic) vein, and eevondly of raduiinf
oohimna of n-Ux lilHoj^ uf> the ra<liiiuti); meabed u( thb vaa<^iilar ntlwark.
HtiicR in aaectiiiu of a hardened and |ire|M»«<l uninjected liver, !n wUdi
iIm) hloodvtsw)* unj lar^lj emptied, (he arv«M of the riectiom of lohalra iif*
indicated hv Ihu radicidlv o'livcrKing uolumnx of i;clU, and fac<^>r-'
the animal cmploywil nrr more or lew diatiiK'tlv marked o*il by thr - j
coiinoctivr ti»iio. in which may be aeen here lunl (here the liiminn <>l lii'
larger interlobular veinx. In h>hu1m in whitrh the M^tOD lirix |ini««l
Ibrou^'h thf miildlc of ihp lobulfi, tli« lunuin of ihi- orntral inlmlobtilar nio
will also lie vinihle; hut iitl«o tbu soOion tiill eiit n tobuln m HUparlidally ai
|o miss llie inlrnlobular vein allogiether; anil it ts only whci) the eei-tion
happena (o paw throu)»h the middio of the h>btde in the plane of t)ie lonj
axis of the flask, that the origin of the intralobular vein in the midille nl
the body of the flwik and iu couiw along the neck to tJra euhlobular vein a
diMplarod.
S448. If ihe section be extensive enouKh there mar he seen here and ihtn
■H-clioiM of the portal vein, hejmic arterv, and bile-duct runninj; in iiliiwoti'i
oiipaule. St-iti'iiiv or the branchea of tlie hejtatie vein Ibrnied iiy the union
of nuhlohular vi-iii.-> may alao be keen. Theae may be reco^ued by iht
abaonoe or by the extrcnw iifHniititMt «f any i-oniiet-lire-tiiMie wrapping tn
th« win, evftn in the <:nM of thi* larger bntni^ben. The widl of the vvin. l'>»,
ifl very ihiu, and ronnisU of hur<lly more tluiu th<- Uiuiea intinia nvitiiig on *
thin conuec[ivc-li"Bue biuia. moiuHilar libro* biriujj .10 verj- wfuniy (liar ihf
tunica media may be mid to bi' ab«cnl.
The wall* of the |H'rtnl vein, on the contrary, are lliick and mu^cnUr;
tlie trunk is more abundantly siippli^vl with muieulnr librai than any xliitr
Tein in tlve bodv : and lbt> liriinche« wiihin the liver aru, in diinini^hui)!
degree, thick ami muKutar. This in intelligible whco it i« remembered thai
the bloud ia distribiiied into capillaries from Ihe portal vein »t fhim u
artery ; and iudeed it has beeii mRintniiMd that (he portal vein is snhjei'i t>i
rhyllimic contractiona of its walls, as if to anist in the iMumgo onward o{
the blood. Neither in the trunk nor in ibe branohea are any valvoa prewM.
and ihuae are abo ab»enl from the branches of the hepatic vein,
The branches of the hepatic artery are ^'ery much smaller than the
bruichea of the jiortal rein, and even much smaller than the hranehea uf
tha jKirtal vein, and even much Bnialler than the branches of tht< ltili-dLi<:i
io mmpaDy with which they run. An they proceed in their courw tlii^y
HUptilv iho wall* of the {H>rtal vein* and of the bile-ducta and the tmb.-ttaiiiie
of tiilaton'ii oJipHite, and eventually dtwhari;e their blood into the portal
WdIoIa. It hat bei-n mainiiiiued thai •ome of the finer braui^liea run
directiv into the vawulnr me«hwork of the marginal parta of the Inbulo,
but ihU is dispuu^,
$ 449. '/'Ar hitf->lu'*t. Th<! larj^r bilo-durtd, namely, the bi'pMlic duet
leading from Ihc lii'er. ihc cyiittc duct Inoiltog from tlie gallbladder. aoJ
the common bile-^ioct firuM-d by the junction of the two, have the onlinory
eharaclers of large tfbind-ductii. An epithelium of o<himnar eelU runta Mi
n connective-tisHiv Dons, and ■»> constitute*! a mucous menibranu; ifaia i»
supported by a well-dox-elonrd nuiscular cmt, coitsisiing of a thicker inter __
layer of circularly dispoteil. and a Ihiiiner external layer of lonijicudiual^
tIiB|Ksed, plain muscular fibres mixetl up with a gooil deal of cuDueoti'
THS STRCCTUKB OP TriS I.ITEEH.
689
»
»
tiM(i«. Tli« walls of ibe /aD-blddder bave «weniinllT Ibe same structure.
Botli tbe Ksll-bladder aud tbe ducU are capable uf oair.vina; oul perietaltie
coDtrnciinDB uf tlieir walla, by the help of which uhen aee^d (i 25:i) tbe
ramd flow of bile luui Ihe inleoUne u«e<curei1.
Tbe bile-duclA within (he liver ar« alvu niiuitarK- vonilituled. iheir walla,
of Gouree, beoontiug ibiaiter aiid lew mtitouiar im the tube* flimiuUb i» »\t/e,
wit) the epilheliunt becoming eiibiottl rather than columnBr. A character-
Utic fcttture uf the «iimller bil^ iliiuls tt.« they run ia OliaMw's oaj»ule U that,
unlike tiie ducta of miwl ^lurxl*. Lhcy firm froiuutil aiKuHomailCd.
Tbe epilheliiim of the •Iik'Ii' c-miniii:) timDy gublcC-n^lh. niid in tl»e u'alU
of the luriccr iliicL'> aihI of Uic gBll-blndilcr I'liinll miio»ii:i gliiniU am jin^Mnl;
a»d ill tbe smaller ducts lh«c arc apt lo be «inip]iticd intf> mere pit* or ahoH
(li-|>r«)Hion9i of tbe niucoii* lucniliriinc.
riic ■itiall tormiiial niiii#tomu«irig hili--diicti!. uow coiisiHiiig of hardly
more than a cubical oplthcliuin n-st^iig on a connective- 1 twuc biui*. may Im
Intceil to various jiointMof the ninrgin of a lohiilc, and tborv went to end
iU>rupt)y. Juet before a bite-duct thus ends or teems to end, tbe cubical coll*
l>«e<>iDe much Hatier. tbe lumen of the tube, however, remaining narrow;
■ud tlieu Ihe end of tbe tube seems blocked up with the hepatic cellit of tbe
lobule^ To uoderataod. however, tbe nature of (hie jieculiar ending, we muu
returp lo tbe bepalic ceiW.
S 4S0. The hepatic cells lillini; up the meeheaof the Taacular network of
a lobule are relatively lar^e (20 to W h in diameter in man) polygonal or
roughly cubical oella. Esoh contains a relatively Inrj^e ruundeci nucteoa,
and in not a few celU two nuclei may be seen. Kach cell is parity in con-
tact with iu neighbor*, and imrtly abuta on a bloodvessel ; for there U prob-
ablr not a cell in a lobule which is not in touch, for some part of iu suriiice,
with one or more bloo<)venela. Where tbe surfaces of two eells mee4, Iheir
BubaUncee are Id contact, that in to mv, thcK is no centeul •ulniance between
tliem, iuk) the external layer uf cell MiilMtanci-. lhou|ch it may at tii»«s nt all
evenUi diflcr fmm the more iiilcnial cell Kulmttimre, i» nut dillervutiateil into
a distinct meiubmuc or culicli-. Wlirre ihr surface of a cell nbuts i>n a
bliiuiIvnOTid the substancn of the (^:I1 i* gcnerallv Miparnled from the wall of
tbe vuocl by a lympli-spacw, which in connM-lvd with tbe hepatic lymphatic
veM^.
Tlie cell subatance ilsc-lf. as might be uxpcctud from what has been already
ur^ed CDDoerning the melaboli«iD going uo in the liver, privcnto u|)|>e<inin<!C«
wbicb differ very widely according to circum«tanocs. Suniotinics ihv cell
eiilwtanco appears denee, compact, and of fairly unilhrni texture, though
more or less granular; the whole coll is then of ralntively small bulk.
8o4netinK« tbe cell suLetance appears large ami bulky, owing Ui iU being
lareely loaded with a subAance BtaiuiDg red-biuwu wiib iodine, nhicb we
slinll study in detail preeentlf, called glycogen. When such a ct.>ll i* har-
dened and Uw glycogen diasolved out, the cell aubetanee apjtears to be so
completely riddled with vacuolee as to be reduced to a mere shell, surround-
ing a loon irr^ular nctwortt except imtuedialely around tbe nucleus, where
it is more solid. But it will be beat to reserve tbe discussion of these
changea in tbe (xlU uutil we have studied to some extent the metabolic
ebaagea which take pluire in tlie liver. We may iu1<l, however, tliat verv
frequently, eipeciatly iu certain animals, the hejiatic cell is crowded with od
globules of variouH »iaai; ihwe are at ti wen no numerous as completely to
llide the nuclcu*. which oannot be seen until the fat bai been removed.
§ 451. Where tbe side* of two henatic cells are in contact, careful exami-
nation with high powera of the MncriMm|)« will «IWn reveal, at about tbe
middle of tbe liue of junction uf the two «id«f, n minute hole, a tentli or
Vkockssbs or TiiK uuur.
Kiw i>f iW ilijiiitcter ot* Ihr evil, wliich, iirmnlitig (o wtme nbeervvra, (■ Itw4
with H (]elicat« nitirtibr lining. Thin )i<>li> io thv wflliMi nf n ntinuU' OUmI
pnj^iiii; Iwtweeo the iwn rrlU in tlx* mi<l<lk lin<t of l)>«'ir nf>iM)M><l ■iirram
A ino'kl of it miglii Iw mmlc <>n twi> niiiitll bl<>ck» of dinlk by cuuinsi
narmw wnn cJn-ulnr ertxivc <t»wii ih« ini'MIr of one Kids nf eaoli bitick, aal
Utcfi biioffii))^ thvMi ino niIm iiil't lu-ninili.- t'Diiuicl.
We Iiiiv« nlrenily mttl lii 4171 ihnt iho bltiu M>l<iring n»ll«>r, RodiuM
Bulitltiii<li;[<>luti.', whw> iiijfrtw) itiln (lip mne i« uxen^-)! by tlw' livrr M i«tI1
M b V iIm! ItKliiry. If thi- iniiiiial )><■ kttkil ut nit «)>|im|>riiii<- iirii« atttr ihv
injedjoii itml iIh- livrr litirHoiii-il atxl prvpiiroil, m«Uoim uf llti- liv«r will, n
sutxvKifii I •jx-cliiti-iiji. n>vptil a c\-(v nclnork of blue tJiin linm Irnvrnliic
the wh'ili- or fiii-h "f lliv lolKiIcfl, The merit» of lli« D«t«<>rk nrc df nhmit
the widlti of n hi'iuitic cell : nwl upua exiuniiuUion U nrill br foiiml thm ik*
etiifitjr riiiniilc li'ik«, jiiot ^pukeu of lu seeii in the HMrtioiu of « liver profMr*^
in the unliiinrv nny. nra ik><>' Iille>) with lh« bhie pi);n>enl; tlic miiiiiti' cnnal
ftf M'hii-li i-iic-ti bul« te u section is n part of a network of niinuto.
|»a»iii^ lii-(wt«D the cells in varioiiH ditwiionb all ov«r the lobule, i
»my hi- inii-nl to the Mnv t-f thi- lohnle, ami ut VHrious iMiDla of Lba n
the hliir lin« between iliv liei>aii<- oviln will l>e »een lo be oMitindolU, '■
Ikrg^r •ttianlity «if llie Miae blin- miih'niil occu|>riuit the lniDe«i of ou« ol' tiir
tuintilc hjle-iluda m it atHiUi uii itic niniyin of tti« lobule. TItese miiinU'
cnniiU Nii'. Iliervfore, ciintinuoUM nllh the bile-ttiieU; th«y are the tcrminif
liaiiiK of ihi< bile-<luna within the )»l>ul<v. nod imlM^ mvi ooly may thry U
injiK-iol •liirini; life with :-H)iitni uttidiinili^iliili-, IhiI injci'tiuo maivrini niny,
though with <lilfii-ulty, Ih- drivi'n inti> th<<ni biu-kwanl nion;; llie bilu-ducu.
TIh'v aiv s|H>kMi tiC iM hilr ciiiilt'trif* ; tbv naini-, )>i-rb)ip<. is put a ittf
dmirnbJe i>i)i*. btit it ha* brrn eciienilly ndii|>iiHl.
We Miid juKt WW thkt ouch he|iiitic wll toiichwl u MwMlvnxel by « l«Bl
(llie of ilii viirAuHii. wv nmy now aild ihiit eiicb hi.-|iiilic cell ban at leMt am
Bide, aitd ^■ni-tnlly mora ihnii one aide, gro>iTed to fonn a Uile Mpillwy.
■fiiire eiicb nidv thus gruuve<l is in coaliKt wilb ihe corrMpowlin); aloo of*
neiKhlMinng irll. it <-iiiinot run alon^ide a bloiHtveswI. Hence, betwvOR ■
bile i-ii|>i]liiry and a bloodvewel BOme umuuni of eel! sulntaniv in nlwnn
inter|Hu>it. The relative jxintioa of the bile cnpillarics itnd l>I<MidvnMli
may ho illiislraleil by takinv a cube and Ainverlin^' it into u (loly^nu by
Iwvrllint; diiuu the an|;leit of tlie itidea. leaviuj; in ilie Knil instance tiinw in
tbe npiier and loner fucot untou^^bed. The bloodveaoeb mny then lie etiif
•IdMxsl as niuninj; <luwu the bev«lted edge*, while bile i'ai>tllarim run aUng
the middle lints «f the *idM. Two vuch cutm placed end tu ind rmeIii
rc|tment a thiu •mall iidet of celU in one of the smaller sbortcr radial
omwIhii of the VAM-ular network: ni>d tlwo the anfl« of the upper and
h>wcr fitce of the ctnjuiiim) mbe« woul<l have ubo tn lie berdleil for iho
craai-barn of ihu neiaork. Frocjiiontly, no we Imto aaid, Uie cells lie lw<>
abreaal in a rnenb of the vawular neiwork; then, (if OMirae, in the model
the ancles or the iiurf»cc« in contnct wixild not bnvc lo be bevtdle<l, nince im
bluoilvtOMk run between tlicm. If la^wral ituch bevollol niVirt wcrw huill
at) into ft model, it «»tild be Bcen that the ncttwork of bil iiiillimw runs
nhw^' ttw- niiildie of the surfacw betwi-en itw bluodvi-^-l". forming n.wUI
iioiiilii wlMrti- wlls are in muilact with each other hy ihcir Kiirfiicfu. aod
Wvin;* «oiiie amount of cell substaiit-e between the l>ile c«|iillary and tba
blo<Nlv<i>«i-U. Tliid at leatl tuay he taken ne (he typical arran^'nu-iit. when
the network of liile i-apillariea ia moel roniptex. Itni ninny pclU have thi*
lumen of a title eaptllnry on une side only ; and oivasionally a Ink- cafullury
if H<en ill section ul the (Mint of cnDvermenee of three «elU nft«r the fa#Uian
of nil ordinary nlwulua.
tVtieii a lnl« iliitt Hliiirn »ii tho nmr^ii of a li>bul« lli« luineu. m «ra h«v«
jireviodsly mM. wci-nut wiiMi'iily to ■Niitifl li> iiii end. The flattrtipd cells
iiiiiiiit iht (lurlule nr ((-rniinnl iMirtion of tl)i< duct »ii<Menly L'lmn^'C iiilD
lantt^ h«|iiUic Ci^)l», iiiatviiiMl ivlU at thv li>liidc. ultii-h n(i)H-i)i' l» He mm-
pleti^ly ill miiliuil wttli mch ollii-r and to liloi-k u[> ititt diii-tuk'. ]tul nliui;;
iIh? Htilm 111* iIm-h! iiiiiri;iiiiit rvlls as nf all llif <'lhcr i'l-11ii (if tlit' I<il)ii1« run
bilt ciijiiiliirif^. mid ihew are coDtlnumie on ihr i>pi' »ide with tin- luiiii-ii of
itii- durtul«. tind on lh« otlier linnd wiih ihe ndwnrli of iW hil<- oa|>illflrt^
iravi-niri^ iJie lubide. In ihc ducdile ilwtf the liimcii n »)llt:h^ i-yliudricul,
and nf wRiv HKc, it suddeolv divides into much umaDcr ]«i>iNi)rn<. ami llie
C«ll» lining; three tiriialler braiicltin]; iinuMKra are uo longer frimply t-)i it helium
eelli" linitiK a diirt. but i-»nt^lex hejMiic cella.
It wotild np{iear tl>en l)ial ufl«r nil the beimtic cells nra rmlly ecllti lining
tW Irrminsl secntting portiotiit of the dticl, liniiitf we might nlmiH>t Miy lliv
alvi^di, but oMiii;t on thu oiii! band to the ilistribution »f bliHidvciwI*. w>
(UHiri-Nt frnm that which obtains in the ah'enli of otiter glaiidti. un<l <in thr
olbvr hand to nioilitimlionj of the heiMilic celtn. due to their being rngaj-id
in other ruiirlion* (liun that of >fcreliu){ bllc, th« relations of the cl-IIs to [hi-
lumian of ibualvmdi li jivculiar.
Sn-iiMc or Litxa of Faui. ILixuliiv.i
TW ilpi;* hkum lb* iiihiiUr •inirinr* »r ihu Itivr. At •■ ■ liihule h trtn Ib Iniinmr. ol h In
lonflrmniinl •Ritisi; '. Imncn i.t lobiilr; Ihv Ur« HMItMI Kfa olnlar »o«.kiiil IWniUiBhuii nti
InnMtoMri't imacM iiuiut* ; lb«uq|«riDn*«H(ibM)'OD<ii|ilc>lli]r tl;ai«<n.
S 488. Ill tl>e lower anlmnla lb« i>lnieltir« uf the limr in vimfdiT, and a
bm-f 'liiwriiHtiHi of tlH- frog'ji Hvit may (lerbiipt a«-i»t tonanl tht: oumjirc-
h4-iMon of ihi- nature of tla- inamniiilian liv<T. Tim Hvtr of a frof: lu wen
ill a •i-^'tioti apiiean l« 1>C aiadc up of a niiintHT of Inbulcii. which ntpi'alcdiy
n^A tinK bmiirh but ahu) nnii«torii<w<- I'Kig. Hi-'l). and iimmn; which run tho
ca|itllart- bhioilvcwi-ls uniting; tbe htnnchts of lh« [Hirfal nilh thos- of the
be|uUic vein. Thwru ia no very obvious ilivisiun into lobules ; indeed, la a*
^
fi72
THK HETABOLIC ]>ROCES9BS OF TUK HOUV.
6i.-c-lu>n uf xninll iiro« ihv luliultv appear to nui irrcgiilitrly : iinviTttiflni
tWy Itavu n ilrfinicv arrnngdniiutit nround tbu limmJMV of iIm- licpHLli' vein.
Binh !»iii;iiiiiliu3l ixttA tranavene soctions of ouo of ibMc tubtilibi nlitiw tltU
it i« limxi «-!tli largt^ ircdgo-Hbapetl oelU. leaving a very narn>w. nlmiMt litmir
but still (li«(inct luitieD. AixhidH th« tubule is diamweil n nelwurlc nf irapil-
larws, an<l, ns in the alveolui of an nrdinarr glnnd, tbe bloodvcmol is wpa-
mtod from the lumeo of the tubule by the thicknees of an euiire cell. Eadi
coll pomaMS a n»uD<I«() nucleus which I'kh Id the outer part of ib« cell DBBtvr
to tbe bloodvewel than to ibe lurae'n ; and we mar mention here, though w«
aball return to the ixiinl later no, that th« («li tiuhstanee <t>niains a iiuoiber
of gmnules which :ii>e sDmelimea scattered thruui;h<>tU tliv cell and sMneUnM*
Aggregated near tbe lumen. The lve|Nitic cell of Uie fr<>)c rej«atd in fact tbe
main cbaravtere <>f tbe »ecretini; <!ell of an ordinarv Kland, of a iKini-n-aitc
wll for example. The tubules, nwrwuer. whi-n tniee<i are found Ut cud iu
ducts, the (UKireltD;;) hojiatic cAU xuddi'iily chiuiiipn); to onlnoal aud tlmi to
ooluoinar (condintinu;.) c-1Ih, which in thi- larger iliieu bear cjlia. In o[h«r
words, tbe be|Milie lul>uU-9 <>f the frn^ un,- alviiili, diDvring from tlw' alveoli
of an ordinary utaml, in that tltey rV[)«at<iUy aDa>l<>m(MC lu' well a> bmiicb,
nnd in dial lh«lum«n w very narrow and, vince it al«u braucliua aud ansMo-
ouweti, furmH a iiotwork of lin« iwimgM.
From a liver ttuch a> (hat of the frog tbe change to the arrang»m«nt of
tbe mammalian liver is une of degree only. The branching and aiuulorDoenBg
of tho inbulfti is still mufe frequent and complete and the lamina of the
tulmlcH utilt tuirrower, so much m that each cell, ft> it were.takcs |inrt in eer-
eral mbuW, nnd tbe network of the lamina or bile capillaries is so oloee set
that lh(! mf«itv« are of about the aanic vidth as tbe hcjintic cells. The blood-
TBWcb', moreover, are more abundant, and by the estabtiBbiuent of an arrange-
nent whereby inter-lubular i' portal) veinlel« send captllariw lo i.'ou\'er)[e
ladially to an intra-lobuliir I'hepaiic) veinlet, the bepatie eu1»tanee, instead
of as ID the froff beinjj ilitlribuled more or leu uniformly, a dividod into a
imiplKtr of small areas, the hejwiic lobulee,
$ M3. Ooncemio); tlie iiervee uf tbe liver we »haU aay what there is to be
mid when we come to cumnder tbe action uf tbe ucrvoua ty»leta o» tbv
bepatie metaUilie pritcvs»t«.
With lymi>halii.« the liver la w«ll provid^il. Within tlwt lobule lymph-
spBC4« exist betwtvn the wallH uf the vascular itelwurk and the imt«r margin*
of tin* bupatii- r^rlln. and at the Hn'u nifercnci^ of tli« lnhtile thcMO SpaOM oMil
into definite lyuiphatiir vowel* which run in the conntvtivu liwae Mparatin^
Iho h^mlcH and furming the Itcginning* of (ilimon'fi cafMulc. The lymplinlic
vcMttls lying near the nn|>«r surface of the liver find (heir way along the
linincute ul the liver to (lie tlioiaric lymphalii«, thwc ooniiog from tbe right
■KM iMUMniig f ) the right Ivmphatic trunk : all the rest of the lyntphaticB pass
onl along tnc porta) canal and fall into the abdominal thoracic tluoL
From (be details given nhnve ne may infer (hat ihc liver is in part an
u^linary secreting gland. The hcpulic cells living on tbe blond brought In
ihcm manulacture bile, which they diKchsr),'e into the nnri-ow luniina of tlw
minute bile capillaries, from nhcnee it llr>ws outside the lobule alon;; tbe more
open paawgcs of the bile-ducts. But the blood -supply is not only out of all
r portion to the demands ot mere becrelorr work, but also is peculiar in sn
that the blood roaches the liver laden with many of the products of dijien-
(ioii. This would lead us to infer that tite hepatic cells are, as we have
already suggeeted, al»i lar^rly enicn^l in Htthdrawin^c nulMaucc* from the
portal blood, not for tbe |>ur|ioie of Bimjdy furtuing bile, but in ordrr that
other subatunces, or the same subtttanoes more or lets altered should be ailded
or 01.
678
I
I
I
I
I
I
to th^ liln^d III' the tiepaiic v^iii hikI >»• iliMnliutiHl (lir-iii^lmtit (lie bcMlr Tor
tli« bulj-'s Miv. Aini we hiivc exjieriuiciUiil cviil'ini' thiit eiich n work is
curried oa.
Tub HufnmY or Glycooxn.
S 4M, If the livor »( n on-ll-feil miininl be irmovol inini«'liat«l^ aAcr
dpnlh, rupiilly divi<M iiiin «n>nll piccm, tlimwii into builiDj; vriii«r, rubbed
lip aDd builfd, ■ deciclioii tniiy bo obuinvd which nAcr c-Arvltil n«utraliia-
tion iumI liltnilian will be toli^rably I'rra I'roin pmleid runlter. Such a decoo*
ttoD is r«mnrkJibly opnlewvnl. mitky in farl in appearance, mueli more »o
than a similar decoction froin niuscto or other tisEue, aitd rcniaioit oimlMoent
even after repeated filtmtioD. Treated with iiHtine, the aututioii turn* a
br»wwiali-re<i. poH-wiDe red color, not unlike thnl given by dextrine wlii-n
iodine io added ; tlie color diMppeare on irarniini;. but reupjieur^ I'n ■.■noliu;;
iircividisl that not too much proteid matter hiu been lelV in th« Kotutitin.
Fri'RiH with Feblinjc's Huid «r other teatA fur mifpr. the ttolution ix fouiu)
to coiKaiti a ^inall and variable, but only a siuall, (piantity »J' aiigar.
If the volution be ex{)<>-ei|, pret^rablv in th« warm, to the action ofiwliva
orof aonwotlwrainylolvliefi-rtiient.or lie Ixiilcd with dilute add, the Opalea-
cenoo dimppean ; and lii* now clear tmnnparcnt «)!ulii«i giv« no longer the
port-wiiH) n«cLion wttb tndine. T<»tird, inoreovi-r, with Kehliiig'tt Huiil or by
other nimnt it in now found to contain a considemblc i|uantily of eugar.
If alcohol br additl to tht- «[ial(-;!<-<-iit onliition until the mixture coniaine
tW per cent, of the alcohol (previooo t-(inc4-nl ration by evaporation being
desirable) a white nnwrphoutt pn>cipitale is thrown duwn. This precipitate,
removed hv filtmtion, boiled with an alcoholic solution of potash in which
it ia iuaoluble, but which dimolven and destroys any prutei<b which may hv
prwent, treated with ether to remove fatly impurities, and washed niih
alcohol may be obtained in a pure condition. It then appears aa a white
anior|>hoii« piiwder, fairly soluble in natcr, but alwavs givinf^ ri^ to a milky
opal«-«c<'nt Aoliilion unless an excov of niknii be pn?s«nl, in which <-u»i> the
opnicweuce may be slight or absent.
The opaletceol solution of ihii purilicd nintcrtal gives a port-wine reartion
with iixUne. but no reaction whatever with Pehtinp'ti fluid or thi- oilier AUgnr
iwU. Trmied with an amylolytic ferment or l>i>iled ^1ith dilute acid, the
Botutioii, like the raw decoction of liver, loses its opalescence and tta port-
wine r««cti'>n with iodine but now gives abundant evidence of the preacnoo
of (Oigar, dextroM, if boiling with acid baa beeji emphiyed, mallg»e clii«fly,
if an amylolytic frnnent hiu lM^en uned. If (jiiantitative deterininutionil DO
employe)) it will be fiiund that the amount of nu^ar uhlaiiied in pnipnrtion-
ate to tlic amount of (he while (Hiwdi'r acted U|n>ii ; in other nord* Uie MUh-
utance forming an opalc«c<.>nl solution i.* converted into »ugur, the nolution of
which 18 clear, Obviously the nuhatance i« a body allied to the ■Iiifch : and
thia ia confirmed by its cleincntnry c«mj>o«ilion. which in found to bi- C,ll„Oj
or aoine multinle of thU.
Hence thi» nody is called qfyeo'jfH. And it Is nhvlouti I'nui what has been
sialyl atxive that the liver ol n well-fed animal at the moment of deoith con-
Utin.i a coni<i<lerable (|Uantity of givi'ogrn either in a free stale or in Kich a
condition that it is »et free by subjc'tiiig the liver to the action of hoitiug
waUrr. We may add that il occunt in tlte liver in the hepatic colU, tor the
reaction of a jMtrl-wine color given under certain conditions by the hepatic
ceils (S ■!■''") i» due to the preseme of glycogen iu them.
S4M. If the liver, inntead of being treated immediately upon the deatli
of tliv animal, is allowed to remain iu the body of ibe dead animal for aer-
674
TUE UETABOLIC I'RUCKAitK:* OK THE ilUI>V
■U lUIB
rntl licMirx, 4V|H'ciitlly iii » wnnn pinoe. buforv n decoction i* iiinil<^ of il,
iJwicU<iii will bir r»iiu<l to hnvu litllc or tio himIoociim, to )»• iiiiitr or nnirlv
i|uiU' cit»T, lA giv« liulu or ou ]M>rt-n'in« n-iu-tidii nitli todiixr, hut In i-ontaiu
It ver}' o(>ii«iilvrubl« qumicitr uftugnr. A* «<■ Imve Miid ulMJvr, \he dt^itctinii
even of it livi-r tdki'ii :iiini<^lintfly itfttr <K-Ath p-nerallj conUtina aotne little
sugar, itiiH the •iiiiintily <it' xiigar in tli<.- liv^T ii|>])viir>. ii» a rule, to
Mcitdily HlWr il<'iit)i. Ihe kidihidi of glycogvo tjimi uniting m th« w
The nipi<iity of the <liniiiiulioD of glycogen itn<l thv rate of inervMe of
vary much uixler vnritHii! ciroinutitnt'es. Moronvcr. lite dcrreuae of the oiur
and theiiicren«eof lbe<iihi.Tnr«not ulniiy» strictly proporii oh al : nnd, iiidtivtd,
some obeervvre bav« insisted ihul tbrrr is mi rclatioD between the tw-u pn»-
cewes. N evert he Ices, the hnwd fuct renittiiie that if the liver uf tliv miim
well'fed anininl be divided into two halves as mod as possible aAer dMtb,
uud nue liulf thrown into boiling water immediately, tvhile the otber half it
li-ll exposed to some little itarnith ior several, suy twenty-four buMrv, iJk
de(N>ctii>ii of the first linlf nill contain much >>lyc»}^n nnd little atijcar, while
(liiit of the wcond half will ooniaio little )(lycii-„'en and luui-h adj^nr ; and thia
furl may be taken, until (lie ountrary is pruved, to ahuw that the ftly
pruieiil in the liver at the moment ni' death is jjrailunlly after death by
neliini »r other omvertetl into titifrar.
The Hi-lioii i^ that of some agency whiMe activity is deatroyei] by the lem
iwraturc of lioilinfc water; 1m!0(,v the din-dinn-t re|)eatedly ^iveii iiliitve [u
tbrun' the liver into boiling witt^'r. Thi:i nalnmUy siiggnt* tim prcMrnce in
the livi-r <>f an nniylotylic tVrmenl. jliil not oaly hiivc attompis to Uutaie
fruai tl)<^ liver mi iimylolyti-.' fcriiK-iit fiiiUij. in llie baiiiiK of miwt ubaervvn
at l<.-4»l. btil the <-xm!t natiirv i>f the sugar which npjxMrK idiows (bm the
cbaiign ii> not alI<Tl<.«l bv nn ordinary aniylolytic ferment. In the <vwuf
thu iiiiiylolylie fcnuent of' saliva, luuiereatic juicv, intawtiital juice. Mid indMcl
i>f all oth«r amylolytic aniiDuI fluids, the sugar into whieh siareh ur glyou-
geii in cunviTteii it maltiMe. Nun-, Ihe sugHr which appears in the liv^u- aAar
death is dextrose, identical, as far at least as ean at pmeat be made mil,
with ordinary dextrciM. We are led, thervfoixt. to infer thai tlie i-haugr of
glyoogeii into sufpir, irhicb appears to go ou afler death b ciirrii'«l uoi by
•one aetioD of the liver, pntiahly of the hepatic cell itself, which is dime
away niih by a temperature uf 100' C, but which is nut the anion of a
ferment capable of beiug isolated.
it4S6. We have used above Uie phrase" well-fetl" animal because the
aniouut of K'vciigeiu prewmt in the liver uf au animal at any <in« time a
verv variable, and Mjiedalty dependeut on the amount and nature of the
f<>Ml previously taken. When all food i.< withheld fntrii an imimiil Ihe glyoo-
geu in the livcr diminishes, rajiidlv at finii, hut morv nhiwly iiftvmard. Kvea
after oonH' davi' ntarviiliou a itmall i^uantity ii> fr(!<(uetiily itill found ; but in
rabbits, at all events. lh« nbolv may eventually dii>iipiiejir.
If an animal, after having hr«n Harvitnl until its liver mar he assumed lo
lie ftv«, or alinixtt free, fmm Klyi'ogen. !k' fed on a diet rich m osrbohydratw
or on one conaimin;!!; esclu)ii\'t-ly of cJirbohyd rates, the liver will in a slmri ticnc
be found locontaiiiavervlargeiiuantity of glycogen. Obviously the prnviia
»f earbiih yd rates in f'Hni lead» to an accumulalion of glyco^^u in tlie liver;
and thio is true both of slitrch and of dextrin and of the various tiirins of
sugar, ennc. grape, and milk sugar. The ellect may be (jitile a rapid one. fur
iclyoigen ban lin'ii founil in the liver in cunniderable ijuautity within a ten
Itvurs after the introduction of sugar ioio the alimentary cauat of ft »lurviug
nninuil.
If an animal similnily Ktarveil be fed on an exclusively meal diet a ce^
lain amount of glycogen is I'onud in the livdr. This appeani I" \w wjiecJnUy
J
Tilt: IIISTOHY OK (iLYCUGEX.
6-S
Ihe cMae «il)i tli« lUif^ (pratMibly uhli nlher cni'iiivuroufl aDinwU aUo\ nitil in
e«irli«r uorks od IW Bubje«t ()i« cniiMaDt [m^cure »f j^lycogeD iti ilto liven of
ckifM ied on meat iraa rejfardod a§ an ini|H>rtant iiidicHtion of tlie t'lirniniion
vtiibin t)i« IxkIv n{ non-niini^eniKiH fW>m DitroK^nous maieriol. fiiii in ilic tirxt
l^aoe, the quantity ol'ytvmgeu thus sloreii up in tbe Itvi>riu the roiilt af a
meat diet ia much leas liian that which follciwa ufinn a cnrhu-hrilnitv ditrl ;
and in the lecond place, unliiian' meat, especiatty IxinH-'fliuh on' whii-h <l<>u;8
ill Mich ex peri men t« are usually t«<I, ctintaiii* in i(»eH\§ •>:>)» cvriiiiii uniKiint
rilber of Klyoogtui or anmv liirm of AU){ar. Moreover, uli«D nnimali are Ted
■Kit on meat, but OH puntieil |>ni[«i«), such an tihrln, cHAeiil, <ir albumin, the
quanlilv af gWci'gfu in tbe liver be<t>iu« ittill ainulli-r, though, acvoriliiiK lu
ttw*i itiwrven, reinuiiiinjE greaK'r than during star%-ati(in. We may iiifiT,
ihdreiure, that jttirl of tht- glyc»ireii which Hii[)ean> in the liver after a meat
diet i« really due lo carlKihvdtMt" iiiaivriiil!' pri-^cnt in iho m<-iit, I'art. how-
ever, would apiHMr U> be llie roult i>f thr actiiid pn>t«-id fniiil ; and we have
Mmilar evii|<-ik-« thai g'taliu uikcn lu loml lends (o ibe formaiiuii of some
jtlyco^'en in the liver. But in ilii« rt^iicct tln^c nitrogenous subatancee fall
lar t>hort of carhohydrati- malrrinl,
With regard to futi^ all ubm^rvcrs are nerved that thcce lead to on accu-
mulation of glyooRon in the liver; an aiiioMl frA on an exclusively fatty
diet haa uo more i^ycogeit in ita liver than a starving animal.
Hence of the thie« great claaes of f-xHl-etuffs the •urb'^hydmlex Hand out
prominently aa the »ubManct« vrbich taken »» ro<>d Iciid lo an aocuniulatatina
of glycogen in the liver. We may rcinnrk that the greatest accumulati'iii
of glycogen ia efleoled not by a jnire carbohydrate din, but by a mixe«l diet
rich m csrlxdiydrateB. A ijiianiity of carbohydrate mixed with a certain
|>n>]K)rtio» of proteid givei riae to a larj^r amount of glycogen in the liver
than tlie tame quantity nf carbohydrate ;;iveii by ilaelf; niid it in [Hiwrihje
that the presence of an afipropriate quantity of (at still furtlier iiMii'ti' the
actruinulation. But this result prttbaMy depends, in |iart at tea;>t, on the
fact that, though lUflerences may be met nith in ditTereiit animaln, a mixture
of the several <-laM>eti of f>HK)'iitiifl's is more readilv diitealrd, rrsulling in
more nutritive jimti-rial ImHiik thronii ujhiq the blotKl. than is a mnil consist-
ing exchisivdy of <in<- kind of fo<id-ntul1' alone.
As far HI wtr knon- at pnw^nt the glyeo)j^-ii which tbu^i appenn in the liver
as tbe n-«ntt "f lii-ding i-ilbcr with any of th« vatiouH forni.-> nf ciirbii hydrates
or with |>rotei«l». or nilh Mber nulutanci-x, i" of the same kind and presents
the same cluiracten> ; at h-iiPt «<■ have no evi4lein* to the cnnirary.
Tbe eloriiig-up of glycngi-n in the liv<-r in aUu lullucuced by otber circtim-
stuueen than the taking of focid. For iuntancr. in the frog an increase of
glycogen takes place during the wintpr nuinth^ In lint suiiinier inonihs lite
liver of a frog will Iw found to conlnin very little glyco}.^-n <Fig. 164. C ,
unlees the uniinal has litvn iinvisually well fed ; whertMU a liver examined in
midwinter i Fig. 1K4. A ) will hi- found to cotilain a (.xiiisiderable quaniiiy,
even though no fiKiil has been taken for months. In such a case the maieriii)
for the fiirnialiou of the glyeogeti in the liver miiaC have been furiiivlitd by
aome pwrt of tlw body of Die frog, and could n«t, as may Iw the cast- wlien
a meal leads immediately to an increase of givcogen, be supplieil directly
from tlie fnud. It seciiis as if in the summer the frog lives up to its capital
of hepatic glycogen, sjwnding it as fast almost as it is made, but ihat during
thi' wtiitiT a iguaniiiy is funded to provide for tbe demands of the late winter
iiud imrly spring.
This winter storage of hepntic glycogen Ja the frog seems closely (Ic|wii*
dent '>n temf-crBlure. If a winter frog, whose liver is prcvumably mure or
less hjAiU'd with (flycttgen. he expoeed for some time lo a leiupimiure of '20*
&7ti
THr. MRTAIItil.U' PRKCESSKS or TUB HODV.
itr n little Itit'lKr, ihi* lii'er will alWrwiinl [»■ fnmul in mntnin litcto nr im
glycofcen (Fiv. KU. It); and ci>iivtn>rty, if' it ■iiiiiiiiit frn); Iw pX[Hi*rfi m
iiiiliiuely oifliL glv<-up-ti, thdUfcb not in nny gn»l t(aBnltly, hcij^iia lu be
i>U>r<Ml ii[i ill the tiri-r.
rn. IM
'^^Bk.
B
V
»f\
t^
>V*^'«
iir*
c
Thm« tnutt» «r TNB umTK cnu M TKK PiHa, j.nmiuT.i
A.nll>nrtilnNl)w«uii, Ttekmi (nm • toK dsrlac wIMs- TlH«U>krwliu(*.ii»d|iKKi;Mlcninuln
KNMiMcl ■niiiiiit ilic Iliidcii, tlM) homi !(*■>''<"■• onlortODW uf llwnllt tabic I'Wir ■ul»|ii— il rf
Sljroorni. «rluc)\ uu I'nsviii lu I'oiul-knlilc >buaikiir*. Tbu ODlcr cum OMUUinl enuMfOW •«
Riobulo. •lumn u ilHik >i«iii; lull, urttnled la Uitlcil. IbuclU Khilnln Tw; ntxtt K, n:U>|U>
IB tljFtdKrii. Takon mno n ulslcr n\« which hml b(ca >:«|4 nt SI' <.'. fcr Mi d*}i. n* oetli CM-
lBlnirw> lliitv (Ijrvomu aail Ite i>oMld tnnnlf* an dUfcnHl UtRnwhout ihcpelL la t iilmmt
ftocucllM ui|xvipl<lt ihnnilI>>tauhlpK«ciita*MrilM(tarapi|iCBmii«, o.iiarNdMlli. luat
bOMkniiuawr rni|iUI«c«lMiKftri. Tlia MUtB»ranallan4alni»i tPM> flvn rlfcocMi. TiMfM
tdiltnnnlMwcilUttnedltiroa^ioiitttaoo*!]. AtlihvanictiDcntHrruhtnSciMdliil p*r«(iu.anto
MM, and wc Oak n to tb* •■•». or oiwtlr to tbv wmc. xailc,
(407. Uefore we alU^aifit (o diacusB further bow food aiiil otlivr cinium-
■tancea thiia nlfcci ilic utyo'ifiti iu (he liver, it will be det>irublL' to take up
the matter whidi ve lirit on tuie Bide iti $ 450, vix., iIm- consitlvntioii of tM
liialAluvical oliHUjint oceiirrinR in the be))atlc ctllii under ^'arioat c<>»dition&
It will lie ei^nvriiwnt ft bvgin willi tlie cells of tbd mora distinctljr lubulu-
gland of lti<! tntit-
In a fnig whieli bM not b«cn aubierlcH to any imrinl trmtnioDt the cell
MubMauce of the hqiniiu cell {ef. Fig. ItH. A)'will ^nenilly be finind in
r»ntain linlg^d in itnelf throes kiiid* of malerial. the pivfnio.- of which, if not
dim-ity rerognixablf in the rmli ci-ll, mny l>« demonstiKlcd Ur the uw of
varimu mgrittK. In the firft |)luci.\ oil globuin of variuhle liu and in
varinhlr anuHint ar* •CBtl«rrd timitighout the t^ll; »nniciini«e, as we bavt
alnrady mid, these arc extremely nliiindiiDt ; but tlien- is otherwise nothing
vrry Kitrniit ubout tlntt^ I'm globules in the hepatic cell to demand an^ dn-
Oimion coneerninf; ihcm apart I'loiii the general difcuMtoti on the formatMii
of fnl, into whieh we shall eutcr later ou.
In the wt-oiid plaoe. a uumber of sBiaU discfelr gnniilt^ may be Keo
IndgH in ihc cell «ul)«t)iii<v. llten appear to be tit' a pruteid nature anil
Y nr iiLVC<
l^niool nliiiiiilaiit on tlw iunvr niili; ol'llic vdl QMr ibe lutui;i) of
t k)r bjlr |<aw*p'. TIh- |in-M'nc(i of th«'i> griiiiuUw in clwtdy d«pendeiit oa tlic
imlT "l" lliL- ■ligwtiir ('mcmMw. Thty <liiiiitiL<)i nlicii ili((™iioii t* g<"ii>^
lUiif ■ccintiuUte Hpiiri nrirrwnrfl. I'lilliiig soidi- oTtiin ili-tJiilx, wu iiihv
IT thai thcw grnmilcf behave vrry miuh like ihr grniiii W in itn albumiiiMis
Jivary c«U. a iMDoimtie <.-cll, or a el)icr^iisiri<^' ovll ; iind nu ms)' probnlily
~ij oMidude thai ihey, like the gnuiulec in lbt«(< celli^ nro tti Rume wiiy
nrncd in the tommtiiii uf tbe aecrvtion : dial », in their cnev, bil«.
Ill iIm> tliiitl |ilac«, the cell txtniaina more. »j)ecialh' in ile outer paris
^■tMrrr itie blnodveaeel, anay from Lbe lumen ul* the bile pawjige, m variable
-^■.jUBiitiiv of luaicrial which tlifito from the urdioary cell lubetjiuee in beiDf^
^ysliiM- mhI refrvetiro, ami bettoeslany looking, aod in fiaining port-wiue
red with iudine, iaslcad of browuiin-^Ilow, an dues ordinary cell subHtauoe.
Till* aiBtcriftl it, though with aoine bttle diHieully, eoluble in water, and by
tU* aiwiiT nuiy be diiMolvdl uut froui the c«ll. When this is dune the plaoc*
wUcfa h occupied apjienr tut vaouolea <>r guy* of various si^ee limited by bnn
af Uiecvll tulMUnce, Hhicli thus Ukw on the form of a netnurk, the nMabos
of which ani wid«r n»d more cuDHpiLtiotis in the outer part of the cell, in
tdich tliv hyaliuc inat«ri«l iras prcvioualy inual abumlimi. lu the inner
fan of ihc nil «fa«rc the hyaliiK' niiili-riul wu •euity the cell riutwtance iii
■ora dflMBt Uld ovta in the <>ult-r imrt u >liell of mom ilciuc, laa reticiitnle
nrll sittntnnco nflbrdv a dcfinitr otitiiiK^ to the cell. There can be no doubt
liiMl thin hyaline material ia cithrmctiia] glycogen, such umaytx; vxtr>rle<i
from lbe liver, or, lu mvau more probable Irom its ilelicient liobibility, gly-
OOgga ill some more or lees loow oombiniitjon with wmi.' other body, n cnrn-
biomliou, hoirever. of aiicb a kind that the iodiiM mwtion makc« i(>clf fell,
{ 4A8. The above may be taken as n guieial dncrijitioii of a cell in an
ordituu? mtiditiou. The question now nunea tiefore us, What cbao)^ are
bioui^hl alxMit by various foods or by the nbsenec of food?
If a frug be largely fed on a diec containing A tarR« <iuaiititics of carbo-
liydrmtAr, the liver will be fuund rich iu tcl.vogen, and tbe cells nill pn»«nl
tM fullowini; ebiinK'tem: The cell \» relatively large (</. Fi);. 1G4, A}, itnd,
mm it mm, twollcn ; the cell Milmtanoe b largely occupied by tbe hyaline
nuibrrial juit *|Hik<-ii of, uA|in'inllv in its outer part:*, ao that iu sectiout prv-
Hunl and muoiitrd in the onlinary iray in wnidi the glyoo}{eu ban betu)
diteolvcd imi. iIk- ^reati-r fxirt «f tbe coll consisia of U loose open uvtwork of
ban of Moinnl cell miImjuiic with wide mesbea; a certain qiumtity uf more
•otitl. gtnerally gmnulnr-ltHiking oil ■ubaionce occnpEes tlie port of the cirll
iwrral t)i« Inmeu. and n thin »hell of erll «ub»tancv forms an envelope for
(bo rest »f the cell. TIk- nucleus is liirge and distinct, but ihouKb chongnt
ia tbe nucleus aceon))<nnyiii|,' chan}^-> in the cell mbxlnnov buve bm-n
dMCribed, tbcy are not sufficiently im[H>rtant to detain us now. When such
aedl is seen in a perfectly frosb state, the hyaline refractive material i which,
t»« Dee<l hardly say, givca a marked reaction with iodine) often hides tbe
Uiiclvus aud the greater part of the cell substanee proner.
If, on tiM ollter liaud, the frog be Inl ou a proteia diet free from oarbn-
Hytlratos — for iastanee, on librin— the liver contains little or no glycogen,
%tKl tbe be|iaiic cclU ore not only much sniuUer, hut present an ap{>eanDi'e
%enr difllnxnt from tbe above (i^'. Fig. t*>4, B i. Little or no bvaliiie material
Sa visible, tJie cclU give little or no port-wine reaction with iodine, but only ilie
UMwl b row u-yel lour proteid reaction, aiul in»|iecitiicns prepared and mounted
tin tbe ontiiuiry way the cell aulwtance npjiears densely granular throughout.
L«tlv, if lbe frwg be »tarvcd, and if to tbe elfecls of stnrvniion there lie
■iUl«d lliose of vxfHHurL- l» a bi;;h tetuuerature (20"), by which, ss ko have
a(«o, tbe hepatic orlU are uiuckedly aifected, the liver is found to be free
87
578
THE METABOLIC PROCESSES OF THE BOBV.
from glycogen, and the hepatic cells tfl be extremely small (cf. Fig. 16-1. C),
only naif the size or even lesa of those of the weii-fed frog, btit othenriae
much like the celU in a frog fed on ])roteid material.
§ 459. In the mammal changes iu the hepatic cells similar to those jtut
described as occurring in the frog have also been observed. When the
animal is fed on n diet rich in carbohydrates, and when, therefore, as we
have seen, the liver abonn<lt in glycogen, the hcpalic cells (Fig. 165) are
larger (so large tluit thev have by some authors been described as com-
pressing the lobular capillaries) and loade<l ivith the the same refractive
hyntine material staining port-wine red with iodine. When this material ii
dissolved out a coarse open network of cell substance is displayed. THk
Flo W:
Yic. ir*.
Sxi.TluN uv Maxxai.ia:! Livek Kkh is <Jlvcii-
r.tN. lI.AM.I.EV.t
llMiilc nfM si-CPiiiu-ii. cljTi)[iCii iiul .11.s*"1vli1
nut.
Siy.TLlIN OF )l.lM)l*r.TAN I.IVEB CONTAIKIini Ln-
TI.E f.lt Nil lilVIOr.KS. it-ANCLEY.}
fi«iiiti' Hoiil sfN.'^'iii^'^u, The L^ranulefl are nM
\n'll jirtsi'iTfil 111 Kime of the coll*.
most marked ]ioint of difference lH>tween the mammalian and frog's hepatic
ce)] under these conditions is that in the former, the liyaline, glycogenic sub-
stance is gathered at first centrally around the nucloua (not more on tbe outer
side, as is tlie case in the frog) unil sprciuU from the centre toward tbe peri-
phery, always leaving on the extreme outside a somewhat thick shell of
cell siibstiinoe, which in hardenctl and j)repared .'iwcimeus may strikingly
stimulate a tbi'kened cell-wall. We may iid<l that in an animal thus led
the whole liver i* very liirge, and, iia it were, swollen ; it is also soft and
tears easily.
In an animal fi'd on proleids alone, fur Instance on tibrin, the liver fre-
(jut'nllv contains ^omc glvci')gen and the licpiitic cells contain a small quantity
of hyaline, ;rlycogcnic materiiil. As in the cninvspimding case in the frog,
the cells are comparatively small, and the cell i^iibslaiice appears finely and
unifornily giiiniilar.
In a starved mammal the liver is small, dense to ilie touch, and tougb;
it <'ontain,-' a trace only of ;^lycoL'cn or none at all ; llie cells (Fig. 166) are
small, as it were shrunken, and the cell substance, which gives nu port-wiae
reaction, or a mere Lniee only, with iodine, is still more finely granular.
S; 460. The mieroscopio apiiearanw's just described show, and indeed
general considerations lead us to the same conclusion, that the processes
taking place in a hei>alic cell ;in- very eoni])lex. In the first place, the con-
stituents of bile are being formcii and dischargeil into the hile passages ader
the fashion of ordinary secreting gland. In the second place, a forma-
tion of glycogen is also taking ])lace, and we shall have prest-ntlj- to consider
briefly the relations of the one process to the other. In the third place, U
is esiMH'ially indicated by the somewhat peculiar ell'ecls on the hepatic cell of
■t cxclilfiively [iroleiO in ualiire. olh«r proceSMS, similar pcrhnp- to tliw
. iriiintidii "f kI.V*."OR^". hut i">t nwultiiig in the ttUarajge of any cnrlv>l)y<lriit«
lUiiU'rial, und denriug poanblir with pnteiii DtibslAncea. aUo take phioe.
Hciicc llie«xacl inlefpretstion of all the ohaoKes whioli may be obierved
l)i-iMii)Cti exc««iliai;fly ditGouK.
tA>avinK the proi-eMes of the first an*) ttiird kind whollf on oq* Bltle for
th« itrwent, and Donrminjr our nttenlioD entirely lu the glycosfeu, it i» obvious
that tbe li«|MLic cell manulaut^irai the glyiv>^en in aonie wkv or other and
hxIgM it ill iK own subatanee for tlic time vrry inuvh in the wav that a
wcrcting oell wanufacturaii nud hMljtei) in iiaelt' for ii tiuin raat«ria) fir tbe
secrvtioD vrhidi it h ubuut to puur forlii. Then- i» thix difliiniice, that in
Umi odo com tiw malarial »f the vpcn'tion. aTtf-r iin'htr^iiin:^, n» wc havn Mtcn,
more or Ian dinngv, !# cmt out into ibv Igm^n of thr hIv<-<)1ii«, whcroai Id
the other CKH' the glrmiD'H, which rnuot iindori;<i citiingo. sinoo it may ba
made to diKiiiiiK«r rapidly from ttu' h<-:|>iUii.' ovlf. u not when chaiigod CAtt
out into the bile pawag** ; it must lhi.-)er»re bo wut back iigiiin to iho bliKid.
461. \Vi- My " mnniiftietiirvi the glyeogni in Dome way or oth<;r," und wu
have DOW to in<|uir« what we know ouiiconiiiig the iinture and the wveral
Hef» of this mtuiufiicture.
We have already teea thiil the presence of ({lycogen in the liver t*
eepevtally iRTored by a carbohvdrntv diet ; and in our studies on digestion
we have seen rvuMn to think thni a very lar^ part at all events of ihv
carl>»hydrato material of a meal i^ absorbed as sugar by the cnpillnries of
the intcatiue and carried as stigar to the liverr in the portal hloud. Hence,
it seems only reasonable to conclude that the jtlyeogeu which makes its
appCNianee iu the liver after an amylaceous meal arUei^ from a dire;:t con-
Ttn>iou of tlte sugar carried to Ihe liver by ilie portal vein, the sugar
bocunting ihrotif;!) s^nie autimi of the hepatic cell aubstanee dehydrated into
' icugeu, or animal starch, as it lia^ been called, the ;>ri)m-«t l>e^in)c a reverse
Uial hv which in the alimenlary caaai starch U hy(l^al<^•l into sn^ar
Ihrtiu^h tite nclii>u of the salivary and inncreatic frriii tnti. V'c^table cells
can iiudoiiiitciliy convert b>>lli starch into sugar and mignr into Mareh ; and
Iheni an: no it priori argnuietits or p'isitive fact-t, which would ]<:ad us to
suppcM tliai tlie activity of animal Mvinjc sulkstance cannot noconijiti^h the
Iflllnr an well as the former of tlieiM changes. Wv arv <|iiiic igii iraiil, ir. is
true, of tbo exact way in whloIi dther ibe liyilrnlton or the dehydratiou Is
edertetl by living subslaiiue; but we are U'lriiiUy ignorant of tW exact way
in whii'h an amylolytio ferment eflr^t^ ilie hydration of starch into sugar,
which it cnrrim out with *<> much apiiarent ease. It is not a grent assumption
tn MipjinM) that the <^>ntinually ciuinginz living snlMtnnoe, which in its
ehangei is continually giving out energy, has the pswer of acting on in<>!c-
ailea of starch or of «U|jar in contact with or even only near lo itwlf, and
»<> of hydrating starch into the sugar or of dehydrating sugar into starch.
Tlic latter process may lie a more difficult one than the formsr, but not one
boyoud tlie power of the living ■iibstnnca. We may fairly luppiss tb.it a
<|Uaiility of sugar in whition preicnt in a vacuole, for instance, ot the hepatic
cell snltBianoe can be, by Mine action of the oellsubstani.-e, converlol into
glyc'igen in a eolid firm, filling up Ihe vacnole. Again, as wa have inci-
dniiatly mentioned, sugar iiijocied into the jugular vein readily give^ rtw to
»ugnr in tbe urine: but a very co[i«iilcrabIe ((uaucily can ha slowly inject«d
iulo the purtal vein without any appeariu^t in the urine. This suggests the
Mm that the liver, so to speak, catches the sugar as tt is passing thruugh
•Vtt hciHitic capillaries niHl at once dehydrates it into glycogen.
Similar eonsiderations may iilso be applied to the onse m«nti<med above of
the npjwnTaiiee of glyct^u iu the hepatic cells of winter (fasting) frogs
Doe
iBr
680
TIIK UKTAHOIJC fROCKSSEH OF THE BUDY.
We hsvo mson tolhiiiW ihnt 8Ue»r inakco ila uppauiincc ni- a product nf the
iueliili'>]iitni uf various liiviicK. Tltf sugitr tlius armae tiiKltiiK its wnv iiilu
blood may be ntde use of at once pltewln^n?, conv«rted B^teedil.v tor iiutaaw
inio carbonic acid «nd so ^i rid uf. Hut we onn tvadiljr )iDattii>e 'tuit uttder
certain circuDuinDot^, ua tor inotaDce when the aclivitieB of the ouitanl w<tR
leneoed by a Ion temperahire, it was Dot so made use of and reniniii«d id
the bloo<l. If so it would in the couree of the ctrctilalion lie ntrrieil to ibc
Liver, and mixbt be at ouce laken up bv tlie ItefHiiic c«lU aud ooiivrried into
Slyeogen ; and theee mijiht be sa active that the blow) waa never at aay
nie allowed to remain loaded willi augar to such nn ext«nt im to pamiit ■
low> tbhmuh tbe urine.
j) 462. f'iH>ii such a view, tlie carbohydrate taken as food would be coa-
vertod iiilo glycogen by tbe agency of the he)Millc coll, without lU any tiiM
tiroomiiijt an lute^al port of Itie living i>ub*tnnc« of the ooll. 8<ich a view
mav be Die Lnie one; but it i» open fi>r ux to look at tiM mattcT in nnotbef
ligfil. We way piiHh ulill fiirtlKT the iiniilog)- Initwoeii the glycogen of the
be|<ntic cell and ihv miiu-nHl with which u erarctiiig evil is Imuled. lu deal-
ing with socrction we eaw reiuons fur regarding mcJi a body aa mucin to be
a pnKiucl of the inetjtboliHm of tbe cell substance of the muooup <.-ell : aixt
we may HQiihtrly regard glycogen, or sugar readily ooDverlible into Klyoi^Mi,
or at least some or other carbohydrate material, as a normal produit «f ih«
metabolism of the hepatic cell. We may ihua conceive of the heiwtie cell*
as being continually engaged in giving rioe to carbohydrate material in the
form either of sugar or of some other body ; aud we may aupfnae that under
oertaio circiinistanc». as in the absence of ado^uate foml, the carbuhyilrau
material thus formed is at once discharjiied into the blood oT the hi'imiic vetu
ibr the general use of the body, but that under other cirvumiianii'^. a* wlien
an amylftceoas meal has been taken, the immediate wants uf the lymooioy
luring covered by tbe carbobydrntee of the meal, the cMrlMihydniti- nmductt
of the hepatic metabolism are stored up as vlycogen. V'nd^r siirli a vie>
the sugar uf the mml i» ii--ed up aomewoere Tu ijie Inxty, and the glycc^cn
to the Ht(ini)ti: of w liicli iu lli<! liver it j^iven lice comf« direct from the hepatic
Wibatance. And a •imilar cxplaiuiliim may bet given of the storitiff-iip itf
glyengeii in the liver undir ouch ctrcum*liuim< lu thuee of the winter tiof
previoiiiOy nn'uiioncd.
We do not pummu at nn-scnt ex {K-ri mental or other evidence of so dov a
kind a» to enable ue to decide dogmatically betvcen llicae two viewii ; we luv
limited lu very general indications. We have seen that prateid fixicl, though
iu tluH re«|>ei-t (ailing far below carbohydrate food, does or may givr< riae to
a certain amount of glycogen in the liver; and gelatin seerati Ui haw tbe
san>e eSecl. Further, in ceriain cases of the disease diabetes, of which wo
aliall liave to speak presently, and which is cbaraeteriitetl by tlie pr>Meuce of
a large amount of BU);ar in the blood, sugar continues to be fonnc*l in larn
(|UBQlity, even when the diet a restricted to ]>ruteid and fatly luuttcrf, ul
cHrtM>h yd rates being excluded. Mow in dinbelva wo have reaanii to believe
that tbe laKe quantity of sumr in tbe bloo<l ia acconipani<.-<I by a lam
deposition or glycogen in the liver, and iwieed in other tifxncs ; lor in ibe
tnt ouea which have been examineil suHiciiiitly KH>n aOcr dcalh, and to
whtdi owing to itie niddennew uC the dutth, there niL> no opportunity KiT
Mored'Up elycrigi'ii to dinaiijii'ar. a very large ouiinlity of clypo^Tn hat lieea
fiHiitd in the liver or iti onic otlirr organs. Henc« the jiTieiiimtena of dia-
betes may be taken a* ithoniiig, in n much ni'ire striking mnnuer than do
any experiments, that protcid material tuben as food may give ri»e to lte|)atio
glycogen. And thi* at lirst sight Meme to alford proof that the liejwtili
Slyoogen is a product of the metabolism of the hepatic cell, the activity <if
le cell being Bliniulaled as it were by the presence of the proteid fiuKl. But
THE HISTOBV OP OLYConKX.
081
I
I
I
ihe |in>»r is not cogent in (he fact of our igiioraiice »f the metabolic chnni^ea
which th« protvid material of fixnl uiiitiTfiiom In tbo Ixxly. An we shall insist
ii|H>» ill more detail later <m, )>r'jt<ri(l iimti-rial in giving riiie to urea thmwi
olf WMuewher« in [be timly a lar)^ <|iiuntitf of » ciirboii-coutiiiniog radick in
•omo coinbiDatinn or other; l)i« proicid fontaiii* liir more vnrboii Utao b
D««de(l to uuiie with the nttrogito li> form urea. We i^hnll ntx that th\*
excess nf airlxm linn n icjidiiicy to apjieitr in the form of Ikt. ))iit wc may
readily mippiim that it might tempomrily us a preliminary proceM or under
certain oirciim»timrc». t«ko on the form of Nifnir. And we may fUrtlicr sup-
poae that tiiii augnr it formed out of the proteid nut in the liver but in some
other tiwue, in muecle for instance. Jtui if ho, hep^uc glyco^^n which u
ihu revult of proteld fiiod, may alter all be formed iu the liver by simple
dehydration of au^-v foond elsewhere, and brought to the liver in the {tortal
bhHid.
Wo cannot, we say, at prcwnl deride Iwtween tli««e two vien-a ; and indeed
it may he thai l>oth views are true, nr ratlier that the true concepliou ein-
briices both view«. It may be that the normal nietaboli«m of the Iiepatlu
oell diiea priKluve a certain amniiiit of carbuhydratc mati'rial ; but if »j the
probability is that the exnci I'-irm in which that carh»hydr«lc apiieant in the
Arst inmance In tbi- la b<i tutor v iif thvot-ll u iK>t that of glycogen, hut of aagar
of »ome kind or oilier. nn<) tliat iho convcr«ion into glycngcJi is a milHiitltary
act for the inirp^Ho of retaining tho carbohydraTi^ iiialcrinl in the gnwp of
th« cell. ]f thi* he (he case, then until it has boon iihown that thert' is
siinietliing peculiar about the augur ihnx produced by the coll itwlf. hy virtue
•d* which it alone can be converted by the celt into t^Iycot^en, we may I'airly
infer that the c«II might alao convert into glyco(>en ei]i>ar pacing into the
iiitonticcs of the cell substance from the portal capillariee.
S M8. We may now turn to another <)ue8tioii, the answer of which is in a
measure dependent on the one which we have jii»t diauussed. Whnt is the
u*e and purpose of this hepatic glyivi^.^en? What ultimately becomes of the
glycogen thus f»r a while .iton-il up in the liver?
One view which Inw lieen piil forward u as follows: We have evidence,
•a we shall preaeiitty U-ani, that u gn-at deal of thu fat of the body is not
taken u such in tlni t'oui), but in constructed anew in the boily out of other
•ubstuoot. Etoth carhiihydniKs and pnitci<U, taken In exceas or untter
GcrUdn Hrciinutancr*. lea^ to an accumulation of fat ; and ve have reason
to heltvve that cnrliixlvdmUv un the one hand and the carbon-holding por-
tions of various protcids on the i>tlier, may hy some proccM or other be oon-
rerted into fat. And it bus been sugetsted llint the glyuo^n in the liver is
a phitM of a constructive fatty melaMiiam, that it it material on its way to
bvcoatefat.
The pKsitive evideiiL-e in favor of this view is very scsniv : it is almost
limited to the facts that fat, sometimes in very largo qiiantily, 14 found in lUo
hepatic celU, that while fat itiself ukeii a* food InwU to uo increaw in tli«
hepatic glvcojien, oarbohydrates. which are edpecially fatteninj;, are must
active priMlucers of ^.'IvcoKon, and that the fi*t pr«Knt iu the hepatic cell*
mvatt to l>e increu^ie<l by suoh diets as natorally iucresM the glycogen in the
liver. Kn eviden<-i^ ha« been itflbred as to (he occurrence in toe hepatic cxM
of any nf tho several rH«\a of tlie conversion of glyooiien into fat, nor indeed
hMH it horn suggested what ihu^ustttjKt are. The view indeed is almost ox>
cluaively hoMid on the supgxiMil jir<>i>f that the hlood of the heuatic vein
contaiiw durinz life no lujfir, or nt lea.Ht not more than does toe general
blood or uvei) tn« blood of the portal vein. From iWn it is inferred ihit tho
glycogen in llie liver is n»t 1o«t t<> the liver by becoming converted into sugar
and w» di»cfa«T:g«(] into the hepatic blojd, and tbereHiro muM bs converted
574
Tilt: UETABOLtC ntOCKSHICS OK TIIK ItOPV
cnil lioiini, «!»|><'ciaily in it wani) pliioe, Iwf'irv n Jecocltun w mmUi nf it, tH
iJi.^MicUun will \h< f'oijiul tu liiivi' liule nr iiu ii|Hi)c*eenee, to he quita' or iH«rh
quite virar. In ^ivc litllc or no ixtrt-mini- n-ai'tioti nilli iotJiiHi, mil to iiinlJiiii
K very oun^iilcriilili' iinniiliiy of otignr. Aa <Htr Iiiivl* nuiil hIkivc;, th<* il«-(VK.-ii>iii
tmu of II liver lnki'ii iiiini<vJiikU'ly iifl«r ilfiitb ^'iit-rally <<(mtui»N iHtiiir liul*
sugar, iind lh« <|iiiuiiily of iiugiir in tin.' liver tt|i|n'nr», mi h tuiv, I» iiifraur
ol^adily iiAtr <li-iilli. ihc ■moiiiil uf jilyro^'ii ilmuiii*l)i»g at lbs midv lime
Thv Ta|)iclity nf the iliniiiiiiliui) «r iflycogni oikI tlir ntti itfincrcuB uf eujtir
vaiT nititti uixlcr vnriou» circiinislnitcoi- Momnviir, lliu (ImrriMtM.' of t.he oaf
and the increase of the olhcr nro nut nU*ii)r« Mrictly ^ir-iporlional ; and. indenl,
some obnrtvn have nuMstcd ibat there » ni> ri-Iation bi-twtivM tht? ;wo pro-
cenee. Nev«rthel««B, th« hmad fact romnins Ihal if tin.- Itvur of the aanw
well-fed aninial he divided into two balviw n» »oun nc piiwihie after dmlb,
and one half ibrrtvD into hoiliu); water imiuediately, white iImi i>iher half it
loll ex]Mi«ed to aotne little warmth for Mvernl. say twiiiiy-lour hount, tb>
decoction of ibe flrat half will ooiitaio much ((lycogoii and litllc 8U);ar, whilt
that uf tl»e second half will ouniain little (flyco^'en and tiiuuh ftU)pir ; and ibi*
liict may W taken, until the cunirary is proved, to eliow llial the ti'ycutrm
jint:iciit in the liver at the uwiueut of death is ];radiially af\er death by nomn
iK-tion or otJMtr converted into »ugar.
The action la that of ai>me ajt'iifiy vrhow activity is destroyed by llie tem-
IMTHliiro of loiilitijk' water; beiici; the <lirertiuiia repeatedly jciv en abuv^ to
thriiw the liver into iKiilio); wati-r. This naturally suicgeata tli« [ircwnoe in
thv liviir of an uiiiytolylic fiTim-ot. [tut not only have atienipis to iioJaie
fiviu the liver iiu iimylolytic frrnn-nl fuiln), in the hands of iiitMt obsei*V«n
U lenat, Ixit the exact nature of ihe vugar vrhieh Hii)iean *hotti< tlial tbe
I'hati^- M not n(li-cli.>d bran ordinary umylolylic fi-rment. In the ease of
ihe umylolyiit- Ivrment of saliva, iitimn-atio Joioe, intentinal jnicr, iiitd indesd
of all other ainvlulytio Mlimal llui'U, Ihv sugar into which utandi or |{lye»-
cvn is convertetl is maltaai. Now, tlie sugar whidi ap{irani in ilie liver after
death is dexlnine, identical, as far nt least as «'Jin at prescui be made uut,
with ordinary dcxtro»e. We are M, Iherefon:, to infitr that the ehiu>][« of
h'lye>>;^'n into ^iigar, which appears to f^ on al\vr death is carried out by
Bouie ftftion of the liver, proiiably of tne hejMtic cell itself, wliieh is duns
away with by a lem])eralure of 10O' C, but wlitch is not the aelicui of a
fenoent ca|vahle of Iwinj; ijtoluled,
$496. \Vk have used above the iihrase "well-fed "animal because ihi
luuuunt of i;lycDj|{;vn present in the liver of an animal at any one tiimi i*
verv variable, and especially detiendeiil on the amount and luitnrv of the
fiMNl previously taken, When all I'nod U withheld from an animal t)Mt uiyoo-
jlta in the liver ilimiiiiBlics. rapMlvai 6r<t. but inure slowly a(\<^rwiirf I. T.vua
atWr Kome days' :i(arviiti<>n n nmnil i|Uttntity i* fre<|uently still Imind ; but id
rabbits, at all evenio, the whoU; mar eventually ilioapjiettr.
If an animal, advi having btinu starved until iln liver may be iMsumed (o
bo free, or alnioHl frwi, from (tlyoogen, be fVit on a dirt rich in iiarlxihydmtta
oron uni< ooiiiii.ittng exctuvivdy of rwrhohydniloe, tW lis'Or will in a short tiiae
he found lorontninn vcrv lRi>n^(|iianti(y uf glyco^i. Obviously the prosenos
of rarbohyd ratios in fixid ItaiiU to an nccuniiilation of ulyeogiMi in the lirer;
and thi> is true both of Htan^b awl of dextrin and oi the various lorias of
sii^nr. cane, )!ra|>^-, and milk <mgar. The efli-cl mav Iw ipiilr a rapid one, fur
tflyuop'n hiM lieeti fi)ond in the liver in conniilcrahln •joiinlity nilhin a few
boors alWr the introduction of suj^rinto tlm alimentary canal of a starviny
animal.
If an unitniil similarly i>tiirvtHl be fed on an exclusi\'ely meat diet a cer-
tain amount ufglyco;^n is found in the liviv. This apjiears to be specially
J
I uitli llie <l»)^ ({)r<il»i)ily uiili <itlivr (-iiriiivonmiiiinitiiitl* iil*»\ iiiiil in
Kurk.i oil ibtHuliji-i't iIkm-oiikIiiiiI [.nwwtci.' "f ({Iv<'<»g<tn in the livcreof
<ii>^ M •111 ni««t nni> rv^iiniiil hi> mi iRi|iQrWfit tii(ii<:iit)<>ii of Uii* tnriiintiou
within lliv WIv of Diiii-iiitrnjirii-nii' from nilMp^oii* m(it«-riiil. But in tlie tint
place, llw <juniilily oj'glyi-ipp-ii lhii« stoivil ti|> in tlic livi.T t» thi- rwiilt of h
nieiit diet w much I«w ihsn [lint urhidi fotliKr* u|ioii n iiii')K>-hyilnit« diet ;
atid ii) the eec»nd I'Iiim, ordinurv nx-iil, tvpcriHlly li<>t«i-th>l) on which di>|i:9
in such exp«rinient« arc usimlly hii, conliiiiii^ in itm-lf^^ l>2.i ii c^riuin aniinJiit
dcher of glvcogen or mmv Ibmi «f' Kiii^iir. Mon-ovcr. when unimnla are fiMJ
iKit on mem, but on purified proteid, ouch »» fibnii. oii«(>iii. <:ir albumin, the
<|uan(ilv uf ^lyco^^^D in the liver becomes still sinnllcr, though, flccuniin)^ to
imMl observers, reniainiDj; grMt«r than during iitarvaiion. We may iiifiT,
lh«rof<>re. thai part of the glycogen which nppoar* in the liver after a meat
dwt i* n.-a\W due to carbohydtau tnnterials prewnt in the meal, Fan.hiiw-
<'ver, Would ap|>ear to be the result of ihe actual pnrtcid food ; ami we lunu
•imilar evid<-uce that ^laiin lakea as food leads to Ihe formalii>ii of m>mt
glycug«n ill tb^ liver. Uut in this respect these nitrogenoua sutmtamH.'? (M
far ttiort (»f carboUydnile niatarial.
With rsgard Ui nits, all i)l»ervers are Sf^md tbut these lead to no rocu-
iniiInli<Hi "f glycotteii iu ih« liver; au animal ted on an exclufii\-ely fatty
di<-t liaM no nmm KJyi^ogeD in ila liver than a starving animal.
HcDoe uf tli« tn>«egnat olasKs of fuod-Bluflb the cMrbiihydriili-n iland out
pruniiiwDlly a* the niHtancoi whiob taken » food lead Iu au aivunuilninliim
of f;lyoogm in tli« ]iv«r. Wi? may remnrk that the tfreattvl accumulation
of givcngi-n i* ttflhTicd not by a pure curbii hydrate diet, but by a mixed diet
rich in a>rlwhydrat«. A i|UBniily of carbohydrate mixed with a certain
pro]M)rtiui) of protoid give* ri»u to a larger iimuuni of iilyMigcn in the liver
than the saroo quanlity of mrlioliydrate i^ivei) by iu«lf: luid it is praiiiblo
that the prwcm-e of an appropriate qunntiiy «f I'at Hiill fiirth<-r amiets ihe
accumulation. Itiil lhi« nviilt pmbably <)e|>endi>. iu )>»rt nl k-n»t. on the
fact thai, ihuugh iHH«-rcRcx« mny lie met with iu dilffri'ut iuiini:iU, a mixture
of the several i-liuscs of fiKidstuHlt i;> mon; rrudilr dif^rKicd. ivsuliing in
niore Dutritive niiitcrial l>cing tbn>wii upon ihi- bhHxl, than is a meal consist-
ing axolusivety of one kind of f<>od'RtuH' uloiii^
Aa far as we know at present the glycogen which |]iiit> appears in the liver
W the result of feedinj' either with any of the variotii> lorm* of carbohydrnlni
or with proieids or with other ^ubstniicvs, is of (he Name kind and prettenis
tlw Name characters ; nt least we have im evidence to the conirarv.
The sUirioff-np of glycogen iu the liver i« also inHiienoed by otiier oircutn-
(tttneot tliaii the taking of food. For inslanoc, iu the fm^ an increnne of
fjycngen take* place during the niuier month*, hi the summer months the
iver of a fnjjc will be found to contain very little Klyco(|eD (Fi>j. 164, C ■.
unltM tW animal has been unusually well fe^ ; whereas a liver exajniuMl tu
midnintcr I Fig. iti4, A) will be found to contain a ooiisiderahle (pmntiiy.
oven ibougli uo f^iod bos been taken for months. Iu such a e&te the niat<-rial
for the foriuatiou at the tflyooeen in iJie liver must have been furui»bt,tt by
some |iart of thu body of the frog, and could not, as may be tlie cn>e nbeti
M nwal leoiis immediately to an increase of clvcojien, be HUpplit-il directly
from the food. It netMn* as if iu the summer tfie fro^ live* up i<i it» ciipitill
of liejMtii; glyt'oin'o, xiH-iuIin^ it ta fast aliniist as it is made, but that during
the winter a 'iimulity is fuiidir<)tu provide for the demnndi of the lale uinlvr
and early spring.
Thii^ winter Kf >rage of bcgwtio glv«>{[cn in the fm^ •ecm* closely de|>vn-
ilenl «n tempi-rnturc. If a winter Img, whoae lii,-«r m pi-nxiiuabty uk^tc nr
l«tn loaded with glycogen, Iw expoaMl IW iti>m« time to u lempflmturv uf 'iO°
884
TnS MBTAIIULIC l'B0Cg98K9 OF THK HOI>V.
«ii4l «l) we nro ixMiiiK <» that name of th« ninUirul It ntnr Hmiiuc luf EmU,
«ii(l ibal thr iittT which il miiv ni»kif of it in lu iriHriiifiitrdire fut.
$464. Gl]'rn|{i>ii U fiiunrl m otlivr pnrbi of lti« iHnly iltitii llii* Itvvr, MkI i
ti|ii(ty of t)K' rnt^.-i rcliKinj; lo tlif prcw-not nf givtvwou ii) otltt-r tiarua trill
1h>1|i 111 (i> ti tnic <vmiv{iliiiii iif tlit- [KirpiiM* of tat> mf<kt\c elycn^Mi. NuEt
to iW liver, till' KkiOotiil niiiM'ltiw iiri' jicrhiipx lh<- mmt ri]ni'|>icuoiifl glyoOfno
liolilcra. S.) l>i-i|ii«nilr in gly<»)r''ii f.iiiinl in inii<«ti- ihiit il may ho Kf^rM
It! nil oHiiurj though iwt itn iiivurinlilr ron*iiUi<.-iit of (hm tWue: indvtd
il tiiiir aliaoft be concMfn-il ii» u oixiMiliinit of nII oniitriu-tik- tiuuM. TIm
(]iiiintitr \*iiri<:« very Inr^ly hi>ih in the diflerant maotW of the Mine aniaal
nnd oormponilin}; miiRd«i of ditlt.<reDt knitnnh. It diMpponre. ii('o»nliii|t
to •om« obwrven, rmillly npoo »tnr\'»tioii, even Itefore llii' hcpalio i;ly<.-u):«D
iM9xbauflt«d; but nil ikliFprrere are not agreed on ihis )Mint, aitd in soiiw
mu«cles, at leasl, il a])pears to be rctHined for a very loiiit tinie. It it aai'i
to be iocmued in (|uanlity when the nerve of the muscle is divided, and th«
inuicle tbtia brought into a stale of quieacence. On the other hand it
diniinishea or even diaapp««n, beiiix appareutlj convertei) iutt> tloxinve,
wlien ilie mnscle enters into rijpir mortis Some ohMrrera hnve toiind that
it diminblies during leuinus, and maintain that it. after coiirerdion into d*i-
trose. is nweit ii|> in tlte »ci uf ointniclion, forniiux tlmxigh its oxidniioii tW
imnkediatc supply of ihn euerfty set free in the ivHilradiim. lint even j^nuH-
inv tbat the glvcoi^n in a mu«('le may l>e dimini*hi>d duriiiji jirutnn^
labor, ft caaitoi W uiimilted ihut tin? oxidntion i>r itlbttr chemu-Hl rhange i^
glyoogen it h nm'MMtry jmrl of th<- onliiiarv nHrtabollHin uf a itiiinnilnr ma-
traotlAn, Ninrv niniiy miiii'lri< wlmlly fni- imni gtyno^-n are |M'rlectly wall
nhic t« carry on loiig'i-imlinui-'l c>ntrni'lii>iii<,
Anotlier view of the uw of jflyco^u in muticle is mijj^i'sIiiI hy (he 6m
that uiidi'vclopoil cmbryonii- iiiui.('li?> are (iccnlinrl}' rieh in glviMgcii, In ■
young embryo, at the time whi'U thi^ miwcular subaUtnco, tbotigb undergiMBg
Mtriation, is slill largely " pmtoplnsmic " in nature, the quaiitily of glycogw
prearnt i» enormous: >t fnxjuently amounts to 40 per mnL of the ^)rv mat^
rial. At this period the hepatic cells are immature and very little glyeo;^
in present in tlieni. Ijiiler on, as llie muKlee become more wholly 8trial«d.
the itlyeogeD largely diMp pears from tlie muscle, and very aoon afterwanl
liegins to l>e stored up in llie liver.
The meaning of this can hardly he mistaken. The glvcogen in the Emma-
ture niu«c:)e h a Hon of carbohydrate material, laid di^wn on tlte s|mK. uhI
ready at onee to lie used in what we may prubably eall the fterce ineMboUe
MruEKle by whioh the simple protoplasmio cell aubatanee uf the rudtnMHt
of tne niuttcutar fibre ix intust^irmed into the highly diDerenliated striated
ODnlraeule suhsiaiHe. And ne nhall pmliably not err in eAi»idering lb«
glyeoRtm of tlie inatiire muscle fi hobi a similar [Mtition ; it ia carlwhydrM*
malertal Motml up <>n the vjxit, a local branch ko t« s|ieak of llie great carbo-
liydniin bsnic. It i> diwlined t4) become part of the eontnictile aillutanco,
atxl a* Mtrh will coutribnte fi the energy Ml free in a museular cnntrae-
tion ; bill its euMvy b only available in ttiis way afti-r ii hii» undergone Um
neoeaMry nteiabAium and become tnrt of inusi'nliir suhiiance: it cannot br
fire<) <i€ in a wntrnclion while it lies as raw glyc-igi'ii. or even at d^-Kiri"?.
in the interstices of the muMuIar tibre. We have aln-adv tiHT} iliscuu
in part the metabolism of " coutraotile sulMtWico." and shall probably ngi
relum to il Inter on.
i 4fiS. Olyeogen may alsit be found in oonsidenible i)iiantity in the pb*
centa. Here, as we shall see in the latter part of lliis work, it is laid dnwn
in epithelial cells which lie on tlie boundary between the mnleninl and die
TBK UISTOBT OF OLVCOflKX.
58S
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feul tiin»«. Am) Wrc Iim> there can bu lililc iloubt tbni it iii a More of
cwbohyitrnte tiiali-rm) tor tin- oourinlnnnit "t'thc fftii*.
It linn «li"i liwn fuiind in Icucocytw. in oiiMiliijp- fiirpiii-clw. wpccinlly in
lliodc liirg« rii|ii'iiy gfwing iind mpiHIj- iiiiiltiiilyin^' i-nrtilnpc oorfiiutdw
viliici) lie in till? iHilor xvneof fiiHocbondrnl owifii-iition, iiiii) In ntbi-r oituu-
tion*. In casM of 'linbet<v, whcro the body is wverlondwl nilh oiirbohv-
(Irate mat'.-riitl. it hiv been found in considerable qunntity in the trslio. iii
tlif bruin and cWnhere. Its occurrence in these Hitiiatioii*, and under tbo"v
('iroiin^tnnccf', may be feKsrd>^ b# addiliimnl evidence of the truth of the
view which vre hai'e expounded above that the main purpose of the dcprwi-
tion of glycoi^n h to aSurd a store, either ;;enerat or local, of carbohydrate
material, ^ihich can be packed awar niihotit much trouble go ling tg it
mnains glycogen, but which cnu he drawn U|wn aa a source of soluble cir-
culatiiig BUgar whenever the needs of this or that tiaaue demand it. Ii thus
forma a verv complete analoguo to th« vegelabk starch, and fitly earus the
oauM of aniDuil Mtarcb.
We have »<>me reastons for thinking that there are wreral rarieliea of
glyo^n. and that llxi gtycogi-n which vxititu in iiiihcIc ii« not quite identical
with that which oceure id the liver. Inde^l ihcrt' h'i-iii I<> tic ioiemieilinte
stn^cen Wlwn-n glyc<^n and March or dextrin. Tin- phyiinloj^ii-al valiii- of
tliese dillrrvnci^ii has not yet, however, been dourly clcif^rtninrd, an<l. with
tltb vautiiin. wc may continue to »{teftk of glycogen m a single ruImuuicc.
Dittlifitv.
!l 466. Natural diabetes is a dlsence chBrac(criH*d by the appearance of a
Inrg4- i|UBiility of wigar in the urine, due. a# ire have already said, ti> the
prawiicc of an abnormal 'pianlily of su^r io tbe blood. Into the [Mth-
ology of the various forms of this diwase it is im])o«ihle to enter hero ; but
a temporary diabetes, the appearance for a while of a large<)iiaDiity of sugar
to tbe tirioe, may he artificially produced in animals in several wavfl.
If the meilutia oblooj.'aiA of a well-fed rabbit be pnuclured in the region
which we have previously denorihed (j 176) as that of the vasomotor centre
(the area marke<i out as the "diabetic arvn aj^reeing very ol'wely with that
defineil BA the Taaniuotor area), llwugb (he animal need »i>t neoeasarily be
in any other way obviously aflt^cled bir tbe ojieralion, ila urine will be found,
in an hour or tiro, or even Itw. tu he inereiued in amount and to ooatain A
CMiiuderahle ipiantity of sugar. A little later tbe ituantity of sugar wilt
have reached a ninximuni, alUr which it decline*, and in a day or two, or
even le«i>, tht' urino will be again jx'rfrt^tly normal. The better fe<) the
aititual. or, more exacily, tht- ricluT in glri-ogcii lli<- livi-r, at the time of the
oijeration, the gniati'r the amount of sugar. If the animal ha previously
i>larvifl Ml ihat the liver mnlainn little or no glycoj^n, the nrintr will after
the oiM-ralion c-niliiin little or »o «iigar. It u cldr that the urinary nugiir
of Ihin form of artificial dinhctcn comet from the glycogen of the liver.
The puncture of the modulla cau«e« wich a chnngc in the liver thai the prtr-
vioiuly etiircl up glycogen disap]>cnnt. and tht^ liloiid hucDmc* loadud with
sugar, much if not all of which paixcs away by lh«< urine. In the alMCnc*
of any pn>of to the contrary, we may assume that in this form of nrtilieial
diabeicfl tlie glycogen previously present in iho liver becomes onvcKed into
sugar, jitsi as we know that it doe> become so converted by p^wt-morlvm
cbangea. Tlie glycogenic ftinctiou of the liver i*. therefore, subject to the
itiflueuce of the nervous aysteni, and in particular to the ioHuencc of a
rt-gion of the cerebri i-sfriual centre which we already know as the vasomotor
cvotre, or at lenit i>f n pari of lhat region.
58U THK METABOLIC PKOCKSiES OF THE BODY.
Before we atlenipC to discuss tliid itervuua iiiHiience we must sav a few
words oil tlie nerves of the liver.
^ 467. Tlie liver U supplied with nervea from the hepatic plexus, ivbivh
passes into llie liver nt the porta ami niuiiing in the jwrtal chiibI with tbe
hepatic artery aiul ^rtal vein, ia liistribued to various parts of the orgiui.
This plexus, which is the only nerve supply to the liver, eonskta partly of
medullate<l niid partly of iiou-iiiedullateil fibres, aud is an extensioti of tbe
great sciliir plexus already often mentioned. Into that plexus as we have
already seen the right (]Kisterior) vagus sends the greater part of its fibres,
and in that ple?[us both the abdominal splanchnic nerves, major and minor,
end, on Iwth sides of the body. The letl (anterior) vagus forms slight con-
nections only with the solar plexus but sends ott a very distinct branch
directly to the hepatic plexus. The liver, therefore, has nervous coiioection
with the ceninil nervous system by both vagus nerves and by the (abdomi-
nal) splanchnic nerves. Besides this other nerve-fibres find their way through
the s|>lanchnic sym|iathetic chain, or possibly otherwise, to the solar plexus
from the spinal cord without taking part in either of the splanchnic nerves;
and these may perhaps join tbe hepatic plexus.
Concerning the destination of the fibres of the hepatic plexus within the
liver we know little or nothing definitely. Some undoubtedly supply the
hepatic artery and its branches ; but we cannot at present say what propor-
tion of the whole number of fibres end in this way. Some again are destined
for the bile ducts, luid before tlie plexus passcii into the liver it sends fibres
to the gall-bladder ; ibcse ]>robnbly end in the muscular coats of these organs.
Whether any of the nerve-fibres end iti the remarkably muscular coats of
the ])ortal vein, or whetlicr, ad theoretical reasons would |>erhaps lead us to
supiHttte, some are cimnected with the hepatic cells we do not for certain
know, though some olwcrvcrs have claimed to have traced nerve-fibres
directy into the hepatic cells.
S 4o8. With regard to the exact nature of the influence started by the
puncture of the mednlhi, and the path bv which that influence reaches the
liver, our informaliun is at jiresent vorv imjierfecl. One thing seems clear,
viz., that the inHnence in ipie^tion is not carried down by the main vi^s
trunks: for not unly has the section of both these nerves in the neck nn
marked eHect in the way of producing diabetes; but the "diabetic punc-
ture" of tbe medulla obli)ugutii is as efticieiit afh'r <livision of both vagus
nerves as bcf'nre. ^^(.'eing linw cluse to or almost identical with the vasomotor
centre U the dialietic centre if we mav use the phrase, it seems natural to
supfiosc that tbe undue conversi<in of glycogen into sugar which follows tbe
puncture is the result of some viiaomotor ilisturbaiice in tlie liver, for instance
diliiliun of the hepatic aitery. But we Inive no clear proof that this is the
true exjtlamilioii. ami. indeed, it' ihc |dienomena arc the result of the failure
of normal vasii-coiislrictor iuipulscs, thiBe impulses do not reach the liver
by the Irnct which we .•should su|)posc iheiii naturally to take, viz., from the
va'^o -constrictor regimi of the cord through the splimchnic nerves, for division
of the i-planclinic ncivcs even on buih sides docs not cause diabetes. More-
over, that the effects are nnt due to vaso-dil,itor resulls is shown by the fact
that strychnine poismiing |)Miduces diabetes in frogs, and produces it by
rapidly hu riving into sugar the hejiiitic store of glycogen. Now in strychnine
iMiisiiniiig the liiiHidve^^sels are constricted, not dilntcii. their mu.-icular fibres
like the skeletal muscles being tlirown into contractiiin )>y tbe action of the
poison.
The vascular relations of the liver are it is true peculiar, the small hepatic
artery contrastiiit' with the wide portal vein ; ami it may be tliat the dialwtic
effects are contingent not so much on the absolute account of constricfion or
TUB HISTORY OF OLVCOOEK.
687
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■liinliuii or lh« liF]>iitic nrtm-, ss nn ihe neUliim of thu How tlinxigli that
nrtcrr to ih« lloir tliri>U);)i the portal veiu. lti<)ved, in xuppurt of this riow
limy Iw adduced the Untement tliai seclioii of bath gpUitchnic ii«rv» uot
uiilj does iiot CHUBC dinbi-ies. hut ptvvtnu ihi* iimiiiiI rlTccli* of tlie dinWtio
puncture: sod Ihia hiu be«n inti^rprvtcd lut ithowiiij; thitt l\u: incntmni |M>rlnl
Ruir thus induced counierlwiliitKxv tlitr (rifi-cL-i i?f dilation iif tho hi^Mtio itrtery.
Uiit we have at present nu exact iiifiirtiiali<iii. Hiid liictn i» lu y«t Dothing
distinctly to DdKiitivi: the vi«vf that in thin arlifioini iliatwli»i tlw oervmi*
iaSu«ace is bmu^-ht in 1i(«r <>n thi- hi.'jintic ixll itMrlf.
Tli#re ai* siHoe facta which noom In »liow that the path of ihiit nurx-ogit
inRiieiic« uti ilH •any Ut ihe lircr from the »pinal uDhl pnmn ibiMiigh tho lint
tbonidt- (ranv''""- fC«iiKltt>ii Mt^llatuin ; hut hoirU mchn the hepntic pluxu»
from thL-> ^«ij;liuii a vriv>ll_r unknown.
§ 468. A icuipfran- 'Iiaiict<» may be brouj^lit about by tho admini'lnition
nf the viibManit: phloriiUin. This, however, i» a gluoo«ide, and part of Iht-
Rugar which apprnn in the urine, vft«r a deee of it, may como airect from
tbc drug iucll ; Uil tho quantity of tugnr dischnrK^d U too great tu bo
BCC4>uo(«] for in thi« way. and eimilar iljflljetie effecld are produced by iho
ndministrntion of pblorctin, a derivaie of phloridzin, not a gluooside, and
not giving riac to sugar by its own dectmi position. The »U|tar which apixuire
in the urine after u dose of ihia aub^taiice atietoa to coqm: in part at Irast
from the hepatic store of )>tyco|{en when that b present; but the druj{ will
fpre rise to Higar in the urine of starving aniraals, from wboee Itrere land
other tisMiee) glycogen is prwumahly ahwnt. In iiuch camr^ the drug appears,
in wnio way or oth«r, to either >tir up th« hepatic aWa to a maniifactufe of
of sugar (and this fact is worth remembering in relation to llie discuKUvn
which we lately entered into ( j Mi'l), na to the nature of the formation of
glycoiien ) or to produce sugar out of some of the othor tiaMiw of the bodr.
Artificial iliatirtts u alwo a prominent Kvin|ii<ini of urari poisoning. Tnis
■• not due to the arliticial rapiration, which i* had recourM) lo in order to
k««p tbtt urariaetl aiiimaU ativo : Wcmisc. ih<>ii);h dimirbitnott of thu renpira-
tory functions sufliciitnt toiiit^TfcnT with the hcjuilic circulation may pr(«luce
sugjir I» the urinr. arlilicinl rci>;>irnlion may with caru Iw carriwl on wlthiitit
any Kugar making ilM apjitatmncc. Moreom'r, urari cniiws diabotM in fmgs,
althotigh in IhnH- animals rapiration c*B ht sntisfactotily cnrrinl ou willioul
any Milmonon' rvc^piratory moToments, Th« exact way in whidi tbil form
of diabeUv is brought about has uot y«t been clearly made out.
A nty rimilar uinbet«s ia Been in carbonic oxide poisoning; nnil is one of
tbe reaultB of a sufficient doae of morphia, of nmyl-nitrate and of soino other
druse.
Tnere can b« no doubt ibat in diabetes, arising front nluitev«r eatuo. lli«
wigar appears in the urine because lli« blood ooatains more sugar than usual.
Themtem can only dispone (either by oxidation, or as seems more probable
in other ways) of a certain <piantity uf sugar in a ceriain time. Su^ar
injcct«<l into the jugular vein r«api)««ra in tlw urine irbenever tiie injectioD
Ix-oomoA so rapid that tbe percentage of sugar in the bloul reaebee a oerlwu
(Inw) limit. Su^jnir in tbe uHnt nienii.'t an exoras of sugar in the blood, How
in natural iliatteti-n lluit excna ari:4--H litta not at prment Ik^d clearly made
out. It may be that some fornin of dinbeuw rtMnible the artificial diHl>e(ee
juu dMcrilH'd BM rcnultiiij^ from iiunctun.* of the medulla, and arise from a
t<M rapiil conversion of tlKi lupatic glyoogen, or from carbohydrate nuilerial
(iiilijig to bo Hlored un a« givcogi-n.'or from lui exoonlvc nuinufa<;ture of
(■arbonydralc mnlerial by the ^'jiatic cells. All fornw of diabeies, however,
cannot Ik Mtisfnctorily exjilninvd in ihiHnav; and it lias U-cii fiiggtsieft.
iboiigh adeijuate proof has not yet been su|>plie<l, that tbc sugar of diabetes
383
TDK METABOLrc PROCESSES OV THE BOUT.
IB iif a peeiiliur nnlurc! iiml iici'iimiilittfti in tho bluod because It U i
ncderjto ibi^i- i-hiiiigi'Ti, wlintrvrr thr_r Ix!, vthioh facfull ttic uurraal at^ o
the blKod. VVi' c-Jiiiiiot b<'ri> iliscuw ih<- subject in dcuil ; liut tb«re ia taoes'
to be Miiil in tiivnr nf thv v'luvi iJiiil th<? witrom nf the cxi'M* of Kiiipv >>> tks
bloiHl iniij 111- varimit, fttiii hcnt'o ihnt Kcvcnil ilistiiu-t variclicii of iliab«c.«
urny exWu In wvcro cyiwa of dinbolcs tlip nbcrrniit nstiirc of ihi" mrtaW* i
which is piiiig on in mmv or other of the tissues of thp body U «b»wii bv ifc-
aiiliriirntRi? of iibnorniiil sulMtMncce in the urine. Thus arctone )■ frctuatt 1
IirowoDl. nml the liila] iRsiit? of certain cSAes has been attribnted 1o pnuoniL
<Y tbiit snb§lnDce: uxybutyric acid and other Tsrious orffttoic, chictlr vol -
tik, nci<ls are also Bometiaiee prttenl. But in re&ped to theM Moa otW
abnormal bodies vm are not at praeul clear wlieiher they are lik« the mtM
itself the i>roducIs of an abnormal inetaboltam whiuh ia the root of thcdS
esse, or whether they are senondar)* products, that ia Ui my, producU of ltd
general diiordered luelabolbin iiiductd by the ciMi«iant [ireMnce in the Uoc:
nf an excew of suear. We have iilriNidy in diKcuwng the formation of g1~
cojien cnllvi) ntli-iilicm t'l the fnt-t that in vevore cane* of diabelea the Riga
luUBt havi; a noD-aoiylarrotM Honri'e; and the fact tliat'the urea b incKUa
(and that Uh> in mjniir ctin* in rnlio with the xngar) in dinlMtcc mggeaU tba
tliu Mignr may ariH- from protcid* wbirh havo been »|>!it »p into a nitrt^c
noUM (urcn) nml a ni)n-nitr(iK<.*>i<>n« moiety, and m point* out the utny in whic
priiirids may be a »oiirc« of glvcogeti.
A« a Mrl of converse to diabetei* we may mention that th« adminLitratic
of arsenic in suttic-ient doses or for an ndc^juate limfl pr«Tenu nn oocuiauL
tion of glyeogen in the liver nud apparently in the body Kenornllr, wbater*
he the diet need. The presence of the metal in the hepatic cell Mcmt C
prevent the cell substance fruai manufacturing );]yco;{eu either from rarl>uhj
drate material brought to it, or out of its own aubjlance. A* any other lun
of convenw we may iilNn ttnie that the ad mi nisi ration of f^lvcerin, eepedall
Ifaraugh tho aliinentart- canal, dirainiihes th« elTcel of the diabetic puncLun
or of morphia or of i^lhor poiyniiins, in hurrying nn the heiMUic uore of gl]
<!ogen into fU|fnr, and thus diminishes tho sugar in the unn«; iht present
of tb« glyoerin in the hepntic coll np)>onrs to he in Home nay a blotlraui
to lbs conversion of tho glycogen inln sugar. Now glycerin injected inl
the altmentiiry canal nf ■ normiil animal leads to an ioenMc of KlTOoeen '■-
thri liver; and the viffw v«ry naturally sugeesCs itself that this iiicrMi
arising fnm the glycerin is to be explained Dy th» glTcerin inbibittns i
si>me WAV a nurmal conversion of the glycogen storo into atignr whicb
continually going ou, and thus increasing for the time that stoic.
The Spleejc.
4
ooosidn
S 470. Tt"! tirwrliirf of the t/ilffn. We may uow uke up the coos
tion of tite forraatiuii of the cunnttiuenM of bile, a matter which in dealii
vilh tho scemtion of bile (i 2't7) we pu«ipoued. Of these conatiiueaia t
moft important are the " hile-snlts" on the uue hand, and the bile pigros
on tIte other. \Vr< will lake Ih? latter first; but linoe, as we have alroat
naid ($ 26\ Ihf bile pigment, bilirubin, apiwars to be derived from bvin
ghibin. and since the sple«D seems to br especially concerned in the cbaag
which hiemoglobin undergoes in the body, wo must first turn to the structu
of that organ.
WhflD a ffMb spleen is cut across, the whole interior witbin tha we
delineil coal or eapnih (Fig. 167] pre<eiits the nppearanoe of a dark-r
spougy maaa, iraverwd by irregularly disposed paler bauds or tmbrfnlir, mt
A
Thix NtETKni or ^rtttx-nxf. iiiiiiiiv >(ji<.MriiiD, ^ikmiwu in* Mods or <hciiii» cif
A HlUI.L V'KIM IN TDK IHIKHtlKTa OT Tllr I'l tr.
r, lb* Uin. flUtil Willi liki>il>nir)>u>i*l<ii. ohli-'b an lu ounlliinlt;' hIIIi nltwn , M, nllltiK u|i Hid
t<BWntHm «J llionlinimillMiiei'r llw imlp: v. wktl dI IM kLd, The aliiidttf Uallo iinittp* IM
'(■d liliwd roffaiicla *fi> iMkooipiiKle*.!
IftH.] Tlie rciluesB is obviiiiBly <Iii« to red blood -carpuecles ; and h U clesr,
U Ui« (lulsei. Ibat the 8j>l«en ^isoetaes au uuuaunlly large supply of bluod,
wliicli iDorcover seeina to be diApoeed iu an uaiisuni uianii^r.
MO
THK UETABOLIC rHOCKSSKS vr
Whet) Iiv II kirvum of uonnal oaliiw solutioD driven Ihrouj;)) iu vcadii
itiHcIi IiI<3(h) dh in iHtHible U wa^-Ued away from itie Bpkeii. aiiil tlw urxui i*
lUlbkciiiciilly hunlincil in itiK unual way, prererakly in a diad^iidifl iHiiiililiuQ,
Mclioiw iwt^nl ilii; I'xllowing ftntureH. Tlie njMule omaisbt nf an inttn
lltrer of c<iiiiii>clivc liwiiii- <Hivi-nH.I nilli l^[IiIlll•lil>ltl [ilatva, rurinirig il)i> |<rn-
li>unl emit. »ii<l poHtinuDUM irith ihJN aii inii«r iliM-)>t?r Ikvit, (iiaipntvd of
(■uLn«ctiv<t ItMim wilh nrlitorkx of rliutriv fibmt, and (■•>tiiniiiin>; a (.-cftiie
Diimbvruf )>iiti<lU-« of |>lititi imiiwrulnr li)t>Mf ; tliU ilifiMr layi-r uf tfa« ckii-
Hilv givra otr roiitKlcil »r tlntu-ititl bnmlKii ol' tbo Minif iiiiliire as itsM,
wlucU p«M ill all dim-'ti'ino inUi t\w iuli'nor of tW urgaii, braudiiii;; and
Biiastomoeing fr(i«ly; ihuM am Imt d«ri;liii)rd lowanl tb«> side nf failm
«hcr« ibe bninchue of ihc spli!i)i<' iirtcrv miiIi l)i« spKtiic iit-rvea eulfr, bikI
nbeiii'o th« Hplcoic voiiM iasiie. Tbc inndo of brauoliiiin is irre^lat. nutl
the braucheH vnn' in Biz«, Inrx^r tralwciitii.' (nviiig Hm Io Biuallor unii«.n
tluit llie ivhule interior of the or);an is ilividml iiilo a labyntiib of irngaht
coiuiuuDioaliu^ chainben, uhicb cunuiu in the freeb state the «jilwn-pu)p
ineniioned above.
Tbe basis of both csiwuId and trabcculn, email aod jfTtat. in coimeclm
tiwiie well funiiahed with elastic eleoMots. Iu some aaimxls, a» fur itutaiK*
in ilto dog. tbin Wxiii is n ridilr pn)vi<lud wiib plait) muscular tibfvs, tiat
IkiiIi trabeciilie nml (V|iiitli! On its d«ei)er layera) w«in to be almnet eiiliret)r
cx>in]HiM>i) (if miiitciilur tbvae. In other aniinalt, in man for iii«taii<.v. tlir
mkui-iiUr i,-l(;iiiL-iit« art! itiuuii more scanty. TImi cu|i$u1e and tratuvul*.
amall and great, thu.'« fona a spim^like flramework, which tieiitg elaalic cm.
evon in the cium vhtm tbe muscular fibre* are srniity or aliseDt, at oa*
nomvnl be diKlotKtvil mi that the chainbcK are rajHiriiiuH, mid at aniilliw
moment ciin by virluv of iUi itladticitv vlinnk mi that ib« «liainb«n iu» rv
dui'c<l in Kixo. In tbe niiimid* in which niiiixitliir fibroc ani abuudani mill
en-amr variation* of mim- are ]Hj«iible. When tbe musoivs are relaxnt. a
disli-iKline force, nuch m U fiiriiiilicd by ttie |>r(wun) of tlw blood^alnuw.
cMa Bircir out the franicnork to a very great bulk; and an adet|iinlr ihhi-
tnctioi) of the muscular Hliri'ti <.-An in ttirii ixiuwu) tbo snon^-liki' man inU
Very small itimoiwiimfi. A* wc glial! pitvently eoe, rhytnraicnl ur i>th«r eon-
Iractioiia of the cn|)«i,ilv and trubccul:>r Inbvrinlh. in nniiuab in nliieh iJiM
are Ur^fly muscular, do produce remarkable and imporUut varintjoiu b
the volumi* of the spleen.
!) 47L This s|)i>nKedikB framevrork of camule nnd trabeculn mniinda dm
of the structure of a lymphatie gland, ana the raiemblanoe is carried Mill
furilier bv the ebamben of the labyrinth beinx occupied by a ririirulu
lU'idiDcation of ooniiective titaue. Itut the rmeinblnnce is superficial only.
The chuiiiberM marked out by the LrabeeuUe of tbe »ple«-ii am wholly Imgv-
lar; there is iiul, as in a lymphatic yland, auv dislinclioii )K'twc«n a iwrUX
with large radiating cliamben ami a inetlulla with aiiit>lomi«in|[ luhuUr
(^hambere ; the trabccnlte are eloteit loHard tbi- hilu», but olhern-iao one nait
of the spleen, il> rvgardn the arrangement nf tralxMrulie. is tike any ulnar.
Morvover, the n'lii:iilar tJMiue uvcujiying the chainl>en> shoirs n<> di.-<Uiie4loa
WiHceu lyniph'sinus nnd f<dliclv, is not exactly like the line reti<-iilum of
the one or the coaro: naicuhim of the other, but of a nature distinct from
Mcb, aud hiu> MO vpenal nonuuction with lymphatics, but has peculiar reU-
lio«» to the minniii lil<H>dviiwl«.
Except at the while «|ii>ts uceumed by the \falpi;;hian oorpu*olo*, of whidl
we will spi-ak pnvmtly, the epreiiie rcliciilnm is somewhat coarse, c<>an<«r
than ordinary iidcnoid lii^uo (^ '2^0), and over a large part of the 8ul(H>n is
miidc up of brflnchc<l nucleated cells, the brancliiw of which are roMnbrnnuui
and fiauge-ltke nthcr than lilameutous. Tbi«e flan:^ of uciglibortog oelU
TU8 SI'I.EEN.
mi
In wilt) <-iu'}i niWr, iind tlili* fona n liilivnniliiiir nrlwin'V, llu' wnllii (if ihi!
iK-
iif which
■I not I
It ot
iliir «hn'li>.
I
]»niuagc« 111 winch arr t'>rnu-<
In .""nut pnrlji nf llu- «|iln-ii, howpvcr, iIkdw fliiiii;f-lik(' |>i-<ic'-wm un- rcpliioitl
!»)■ lilin*, mill, thi- Iiwlics nod nticlpi nf ihc ciHi*.iiliKni. ci-tts Mnj; nrv, Ihe
reliculiim i>p)irar« a^ n more oniinnry reticulum uf tine til)rL'«.
T)i<- )Mire of this reliruliioi, vrhrtlic^r flang«-lik« or li]i«nirntouif, nre at Um
cii}^ uf itie trnbcculft continuous willi llic siibslanM' uf die trnl)ci-ula> ; tlie
Bnitill«r iratKcuhe brvak up into llie r°liculiim. and llic Inrj^r trabectihe are
iVingeil will] piuecMee coniinuoiis wilh tlio bars ut' Ihe reliculum. Tbiia the
ruar^r network of the trabecular Bjstem is continuous vrilli ibe 6&er net-
work of the reticulum.
Tlie n-ticuliiu of iJie Irmphntie ^land coiit4iiR«d. it will be ronwuibered,
Hde^ fliikj, Icucocyteit, ibese betn;; cruwdeil iu th« folUck ami more Kpnne
fn the lyiuph-fliniuL The ajtlenio rHioulum nUo conliiintf )fiu<!iicTU-ii, but
tIiiT.-H! arc thrown )Uli> tin: iMU^kgroiind by the ]iir)i;i! iiutubnr nf nxl eorniiMilM
\ritli whirh ibo mt-Tihc* of the n-tiouhirn are crowded. The n-ltculiiiu. Id
fact, i* lllli-d with blnod, ami [n'CiiIiiir nrniii^ntunl* qx'iA bv whii;h ibe
blood gaiiti ud^vM to the ii{inc<4 of th« rdiculiiin. Wluii wii »ii)ke of above
M"»pT«on pulp" rxprvsuiil from the friiih opWn ci>ni>isl« ot fragniunlx of
the rMicnlum, to^IxT with tli« ml and white eorptiwlv* occupying the
HMshai of that rr'ticulum.
1} 472, Tlifl apleoic artpriw entering thi> iiiileen at tbt- bilus arc in Bomo
■niiualf ut lir«t mmxirtnl by ihi- tnilwculu;. id«n<; irhirh ihi-y run. dividing
as ihpT go, but the l>nincht»< iil laxl lc«v<- tlu' trabt^culic iind nlungi> into the
rvliculum. [n other nnimnl" the artcrire run more indepondcnt of ibe tra-
licculie. Ae iher leave tlw trabeeula-, or toward their Icrininalioiis. the
email arteries are apt to divide into pencils of small twigs. In a similar
manner the veiu« may be traced back along the trabecnlai, small and great,
along which they ar« gathered up from smaller veins uf Uie reticulum ; but
Ihe veiuB do not run in the reticulum as distinot Teaaek to the same extent
that tli« aneriee do.
Ill the reticulum tlie minute arleriea, according to most observers, are not
ci>utiiau-Hi» in the u»ual manner with veins by meaina of cluwd cnpillariefl ;
but a pm-iilinr armigemenl i» met wilh. The epithelioid \>\Mei furuiiug the
mnillary wall, indlead of bi^iug ceraeDt«d together to fi>rm a continuous
tul>ulHr ^hiriith, are si-jMrate from e«ob other, i-onie asunder as it were, aud
lb»« allow ihi! lumen of the rafiillitri' or rather of the minute artery to opeu
Dut into tin' .■<pl<-nic reticulum: iixlt-'fl. the epillielioid plat«« uu longer retain
their oimph: i>plindlp ihupi-, hut bivDrtiing brauclieil and irn^utar are tmns-
lornKd into the ci-ll* of the ix-ticiilum. In thi« way tht- ohanncl of the
liliMxlvi-tM'l beeomi^ coniinuoii.'t with lli<- Inbyrinth of llin splr.iitc reticulum ;
ntifl bv a convert.- jiriicCM the :*Him; Inbyrinth ix made i^intinnou" with the
plexift'rm b<^i»ning" of >mall veins, tlw iw-cullci| vcniMw ginuvus, which end
in ihc; veiiu* running along the irabivulie.
Thus ibe blood Howiti^ along the «iilcnic arterv twiaiicii from the o|>un
of the minute nrti-riM into tliv aplenic reliciifum. and in gathoro'I up
the reticulum into the »p<m nioutlia of niinutv vcimt. When the cn|>-
le and Irabcctila- are in n relaxed ivmdition a n<i( inomMilurable portion
nf blouil thus oecapcs into tbc reticulum and tarriiM in the raiMhta, wlier« It
nudergoes olmngra of which wc nhall prtMtutly sjxMik; whc«i the cii|i«u]e
un<\ irabeouUe are contracted and ibninkcn, tlie blood Hows in a niura
tliret't manner through the narrowed channels from the art«riM into tlie
vclnn.
§ 4T3. Tlie lymphatic ves^Ii nf the dpWn lire not very numerous. The
cn|i4ult> and the trabeculic coutain tymphiitic pIcxiiMi ojiening into lymphatic
IJU2
Till: MKTAIIULIC PKOCESKSS tiV THE ItU 1) V
Irutika, wbiclt l<;av« l)i« lilliia uhh thu bti)odv««wl«. Tlicrc i*, hunrtm, >
roiukrkniili: lympbatic iluvclopfDciit, in tho form of n t^livuth of silenMi
U•MI1^ wliich n<v»iu|iauic0 cIh' ariuH^i Air moidi.- (tiauticv mt lIk-v lc«ve (br
tnil)rculii- tuiii with whicli tlw )yiii]>bHlic vifw^ln of the tmbt-culw nre wo-
n«ilt*(I. i^i luiig n» tii« aruriet am runiiiug «]oiig Uiu Irabeculia:- tbts wteBoU
flwaUi il«ilii«r nb<wu( or extrvriKiIvKaiity: butMth«iiDor arterini bntac^
plung* i>iU) the nticiiluiii, it is wi liu-nNUH-il iii btilk nt iiitorvalB, util f*p«
dally wh«rc nil Rrtcrv in 0)vi<liiig into two, iw (o form an uval or :'plirrind
naas Twble to lh« D«kwl vyo, mid coospicuoua from its color Ihvjiu*^ ibc
Adenoid tinue, crowdvi] m usual witU leucocytes, appwtra vrEiite or i.iil<'rW
OA oompaml niih the darkixtj eplcen^ulp. Th«i>e, in fiu:t, an- iltii Mai-
pighiaa corpuscles spokun of nlwve. Each Mal|ti);bian ooqiuarlc if a lamt
or leas globular mass of adciioid ttwue, crowded wilJi leuoooytiM. >]vv«1ii)mi1
Lp the aav«Dtilia of n minute artery running in the splento n-ticuluru. .\i
K rule the developmeDi takes pUce on one side of tlie art«ry, so that tht
rouivded MalpiKhmo eorpusele seems to be sitting on tht^ nrtvrv. Soof
tini«a the develujiinent lakes place mora or lec« nyularly <iq all sldei iif
ihi^ artery, no that tJie artery ap|)eara to nierx^ aud run thruugh tite imiwM
miuK, which t* then called not a MaljiiKbiaii (uirpiivcte, but n "br|Mr
pliuc s{Kit;" and not infre<iuently the artery dlvidca in lh« laidilte of uie
maM.
The adiiioid tiwiitr. as eli»e»h<'re (5 -t»0), is compoard of a fine reticulum
crowdi-d ii'ith Icuoijcyti-n ; ilu- corpuscli', in fact, cJoecly rraembles a wilitarjr
!;luid of the inicaline or a muiideil nuuut nf thr folliciitnr subsinric^ivfa
ymphatiu glaad. Bui it dilfrn from thvwc dnictiiro in nut bring ra^
rounded by any dintiiu-t. lymjih »inu«; at the i-i renin fore nee l)ie iruc iidNoU
tjiseue ymm!* vuildvnlv into the ooaiwr aplcnic reticulum. The arivr}', at It
puses through Iho Miilpiahian corpuscle, nv«s otf to it lino bniurhi-«,'wbkh
form a capillary- iiclworK through tlx- adenoid tissue, aud at the i-iroi»-
fereiice open mit into the htliyrinlh of ihe splenic reticulum. Thaw- ]ktl)-
pigbian corpuscles are so nuineniui that in a section of a fresh aoniinl »j)ia»
tlie dark-red ground of the splenic substance appears quite mottled by
KaaoD of the white dots. Hence no incousiderabte portion of the blond
raadiing the spleen finds its wav into the nieebee of the HpK-nic n-iiculum
after junwiiig throuvh. and, proliabty after acting upon, aiid being umI
upou by, Iht- adeiiiiid liMue ol a 3[atpighian corpuscle.
Wlial b known a» «uyo iplrtn is so oalletl becaiiae the- Malpigbiaii oorpuv
clcs become en largetl and traiu^arent, incoiuefjueiiceof t)w IrucKwyti-euDdei^
going "laidaceuiu" tiegeueration ; the nuiui^ chiuijce may ab»'af)eot Um
adenoiil ^•ni« of the small arteries and may even Mprtwl to the itplecn-pulp.
^m. The nervn of the Hple«ii which [iiim< into the oigan m the hih»
with thv bloodvevelK arv derived frcmi the »)lar plvxus. They eonj^ist rbteSy
of non-indullnti'd fibre* mingled with which arc n few ntetlulliiled fibrea
Their tenninatiotiji huvr not Ix^cn ns yet exactly made out, but white many
presumably are distributed to tW hl<)(id\'e(«cls, there can bo little doubt ibM
some end iu the cnpsitle a»<l triiheculic, at least where iheee contain muscubr
tia8ue,atid thus bring the contractions of theee structures under theguidanev
of the special nervous «yKlcm.
The ceutri|>e(al courM of the fibresof these splenic nervos has not yet beta
made out delmitely ; we may perhaps snfelr conclude that the majority arc
derived, like the fibres distril>uted to the iKighboriug Abdominal orgaas.
from the dorsal spinal oord. That the vagus also eoniributes fibres i» very
philmhle.
it 476. When the sa-called spleen pulp b examined under the mi^^oM^«M.
it is found to consist, besides toe bnDcbai oells and libres constituting ill*
ntCmluffl. of fvllf vhlcb may be detcriliei) as pnrtlr recTmrS^^^^Bd pnrtl^
ibilc curjiuKln or l«iim>cyUft. We »)>ukv at llie tn«>lii?< nrVBRBuliim am
iMRiSllnl with liliiDi); btil it U olivii'Uii ttiat itit? corpu»clm of the blwid
■M BOirr Inn n'liililv ibroiijfb thu lubyriiitl] than iI<m-« llic tltiiU plaxma, add
tkn imce a ranopotnilinn of tbu c»q>u»'li3< tin ci>ii)|>iiri-d wilb iho pimmii
BWl takv pUc* Ui tbo iii(iih«*. Tliv nuiiciilA of Iho inttthcv cnniiiit. pruiwrly
^Nakloc, bo CTillrtl blmx], but arc riithvr uggn-gulioiu of rorpusclw wilb a
nhttnly anuUI i)uiLotilj of Huid.
n* whit« corpuKin or kucocylra are very vnriout. ^omc aro *inal), lik*'
itvtMCorTinarBlviiiphaticigland.lhvuGll eubctiuic« bdiig WAnty relatively
U the nucli-iia Oltivn nr« indHtingiiiflmble from the ordinary white corpus-
*im of tltc bltMxI. Olhcn ngaiii are large, twice as largo at an ordinary
■•hit* corpitacle or {-vui lai;^r than thi«. |iuM(«b nion> than one ducIpu*. and
raiDUio in tbcir rt-ll subftanni nunicroiis refractive. {>Ale ydlow or colorlcas
jcruiuli^ ^me of thef« lari^er ti:irnis. which lik«r thcothcrieihibit aiuoibaid
•OOTWMtitB. and are often irregular in Ibnn, ar« characterized bj- the pr«s-
*tae> in Ihcir cell lubeUiDce of red corpuscles. somotiniM in almost a natural
CVkndition. Foiuelimes more or Icn irregular id shajx? >nth their red bitinu-
K*ulitJi cbaUj^og iuui the brovtuer btetnatiii, and fometimcs disintej; rated iuto
* uaM of brown sraDul«L The fluid or plasma in which theKcelle float
*1m mataina beucKa Dormal red corjmsclaa a certain number of red corjius-
*^lca in various stages of change, a» uell as [li^ineui i^raiiulea which ap{>ear
^ be derived fVom bnmoglobin. Ubvtoualy a certain number of red rariius-
Clu do umlergii cliiinge in the upleeii, but whether the change in mainly
c&cted in ihi: ctrll >ub<UU)ce of tli« celU junt nienlion«d, or takes place iu tite
plnaaim,the pnnluctH nfdiHinlcgratiuu being «ubiH>i|uei)tly taken up, iu atnu'-
boid fashion, by the cells in <iueKtiun is irnt as yet clear, lleaidea the above,
in ibe sple«u uf youue auimaU, nudeutcil celU with hieiuoglobiu holding
■ cell aubtUDce, luemaloblaala (xit § '27), huv« becti d*wribed ; thtae ar« tatil
■ tw npl*^ si"! iu the spli-cti of adults aflcr very great low of blood.
B $476. TftfmuTfinrnUo/llie^lffn. As we have already stated, ihevolufue
Htof iIm »]d«D if fubjccl to considerable raniition».
AfVcr a inral (Ik splc«n iucrvaws in size, reaching its maximum alxHit five
iKMit* aHcr the taking of food ; it remains swollen for some lime, and then
veCurtH to its normal bulk. In certain diseases, aiicb as in the pyrexia alten-
<laat oa certain fevera or inflammations, and more es|M>cially iu ague, a some-
«b«i similnr temporary eulnrgemeni takes place. In prolonged ague a per-
maoetiL hy))enrupby uf the spleen, the so-called ague-oake^ occura.
The turgesoeDce of tlie spleen seems to be due to a relaxation both of iho
small arteries and of the muscular timue of the capsule and of the tnibeculu-;
lube, iiifact. a vascular dilation accompanied by a local intiibitiun of the tonic
ooDtracliun of the other ulain miuuiular libre* euiering into the structure uf
tlw Dfgaa, tbe latter, at all events iu «omv animals, heuig probably tlie more
Important of tbe two. AikI the rondiliiin of the spleen, like that uf other
vaM*u]ar areas, Bppean to be n^gulutctl by the central nervous systein, tlw
digaativw turgCKeucv being fairly comparable to the Hushed condition of the
pDCfens and of the gaslnc mcmhnine iluring their plianeti of activity.
Tba a|>f>li<«tioa of the j>lc(hyxm<i^'r»phiv method to the spleen, carried out
■• tbe way which we dM^rihcii in *|H-iikiug of the kidney ($ 41 1)^ enablea us
la study iiiur*^ exactly the variations in voTumi- which the organ UDde^[0<a.
A "spleen curve "(Fig. 1IJ9) taken in tbe same way ib> a " kidomr mm**
doat iH>l. in ibe dog at all events, show variations in (be volume of tW iplaui
(iorreB|wndiDg with the pulse waves. Tbe kidney curve, n» we have iMti
(.$411 i.irives clear indications of each heart-beat, but the spleen curve shown,
the buyer waves of which we shall sj*eak directly, only uudulalitMia
a»
004
THE MBTAllOt.IC PROCK9SK8 OF TUB BOUr.
due lo tbe reaj>irMory movenienis ; him] (hew, iilwiiyi wry flight, atv
times not vbible. In other norcU, th« »|>lccii iloo* iml cxiuux] vrith lb
Incraaae of blood-prcMure occurrinjt in llie splenic urtrries wrttr e«ch Ixwt-
Imil ; ibis niar be due to tlw mueoulur coat rMisliD|i; espntition. Murravtr
vihea llienipplf of blood to the^>leea ia wholly and ludilciily cut i>d', ■> br
cbiinjiinfc tbe aorta, tfa« spleen curve sinks very slowly, ^howinu titat Uwi
vplLf.n i* iliinioiftliiug in volume not suddenly but x'ery slowly. Tfae pAlk-
way of the hlood through ilic Hplenit: reticulum is peculiar ; ami increase af
d«orMue tn lh« voluniv of tlit- upli-^n means more or lo*^ blood htid in tbe
aplevn pulp, not nccenuinlya rreatcror ban flow of blotxl ihrougb the otj/ui,
Of RiM-ital intorot arc (be i*t^- (low viiriationa of volume whitih, beaidcf
ibe rMpinilatory undiitaiinnx, the Kphtii i-iirve usually shows, na aeen ia tiie
figure. Rhythmic con trad ii>iu> mid cx[)nn)>i>>ns. Ihou^h not alwayv pnsenl.
fmiuently make their nppeannce.eacbcontraoiton with ibi fellow nspawioo
luting in the cai and dog iibout a minute, and n'ciirring nith gmat rrgn-
p»>. iw.
mmm
Kotuut Sruas tvavc wmt» (hw. (Rotj
Tlx >i|>|i*<' curv* U 111* )(<1c«n pum aliouliii tliv rliriliunc •NinlnuUoiv ui-l eipaariona; IM
mumUh <•■*•■ >re doc (o llie mplmMfr nwrciiwau. Ibi lowrr cam ii n>s tilrml iiiiwni aim.
andlh«|i«iiil«(rfi<>i"Hi<(ii nirr* «an«<taadi In ilmi- (•> ilxiiiotiii tufiliit h)nod-|a<eauni wni.
TiM mark* n» ilio ilmr o'lm tola* IihUmu naciadt.
Inrity fof a long time; itnd liiwiiU-* thew the volume variee widely from liflW
to (inK. Tlicre can be littlo dtubt but that tbe rhytlimic vnrialioRs la
volume are due in th«ee animals to rhythmic contracti>ins, vritli ihtcrveniit;
rolniatloaa. of the mnscubir trabeculte nnil ca^isule: tbe slower viiriatiow
are also probably due to tbe saiue cnuso. In many animals th« ooutmcciliiy
4>f tbe splenic tissue is riiowo by the whit« lines of ronstrictioti which nppoar
when tlie electrodes of an induction machine in action urudrnwn over i(»siir-
fii«e; and similar tinea may be produced by mechanical iftimiilation with lbs
point of a needle. Ho tliat the spleen iu th(»e animaU may be cotundeivd u
» muMular orf^un, uon exoandiii)i U> receive u Inrgrr omuitily of blood and
now contracting to drive clip l>liio<l on tu tbe liver. WJim t\\v tnii«culnr ele-
iiivulit are scanty in or ulu'iit ('mm the capMik- and trnlwctiU-. th(^ cxjinituaD
and cool riK-ti oil of the wlioU nr/iiti mu^i dcprinl nlntic or chicflv iiu viuia-
lions in (hn width of th<r xunplyiii^ iirtcrii-*. W'f have evidcnie, nKirvover.
that the inii.-wtilar activity "flbcuplccn. whether of tlw miis'-iitiir capsule and
trnlN'ciilii-and artcriiii ctmibiiMol.oniflht' latter alone, is under tbe doniinico
of the nervous syslom. A rapid coiilraclion of the aplocn may be brought
nliout in n diniKlt manmrr by Mimuliition uf the »planchiiic or vagu* nent*.
or in a rvdcx nmniier by (.liiniilation <if the ccnirnl oud of n wusory ucrre;
it mar also be ciiusod by stimiilution of tbe tuediilln oblongata with a gal-
"COSSTITI-KNTS OF Illl.fe:. 595
lonraot or hy im-iuunf lufihirxin. Thougti thvn]»tkr luw unt yut Ikvii
S' worit«d ou(, w« buvc iilrVBiJv nitlici«uitly cloiir lodicnliutia thnt tin- tluw
load thruugli Uto spleen i*. tlirougli th« ngitnrr af lh« Dcrvnu* mvbiimii.
_._.Nid to UKM clinngin^ ii«Oi)«, At ono time a nnM i|iitintilv "f bt<jo<l it
pmiag Ihnxiiib or in twin^' ticM lir the orgiin, niiil th« oietAUoltc cliMigM
wUct il iin<li-r|^<x« in llic IrniiNt urn cv>m|>nr«ti«-4!ly sliKlil. At niiolhvr time
■ brger ouODiil}' »{ lil<>»<l mura lh« orguo, and ia lot ioow, s-i to s|ic»k, into
t^ *f4Mitc T>ul{i, then? Ut iiiiilt.i;go mora profound chani^, and afterward to
b»^K«<i) by lh<- rhyltimic eAOti«oli»n* of Ihe iiiuHCular trabeciilie.
U it furtlier obvious th*t these cbongea going •>□ in the spleen iuii%t have
■a iiBponsnc influenee un tiw chnagas going on in the liver; it cannot Iw of
iodlApRDce In tbe liltor organ, ivhethtr ft relatively muuII quantity of blood,
fvlatively little diaiiged, rtacbee it from the spl<«u. or whether it receives a
i«liUir«ly lar^e quanlity of blood, profoundly altered b; the cbangea which
it ha» UMefK^Ho >■> tl>e spleen pulp.
$477. Thr ehemienl (ttntlil'itnU of the spleen. Besides tlte obemical bodiet
whid) one would expect to find in a rusoular. muscular or^an full of blood,
ib« tpleeo Gotiuiiu bodies, lodged apparently iu the spleen pulp, which K've
it •fxcial chemical characters. One of the inaj.t important of tliase isasiiticial
l>roiei<l of tbe nature of alkali-albumin, holdiu]^' iron in some way peculiarly
ai^wxiaMd with it. The occurrence of this ferru^^iiious ]>roteid, ucouiujiauied
aat it if by several peculiar but at present little uiiderslood [ugmaats, rich iu
carboa, which are partly present iu the cells s|>oken of above and (lartly
dajowU-t in the bfanched cells of the roticulum, appears to be coniMoted sritJi
tlMtfaiDgm under^ne by the hiemoglobin which we sliall {»reseut]y discuss.
Ttu imrgaaio salts of the spleen, or at least those of it^ anh, am remarkable
fnr lbs larK« amount of both *uda and pkosphaies, and the »(nul] amount of
polsih and clilnridei which tliey contain, thus diflerine from those of blood
ofirpuitclc* on |Im> oii« band, tuv.1 frum thow of liloiMl-sernm on th« other,
Buipcrlui|» ihc ntiist Mrikingfontunsof the »plci*n-pulpis ile richnnu in the
*4>^llcd extractives- Of these the mo«C common and plentiful are succinic,
A>rvic. acetic, butyric and lactic ncidfi, inoeil, leucin. xuntbin. bypoxantfain
Knil uric acid. Tyrosin apiuirently i« not preeeot in tbe perfectiv fre^b spleen,
^bmi|^ leucin is ; both are f •unit when decompiwtion has set in. The cou-
•Cant iireseooe of uric acid ia remarkable, espedally since it has lieeo found
w*B ui tbe spleen of animals, fiich as tbe berbivoru, whose urine omluius
ooae.
The riehnev of tbe spteeu in th«e extrac-ives id an iodicatiun of tbe im-
portaDif of the moiabolic events uith which tbe orgaii has to do; but it
will be nton profitable to discuss what lilies on in the spleeu in connection
*ith tbe metabolic cliaujt^H in the ixbcr partji of the hi^lv, in tbe liver for
inat anca. than to attempt t<i Iny down any o'lcalled "functions" of the
■pleen. When wv cmlino our allCDtion to the oplnen ilMilf wu \«&ta very
lo-tle; thus the whole ur^fau may be Miccwtfullr removed without any very
'^tsirioas cliange* in ihv ec»»omy rcMulling. We may rcluni, iberalnre, to
lJi« lUtcuMan of tJi« formation of lh'< bilirubin of bite, and of tho changes
■ncltr|[r,ne by hnmogloliiu, with which as wc xhnll ft* the Dpiccii is oonneoied,
•»«! ahicb. tDoraovcr, has to ilo with the lonnstion of other pigments.
Tbk Formation or the Coxstituh-vw oif Bile.
, S478. liUe pigmenlt. Al\er extiq«tioa of tbe liver no accumulation of
*'** ptgnietil or bile mIia mkes place in the blood. This is well aUowu in
s. which survive tbe ujwrutiou for some coiuidenible lime ; but the same
606
TBBin
PROCKSSKS OF THB BODY.
results have been obtained id birds (gecM and daoka). Them ma \» m
doubt, tli«refore, ilmt Uiese ftubstaoonars formed in ihft liver, bikI BOttiia|^r
vllbdrawn from the blood by the liver in some such irav mm wo faATSKM
reaaon to think urea is withdrairn from (he blotxl by tlw kidney.
Wbeo tbe plasDW of dmilotiug bluMl U made to vouutin limm^lohin
detached horn tbe oorpusclefl, bile |>if[m<-Dt frecjueiitly nuiki-H itM MpinaiMa
in tlie urine. Tbe preaenee of free hgtmnglnbin inajp be obtaitimJ by injact
ing into the veina a solution of hwmDgluliiu or hloo<l niiwle " Uky " by fno-
ing and thawing or by the addition of a «niall qunDtily of bile 'ralta, or In
Hnijily injrctiug into the veitia u ituuutiiv of dtrtilM wat«r or a mhlU
<iuantity of ether or chlorofonn or »f bite miIu, all of which tend to " break
u{>"red corpuaclea and wet frc< bicmoginhin. A similar mult oocon io
]>oiiM>niDg by certain dniifs, Ruch as toluybndianiinr. Under theoe einun-
alancea not only dm^ bile pi^mi'nt. binrubin. make its nppearanoe in tb«
urine, btit tlie quantity of bilirubin secreted by tbe lirer ia Incruued.
Obviouvly (he nreMnee of difaolvcd hvmoKlebin in the plnsma of the blood,
and, prexiimnhlv more etpecially of the olood renchitig iho liver by the
porisi vein. leaHii to nn incn-iiwd formaiiun of bilirubin, which takea'pla<«
ID luch II manner (linl the wliole of the bilirubin so formed does not )ia»
into Ibe bilo but |inr( is retnined In or thrown back Into the circulation and
apneara in the urine.
We have already raeniioned tbe cbenioaJ connection between hatmoglofaio
and bUirubiti. Hjemogtobln, aftfrr the delachmeot of ite proieid oompootM
beeomee btematin (CKl(„N,FeU, i. Uy treatment with aulphuric acid tr
otherwise ($ 352), hnnjaiio mny be d«prive<l of its iron ; and ibia iron-ftw
lupntalin (aomeliniee called harDiato]x>T]>hyrin) u laid to have the ounpcij-
tion (LHsN,0^. diHering frtim bilirubin only in ita oxTKen and hvdroM
(C„H>'.0,+aH,0— O=.CVUN,0,:i.' Moreover, In old blo<«l cloie In th*
body tbe hwinoglobiii of the clot bee»m«a in time transformed into un Inia-
frei- body which baa been callc<l hwmatoidin, liut whleb Uith in c<iii)|>ueiiii)D
and reactions appear* to be idenlieul with bilirubin.
Thoe tereral raets lead iib to the oincl union tbut the bilirubin of tb» Ink
ia eimpty acinie of the harmoglubin of tbt; blixirl tranaformrd l>r the ifaroir-
ing ot)' of lis [iTOteid nud iIn Iron cum in men tn. Tt i:< nuturnl to auppoee that
the irnn^f'irniatioii take* place in.nnit ix iiflVclcd by, iheagtney of the hepatic
celb; nnil I hi* vlrw io vupporied bv the fuel that the hepatic rvtk are dianv
terized by rnntnining ct^rlajn jH-nilinr inm conipouml*. When all the blocd
iii carefully waohed out of the liver by Injection through the bloodveaoeK
by which means the remuiuiiig bite i» got rid of at the same time, tbe befwlie
substance is found to cuniaiu a small quantity of Iron, sufficient to give the
celts a diffufcd dark color when treated with ammonium sulphide; lb*
exact amount appears to vary largely, but the eausea of the variation hat*
not been delciniined. That this iron is in organic oombinallon ia indicated
by ihe tact that «ltb iratiiasium ferrocyanide and aulphoevanide tbe bine or
red reaction ia not obeerved until alter treatment with bydrocblorie add.
Apjiareutly there are several snob coni{><iuiida, of a proteid or nf a uuclcto
(§29) nature, Irom some of which ihe iron is more easily removed than
other*, and these compounds appear to be preseni in both tbe c«dl aubMaare
and (be nucleus. It utill be remembered (§ 244) that bile contains a dit
linet Quantitv of iron, which proltably lina its origin in Ihe iron thus act frtr
from oirntoglobtD and retained in the liepatie cell; but il does not foll>i"
that all the iron thus set few mak<v iu way into the bile; and, indec*!, the
(juantity of iron discharged in tlie bile iu twenty-four houra is much sniallff
■ Doubttot tbo (Mmuta te WMraUa ginn In 1 3av
TOK rORUATIOK OF THK C0S9TITUKST3 OF BILB. 697
innc the »afiie period. Ap-
!mv6 N<ime iillier work lliaii
I
I
I
chftn lli« <|uaiitilj' cnloulnle'l to be set fiea in the formation out of hnmo
glolrin i>f th« quantity of bilirubin ilinc-bargt"
|Mren(lT the iron cumpouExiii of llir hi^iiutic ■
the Miuple (tiMhargc nf iron into the bil«.
Ttie fan ti»oulion«c) nbove, that th« pnMnoeaffrce liamoslobm in the
bloix] Iciiila not only to nn increatn of bilirubin In the blU, hiil uIho to U»
praieiKe in tliD urine, offera aooM dilliciilliM ; for if tliu bilirui>in be; f^irineil
out of hieiDOglobio by nn<) in th<^ hv]""'^ <^'''- ""'^ vwM cxpwi to fiitd ibnt
the whole of it passed into Iho hUc. iiad thut it could not appuiir in the
blouil and »o in the urine uii1«h reahMrption from the bile powMigoa, due to
ukMiinictioi), took place; and tJiero is no evidence of any euflicient ob»truc-
tion occurring in thete cnet. Indeed the presence of bilirubin in the urine
in them omm has beeo argod by some as an ar>;unieol that bilirubin m
funnel! in the bloid or at least ebevhere than in the liver and is simply
L-xcreted by the licer. Not only, however, aa stnCed abnT«, is thore no Hc-
uiimulatioo of itile in the bioiid al\er exlirpatioo of the liver, but that oper-
ation prevents the apiiearance nf bilirubin in the urine as a contteijut^noe of
iliu prcecnoe of free hxino^lohiH tn the blo<Hl. The phenomena in ijuesiion,
therefore, do noit disprove lliat tlie bilirubin ia furme<l in the liver; tbey
iiiay be taken, however, to show that that funnatiou, viewed as a Decretory
act, U peculiar, since the hepatic cell appean under certain cireum^tances M
discharf^ ils product of secretion into the blood or lymph as well as into the
bile paMaire».
M78- We may awun>e then that the he^Mtic cell ha* the power of split-
tin? up llie li.-vmofihibtn brought to it, and of diiichar^iri]^; part »h biliriditn
H'bde it ririuinn fir a linit; the imn uoniponent in some orjifaiiii- riiriihiiiation ;
and, if we fiirtlHtr a^omt! llial it work* upon thi* (entire htcmiiglobin we may
premme ihul it niakts mitnc niilMX'iiuenl um; of the proteid cnm^Kinent. But
are we juxlificd in (umiming that the whole work u dune bv thr liepntiu
cells? Are we to cont-lude that hilinihin is maniifiichir^^Kl by *(ime act of
ll>e hepatic mils which includeK not only tlie conversion of hiemot;l'ibiii into
bilirubin, but id]U> the extraction «f the hicinoglobln from tli<' red corpus*
clwati ibMMt are Htreniuing slowly through the lobular hepatic cnpillarics in
oli)*n contact with the hepatic ooll»? Now, as far tu we know at prownt,
hwnioglobiti can only be set ftve by lofans of a disintegration of tlio CJirpus-
cien; we have no instances of * corpuscle parting with some of its hamo-
?;lobin and proceeding on its way otherwise unchanged; and we have no
itstolngicnl evidence of any disintegration of rc<l corpuscles in the liver cor*
reejwnding to the formalioo of bile. Xor can we draw any conclusion from
the result of a comparative enumeration of red corpuscles in the portal and
hepatic blood, for tiiesc are loo tni^ecure to rest any conclusion upon. On
the other baud, as we liave just seen, the nreeenfe in the plasma of [he blood
of h.-etnoi^'obin in a free oondilion is |>eculiarly potent in exciling the forma-
tion of bilirubin. The evidence, therefore, \a very strong f^kr the view that,
as far as the formation of the greater )>ort at least of the bilirubin is oon-
rerued, the action of tlie hepatic <^ell i* limited to converting into bilirubin
the fret? h;enM>globin oflrreil to it by the portal blood.
By wlini means, under normal coHditions, ia the presence of that free
hwinoglobin ttecured? We liave seen reiuMin (§475) to omelude from liisto-
logiail apjtearaneee, that a certain number of red corposelee iinderfia change
in tito uplenn pulp ; and it seems natural to inlbr that one duly of the spleen
fa to Mt free lucmoglobin fnmi the corpusclcn aud thus, thrduxh the splenio
veilit and so the portal vein, to supply tlie liver with material for bilirubin.
But ihiv cannot be the only source, since the wicnitton of bile continues
; aftrr CftirpatioB of llie ejileen. There mtist, thcroforv, l>v other ivgioDS of
18
TOE METABOLIC I'I10CBSSE9 OP THE BOnT.
lli4- liiiily in uliicli h vimtliir i'linn([e of red ctiipUKkB is (C"<i>R on
\itra Miirgi'KU'il (liitt llie red uiimw uf Imiikh is ane of tbrav ; l)ui fur
iiifnrniuiioD on l)i««e jiointit i* DfMkd.
Amupiing that iimirr ixinrnl circumMnDcrs the chkf supply of niMm)
ti*r ibe Bianiitkolurc of biliniliin i't)ni<v from iIk- i<{ilc«t), ihi- iiiirritinii mriftt,
Pi^-ii llint nitilcrinl It'nvc iIm* xpli'ii) in iW form ol Iiieini>)[M>itt, ur tlwsi ibr
«|il<-i'n Mill rurtlifr a"ti<4 in l)i<: imitltT, tiy t-Ki'ciing ioni« fKliitiiiiHrv duinc*
in Ihc liiciDOglvbiH. by convrrlin^ it I'lir inxlnnri' inUi a nrolriil Ir*« fuenuta-
lilie lui'h? And iheranM! (iiiiTiion imiv Iw aUi applin^f l" llic ollior ti«u«*
which may fimilnrly piovidv niittcria). Our knoulrdgc iit At pn-cvnt ifusA'
cipDl tu furui^h a eulisfiicloty niii>«i'r in mi< )i ii ijumltoD.
We tns}r th«n go to tar as to My that tin- l>ilinil)iu of thv liitv t» drriid)
from (be buniugTbbin of lli« blood, and ihal ihc lafr flagvt of ibe irauf
fomiation, incliidiD); the d)«cbarg« of the iron of tbr b:ininliii coii>|K^irr>i.
lake piflre in and by menus of the beraiic rell ; but much l'cy<ind ibio to k
piMcnt iiDceriaiD. It oiunt be r^menibertd loo that, though alli-r extirpalion
vf the liver ud accumulnlioii of bilirubin takes place. shoMiti); that llx
bilirubin is formed br the liver and not eleewl>ere. ytt the whole (■bnu|,>r
from red corpuscle to oilirubiD may occwionally tnhc place quite afiart livia
the liier, as ^l^o»^ by the prewnce of hKiaatoidiu in old blood- clots.
$ MO. The j'ormalioH oj th« bik-atid*. Aboul ihis we know biIII Ich.
TakiD^ fclycocBolic and taurocho!ieacidBa» I h« typical bile-acids, recacDiib|[
(!i2-lti) ihal tlitAearire Irom ili« uniun of cholalic acid with glyciii auirtauria
tctpectively, and renM^nihcrinj; lhal lauriii is found in •erersl tipMic^ and
llial i;lycin (»c« $ -l'J& though not an artual «inBtitucnl of any of iIm' (iNaica
niusl c«rlaii]|y art»« in li»ue ni^lalKilirm. »■• may c^mclmle (hal ibc cblef
work in lhi» n;H{iccl of the ht^jwtic cell i« to iimvide cliolatic acid, and In
rA>K( the conihinutiun nilh glyctn and laurin, tlinugh |)ci(«ibly Miniv ntoouut
at either one or t)ie other ol thcxc hoilin niny ba tuniinheit by thv bvpntk
aabuance iiaclf. Aa to bow cbolalic acid ariwoH out -jf tbemetnlxdwoi of the
h<.')<aiic cell, no know im more than «vc d'l about tlw- formation of kreaiin la
niutwlu orof pqHiin in a gaftrictyll. Wc are eipially ignorant about tbr
origin of glycin and tnurin, and cannot explain whir in i>uc animal glycti-
cholic, and iu another tnurvcholic acid i« prominent lu thi> bill-, Ihoueli lb*
iwu bodita, a» abown rspecially br the prevoiice of sulphur in the taunt), arc
widely dinertnt. It ha« been ob^rvcd that ibe pieeence of bile i» the ioK*'
line ecenui to excite llie liver to incrmsed biliary- action : vince the bile-add*
ore rapidiv changed in tbe intestine and the cholalio acid speedily ali«r«il,tt
f«vnu probable ihat the increoK^l biliary nctivilv is due lo lite ab^irtitiuD vt
the glycin and taiirin rps[>ectively. From whii-n we may conclude tliat llw
pteM'uce of lht»e btidies rtita up tbe hepatic cell to an incrcaw^l fonnatioii »(
cholalio acid.
fi 481. As a jfenera) rule the ri>rmation of bile-acids rune parallel with tbr
fonDalion of bile pittuent. an inct«i»e or decreaae of bile m«auin); mi io-
CKweordecreueof l>o(h ooiistiluenta. But there are some fnctv wbich smm
U) abow that the two action* may t<edi<-HH-tak-(l. The couililion or MyniptiMK
known as "jaundice" ia ewviiiinUr an excc^n of biliniliiii in the blwd.
whereby the lisBun auch »» the vkm, and \\\f tliiidi mich aa tlte nrinit nrr
colored with tlw yellow pigmt-nl. Inmuat of the niahidinxif which jauiKiin*
IN a svBiptoiu, there im fvidcn<« of nn i>b«lrucli(>() t(i tho flow of bile throuek
tbe bile pataaf^ ; an<l the jirtwiici^ of bile in tlie bloud, and bcticu in tW
tiMdeii at lar^e, iti in aueh c»M« due to the fact lhat the bilr after accretkiD
by ihi? be|>aiw celU n reahxirbwl fnm* the WIe duct* {*vt> § 257).
But in certain nvra nberc jaundice is a promiiicni xvmptom, no evident
■>f any obstruction whatever to the llow of bilr can bi' obtained. Thia in the
URKA AS^» OH MITHOOBKOU9 MBTABOHSM.
590
car« in (lie jauni)ic« of yellow (ever iiiul of a peculiar n1lk<l innlady kuowii
I
»
M
Jte veil
air
I'tiv of 111'
iIk-
there :
1 evi-
I ^
■ or I
wk
il«Dce of ail accuiuulutiixi in iti« UIumI nr dsonhert! of Itiltiaciitii as iWri! ia
of bile pixuieni. And in the iilM<Hir« nialtnly knonu as simpli! ur iiHojual liio
jaunijire, in nbich thuii^li tlu^ iiimiomiMil coiKlilioiiii iirc iinkiKiwu liicn in iit
least uo vigil of iibHlrLKrlioii, tbu \iriiii; iIkhi^Ii Igwlcd nilh bile pignieni i»
Mid to ronlniii iiu bilc'iicid".
It baa itrt'ii KU|ipiiM'<l ilmt llu»c cah* nfli>nl proof lh«t the bile miiy tie
fonuetl olMfniii-ru lliiiii in ihi; livt'r. In fao; huwi-vcr, of tho nrguinciita
hr(>ii|j:)il foruoril in llic prcwiliiig paragni^>h9', lliry ouinot l>c nccupted M
pnml thul ihi* iiorninl (ormiilioii "I" bilirubin i» so oarrinl on ; nor ia tlivre
Uir cvi<l<-(ii-v lo «hiiw ihnt in tWc nu>c» bilirubin is foriucd on a plim wholly
dilwKail from llu; noiniHl, AuH n tiiRcrcnt vxplnnaibn seninii gKMtihlo. Wit
may m\i[«<i>v tbnt in ibno cutm the molubolic arlivity of ihc bcjntic ccll« m
TOoiJitin), iiii>l. further, ao niDdilie<l lu while atfediiit^largrly the fornintion of
bile mIIii ami ollwr functions of the bepatic <-ol)s only pnriiatly to nlTc'l the
fonniitioti ami di»ch.iiy« of biJirubm, to aflect indeed its dii^bari^ rathur
than ils formation. That in ncute yellow atrophy the ftincliuns of the cells
an- grcutly alfected is not only indicated by poM-iiiuitein histological appeor-
nnces, but in also shown, at we shall itm^iilly have ocoasion to point oiil, by
the iiubstilHlion of leuciu and lynuiin for urea iu the urine. We have
already ouiumenteil on lli« faet. that therv \» something {leeutiar in the action
of the hepatic cell iu »«<:retin)c bilirubin inasmuch ax the bilirubin iu formed
may, UDoer oertaiu circuiuMamr^, in iwrl |ia^ fmni the cell itadf into ihe
blood instead of into thi- bib: piivxuv^fji. And no niuy |wrhap.t explatii the
jaundice of tbL> diwiiKn umivr dix'UMioii by HupiMwn^ that the morbid
obaitKes of the bejwlic wHk, while arrmtiiiK thn more diflivult luclaliolic
labon of the ccIIm, mich m (he formation of nil^ucidit, ilo not put iin end to
the lih'hlpr tiuk of turning htcmoglohiu into bilirubin, though m> aHL-vting that
priH^rx iiIm) that ibc bilirubin |aiwws into the bliHid innlcad of into the bile
paaMSRi. In <>lli«r wunlit the formation of bilirubin !■ an act indrncndciil of,
aiu) (lilli-n'nl from the more i>rdinHrT secretory nclivity of the cells.
$ 482. The question niny be asked, 1» ihu secirlion of bile independent ol',
or in "ome way or other connected nilh the glycogenic ncliviiy of ihe cells?
~^g this wc cunnol at prt^eot give n delinite answer. Iu sonie of tbu invcr-
irala ihc cells in the orftaii, called a liver, nhich manufacture glycogen,
are dietinct from thoao nbicb secrete bile or other digestive juices; and it
taight be inferred that in the vertebrate the two actions though lakiug plactt,
OB the certainly do, in the same cell, take place apart and distinct. There are
Ikcls which seem to indicate that tlie two are iuiioiately connected ; but we
have as yet uo exact knowledge coucernin;; the matter. It has been urged
that the portal hlood U diivAy coni.-erned with Ihe formalioii of glycogen,
and (be blomi of (he bfr)ialic artery with the :«eretion of bile ; but there ia
no sdeijuale support of this view. It must he ren»i!atbere<) moreover thai,
in addition to ihe furniiilion of glycogen and the »ei'retion of bile, other
metalxdic events. cf[)«:iiilly aiR-cting proieid or at Iciua nitrogcuvuus ctiii-
atitueiitE of tb« body, nro also taking |ilace : and to ihcau ne must now turn.
0» UUKA A!lt> ox KlTHOUEXOtn UbTAW>I.I»U in GKXUItAI..
rj488. We have seen thai nilrogenous proteid material in some form or
ntberenton into the conitx^ilion of all Ihe tissues of the body, and wc have
l^irtlHT n^vn that it in m> conspicuously and constantly present wherever
living buImiaiicm are munifeetiug vital euei]pcs aa to justify the concluaion
THK 1HTABOI.T0 PROOES9R3 OF THB BOUT.
tlial ibe cbansw vliioh it uuilerf;<i>ea are in iHiini; way encntitil to the maiti-
ftaUUion of lon*e en«rKira- ^Ve liave wen, it in tni«. reRMn ti> iliink thai In
BOoie tiasuee >t leaM, in muwie for inalaiicv, ii largr part iif the cnrrgiF' Ml
tne during ectivil)^ prei^xiMed as taleiit enervy and liait itn imntedititt nunrcv
not in pmteid (nitrogenoui) but in iM>in« otJin- (MinxUtiHiiiu of niiittclv; itni)
Indeed, aa vre sliall see later ixi, the ([ml«r jmrt of tin* wlii)k i,iMrgT i>f the
body muM be iv^rded as lh« eotfrny of mrbiui ottinii'Minds and ni>l n^
nltro^n compounds; but Ihiii t* miiu< ocuiMxtmi with the vi«w ihnt proU-iil
iiiiitrrial ill mnni- way or utbi'r a^Moiiliiillv int<'rv(>n«» in. wo mav |HTlin{w go
•0 ftr w wi way dim-iM, tho cltan|{ca by wltich in th« body taergy is sd tnt
in tb« tteculinr wav which wc npttik of U living.
We nave tu^ni tfiat at all rvi-iitt tli« grMlt«r mrt of ihp pnHeid material nf
Um) foot) rntvn the I)!(hhI ax protrid nuiKrial either ns peptone »r in suax
other form, tuid ih currit-d a« proli-id material to the ti«iie».
Wc linvc mvn that lh<> iiilrogi-n of nroteid innlerial IcavcB the body f»
largely in tlw Corm of nrt-a. that the other nitrogenous exfrelioaa may for
the time l>r left i>iit of eons deration.
And lastly we have wen reasnn to ttiink that this urea vrhieli leave* ibt
body in urine is brought t? ihe kidneys ne urea iu the blood, the kMoey*
Ihemerlves apparently having no Kiweial power of forming ureit out nf wmr^
tiling which a not urea, hut only euntributing l» the geueral slock nf ami
by virtue of their own proteid nielaboliiini. We have non' lo study ihf
little we know concerning the steps, by which the pruteid material of the fowl
and of tlie body is converte<l into this urea of the hln«d. which is the Mmiree
of the urea of tlie urine.
S 4M. In Ihe first place we may lake it fur granted tliat the urea carried
to the kidney in the blood, bad an aniecedent in sumething which wna not
urM. We can hanlly iHipp(»e that the proleid <'oiiatilueui of Itvtirg sub-
stance, when in the course of 'i\* nieiatKilinni it ceiUHW to be proleii*, bmlu
up at on<% into urea and into n«n'»itrog«tnouii bodies. All we have leamt
COM lo show that what we call melahi>ti>im is not a singto almi|>t change,
but eonsiet rwentially in a wriiv of •liangiTit ; and we may safely conclt^
that proleid malerinl in Wcomini^ iirnii pnMW lhro«igh phases in nhich the
nitrogen exists in elicinicwl ci>mh mat ions distinct frum proteid material on
the one haikl. and urea on the other.
In the second place it is exlren>ely proliable that the seric" of ehanffea by
which pruteid mnlerial l>ecom«s urva, is not the same in all tlie tissues and od
all Dcosions. We should naturally exjieei to (ind the proteid material ful-
loving diflerent lines of inetalmlism in diflerent plana or under dilTerenl
drcuDMtancee. the dilf^reol lines all converpng lo the same body urea, bfr-
cause for some reasons or other urea appenrs to be. iu the main, the mmrt
GonvMiient form ui which the nitrogen can leave the blood and the bo<ly.
Wi! i-hould accordingly eipcot to tlud. on the one haii'l. various nitrogv.
ww» hodiis n^ulting fntm proleid metabolism in various parts of the Iwujr,
and. 'in tlx? othtr hand, arraugeineaita by means cif which these various bodies
were n-ilm-Ml to Ihr niinmon f<inn urea, preparatory to tlietr discbarip' firim
the body l>v the kidney. An luiual oWrvat ion ns far an it goea SUpjiorta
this view, tli<>U|.'h iiur kuowtcdgi' of the whole miitti-r ih very imperfect.
§ 4SS. We uiity turn our nttenlion firxl to the m<'lali»1ism of tlw skeletal
muscle*, siniv tlii.r*e n>pr«M'nt, as far as nieot i|Uflntiiy »■ coni-erneil. bv far
the greater part of the proh-id capital "f l\w body. We miiy siilely infer
that they furiit«h a large part of itie un-a of the urine ; though uiitloubteilly n
small iiiiLw of (iwue might by reason of its more ra|>id ntetaTiolisro xxirk i>ver
a greater c)uantily of proleid material than a much larger mass with ii slower
niutabt^isin ; yet we have no reason to think that the proteid melabulism ttf
ON tTBICA AND OK XITROGKSOt'A MBTAB0LI3H. 601
skdcml mtiKle. obscure tlioii^h ii ■« in iu oattire, is no slow at to neiitrnlice
\ht |>rohit)>l« eir«ct of (lie ^rtAl bulk of mtiwlp existing in the body.
In (Jralinx miih die eWnibitr}- of inti>eli> (5 62> we »»w that nrvti, bbtc \n
the cxreiXiotiiil insiaiic-oti .if (HTtiiin nrliUginnus fi»hM. wm coDtpicuoue by
its nbsence from ibe exlnict of miiHclvi. wh«rcii* a rerv appredftble ijuaiilicjr
of kmttio wail Juvariably prcacnt. and, inilecd. n-iu ll)c promiueul nitroge-
nous cr\-atu[|iii« <'Oti*tiUiont of that pxtrapt. It scorns <liliiciill to rraiM the
concluHiou liiat kmuin is the main normal nitrogenous product of the inuU-
boliiini of Kkvlrtal miisclca. If we accept this view, then upon tbo fitcl of
the prexrnci? of krratin in. and the absence of tirea from, th« muscle itm-If.
yr« may twsc the conclusion that while the muscle pruduoee kreatiti ai< iia
nnUH'colenl of nren. ihe kreatin so prortooed is converted into urea in siime
part of thr body other than the muscle iuelf. Kreatin oa we have already
wen may he easily split up. and we may prt)hably wilh safely awume is split
Dp somewhere in the body, into ure* and MrcoetD. But samocin does not
appear in the urine as such : hen(^4> the t!onv«rsiDn of kreatin into (pari of)
the urea of tlie uriite entails an wdl the further i^mventioD of saretwn into
urea. Now aarcnaiD a* wm have aMn i* niothyl-{[lycin ; wn may reheard it for
our present )>urprM« as *imp1e clycin. and heniM! ihn total cimcinrsion of
kreatin into nn'a entails the convomion of ^lycin into urea. Tbi*. however,
I doea not offiir any adfliiional diflinilty. since wo know from direct oWn-a-
I lion that slyctn iiitnKlnoil iiiUi the alini(nitary i»nnl does not reappear as
Lwpb in tAe urin« hut pr>Hliio>sa curreaponding increase in the urea of the
Vinne : fmn which we infer that glvcin aluorbM fmcn the aUnientarv canal
W ft somi-whero in the bo-ly i-onverled Into uresu We shall speak of this con-
f ven>i<in hilcr on. and nhnM then k« thnt. as far as urea is eoncenied. glycin
(amido-iuvtic acid) and Karcwin ( methyl*(-lycin. niethyhamido-acelic acid)
underg<> the same chanKe, the amide moiety in each case being oonverli'd
(into urea, while the non-Ditrogtsnoua moi«iy is oxidised and thrown xlf.
.Meanwhile we may stale the eniMlauon at which we have provisionally
arrived, namely that the nilrngenons metabulism of mtiw-le probably gives
rise to kreatin. which in ^ime |>arl of the body other than iiiuadi- is proba-
^y split np into urea, ready fur excreli'w. and into aarcwin which also,
Hnewhere in (he body, is fviriher onnvertcd intii nrea. Xntl hearing in mind
the large maw of the skeletal miiacleii. we may furlhiu- conclude that a large
portion (if the ut«« leaving the body by the urine is formeil in this way.
S 4M. We must not, hiincver, leave this statement wilhrnil referring to A
tlEfficnlty. Krealinin as wo have seen is mt frequently found in urine as to
be reiiirded as n normal conilitutinl, nt all vvenis of human urine; and
krentinin is as wc have eerii the urinary tlirm so in sjwak of kn-atin ; the one
body easily cliimgee into the olh^ir bv the iw>umptiou or removal of H,0.
Tliis eiiggesis the question. Is not the krenliain of urine the reprmeniative of
Ihe kreatin of the muscles, wbicli la thns excreted directly without iindurgo*
ing the ohani;e into L'rca just discuswd? In answer to (his «-c may say in
the first pla<« lh»t the ijoanlity of kreatinin in the urine, though variable U
aniall ; we may put the average at about 1 grm. in twenly-four hours. Now
mttsole conlMiiis from 0.2 to •) 4 per cent, of kreatin : and ihla. taking the
total muM-le of (he body (to say nothing of other sources of kreatin which
»we shall mention prftiwQtly) at about UO kilos, would give 60 to 12') grma.
kreatin as preanol in the muaoles of tlio body at any one moment. We can
biinlly supp'Nte thnt the melattnliatn of muscle b ao slow m oat of this stock
only Ut provide the 1 grm. of krealintn in tweiily-fnur hours. Moreover,
the kreatin in urine \-aniMhe* during ntarvatiou, h very markedly increased
by a diet of tlesh which ronCain* kre.itin. and is not increasuil either by mus-
cular exercise (which, however, would only indireolly aifect nitr<>geoou8
■ cuiar ex«rc<
003
T t J K
OF THE BUDV
mrtnbulwrn i>f niuKcIr) nr by *tu-li iMiicltiioiu, ferer fur ioHtancp, as iioMI
incrcHM tliu uro of uriiii- by iiicrraxii'jc tbfi nilmf^uouB iuc(Al>i>li#m of iiHit>
eUs. We infer, thcnforr. [|iiU tlio norm*) prvseDoe of krratinin in uriM >•
diH to the direct nd id in is t ration of krcMtiu pnoeiit in a inomiitl) flroh <ti«t
■nd has ootliine to do witii the miMcitbr mvUiboliiiro of tii« iudivitlual ohu
is K<-reling ibe KrcntiDin iu lib urine.
The fact, iMWever, ttiai the kmiliii prcw-nt in tl>u miucte of the fu<<d ao<i
absorbed from (h« atintenlary canal don not uiidcr^ a 4-biu)g« into urea but
» excrrled hh krcalioin, thul i», virtuallr ii» kn-atin, warnn iii to Iw vtuHul
ID adopting' the conduniiiti arrived at afiovo that th« knuitin produced bjr
muBciilar tti«tH bolts III ill the livio); body is a oonapicuoue aDt«cci|«nt of (he
urea of llw urine. It in difficult lo see why krcaljn pnwiiig into ibo bluud
of the capiilarita of tliv jnutulc shouhl be changed into unm wnilv that which
pooaeB into tlie enpiltaricaof the portal system is not; for rwuona which will
MttpDareut pr«Mutlr we Nhoula ratlker expect that the latter Iwing; mure
direcuf expoaed to the iiiflnenct of (h« liver would he more r«iidily and nrnre
emnpletelr converted than the former. Iuil«^l, the (|ue(-tiuii liiicv* il^lf
upon iw. Is kr^uitin arti-r all the niitiiral inniii prfxluci of tin- iiitrogvuwa
metaboliMi) of lunnclv? li it pontible that in the uorinal mctdholtcin of ttir
llvinz muwlo the nitrogen kavec the oiuwularwihwiuiof and |uu»i;e into the
blooJ In another form, as twme BiibrtAmw not krcHlin, and that it is ns ik
RiiiKvIo <li« that krcatin is l>>rm«d, juM as the wltd inyioiii U uuknowii M
living fibre but makes ita ap|jcaratice in a dying one? We have no inwitin
eviitoiii'e, liow<.-vcr, (hat this is ao, and mranwhilo may (xtiitiouo to su|>part
that krciitin is formed, and that in conHMiucoi-e krcatin i* n i?on*piini<iui
antirctvlcnt of urea of the urine; hut we must not rcgan.1 this im proved.
S 487. Our knonledfce of the nietnUiliHin of the nervous tiwuco is, ua ws
have seen, very imporleot (^ T'i). but the presenL-e of kreatin in (he ccaunl
nervous system lc«ad us ti> infer l)ia( the nilri>t;oiii>u8 mecahulism of Uto I,**i|^
aubeiance of nerve eelU and of the axiHeylindtr of ikervetibrvs. ta i^Jm
brotid leaturea )denti<al niih that of mtiM-le autHtancc. The mass, liowt-vo^
of the nerve cellii and UKiK-cylindvr* of the body, uU put tO);«llier. U nnisil
OOlDparKl u'itli (he maw of nkektal nium-b: moreover, the ener^^ ^t lrv<' >>,v
(be metabolism of a ina«» of nervoun innllor llimi^h " higher" in (|ualiiy «
leas in iiiiantity than ihnt wt free by ttif nH-lnboliKm nt an eijual lUMti ot'
iBuacle. or in <>thi'r wnrdu ilN ntriaholiani in ]c*k rapid. Hi-nii; wc nmy prcibi-
l>ly cuntiidcr the niclaboliiim of the nervou* sviiivm as a men.' ndditlua to
that of (ho iniiM^ular lyelem. at \vwt as regards (he })oint on which we ate
now dwelling. Tkc amount of nitn<getinug mDlnboli^ni faking place in eon-
ncetivn lUnue, cartilage, hone, and the skin is probably still IcM, and lor tiui
present purpMHf nvnis no special discMKSion.
^ 48B. The nitrogcnons metabolism of the glands, however, more pwlicD-
larly Ihnt of the liver, does deserve epe«inl consideraiion : and we inny <l
once turn to a (juile ditferent aspect of (he >]ue9lioii in hand.
When tbe rate of ditchar)ce of urea from (be body is oliscn.-eil during •
period of some leD);[b, especially under varied circtimntaix-es, the direct eflect
of nitroRenouB food hecomea most striking. W« have already Miid. and shlill
■nin return to the puiut. tlmt niiiirolnr rontra<-tion docit not directly iitcreaM
Ine output of urea; the diwhaic^ of uirn for inniance i* not ncTwurily in*
ereaaed by even great Imdily liiimr. T)iv iulroduition, however, of even*
n\»\\ (|uau(ity of pmicid maierini intu tbe nlimeiKiiry rnnal at oiKe incnaad
(he una of the urini'; und in ihe curve nf the diselinrge of tires in th'
twenty-four houn i-nch lui-nl in followed by a con»picuouB rise. T1»e nbforp-
lion of prctteid uialcrial froni the nlinienlnry canal is followed by nn inini<^
diate pruportionnte inercafc in tliv i^uautity uf urea which is ^«cmed hy '^~
nENOLS MKTABOCl
kidtmv. am) Ihnl nf w hav<t *ttn mnuw iin iiiciimM! in tla- iir<.ii hmnght to
tbe kiAary hr the rciinl iiri-ry. Wbnt » Ihu origin nf ibii- mtilitiutiul iirt-iit
Two vipwa prrfcnt iheiiiMlvi-n, On t\tt> nav hmiit since wiiui- [MirlifjD uf
Ihe (iroirii) tnikluriat nf pvcrj- nn^l. at hII tvcnis of rvvry iiccrtvHry mrnl,
B*>*»Ui n-(>air tlic )>rutci<l wmIc (-i>ntiiiui>Ily going ftU in the pnriBoJ'thv bod/
whtn pniK-iil nict>lH)lif<ni i* lakiii); pliicf, ne mmy supjxxc ihnt the pmitDCO
■^r an extni ')uaniity of proteid malcrinl tlirunn upon tbe blooil mim ihv
'■••wl Bcl> M elimiiluH to iW ti»i>i]ra, lu lh(> niiiBcles for insliiDcv ii« well ns
'Mtirrv. Blirf ihem up to ineTCHMH) ititmin^iioiis mvlaboligiii nnil lliiis prodiii'es
■1 iiif r«aw of vneriiy, i-lii«tlT if not eictliifivcly in (he Ihrni of Iicat, atcoDi-
p(Ui)«d by an incre*iM> of tl)Q antercdrnts of iifca and so of iires. In otlicr
Worda tbe increaee of urea in quratioo is tbo result of an iDcrease in lh«
Btrfleral nitro>;raoua metabolism of the body.
On th« other band we niav suppcee that in order to prevent ibe whole
l>o«Iy lieiDjc eDcumbered "iid il, this exrew of proieid fimd maierinl ia, in
•OOie special |)art of the bndy. aplit up into a nitrogenous and a ni>n-nilrci-
|t«rDcius ntuiety, and that, while ih« latter ia stored up ue fat »r KlreoKen. the
r«>rnier m at imce coo\'ett»d into nr«a and ([oi rid if. We liave already
< $ ^4U> »een ibat a step in iliis dinciioD may take place while the fund U as
A-rt in tbe alimcuiari' <.-anal: we have secu ihni punrreiuic jiiii-e may rarry
pan of the pmteidri nu which it aits beyond tbe MAjte of alhtinioae and |)^p-
Xontt, and rtduco that part into Ifuein, tyroMn.and mlivr lii)di««- We do not
know. n« ne have idn-ady Mid. In what extent thin more iirufound dijiieitioii
btv iMincivatii.' jnirr ilocx nctually tiilcf plnn- in tlie living Ixidy ; it may take
f >1ni*v I') a vrry kH^-IiI cxlrnl and it may under <'<'rliiiu (ircuini'tuui:*^ take
|>lai:v to a eoDHidi-nililc extent. But in anv onir it ilkiiitratm the wnv in
^■htcli n Krmewhnl ziimilnr dieruplioii of priiteid material, a disruption wliicli
niav l>e hmadly di-M'nlK-d as a splitting titt of the proleiil into a nitrugeitoiis
mtna a non-nitroct-nouii moi«ty, may lake place somcwhen in the body and wO
ttad U> ibe sudocQ furmntiou uf tome antecedent of urea. The auteccdcnt
SDkT be leuciii or may be some other body or bodies.
In support of this view may be urged the fuel that such bodies as leunn,
xtycin, B(paraj:in. nod many oiliers when introduced into the alimeolary
•««iial are transformed into urea. When these bodies are adniinistered in
Dut loo K^v*' quantities they du not reap[>esr in tbe urine, but the urea b
proportionately incieased,
i 468. We liave >eeii rernum l» think tlml tbe proieidi uf u meal are
■btnrbed nut by the lacteah but by the portal bloodveswh, and such bodies
Si Ivurin probably take tbe same eourse. Tbii* heinK yu, all thtue bodies nan
thfuufth the liver aiitl are subjecte<l to sticU iutlueuof^ a» may be exertea by
ll» hepatic rellit. Nxw. we have nu |iusiiive evidence that the liver does or
OUi exetl such an urtiuu on jirolud mBlerial itself as lu wparate u relalively
nnple nilrcigra cuniiHiuiid from the reiuniuin;; cunNtiluenlN, leaving these to
ibrni a body rich in earhon ; we have no jHicilive proof that the increate of
Jtmi^id melalioltMn just >{»>keo of as leading to an increnw of uren tiikco
Clar« in the liver rather than in (he liM>uti> at large ; trc may go ao far per-
■ ps a« to Minpeel that it i* largely »r wholly confined lo tlie liver, but we
fa«va no convincing demon >it rat i'>n. We have, htiwever. a convergcnm nf
•vidcnca that the Inol otagi' of iIhi proers^, nainelr, Ibe oonvereivn into urva
*>f SMRinor other pniduci of prolcid metabolism, nfiich though allied to is not
^XBrtljr urea ilom oeeiir in the liver. In tbe tint place, a larg« ouanlity
*^f urra (retn* to lie pnMwnt in the liver of mammals: in this resticct ttie liver
f^rv*«iits II strong eonlnut ti> the mufcleo; in the liver of birds tbe urea la
*■ wpre»enied by urates. Moreover, nhen a stream of freeb blixiid is pMnd
everal Itraes lhroii;;h the liver of an animal recently killed, the pereeHlage
604
THE METABOLIC PUOCKftSKS UF THE BOUT.
of urea ill the blood sn need is found to be d«ciiiledlr inerenaed. Thi«,h«i-
ever, does nut prove timi iiruu in formeil in the fiver, si nee tiie tacnwd
(quantity of urea in the lilimt wliicli bad been eirctilHled raixbt bavc bna
simply urea wlitub liiu) lieeti wa^beil out fWtin tbe liver, wbvre it hwlfir-
vimily been mayiiij;. Still iia far ob it goes il in suggeMtve. In tbewoitid
place, in cerliiiii ciiwm of a furmof diaeaae vf tbe lirer known tu arule iyIU*
iitiY>[iU}' in u'bioli tla- hi^imtic o^li are *o cliitii|i;e<l tliat tbeif fiiar!i<ru»l
iiclivily In lur^roly ilim!ui»Ded, Uic urea of the urine uot oiily undi-r^gnn t
very inttrkt^l dwnate but an(«ani to bu mplaced to a very \mrgp •Mwil bi
ii;iii-m. Tin* fiiet Huegeata thnt Ivucin (iind not for inalAOoe krMtls) wtlii'
chief imnii'jiintc prixlurl of the nitrogiMirouK mctnbolism of tlit bodT, mi
thnl tbo Ivuciu thtit priHlucc<l i« in u normal rI«c« of ifain^n ranvcRM iau>
uru« by the livor. And in ihi« conuuclion it may be rcioarkwl Ihit M
only it Ivucin found In ntuirly nil iho liwue* ufkcr duilb, especially in tb«
tiandular (tMut«. bnt nho ap|Knn> wilh striking reodinma in alnualill
cieompoeiliona of prok-ids, ami i» moreover a product uf deeoiDpmitioa oT
Eptatinifcrous subntnim-e, Without going, bowevof, to far as to ooDclaJe
Uiat loudn is the chief tintoccdimt of uren, wo mny tjik« the nbovt obaatfr
tioD as indicating Ihnt the normal liver has. la tome way or other. the ptnr
of converting leiicin Into iircii. If this be so we mny aim vejiture to lUfpM
Ihnl when inch bwlie? as Icucin, glycin, etc.. inlividuced ioto the alioieiUarf
cnnal apiienr in the urine nt iirca the transfuroiatioa bax taken place inlbf
liver. Ihe body tyrosin which «o often accompantu leuciu, belotigiof; y il
doea to the nrouinlic series, stands on a diHerent footing from leuciu andlbt
lik«.
§ 490. The CmnAformntiun, however, of louciu into urea nuses a new jn'ml
of view. Lcnrin. us ne know, it aniiilo-caproic acid; and. with our pfwrni
obeinieal knowledj^, we can cjjueeive of no other way iu which Uucin no
he converted inlu urea than by the comjilel^ n.-duction of tbe former la lb
ammitnin condition ('[be ciiprolc acid rivldue being either olaburat4d inttt
fat or oxidixfd into cnrbi)ntc ncid) and by a re eonn I ruction of the laltei noi
of til u aniRKiiiiu »ii f'lrmiHl. We have n ><>nii! what jMmllel com iu gljua,
which is amiiiii-mielic add; here. tin*, a revon* I ruction of urea out i>f «
amnionin phnxe must lake place. Moreover, when aromoniuni chloriik ii
^ivcn to II dog n vary liirite (xirtion mi|>pciirs lu ur«a, i. r., there is an
increase in tbe urtaof ih« urine corrntponding to n largo porlioa of lbs
nitrogen conlsiincd in tin- iimni'iniiim chloride, .\ud in ihe case of oilier
nnimals al»i>, inih-cd iif niiin hiiiiM'lf, there is evidence that soinoirbere in the
body ammonia mny br cmverii'il into urea. Henco in all thoM casee where
ammoain or ammonia {-uttiponmU niv chnnf.'cil into urea, tbe last st«n at all
events is one of synthcsi* : and thi* suggt^t* the jioHibility that in Ute ordi-
iiarr proleid niciubiilisin nlsn, the itownwiird katabolic series of clianM
may linisb off with a Miilliclic t-D<irt. the last ttaj:^ of the foruer being Ibe
appearance of iiu iiminonia ciitnpinind which is eubaequenlly reconalnJCIed
into urea.
'lliis synthesis, like the trans form ution of leucin and other bMliaa, proba-
bly tjikce place in the liver: and in support of this view we hare a certain
amount of experimental evidence. Birds may be kept alive aAer total exlir-
|Mliou of tbe liver for a longer time than can mainniiilB; and wlivn in geeae
tbe liver is removed the uric acid (rrprv^enting in thew animaU tlie urea of
the luauininl) is lar;;ely deoreosed. while the ammonia of the urine ia lar^lv
iucraaseit. Aller the removal of the liver also, leucin, glycin, and otber
amide* or amido-aeids administerod t»y ihc alimentary can>l no longer
tncreaie the uric acid of the urine, as they do in the intact animal. In tbeae
■nimak, tbe synthesis of ammonia eonip»un<I* inin nric acid, nbich la parallel
IMP ON NITBOOENOUS HETABObl
»
\
I
to th* ayntbcna into urea occurritiK in the mammaT, ««etii« to take place in
tW liret, and we may inter is to »>ini« way or other elTeoted by the liL')nitic'
oelk.
As to tke exact way in vrhich ammouia, either as such or in form of aD
taode or aiiitdo-aci<) chaiiKra into urea, we have no certain knowledge.
AnnoDiuni cartiuuate. we kuow. is readily lomied out of urea bv ulmiilc
hTdratioo, and we may iumciiir that ilitr living- organiain ghd carry uiit the
merae ptttcvm mad dehvitrati^ amiiiuniiim cnrlKMiute into urea. There in,
iMWerer. a ceriaiu aniuuni nf i!vi<li!nce that o<>t aniinr>nium ojirtHmute but
Hinoniuiu carbaxuale iit the iuin]i.'<)iale aiileci.ilvnt (il'urui; lun), indeed, nut
oTthe body, by elect rolr tint: a Nolution of nnimnniiim carbaninli- with alt«T-
Uliug current*, a certnni aoioiint of urea may W arliGcially protluocd. But
lliu ia a maitrr tiui iib«<^ure to Ixi diiM'u»<«l Uvtv.
$ 481. {'n> aeii!. Tlii*. like urea, is a uurmnl conRtitncnl of human nrinc,
•ad, like art*, ha* Iwen f'xind in the bliHxl, in the liver, and ia thv vplecn ;
it b a ooiupicuouK voui^litiicnt of an exinict of l4ie latter nrgnii. In tomm
anuDala, hucK as Itird* and inn*! rvpliles, it take* (he place of urea. Tn vari>
ous diaaaaw the uuanlity in the urine is incrtiM.il ; nod nl tiineit, u in gout,
nrie acid ■ccumulatc* in the blood, and n deposit of unitos lakc^ place in the
tiMUM. Sine* by oxidation a niolocule of uric acid can be iplit up into two
moleculntof urva. luid a moleculflof M>tn« carbon acid, uric acid is commonly
■ookra (if ai a liiw oxidiHnl product of proteid nictalxilisin than urea. But
UMir ia no eridenoc whatever to «how thai the former ii a necesmry ani«-
«idMl oT Um) latter : ou the contrary, all th« &ots known go to show tliat
•^ Bpfwarance of uric ncid is the result of a m«tabo!ism slightly diver^ng
from tJint leading to urea ; indeed, it ia |>robnb]o that the divergence occura
toward the end of the scries n{ changes, tor urea given by the mouth to birds
mppean in the urine as uric aeid, and. conversely, uric add given to mam-
■Bwa appears in the urine as urea. W'c have no evidence to prove that the
«suae of the diver^ooe lice in an insutticienl supply of oxygen to the organ-
iun at large ; on the contrary, uric accid oceura in tlie rapidly breathing
Urd« a* well a^ in the more torpid rcplllcd. Nor can the fact that in th«
fnc, again, urea replaces uric aci<l be explained by reference to that animal
bsTing M large a cutaneous in addition to its pulmonary rtapiration. The
fina] cauMS of the divergence are to bi* wnight nither in the fact that una
h the form a'lapird to a Hind, and uric ncid to a more mlid cxcrenK^ut. Nor
■I ifatn in man or ihv mammal anv aaiiHfni-tory [>hy»iologtc«l or clinical cvi-
dnet thai en incrcoiH' of uric ncid i.-< ihii romlt of deficient oxidation. The
abaolulc amount of uric acid diiw^barged by man and ila pri>|>ortion to the
uiM [laswd at the Mine time varivs a gootl ileal. There is no pomtive cvi-
dnice that the (jtianliiy cxcreled is neoesiaril^ iucrcwHHl by nitrogvnoas
diet. ualeM some dixirder supeivracs; indeed, it in nsecried that botn abw^
htaly and relatively to the urea the (quantity excreted is greater upon a
Bitzed diet than upi*n a heighly proteid one. Alknlica in the food sn'ni un-
doubtedly to diminish it. ami alcohol, at leant in excen, to incranse it.
Su far from eonsiileriog uric acid «a a lem oxidized nntocetlenl of urea we
ought |ierha[ia rather to regard its appearance as a result of a synthesis in
%hich urea or some allied body takes part. As we have nid, uric acid may
b« fiimitd BTtlthctically by heating together urea and glycui ; and it has
tborv reveutJy been similarly prepared from variuus allied bodies. As to
where or bow «ich a synthesis is eifected in the living body, we know littl«
oc mrthlng for certain, and ean only make conjectures. The constant pres-
•«« of uric ni-id in ihe spleen, however, and the frequently noted connection
btlwern a rite and fall of uric add in the urine and Tariatioue in the volume
and therefore preaunuibly in the activity of the spleen, euj^geet that the
WW
Tin
*AnoLIC I'ROCRaSKS
chttDjn niiiT 1m) t>nMif!;lit iihnut in thiit »Tgmii ; tmt it intut ht nmttiubuti
llillt in birds and r«titilc« the rormmioii nf uric lU'ti) mcrir Io Iw BtKeCnl In
tlie BameorniwMlnat of urpii kikI in nil iiiin]i>g;i>ii* nianiiftr; mail lUu nrgv-
aveata which «"« h»ve u«ei) concrniing the fnrnmtitin of iin>n in the liver ot
roammalB, niar Iw njtpliod U> Ihr furRintioti i>f uric xoid in Ihe liv«-rs of liinb
and re]>tilra. It a more |initMbk-, therelurv, thtit in thu toninnml ijie turn
to uric acid mther thitn um is givm in tli« liver, liM apken, however, pa*-
aihiy playing il« part nlen In the mnltor.
$'49S. Of the meaning of th« nppcarnDoe in llw liseuee of siicli iKHlka w
xnuUiin, hyjKixiimhin. gtianin, nnd the like, and of tho esact nature uf tht
laelabolism which gives rise to them or whieh tlwy thenmlvee uD<)er)io, we
know little or nothing. The preNOce of ihc^e several bodim inny 1m taken
a* illuNlraiiDjr the eomplex ana varied nature of proteid nie4nl>oliaiii to wliich
we n-ferred aoove- Urea is the chief eDd-priKluct of proleid nKtaholinin. hul
thitt 4-nd in probably reached in aeve**] ways; ao thai probably a very Jtir^
tiiiiul>r-r iif nitroK^noia chemical substances make a monieninrv np{K>aruee
in tl»<- IxhIv. Some of theee fail to become urea, and either wIiVhii nr aflar
fiirdier ehange make their appearance in the urine. But we do not )(ai>w
nhelher tlivir appearance » aecidentat, Ihe nwult of iraperfbct cbcBUcal
mnehinery; or whether they, llwugh HrnaH in laianltty, tterve miuo apodal
ends in t^c economy. I'eriiapa sometlmei^ or with mhuh at tJieui it \» tltb out
eaae, at oih^^r dmea or with othcn il i.'t the other caae.
When proteid material undergoes outside the iMxty, either by tlto actio
of trypsin or a» the iv»ult of dAcomponition or under tlti! inflHenoeof ehcuictl
ai^entB. that chan^ bv which it i* converted into leuciii, the leiioin which
appears in s<ime coniiidenible (juantilies, it nccoiDiMtuied by tynMn, whitk
appenra in (mailer (piBnlilii,« a* well lu* by other iKxIin. TIm! aloiiMt coDstaal
appenranoe of tyrosin ax ii rceuU uf the decompocition of protmd material
leaids one, as we have previously siud, to the conception that >orm rvpnaeula-
live of the aromatic scries enters into the constitution of piutaid subMaiiM;
aud it is pomible that the hippuric acid of flesh-eating animals derives ib
beniioio ncid constituent from thisaromaiie radicle of prui«id matter, TymMD
itwlf does not appear in the body as a normal product of protcid nielaboliscn,
and we are therefore led to inJer that in proieid inetiiboltsm the aruroslic
riuticle takee on some other form. WhotJier, as in lymnn, the amiaatie
(phenyl) nucleus is associated with an ammonia repreeeolative or no, we do
not know. Bui if it is then, since neither lyrosin nor any similar Imdy iti
mnftituent of normal unite, the ammonia constituent is some when; iIIhmo-
ointed fnmi the phenvl one; aud while the former t-onlributes to the Jt-wk
of urwi, the latter is either disctmrtieil by the urine n» hippuric acid, having
iiK we have neen eHeoied in the kidiiev n new a«>ociation with Ibe nmmoiiis
n-prtsenintive glycio. or leaves the Wly as one or other of the uriuary
phenyl mmpnunds, or |>oMihly rihv txf u>:idix«d ountewhere into cnrbonW
acid iind water. Our knowii^jj^'e on thin fH>int b limited, hut ire have ven-
tur^il to n-fer to llut point *iuce it further illustratM the complexity of pro-
teiit niHabolinm.
§ 493. In Kpcakiiin; uf urea {i 402) we allnded lo ila retaiiuni to iW
oysnogen con)|Miuii(tH. Bmrln^ in mim) the fieouliarly large nm<'i.
energy net free iii> lieni during the isomeric trnn<'formaiion of many r> :■
compounds, as well iwlhr liirge store 'if [Kilmtiiil <-iK'i¥y existing in iv«iAi^>ai
itself. i!ie heat «!' comhiisli'in uf which is Vfry laiye. and cnlnuitiiig ihiM
properties with th'i^ "I* uniiiiuniu and tltr iininionia eonipounds. we cani»al
nelp Wing tcmptod toward Ihe view (hut in the nctnni living •tructure the
nitrogen exii-ts in the form of cyanogen compounds, nod lltitt in the {Kfeage
to dead nilrogenuus waste, during which energy ia Mt free, Ihe cy
STHUCTCKKS ASD rROOBSSKB OT OnSOlTBB KATl*BB.
I
>
ooiupotiitil cbangn to tlie amide or other nmnuMiui T«prtM«Utive. And
thoiv iirv UTcml I'aviH whkh Ini'! Bupporc lo such a vi«w, auch its llie prw-
«noe of »iilp1ii>cy«»ira iu naiivii and urine, wbicli we niiiy look upon lu n
wort <if lenkngc ofarunogm I'lU-ton, lh« artifioinl proHiiciioii of Icreatinin out
of pynniulc iitiii Miri-ixtin, mnd oth«r facts. But (lie iiiattor, ihuii|;b h deMri-eg
lo hi- l>»riM> i» ntiDd, iit um obacur« lo be dweJt on liere.
§ 494. \Vc may unv 1iri(-lly »uni up (be varit^ <lieciiMi<>iia whivl) have
occupied us iu tln' iimiciit tMvtion.
Urea i* tbc nmiii wid-prmluci of pniWtid iDMaboliitni. I'liliico hippuric
arid aud soiDe otlier conelitiiitnlBi iif urin«, iirva in niuply i-xcrvldd bv the
kidneys, bang brought to tbrin in llir blo(id, thi^y apfxiri'iitly. beyond the
tiinple ai-l ol excretion, doing no more thnu nicrfly cnntHbucJD^ [<> the
stock of uren iu bo far as ihcy iiru ruimm of ])ri>li.-id material uiidergotiij;
proteid metabolisni as part of ilieir svnirral life. What are tbe in)m^iat«
iinteoedenU of una we do not dearly know; t»it it is probable that they
are not one but several and indeed p^raibly ninny. We have reason to
lliink that urea may be furmc<l out of amid(« or amidii-acidt>, or out of
ammonia itself by a synthetic nrooc^s; and we liave indieatiuoa that this
synlbe&is iieflected in tbe liver oy ibti agency of the hepatic celU. But we
do not know whether this avuthesia bear« only on particular r it ro^n- holding
Mibnanoa of food or of tW body, or whether it comee into play in the
uomial inetabultsm of proteid material. If ibe krealin whirb in to coo-
apicuoufl a riiiuiituent of muacular and uervoua structural ts a Mtjigc iu tbe
air«ct linu to iiri-a, then the synthenia would aflfacl only tli<! wmvHtni which
tJ»e krttttin in becoming urea wets free. But we have sneii tliat it is by no
means clear that krentiu iw M«h a atage.
Tbe evidence as far as it goes tendii to whnw that the inriabi>li8Tn of proteid
is very complex and varied, that a lar^ number of nitrngon. holding auk-
Ktanoi* make a momaitary apiK-aruDcc tn thn IhhIv. taking origin at this or
that step in the do«ii<vanl Hliur> <>f katabolic m<^lid>oliiim and changing into
aonM-thing el»o at the next atcjt, and that lliu prv*«-nci- in various jMirta of tlie
liody and even in the urine, in small 'juaiiliiitv, »t' ki> many varied iiitrogenoaa
eryiitallini- i>uli*taiiGcs, fonuiog a lari.'c part of what arc known as e-itraclivee,
has to do with ihie varied metnbolinm. IVmtlhly the tranatbrniaiions by
which nitro^'en thus panes downward take place to a certain extent in sucn
- organs as the liver and the spioctii, which are remarkably rich in theae ex-
I tractives. But the whole slory uf pnvleid melabolisni eondsls at prevent
^^MBlly of guraaes and of gaps.
I J*
Os SoatE STKt'trriiKBs axo I'kocbssgs ov 0B4ct;H8 NATirRi:.
S 48}. ne Iht/roid b<nly. Certain structures of obwrure nature, but |>rut>-
nbly connected in auiue way or other witli some of ihe metabolic procCMtes
ill tlie IxNly. ana oHen spoken of inider the unduairahle name of "ductlcM
glands." .Sich are the thyroid body or gland, the |>ituitary l>ody, tlie
ibyuiiia, and the xuprarcual caiisulea. These dillcr from each other »> msen-
tially that ibe only plea which can bo urged in favor of oiin«idcrtn^' them
lujrother i.i convciiiitnce and our ignorance of ihf ir nupcrtivc functions.
IT- Xbi< ihvh^d IkuIv is tite one of the group nMwt deserving to )<c ciillct a
ad, siui'v ii, like tltc lung», ariwea asatwodobed diverticulum fn-m the
ventral *urtikcc of the antctior ^Kirt of the alimentary canal, and at Arst,
like the lungs atf ■, huhnv<s as if it were about to iMCximc a double racemose
gland. The tviniM^ciion with the thnnit, however, which should have become
a dud, U mxin olillleralcd, and tlie two lobn, united with cacJi otlwr by au
HDS THK METAHOLtO rSOCESSRS OF THE ttODY.
iNhmii* ncroffi tli« tniclKii. Umv nil tnocs of nny lirnnchini; duets witKin
thtlD. nnii brormio tranKlttrmo) into niBM« of iBol«t«d ituctlew alv«(ili bcui&d
tofMlii't K'itli r»tiiiccti\T tiiHie.
HtiRci-, vrhiii It MMkhii w lnk«n through a hardened and prppareal lol»a(
Ml niliilt thvrtiwl, whikt ta «e«it is n liinitiDg cnpaule of oi)im«ctive tlMur
eendiiig into th« interior numorouB septa, which Hirruund and fte|>nrBt« frnn
euch other round or ovnl sjuces, the sections of the ieolaiwl aIv*Mi|i. [PiC-
170.] ThcM are of variable siie. some being visible to the nakul eye, aiM
each is lined br a sin^'^ layer of low columnar or cubii-al nucleatnl vHk
reating on a btsement niembraiie. leavinj: a lat^ cavity, which lo (Vtih
•pedmena is filled wiUi a glairy fluid. The ix\\» present uo apcciKl diw
ftotem.
ini>.im.
I ' ^I'^.rT,'^^.
BiuTuo or nia TiiTMtb Guini or a Cmut,
Two n>iD|ilMa ndcloi aQi] poillulit al cihcn ■!« rvi-iwcnled. Tlw Tiii<rlll Ma Mlljn(||h •-•UcM.
1>hl(?bBl*narra|ilwih*liitvnlllfNl>|fln», In 1t« uMdM o( am of tlw (pMNRMMSTCSat Wierii
put obll-|Ui>lf , ind ciiM lo II !•• rlatiM tvIL IMse*n (tieiublcal ((itkHlllD C«n«mall»r«>nill>*
lymph ruriiiuclr* nn htrt anil Ibm ■«n.|
The Mpla of eanitcctiv« litaue, fairly rich in elastic «1cnicni«, but reinark-
■blj Irm from Mili|»iMT tiMue. cuiitain numerous bloodvesnU <(criv«d from
the wperior and inferior thyroid arteries, the braiinhn* of which. relatir«ly
lafve and rni-ninilly Hmwidmiwinp. end for the mont part in mpillary Det-
woras round tlio alvculi : from Lhe«e i-apillari4« and lAuwi of tJte eeuM ibr
blood is gnth«rcd into veinn iilifi rrlaiivtily largi', which, forRitDz pleXDaw
on the surface of the oretn, vnd in ihi- itu|><-rii>r middle and in»nor thy-
roid veins. The thyroid tKiiiv \* tbiiv ftirnixhni witli im abundant anpply
of bkod.
Tbe sepia nbo eontain a rer^- larve number of lymphatic vessels, whicli.
botli on the surfacn of (be organ and along the mpia. arc arranged in |des-
nB«e of anaslotnoeing trunko of ii>R»ickTubTe ai». .Small uodulea i>f adeDi^d
tiMae are al<o found iu the sepliL
The nerves of the thyroid body are also abundant. They are, in man,
derived cbieBy from the cvrvical sympathetic nerve, paasin;; olf from tlie
middle and lower cervical ganglia; their exact terminations within tbc xrgan
b not known. Fine GIatnent;i are also said tu be given ofl* to it from the
external branch of (be ntperior laryngeal nerve.
The "acceisAry" thyroid bodies ollen found are of theMiae nature a« tbe
main bo<)y.
Very IVeiiuenlly, to freijuentty in the adult m 1a b« of almoM normal
ooeurreace, the alveoli contain not simple glairy lluld but u more aolid clonr
STBCCTDBK9 AW PB0CK8SXS OF OBSOl'BR NATUBB.
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I
I
malt-riiil. cnllixl "rolloid :" tbiii gviivrnll^ «[>|)(^iin> in tliQcciitrft ofiin nlvvnliis
Mwl laity Hll up the wliolt! Itinuin ; iKTu!nnnlly morr i>r Imw chnugcti e\i\-
(hrliftl itrtlK in»y 111' wM-ii lying bvtvrmrn it nml the lityer ol* ovIIk ratting on
th« tMwmiMil mcmbraDC £xtrxvaMilu>iu of blmid into tbo nlrooli nra nImd
■ not ijni»niin<in.
Tbv tbyriHtl body m ren* opt to bceomfl enlarged, MmnimfK enoTmouMy
>o, and is ihvn *pok«it of m goitre. The mlAreem^nt may be iltip »ini{ily to
nn incKuae in thv number t>t otlicnriH fiiiny oormnl alveoli aikI mpta.
But very often n number of a)v«oli bccomo more or lem conltuent, forming
« cyst: itnil at times the whole inland ap|>eiiT« to be compueed of n aiimber
of cysts of vArying size, frc(|ueally lofidwl witb "onlloid" materinl. There
ifl also n form of goitre id whioh the enlartjeiuent is chiellv or even exclu-
sively due (o an increase in the vascular BUi>]>ly, the bloodvMeels being
abnormally distended : and this apparently may occur without auy structunu
(^hangee in the walh? of the blooaveeaele. Somelimes. however, the arteries
aoder^ Mwuriatnal enlnrgemenla, with ohangce in tbdr comu.
The gluiriDMB of the fluid coDt«nia of the alveoli has genentlly been
attributed to the presence of mucin, and thia body has sAti^ been said to
kas'e b««o found within the lymphatic vessels rtinning in the itepta; but
some obeerven have urgeil that the material in ■(ueMion ia not true mucin,
but a peculiar form (or fonn)>) of [iroteid *ub«itanc«. The " ooUoitl " mat«rial
eo frequently apix«ri»g bus uIho \it«a ruganled as allied tu mticiu, but \i»
exact nature ban not as yet b«cn KutkliKclorily determined. R<Mdai tbcM
•pecial nabMauon the alveoli or cysla uIho contain »enim- album in nud
etobiilin. Tbe "extractives" nf thi; thyroid amiear to omiaiu knMtin or
lireatinin In not inc«i»id<^rnblc i]Uitntilici«, XAiituiu, and lactic (paraloctJc)
icid : gtiauin is snid to Ik abMcnt. In htrgv and old cyM« ebolertotiD is
•omciinM pratent ; and wlwn, us ol\cn ha{i[>nD8. cxtrnTaantioaa of blood into
tlw cn4ii luivi^ litkcti place, hatmoglobiii, or nt n later stage liwmatoidia
(bilirubin) has ti««n fotiRiL
S 486. The larga mippJy of blood to tbo thyroid suggMlii the id«a thiit the
ornii is the leal of sonu of tlic siilwidinry mctjilMlic procema to which we
retcrrvd iu the laat aection, and thin view in supported by the presence of the
cxtruGtives just mentioned; but we hnvc no detailed Icnowledgo of what
actually goes on.
The pnneace of the peculiar mucin-like body in the alveoli, and the teu-
dencr to " colloid formation," further suggest some relation of the organ to
the lormatioti or distribution of mucin ; and this view has derivMl a c«rtua
Hipport from some ezperimenuil results, hut theeo, though numerous, have
proved neither uniform nor accordant. When in c«rtuiii animals (mookeya,
dogs, and other caniivora, and the same baa been observed in man) the glaud
is extir|>ated, even witb the greatest care, the operation is frequently folTowetl
hy the oci-iirrence of pe<-uliar nervous sympton».such oa muaoular twitchinga
nod tremors, siMunia, and even tetanic convulsions (more especially obaervM
in young animaJs), aecompanied or succenled by irregularity or failure of
voluntary movemeiita; mbaequently there may ensue varied symptoms
which niiiy Ik^ dnrrihed under the general term of disordered nulritition,
ending evuntunlly in death. In a certain number of cases, however, in the
above kinds of uniniul, no HcrinUH Hympioms follow, even the total extir|>ati<>n
of tbfl organ iirixliiciog no marked dlect ; and in rah bits and other herb-
ivArous niiimaU removal is raid uttrer to t>e followed by any of the abovo
re«ult«. It has been urgcil ihiit the symptonia when seen are the efibcts not
of I he mere absence of tito organ, but of mischief set up by the ogwrattoii In
ndjoitiine «tructuros n>or« nj>ecially in the laryngeal nerves and vagus
trunks ; uut this do«a not sceni a valid explanation. If, aa suggested abovo,
an
610
TOI
tltuLlC PROCKSSES
BODY.
ogiMJn BWtNlHlio proccwM nro itornuilljr going on w t liv <irgan, ims msy dirlf
■UpptM tlial. is tbe BbMuw I'f the orgnn, th<^ iiik'rTii|>tiou of the DoriMl
■equMtoe i)t cbtmicHl chnncv woniil tlirow ii|m>ii iW cirmltttion wruk
itnuigi) «ub«MfMWBwbirli,KCling lik<- n p<>i»i>ii. might pnHlucr the iiar voua synip-
tome, Ihrow Jnlo <)i»ort]fr thi.^ oiiiriCii<ti of vnt'unit Iimum, u)c) finalk ttriug
alwiii ileiitli. W« mar farther ex|>)uio the cHt« whorv STinploiaa are aheeot
1>y Bu|>po«iQg thill, for Mine rciieoD or other, " thiii^ have lakea a diflbnut
luni," the pnrticular |HjignnuuH sulwlancca have Dot [uad« their apiiearttDit,
but iunocuous ohm havo talcm their place; aad we know hovr dli|-lii *
ehw^ in chemical coinixmtion maf turn a poison into an inert bixlj-- 'Hiti,
of eourse, remains a mere euppooUon until -m can eisie what tho ^xtM
inetBboliv proceascs are. sind Dsme tbe substancee vbich work ihn miM-hi«f;
but il seema more reii»>»able l« accept such a proviaiona) BU|i|iuMitii>0, tliaa
U) conclude tliat the thyi\.iit may be removeil uithiiut )inii]u<-tMg aajr aSott
whatever i>u the urKiuiiHm. An atiimal without ii ih.vr»iil may npfxnr pe^
feotJy «r«II, because the eircunutanoee to which it l:< (tx|iiw<l <}o itot ha|i(Mi
to teat the ini[»erfection from which it ia realty !<ullei*iiitc, ju«t as a roaB'i
inability to itnim may not be apparent until lie hap|K-nii to Ail) into Um
water. ' Tlie MEiimaU which do succumb to the nperatiuD of rem^iral »f tb*
organ nr^, for WMne reason or other, put to thi! tiH, anil ar« found wanting.
The very dlflcordaiiGfi of the experimental niiult* )i»intH tl>o physiological
mural toal the pheuomcnii iiiiich wt- uiv an yt able tooliwrve flirot, Mit
vcrc, a mere :<iirf»ce c<ivcriug iiitricute j>rucc«»cs at prevent wbolly, or Dtail;
wholly, hitUli-n from un.
The above vxpvrinK-ninl rcHulu receive additi'iml interral and at the eatoa
time support from clinical experience Tbe co»n<.*clittn between satire aad
cretinism — ibe latter clincaw! Iieinfi:, broadly spetikiug, a rvsiilt of iiaorderMi
nutrition (clling iarKrlr on ihu iicrvitu* nystcm— line long l>eeu rec>i)fTii]U)<l ;
and attention has auo DC«n called to eocav tic between dlwatse of the tliynid
ami a morbid condition, known as myxtedcmn, in n certain number uf caaa
of whicb mucin or a mucin-like bodr boa been luund in Kreat exc«« in tbe
akin and in other tissues. In manKey* the removal of the thyroid low, ia
•ome cases. l>eea followed, besidee the symptoms mentioned above, sumr uf
whidi rtttemble those of rayxoedema, by an accumulation of mucin ura
nmcinlike bmiy in the skin and various tissues. Il is very dilHcult not to
connect thi^ with tbe fi^rmnlion in the thyroid id* colloid nvatiTinl in the mn-
Wntf of Ihe iilviWi. Hut we know so little a&oul tlie nature of mucin awl
its altiiii, abi)ui Ibeir real relations to more ordinary prateid nulMtanoea, nnil
obout the giart tihich they play in pbysio logical ]>roceMea. that any views «i
to tho exact connection belu een the nreeenoe of mucin in the tt«»iie^ at laq^
and cbai^ca taking place in tbe tnyraid muH( be at present to a lur^
ext«nl iq9eculnlion.
Thv large vascular supply of tlic lhvroi<l, and tliv |>lM-nomvna of a dbeaae
known IW eiophtlialmiv goitrv, in whie^i vascular culargumMit of the thyruid
is ansociatol with cardiac syraplom« and other vascular disturbanoea, eaped-
lilly of the bead, have oiigKefled that, njiarl from mutnbolic pninmiii. tbe
circulaliou in the thyroid nmy, nerhajn in a more or Um mechanical way, bt
c(>nnF<-ied with niid inl]uen<« the circultttioD in the brain. But tbe exMi
nature of thix inllui-nce bus not bcc-ii made clear.
S 497. Tftr jiituilani lioJy. Tlie lower. |KMt«rior, lobe of this orj:an r«M<ia-
biw the tliyroid bixly < ibo upper, anterior, lobe ts of qnile distinct UMUn,
beinK really a part oi' tlw rciilml nervous system), in as much am it 1* n
diveiit<-ulum of the nlimenlary eiiiial (nnnieiy, of the mouth), which, instead
uf lM!cumini> n branclicd inland, is converted into a uiaae of round, or oraltCr
cylindrical alveoli, Htpaiated by septa of vascular oonneclive liaaua, Tliougli
STnUCTt;RES A*rD PR0CER8EH OF OVSCPBE KATCRI!. $11
In wme iiirtBiicw tin- ulvwii ol' the pituitary body, like lliose of the thyroid,
pOHMM H liinif'n. which, n>areov«r, mnv hnid more or km " collnid " cont«aU,
the ninjnrity nrc solid iiiMeeR uf epitlieliiil celU. The cells, which are
cnluiniior or polylM^ral, pr»ent no epecial characlers, expepi. perha|>«, ilml
bvtv((«n tlwt iiHinl cpithi^lisl cells are occaBiunally Touud spindle !<haped celb,
itppurcntly of nicsohliuik origin.
C»nccrniii)>; (be jirooeases which uke i>liu-e in thwe alveoli, aud tlt« pur-
poses of ihe organ at a trhol«, we know abwiulely nothing.
S 498. The ruiimrrual boditt. A (»mmiii«lian) Niiprvnfflnl body, wheu
ctit iKn»8, is seen to c»n»iiil of Ino dutinct partK, an otitcr thicker mrft'oi/
part, of yellowish (v>l»r. Htriated radially,
and an iuuer ihiniH-r mrtluUarg {tart of t ^
darker color. [V'tji 171.] Attlie<kprM-
Mon ou tlie anterior .turfacc <-«ll«(l the hilun,
whence isMie- th<! coinjiaratively large
Fiipra-re«fll vein, lh« curtvx thin* away »i
thai the ntcdiilla mmm li> tho (iirlunr.
ThcM iKo partH, cortex and ini.'<lullu. arc
not, like tlie cortex an<l itiixlulla of a
Ivnipbatic gland, dttrrrenl arraniii-mf-nlB of
the fame inatrnal, hut arc of <v>M-nlially
diflcn-tit iiuturc, and, ind*'»il. areofiliHi;rftHt
origin. Tlic medulln if derivvxi fmni, in %
mod ill cat ion of. «ynipatli<^iip g^iigliii. while
ihc orlex is de.rivr<I from niamw of mew-
lilamic (tIIn iturn>»niliit)i: the Krnit blowl-
vOtteb; and in ifunii: animals the Ino form
^ wholly ec|iunit>- bodice, The si>-i;nMed
■■jaeiMory supnt-reiink am ronipoMMl of
^^^rtcx aloiin.
Thfl wholt W]gan it surrounded by a
ai|Mdo of connective tissue, free from
muscular libres, and not very rich in
daMte elements. From the capsule eeptn
pass inward and ftrm n frameworb, the
caviliee of which are filled by cells or
(•roups of cells dilterinc in nature and dif-
ferently arrangni in the cortex and in the
inedulU. The middle larger juirt of the
cortex is composed of eomewhiil long solid
oolunuiB of polyhedral cells, lodged iu cor-
retpondinjK iDesho of the framework. The
columns, whieb arc three or four cells
thick and several im^Us in length, though
•utDewbiit irregular and varying in sixe, do
not anastomose. Wiuj; irhitUv m-piirated
frotD each other by the Wn uf i-oiitio-tive
Umne, and jhimcsh no central cavity or
laiiMn. Tho liloodvoHeU, which an.- ubundant in tlieie bars of connective
tunic, do not [lenetratc tlir riitumii^. The i:ell !<!ilK>iance of the cells is of a
yellowish color, ol\cn nmiaining vrllowixh oil gloliulet, and podseKU a clear
round nucMio.
In the oater jiarl of the cortex inimc«lialGly underneath the capsule il a
thin xone in wl)ii:h tlie gron^w of cvil« are not columnar, but niuudcil and
irrvgular : and ngiiiu in the inner part uf the cortex abutting on the modolla
I
VEsnc-ALtliicriaMMSi'i-r.i'iKoii. Bonv.
(XU)inincil I
I. uvtimi tilManc* : !. miNlulliur i**^
•ud»: a. ttiBtnit : i. tail* ilunuiraliMi :
r, kin* CuiitcuUU: <l. »>na rotlculiirUl
I. ■raaiaof innlullvr orlli i /. wMioii «(
« latp tdu).
612
THE MUTAIIOLIC ftlOCESSKd or Tlti: hout.
fs Hiiolhcr thiit uinr, in whicb (bi? columnar arranj^iuenc is ]>. I!«
being hvn ttitpoeeil in a necwork of thio cor<l», and tbe it><ltviilii.<
Inrgv «xteiit Mpttrated tnm eadt utiier by delicMt« citniiuuutiuiu ut ih<-
Goaner connecUre'ttmte MpU. ilvm^. the ntaiu mediau [mri ut' tli« iom. \
which frutu tbe ]>roiuineiit culuiiniar arraofcenient kppears atriuttsl m <
is oflcn caIImI tbe »>ua fascioulala, the thio outer part the xona {{luioviui -
uiiil tbe thin inner p«irt the xuoa reticularis: hut a» fkr «• Um vRwniial
diaractera of the cells are ooncem«<i, all tbe iliree touea are alike.
Tbe medulla also oonaiUs of eells or fjrouus of oelU lying in ihv innihni uf
a connective-ltwue nvmework, but the c-ens are of a diflereiit iinLura fnin
thoee uf the cortex. Tbcy are irr«frular and often bnuit:h«d, and their ivll
aubalauce. Ihousb it »i>Ri«tinie« <<un(aii>a )»|froetil, t* fctinemlly rtcnr awl
tnnaiMireinl. Tbe iDedulla. ninreover. ix furtbi*r diMtinguixbticl fmni the
Ctiriex bv lh« ubuwinnt Mi)i|ily »f hl<iodv««wla and of tw:rvM.
The cgIU uf tbe medulla and of tbe iuuer wn« (Una rrtkularis) of Ike
cortex lire very aj'l Ut untleiYu cbanue after ilcatli. anti to boocMB* diffiuMtt.
TIm- artrrini winch eoni^ mmi Oii^ mirta and friiin tbe nati and phnaie
artertisi jiuni inti> ihr organ on tb<^ Mirfacr, «n<l travcrfinp ihit ourtox, stipgrfr-
Ing an llu-y go both (!H|niilt' ami mrt^x with a niudi'nile iiiiiulfer of vematt
end in the imxliilla, ibc (1)nn^(rIiv<^li.■«tIe luire of which Ixuir uuniorous Uup
Y4DOI1K MniiK-i>. into which (be cn[>Illariu' jxiur their hliMxl, and fra«i wMek
tbo bUxx) is gnihcrtMl up iiiiu Uie »uprn-T«wl vein.
A targe nuiiilHT of ni-rvut, M>nsi»ti»); chicAy of tnediillated libtva fnnu the
Bolar )>lvxuH, thi? renal i>lexii». llie j^renic i)urv«, and the Taf{ii», iwmi iiiio
the ■u{>m-r?iuil body at the liilno tutd od the under surface, a»d foruiins
iiunicTouv i>trxu«(«. coarae aw) (inc. some carrying tunall groups of nerve
cells, end chiefly io the in«lult.i, though eome pnaa on to tM ooitex. Tbe
ullimatc endi»g« are not yet knowo.
The lympliatimare fairly numerous, and form plexiiece in l]w capsule and
in the connective timue of the framenvrk ; it is staled that the lymphatic
veeeeU surrounding the groupsof oella in the cortex commuoinile wrilh spaoca
between tbe celts.
^490. Ueaidce the onliiiary proleid and other chemical ooostitueiils. the
snnra-reaal body cuulaina sume subsianoe or eubstaDcea posseasin); atrikiag
color reaoUous, ;;ivinf; a dark-blue or darkgreeo color with ferric olilori(l<^
and a cannine-reil tint wi[h variiHia oxtdiunK atrenta. Tliis Bubstanco f whoa*
nature is not exactly kuouu, and which is confined to or most nhundani in
the tuedulta) is not aoluble in the ordinary aolveuis of pi)ctiM-iii», MUch ai
alcohol, etber, ehlorofonn, etc., hut is readily soluble in dilute acacb.
Anionx the extractives, hippuric aw) beiiioio acid, and laurodinlJc
or taurin have been found, out it ia not certaiu lluit tbaie are ooroutl
stiluents.
i 600. Hotiie of the bitlolojctcul featumof the Hupra-renal bodii.>«. nanal^,
the rniujiv of <fll!i aiid their abuudani bhHKl-:>u]>ply, NUggeM, on tl» u
hand, that imimrliuit metabolic prooeNteit taki- place in them, »ouo of wbl
are prulxahly connected with the history ot^ tbe pipmentv of tlio body at lat].
On ibe other band, the unusually larev norvv-«upply, and tlio durivalioti i
part of the bixly from tbe HyiiijiathKic ganglia, iiuggmtit pciiiliur nervo<
cuiiiiectioui>. And tlte organ has often acrvcd iii> a i>(iirtiug-|K>int for specu-
IntiiiiiN in ilii»e tw<i direction* ; but our exact knowliilgi.' i>intx-riiing tlieni is
very liniiu^j. The nwulli> of cxiierimeni have taught ut lilllr; cxtirjNUiou.
for cxBiuple, has been otleii followed by the doatli of the animal operated
ufHiu, but the cause of tbe dvvth ia such c«Ma b br no mtwuf ciMtr.
Oiw fact, glinftd by clinical cxperieuc«, n tlie only real ilvin of knoMledgu
which we poneat. I^iseate <tf the supra-reual hwlio*, H|>|)«reritly lulicreular
CESSES OF OBSCl'RE NA
:iti^ W)cii)niiit! in l)ic in«liinn. is »n oflrn Rsmctiilccl with n o)ianK«
: III' nil)) iin incrvDiv "f ihv iiii;iw(il nf tlic »\dn, " lin>nxv<l vltin,"
II ■■.!._. -_ J .:. I... _ .1... ._...
Id I
in
** -\ 'Iwi'iLM-," ilvnl ■Atni^ cotiiui'tKiii Ix-twif'n tin* iwo iiiiikI oxwt : bni
tli< . ^itiku of ihr cliain nrc ■>> yet iinkti'iun. Il t* trfiiitfin;; to iittm-
ci»tt> Uiu incn-AK of pi^'tn^Dt in Ihr l>r»«z<xl wkin with the i-ltruniof^ii or
cotor-jHtfliliDi; wiMatitv xpokra of ahov* ; l>ut ve linvc no wnrmnl lor 'iimig
•O, Hii-li for inMiiniv iif miT indiwtioij iif lira boiwi>oii tlm >ii|)r»-rt'iiti] Imliea
mM*l <-h«>iig(« 'iihnr in hvmoglobin il^df or In bilirubin, which tiro bmllM
^•w l\»vt niuon to rt^n) more pnrlicitUrly ae molWw of pi;,'m<.'m. More-
'OVcr thv brt>Dic<l #kin is oiilr oti« of ihr sTi»i>lomB of AitiliBoii's ilUeuc,
fiulitrv of tiiitrili'Ki and Derroiis •^'mptoiiii' being dso present.
§ fiOl T%e Ififfmut. Tliis, thniigli il ai-iM'# in ihc embryo as a paired cnt-
f^iwth (Vom tite epithelial walls of a pair of viseeiml clefb*. and thiiH begiits
a«0 Dii e|>ttbetial structure into irhioU meeoblastic elcmeiif» subseiiiieQtly in-
Cruile, soon )Mit« on such cbaracters aa to appear eswulially a Ivinphatic
Mtrueture, am), in(l(«d. mi^ht be regarded as & ])art of the lyiiiphatic syTUni.
It DDDsiMs of a capMile of connective tissue, plain muscular tibns being
■tlnrai. u)d aepta or tni>)eculie of the rame nature which divide the orpin
Ibio ■ number of irreunlur inure or leea orliDdrical aiiastoiuosiuK follicles or
lobulw, and H'lid finer nidialiii^ septa inui the interior of each lobule.
TImm li>buliv preiL-iti iIk- Niiut.- characters ihrouxhout the whole lusai of the
«TrgMD, there noi U-iug, an in a lymphatic j^anil, any dixiinciion tietween a
cones ami a miilulla of tbe wliolc- body. The words are, however, applied to
«Mb lobule, to i]i;>liii;:iii»ti Uie central from the )ieri|>lK'ral {larl of ih« lobule
Itaalf. Both the ii>-ntrat ineilulUi aiid tbe pcrijihcnd cortex of tiach lobule
Oootitt of a frnmcw'irk of ri'licular conneclive liwuu, which in the cortex l«
idoiticfil with or closely nllicl lo adeuuid tlvue, but in tho medulla in
eoaiMr and nior« open aiid to a Inr^er extent conipn>ed of branched aniu-
lOBoaine epithelioiti nils. The mvslint of the cortex arc cruwdci) with leu-
cocrlea, out these arc much lust nbumlant iu uxd mora etu ily fall out of tbo
nmulta, so thai in sections the medulla appears inor« tisJDS{)arenC tlian iIm
wrtex. It will be observed that this arrangement » almoet the reverse of
IW obtaiainK in the atveolos of a lymphalie gland, in which the liner giant)
•ubitknce wiin its adenoid tissue crowded with leucocytes in plaoe<l iu llie
centre, aurroundeil by the more open network of the lymph sinus.
The bloDiIvfnels of tbe thymus miininK alunt* the nciiia form capillary
ottworkt which. thou)[h closer and more abundant in the curticul than in
til* neduUary [lorltoiis of tbe lobulea, have no such »|)ecial arrenj^ement lis
obtnttis in lymphatic glands.
l.ynipbaiit? v«»e)s, abundant in Ihe capsule and septa, are uuduiiblodly
(a oMOOedton with tl>e nuluiancv of the lobules.
Tbt nmlullary Milounce freifuently contains brnJieA, known iu"vuiigc»*
trie cwiBuW," nctuof oonvL-nlrically iIiajhiims! nucleaied flattened epithelial
*~'r (-tiilhi'ltoid oellt. Thev ap|>ear lo nriM- I'rum a pmliferalion of the ttpi>
% lielioid cellii lining riiial) bliKnlvciBL-lH, and have beein suj>p'i»e<l lo 1>« e<in-
X&«(.tcd with the dvgeuvralire cliango by nhich, with obliteration uf the
~W iiiwb. the whole orvan ilwindlrs away wion afl^-r Iiirth.
IS 0OS. Prom ll»c ih^vniiin ihcre may lie cxtractetl by mc«ns of saline mIu-
%Joa a Ibrm of globuli'n or n prolci'l allied to t:tobulin which, like Ihe eornh
^ipnnding' lH»)i«n from lymphtilic glands or from leuoocyles, seems to htn
Wonie ■pi-cial relations to the foniiuliou of tihrin. Thus, as has alreatlr bca
Vni<) (it '£"2), a s<ilutiofi of this globulin like bridy from the ihymiis, injected
%nt» the veini' will give rise to extensive intra- vascular cloltiiig.
Tbe tliymus. like the other )KM)ies on which we are now dwelling, is alao
I
I
I
eu
THE MKTAROLtC PROrBSSEB OF TltK 11«t»Y.
rivli ill cxiriK-livra. TlitrH xnnrhiii, hyjwxnntliin, luciii, tnotie, suvciitEr
iAh«t iid<li' liiivi- liecn founil in ii.
Bui of wh«l renllv taVrc jiImcc in the IhhIv we hnre no exact knowlalu
Btncf lb<> thymiiii is Wt dcvclnpetl bcfitrv birtii, (tisap|)earin)^ after l)irtlHL_
» nUi whicti viirics much in diflercnt iudividuab «tu Mill morv in ditrcrenC'
kiiidi "f iii)imiil0. nod being eventually i^plaoed by Tat atitl contiectivn thmie,
it u obvious (hnl its chief functinna are in aunie vny aesndaied ««illi evi>»u
taking |>biCL' befare birlh or in early life.
Tub HbrroRT op Fat. AmpaeE Ti»cb.
( 003. Gl»tiulc» (if fat nf rariotu titn make tHuir aripeaninrv in (hi
alenwniji of mivit nf tin- liN^iH^, in iniiM-tilnr ftbrra. in vfiitbfliiit ix\h.
nerve c«IIh, iu li-tirncyti-^, ami w> on ; and tliv medulla of incvtullai(?d a».
tonim* largely of a ]>fculinr fatly innirria). Boiidct ihifl, ivrlaiu cella
oonnrctiv Itrmv at various liniiti, and in varioiu placw. b«ci)mi- to loati
with fat tliiii groiif* of the evils l>croii>e ]>nic(i('att^' mwwn of laL Uoam
live tiMui' tbki« l<>iute<t with fat is called adipose titHue; lUid inamM ot m
|Mie« iMue uf all innnm-r of sixes aad of ehnpiv ailapted to th« Mvenl situa-
lions lire found in various pafls of the bndv. Xaoy of th« inUrnnl orpuia,
n»tm' <v]>ecinlly the kidneys, arc wrapped in adipose timne; but th« larywi
depmit is one lyin^' in the subcutaneous tissue, § m, tometimcfi called tf
" pannii-alus adipoeus " ; and a " fal " body is distingutsbiyl from a " kaai
body chiefly, l1i<mgh by do means exclueively, by the amount of subcuiaDeoin
■dinnee tiuue.
Of at] the Itwues of the body Hdi|M«e tissue is ihe most flurtuntiu);
bulk : within a very abort s|Kice nf lime a lar^'e antouot of ndiptiM- titui
may dtsapjiear, nrnt nilhin an ahuoet e<}tia1ly short time the i]uantily prei^n
in a bodjT mar he wevi-rut times multiplied. When ton much or too lit
food is given it i* thi^ .itilKOiiuMit adijMise tisue which lirst and nioai rapidly
incrcwea ArdecrcftAn in bulk.
iSOi. A mnall |>i<-CH- of adiivHif ttMue. exAniin«i1 under a low [oiwrf,
ap|K-aR> ti> be niRilr ii|i nlimwl entirely of n>undvil muMi'M of highly refrac
five niatcrial, ctoncly jiavkcil (ogeilier. l'b<«e ronTKU'd mnoM-ji, which ntain an
inU-iiK- black with iMiiic acid and givr other rvjuHiom of fal, am arran;
in irrej'ular lohulot; between the lobule*, ami Wincen the iudividui
rouiidod maases, mav bo acaii a small amouut of fibrillaMl ooiinMtive ti:
carrving bloodvoswis.
When the tissue baa been hartlmcd and stained, and the fill haa been
moved by Biitveota, what wna provioimly only viiibk' w a rounded man of
fat i« nuH seen, under higher {Hiwere, to l)0 a cell, but a dcU noarlr tl»e wbule
of the cell Kuhstaiice of which has become (ranslbmicd into a Ringlo laifie
vacuole. Over the greater part of the cireiiniference of the cell the cell
nihrtnnce is reduce<i lo a mere thin slietl or envelop, or cell membrane, but
at one part a thicker disc-like renmnni b secit, and in this ia placed a
roiindiH) or ovnl, often llatieited nucleus. Bedneen tlieee f'xt-etHs may be
•cm a few huudl«« of connective lisme fonning a eeanly looee network, the
rounded mrahiv of nhich are occupied by llie fat-celts, the matrix uf ihe
bundles amicarinv at pliitH-a continuous witli, or adherent to, the envelii[ii
of the cetU: onliimry omnvctivi^-tisiine corpusciee are als^i here and there
nreeeni. thouEh ran-ly viaibl« between ihe lareer, 50u lo I:I0", fiit-cetl.-<. In
injected sjiecitnens it i.i further Keen that Ihe connect ive-1 issue iiiefthworl
carries smiill blood vcswl.-, which form capillary networks around the groii
» an
tuiH
TITK HISTORT OF FAT.
(lis
(tH-<v\\» ntiil even iiniiiiHl iwlivliltiiil c*ll». After ilcnili, upon iixiling,
th« fitt in lli« fitt-ctOU may tnlulily in irrvBtnl*.
It i> <t1tr)i>ii* thni n /iit-ofU in a etil. I>c)fltigiiig to rooiMicUvo (iMiti*. in tha
c«n «tih«tiini-«nf which fui hn» liocii ciilloctod totuch im cxliiit llint the cell,
which incn>UM« liirgcty iti hulk thiHng th« pmcvM, is «lin<«l nh<illy tnine-
fonitcH into n \iikrgc vtu-uok* tillcil nilh fut. the evil sul>^4iiii:« licinf; reduced
to n ihin envelop of the viu.'U(>le. thickfineil iil one purl where the micleus,
thrust on oik Hide hr (he piiilhenn^ Tnt. 'a pliiced. Adipose ti»ue b a cul-
lecti'>ii Df fucli fiil-cellH held t<i;;ether hy a meagre t|iiiuitiiy of TMcdUt con-
nenive lisf^ue.
Ity sUidyiiiK the devel(>pm«nt of s'lipose tissue in the «mbrro or elaa-
where, we ma<ir trnep i>ut the •Hp\^ <>!' the forniMioii of ihe fat-oel^^. In tha
emiiryo, in n aitiiiilion whi^r^ adipine imtiti is about M Ite formed, Ihe con-
nective tijuue i» wen to onilutn n ntindwr uf ttniiill ntieleated nW^, nxitnled
or Minii'what irregular in fiirni, the irell HuMance of which at lir>l iirt«(!nid
■10 ii]H'ciiil chimicien', imd coii(nin« not iimrc thiin what nmy he called the
onlinary lunotmt of fiit ghihiilco nr Hplivruli*. Very un»t, however, ilieiic
miiiulv drops or 8pe<'k» iticreUH- in nuiiiliiT, the cell oulwtanue at the ttanto
time inrrenMug in bulk nhilc n-niniiiing round or iHt-uminir inon> dixlinctl/
eo, and the eninller dmps nil) to<,'elher into larger oih-s [l^if;. 17'2]. This
nr^^
l>ir««fn<>N or fit m OMtfiK'nvR-intri: Oiua
I olili ■ n-.vr l»liil«d bwlnifilsU In tu firolopliuiu : /. »etll wilh ■ ilnitlcUrt" «ii<liniMnil
■ilnalu ilnic* ./". huiuanr liav Uriii' <1(<>|ii ; w, (maulkr or ptumwecdl. i<i-l )nl vll>tliillu(i«ny Ikt-
■lepoMooi t,l, IUIocan>Hiln-aMiH)*ur|iwrl«i *. r, mtwMk «t n^lulM.]
gctot on : the fat inermning in <)UBniily cunlexom morv and more, and the
cell, u a whole. bpcom<w larfiier and larger, the cell ^uWance at tin-t keep-
ing up in bulk wilh (he ineieasiiiu: Ikt, but suhMnueutly cesHiog to iiicrcaHC,
beiuK ap|>arently uae<l up iu the formaliun of the Alt. Thus the ortjfiual
small " nrotoplaamic" celi in at last (raii^fomted in(o the larg^er fnl-eell, nil
the fal having mn t>r;{ether into S veeiclo ibe envelop of which, ihick^ni-il
on noeeide to carry the nucleus, is tVirnisJied by the reaiDunl of the cell
substanee. In lome ca$«s, the nucleus iostad of heiuK pusheil early on oiio
aide, remaiiH ceuirul th->Uk*h the ciiUei-lion of fat has liecomc coiiHiik-rahlc ;
it in. however, eveniuully disj'lacvd. The whole proccm ap|H-Hn> very similar
tlte d«pn»iti<in of mucin in t)vc cellit of a inucoiM ^land, ii '2^-i : and we
njr h^ ana)o);v iulcr tliat the liit-cell bentmeii a fiil-eell by the cell luaim-
eturiDg fat m aome way or other, and depwitinf; the mt i«> fumied in
iDteratieea uf it» 8u1»tani-e. The mwi Mrikiii); Kupcrficial distinctions
016
TnK METABOLIC PBUCKBSKS OF THE BODY.
seeui In he that in the toacoiie cell tlie cranulea or apbemlcs remain tViM
williiu the cell, lieing sepamted by bitra of cell substance, n hereus iii lim
cell the ((lobulce, as tbey form, run m^^ther uutil at laai tli«y uniie iiit«^
BiD^le inasi; and further thai wliile iu lli« uiucimu c«II, evi^n whiui nk^^
bcavity luftilml, a relalively lai>.'e anKiuQt of active cell-Mhctanoe Mill f^
utiiiiu, ill the Inrcell a mere reiunant U l«ft and tliat chiefly •urrattuitf^
the displaoed nucl«uit.
.Some ob»erven are of uinniou that the oelb bolougiuK to cooiicrtire ti«a«
which thiiM hi-ci>m« fut-vclU of adipoae liisue beloii2 flxclimively to iht ki^
which w(! i<{H)kv nf ii» pliuuna cells. § 105; but tnii i> dMihtful. OUhr ,
Bguin, while mjinitling that the oella which beoodM fat-<.'vll» rMenbltin
apiHAnino.^ ordionry ountiective-tiMue ooqnuclet and luav lik« thi'ii U
branched, hclicvit them iitivorthehm lo coiUtitUtA a >tiecial Iciud at c>ttM»
tire-tifuic corpii»clo. being Ivil to this view by the Coct. that though a^ua
tiasue i« very gt'iKirnlly (l!«trihutc<l throughout the ooonective tiaue oTlU
body, it is apt lo np|ic*r in |iarlic>ilar nitualions, rather than in otb(n,aJ
in some tracts of connective limtic never under normal circumitanon taSm
its Hppcaratice. Others again nininiain that, nnder favorable circuautiacai
anv counectivc conxiscle nmy Ix-come a lat-eell.
The fat in the luterior of bones formiu); the yellow ntarron appMn b
have the same general sinictiire and to be formed in the same way a tk
riM of the adipone tissue.
$ SO0. The fftt ihuB deposited in a fat-ceU aooner or later dbapiwan. it
M nut injected bodily into the aurruunding lympb-epaces of the ci>ii»MiiR
tiMUc, but itaasoa away eillicr into the blood alremm or into tlie lymphaiia
by tODie pracesaea not as yet fully understood. The shell of cell xiUuDa
wnich forms the envelop of the fat-eell is probably of adilTerwntiartil oatan,
and may have properties which assist the orajie nf the fat: but im lUi
point wo have no exact knowledge. The dtnjtpeanuioe of the (at a|fW
to takv place in two diB'erent way*. On the one hand, and thi» i«ru|ia
the mon? onliiuiry mcthiid, the fat gnulufllly diwippcar*. tittle hy littlt,u4
the rounded distended ve>>icle gradually nK<iinif« the chBrvclerB of a c«mk-
Uve-tiBSUC corpimele, even of a liituiched one. On the other hand, atjwcitlf
irben tlie diHijipenmniv in rupid and total, the Kpuee previously flOfiupted wf
At beeoioes fillixl witli n ele-ar Huid nvenihling lymph, the fat vesicle bmf
tranalbmeil into a tytuph vesicle. Thif^ condition, hnirever. is tetii|Kira>7
only, the lymph is EiibseiiucDtly aheorbcl and the vesicle shrinks. At xiam.
the emptying of llie cell, ivhetlier by the one nioth'jd or llie other, ia fiilluW
Ity a rejiivenesceuee of the cell, the nueJeus by division gives rise toatiwJ
nuclt^i. am) the cell <lividc8 intii new cells, each of which may, under tfic^-
prial'* condilions, devdop again into a fat-cell.
i 506. 1'he fiki thus lodged in adiiiose tissue varia aouiewluit in ciirafo*^
tion in various animals, but is chiefly compoeed of oleiu. luklmilin.and ft«M
ill varying pruportiuns, witii small quantities of the glycerin oumiMUiiikif
such ^aiiy nciib us butyric, capronic, ca]>ryiic, etc., together with a ItBtr
lecithin and chokMeriu. 'i1ie " fat " of one animal, that ix, the fat tbut ««■
laini-d ill aili]K«e thme, diflerw from the &t of another animal |>nrtlr br tl*
preeeooe of ntore or Icm of one, or more of ihcvc leas nbondaiit fau, M
ohicdy by the projKirtion in which llie three main fittc, olein. paltuiliD. sai
rin, arc n«i<eetively pruMtnt iu tite mixetl fat. The melting jHHBlt sf
thne thrre fal» Iwing dil1<Tcut, the nK-lting juint of the fiu of the bod* aS
difler aoct>nliug to the relative pnifHirtioiu in which the three are prSMt.
Thu* the subeutaneouc fal uf man mcll« at Imm !•>' to H" or higher, iW
hit r^'uncl tlie kiilney being flrmer and not melting until ib" ; the fat ot |k
\h% inrlta at about 'J2', that of tl>e gooee at about '2^i'. of tlie ox at
THE HlflTOBY OF FAT.
617
I
I
I
I
-to**, And (if (III* flfaeep at oO'. ili« Itte nT(iii>iant tut of I)k- tiini) iintl dog
cvnliiiniiii; rebliv«ly mure oleiii tlian ihui of die ox or ut' the »ti«ft).
S 507. Wlwn w« conie to <»iiEider the <iuesti(>n, Ily whnt i>roccMn does
thv fdt ntttlcc il8 apfieflrnnce iu (he fat-oetl J y^ ar« brought (Ace tn fnof with
oiiicli the same kind of probi«Ri ns that whicli occu|)i«d u» in d<-aliug with
Ifljcoeea. On the one uaiid we may suppose that the fat is bruiight to th*
fat-ceU aa lat and ta iu 8oni« way taken up Ity the cell and deposited in the
c«ll substance with little or no olianiie. On tJie other hand, ive may stippoee
that thi! fat is iflaiiufai-tured by the fat-cell iu e»iuc nuch way as mucin or
fW![Miin is man u fait unvl by a nim-uun or a );B«irtc i-cll. out of and by means
of ila cell subntaiKe, and that the p^><(^t^■« uf lutteiiiiij;, or uf produdog fnt in
fai-VRllu. conHi.itN mMntially in fiiiliii); and wi buildiiigi up the cell >uliiitanoe
nhidi MulM«<iu<!nlly bpcialcM down into fat, and ilm-i not i-cmidst merely in
bringing fat within n»di of the cell. Which of thiwA vinw« iii the true luie,
or bow fur »Tf holh iImm opentionit oirricil 'iti in thi- animal body?
In HupiKirt of thr latter view it may )>o uritrd that, not only ibv more
complex livin<; MilMlnnoo, btit. ii« wv hnv<! nion' than nnre. urgttd, the fimplur
proteid I'lmnUtiM-nl of living mihcliuici- obviously contiiin* what wc may cnll
a fatly nuliclc. «'> that wi' might expect fat to be formed out of it» me1«-
IxJiitm. And as a matter ot Inct not i>n)y in adijHwe tissue, but i» every
part of the hodv, living subitaniT is continuoiisly ^nvini; ri«e to and tempora-
rily dcpi«iiing in itself tome amount of lat. and in irhat is known as tatty
degeueraiion ibere Bcema to be evidence of the formation of fat out of proteitl
material.
On the other band, we have traced the fku taken as food, and found that
tbey pass with com|»nttively little change from the alimentary canal, chietly
throughout the intermediate paseage of the lacieals, into the blood, from
which they rapidly diaappear alter a ueal. We might infer from this that
an exceOB of fat thua enlerine the blood would naturally be diapoaed of by
bedng nmply stored up in the available adi[Krie timue uith»iii aiir fbnber
chaD)i:e; we can ima^ne that the ful, not immi-diatdy wanted hy the
economr, paaea in some way from the blood ti> the ci>ntiective timne (the
while biood-ouqtttaclea nbiiOi a)>j)ear loaded with fat atli'r ii nu^\ pomihly
acting as intern»edianc«). and that the connective tUaui.- oirpum'lvs swaltovr
the fat brouftbt to them uA«r th<' fii-ihion of an onueba, not digextjug it hut
•imply keeptni; it iu More until it niii wanted elwwhore.
What di> expi-rimvuia teach on lhii> miitU'r?'
In the finti place, it i* evidiiit tbnl in uii unimal fattened on ordinary fat-
telling fwHi, only a iinall frui-tion of (he liit st^>Tiil u|> in the body can |watii-
biy ooine direct from tbe fat of tbo food. Iy>ng ago in opposition lo the
tSsws of Dumas and hiM Mihool, who taught that all ci>n<it ruction of organic
material, that all ncttiul manufacture of living suht>(ance or even of it#
organie comtitui-iils, was conlined to vegetables and unknown in animals,
L.iebig shotted that llw butter present in the milk of a cow was muck (greater
tluin iMuld Iw acconuted for by the scanty fat present in the giMS or other
fixlfler «he consumed. He aU-i iirgeii n» an argument in the same direction,
(bat (be WAX produced by hecH. which though having a diiri-reui compfHiiiim
from fat nuy ue used as an aunloi^y. is out of all pro|Hirri<in to the »nx or
allied budiea contained in their food, coniisiing as this duvi chietly >>f .-oi^r.
And it lias since been shown i» many navs that, in fattening,' HiiimaU. the
i'nt accumulated iu the body cannot lie accouuteil for by the fiit nhicli has
twM) taken in the fowl. It lui» \>n:a ]>ro%'ed hy dim't analyxi^. Tliuii of
two young |Hga. aa much alikt^ u^ jKMMble. of tlw utinw lilier, om- niic killed
and analvxed, the anHMinl ol' fat in the body tmng among oihci' thingii deter*
min<-d. The other wan futtenetl for a certain length of lime on fivnl whoM
THE URrAnoLK^ processks or tub bohv.
oonifNwdon woe kuown. and theit killed wkI aiialiried. it irns fotiixl it^^
ftweviTT 100 (>ans of lat in the fooil 472 parts i<f fat were sioreil uii in ».|,
body duriii(* the fiilteDiuc period. It it) i;l««r UuU &l may he fvritiFi) in \^
budy out of 3»mething uliich is nut fat.
$ 509. Tliere ara twg iH)a*il>l« >oiirceii of ihu maniifacturad fiu. Tbf t^^
boiiydrau* of the foiid rorin on« itource. In trattinj; of digeMion (i 2"^
we referred to the pombiliiyof cHrbubydnile* during dijcuMton in Uik aliinn.
tary canal liuoaniing; by fernientution nriivcrtiiil iiiii> liutvric wid ; and «i
(UKgWted that higlitr iiml mon- coiaplox miMii)>cn' cif tlK) Mcnvfiuiriol
Mnu miglii. bi- <il>tnii]<il oitt of carboliydnttaa by cuinowlMt aBalnnn
changes, mrmi) im, hnwvvcr, not in tbe nliaMnurv canal by m- i
foreign orgiiniwd fcmicnl*, but in tbc tiwuw tbrougn tb« iiclit-ily < n
tiwnt^ Chi^'mwlvt'i'. Wv ciinnnt na rot inioo out the Kiept nor cao vt
dctiniUrly piiiiit to uny pnrlttrulnr tiwucx other thtin tho fnt-odls ibtuutlnt
m the ivatf »( on/ Kuch changt«. But ihvi'o can be »u doubl thai oub*-
bydrato inst«rinl duva in Bomo wiiy or olb(<r give riso to Int. A rarb-diydiM^
diet is iho kind of diet niiist cfScacioiu in producing nn nccuniuliUioti U hi
in the body ; sugar or starch, in «onu* forni or olfa«r, is alwayt a )irj»
GonslitiiODt of ordinary fiitteniug fbod».
Another source of tnt is to be found in the proteids. We hanMCal^
the urea of the urine pntctiunlly reprewuia the whole of tho nitrum* ■biek
pHBH through tbe body. Now in any given qunntily of urea, tBetniOM
of OAfbon U far le«H than that found in the ijuuntily of proteid CMHaioia;
the same amouni of nitroj^'ii. Thus the peroentajKe compooition of thttw
being ra|)t;eiively,
Sulpter
1.13
100 gnus, of urea contain about ns much nitrogen as 'tOO gtau. of praletd;
but the 'iOO gnus, of proteid contain \'-i3 grniiii. (15!t-:!0) more oarboo thu
do tbe ItH) gruis. uren. Hence the 300 grms. of proteid in [laasin;; ihniii^b
the body and giving rise to 100 grms. of una. would team's b«hind 13S grni.
of carbon, in some combination or other : and this surplus of carbon, if the
needs of the economy did not demand that it should be tuininliately «id>
VL-i-td inti> carbonic odd and thruwn olF from the body, might be depooitetl
tonicwliere in tbe form of fat, It ha» been calcuUie<l that in tha way 10(1
SniK. «f proteid food might funiidh -I'i grms. of fat. We have already «eea,
treating nf tbe action of the piLULTeatit- juice (§ 249), that tlienr U evi-
dcDOe of a fatty element (vin., leuciu, which in amido-capn)ic acid, and to
belongn to the fatty acid seriea) being tbmwu otf from the oimplcs protvid
comjiound in the very pniccaa of digestion : and though, a^ wc havi* Miid. ire
have no proof iliat tliiii action nf panci'uatic jutce tabtB phic<> largely in ibe
Dornnil boilv, ito viiluit il> nn cxampli; i* amu-. the lesa important.
Soma obMrveiv have pnshtHi tbii vi<-w nf the poiduclion of (at tnit of
pn)tai<b so far os U> inHU>t Ihnt all tbn fat formed in tbc bmly arimw in thi*
way out of prot4.'id nmtiirinl, and thiii when carlMhydrati- fii-iil giw rbs to
the formation of fat it duc# ki lir ^bidding from oxidation liw i.'arbon mmety
of the protoi<l food taken at tlK> xamc time and tbiu ixinnitling it to ha
Stored up as fat. The carbuhydrnic it»elf, they ari^iie, never becomw fat but
its iwesencc allows fat to be formed out of protuid inatorial. Thw view hac
obnoualy a very imporlanl eo'rtiomicnl twarin^. viiice, if it Iw true, it is use-
less tu inorease the carbohydrate miiteriat of liiiid for the purpose of fatten-
ing, unless a siithcient pniportioo of proteid material bo given at tbe
time.
t^rtntl.
llTlLtnp!!!,
Oijsen.
Nnraavn.
Urea
2t).00
6M
36.G7
4i(.«7
Proieid
dS
7.30
33.04
lfl.S3
TIIK lilSTORV or FAT.
619
I
Tli<- view homovcr liii» Ihi-ii iimvcti to be uDloiitiblo br tc-vt-rnl mvo«tifrK-
oil.-' (-urrinl out uti iliirrmii atiituuls. It h)i« bcrii kHowh iltnt itii nnimnl
rii|Hi||y tult^notl on ■ dk-t coiMisliiig of protcJds with much c«rl)o)iy<ln>(« irill
HUiix- up far mwn fiit, thiui (-mi puc^ihly b? tccounlcd I'or bv ti».- pri>U.-t<Iii of
thr ■!)«. Thus ■ do^. iho flit in whote body hud beeii reduced to ft miniitiuni
bjr starvation, wiu fed for a ncriod on measured iiuuntilies of proteids iind
enrbobyilmica, and killed. 'I he nmoutit of fat found ullerdenih in hix body,
mnkitij; full nllowiinoe for tlie fat which remained n>W the slarvntion and
for ibc fat nccotupanyiuK the proteid^ in lh« meat };iven ah food, was futiiid
lo be lar more than could be supplied by the carbon in ihe [ir<itei<U of llie
fomi. «Tni nupposinf; that every ^oi nf thow prun-ida whirh itid not )p> to
mnke up tlie iocreaae uf the prot£i<] "Detih" of tin- body taking plaie during
tl>e fattening waa axA fur the piirpooe »f fiiriiiing fnl. Simihir experiment
on ^eese ai>d pi«^ have led to simdar nvnltn ; ami if fal he fi>ri»e<l in thiit ,
way in the IxMliiii of <wrnivora and nmnivoru. wc may Xir. sun- that ihti Ktnie
holdii kixhI for thu bixlio of h«rbiv(ira. Wr may therefore- ooiirluile thtit tilt
teai) Ik miinlructcrd in the body on tl>i- one luiiid out of proleid material, and
on the other hand by nomc direct convention of cnrbohydralM.
S 609. Ii ifi *'k-ar then that a const ru el ii>n of 611 iloctf occur in the Ixuly
S'luifwlien.'. What limitB run wo plaw on the degrco to which thii" wimrtrun-
litm in rBTTtc<l7 When the fomi containe Miflicient actual tat to act--ounl for
the fat ftnnil up in the Imdy. does any comtruction of fat lako placi;? In
tJM Bret place w« find llial wht'n the fooil contains abnormal laii< <uch an llr(^
Dot pn-aciit in the body, snennuceli for iiislAnce, or orucin (from nipe'lpre.d
oil), them Ikla are not to be found, or are found in very email i)uniilitv, in
tho fat whidi M Stored up in the body a^ n eonaequence of a larife supply of
that food. In the Becond plaeewe may call to mind the »tatement previously
made, that the compoeiiion of lai varies in difierent animals. The fat of a
man dilfen from the fat of a do^, even if liolb feed on exactly the same fuwl,
fiitly or otherwiw. Were llie fat which i» taken an food si«re<l up as adi-
pose titwie directly and without chanjte, recaurse being had to other wourec*
of food for the coiutrudiim of fat only in ctM^ where the fnt in tlie food wna
deficient, we *honId exiM-ct to tind that the nature of the fat of tho Ixnly would
I vary Rfeally with the AhhI. .So far fmui tliis being the ca«i, tlirect exi>eri-
tuentH abowitthat the fnt of ihedn^ in, ak far a» composition iicont^niiH^very
lanrely independent of the ri>od, tbnt tho normal const ituciilsi of fal make
tlKir a^giV'arance very much iw ufual, and in very much their appntpriate
pro]wrlton. though their profiorlion in the fooit niuy largely vary, and ibouvh
some of ihem may l>e wholly ahM-ni. Thus in one cxpcrinj«tt the fnl of the
bodyooiitaincdou<Mtdernble<|iiantitit«of stearin after a diet free from slenrin,
■od in anolber preserved the normal amount of olcio after a diet free from
oldii.
Of course it is quite possible that in such cases as tbeee, ihougb tlieslMriu.
or the iddn, when ab<ent from the fond, was in some way or other constructed
anew, yet at the same lime th"se ooiuttituents which were prcaeut were simply
IHiTL-x) up: and the small ^uautily of erudn prevnt in the fal of the body
after feeding on erucin must have h^u dinvlly atore^l no. Ho also, when ail
nnimnl is rapidly fattened on a diet cmiiiistink; of a small quantity of proteid
•nd a ltiT\^ iiuaniicy uf fal. the amount of fat slore<l up may l>e too great to
havo nime fnmi the prott-idri i>t' the dii-l, in which eaaa we may infer that it
WM the actual fat of the f-'od _.>imjtly deiKuiCcd in the fal-cc!l.-< of tht? body.
But «v«-n in thii* an^; lU" more ilistinctly lu the olhent, it ia also o|>vn for us
t<' sapiMiKn that all the fat taken a< food was tn some way or other disjKwed
' it, mod that all the new fal which ma<le iin appcamnoe was conslructeil anew.
620
Tll£ METAHOLIf PttOCKflSKB Of
Ami Ihe latter vi«w t« mora [wrhH[iti in Imnnony wiili lUc h!i>i<>l<ii;inil Udt
jin'vioiiBlr ineiitiuiM'il, iia irell u KtimHirhvl ■>}■ (iitivr •.>irisiiiernli<>tii>.
At tite present, ho wo vcr, w« Rwylx' wiiii'iit witlitiM- l>>ll<>wiiifr ooncliuinot
I. Fat is actually foniH^I in the animal bixt)-, and cl>i> Ikt pmwnt «t aat
Rioiueol io ihe boclr i* doI ucliiaiv«lv, if at nil. >iit nwrxly *t(>r<n>) ufi fmn
tiie fat of'tlw fiMid. '2. The nr^ii vlpntcntja of iIk iMnly-fnrtncH fnt ntay h-
e\ipi>lie<l i-iiliiT from c»rlv>hyilrate fnoH. or ftxwi ll>c wrboii i«ur|iliiit of (ii*-
t«iu food, itr from fiiU Utkeu at TocmI whicli ar« itui tlto natural omuitituuit)
of ibe hrxly-fal. 3. Tho fnl tU>mi up npprare M fat gmiiulM or dm^
dep()6it«il i» tho G<^l etilMtance of cortAin crlle. aod the increme of th« fat a
tlio cell* t* uocompanii-d lirsl l>y a gniwth, ami MihmquMttlv by a oonwni^
tjoD of tbo cell ButMitaDoe; btit^ aa in the analogmwcaw o^ f;lv'»!r«ti. ilirr>
it IM> complate evidence to »liow nhMher the fal granule* irhicli appear ir*
simply depoailed by the cell gtibatance in a more or Icea niecJM>i)ii^nl maawr.
without their forming au iulei;ral portion of (hat cell eub»t»i)crr. tho diicf
' t)ic mniiiifacture of the fat liavinf; been |^e throueh «leewb«rh m
ility itrbe from a lireaking up, a fuoctiooal metahoTiMU of tb* OtH
«ub*iaiii.-e of the tat-cell ilaelf; the latter view ia uu the whole however tW
ini>ni ]>rob<ible.
Thr Maiiuary Gi^xd.
S ftlO. 8itico milk i« a nerretioii, ani) indeed an eicretiun, the mammarr
glaiid ought mA to )m: cIimxhI ut n mcialtolio tiMue. ii> the limited mewilaf
we are now attaching ^> iIumc wi>rtUL Vet the n»ctatM>li(- piienoroena gtrlnf
rise to tW M'crrlioD of milk are ao marked am) dirtimt, have ao many analu-
yiea with the purely mntaholic cv<-n1« which tnke i>lace in atltixiao tiaaucaaJ
ao alrikiugly illuNtnitc m<'tiib<ilic cvcniii in genvrel, that it will bo lunni ooih
venieni t« coDnidcr the matter here, rather than in aiiv otlwr <s>Dni»cUon.
The mammiiry gland, formed tike a sKcnt gland, o> which it inny be mt-
■idered an oxireme developfHeal, by iin ingrowtli of tho Malpighiiin larirr of
tho i^>idermis. ie acomponod racemoM gland, cDitatructnl alb-r tlm fn-neml
plan of »iioh a ^land ai>d tbiifl oompoeed of branuhing ductn, cndin): in
MCrciing alveoli.
The whole arg»a isdivtde^l by conneelive-tinue m|Mii into a mirot>«raf
lobM, in woman alxwt iweutv, each of whioh poeeeMts a dtatiuel •lart. umo-
iug )iy a »>p<trate urific« on to Ihe nipple : the Ktnod
in (hot is uot a single itland. htit Mvi^rat itLmiti
IxMind t4)f[elher. IC»«h lobe U (\irthcr divirled bf
eonneclive-liaiUe se{>ta into smaller loliee, and tin
diviaiuu it repeated, the la&t diviaious niarkiuK iiut
Muall mtuHW called /n6ti/<'^. [Fiji- 1*3.1 The nuiia
dnct Mipplyinjf a lolw bratiehej into a numlter of
uuall ducbi,4«eh of the ulltnutedivbions »f wtiMi
undit in a lobule.
Witliiii the lobule the duct divides into u oumher
of rvluiively wide tubule* wbieh pursue a warv or
oven tn!fi4?<l oimne, aitd liear drep latn^ biilEiBff;
tbi-Bv nr.^ held bi|fether by n rtnn|»T«tively lAigu
aroounl of ooiinecltve liMU«^ Ilrncu in a motion of
a };land, oaoh lobule appean u> U' eonipoaett of a
nunber ot irregularly ruund ipfloee or abitoli, which are tlw eertiona of the
tubules and the bulgings, ami which at some pnriA of tl»e aectlon apjiear bi
be eloeed s|»aeee and at others to communicate with oadi other, or witli a
pasn^ in the centre of Ihe lobule leading to the lumen of the iluct. The
lino. in.
■'JJ - \ii^M
TnaMATiDN or rcMtHdt o»
Uii-K-iM CT III a Ckiwnui 0*
PMi.tnia (trromauiatmitei
IBjedtoa ; •ntuaed t luntk)]
iv ar-ANfi.
R21
nppi.'araiicce thus prcsonicd, nt lonM bjr h Hiickliii|^ glnnd, o^ntrmi iiiark«<lly
ivilh ihime of aa ordinnrj i;liii)i), suci) as the Mibinaxillary, by reason nfthe
iMge alveiili iriili lh«ircutispicuously wiije lumina. often oooupied by remains
of the milk.
The ductH conuets of an epithelium, rostio); on a connective-lUaue baaia
which in the c«ee of itie main ducia ia etrenf^hened with loDitiUidinally dia-
pcMxl plain muscular fibres, coiiiiimoiiA uritti the musmilar fibres present in
tJi« dermis of the nipple. Over i]>e gn^ter part of their oourM the diiciM
mn lined itiih a single Layer of columnar e|>itbelial cellH, Init ul tite in'iuthii
of tbe amin <)ucta un ilie nipjile (boe paw into an epidt-rnilit nf mora titan
one layer of Hattened fnWs. JiihI liefore opentnjiC on tii llie i)i|>tile cju-h nutin
duct i* widened into a llaak-«)ia|ie<i (.'ularjjiement. At lb« teriiiiijali<ni of tlic
small <luct« iu the lobulM, the i.>oluiiiiinr irpithvliuw ih said to u'vr plsc« to
flalteiicd cdb, m thut lh!« |iart •>)' the duut iniglit be vidl«<) a duuinle oorrfr-
epooding to the ductule uf a »alivary kIidiI.
It SU. Th« apiic-amneeM jinweiited liy the atvmli dllTur widelv according ne
the ^land i» oix* whii'h Ih being used ISir xurklin^, ur '}* one iii a rrating or
duTutant cMHlitioii, that t> to Miy, lH'r»n- any [irc^'iiancv at all linH tnk^-n pTaco
vr in tbe inlerval bi-twM'ti two HUckling |)cri<iilH. In thi-nucklitij; gland fach
alveolae oiHuiii I » of n bawntent inomUram-. prHH'iiting ibi- ii*iial chuniotera,
lined with a nnglc lavor of colla leaving a wide lumen : but the ap]>eaninoee
preeented by th« ocIIh differ from tini<r to time according to cimitnMMwee
and are not the snn>e in all the alveoli nt tlic same time. We may, Imwever,
di8tint;ut«b two conditions which, tinop they aeeni to correspond to the loadetl
and discharged condtliouN uf an ordinary gland, we may call the lo«<lcd and
tbe di8charg«d pbaee respectively, conditions iatarmediate between the two
being met with.
Iu tbe diBcharml phase ibe alvetdns is lined by a layer of low cubical or
eTGD flattened oeUs, so that the relatively large am of the alveolus is almmt
wholly Dceu|ned by the tum«n in which fiome of the conslituenis of the milk
nutyalill be retained. Kaok c«ll oonsisU of granular cell substance in vrbich
in placed ■.rounded or oval nucleus. Sometimes the free edge of the cell is
jagpid and uneven as if a portion of the free border had been torn awav.
Ill a fully loaded phajie ili<^ snpearanoes are very difiereut. The itlveoliia
' is now linul irlih a layer of tall eolumiiw cells projecUng unevenly iulu the
luniDii, ihv outline of which ia correapODdinely irregulsir and the arva of
wlikli is mueli n:<luceii. While the broader base of each cell rwu on the
bcaeDMnt membrane, the other end, fuinical or irregular, riretohes towan) the
oetitn< of the lumen, [natrnd of one nucleus, two or even more are now
present, tiw well formed and normal being pUoed nearer the base, and the
[ others, ottxa tthowiug signs of breaking or degeneration, nearer tlie frev end.
8)>mc<lmw oDtistriclioiia are seen whereby the free peripheral portion of the
cell, including one or nvore of the nuclei, is apparentlv being !«i'{iamtcd from
Ihi! basal |H>r1ion in which the reniainiuK nucleus i* Imaged ; and occai(t[>nally
•rtions or j'rw^'mentM of cells, nucleated or nueleualeHii. may be mn^i lying
II the cavity of the alveoltu. In thi- cell substance, <s)Hxrially towan) the
free border of Ihr cell, are numerous oil gliibutM ef varioiiH him* ax well as
granules or panichti of other nature; some of the lai^^r oil globulca may be
seen projecting from tliCKurfnceas if about to he extruded from the cell; and
in the cavity of the nlvoliuioil globules with a thinner or thicker coating of
cell MtlwiaiK-v are I'rcipKiitly prtaent.
Betweeu n»ch a fully loatkd phase, and a completely ditchargcd phase,
TttriMM ioteroMdiate condiliun* may bo obaarvcd. tbo cells being of greater
or !■•• height, ooataining laiv uuclvtis only or more tlian one, the cell sub-
C
S22
THE UETAUdLIQ fKOCBSSBS VF TIIS UODl
■laiKe oi-cu|Mecl uiih n fiw or witli ntiuiy <iil g)ubul*> iidcI otJivr gnuslA
ud iIm Ttw bonier itiorc or less JRg^ifMl-
Futtin^ llieee facie ia|tc4l>er we tiiav draw the fullowin^ coiii'lti^iixi. wUrt
(• aii]>p<irlc(l br Alber evitieiKv. ati to the chnngee in iIip pl>ntl whi^
character i He tl»e load! ui; mid lliedi^M.-lini^'e. UurJD^ Inndiiig the Ifiw fliiiln«J
eel! (if tbe dbcliorx^ alveolus ^'nin-^ m|>idlj, cloni^Bling into the cvliodriol
lunn, and tlie nudeus giv«a birth to two or more neir midct. MMii«hil)
KClivc iii«ta)Mli)>Ri is f^ns on in the cell tiub«t«nce, depoeiia of fat as well ■
of oth«r fulNilanvoii an- (alciiiji |ilaue. Ily vhta seeins to be of the luiiiinof
an unitrhoid innri-iueiit, Nome o1 the oil friobulee (and potnibly othtir tnailcrf)
arr fxirii<li-d fniiii th« i-cll, miid) in the name way tbut an aiiKi^lM (■stnxla
il« rxcrcmeot. But bc«Hle« ihi*, a diviaiuii of tbe cell, ihnt ia » nr|Mtnit»<io J
[lari i^f ilin^ll sit balfiticc with nu included nucleus, la lcei» jilai-*'. the ilnuirbMr-
cell thuH (liriiwii off jmming inm iIh- iilveidti* l» form |Mtrt iil' tbe milk ; <*
n tiiiddiiig III' ibf cC'll ooc'iini, |>ari of tbe cfti without a nuclr'us lioiiif mhU'
liirly caet off and iiiidiT^"iiiff a Kimiliir fiU'-. lii olhcr mi)r>l!> the i>«>«-rrtiii(
cell gixiwi., liHuh iiM-lf wiili nielulHilio |inidiii'tJi, luid wlim lotidi-d i^i-ri^ of
binlily pun of iiwlf to coiiliihiiic to ihetecrvtJnn, )>Mit of tlw ■< hu
alway» reiitinin^' a niicIiiiF. n-nminitig behind lu order to M-i'ur> ' tit
Krowib aiHl further Hivrclion.
I1ie trecreli'in of milk diHtMH from »uc)i n Hccrtftimi Or that of anliTs, aai
approacbo the formation of wbuni (^ 4:tK) intuniiiich ax tho traiiafarnwi
c«ll substance ie shed bodily to riinn part of the milk. W« mv fi>rm pan
of the milk becnuee thi« RtoBS ntodv of Kvcrctioa w «eooni|uiutc<l by the mm
ordinary mode. The cefia are at llic t»nw tinw in ibe uioro ordinary war
dUcbar^u); into the luines, water holding citliito and other confltiiuent* ii
solution. And the peculiar fcalurceof niiik, j» we bIiaII we presently, cvm-
f*)H>Dd to ibis double mode of secretion. I'erliam, however, we ooj-ht nut u
call it a double mode, for tbe one luethwl really {MtaNa ineensitily lotci tbt
other. The discharge of sodium chloride in solution from every kind o(
^laud, of mucin fVum a uiticous ffland, of oil globulce with n pnuid
envelop from u moniiuary ({land, and laMlj of nucleated tondifd cell nib-
■latice from the mammary gloixl, preaeni so many different phnew nf the
Mme act of net'TVtiun.
ji 612. The dormant rcMing mammary gland, that for tUKinnn.- of lo
animal nbich bua never bi^t-u preitnani, is much Hmaller than a 6iickline
gland, owing to llie alvi^di being liotb muidkr and lea DUUH'nxi*, Kacii
Mlrc<>lm moreover is nut n mvity lined with a i>ingl« layer of efiilbetiiini. b«l
a aolid cylinder or iiiiwn of iHMnjiaratively nmnll, roundeil iir (mlvhcdnl
cells. So long lu iirvgimiicy tlon not occur the growth of tliiiW is escHcd-
iiiglv xluw, and tne produclH of sucb metidioli.'>m iw goui on in ibein air
ciirrtcd away by the blood, so tlint under iiorniul circunMiaiioM no sectvliun
tiiki« place.
When jin'gnancT iKcura m|»cl growth of the mamma takca place, atitnem«
Ditw alveoli Wing formed by budding, but nil for a lime rvmniDing aolid
oylimleni of rvllM. .\t the approach of the birtb of tl»e uflsprini;, th(
oentrnt celli' undergo metnWIic changes, ee|iecially a fntty tranaftrmulios,
and either Iwforc or at^cr birlh ar cast off, leaving a single layer !>■ line tli*
olveult and to carry no the work of secretion as described ahuvi% It is
generally mp[><:ccd that tbcsi' shed oelU supply itic Mmlled ** cobiurum <ar
pusolee' charactorift.ic »f the tirst milk, of whii^li we shall sjieak prcMUtly-
At the end of laclntimi no abwrptiotl of nhhq of ihe alveoli take* plMe;
and in old age still further aliaorplion goes on with great dituinutioii of | '
luiniuo.
i SIS. Tli^ cntiDcrlivc tuaiie, Juiniiit! lu^-ilier the lubules of various eues.
Mtrriiuailini; t)i« tdhiile* and ruiiDiii); in be(vre«D ihe girojediii? bliod eii<b of
liw «]Tmlj tsjtliin tlie 1ii1hi1«<. is rich iu bloadi'«Me[s wliu-ti funu cnmllury
Iunirorkc ntuud llw ulveiili : u nlan carries a c<vmiilcra))l« iiuinW of
lyni|<b»lii- voaela wliicb ariK- iu lyiiij>b ^iiaoM arouuil the alveoli aod else-
«bef«. L«ucocyia are uuiuerouit in thb i)|)ac«t) of tbia (xmuectire tiafue,
and aame of tbain may make llieir way tliriiutfb Ihe basement luembriiue
aad between Ihe aM-retiDg cello into tbe caviliea of liie ulvculi ami »> appear
In tbr milk.
i 514. 77ir Nfliure of mUl. iluiuaii milk baH a upecillc (irarity of fVom
l<y21f ((> 11^*14, aitd when quite frettb [xiMenMit a digblly ulkiallue reaction.
Ii apmlily becomva add ; himI ron'H milk, even wbt'ii quite freiih, i« Nome-
tioMv pli^htlr und, tb4> cbiinice of ranvtiou tukin)( pliu-e during the ola^ualioD
%l Ute uiilk ID the nuimmary ducu.
TlMCODStiluGiil' of milk are:
1. lyoieiitt, tit., coado, and an albumin, agreeing in it* general fcaiurc*
with ortliiiarv ronim-nlbumiii, but nbicb, >iiii'e it i« raid to dilli:r i>i>(iien'hat
» ita Mlubilitint nod rotary {miter from xKruui-alhumiu, bnx been called
iilrfnfhiiaii'ii Tbo cawii), ni> ne have iioi-ii. .^ '207, uiidi-r;(o«^ through the
•etiuu of reiiiiin u ebangv whereby in*ahiliki i-umiu (tyn-iii i mako it« appear-
snc* and tiM' milk >• cunllcd. OiKin may, huwever. lie prcoipitutcti in an
ltDobaaKr<] form by ^iiiumting milk irith m^iilntl faht, <tr by tbc i-arcful
wUitiMi of acetic acid to ililiitnj milk, or by lirst luliliiiji; to the dilui«(l milk
S alight ((uanlity of lu-rlic ucid and then pnwiii); through it a strcMD of
^wrbraic neid. In the liltmte tl>e prcwnco of the iactulbiimiu, which oocun
tin Sinftll Mild variable (|uim[iti<«, may be ■luiwn by euneulation with heat, or
Lv pracipitjitiun with putaMiuni fiTmcyanide, cte. lo tlie nroceai of curdliD)^
tiw vawin, v stated in ^ '207. a))pear8 lo b« not simpljr cnan^l into tyretn
Iliut to he Bptit up into lyr^in and into another nroteiti, which unlike tbe
lactalbumin b nut couf-ulatcd by heat and vhicb u|ipeiu« Ut be allied lo
p«|*Uine or albumoae. ThU or a similar |>epluue-like body has aUo Ifcen
JuutHi in small (juantitieii even iu milk wbicb has not curdled ; it hait l>t<eu
nlln) lactopfyjuD. The lacutlbumiu, tbouub eoaj^uluted by beat when
iwlntLil. i.- not ao coi^lated aa it exiAt in the natural milk, the alkalinity
of the milk, which u inereuied by builing, preventtne thi^ .Similarly
rueiD, tbiHieb eoajiulaled by hejil nbeii ilmply aiispended iu water after
bring pn-cipilaitil, i> out ooagulUed by hcM wnen it exiMa in a natural cun-
ditivfi IU milk -. iu tlMve re«pe«U cnaebi behavea like alkali-albumin, which it
RMinbliy in otbtr fi>atur« nlwi. Hcuoe milk when bulled does iMt coagulate
at a whole, though in the Mi{>o.rltcia] layeRri expuHMl to the air changes t4ike
plam by which a film or fkin, dcrivi-d chiefly from the albumin but jwnly
bun tne cauvin, nppvnry uu ibt; Nurt'iici.-; if tbiM be rcmoveil a fn-^h [H)r[ioii
uadergnea the same chaii)cr. The jieculiar Ixnly nuclein which an wc- have
KQU, i 'i9,nn complex nitri)genou<( bmly dilTcring in eomjioiitiun from pro-
tei4ls, is also present in milk in small ([uiinlitim, and aonmling to mhou
sbaervcre ts (imply sm^Rdinl. nut mlly in solution, or it in aoinc way
peeuliarly asenciated with the ciuMn.
2. J-'ait. Thcw arc, in the mniu, palmitin, stearin, and olein; bat otltvr
&la, supplied by butyric ami otlw^r fnliy acids in combination with glycerin,
accAmpany the above in small quantiliva. In (his respect tbc tat of milk
reaemblet that of adipose tiwue. I.ecitbin and cimlnteriii are also prceent
iu very «nuill quautitv. as well as a yellow culonug matter. The fat prownl
iu milk differs in drUi^rcni animaU as to the relative proportion of olcin,
|»luiitiD. and stearin, and as to the kinds and relative amount of the other
Dlier fai£.
4)34
TKE UKTABOLIC I'ROCKSBXS op Till
TIm lutxtiire (>r lh«tie fAt*. fliiM nt onlinnrv trinpiinilum. Is pnanu m
tMUiral mtllc iii tltc form of gl<>t>iili!n of various iiijtiv but for tb« meat pm
oxcediii);!)' tftuutl (iii man frim 2y<< to .^m). Milk is in IVict a typical onul-
i>ii>ii, mill it t> lti« |>TVM.-ucc iif th4! cwtcin iu tho milk which briiip mhoitt ibt
«niulHititi. Some ()lim-rvi-r> miiiiitiiin tliMt euch f^obul* <if fat ia surrotitiM
by an fDVdlitp or mi'mtirnnu uf Kolid iin<li«M>lv«() raaeia ; but. chougb W-
<ltiul>liylly even wlivii ilit- lilt is rvniovtil from ihe milk raoli Klobulv nimaiai
Mirr«tiit<t<.<<l by n lityrr nf milk pIusmK, if we may to call it, rioti in owtn,
tU»n> »K- iio Mikijuiitv rMutuns for tliinkiiif* tluit the cweiii uctuolljp form ■
Rwnibniiii'.
On iiMndiiig ii gn*l (]<*l of th« fat colleota na th« top of tho milk id th*
form of creiini, but iu thi«. (u in the butler wliicb ia furnted fnim It, \ht
globtilcs are ftill discrete', eo lon^ nt I«a8t lu tlie butter is " fnsh," By ti>
UK ttf a ireotriftigal tuaohine nearly the whole at' the &t may b« Mtparaicid
fh)in the plasm*.
3. Milk at^r or lactate. This is very afit to un<Iergu fi!rm<-ntJUton iutu
lactic add, through the o^ooj of aa oiYaiiiMil femtcot; thu niiJk ibiu
becooiefi aour, and the eai^iii ia )ireei|iitat»l iu a lloccuk-iit furm whcai thi
acid i* itroduced in sulfidenl ijuautily. Siuci- ihii chaHgi' will take plaife
vv^ui when t-rer^' tare is taken to exoliMle g<-rm> fri)m ihit mtmiHplK-re bavinf
aoona tu tbi> milk, the nrganiieil fonnonts mi.i;>t bv pr<'m-Mi in the milk in th«
duct» of tlu! glaiKl.
4. -Salu. ThouK)> trnoa* of umt and kreiuiuin have bvM Doted by boom
obmrvpn, thccxtrnc-iiri'sof milk.lH-yond tlK^K-^rithin iinJ choloal«riii already
mentioDMl, aro insignilimit. Thv calts nrv of mora im[Kirtai>cc : the«e sic
obi«fly calcic photphnlc, of wboM> function in thu procum of cunllitiK we
spoke in $ 'J07. and potuaxic and sodic chlorides, with n small qunuiityrf
maKae«ic phosphate. Sulphates appear to be a been t. A small <|tntiitity gf
an iron salt ia pmeni.nnd traces of nilphocyanidc have been obsarVMi
Besidee the pho«pbr>niB in the actual form of pliosphalea, milk ooBtldliaa
furtlier ooindenwlfl quantity of phosphorous iu ilie protetds and in llie
uuelein, as w«ll aa aoine sulphur in tJie former. The iDor^anic ootwtltuanu
of milk mar, hroadly speakinf;, be said to difler diDtineilr fnmi tboM of
blood, and to much more nearly reeemble those of tlie eniin' body.
The ompiMitiou of milk id tbe same animal varies widely nom time lu
tiu>e. and Wide« uiider)|Cieji marked ehengcs during the periixl of laotali-m.
Tli<- retaiivt! ^neral compoaitlon of human mitk and liiat of the ooit, tlw
mure, and lh« biU-h may perlinps be Nbiiwn by ih<! follnwing tabic : but i* it
difficult l<> draw na average, Mnoe the individual analvHC^ given differ M
luucb : the lifcurvs givau for ducin and fat in the milk of thv hitch may b«
unusually bi^,
Anrifjfe Cotupoiition </ J/i'U' in D^ercnt Atiimai*.
a'tnui.
OW.
Hm*.
mica.
Ca»ein. «tc..
4
4
S.5
10
Fat*.
2.75
4
t
10
Guitar,
StnTts,
.5
4.4
6
8.8
i>.23
0.4
tl^
0,6
Toul golide.
10
la
lt>
i*
Waur,
90
«7
flO
78
The <|uan(ity uf railk secreted by a woman in twenty hours at tho hi-iglit
of lactation has been calculated at 70ll to MM) cc. A good niiluk cow nil)
f ii>Id about 10 litres of milk per diem.
t 61ft. (.Woofnnn. Thix i.i lliv tiiiiiie fi'ivm to the milk »ecrele<l at rlx- ht-
[inntni; of a [icnoil of Inoliilioii, jij^t iiefi>rL^ niti) fur Mime i\an uIUt |iurtii'
riiiuo. Tlin milk (litn-n> Trom tlii> ■ul)i«e(iu«il milk in microttGO{Hcal cbnr-
•ct^n Kiwi in rbcmti'al r<iiu{Hi><itiuii.
WImu ordinnrj milk \* txtmincd uiidrr (hv roicroM-npv hiirilly itnylhing
i* M«a b(«i<Ii-» ilii- Tnl globulM txcrpl u vcrj Tt-w im{>rrrri.-t ci^llii or piirliona
nf ckII*. eoDsuting of nil subetniitv mon- or U-m lomlc-il nilh fnt aiitl coo-
-iMttiint! Hometimce » more or Km HlUml »iicU'Ui>. A fi-n minute ^niDulM,
khoii^lit h_r some (u bo (wrlicln of »ui>pen<lr<) cBocin or nuclcin, nrv, however,
'•Uu V IN hie.
Colostrum, od tie other b*nd, contains « Inrgc number of n-lls or oor-
ptiaclv, nbich have been culled "eolottrum cuqntFcled." Some of theae
eluoelr memble leueocyteB, otbere are either cell* of iibout the 8aoi« riie,
ruMnd or irteicular, and [lOMefcing a nucleus, often mifsbkncD, or are meretf
|M>rli>inA of cell aulMinnce wilboui a nucleufi. In nil of tliem llie ctil aub-
«Csoc« mar !» loaded with fat j;lobul«« or amy be fairly )>*e from fat. Sonic
of tbnw colls a|>|>ear t'l Ix- undergoing dittinieK^Btion : »<>m« niny at a fnror
Mble tetiiin'ruiiire cxliibil alow amoeboid movements, and muHt then at lout
be KnrdFd m living.
CoTr«tram uImi difl<-n> from ordinary milk in containing not ^nly a Inrg*
qnantiiy <>f al)>tinilii > tacliilbiiniin), but al»u a decided amount of globultn.
In eott—quewcB of tbi*. coloMtTum difTcni from milk, inaamueh a> it iadia-
tXiittUy eM|nilKt«d by bm.
Aa >iati.-d alK'ie, ilurinjc tbv rapid growth by wbi<-li i1k' ^'Innd i« enlarged
ttr«mrmt«ry to Inrlaiion, the alvmli nrt> at Mrnl Milid niu.vK^ of cells witb
litue or no lumen, ami a liim«n is r«tnlili>>heil H<il)w^(iifntly by the discbarge
«f 1^ crnirnl cells. It is UKially su|ip(iM'd thut llic ci'lU to discharged.
•DOM uixlerguin); much, o4herB comiwralively little, chaiigv, supply the
«olaeiram eorpu»cla just spoken ol, and at the Mime liniv funiish the
alobulin and eieen of albumin also chanicl eristic of colostrum. But thia
m Dot certain. The alveoli at ibis lime contain peculiar cells reaeubllog
faloatrum curnuadea except tbut ibe^ are tne from fat : and it is auggeMea
ihu tboM b«ing diHeIiar>,-cd uml taking up fat in amtsboid Ikahioii beoone
eDlMtmin corpUKclex. Sume regard the eoluetmm curpuaclea as aimplj leu-
IContM wbidi have similarly inkeii up Ikt.
f Slfl. Tbe awmmarv gland is pmctil Ixitb in the female and the mule
cfaild at birth ; and in bcilfa lexes at iind fcir a ffvr dnyH al\er birth U thrown,
n roiuraon with all the olbrr w-creting gland*. iuti> wcretory activity, and a
HDall •(iiaulity of milk, the "ii:itehn>' milk." ao culled by the Oeiinang, is
■lisdiargeil from iho nipple. The milk rc«cmlde» in nil cafcntinl f^Uim
tlie milk of lactation. In both wxco this initial activity soon pniMS off,
the xlanil in the female fonher developing at puberty, but in th« mate
remaining, save in exceptional eases, in its infantile condition or somewhat
r«lrm[n»linu;.
i in. The tirreiiim »/ ntitk. From nhut bns been already said, it b
obrfous that Ibe secretion of milk, while rewmbling the secretion of tlie
other accreiiiig glandH uhich we hare studied in liein^ eseenltally an activity
of the cfNtbeliitm cfIU lining Ibe alveoli, neve rib el iim pn««nu ivrtun in-
lereMing foatun-n ijio-bl to it^olf. If the account giwn in ^ >31] be a true
<•». miirpbolugieal changts in th<' cells are more pmniiDetil ihaii in the caM
of uthrr L'tandv; and wl- may iiilt-rpret the npjtearani-eH there related aoni^
what nx I'lltown : \Vbe<> tlw <Ii>chnrged giaud with it^ low epithelium begioa
Uir Work of lundin^, the celt* dixtiiiclly " gntw." Their cell tubstance in-
iHiawt in bulk, awf elongaliug timjceti int" the lumen of tbe alveolun. At
r tlu «aif lime tbe nucleus divi<K-« aa if ihv ctll neru about to give birth tit
40
TnB J1£TA»0LIC rROCBSSBS Of THIE 801777
new oelh: but at Am, M nil «veniH. Ud itivisiun of Uie wll Hubauuice
place, mid the uen nuclei lie JinWiiOtMl in a cooiiihmi cell Ixxly. '
subaUnoe inevnwliilt? (lutit ua ftecnuorv aclivitv ; it (}i:|HRiiu ju ilix-if lualeriil
to form milk. Tbi* ilvjKxiit of Tut ia cuiiKpicuouii atkl ctmily n^oftniiiil, liui
wi- may furly iufcr tlinl llw (itbcr I«m voaily ilixtiiiguiNhed jiroLoiil und (uiil>>-
bvdnU« mntc^rinl:) are dujuvitei) id ihi; c*Al vulwuuive in a niniiliir lisuibina,
"ftiftn fulloii's the (--jiflimi of llw prvpiirvd material, and tliix ninT tmko pUo
in ono of two way*. 11m- oil f{Iobult« of lat may l>c proirudis) tmni tbv crW
eubataiioo mucli iu the miiik' way tluit an anuvba fxttudw it» i-xi-rotntnl, as.!
E«ribl;^ otlwr c-oii>titiio»i« nf milk may tic i'j<-<-t(->l by ii utmilar metbod.
lit, bttidea thu. tbo dHrrrwl oiiU divtaioii now tukta iitniij in a eomewhat
imperfect fnsliioD, m that jwrlioiia of ibi- >M ovll ciirr>-iti^ nuclei nilb ihtn
come aaiinder from tlic reet of tbo cell in wbirh a nurlvux ta lell. and In
loose in the lumen of the alveolus ; jwriions of oolt su)>»taDoe free Gram
uuolei appear also to b« ciMt oA'. Here, in the lumen of ilie «lve<iliu, they
rapidly under)^ chan^ ; tbe cell aubetance is allerod ami ilissulvexl. awl iU
load of prepared material, probably UD^IeiKoiD); in lite nci some ftinher
chaiwe, is mt free, the nuclei aleo undergoin); cliaof^ and t>ei.-iim(nK ulli-
mnuJT broken up. Hence the oonsit(uent« of milk are provide^l for, »>!
only an in o4her i;landd by tlie inalenal wiib ulii«li the cell loaiU ib«lf Had
MiljWHiuejitly diaohargea into tbe lumeu of tl>e alveolua. but hImi by the
actunl ■ultntance of part of the cell ilaelt The i.-liaracteriatjc nuclniD of tb>-
milk biw ibn* it* onjiin in all pFobability in the uliwl nuolei of th« iccreuag
oalla, and we may {>erlva|u iufir that tbe mill more characteriMic caatlll
exiat' ill milk in Inu form of <-n.ii-iii and ni>t uf iconic other pruteid III iniiM
aueDC« of thii' inliirvealioii uf lh« mrUml cell Muliataiioe in the fonnhtiao of
to milk.
It ia hardly iHrrt-Kuiry to add thai ihn* bodily coniribulionauf ibaMuratlng
cell Ui the H-cn-lioii nrv iiiHXimiiatiii'd by that mora nnliiiary |iai1 of wi^rflioa
whi«'h fiiniiii't« in lli>- How o) lluid mnluining various matlvn in Milutioa
Ihrotigh Iht celle inu> the alveoli, iba general oompoailloB of tbe milk being
thus secnrml.
I; A18. TheM-crotion of milk ilion would npiivar to illitxirat^-, even tnori
fully and dearly than do other glands, ibc tnith on nhich wv hnvt- ao oftca
iu^ted, that n secretion is eminonily the n»iilt of th« mxliitiolic actiritj of
the aeotMuig cell. The blood is the ullimiilc souroo of milk, hut it beoonec
milk oulv IhrouKh the activily of ihe ct-ll, and that activity oonsista lar^y
in an>eta*bo1ic manufacture by the cell, and in the cell, of the comrouu thiajp
hr(iu)chl by the blood into the epeciat things present in the milk. Kxperi-
meuud rttfultK tell the same tale. Thua ihe (|UimtUy of fal present in milk
in largely and directly increased by proietil, hut not increased — on the co«-
trar%- dimiut^ied— by fotty food. This effect on the mammary f;lnnd ia
particular is in aooordance with what we shall preKntly learu to be tlic
geiK^nil elfi^ct on the body of proleid in conlrast to that of fatty food:
protei<l fixHl kociuh to iurivaM the eeueral metabolic activily of the IkmIv.
white fatty foiwl leiida to leMCti iL Momiver the pniteid food bcouh m-tiially
W funiinh (he fal ; aiid we liavr ulrondy >u {[)<•'' l*i I a maimer iu u liich )>ni-
beids may pve riae to fal. Tliul tltv lat of tint milk nred not ni-i'ew«rilT
oonw from ibe fat of the food ii> rliown by the follouinj; (.-xin-rimeut ; A bitcn
fed on mimt for a giv^Ji periixl gave off morr fal in \»:t milk than she oouM
ixwaihly have taken in livr f<H)d ; and thl* mon'over tiuik place white abo WU
gainini; in wei)|:hl and " Inyine on fnt," »» tlml ^hr ooiild nut have supplied
the mammarv gland with liit by i<imj>ly tniiwlcrrin^ tiit from (he ttore pre-
viousty exiftling in ihv iidi[i<i»e timu« of her bmly ; sbo nppun-iil ty oblainei
ihe bt ultimalclj' Ihim tlic proteids of her food. Aim) the hiiiologicul
ahovf favor rliv virw lliitl the lorRintioii ff lilt out oP prot^il* iti »ilu1l
tmkm fAnei in llif tx\U of llio iilvmli. Thi- PXiioriniAnial l)ii:ai m noil
«u thr hi>iiil<tt.'WI vvtilviic* gom to riww Uiul till- fnl nf milk i» r»niK-<l in ttifl
<.'<-l1 Biiil l>_r tltu (vll, mill i« Doi timplj gathorol out of tlio blood.
Th« cuvin ill n similar way 8nn» to tw fnrmvd l)y thi? action of Hif calL
It cannut b<- gxiht-ml oiu of Ibc blood, eioce the blood cootsins no n»)
~n: it iniut be funiica) in the gUnd. Soma observeri< hsvc tnaiitlnined
•hat when milk is k«|>i nt '-i't". ihc ctwtn is incrcmod through some ferment
»(Xti>« taking pliu.-e in the milk itielf : txit this sceius not to b^ the ciuc, and
tt>c furmalioD <y( Msein miisl he regarded nt the neult of the action of the
cell. £vm the albumin present nppenrs to he uot ihe ordiiinrr ■enim-
mlbumio siniplj paMed fraoi the hl'iud ihroitgh the cell into ihe tumeo of
th* atvetdus, bol the stightlj different lactalhuiuin. We may iwrbups re^rard
the albumin u leas difficult to mHnufnctiire than the ciMein : and we may
vxplain ibe laoi that relatively to the albumiu the i-aaein is le^ at the very
berinuinK anil otpecinlly towanl the «Dd of lactation, hy supiKiaiiiir that the
ttU hat iu [Ik lint cane not g^t into full ir»rkiu); onler, and iu UMMoond
cMa a waniuK iii fin-vr. The peptono like body in milk, though «mall in
aunniity, is a furtlH-r indieati^m of the pruteid metabolium tukin); pluoe in
inr ffll.
lliat the niilk-KiiKiir, liu-litw, aUo in formt!*! in and bv the cell, is indicated
fay llw facia that it i» fiiund in mi other ii^rt i>f the Wly, ami tha! il.i prua-
ncw in milk is nut dept'ndvni on (■■irbniiyilnilv foml. fur it in nuiintaini^l in
ahundanoe in tb* milk »f nirnivuni nhi-ji ih<w are fed excluitivoly on m«M.
M froa m poaiibl* fmm any kind "f iiugtir or glyco)£«n. A glvcogtn-llko
boily ha# iiKircovvr lir^'it dencrilx-d iw existing in the cells, and it lii auggMtod
that this UhIv i* lli« antccodcDt of Ihe Inctow.
We thuD havo evidenco in the mnmnuiry gland of the Ibrmation, by tho
uetsbiilic activity of tho wcrcling eell, of the repruentatives of the three
jtmt cluMS of food'ttul&i. urotcids, fata, and carbiihydratn. It is, of course,
■{uite tnie that all Uie cell Itua lo do may he simply to turn aside into the
•pccial cAiteiu, (hit. and lactooe the seneral sumily of proteids. fau. and
(iu^>»hy>l rates brought to it in the blood, without these over het-oming
actually part of the cell, the formation of Ikt nut of pr»Ieid «[>>'>keu of
above takins place in •ome other part of the bcxly. Still it b ojien for us
tanippoae that iliey ate all three formed in the cull iiaelf out of the com-
pralmwlTc living cell 9iilutim<-c. [f ne accept tbr lutlcr view, we may look
apoa what is taking pliux? in the timmninry cell iw n pictuni: of what U
roiiig on in vnr><Hi> living liiuucs. If the fut of the milk wore not ejected
from the maiiimiiry celt, the mammiiry gland woiilil become n iniua of
•dipoae tiaine, <»|>in-iiilly if, hy n sliiihl change iu iho metalioli»m. the pro*
duction of jal wen- exaltnt at ihu exiiensi' of the production of casein or
nilk-sugnr. If, again, by a similar flight change th<i milk-ntgnr wei«
accumulated mther llinn the tut or protcid, «e should hnvc a n-jtiit which,
hy an easy step, woiil^l bring ti« lo glycag«nie tiflHic. And, lanlly, if the
Eroteid aecumulntion irore great«r than the fntty. or the aaocharino. Ih«l*
eing carried off in some way or other, we should have an image of the
natrilion of meh n tiaue aa muwle, in which tbo protaid constituent >• in
«xceeB of the other*.
( 516. That both the secretion and ejection of milk are under the control
of the nervouA systetu is shown by common experience, hut the exact
nvrvm* mechanism bu not yet been fully worked out. While the erection of
Che ni|)p1e oeoaea when the emnal nervod which supply the hnsiiti aro dividiil,
Ctic M-cn-iiou euntinuuB. and is not arrested even when the lympattieticaa
^lell lu the spinal nerve* are cut.
CHAPTEK T.
XUTRiriON.
TllK STATWTltM OK NwilITIOK.
^£30. The preceding chapter bos ilionn at how wholly [mpOMtUe i^Ki
at preeenC lu muster the metabolic [)heooiuena of Ilie body, br atleiuptiii^- to
trace iiut fcirwnrd or backward, the several chaiigea uiiderfpine bir the in^
vidual louBliluenl* of the Ibod. the bmly. or the waste prcxlucla. Anothrr
nx-thml h. however, open to us, the itatUticoI Diethud. W« may ttvtrtmis
th« t»tal income and the total expeoditure of the body duriug a gimii
iH-rivd, and by conipariu^ the two luaj' be ublu to draw co»cliuion» cnnccni'^ ,
ing t.h« chaoges which muni have taken iilac« in the biidy while the inconiej
wnM being converted into the output. Many newarchi-s' have bn-n cariMl
out by thin methoil ; but valuable uii are llie rntultjt nhioh hnvo tH'Cti thertby
gained, thrr must Im^ received with caution, juuce in thiit metb'x) ut' inijtiirj^
asmall ■.'^rnir in the dalH may, in tbr [injcettn of calculation and inferetieC)!
lend to mtwt wrong ciinclu»ii)uii. The grc«t udc of nuch iniiuirie» is to eaf
geat tdcii*. but the vicw!" to which thtty give riw iieetl to be verified in other
ways bcftin- they can iicciuin- real wfirth.
Vompogilion of lh( milmnl bodii. The first datum we rMjuire ii a knowl*
edge of the cwmpoeition of (he body, a* far as the relative proportion of tbv
TKiious tissues is concerned, lu the human bodv the pni|Hirtioiis hr weight
of Ihe chief tiwues, in the fresh state, are probably somewhat aa follows:
A4ull nuui. Nsaham lalqr,
fernni. l-MnuL
Skeleton 15.9 17.7
MtisclN 4].« U.9
Thoracic vincera ].' S.0
Abdoroinul vueera 7.2 11,0
F-l 1«.S) «,„
Llrsin ].$ IS.S
Ao toaly^ of a cat ba* given the folkming nwult;
Muiwlm and tendons 46.0
BonCT H.7
SkiD 12.0
Mesentery and adi|>o»e tiun« S.8
Livtr -LS
Dtoud (racitpinji M Omlh) ....... ftiO
Other (irgani and iiHiues I8.T
One point of importance to he ui>tice<l in these annlrMS is that the skeletal
muaeles form nearly half the body ; we hiivc alrradr seen ■ $ 38) llinl aUoal
a ijuarter of the total blood in thv budy is conUined in tlwn. and h*r«
yiTKlTION'.
^■mSnaif ft 4&S) imiHlod that « Inr^ part of tlie itivlubulutm of llio hutly !•
outin on ill th« muiicln. Nosl t«> iW mmtclfu' we niunt plnt-i; lli« liver, for
tbuuf;h Ikr Imv in IhiIIe (liaii ih<<i|i, il is Rubjrct to n vi^y nctivi- mctiiiwl-
iin : ifaifi i* minnitvd tiv the l«cl ttiHl il Htonr may bold about a (jiiartirr of
tW Kb<ilc bliHidTand n also indioMcd br tho Dum«rou» fncU brought IwfoN
o» in tlK- prvcwling chontcr.
$d31. Tlir tlarrinrj My. Bt-fon; altempting to stiidv the inHiicncr of
Ibud, it will 1m- tiKlui to AfCi^rtkin whnt changes occur in ihd h'xly when nil
fnal if withheld. A <-nt of known w«if;ht wu starvfyi for ihirtci-n ilitTx.
Al the bf^nning of iho period the body was prcsnincd to have the eonipoM-
lioa ^ven above; nt the close uf the ]>erio<) a direct noalysis of the body
WW niaile. From thin it aiipeAred that durinj( the huni^er period the cat
kail I'let 7-1-I f;ranii)ieH of «o[id nialertal.of which 246.8 were fat and l\>i.'i
auacJe, the retnaioder bcini; derived fnxn the other tuau«s. The [lerceut-
•gn of dry Mtlid matter liwt by ihv inure imporlaut tinu«H dnriiig the period
««n aa follun'* :
AiliiHiM titaue 9TJSt
UdIvm «S.l
Lirer &t|j6
MmmU* 90.2
ilhwd lT.a
ilrvin atui njnMl mrd 0.0
Thu» tite l<Mu (luring' slnrvntioo fell muHi heavily on tho fat. indeed DMri.V
the wholf iif thifi diaajifx'nred. Nnst to the fivt, (he glandular organs, the
tiaauea Hbioh we have «(wa to be eminently metabolic, suffered m<ial. Then
eurae lb« niUKlM, that ia to aav. the ekeh^ul ninacltt. for the linu in the
heart wa* very trilling ; ohviouily lhi:< orgnii, on Recount of it» iniportjiDoe .
in carrring on ihv work of the economy, hub spared m much lu powible : It I
^WAft in fact fed on the mt of the body. The «ame remark nnpliea to the *
br«in ami spinal cord : in order thai life micht be pralonroa as ranch as
possible, theee imprirtanl orLtmH were nouriahefl by material drawn from Ims
noble orfputf and ttMues. TIil- blood aiifibred proportionally lo tho gonoml
IxMly-wasle, becoming gradusllv lew in bulk but retaining the same ipccilie
Hmviiy ; of the toul dry prmeid eiiulituenu of the body 17. •! Jicr cent, ww
loM. which agreea very fli»elv with the !7.6 ner cent, dry lUHterial (almoM
■wbolly proteiil) lust by the bliXKl. It in ivorliiy of remark lluit the tissua
in geoenil l>ei.-iuue more uaiery than iu health. Similar obaerrationa on
otb«r aniiuii] have led to similar reMihii, the chief dJ^oonlaDoe being that in
•OOM cumb the bouce have iuiremi conaiderable Iim. in oiliera comparatively
Uttlct. We might be ineliiKn) to inlhr from tbv» <liiiii the ouncliiaioiiB tfaM
nieta Ik ilium in mmit ni-tivo in the iidi{Kitc limur, next in *uch nietaliolio
tMHic* a« the hepatic celU and (]>lecD-pulp, then in the mufclcs. And to on;
but we hare no warrant for ihnv conclufrion». BecaiuH- (lie ln« of cardiae
and nervmis tistue wa> »o «iniill, we niiiRt no4. therefore, infer that their
BCialHdi^m wm feeble; they may have undergone mpid tnciabolisin, and
yet have been preaervod from lue» of lubstanee bv their drawing U]xm other
tiwica for their material. Tlw grml lorn of adij»wc tissue is obvimi^ly to
be explaii»ed by lite fad that that tinac laceBenlitillr a storchoUH of mate>
ml. and the eimilarly ^n^ui though Icaa loM in the spleen and liver indi-
caiea, u indeed the facts recor<^ in the previoas chapter suggoat, that
ihaw organs too serve in part a^ storehouses.
Daring tbu starvation period, the urine contained in the form of urea
(and that practically represents nil the uitrogeu of the urine) 27.7 granimea
i
XCTBITIOS.
of Dhrufiti). Now ibc ninnuiit of mii»cle nhich wan Iwt diiriii^ llii: {xtriixl cun-
tuinctl lUMHil 1>>.'2 »f (litro^i. Thii*. more lliaii linlf Ui<' nitroifi-ii of ihr
output during the nUirvatioti |>rrio(l imii^t h«v« v<>iiic ullimalrly rn>m tbo
Riclftbolism Df niUHuUr linui-. This furt ire haw nlri-nHy fiHsi in tliMur-
eiiii; ihr hietury of uren and bIihII have occn»)on to Rinki; Tiirtbtr u» of il
horeafier. Tbt< amMiiit uf urm excreted |>er dieoi 1>r« l>een ulwervnl in
some cfls«8 to fall verv rapidlv duriii|i the firet dwj or tvo of BinrVAliiiii. ami
tlicn lo diraiDiBb gradually, tfioup;!) ufleti ehowiiiK w>Qsidierable irreitulantlis.
Ill i>ih«r cases Da such lar^ initial fall haslieea obeerred. Ii is ruiwt
marked in aDuiiala which have b^a well fed beforo Ibe liej^inoiiiK of ibc
UarTatiou, eepecially in tboae vrhicJi bav« had a ricli nilro^Doui diet ; and
llw diseharife in ihese caaea of an extra quantity of urea iu tlie Orxi day or
two is dhviounly conoected witli that iiuii)ediat« eiR>it of TimmI on the exi-re-
liiiu of iirra to ubicb uc bav« alr«iuly (M^^^ reftrrvd and (n wliii-b we
skitll have to returii id 8]>«itking uf wbal u known ww " luxu»-tii>uituni|)tiou."
OmparUon of Intomt and Output of .ValfriaL
i 582. SlHhod. We have i>ow to inquire how tlw eleiuenta of fixHl an
dial rib II led in the excreta, in order thai, fmiu the manner of tlie dbtrihu-
tion, w(^ may infer llie nature of the interinediat« aiaget which lake place
within ibe body. By comparing Ibe in^eslu with lh« excreta, w« iball learn
what elrmcnlH lia\'e been retained in the body, and what elemenU a|>|H-ar in
the <'Xcri.-tii which were not [>resent iu the foixl: from these vn inny infFr
tliv cbuDgra mliich the IkhIv hai umlergime ihrougti tin' inlluencv of ibn Tikm).
In th« finti phict', the real inooiiti^ mii»t Ik rliilint.'ui'tici] from llic npimrvnt
ont by the ttuntinctioti of the feci'!>. Wo lutvi- rHVu tliiit by liir ibe Kri'-alvr
part of the fccw i;" undigoHi-il inHltcr. i. r.. food which. tbouKh |>'"cird in the
alimeiilnry caiinl. ban not rcnllv entprrxl into ihe l>ody. The >>lian< in till
fet^'i'g lalicn uji bv nmltr-r nbich Imit been cxcrelci) frvm the lilootl tolo llit
alimenlnry cannf. i» *i> xniull tlwt it may be neglected : cwrlainly, with r«van)
to uitroi;cn, the whole <iuantitr of this ekmonl which is jireecut in (be fvcw
may be rcf^jxled n» indimiing dimply ondigeeted nilrogenou* matter.
The income, thus oorrcclol, will <wii«isl of so much nitrnf^n, carUin.
hydrogen, oxygen, sulphur. pb<«pbonis, saline mailers, and water, contained
iu the proteids. fata, carbohydratee, snlto, and water uf the food, tovetber with
ihc oxv^trn absorbed by Ui« lun^, skin, and nlimenlary canal. Tlie output
may be rejnirded as consiHtin}; of \ I ) the reBfuratory pnxlucU of the luup,
•kill, nud alimentary- canal, conniAtint; chiefly of earl>onic acid and wiu«r,
H'itli xmall iiUBUtiiiee of hydro}^u and carburetled bydn^een, these Iwu liUlcff
coming rxi-luiiively fntin ibe alimentary canaJ : (2) of perepirHtion, coilliil-
{ui; cbirtlv (if water and ealid, for the ilubioua excretion (!«e s 4;iU) of una
by the »kni may Iw neglected, and the otiter or):ani« conntituenta of s««al
untount to very lillle; and (3) uf the urine, nbicb in il-wuiihiI io contain all
the nitrogen really excreted by live ImkIj-, biwid^ a lur^o ijuaniitj of utlino
matter* and of nnter. Where great lu-i-urauy i' rei]uire<l. iJm; total nitiD^cB
of ihe uriiK' oiifjbt to lie determined; il t« maintaiiH^, however, ihnt no
error? of wri^tnn tmjiortaiico ariw wlieu the nrvu nione, aa deiamtined br
Ltebig'n nielbod i ubicb was largely ui>ed in the ri-»eiircbr« fitnaing the haeu
of ibe premnt dikcumion), in taken as tiw inewuni uf tl» UAal otuuititr of
nitrogen in the urine, sinc<\ in ibio nM-tbod, <itbur nitnigeiwiia bodica lieeidw
urea are |>recipilal«<l, and m> contribute lu ibonuanlitalive rntull. It hna
been, and indeed Mill is, dcWted whitbcr the body may not suffer loaii uf
nitrogen by other cbaniK-ls tbnn br tbo urino and fec««, wheiber uitrttgm
may not leave Ihe Ixidy by iho skni, or, indeed, in a gaseous aUle. by the
Iinliini-r »i' ihn coiiflioting cvid<-itr« weni*, however, in t»\X
__ Tim tbat no «uch Iota inkn iilncc. It would n|)ucnrlhBt though oitrogen,
xhr |iivul, M i(> f]trak. of tlic i'fi<^mi(al cluitigM nf living l>riiigs. fvrniH ao
I«rgi? K porlioD of lh« a(mtie|>hcri-. iimJ. moivovFr, is phvKiciilly iltHiiMKl
vbrouf>h ihr boilin of bnlh |i|nnl« nni.1 iinimiiU. tVre nilri)^en n ul' tiiichiimcal
to diher uf thvat. It enters inM iind rcninins In ihcir bodicv us nn invrt
I
*ul>'tiinre. niii) the ninn);en iihich lenves a piiiiil or animal, in a gmraua
»liitir, is «iiu]tl_r a part ol' the fame inert supply, and does not cone m>ni the
tirrakitig up of ibe nitrogenous etlbalnrifee of the borly or of tbe food.
Of ili««e eleDieols of the ineoiue and output, the niirogea, the carbon, and
Ihe fre* oxj'gen of respiration are by far ibe most iinjioriant. Sioce water M
of UM to (be hotlv for merely mechanical piirpoaes. and not solely as food id
the Mriet aenae of tltc word, tbe hydrogen etenieul t>eciiniett a dubious on« ;
th« tuipbtir of the tir^eids aud the phoitphorus of tlie fnt^ are inH^iitieiint
In amount ; wbih- llie aijine nintten stand on a wholly dilli-rent fiKiling from
tlMolbvr Mrt* of food, ioaainuch iu> tbey are nut wournv of i.'ncrgy, nod pfMS
ibrougti to* body with conipaniiivcly liitle ■•liaiigv. The biidy-weight must,
of CoarM. b« oreTully a*ctTlain<^l »t thf bi'ginning and at tbe end of Uie
pftiod. corrrctinii being made wh«rv jiowilile lor tbe fctN«.
Il will bi' frttt tbat the laborof auch in(|uirioi it eoDnid^'nililc. The urine,
which tiiu^t bf rnn-fully kept a^Hrale from the feces, rciiiiim dnily mc«>itny
iD0lt and nnnlyiiK. Any loi» hy tbe »kin. either in tbt; form of xwMt, or, in
ibt rwc of Brmltv nnininU, of Imir, miivt lie r«timateil or nccouiiled for. TIkC
food of tbe perrtjal mtiM Ih-, ni> fur m* poiBihte, iinilbmi in cbaracler. in order
that the analyws of »|N-eiiueiii> tnuy serve faithfully fur calculation* involv-
ing the whole luantity i>f foiMl tnlcen ; and tbio is <-8pccially the en»e wh«n
~:ie <liet n a meat one. ciiice tiorlions pl'monl dilfcr tiy much from each other,
lut tb«gren(eBt difficrilty ol nil li«; in the estimation of tbe cvhnnic a<-id
rudueeir and the oxygen coiisiinned. In some of the earlier rewarches this
or waa negleeie^l. and the variationi ocourting were simply gueweil at,
titrough which veri- serious erron wvrv intrudueeil. Nu comparison of
iaconnc and output can be eunsidered saiinllnetury unless at least the carbonic
add jinxlured be directly measured hv menn.i of a mpiration chamber.
And in onli-r that the cunipiirison fhoulii be really complete, tbe water giwa
off by (hi- xkin and tiiiiK» mtiKl he dirt-ctly meftsured also: hut lliia aeenis to
bo an-Tv dillicull than the diterminatioa of the enrbooic ucid.
' la the plan originally adopted by Kegnnult and Keivel. and followed by some
Mbrr oHwrrrm, (he aniiniil cximiinenieil ou !■ allowed to lireatbe n Unii(r<l and
mesaurtd hi mod j< here. Tlie carbuuic acid, as faat as It In luraied, in lived ami
(tnoved by ■ i-lt'iD)C Hilution of caiulie potash, and the normal percentage of
uygeu in the a(Eii(niphi're i* mainlninrd by a suptilyof thtvssH from ■ gas-liulder,
la ibi* way both Ihr myKcn cooauroed and the carbonic acttl ptoduied arc
Mrtrthi detrrmined. uhilc the coBtlnual snpply of freob oxyseii prevents any evil
rlfrtrla dne to brrathing a i-onflned portion of air. la order, Dowev«r. to avoid all
Cdlile rrron> sriaini: from a (oo re»tricle<l atnioaphore, a dlflVrmt iBMlind has
D adopird by [Vinokofcr and Voit. Their apparatii* coDsists cwtntiatly of a
lanv chamber, capaUe of holding a lusu conifurlably. By mean* i>f a ntcam-
«ngine a cnrrt'tit ol iiure air. mtaiueed by a gwometer. Is drawn tbrtiu^^b lh«
rlinmlrrr, Meaturtd puitionn »l tin- outgorng air are from time lu time withdrawn
Mnd anal^ced ; and from (he ilntu ofTurded by ibeoe Hnalysct. Ilir *mouiitji of cur-
1>nni<' Mriil (iindoibrr jtimlBiid of Maicrgiven olT by lh«occupaiit of the chamber
^uriiiv a given tim<- me Jricfniincd. Tbe oxygen coBnimed » not determined
tiirefiTy ; liBt if ibe total amoaat* of i-aihonic acid and of water given out hy the
Innp* and akin are ascerlsined. and tbe amount of urine and lecn known, the
c|uantity of oxvf ni comumed may be arrived at hy a uoiple calculation. For
«vl(trnily tbe iiilfrreiK'c between the terminal weight, plus all Ihc egnta and th«
\
Iiiitlal wcigiil plus all th« itifwU, vu be nuttiin^ oIm ibait tho wcigbt
DKXK^ii abMrbM (Jiirla^ the pmod, Thtti mclbod in turn, hoiroTrr, iit alto
III (>bjoctii>iM, aincT mmuui <>rri>rii in th<i HnalyM* of lb* mokII tamplw nf air
I'liiplitjod for lh« tbc ilittcrminnlioni nttaio conaideraUe dlnwadooi frheu tbett
arvi mul(ipti«<I m w to give ihv rbani^ in tbc wbole nuM «r air puaeil throagb
tlie aii^Mratiu. Il ae<eiiia, luureover. u(idMlnb1« to lrav« the i|unntttx UMd of n
ini|K)rUnC au eleoient hs osygen to be delermined by iii<liract caluilMiom.
J
Let lu iinsKitie, then, an experiioent of this IcjikI to faav« been oonpl
earned out; tnal the auimul'i' iuitial and terrainal «-eif(hta have baeu
raiely dHerinineil ; tb« ouuputttioD of tJie fuodsatufactoriljknowu tucioaairf
of ill mucb )ir»tei<l, fill, carbobydratM. nalla, aod water, itnd to ctiutaiu »■
mui-li uitm^-ti nud ciirltoa ; tlie woi^bl nf th« fecM and the DUn>i;*?u chef
contain Biici'rtaiiKid ; tbc tiitrcifttii of (be urine dettrniiniHl ; the I'ltrlHiak
ari<l and vtat<-r givvn off by ibe wbi>l« body can-fully niensiirud, and the
Biuunnt of oxygen abeorlH^l ailuuUted — what inl«r{in-tution i:hu Imi pUowl
on tlw raulU?
I^ iM «up|>c«« tliflt tbo ftiiimal bits gniiMMl w id weight during lk« jMriod.
Of what d<fc« w ooiuisl? Is it fat or |>n)leid inntcritu which hiui be«a laid
on, or simply water which luis l>con rviaiued, or sonne of one and iorki of
other? Let lie further euppaw that ihc nitrogen of (he urine puam] du
the period is leas, say by z i>Taninic«, than tbc nitrogen in the fovd Ui
afUr duduction, of vourae, of Ihc niin>gea in the feces. This meaos that
* grnnimes of nitrogen have bean retained in the body : and wo may wilh
reason infer llmt they have been fv<taiiied in the form uf proieid material.
We ntay even go furllier, nnd aaj that tbey are reuiiied in tbe form of Aeeli.
i.e., of muacle. lu thin inference we are ^n|[soiaew bat beyond our tether,
for (be nitnigi^n ruiKbt be ntored u|> aa some pruteid constituent of live hepatic
oella or of wmie mlivr I'tmue : iii<ie«i), it inighl be fur the while retained t
the f'.iTm of wmin Hitmgvuoui cryxlidline UiHly. Itut thti last event
unlik<-ly ; mid if nc \n« tint wnnl " tli«li " to lueau nilru^eu (proteid) holdli _
liviiijt nulmtance of any kin<l, wv may without feMTof any SKM error n-ckon
tbc 'lefii;iviicy of T gnunnin nitmgcii »> lli« ttturing up of a j^inmeK tInfL.
Thcrv utill reniain if — a gTsninitw of iui-reaM- to bi> luioiontii^I Atr. l.iH i
Bii|»pow that tbn total carbon of (he i-giiila ltiu> been f<itmd to Iw y griiitiim
lew llian that of tho ingnt4t; in other wiirds, that y griimmn uf varUin have
b««n Nlon-d up. Some carbon hiu bc«n ittortKl up in the flesh willi t
nitrogen just considerod; this wc must deduct from y. nod we shall tb
have '/ gramma of carbon to aecuuni for. Xow there arv only tnu prti
ctiKil forms in which csrbon can be stored up in tbc bofly — as glywtj^en or aa
fat. The former is, avea in most favorable easra, inconsiderable, and wc
therolore cannot err grenlly if ue con^der the retention of ^ grammes oirbou
as indicAlin^ the layin;: on nf b f;rammee fat. If a -^ b are found equal to
te, then tlie whole chan^^ '■> the economy is known ; if w — va + A) Wve* s
resiilue c, we infer that in addition to the layinji on of fldsh and fat »>iriw
water has been retained in the system. If w — (n -)-&> j^voi a uei;ativ«
•(uaulity, then water must ha\-i- l>een gi^'i^Q "^ at the mtne lime that Unh
mid fat were laid on. In a similar way thv nuluri' of a Ium of weight can be
BBOFrlaiiied. wlieiher of flOHh, or fat, or of walC'r. and to what extent of viicb.
Tli« careful comparison, tbe debtor nud crmlilur aocouul of inciinc and out-
p«it, «nahlM tis. with tlie caii(i<>n« rcmlenxl ni,'c<«utiv by tbc i»aiim|itioD« just
iiow mentioned, to infer the natiirv and extent of ibv Wlily chaugt«. The
nviilu thus gained ought, of cuurfc if an ois-ount is kcgit uf tbc wnicr takeo
ill and given Oiiit, to agne with the amount of oxygen conMimed, and nlav to
tally with the conclusions arrived at conceniing the retention or tbe nvttm
of wilier.
er.
'-' I
THIt STATISTICS Of XtTBITIOS.
083
I
\
Havinj; thus Mudi^^l the mMhud. and ^«eii its woikneu as well na its
etrenKtli, we amy iHiefty review Um rewilu wliiuli liave been obcoiued by its
iti«aii8.
^ OSS. NilrvyeiMiiA aiHiAolitm. Wli«n a meal nf Icjui cncat. as free as poe-
»tbl« {mm tut, la ^iv<!ii to ft dog wliicli had [irevioiisly l>M-n deprived of food
fvT WPW time, mill wlmow bwly, lliitrvfore. in jcmitly deficient in flesli. it
inif^bt b* exptctcl tliiii l)i« laru:vr part <>r the food wDiilii be at once nutted
tip to aiipplr pri'R'ing iti<:liri<-ncin>, and that only tbr Mouilior pari would be
ioiRicdiiitoly n»rk(->i od'xt urea corn-sijiondiiig Ui the pitrogt'nmii ni<;laboli»tn
going Ml io tb» body tU the limo, JDcriMHcd MitnKwhat bv llw iali<>r tbniwn
on tM tCMomy by th« v«ry proMDce nf the food. 'fhw. however, io not the
CMM ■• lir M the nilrogpn of (be nionl in cniicomH ; the liif)^ pirtion
pnam off » urcti at once, aod only a cnmparalively small cgnantiir in
retninecl. If lh« dirt be continued, iind we are enpjKning the mcal« given
t'> 1h' large ones, tlie propurtjon of the nitrogen which is given off in the form
<if nre« goes on increasing; until at last a condition is wtablisbed in which
the nilmgen of the egesta exactly equals tliat ofthe ingesla. This conditioa,
whieh is egioken of as " nitrogeDous equilibrium," is altained in dogs with an
exdiwiveiy meal di«lonly when larg« qiiantitira of food are given, and it is
not flBsily maintained for aay lenKlh of time. The exact quantity of nicKt
reqiitml t>i attain nitro^nous e<|uilibriuin varies with ibe previous condition
of t1>e <to|r : e>:|uiiibrium is fr«(|ueotly atlaioed when loOU or ICtOi) |;ramin«s
of meat are given daily.
Tims ihe ni'xtt etriliing eJleet of a purely nitmgeooufl diet is largely to
iiierease the nitrngemHio nietabiiliim of the bjdy ; and we shall see Inter on
that it iurrenM-A iIk- nietabolium not only of the nitriigejiouM hut also of the
(itbrr (■iiiisiiiufniB of the t)ody.
The ntalilintiment of nitmgeuous e<|nitibniiin doua uiA luean that a body-
<-i|uilibiiom ia ntahli^lieil, that thr iKnlyWdlght neither iucrfiinen ni>r diniin-
iiilies. <)u the contrurv, vihvn the nif-al m-eiiMtry to tialantH' ihi; nitrogen is
n large onu, Uie boily thougli it ia neicbi;r gaining nor ln>in2 nilro^.'cn may
niD in total weight; and tliv iiicrvnM: ii proviil hy emlcnltition fnmi the
FneoRiv and output, and in<h)cd by acttinl examination of the body, to be due
to tba laving on of fat. The amount »o emnil up may be lar greater tbaii
caa ponloly bo aco(niote<t for by any fat stili adhering to tbe meal given lU
fmid. Wb am tlicreforo <lriven to the eonclunion tliat the protoid food i»
Milit into a urM moiety and a fatty moiety, that the urea moiety is at one*
(liacbarsnl, aiul that such of the fatty moiety as is not made use of diractly
by IIm body is stored up as adipose tissue. And this disrujHioo of tbe pro-
t«id. H we bnvi' already (« -1)4')) suggested, explains at the same time why
the niwtdicl so largely and imnie^tiately incneate the urea of the egesla.
The charaderistiG eflect of proteid food to iacrease the metabolism of the
IhmIv is shdVD on other animals beudw Hiv do^, aud not only by means of
cKleuUtioiis of ii'hat b siippoKd to take place io the body, but nUo bv direct
nnalysi;. Thus the analysis of tbe body of a pig. which htui lieen le<l uo a
known diet, ounipared with Ihe aoalyeis with that ofanotbc-r pig of iheMtme
litter, killed at lli« time when the lirat «'a» put on the lixi^) diet, gave a» a
result that of ibe dry nitrogtiioUN muieriul of the footi only nboul 7 |>er rent.
was laid up as dry proteid lUHlerlul during the fattt^niug period, though the
amount of proteid foud nan lnw. Tbia owitfrnMS strongly with the amount
nf fat stored iiu during tbe name period (aee £ A07). nimilar obw-rvaliuna
cftn-ied out no slieep vlxiwod that in ihstw onimala Ihe storing up of nitroge*
lioUA material wa* cvvn lew. only about 4 |)er cent, of that gireti in (lie lood.
Every quantity of proteid material taken into th*' alimentary mnal thus
ap|>enni to atfM-t pmleld mctuholiom in two nays. On the oiie hand it excites
631
SITTRITIOX.
a rapid protcitl motnboliitm giving riM to ui imror<luite. Mid gmcnllT tiitf.
iiu-raiM! uf iirr.i : on the o(h«r hnnd, i( abrvem lo miuiiuiii tbe nfm'n^tbr
oornuil {irotFid mi'laboliam roniinually taking piice io lh« IxmIt. hii<I bdo«-
tniHites IO the normal reguUr dischar^^v of urea. It BWtus very latiitilb)
feiippofce that the proteid which playe llie first r>f thetv two parl» 'u. tvA ntiU
built up into the tissues, dote nol t>ecome ai-tnal liviof; uibeuiicv, bsi aniir-
^oe» ibe changes wliich givcH rise to urea outside the aniial living nbUMt
tD tbc blood or eliewhere ; and we bave seen that under the iuDuoic* </ ibt
pKucreaiic juite ^oiue of ilie prnieiil food may uodergo tbe ffreater piriiif
aucb a chani^e while it in an yut uiihin the lUitueDtary consl. Heno* Iw
sriscti the verv litiliiral dintinclion lo which we hare ulivndy Blhi'leil hctirna
" li»ue jviiteiiU " nr " morpholic protiiidn " which are artunllv liuilliipinlniW
llvinu Hubctuncft nf the tiwues anil give ri«c to urea thmujin the inciaMifs
of living *iib«tancH'. and " circulatiDg proti^da" or " floating pn>lrid*" wtidi
do nol nt any period of their rarc^^r itithiu tbo body becoiDc n» inl^grd |arl
of the living Rnb«1«n(v and by their indaboluni .-H-l ftrt wwrgv wit in ik
wny nl' vital mnnifcxtalioos. but in lh« form of hoal only. Wv nliall lUtna
consider what if the exact meaning which vre ought In mtuch to tbf wonb
" bccnniing part of the living aub»tftjice :" and hence tball d«f«r uotU tlcfl
uny iliKcUMion of the appropriatenen of theitc phrowe and of llie validity of
tbc <Hstiticrion which ihcy I'nrmulatc.
It was Diicc ihotight, at we shall presently 8W erroneously, that tbotadil-
sive piirpoM of proteid food was to supply ihe proteid tissues, and lliU ill
the energy tet free in the body in vital nianifefltaiions, such as movement Uid
the like ait distingiiiHlied from heat, had its origin in pmteid metahDliem. Ik
meiaboliMD of fats and carbohydrates ^ving rise lo beat only. Heficc«lM
it lirst became known that a certain proportion of protetd food appanulf
underwent a meiaboltstn giving rise to heat oulv, witlxrut becotniug faitof
tbo liMiiies. ihin •wcnieil to he a wn»iefiil exucntliture of preciou* aialtrill:
and the mclaltoliitni of thin portion of prolcia Ibod wai accordingly «palUA4'
iw a " luxiiaconHuiDption." ■ wasteful coniumptioii.
Iteforv leaving' Ibis suhjcot wc niny cull nttontioD tu * powibic UuNp
hoiwcrn the hietnry of prolcidx and that of fats and cnrhohydrnlffL TM
iiDtlnrm eomptxition of the blood, which the body n<0Ow ever olriving Id
iDuintnin, probably applies to its prolcidK as well as to ilx other conttilurnb.
We have seen that a sur]ilus of non-nilK^genous mnd-rialN in the bl'Md ii
withdrawn from the circulation and stonfl up as fat or glycogen, and it it
IHMi^ible iliut an excess of proieids might similarly be stored up in sonatkne
or ti.'riues. in the hepatic cells for inKtauco. though from the facts previously
nietitioned it is obvious thnt the power of storage is far le»i ibun in th* cm
of fal# an<l carhu hydrates. .Such a store of proicid matter would n-|ireeefii a
aort of circulating pruteiil, but uevenheleae fur its linal metflWIium inighl
liave lo form an integral part of some living tissue unit.
^ 924. Tfie ffrrl'oij'attiiiin'i nf firtrohydratr food. Unlike tltnee of protMd
food, the ellL-ct» of likts and earbobydratea cannot be studied aloue. \Vb«a
an Hiiimal i* fed ximply ou uon- Diirogeiiou» food, death won takes place:
the foot) rajiidly o-hm* to be digcM4>d, and Hlarvaiion eniueti. We can tbere-
foiv only study the niiiriti^'e eflbcts of thcw subsuinoa when tliey are token
(O0Bther with proteid niatirrial.
»h«n a small (pitiiitiiy of lat is taken, in company with a Bx«l niodenue
ijuantiiy of proteid mnlerial. tbc whole of the carlioii of the toivl roappean
iu the egesia. No fat is stored up: »onie even of the [ircvioui>ly exinting fat
of the body may be consumed. As the fut of the meal in iocreaMvt. a (loint
is soon mohed at which carbon is retained in the body n» faL 80 al-o with
starch or wgar; when the i|uanlity of this u tmaW. tberv is no retention of
TITK STATISTICS OF yi'TRITION.
e.'is
I
I
oa MMii Imwen-r hn it Lt iiicmtM^l bcyoni) a c«rtii!n limit, nr)»>i> i*
:> in tb« form «r fitl ut, ui b «niiillvr vxtvDt, na givoogcn. FniA mid
c'iirl><>by<)ruie)> lhercfi>re ilillcr markmllv from proleid fowl in thnt ihi-v iirc
ii<i( Ml (lUltm-lly pmT0(-4itirc of melnbofiani. Ihis is escee<lio|;)y well bIiowii
in the iTtiillK olvtiiinni on thtt pig previously inentioned. ll was found ihal
472 uiiiu of riit Kvre laid on fur ev^ry lOO uniu of fat taken aa sucli in
llin food ( which cmuistinj; of bnrley-nieal. vie., conlained a v«ry Hoall amount
of aiTiiiBl fnri. while tor every 100 units of the total drv non-iiilroj{«noiia food
indiidinj; fat, ttarcb, c«lluU«e. etc.. no I<^m than 21 unita were retained in
%ltv biMiy in the form of fat. Jio clearer pnxif than this <:oiilii be atTorded
that lal is formed in tbe body out of Homrtbiitj; whivh h not fat. In !; o08
we have already disounMi thu formatioH of fat out »f i-iirbnliydrulea.
At one u))k1iI imagine, tbe pra»en«e of lint or carbobydrutt'n iu iIkt fuotl is
fonnd to decreiue tbe ainouul of proteid material iiiK'CMitry to ■•Mtuhliah nilro-
ftenovs equilibrium. For indnnon, with a dii-t of HIXJ grmK. mc^ul and ISO
Srins. fat, lh« uilrou^n in the e){i-i>iit Iwcnnie equal Ut that in thu iii)i;isia in a
Ofc, in whiMC caite IKUO grinii. meat had to b« given to produce tlie «ame
retiult iti lli« nIkh'Iici^ of fat« or carbohydmtcfl.
>Ou ib« cithiT Imnd, it was fmiod tnat, with n fixed quantity nf fatty or
CKrlv>hydntli- f»<Hl. an increa^c^ of the acoompanyinK jtroteid led not to a akiT'
iog up of tlie surplus <^'arl>un contained in the extra ()URntity i>f pmleid, hut
to an incruue in the ooneuiii]>tion m' carbon. I'ruleld food increows nut only
pr<>tei<I Init also iton-nilrueeuous metabolism. This explains bow an exnw
of proteid food may, by tne inoreoie of ncneral nietaboliam, actually rwltioc
the fat of the tmdy.
Wc have al preeent do exact information coDcierninjf the nutritive diDer*
encea betireen fats and carbohydrates, beyond the fact thai in tbe final com-
bustion of the two, while carl>ohvdrale!i require Bullieient o.xygen lo combine
with tbeir carbon only, th«re btfiui; already MiAicieDt oxygen in the carbo-
hydrate it«elf to tbrni water with tb« hydragen pmwnt, fku require in addition
oxygen to combine with mxne of their hvdn>tfen. Hence in herbivora, living
largely on carbobydrati-H, a larger |Mirliou of tlu; oxv^feii cuUHUmed rvajijieare
io UM carlM>ni<- add of the egota than In carnivora, in which animals, living
chiedjr on pruteids and fat^i, more i<f it Uuivoa the body combined with hydro*
gen to Itinii water. Thin relation of tbe oxygen toihe carbonic acid i^ ol\en
exprtaaetl bn the quotient of the volume uf the carbonic add expired dividol
by tbe volume of (be oxygen eonBUtuc<l, the " respiratory quotient," =^\
which ia in herhivora about 0.9 and in caniivora about 0.6 or 0 7. When a
herbivorous nninml tlarvo', il teeds on its own fat, and niider liieso circiim-
Mances the rc^i pirn lory quoltenl falls into ibe carnivonma standard; and
indeed many ciicuntMancea afleci ibis respiratory quotient. The carbohv-
draUw are notably more digestible ibau tbe fats, but on the otiier hand the
fau contain more p«)teniial «ner{>y iu a Kiveii ncighi. Aa u> tbe nulritive
dtArenoe between Blarcb and HUgur, wt- Know nothing very delioite; it haa
becti ih'>u;;ht however that cane-nii|i:nr i.-i riither more Eattenin^' thnii starch.
S 535. Thf efirH* oj iiflalin a* food. Il in a matter of common exjierieuce
that gelatin will not Mipply (be place of proti-ids n^ a coiutitui-nl of lixxl.
Animal* M on gelatin tugelber wilb fat or cgtrbohyd rales die very wuub in
th« same way aa when they are fed on noii nitrogetious nmtcrial aloue.
KevertbeleH it w»uld ap|>ear, its might be expcclcd. that ilie prewcnee of
gelatin in food ii> not Hiibunl cHi?cl. This nittxigciiuus equilibrium is cMab-
Kahed at a lower U'vcl of real proleid food when gelatin is added. I» a dog,
Moreover, fni ou a diet of gelatin and fat, ihe exccas of nitr<>g«n in tlie excreta
over tfaat in tbe iug«ei« is l«w than when iIk same dog is led on a diM of&it
I
636
WITTBITIOX.
■KiiK' : thiit ix to Mv. thi- gelatin liii> kIk-Iivi-mI fr»m meuttolnn suiiiv pnMtU
CDnHUCm^ulv ••rtliL- IhmIv : aii't thv- cPDitimiHkiu •>r fat »o«iiu also U> IwlearMl
by tlw |>wwiioe. >'f gi'btin. These fiictt iK-TOtin; iotvllifriUI*- if we suppnee ital
]fi.>laliii is rnptdly ejilit up inU> ■ ur«a «n<l s tiit raok-ly i» tbe Mime •ny ibn
wu luivc M^i a cof tain quaiiliiy of prolcid material to W. It b this ilinci
<UMrui7tivc m«ubolist» of protettl matter which gvUlin on take up ; it bcm*
however unable to imitate ibe otber functioD of proteid ntalt<^r, nnrl to takt
piiri in the formation of liviuKSubatance; or id the phraMM)l(>gy of a nncBl-
iiig pniah'Tapb i $ -V^^), it eaii lake tlw place of ciroutaling but not »t tiMt
pruleid. What is the t-aiiK- of tJiia ditTereiice «e fiinnot at pr<wi«iit »*r. •
S 886, J'eptone -isjood. SitK-e prol«i(la are at leant lar(^ly, tu wc have mm
(!i 310), convened into h»<1 ntiHorbed a* p«^|>li>m-, awl iiDce. aa «r« have aW
aeea, the peptone Bj>|R-jin><ltiriii^' the very Mit of nl»orption to b« i«coiiV«tt4
into noiue oihrr form of nroli-iii mntt<T, |Kmibly wMTim-albwiuin. it nii^
M'L-m natural to aiijipim:! thai pi-jiloiie given n* fmMi woubl, u-' far a« iwta-
bolUm n cunrpmiil, play tbi.' onmc juirt nit other prol«idN. NevertbelM.
WHue ob»erv<T# have timii)tnin«<() nitb regani to both peptone* luiil the alM
albiimoM* that, like gcliitin, tbese bodies " can lake tbe place of circulalini
but not of tisMie proietd." On the whole, however, tbe evidenoo guv lA
«b«u- tbut Atiiroaln can ** lav on flesji " when the protetd in their (nod ettoiiA
fjitin'ly of peptone or albumose. A difficnlir ap]>ertaining to digmioa
prevents any large snbAtitulion of ]>eptone for ordinary proteiids, stnei M
might be expected diarrhcea ia apt to l>e set up.
gAS7. The rfcel* n( niiU a* Jood. All food oonuinB. besides the Bubstanc*
|)oaseB»iof; potviuial energy, which we have Just studied, certain raUoe imI-
tera, orKanio aiul inorganic, haviiix in themselvei little or no aiicb |»>leoliil
eowrgy, but yet either abtolutely oeonwiry or ht^ily beneficial to the boiU.
ThcH uiuai luive imporlaut fuuctionx in directing the meubollam of Um
liody ; the xlrikiiitt dintritnition of them in thv tewnw, the prepondenuxwef
sodium atid i-hli>ri<lni in blood-iN-rum ami of poULMitim and phoapbalfls In tbt
red CDri>il*cl«a, lor ini<tani-ii>, m\t*i have »omc meaning: hut at prca«fll a*
•re in ttie dark oin-eriiin^ it. The clrmont phosphoniR ik«iii< n<> ti'M impv-
lant, from a biulogiml point of view, than carbon or nitro^n ; it i« a* aW
lutcly (wenttal for tlie growth of a lowly being like I'enicilliiiin as for man
hini»elf Wo find it probably plnying an iniportani part « tbn conspioi«<U
i-onstittient of lecithin and other complex fats belonging 10 the nortwui
BVrtem ; we find it prumineul in the iwculiar body nuclein ; we Hud it pecu-
liarly associated with tiie pruteida. but we cannot explain its rd/r. \ht
element sulphur, again, in only ancond to phoapborus, and we 6ml it a* a odd-
atttueiit of nearly all proteio*; but we cannot foretell the exact ohaii|pi
which would take plaou in tJi« economy if all the Bulpbur of the fond wV
withdrawn. In the kreatin of the epidermis and iu appendagea, hain, <M.
it \» pruliably undergoing vxcrciion, though its proaeum id thin iHKly nuf
have to itu wit)) the jwciiliar }>hyi>ind charucioni of romemw epitbeliuni.
We know that the various saline matters are Mamtial tit health, ihM
when they arc nut prir««nt in proper pro|iortiont nutrition U a6i>ct>^l. \iop
fed on ftiud. fremi as mui-li a« possible from alt saline matlcre. but othorww
abundant, with a proper proportion of the food-stulltf, soon exhibit symplwD'
showing that the metal>oliam of their tissues, (ia]>ecially of th«ir nutnl
nervous system, is going wrcmg ; ihe>- sufli^r from weakneaa, aoon amoualipt
to panilysis, and are often oarried off by conviiUiona. And innrv or iM
similar derangeroenta of nutrition follow the aboence or a delicieiirv of indi-
vidual aalta. During atarvation thcste various aalls continue to lie diaclumc*'
fVom the body ; in aoiue way or oilier they are carrie<l along in the metabolic
ttreaiu, and ih^ir pretence b in .vume way eaaential to the varioit* mvtaholic
THB ENSBGY OF THE BOPY
687
hmor, ili«y iMiiH) lo be mlwav* iiit^ent iii iliiily Tiiol. In whnl
■ajr tl b tfail tlu'v tliuN ilirwt iDdnbcilisti) ur do ii»t knnw ; wo nrc a«'ur<!
tut tta* pmporlHti anil iMUioui of vnrioux jiroLriil »iib«Uincc» uro clotvljr
llpMHlntl nn Ihi; [>n>wrn«e of Mrlain mil*, but buyiiiid llii« ws know verj
Uubu Tbf inorganic Mlt# nro lhi>w tho imiritivc value ul* which hiu bcMi
|U*djr *(U(lii-<t by expvHim-tit. Iml wr liavv n.iwu« «> bolievo thac the or);>ni«
<ir vxrafnivo. which iirv prcwi-Dt id grantor or Icm <iuantity in all food
tt both vrnUtble umI atiiiaul origin, are no \em cewntiat to the proper meta-
Uic iHiiTilic* of ibe bwlr. The iindoulitod connection of scurvy with lltft
i»rk iif fmb v«gFt*blt> fiwcl. other conditions helping, rosy rierha|>« turn in
fan uu tUts, for lh« erideiiL-c that the <li»e4iHe is due lo l)i« deflcieney of
pstasb alone u not eonchisive.
tjutly, water ha« an HTed on meinbolisni, us shoirn. antunjt other things,
Vf the fni-t that when the water of a diel is iodvased. the urea is inorMMd
to aa exienl Iteyond that which can be eiptniucil by the incrcMe of fluid
bcRMMUK the facilities of mere excrelion.
The Enkroy of nie Body.
The Tucome »/ Eneryy.
S ASS. Bnuidly ^prAking. the nnimnl tmly it n machine for couTertitiE
'Mtiai into actual cnorg}'. Thi- putcntlnt eiwri^' i« ciippticd by food; ibu
■feCtabolisin of the body convert* into (he actual energy of heAt and
Bical labor. Wc have in llie prevnl •oclioii t>r study whal h kmiwn
laws of this convomoii, nnil of the <li»trihutioQ uf the eni^r^y *el
Ccttkatlog all tubaidiary nod untiu|M>TUut Mure«a of «nerev, we may say
t UN ineofue of animal enerfty cunsiftis in the oxidiiiioii of food into ita
'Iprodiivta — vtx.. the nxidulion of pni(ei<i.i. fal*. anil carbohydrates into
irbonic aeid. ami water. A principle laid down bv the chemist teaches
a potrntial enrr^y of any bgdy, cuUMdcrcd in rvlntiun to any cbeniicoJ
luee which it may nndcrgo, {a the mtat when tbo final result is the same,
,bpr that mult be gained at one leap or by ■ scries of sle|n: that, for
nor, the energy set (m: by the oxidation at 1 grm. of fat into carbonic
aiKl wat«r is the Mme. whatever the changes forward or backward with
fat undergo** lH-f<ife it finally rencbcM the stage of carbonic aciil and
r; and similarly, that the energy Available for the body in I «rtu. of
proleid is the energy gjv«D out by the complete combustion of tliai 1
Im the cnerey given out by the coRiplete combustion uf that i|iiimtity
~l to which the 1 grm. of iiroteid gives rise in the body. Taking this
irnir guide, we can reiulily calculate toe amount of potential euvrgr C0JI>
■ml in an average Iweiily-I'our boura' diet, and thus obtain the avera^^
iiicunM of energy. For the potential enetvy of ninat of the aubMoacn
as food has been determined by direct enbmntetric obaervatiMw; umI
wveral determinations, tboueh il>ey vary soraewhat. ngnc sufficiently
1y 111 serve en data for the calcitlaiions in i^ueatlon.
le total i-oinbuitionof thefidlowinKHibstanci-a ha.H given for oiie gramme
•ubdauce the fullnwiug results cxpreMvd in oalorica— tluit is, id
uuilf <if heat :
, fVee fVom fat. -^103 aixl 5.^24. Fibrin. A6I1. E)!g-albiimin, o-'iTS.
lis, taking rnun<l numbers, we nuiy kav that 1 grm. ol proteid material
ntaius /fiiiiil or 5,'>iiO cnlorieii of potviilial energy, aeoonling as we use the
■pr Of higher dctenninatiODH.
633
SCTBlTIOy.
Fai of l>eefnr mutton, 90UU.»36o,!)42;l. UuU«r. 721(7 nr 9192. Apii,
ID rvuud numben, we may auy tfa«i 1 grm. of fat conlaiiis Ktiotii 90<»ntln-
ries.
Arrowroot (nenrljr pure March), 3912. 8larc)i, 4123. ll^llulaw. 4Hii.
Dextrose :)G9'}. Uine Kugar, 38(i6. Here again, taking round nuRilm,*t
AM not be fiir wroTiv in MvidK ihat the pntAiitinl t:\Krfiy of 1 gna.tt{ rtAit-
hy<lntl« mnlvrinl i.-t aliinit -tOdO calorics.
The comhuHliim nf 1 grni. of iimt *ot« free an amount of riivrgr vlilcbhw
been (ii^tcrtiiiii'd l)v dih- oli:irrvrr n» 'JJOB, by iinotlMT as i4(i-'> calurieiL Vr
havceccn (,ji-W>*) iliiil I gnu. of i>rot«itl giv»!> riwJ in tl» body to iitnamna.
tleoci', to obtitiii Ui« eiiergy ol 1 grni. prolci't mulcrial arailable (iir ik
ecoDcimy, n*c must d«luct from it« |mtciitinl energy oiie-third ibepoWniiil
coergy of I griu. uren — that i§. in rotin<l numbcra, TUU or tM)0 calonot. Th
will ipve UB oOOO ^ 700. or .^501) — >H>I>. ibat is, 4:tOU or 4700 caloriea. amri-
joj{ ■• we uko the loirer or hif:lier d iilti ; or ue may take n a nMao 4U0
oiloriea. The data, then, so far, are &« folloirs:
I l-raiiiDie proleid -I-VW calotw*.
1 grunimf tat 9rtXi "
1 grnnimc cnrbohydrnio 4U(fO
The average diet of aii average nmn— ibat is, the at:%ni^e auouiii ofeKt
fixxl-atufT reapectivcly laken daily — n*^ be det«rniiiKd ex[>eriincDlally «
»laii:^ti<altv. TbuB, a mau may aMermine by a seriM of triala tlie did m
wliit-h. while ndtber louag nor gainin;: weight and Riaiiitaining " nitrugtowi
emiilibrium " (5 523), be eojoye (rood health. Or ati nviTage may be tUwk
of a iarge number of dieia u»ed by varioos |)enple. Wenhall have twintlliint
to Ay o? this I<«r Btalisiical method when we come to ji|>eak of die<. For
the |tK«eDi purpose we may use one arrived at experimenlally, nlikbut
will »|>rAk 111 a.1 RaukL-'s iliei, taatx it wan deterqiincd bv a |ibysioli)f;ifl U
that onmi! frnm uliwrvatiomi on biniMlf. It w«« compdn^i] of lOHllgnia.
proieid, 100 gnrnt. iat, '240 grnift. carboliydrate. Such a diet would gire
inogrammniproteid (V'HMJj Idtt.UOO calories
lOOgraniina fat (9i>'>r)) 900,001) ■*
310 srammoi carbohydrate (4000) .... 900.000
2.310,000
If we translale the unii« of lieat into units of w-ork. tlie 2/110,000 graRiine'
dr^nve. or 'Z^ilO kilugraoinio-degree calories will give ua about IMO.OOO.Of, in
rouixl iiiimtren. Aomcwhere about one milli<fii kilogramme-metmi.
Wf may. in pn»iTig, ctkll attention to tlit? faoi that Ibi? pmieide «i|iply a
rclntivcly ixnall part of lh« total eoorgy, and ibut ibe nbarc runlrihuled'by
the lar^ji- niaM of carbohydratet ia Dot much grt-altj- than that belonging in
the much unuiller ijuantiiy of fat. In the averaj^ii dirt obtained br llii; au-
tislical method, in which the data are largely dniwn frnm public itiMttuticnu,
the (cbcapcr) carbohydraliw aru atill furtli«r iacreavvd at ibc eKpeuae of tltv
(dearer n«t«. a change which may tend to reduce tonwirbat tbo total ener^;
but thi« docs not materially afTveC the broad mtult jnM giren.
Tlie ErpenilitHre,
S 529. There are two ways only in wliioh energy « •W f'"*'* from the body;
me«linnical labor and heat. The body losw energy in produting muscular
work, u* in looomoiiou and in other kinds of labor, tn the uoveiDcnta oflbe
TDS BXEROV OF THE HOW.
688
I
I
iu nxpiruliixt tttid siwccli, ud<1. though to ii )inrtllv ns.>ogaixKbl« exttdit,
h<' niovcnM-iiis of tli« uir i>r contigiioiiK boiliut tiy tliv piilMlioiisor Oat
vasnilnr HTRlcm. Thv body loan cnergr in the Ibrm of hmt l>y i:«iidiictii)ii
And radiation, l>y r»piration tind perepirnlion, iind by th« warming of lh«
urine and feces. All ihc intcrnKl work of (lit' iiody, all the niechiinkiil labor
of ilie internal musi'ular niecliamsnis uillt their accoiupanyiDg friction, all
the luolM^uUr labor of the nervous and other tiwura, is coiiverl>-'d intu heat
before it leavee Ibe body. The inoet intense mental nciiou. unaocoiupaDied
by any niuwular man ifntal ions, the nii»t eDeT;geltc action <>( the heart or of
the howeU, with ibe ali^-lit esoeplioDB nieutioued above, the husit^t activity
of the jWcretinK i>r uietabdlic itMues, all tbe^^ eml Biui|>ly tu aiigmeutiug tlie
cxpouditure in the form of h«at.
A normal daily rX|Mmdiiure in iho way i>f mec-bniiival labor van be eauily
<lfitermiiKvI by obm-rvaliini. Wht-lhrr itiv work liiki- 0:1 lli<- form of nalk-
iug, or i>f driving' a niarhim', i>r nf any kind of iiiuxciilnr toil, a ^>iKt day'«
work may 1m; |>ul donn iil abotiL l^O.IKKt kili)j;nimtiii--m»trcK.
The normal dnily rxjicndiliire in th<! way of bi;iit aiiinoi W «> readily
deterraiiM-d. T>itvct (-alKrinH'iric iib»ervulioi)* on ihi^ whole bodv an- nttrudvd
with B» many ditlivtillMs, except in tht- »uc of nmnll nnimaU, (bat ilivir valui'
w UMM^rtain : and oWrvaliun> miuU- by plannu a part <mly of tbi; body, an
onn or leg for example, in ihti ealorimcler. and from ihc data thiin gained
calculating the heat produee<! by tJie wh<>li> body, arc siihjccl to nmny «oiir««e
of error.
The caWimeicrii uiually employed in chemical operation*, in mrJuurinK, for
initlance, the b«Hl ^nven out in chemical vbangM, are uiisuiiable for rt peri men In
on liviiift NU>m>l*, l^ucli are the mercury calonmeler. in which ihe iliemical
aetlon to be ■(tidied t» nmde In tnVe plncr in tlic inirlut of a miuw of mercury, from
tite cotiKcnueot expuniiiin nf which tlimujfh thi^ lirAt taken up the iimount of heat
givaii out ■■ calculat(<d, or the ice otlorimeter in which in a HJiuiluy way the heal
KiTen out la calculaled from tlie amount of ioe melted. TIh- latter hasbren used
7<ir phyTiio><>^ic)il purpose*, but an animal eurroiniiled by ice is under micfa abiior-
■nal conilitionii llial (lie m>ntlii are of little value- The method* usnully adopted
liy phyi><>1oiitKliL ate an rollnwH :
la one method, the water calorimeter, the animal h placed In a metal chamber
iKirr»uaded by a jacket lilted with water. The heat given out by the animal
warms the water in the jacket, and the amount given out i* calcniiiied upon ibe
Incraasa of tbe temperature of the water, Uy supplying (be anlDial witli air
tkrougb a Iork apiral tabe paaaioc throagh tlie wiiter-jiicket, the heat given out
in tti« expltea air is urereoied Inua being lost.
This method may be employed in a Klmplei foiin, when the beat given out bv
apart of the lH>dy. the arm or leg for intUtnce, i> all that hiu lo bo detcrinincd.
Tbe pert in Ibca mrrety placed in n bath «( water, from lht^ cbntira of tempera-
lure of which the Hmuuni fiven out ■■ cakulmcd. And thii modi flea tluu of the
malliod may with dae precaution* be employed for the whole body.
In tto«enlluir>' calorimeiier the cbamhcr in which the body or iiart of the body
ia placed in *urroundeil by, not n wnler-jackel. Iiut an air-jacket, which thua Mrroa
aa aJi air calorimeter, Tbe iDnirumeol consiitH ewcDtially of three couoenlric
cop|ior ryliiidcr!! ; the inner one contains llic animal (or other ouurce of beat) ;
the other one >H-rv«M mricly a* a casing to protect those inside from changes of
leniperntiire due to vurreniJt of air nnd the like ; and the middle one enclonn an
air space bvlweeo Itaelf and the inner one. 'Iliere are niiccial arrangements for
clnsitig the cylinder* after Ihe inir-.^luctlon of the animal, and for supplying (be
animal trilh air for breathing purpoHea. With the ali-jscket. or space between
the iiinef or middle cylindcnt, are connected a manometer and a tbennoDKittr,
Wlien an animal lor other source of hi-at) ia placed in the inner cylinder, the
tempcratunr and the pmuiirc of the air in tlie air-jacket are increased ; aud from
the iimounb of Increase measured by the thermometer and tbe maanmeter the
amuaiit of beat given ont from the animal is calcuUted.
The calurtBHem of D'Arwrnval and Kuhner are constructed on very nimilar
principhw.
64U
XUTRITION',
Vurtuue attempb bavo been made to ascerlaiu ihe aninuot of bni pm
I'Ut by tlic body in an indirect mnnner. na for iostAnoe by cnliniUliaf iV
h«iU ^ven out by the oxidation of the food. Aa iniitn'ortlty luaajvi^
|)tiui of einiply BubtraclioR tlie norniEl duily mecliaoioil exprnidilun ki»
ilie normnl daily income. Thus, 150,(>00 kiloKranimc-nieirM MibtfiQnl
from one million kilogramnie-uieir«« i;pvest(aO.<IOI) kilogramruc-nMnBHll*
daily espcHditur« in ihe form of beat ; L e., Iwtwn-ii [>iii.-finb and <->ii< '
of tlie tola! income iM;x)>endeilaa mecbanieal labor, tbe rcmainin): rii:r :
(ir llve-aistbs leaving iIil- liody in the form of lieau The results ^t«ii bj
direct cal«riinelric oWrvnlions and by otlter nilculatioos gire sonrnhK
higher fi^ret llian t\>inM> ; aiid iiKlecd tke«(< may probably he taken aj imilcr
ruber than over tUa true iimounl. In any CMO they are to be rcfaidal «
fumixhiiig nothing mure ihnii a roufth ftv«niee. the esiict amount *umt
according (■> tbt* inte. the weight, and the ooii<ntioD of the individual, uVrS
as ncennliDK to varintiims in cin-iimotunouf^
$630. T/tf inmjy of nn-chnnimt wrh We bn»e already in irealin^ </
luUBele and clM-wh<<rv [Mirtly dtMUWcd this subject, btit may here ay UiMot
that baa to be #aid.
The older writers, even aAcr tt hnd been proved that the animal bnlym
OODBlmotive, as far as the formation of (iit wns coneerued. still belli lo tbt
dlftineliun between tulrogvnous or plosiic and non-nit ro^uoua or ittyinWt
food. I'ut broadly, this view was that all the nitrogenoua food wuoi to hiil^
up tli« proleid tissnea, the muscular fleah and tbe like, and tliat lhaiiirv
gciioiiA ege^ta arcae solely from llio functional metabolisni of iIhk tlMMi.
while the noD -nitrogenous food was used with e<{ual exclutiveti«M Ibrrajaifr
lory or caluritic [lurposes. being either directly oxidiued in ibe bhwd ur.if
prcaoui ID excen, itured up a» fatty tissue. According to this vivw iht t<<>
cImnb of Income curreEjwnded exactly to the two formx nf expendilDt^
We have already UTV^•*i w^vvrnl objections agaiuHt ibis view. W« have«ai
thai in tbc blood itsi-tf vcrv litlli- nxidutina lake^ place ; that it is the Mbn
tissue, and not tbc ptis:<ive ^loixl-plnmiin, which ia the itcat of oxidation. \Y«
have further seen tkiil pn>icid fiKnl may undinibledly b«. in tbe above aMM|
respiratory and incidciilnlly give rise ii> the vuiring-up of fat. One divUsa
of the view is thcrt-by overtTirowti. We have now to inquire whether tht
other division holds good, whether niusetc and tbe other proleid tbau« ar«
fed exclusively on the protcid tnulerial of fuoil. Hnd whether miiMular
enerfcy cornea exclusively from tbe nictaboli»in of Ibe proteid coualitiientf of
mujiet'e. We have already seen CS 6^) that when the muscle itself a oxan-
ined. we find no proof of nitrogenous waste, bnl. on the other baud, dear
evidence of the production of non ntlrogvnous botlies, such as carl>»nir acid.
And when we oak the question, Docs uiusctilnr vxercise pro|j»rt)i>tMii«Jj
Incieue tbe urea given off by ibe body as a w hole T for this, act'urding to tbe
theory in ({Utation il certainly oui;ht to do, the evidence we can obtain,
though somewhat varying-, i^'ivee on the whole a <lende4lly negative answer.
In tbe majority of observations no marked L-bniige at all in the mnount
wns met with : indeed, in M>Rie cases there was a di^inei decmuo. followed
by an increoM* on thi.- following daya. Some obserreni. howt^ver. fuui>d a
Tcrv niurknl iiicrea»e. and lliia was eepecially Ihe ruae whi-n ibc subject
under oliwrvntion took a lari;e amount of food and uerformnl very severe
labor. 0:i tin- whole, the vailou^ re»ulta obtained bv difforeol obeerven
justify the cunrltiftiin that cxcn'iae by itself, even when severe, doee doI
necessarily increa*e thf amount uf urea excreted, but that conditions tur
obtain in which such an increife undeniably occurs. Wo may draw the
further cundunion that ex|ieriineuln i>f lbi» kind do not supply the riiffat
method fur detemiiniiig the point at ivsue. It must l>e rememb^«d thai it
E KNEROT OP THE B0D1
th* nntclM done which feel ibe iiifliinicdniio 1«I»ir; Uw dKulition
iadm) the wbol« boily are utli^cted by it. li' wo au|)poM«hu)[t part
' or even onlv some port uf the urm li> wmv rnim oth«r than miiiK-uIur mttti-
bolUni, froiD cluuivei in the he[>iilic ccIIh for iiistnncf, we ihoulil cxprct ihnt
thnv cbanceii. aii^ nilh them the amoiiiil uf uix-o <)i»chorgol, would bo
inl)u«Oc«l by labor, nupccinlly by severe labnr.
In no csw hu ■ direct relnti'iii Wlnti-ii lli« amount uf labor anil iini'xint
of urea been olvervotl. More thiin lhi». the follflwing <ixp<Ticncc lumln us
io an ah«uriltty. if we KupjioM- the whole energy of muscular work to iirisc
(nmt pnitei'l nie(a helium. Two obM-n'cn iwiformrd n ccnuin amount of
work (an aM-etit of ii mouiiiain) on a noii> nitrogenous diet, ami estimntml
the amoont of urea uossed during the period. AsMumin^ the urcn to repre-
•fnt tilt oiiclaiion ofao much protcid matter, which oxidation represcaited
in turn ao much energy set frve, they found that, whereas the actual work
dona amounted to riit.026 and 14S.6o6 kilogramme-kilometres for each
oWrrer renpeclirety, Ibe total energy availahlo from prot«id meiaholisni
■luring the perio>) wns iu the cose of the tirsc (tK.flO, and of the second 68.376
kilogram nc-kilometrce. Thnt is to say, the energy set free by the protrid
metabolism of the musclea engaged in the work was far less llmii the amount
neceMnry to accomplish the work actually done, to say nothing of its having
In provide as well for the movements uf respiration aa<l cin'ulation. Their
mivcular energy therefore must have had other sotiTcea than proteid meia-
boliFto.
Tb«toa the contrary the praduciiimof t-arbouioacid is at once and largely
incKMed by muscular exercise is beyond all doubt. One hour's hani labor
will ineroH RvefWld Ifae ijuantity of carbonic acid ^ven oiruitliiu Uiu hour.
Aud in an experiment direeud to tliis point ii wast found that u man in
twenty-four hours coiifturoed 9^A graiunini oxygen and produc«il ]2Ai
gnnmt* rarbonic acid when doing work. »» aguinai 708 graiutuea oxygen
cDiUUmeil and 01 1 grammi-M curbouic acid pn^liicetl when reinaiuiug at rtat,
Uia iiuamiiy of urea fccretcd being in the lirst case HI grammes, in the
•Mond 37 .'J! grammes.
It is erident that Ibe conclunDns arrived at by the statistical method
tntirely corroborate those gained hv an examination of muscle itself, vi^..
that during muscular contraction ikc explosive decompositJon which lakes
place bMis chiefly, if not eicluRively, on the nou-uilrogcnous oonstiiuenta of
the muscle, and ibat it b ibc non-niinogenoua products which alone escape
from the muscle and from the biidy, any nitrogenous products irhich result
being retaiueil within the muscle, or at least within the body. We muU
tbcnfore reject the secoml &» nell aa the Gret division of the views under
dbcnwlon : not only in the mt]»cle not f<d exclusively on proteid material,
but ftko iu energy does nut arise from an exclusively proieid metabolism.
I
Animat Heal.
I SSL The (oiirvfs nntl diglrilnUiou of heal. We hare iilremly (ten that
tht conception of the non-nilrucenous portions of food being solely calorifa-
dent or respiratory proves to oe unfounded when wo attempt to trwoo tlie
hiatory of tlie food on ils way through the body. The same Wew is still
more Mrikiiigly shown to be inade((uale when we study tbe manner in which
the bnit of Uie body is produced. We may, indeed, at once affirm that the
hut of the body is generated by the cbemioal ofaangee, which we may speak
of gmemlly as thoae of oxidaitoii, undergooe not by any pariicuutr snb-
atancM, but by the tissue* at large. Wherever nwiaholisnt is going on, or to
b* more exact whetcvej de^ruclive metabolism, kataboliun, is going on,
41
642
XfTRlTlOX,
lieat is b^iifc set free. In growth and in repiiir. in ihe depumiinn of an
timu-fiul. ill ibe iratiEliiriiuliuii of lifelese piil>iiluin iuto living' (i«^u<>, in tke
coiiMmclive tiiecab'itisiii, the itnalwlism of tlie liodj'. and in the sRuiller trs-
ilietic pniceffiefl »f which we spoke in denliuj; with urea O -1D0^ bmt ■
undoubteiily lu II certain extent beinx »1»urbed and rendered Uteui: ibe
I'litrfcy or llie i'<jn9iruclioii umy be, in |>ari at Icilm, supplied by (he lieot
pn3H.-iil. But all this, anil nutre than lliis, vit.. the heat {>r(«eiit in a jioten-
tiiil form in thi; niibnlanL-Mi thetoHelvai so built up into the iLvue. is lutt to iJw
liacue during iu deAlruvtive metaboliizn ; au thai the whole inetabolbin, Uie
nhol« cycle vf cliaugca from tlie UIUcm piibultiin ibrou^di thi? living lime
bocj; to tlif lifelew jmidiicta of vital arlimi. \» eniinetilly n WMirc-v of Urst.
Ofitll the tiMuvit i>f ihv IxmIj' th'' niuiti-list, mil only fniin tlieir bulk, I'uria-
iii); III iht'y du to Inrgo II |M)rlic)ii iif the whol« frame, but alit» fmni ihc
chnnii'icni of their ineiiilxdixni, niu.nt be roganU-d iii> the chief lunircvs of bnt.
In tfx-nling (^ '!$) of tlio Ihi-rninl chnngiv in miivclc wc huvr )n?*mi ih.ii ia
the tulnl rncrgy CX|H-n'lcd in ii miiwiilur cimtmetion, ibi- ratio of thnt whiefa
ap|)C«r« ii» hcHi (o tlmt nhich npi>cnr> n» rxlemni work '» variable. If
wc take a pro]>onion which !■ sonicwhat higher than the nx-an of tlia
ratigL> there given (unc-tidh to onf-lwcnty-lifth), and nunimc that tlw
energy inrolved in the work iloiio in a muMuLar controctiiHi is almat
one-tenth of the total energy expended, the mt going out lu heat, tfaeti.
Upon the cnleiilation that iW tulal cstenial work of the body is about
oue-fiflh of the total energy twt free in the body, it is clear ihat tlw
heat given out by the muscles, even if we consider only the liesi fEiveo
out when they are contmcting. niusl form n very large part of the toUl
beat given out by the body. And even if. as recent researcbee indicate, ibe
musculiir inaebinc works more econoniiciilly than we have hitherto aunpuaed,
the aniouiil of heal ^-iven out by the sketeial n)UM-le« must stjil remain wry
Uice. Moreover, to die skeletal ruusele we mum add the heart nbieb. nevar
re£ng, du^ in the ivrenty-four houn a» we ba\-e mi-n, ^ 1118, nu incoiuId«r-
•bleamotint of work, and must ^ive riie to no inconsiderable nmouut of heat.
But the xkelotal musoltn, though frtHjuently, are not conliuunlly contracting :
tlicy have period*, at timea long jieriodii, of tchI; and duriu); thesv portoda
of rvM, luelabolism, of n auhdiied kind it is true, bitt Hill a nii.-tiibolrsia
involving an cx|MMi<lituri.' of energy, it going on. This iiuieaccnt metalwlism
must iilso give nse to a certain amount of lieat : and if w« add lhi» amount,
vfaicb in UK prcsetit i>iale of our knon-1edg« we cannot exactly gauge, to
that given out iluring the movcnienli' of ibc body, it is very clear, even in
tbe absence nf exact data, thnt (be nietabolistn of ibe mtiacUw niiist siipplT
a very large proportion of the total heat of the bwly. Tlrey are [wr exc«i-
tenoo lb« Uicrmogeuic tiasuc«.
Next to the musclea in iinportance oome the various eecretiug glonda.
Id these the secreting elements, at the periods of secfettoo at all eveoia. are
in a alate of metabolic activity, which activity aa eleewhere maM give riae to
bent. In the case of the salivary gland of the dog tbe temperature of tbs
saliva seerole<l dunng stimulation of the chorda, noa been lound to tiv aa
much as 1' or 1.5° higher ihan that of the blood in the carotid artery at ilie
aiime time, and in all probability the invteligation of other aecrelin^ glands
would lead to -imtlar results. Of all ibeae varioua glunds. tbe lii-er deaervca
special attention ou account of its sice and large supply of blood, and
becauM it api>eaR> t<i be contiunutly at work. If Uiere lie any truth in the
vieiTB urged in ilie preceding cliaiiter touching the large uud varied metahuUc
work of tlie liver, we niurt eunclude tliai a vfirv large an>ount nf haal ia ant
&ee in this oi^gaii : and thnt holds good even if we make a large altuwano*
for llie variou* aynthvlic anabolic ]>rocea<ct which may lake place aiul ti^
vbirli bm nriuld b* nhMrb<^l nod made luieoi. We And. indeed, tbiit tbe
blood ill ibr hirjnlir vrin ii> the- niirtiMwt in ibe iKidv. Thiin in ihe dig n
ymmrmlnn' of A').'^" V.. hii> licvii iih*i-rvc<i id cbu beimtic vvin. nbilr that
of UN vroa oith iiilV-fior km :W.:U* tu Ilfl^S", nud rlint "f tbe riybt b.-nrt
S7.7* Thr furl Ihnl ibv binod of the he|iatic \-vin in niirin<-r thiiu ihnt of
rilbrr tbe (lortal vein or thr iMTta, nbown ibul the iiicniiniil (tmjK'raiiin' t«
Dtil (lur riinpl^r to thv livrr being fur reniovtNl from llie xirfHCci of (bt^ bodr.
11ie brain, too. niay be reg«ra«d m n lource of beat, siiiec lis tcinperamra
b bifH'T tlwn tbnl of the nrterial blutid witb wliich it is supplied : Iboiigh
front tbe amnlter <|iinntity of bloott piiMing tbnnigh it* vntsols, m well lU
fWim tbe obangtt i" it being lt» iDMeire. it cnniii>l. in this respect, cuiupare
with eitber tbe liver or tbe mtifcWaa ■ source of bcut to tbe body.
Th« blood itself ean&oi be negardod M&Houroo of niiy t^nsidernble amount
of h(«t, rinoe, sa we bave so TrequeDtly urged, tbe oxidiilions or other niels-
bolir rhan;^ mkiug plaee iu it are comp-iratit'ely sligbt. The h«al evolved
ibe iodlHerent limuee, such ns bone, cnriilage and coniiective tisfue. may
[unwd over a« inf^ijinificnDt i and we cannot even reganl the adi|tQse
ue as a seat uf ibe produttion of beat, since tbe fat of ine tut eellB h in
all pmbabitity noi oiidiied in lilu. but eimply carried away froin its place
Kfuon^ to tbe iiwue wbicb siaiid^ in need of it, and it ia in the lidaue thai
it undtrgoes th« metabolistu by which iia lateoi energy is set free. Suiuc
atngiint Afbeat is alao prtxluced bv the changtH which the food uoderfcucx Ln
Ibv alimmtar^' canal before il really enlen the body.
HcDoe, taking ii Mrrey of the whole bxily, we may oonclude that since
metaboiboi is going un to a greal<^ or lea« exteiil ererywbere, beat is erery-
wbtr* being generated ; hut thai, looked at from a qua nt its live jHtinl of view,
Um ma>cW and the trlandular organs must be regardeil an tbe maiu fouron
of the heat of the Imdy. tbe muwlest being, in all probability, the more impor-
but nf the two.
i ASS. Hut Im-hi, while 1>cin^ tbii» continually produeod, is on oniinually
bnng luM, by the skin, the bing«, tbe urine and the fcoea. The blood pass-
ing from ODB part of tbe body lo tbe otbrr. and carrying nnnnth fnim the
limiei where tienl Is Iwing rapt<llr generated, to the tifsucs or iirgan* "where
b<«t is being lost by radiation, conduction or evapiinition. tend; (u eruialixe
llie i(!iii)>eralur« of the various parts, and thus maiiitain)> n "eonHtanl. tiudily
temperature."
Wbeu tbe production of beat is not great lu compared witb Ibe luiti there
is no itieat accumutsiion of heal within tbe body, tlie temperature of which
caiise(]iieui1y i^ but slightly raised abuve that of surrounding objects. Thus
the temperature of the frog, for instance, is rarely more than 0.04' to 0.05'
above that of ilie atmosphere, though iu the hreeditig seasou tbe dilTereuce
mav anwunt to X". !^uch animaU. and tbi>y cuaipriae all claasM except binls
ao<) mammals, are sfMiken of as i:old-b|iii)d<M] ; ihey bave beeai aUu i-nlled
poikitulhermic, ttint in, of varied tempernlure. Ksce|rtioiis among them are
DDt nnrummtin. H<ime diAt. such tu the iiiiiny. are warmer, than the water
(n which tliey live, and in u «pcci» of pylbun (/*. bii'lUcilut) a diffcrotice of
as much as 12' ban li«-u ohwrvi^l. In a Iteebive iltc temiwmtnrv may rise
at til
and
tc temiwr
at tin»» as nmcb tf u< Ait'. In the so-called wano-blondcd animals, bird*
mamniaU. the ton and jiroiluction of brat are *o balanced that tbe leni-
peniun? of the body n-mnins constant at. in round numliers. $5^ or 40^,
wliatovrr Im- I be teinperattire oflbaair: hence ibiwv have l>een called bomoio-
Uteroic, of i-onMant leinperalurc. Tbe temperature of man ii about 37^;
is anme bink il is as hign as 44^ (Hirundo), and in tbe wolf it it Mid to be
aelowM3&/J4'.
This Unperaiure it with slii,dit vnnations maintained tbroughont life.
644
XfTBITIOX.
After deftih tbe generation of he&t rapiilly diiuiiiivhtv, anrl tb» body PpMililT
becomes cold ; but for twine *hurl lime iitirDedintcly following u[Min vvilcmir
deatb, n riae of leiuperuliire may l>o obxcrvol, due to the fact, that while th*
■uetftboliKm uf ibeiiwue U huII ^iiifc on, ibe Ium of beat i« *oiDewbat checlit<d
by the cvitsaiioti of tbe vircubilfon. Thi^ onjH^t nf prououiicnl ligor mortl*
cautcs u luurkeil accuviiiii of heat, mid when uccumiig utttr certain dimKs
ninv ^ivc riw to a very toiwiili-riiljle clcvntion of temperature.
tins iiiviui biiihlv [(.■miHTutii rt- uf niirm-bloodod animals », Jurine health,
miiintaiiK-d, witli iilit;hi vnrimionit of whicli we ahnlt I'rtfeiitly sjwaE. withio
a vc^- narrow imirgiu, n risr, or indcod n fall of much more than a d«grM
above or below the limil given above bein^' indicative of some failure in (he
organism, or of sotoe unutuitl intlueiice being lU work. It is evident, lliere-
fore, that the oiechtiDisms which coordioatc the lo6» with the prudnctioa of
heat must be exceedingly senailive. ll i^ obvious, moreover, lliat the mecb-
ftniaiDS mRV net when the bodily teniiRTitmre i« icndinf; to Hm. bv eitlier
checkioR tbe produotioD or by Buj^meniing the loM of beat, ooiiveraefy when
the bodily temperature is Ifoding to &U, tbey may act by elllier iuef«a*ing
the production or by diniiniflhing the losa of beat. As tbe re;(ulatian of tem-
perature by varialioDfi iii the loss of heal is better knoirn tliau rej^ulation by
variatiouB in prodnctiuu, it will be beat to coofiider tbein first.
g 533. Hegutalitiu % mrtalion* in liM: Heat ia loaa to the body, by the
n-armiii^c of tbe feces and uf the urine, by llie warmiogof the expired atr, by
the I- VII pi) rat iun of the water uf ret* pi ration, by euuduction aiid nuliati^tu fnxn
the ^kin, and by the evajMirntiun of tbe water of perspiration. It liax hoon
calnilali-il that the relative umouutii of the loss by thewMwral chnniiel* arc
a» f'lllow*: 1)1 warming the fecea and urine about 3, or, iicomliiig in oUmis,
6 per cent. By resjiiration about 20, or, iiecording lo other*, nlM>ul ooIt ft
per eenl,, le/iviiig TT, or allcnialively 85, per cent. For conduction and nulb-
tion sod evaporation by (hcekin.
The two cnief means of lose then, which are nt all susi'eptable of auy grtai
amount of variations, and which can be used to regulate the tein{)«.'raiun' of
the body, are the skin and the lungv.
The more air pnues in and out of the lungs in a given time, the grenUir
will be the Iob« in vifinuing the expired air. and in ovaporuiog the mtier of
retplration. In »iich aniiunk as the doj;, which i|o not perspire freely by tbe
skin, respiration is a most important means of re^oilaliog the teni|>emiiin.-;
and in the dog a very close couoeeiioii may be observed lielu-eeu the pnxluu-
tlou of heat and respiratory activity. Tbe ohangea wbieli give rise m this
loM tak<9 place before the inspired air reaches tlie pulmonary alveoli ; both
ibe narniing and the evapurattou are ellected in the nasal and pharyngeal,
and to some extent in the hruiicliial paa^tagiw. Some olMerven liave main-
tained that the li-fl :«ide of the heart is warmer than the riffht, and hence
have argued llial diemical chaiigen leailing to a t^onuderable derelopaiait of
heal tak(« place in th<- pulmoitary capiltiirir*. It would appear, tiovrever,
ihni the righl vcniriclc, owing to it* lying nearer to ibo Hvcr, the high t«D-
peiiiture of which has already been mi-nlioncil, i», in rejilily, rather hotter
thnn the W-H. And. indeed, we have no «! I in factory evidence of nuy lar^
amount of heat being produced by any niilriKinary nieliibolieni.
The great regulator, however, is iindoiibK'dlv the skin; and tbis has a
mora or leas double action. In the lir^ plnec, it rv-gulates the loea of heat
by ueaDB of the vasomotor mechanism. The more blood posses through
tM skio, the greater will be the loss of bent by conduelimi, radialiou, and
evajMiration, Hence any action of the vasomotor mechanism which, by
catuiiig flilution of the cutaneous vascular are8«, leads to a larger flow of
btoo<) through the skin, will lend to cool the body; a ud, conversely, any
*
*
I
iinouir Mtlnn whk'h. Iiy coiutrictliiK t)i<- oulnDcouit vtuMiulsr nmia. or by
iMlilhis the uplmiclinit- vascular ftrau, cnuM* n «niaUi>r flow ihmueli ihe tkiD.
ud KMrf^r nowof IiUmhI t)ir»iigbthe nbdominal viM'ern, irill t«na tohcMlh*
bodjr. In itir Accxiil pturt-, t>^i<lai this), the xpcrial nerve* of nerapiratton
will Hrt 'lirTCtl)- U reeulBlor* of tprnpt-riiliirp. inrrmning (be lo» of hcftt
wbrn tt)i-y momiote, ano liMCninK (he l««« nhcii thry caw to promoM, tb»
HCfMioa 01 llie i>kin, Thr worlttnif of M* hrnt-n-^iUting mochnnwcu ia
vtll tMm in llw ciu^ nf cxorrifc. i-inw i-vrn- miiHcnbir conlrnctinn giv«*
rfa* W hi-nt, Mcrrin- nm«l incrcnM' for iHp tinir (K-ing (be prmliKMion of hf«t ;
JH the li-xlilj' tern thrill lira ran-ly riiKH no much iw n degree cciilii^niili.', if iit
■II. By cxrrviM (h« re«piniti'in )• ouickcneil. nml (Iir low of licnt by th«
lung! iticn-w«H. Th« circulalinn of liIooH ir iiIrq (|ini.'k<-nei], tiii<) the <;ut«-
n«<i(i» voM-uliir nrcH'> lKs:oniing dtlnteil, ti lnr);cr amount of blow) pnmtt
lhr<Mi];h lh*> 'kin. Added lo tnt», the skin pervpim frcelv. ThiiH a Urge
UiMfinl of heal is \tnft to the body, xutficient to neiilriiiii:e rhe nddilian
nuiw«) by Ihe iiiiiscular eontrAetioo, the iacreaw irhich the nioro rapid flov
n( blood (hrou(rh the abilomtDal orgniia niii.'ht lend to briD|> about being
iBoiv ibau fiutlieieuily coiinlerscled by their ftiualler supply for the lime.
The MDM of nnrmlh whit^h is felt during exercise in eonseiincuce of ih«
ItnabinK of the akin. \». in it«>If, a token that ii regidalive eoolitij; i^ being
rarricd on. In u Airoilar way (he applicaliou of external cold or beat ilel^ta
ib own eoda, either iiurlially or completely. Under the influence of external
mid, tlK' culaueoiu veuel* ore eoiiRtncled. and the aplaiielinic vascular area*
dilaiMl, Ml that thi- blond !« wilhdmwu from the colder and ctMiliT rc>;[»iifl to
the hotter uud hi-at-pniduciD^ orj^an*. Tht* vancular vbang« niny be used
to explain tlw fact that ntripjiriig iiiikeil in a <^ld ntn]cMph<-re oOen gtvi-* rife
to a diiliuct ini'rcttte in th« mean ti-inpfmiure of the blnoil, ai iiidicatnl by
a tbi-mioni«ler placed iu the mouth, th'iu^h |Hi9»ibly thi' cfliyl mnylw partly
doc to an acUial incrwiBc of the pMdiietion of heal. L'nder the uiflueiice of
ext«n»l wartiilh.on the other bund, the cutaiieoua venels are rlilatol.a rapid
dtachar^re of heat takes place -. and if the circumstances be such that the body
can pefpire fre^'ly. anil the perspiration be readily evaporated, the temper-
niurrr '<{ the lto<ly may remain very ticar to the normal. even in nn enccHively
bot atmosphere. Tliu», more than a century a^. two obacrver« were able to
remain with impunity io u chamber heated even to 127" C. J 'ilSIJ' Fnhr. ), and
with e««e in one «o hot. that it tiecame painful for them to touch the metal
buttona of their eloihinK. It Is unnecea&ary to (five any more eiamulen of
ihb rejiulatton of iem|>enilure by variaiioos in the Inss of beat; iliey all
readitv esi^aio ihentM'lrea.
5 M4. Tfif pro'luctiou of heat, ill varlnlimu aiid reyttlaiion, Ai we have
■Umdy Mid, the eiact determtiiatiun of the amount of heat produced in tl»e
KTlog Dody b atlen<led with grcm dilficultiiH ; Mill, certain wncluxiimH have
Iwen arrived at Imted partly uii cliri-ct cji In ri metric n)>M'rvation». thv more
nct-nl uncD with imprnvcd calorimeters bcin^ niijeciallr valuable, and
partly on what *«eni to be tnurtworlby deduclioitt from oWn-cd clieniical
clungn.
The rate uf production nf heat in n living biwlv Ik dcli-rmincd by a variety
of rirciiRi»4ancc«. Id the finit pla™. what may iic cjilldl the gcnemi rale of
HtetatHdiim. and » of the pnxhii-tion of heat, vnricw in difTerent kind- of ani-
mals. Of two Rnimai« of ihi:' uiinc hulk and weight placed under the Miine
drrnmnanoea, one " live* fulcr " than the other, melabolixcs lie living Hab-
tlsnoe more rapidly, and »> pnxluccs heal more rapidly. Thus direct caloti-
netric obaorrationi, a> far at thev at present go, show that a man, on tlw
Kvenge, produces more heal, [«r kilo, \vr hour, than doe» a dog, and a Aug
non! thaD a rabbit. Probably every ai^eciea baa wliai may be called ita
n
046
N'fTRlTIOX.
•pon'ic corrtiPiFiii, nrnJ ovorf in<liri<tut1 bis {MTtniinl co<-tfiri«iil of hail-|>n>-
auction, the r<i'tHdrni hvlag Ihv expn-^iim of the inl>om (|UBliti«c proper lu
the living futisOncc of th« f|>pci«« imd of lli« individual.
A litrger livioe body nitl amnnillr produce inor« h«nt thnn n ainallw
living Iwdy of ihe Mine nature, Hince t)ie larf^r Ixxly poaaeeae*. to to
epeak, > gr«alor number of beat- prod ucinj^ unit«. Bui iliis is DMiltmlitfd bjr
Ml oppodng teudcncy. The smaller body, liavioK relatively to its bulk a
larger amount of surface, loses heat at a niorv ramd rati? iliau dues tlie larger
body : Slid, therefore, to maiDtain the balance t>etwe«a loss and produclixa.
■0 as to secure the iranie cousinnt bodily leiaperature 'and, as ve Imve jiurt
seeu, the bodily temperature of warm- blooded animals is n-niarkniily iinifi>nn>,
it must produce heat, per unit of its body, at a more rapid niit>. Ah a nil«,
the gmt«r loss of lieal owing to the relatively greater nirface it no marked
llMt of two animals having the same couatant iKMJily t^mperaturv, of two
mcieeof uammala, or of iwu iiidividuab of the wim« race, weabouhl 4>xpwt
llie smaller one to produce a rt-latively larger ainounl of heat. And dirwt
cahiriini-tric obaervatioDS shon' thiit thU i» to. The *tniggle for f-si«tvnc«
hat rained what we liave ju<t called the specific or penKxial i-ocificiunt of Uia
imallcr aiiitaal.
From what we have »een conccniiiie llie iminodiale cITccls of a niial. «a
•boiild be inclined t» expect that fooa would temporarily increase tho pn^
duction of heat ; and not only U this view confirmeil by common oxperieoo*
an^l by our own acnsiitionf. but (lirei.-t calorimetric olwcrcatiuRS »ffam experi-
mental proof of its truth. In llic dog it hu hern found thai the rate of
pr>.>duciion incnaeea aller n meni, reaching its maximum from the sixih to
the ninth hour, and then iWlining to a level which may l>e ragarded as Uui
teeured by the general mcTabolisn) of the body, and which ap|ioaT8 to be
maintained with remarkable coniOaucy until aller long starvation the
economy begins lu break down. Thua. in some ex[>eriineuts the prndiirtion
at the nintli hour, after au onlinary meal of meat and fat, wa* at a nU*
about 'JO or 'I'l per cent, greater than that at wliich it wun goiiiff on before
food wa» given, and to which it suhseiiueotly sank before IimhT was ajjain
given. It would ap|>ear that if sugar Ih; added to the nwal the rise hctoomat
more nnirkett nt an earlier jicriod, as if (be economy found ^ugnr raaicr 10
consume than fat. Thin, howover, i« a matter which as yet re<)uln» lo b«
n«ire fully workwl out.
Labor, mufculnr work, lias n powerful influence in incttasing tlie tirodtio-
lion of heal. As wc have seen, of the totjil heal produced iu the Body, a
certain portion must alwayx Iw attributed to muK-utar oontrodicKts, which
even iu the most qiiiei hoily arc always going on : in au ordinary active body
a considerable •|imntity of heat must be thus gencralnl. Heiiee, Ihe ninr«
active the body the greater the production of li«il. As we slated before
( !i >^T ), In a contraction the proportion of the energy set free 10 do work to
that set free at heul appears lo vary under diHcreiit cin-umslances ; and the
increase of heat due to labor probably varies in a Gum«|>o«ding way. The
deiails of thi» relation have yet to be worked out, but wc miiy at least con-
c I II lit- that, when a man pushes bis daily labor beyond the i.>0,000 kilo-
gram mc-met res. Ihe additional energy thus leaving his bndr as work done ie
not taken out of the H.50.00U kilogramme-metres given in ^ o'.^ as the average
daily output of heal, but the total selling ttw of energy and ibc total pro-
duciton of heat is at the aauie time iocr«ised. \aA it need hardly lie faid
tbni ilir figures in outstion itive only an average eettniate for a nian of
average biiml and weight, takiiijf an average amount of average fiKid, and
doing au average amount if work.
THK E.VEROY OF THK BODY. lUT
i 539. Tbc tiroduniuii of beat ibus determiueil by thtve M;\*«riil influeneea,
t>{ nliicli are ihrniitL-lvc? rei;iil(itctl b^ tbi- iitnouB iiTsti-ni, U furtber
. _ ^ihiUd ill a r^nmrkalile maiiiu-r. Fur it i:i tii>t wlely br viiHiiUon* in ibe
Mttfbeat ihni tbt- <v>iiMant U-oipc-niture of th« ivurni-liluuilinl nnimal U
amjutaintt]. Vurinlioii!! in tbi' niumiul of htuil ar-lually jft iifnitnl io the
body i-oiutiliilf ail impnrlitDE Im-lur, not "iilv in l\w muinU'iiiincc nf tbt-
nnnml lisnjwralun*, but aim) in llie )>nid)tctuin til' ihf abiKirmallv bigh ur
luK iMup«r«liirvii uf varioun (lii>mM^ Many ii>niiiilcrHlii>nH bare fon^ I'tl
|iby»i<>)iiu;i*U> to (iiiitic<-t lh<' t'xii>tciico of a iHTvuiiii incchuiiiini, by which
dhrtnt itapulmr* arming in thr nktn or clwirlirn- might, thnmch the eeiilnil
Mrrrou* •ymrni. urigiiiiilr oDcrctil impiilnv, wh'w rrtV-ct nimhl Ik; to iDi-mtw
or In iliminivh the nK-lab»li«tn of the niiiscW '>r oth^r nrganx, nni) tltun to
loonMV nr •liminish rbr iiinount of bent grnvrati^l f<ir lb« lime bving in tlw
h«lt. Tho rxivtenre. in fact, of a mfiabcilic or thermogenic tiorvdus
BM«-lMUi»ni, oompairabic in many n«]>ectF In tho va«oinoti>r niccbaniam »r to
thr various creating iiervnns mechanUms. fcrnis in itwlf >' /irhri prutMble,
And tio bav« ozperimcnial eviiteni-e that Ruch a i»i-chnnism dots mlljr
aiit
The viirmbloodcd animal is diBlingiiiali«<] froiD the colH-bluodtxl anima]
by the fad that vheti it in exposed to cold or heat it does nut, like the latter,
bccMiie colder »r holier, as the cose may bo. but, within ei'rtain limits,
BMUoiaina its nornial temjierature. If the maintenance of the temnerature
of the wanii blooded aotmul during exjKiBiire to cold is aasiMea by an
iocreved production of beat, ami in uoi due »mctly to a diuiiuislied Urm. w-e
ought lo fintl evid«uce of an increased roetaboliflm during tbni exposure.
Vr'e ought to find, under these circuBMlaDcet, an increased production of
cai-lx>uic acid and au increased oonaiimptioD of oxygen, Hince it in ui these
pnHliirU, rather iliao to the nitrogenous uctors, on tlte peculuiriliet of which
a* nnoertain sign* of metabolism we have already iniustetl, we mu»[ louk fur
uxlicatioiw of the riw or fall of mctaliolie activity. Of th«c two, tbc pn>
duciion of curhontc acid sn<! the consumption of oxygen, the latter is tbc
tnor«tniportantand Inislworthy measure of mclaholiam,tv|i<-cinlIy when Db»cr-
vnljons an- toade for short period* only al a ttnic: fur, tut nc have nvn in
Inaliogof nvpiration, the exit of carbonic ncid '» more ckiMly dependent
on the acttoo of the respiratory mechanism than is the income of oxyeeti.aiid
carbonic acid can be retainol in loow combinatioo, and to tenipornriiy stored
op by rarious constituents of the body.
Takinjf, tlwo. the consumption of oxygen, and, though nilh le.<e confidence,
ibe production of carbonic acid, as a measure of Diclanolic activity and so uf
bent-production, il has been shown that a marked contrast in this respect
eztsls between oold-bloodetl and warm-blooded animals exjioeed to changes
of leraperature. In ibe cold-blood animal, cold diminishes and heat increases
ihe metabolic activity of the body ; as the temperature lo which the animal
is stibjti^'ted riset or falU, m the consumption of oxygen and production of
cmrbonic acid is iucrcAsed ur lessened. The IxHly of a oold-blooded animal
beliavea in this respect like a mixture of dead substances in a cbeiuisi's
ivton ; Iwat iiromoiea and cold retanh obemical action in l>oth aises. V'eiy
dlffrrent i* iiie liehavior of a warm-blooded animal. In this ca.'ie. witbiu a
lower and a higher limit, cold increiiMiti and beni diiDinisbw tl>« bodily meia-
ho)i*ni, as shown by the ineri-aiR'<l or <liminii>)jei] eomumplion of oxvRtm and
pr<»lurtion of caiuunic acid n> the temjierature falU or risett. lu tboK!
animals itierc is obviously a mvchanisDi of aunie kind, counler-acliug. and
in<lerd nveiciming, (hpso more direct cS!mU which alone ulitain in cold*
bliH^kil animal*. And that this mecbitiiba is of a nen-'im. natuiv, U iiuli*
catvd bv the following facts:
xuthitiox.
WheD n wiiriu-liliKKlod noininl it |>oifii>nec) br iirnri. thi- teiu|H>niliiiv fnlU
ati<l Ui« nictabaliem, iiicosiirc'l by llio consumption of uxyi^ii iinil ttic [iro-
ductHrn or urbunic ncid, sinks alsa ; and thai tb^ Inlter Ji llie (.-auite, not ihc
elfecl, of the foraicr is ehoivn by llie facl tbsi ihe lueinlxilUni continiH* to
fttll lhoii(;b low of btrnt be ]ir«rente<t by surrouiKliiiK tbe animalt witli wra|>-
piii)(n I'l' i.i>tl>.>n wool. In eucli a uranEed anininl, exfiosiire to liislwr l«ni-
Kraturw imKnionU and cxi«iHnro to lower temi)ora»»r» dlniitiiKbo meta-
liiuu; tbi> urarix^d nurm-bloixJcd animal, in laci, WliavM like a ooM-bliMHW
Miiinal. Similar, but iierbapu not Huch Htribiuj; or «i couMaiit r»ult», are
gaiii«<) by division i>l' tlie ui>^iliil]a oblon^ita. Aflor this openttioti tti« t^m-
peralurc of tbi* body ftitikx. and the fall, tbousb )>Hrllv due lii inrraawd lam
of heat by the i>kin, caiiAe>l Iiv iht; dilated vooditi'm ul* Ok cuUinvQUa vtMeU,
i» nliM) n<s-i>iiit)HMit<d by ilimiiiiHbi'd nictftboliiini, aiid >», tb«rclbi«, io part diM
li> diminiibcil |iri<rliK-iion id' Kent. Ami wben an aninu] b in thit cotuUtinti.
CxptMnrv Io hij^bi-r [ciniHTiiturv* incrriiw^ and exiHifurc to Iowit teinpvni-
liinx diniini^hcn ihv bnaily ni<:tnb(di«m. Wo can ncft explain iImw re^lu
by wpiMwn^ that, under nnrmal cniidilionR, tbc lanjclci, which ju we lwv«
Men conlribuli) nn Inr^rvly to ihu totid heal of the body, are placed, by ia«Dl
of their tnolt-r nerve* and the central nervnns system, in somfl BjMyiial coo-
Dcelioii with the xkin, ho that ■ loweritii' of the temperature of the skin leadi
to an in(-r<-«i*e, while a heighleninK of the icnineniturc of the skin lewis to ■
decrease, of ibe ntuti-uUr mrtabnliBm. Furiber. the centre of this ibermo-
laxic reflex nicehntii«m apiteiirs tu be placed siimewherc in tbe nervous syvteni
ab[>ve ihe spinal eonl. When iirari is i;iven, the reflt^x cliain ia broken ut
ils miieculiir end ; when tbo spinal cord is dividi'd, tbe break is nearer the
centre. Whelher w-e should conclude that the working of this reflex
mediaiiism is nf such a kind that culd in ihe ekin excites the centre i<i a
beat -producing activity, or of such a kind thai warmth Io the skin inhitnti
R prcvioustv exialin); autnmittic activity nf ibc centre, iniiy bo lel\ liir ili«
prcMHil 11 nileterm tiled.
Wv may add, ihal the niii*eulnr nietaWiam which ihun helpii lo Rfulal*
lemperalurc iiewl not involvu viiiibli> mniKiiliir [imtnicliuns. Althcaaiuf
time, the heat jgiven out by the musclm will be Icnipomrily iuereasHl at
veery contraclion which may iict'ur. Thus, the shivering which fulluwi
expiffure lo cnld dinlinctl r beljM to warm ibe lindy : indeed, e»nir obwrven
have been led to think llint. in man, this visible elfoct ofoold iditrs a uiiM
iiujHirtaul part in bis heat regubitiun than the invi*ible actinne widcb we Iwre
jllM de>orit>ed. We mav niso add that the regiilntivc nervous mechatitfm
mav apparently be overborne by an exposure lo too great heal or cold.
When, I'lir iustiuice, the cold lo which [he animal is expoeed becumee excea-
>ive, ibv reaction of the ihenuotaxic nervous sysleu Ja powerltw aKftioH th»
direct nctinn <iii ihe tiuues of the depi'csfiiny infiuences, and tin metabolism,
lo^lhvr with ibe li^mperaltire, lunk^.
Tite n'9ult.-< with iiniri Jnsi mentioned «eein t(i show that this thcmiiriuxic
ncniun nit-chanisiii \imiTs chiefly ou tbe skeletal inu»cle4. Whether tbe
glumluhir orgiin* take any |)airt in it. or whether they have a luetakolie
tbennoiaxic macbinerr of their <>wu. nf such a kind, fur eiaiuple. that the
increiut' »f bent proiluclion <lue to f'x^l i* tbe remit not an miiHi of the
initnedinte cuDMiniptiou nf ]>nrl of the IiiihI iiavlf (luxus conaumptionj n^ of
tbe i>r«eence of fotjd, in the alimiMlUry canal or after nliMjrption, etirriujt Up
llie li\-er lo increased nielabolisin, we do not at prc«ciit know.
S 536. Id a number of cxperiinenls it Ixw btwo sliowa that injttrio> in, mch
ni ihow oau«ed by punoturc or galviinie cauterjr, or tkotrical Mimulaiion oj'
limited poTtioDB of the mure ceotml portions of tbt bniin. mav give rise lo
n ^nat iiicreoae of the temperature ol Ibe bixly wiihout prvxlucfng any other
I
I
mHcMl ■jriiiptoRi. Thr incmiito if shouD by tbo increow of nietnb»lisni,
bcmMtl pnaliKlkni of cnrlwrnk- ncitl. mh! incrfni'cd (^iii«tim|)linn nf oxvpwi.
m wtII a* dv dirrct r»lririin*tric obBervulimis, to Ih' iIiip t" an incrcatwa (Wn-
duction of Honl. This Dahirnlly suggv^rii ihnl the [Hjriioits of the brain tn
^BOtinn ronUiD the hypolheLiail fac«t cctiln' jii»t nienlioned, the ic«ioti in
MimulntioD exdliog tlie centre tn iictiv-ity by direct nctioo aii il. iiislcsil of
IB the usual reflex maiuier. The mnttcr hue not, however, ai* yel been I'Imrly
worked wut : aoil indeol obeerrots are doI ii^reed as to Ibe i-xact parts of
llie brain injurr lo which, or EtimulatioD of which, produces lite efleet.
While some place il in the inediiiD nnii boaal portions of the corpus Hlriatuio,
oUicfa niaiolaiu that il t» situated in the optic thalaiuiu. The fact, however,
remains that aa aAeetiun uf a very liiiiitt^d fiorliou of ibe central nervous
mten) may, without producing any other obvious ojlects. w> increase the
Iwai priMiuviiou of the body as Ut raise itie temperature uf the budy »e\'eml
dMTcea.
1 087. By rc^lative inevbanisoui of the kind ju^ dijcuwed the lemjiera-
lure nf the warm-blootled uuimul h tnaintiiiuiirniUitD rvry narrou' liinito.
In onltnary bailth the temiieratun:^ of ninii varia> bi^tweeu :J6^ atwl ^°, the
itamxiriT Jirnibt Wiiig :iG.'i5' aud -il.-f, witvii tbu therniomnler in placed
ID the axilla. In ihu month the readinjr of the thvrnto meter i» nomfirrbat
fO.iA'' to 1.4°) hifthur; in the rvctiim il isVtill bijrh*'r< about 0.9°) than in tbe
mouth. Tho tcmperaltire i)f infantH and childn-ii \» xligbtly higher and
uueh nora Mnoefitlbh; of variation than lluit of adiills, and nOff f<ir[y tmi*
of ago the avenge maximum iL-miHTaiurc (of bi-allbi i» mnu'whrn lower
tliaa beAm thet e{>ocli. A diunial \'nriation, iode)ii.-iidrnl of food or other
dnuBMUaoea, ha* \nvn olwcrvcd, thv maxinmni ranging from 9 a.m. to 6
fJL and the minimum from II r.M. t» 3 a.m. McuHs cniiw flomditnw u
alight oteratioR, tomctinics a »ti<;hl <lcpr«9Hi<>ii, the direction of the influvtH.>a
dvpendiag on lh« nnturcof the fowl — alcohol wkou always to produce n fall,
KxeniM and variaiioiu of external temperniure, within oi>dtnar>' Umiia,
oauae a rery slifcbt clianp, on eeoount of the cofopenMtinK influences which
bare been discuawxl above. The rise from even active exercise duce not
amoant to I"; wbeu labor ti carrieil to exIuuMtion a deprea^iou u( lemji^rn-
lur* may be observed. In iravellinK from very ouhl lo ver^- hut n-ciona a
variation of lea than a decree oi-cunt, and the teRi|ieraturo of inbabitauls
at the tfopica if pnu'iically the miw! us of thivu dnt-Ilintf in arctic rrgimu.
S fi38, Stany of the maladii-ji <if the bixly are charucirriMil by nn incmM
ef tlta budily temperature known ax" fevur"or" pyrexia," ttiv t hi- nun meter
Terr ftvquftntly ruing to 39* or 40% not unfr^iucJilly to 4P, an.i nl lime*
rraching 4^" or even 14'; but thtwe hiulMT ii'iii[>cnit(irm cannot long bo
Inirne aiihout the organtfni failing. And, nn uc luiva nid, any imrt-iur in
BHin of the boilily Icmpcmliirc beyond 3S^. or even hcy-nd '-iT.-t". indicate*
■ooM disturbance. In mo>l ca*r» the rine of trni{ieralure hiu> a dclinile
oltjactive cnMe,BOino local inflammation or sup{Hi ration, or, n» in ijiecilic
ftvrm, Ihtt pmnuco in ibe economy of lotne " lualeries niorlH," of iho nature
nf an ort^niAnl germ or of wime other naturi>. We cannot here diM.-UBs the
oovnrction lidwwn local inHninmation or the specitic imiimiu and the high
lani{>criiturc, but we have iucreaaing evidence that tlie ui|{h iem|)eratnrv of
fever is due, not merely to n diminution of tike Itua of heal, though this may
be a fiictor, but also, and indeed chielly, to an increased pr»ducii<xi of heat.
In fev<-r the prodiietiou of carbonic acid and the couMimption of oxygen,
tJuil is lo say, tlie metabolic chattel of the tiMuea, are incrtuwoil. The urea
also ia inereflswl, ami that in suob a way at to nmlirui the view already
exnraMod that niudi of the hint coniea from such a mclabolitm of the skele-
tal muMlca aa, unlike nu ordinary contraction, dirvctiv involviv Ibe iiitro-
i
600
SVTKITIOS,
getuMs elenieuU. The itiordinnte in«Uibo)lMii uf ihc Ivalr nt Imrxc ihui
c-h a nici eristic of l^ver is slio<ni by the irastin); which it i-nliul«. (^lorinwUJc
ohoerrntiooa also ahair in a direct niunner that (li« productioa of btai U
increased. Of <.-our«e, mere iocreawd produclioti nlone would be inaiifficiait
U' miae the teniperniiire of the body, for it DiiKht be met, up to k vrrt bigb
Hmit, bv a cniupeDiuitiiiu increttse of luas of heil : but in fever ibi* iiim-
itetiiiniiikii t.t uuDiiii^, luid it Li [>erba{M thb abseooe of du« regulatiun «luch
■« iiKal muit chiiraL-tc-mlir i>rthe febHIo cuudition.
In »>inic ii>u1uilift> the ho<lily ti<itip«raitiire &lb distinctly below thv [i*jriual
avornjp^, n-ncliiu^' Tit iimtance ^-^', or even lou-er. Ill such c*»e« there oio
lie little iloubt that the couditiwu id due to dimioishM) ntetaboUuD and
diniintsbcil livat pmduclioii.
One of till- iiiuwl marked phoiiiiini;iiii nf iltirvatinn \* the fall of l«n>jieni-
ture, which tiroonKr; very rnpid iliiring the iiwt day* of lift^. Tbe lowered
iiintatHiltMii dimiiiixhw ifit.' pHKhiolinti of hvnt, ami the lonered leiDjieraiure
in luni »till furthi-r 'liinirii«hr« the nivtabiilitii. lii'lti'il, the lo«r tetnpera-
tiirt! i--< n )iiin'rrliil tiu'tiir in hriu;.'iiig iiImiuI death, lor lifv may be mueli
pr»Ii>iigi-il l>y ivroppirig n starving auimnl in sonie ImuI ounductur, so asM
MODomixv thf bo<lily heat.
§ 639. Kffevt* of ffrnil heat. Aa wp «aid above tho n^lative heat ■?■
chanisni i> iiiinble lo withstaud the fitrain of too gran an externnt heat m
too prulongf^l uii exposure tn a great but less degree of heat. Tbe tempera*
tiirc of the body then rises above the normal ; and it has Iwen obaervod thai
th« tcnipcrnlure is more easily raised by warmth tban doprvMud hf c«dd, at
least when neitber arc very inteose. When eilber in ihii way by exiemal
nrarmib or throu);li pyrexia the temiiemture of the body is raised foiiM B'
or T' above tbe normal, to 4'V or thereabouls. dealh speedily eiltMiai> Tbo
chain of evenle thus leading to death has not been as yet cletirlv tnadr out,
and moil likely (he evenla do not take exactly the .-uue course iii all cam;
but tte shall probably not «o far wnmg in altributiiu <Ieath to ttie fact that
the hi|^h temperature burrie* on the metidii-lism of the «verttl tinues. of
eonie luore than othen, at micIi a opt-ndthrifl rate ihnt their capital is aoon
exhausted. We have tuxn (£ 'i~2) thai loo warm bluinl pruducra dyapnwa
and soon exhausts the metabolic capitnl of tb<- rtvpinilory centre. Too wami
blood Mmilarly hurrieM on the bent* of the bciirt; an explmion of tlie coa-
traclile ^ulutani'e i.i cAch timo preniaturrlv brought on before a sufficient
quantity of cxplu«ive mbstnncx^ ia accumninted, each ntroke lK«nm«a more
and mora foeble as the rate is quickened, the Ih'hUi bi-iv>me irregular and
filially cciuc. lather of lhn«« two events bIoik- and ceriainly both luj^vtbef
are enough to brine the working of the bodily mrrbaniiim to an end ; but
Other tiiMuat beside the heart and the respiratory centre arc suflbring in tbe
mme w»y, notably the rest uf the oenlral ncrx'ous syvtom. Thi^ too, it
being hurricti on unduly in lis inner changes, so that not only coni>ciou«ntaa
is lost and other objective mnuifcela lions ot nervous action go nroug nr (iiil,
but that regulatice grasji of ihc central nervous system on Uie timiM of the
body at targe is loosened, and tumuli lakes the place of order. Wbdher
tfaia or that sign of dieoriler comee to tbe front, whether, for instance, on-
TulakNU take place, would appear lo depend upon the exact Uim taken by
tbe sboornial event^i. A heat-stroke, more commonly known as sunstroke,
the eOMOtial condition of which seems to be a rapid rise of the tcmperitlure
of ibe body, owing lu a fuddi-n failure of the therniutaxic mechaniMii, the
aymptomi var>'. Njuieiinxv thi- heart Biuldenly gives way. at other limes the
nvpirnlorv ceutrc nei-ms m \w more diii-clly iirtceied ; mmeiimes convulHton*
make their appeurauce. but niure i-uniniuuly death tiikei place through a
I
BODY.
<aBMlow coDdiiion of (lie hraiii.ati initiai phtue of exdtement »t'Ui« dentml
armwB >Taleni beiiif> uot iiufrequ^Dllv wiiuewei).
)laninikliKD muscle, it will be remeiulH^n-il (i M). becomes riKid iil alxtiit
Hfi but deaUi probably alwuya u<-'ciin befitrc that hi^ihcr teiii[>ersturv is
mchcd by the bluod. so tluit a nurldeii rigi>r luorlui from beat (rigur calorls)
naihii be re(,-»riM an a faotur in ilealb from exiHAiire to too neat heal. Ilul
ibiMiIil Lliai temjierature ever bv reached by tlie living btiay, all trc Itixtw
lods US to infer that a *u<tileu ri^iiilily of (lie whole biMly would at i>oce put
«a abrupt end to life; to nUfiiniie that ii biimiin budy can truly ri-|^Mer thia
or a hign«r tecnperalurc whdv rrmaiiiinj; alin*. t« Miy iiotbiug ori>h<>wiiij{ no
tokena »f ditfnaa, entail* the mp|>ricilit>ii that KUch n hudy aui diflVr from iia
fellows iu iu abaolut«)y ftiiiilanirnlid qiiidilio*. and yet make Do oth«r aign.
i MO. KJfttU of iftrat fold. The fdect« of a loo great lowvring of the teiii>
]«niture of the bocly, which 'a ^crally tlui result of loo grtM external eoUl
lod nrvly if ever arittf* from internal cnii»» towering the meiaboliwu and
thiH the prodnctioD of lieui, nre in their oricin the reverwo of tbn»o «f a ton
bi^ temperature. Tbe meMbolisni of the tiwucx is low«r<Nl ; and nut only
•re the aaiabolic oban^ which lend to the «etting tree uf energy thna
alAcieil, but the anahnlte changes abn share in the dcpranioii. The " livinv
•uhManev " Gilb to jtiecee Ion neadily, but is uW made up leea readily ; and
could ihia nlBcktoing of meiabolisai be carried en in tlA; acverul ttMUea Kt
a rale pn>[xirtioData to the rate iit which each tiwue lives, life might thuN be
hrou^'ht to a {)en(x-fid end by gradual arrat of the life of cturh pnrt of tUu
whole iMMly. And, indeed, in some CftMi, where the lowering of the tem-
Deratura takn place gradually, »outetbiog like this does occur even iu wntm-
oiooded auimal<>. The dimiutshed nieUibolism tells Aral and cliieflv on the
OWtral nervous fvsteni, (specially on the brain and more particiilnrlv '>ii
Ihoac parts of that organ which are concerned in coneeiouaneeB. The intrmsic
toweriog of tbc cerebral nietalHiIiim ia further aaaiated by a slowing of the
be«n-l>eRt and of the breath, drownincM U sunceede<t by a coodittoii very
like to, if not identical with, that known an xlei-p, which we »liall study later
lai, but by a sleep which inveiuihly ptuaev into the sleep of ilealb. In some
CKM9, however, eafieciallT tbote in which the loweriDg uf tli« teoiperalure is
iiiddeo and rapiil, dlionleni of the nervou* syatem intervene and cunvuluous
like tbnae of aajthyxin are iiroducud.
$ A4L HihfrnaUnn. In the mnjority of warm-blonded luiiamU, the cuiidi*
tiMM thus iuducnl bv oold an- rapidly fatal, and mijreover in their progrcH
Tilj waoa mch a Btogi- fr-nn which recovery become* iniiMMwible. In the
Ktm of aocne few aninudK, »ciit(rml mcmbem of Mveral groupx of mammalia,
• ■inilar di-premion of nH-tal>ii|i»m by cold is of yearly occnrrcncc, taking
place regutsrly a» tin- «xlenial icmpcralurc falls in witiler, and bein^ thruwn
off regularly lu ihc rxlcrnul temperature rises in spring. 8uch luiiniab are
fpokeo of lis hib).- muting atiimalt.
Wb ara not able at present to explain why these animal* behave iu ihia
wmy. It is obvious that lor some reason ihev lack that power of rcactiou
against estemal cold nhU-h, as we have seen, » one of the eh a raci eristics of
toe warm-blooded animal, but we cannot state what is the dilfereuce in their
Mooomy which lewis lo this lack. The " winter sleep " is undoubtedly due
to (l>e cold of winter, and may, in some ca<«e at all events, be induced by
coM protluced aniliciallv in Mimmer: but the system is prediiposed ami
aila[>ted to undergo the ebungi^ nt the ap|H>inied semon, and a dormouse may
Gill into winter ileep itt a temi^racure m winter higher than that at which
it awakes in spriug.
The pbeaoraena of the biWruating mammal mar be described as tliooe
doe 10 a lower rate of mctaliolism, ami Ikcnce to lowered activity of the
MTBITIOS.
tlwuo* in genera). TJie heart beals verv bIoitIj-. nod each heat m »l !»*■£ «rf
but inotic-rale ncrt-ogth; iiud ihe Ureaittt are few. reebte, and far beiwi-m.
KoKfitnaioo niid drculaiinti are thii^ fT'^i"^ ou. but go oD. so u> sp««k. at
almost ihft nlonrtl |)f»iblL- ral* cutinutem wilh (he coniinuaiic* of th« work-
ing of the ecrtiioray. The liieath» an>. lu we have said, few and far Iwiwtwi.
bill they Kuffic^' (a carry lu the tiiuue« the uasl\ ani^xini of oxyj^a irlikh
tliiw Di-ful (iikI I'i carry ulf the .■■mnll iiniouni of rarbontc aeid w hicb ibef
jinKlitcf. So Kiiiall i» ihc n':>iiii-nli'>ii of thi- liMuee that in the dept)ij> uf Inc
wint<T iWp the vniDUK MiHid i« alm<wt en bright ob the arterial, tlie color nf
u-liieli i< niiirlv normul. And ihc «mali aiiioniii of dettniclir« kauMic
changrv w)ii<-h i« goiug oil U Khnwii by a ehniigi' iii the rir«[>irsl'>ry (|UOt>efil ;
oxTgcn i^ liiken up out nf pniprirlion lo the rarWiio nciil i-xfiirttL iDdeed,
it biLx Imth iiWrvril thni ii dnrnmim' arttially tiainril in u'<-ight during a
hil}t'rnnT.iii); |K-rm(l ; it. diiichar^'d duriii); ibtn pcrii") m^ithur urinf- uor {<^x»,
nml Ihc giiiii in weight wn« the excm of oxygen takon in ottt tli« carinnie
acid j;i\-cn out.
As far B» rejmfl'' the other finx-iioim i)f tin- h'Kly all that oau at pnwat
Ijcsaid is that iticwvvral rund.-inicnial nctiviiic» of lherari«tp» ti^HK^. thoiigfa
lo«n-red. arc itill continiiol Tcrv niiicb as usual. The lutiwlM ami Ufrrmu
elements are irritable : indwd. the bilicrDating animal may b« sa'nk<^l thougb
nilh diflicnlty by mlcqiintc stimiihition ; and ik* an instance of tb« funiia-
mentat similnritr of the sleeping ivith the awnke condition, we may My tint
the slowly heatinc heart can. during liiljernation, be still furlhvr slnired or bt
arrested by etimulation of the vagus nerre. The essential fntliirc uf hiber-
nation in fiiet is that external oold ie not resisted by tbe tiK-nnolaxic nervottf
niechaniBm, Init lowers the metabolism of all the tisraea. and thus loners ibe
functions of the whole body. When even in deep winter the liibcrDaiiag
aoiiual is exposed to adeiguace warintb, the inereased temperature awakes tm
tiwncK tft incrcaaed metahutiiini. and ibc awakened animal regains tbe bodily
t«'mp<Taliir4.' and aetpiiri-ii all the piin'cni nhich it jHiwiiai.i) in niidHunimer.
PrepHiatnry to the oncoming nf hiheriintion t.hi- boily lays uti nnusiiallT
large stt)re» of ftit for thi- winter'* extx'nditun^. Many hilx-niating animali
piMftw a " hihermiling gland." tbe Pffl.i of whi<'b iKtrome hnulnl with fat In
the atitiinin and loose it dnring hibcrntition : but in all caMw tbi' gn-at itutc
of fat is in the atliposc liwin- geiicrulty. Thi- liver of the hilwrniiling ani-
mal, at all events of the dnrmmwe, i"nnlains a- eonnidorable (jiiantitv wf
glyoot^n, which may be rcgardc"! u» fjoite cimipnmblc to the hepatic glyco-
gen of tbe winter frog (S J-i6). The fat thns sfired np Wforv th« appmadi
of winter serves as the main snppiv of material for metabolism in the wintwr
elecp. Since during the whole biWmating fieriod "onic nmonnt. al lensl. of
oxygen is at the connnnnd of tlu- tissues, we have no renson to think thai the
metaboliBm of hibernation is fundaiuentally diRerent from the metabolism of
ordinary life, or thai the stored up fat ^iilfers changes and gives rire to
energy in other nays than by the oxidation ivhieh fat in an ordinarv war
undergoes in the boilr. Nevertheless a detailed study of the mvlabolisui of
liibenintioii aceompunteil by direct catoriuietric ohservatioiw irmild probably
diBcloae intcn»itng reriulta.
Ox NfTHITIOSf IS GEXER-tL.
S 643. It mar now W profitable to take a brief Him'ey of tbe varioua
cnnchisiom nl which tvc have arrived concieruiug ihv pmhlem* of nntrilfou.
We hnv*; seen that the several liwinftt, using Ivniph a» a mealinm, livo upon
the blood, taking ujt from the blood the mntennU for, and rvtumiog to (be
be producli of. lh«ir nwinbotUm. Tbi- blootl itself w« luva aIw wen
'litenisbml with Wwti froiu ilii' nlitiicnl«ry ciinnl tan\ with oxygea from
lun^anil to bt- frtrtl Iroin nn»ta pruiliicbt by lucAn* of the Dxcretorf
oTBUM. In this tl<>ubl<' m-tioD tlit< ntir imKcHal nl' ihv Ibod on tliv one buid
nBdergue*. bcln<-vn its bi'iiiL' nlncfil in (1h- mouth nni) its Inkine part in the
■HUUJiflin iif iW tiMiii' wind) ultiniuielv u»ce it, rnniiy iDtcrtncdimio cliauRce
oafritd iw in vanuiu parta of the body, nml the n«4t« products iiimilsriy
uadfrco talcfmediatc thangca belween leuviug the liMue aiitl appearing in
iW uniw, tbc aweat, or thi- expired air.
VTe have lurth«r («en reoaon to think that tb« metabolic eveuts of tiie
hixly take placP in the niuin in the tiasuee. not in ibe bloud stream on ila
my between the bean and tbe tifisuet. Ctiangea, uroiter tu the blood ilaetf.
take pUoe in ibe bloud; tbe corpuscle*, red and wliiie, with tbe jilaarua
undeno like the mt of tbe body, their |>r(>i)er metabolic cycles, and in tliU
MiiM! lih-od inay be calkd a lieaue if there ii any advauitifce in udng llie
|ibra*e ; but, ufArt from these iiiirinHic bhuid cliimgiw, na far lu we cuu te« at
prtaanl, tbe metaboliaiu uDderuoue during their transit alons the bhut]
cbaimela, by the sulelancee whic% are merely carrietl in the blmid tVom j>lave
to pl<KC, ■» nu iiuignillcaut part »f tbe total lueiabnllscu nf lh<i tuidy.
By meiabolium of a tiMue we undci-ituiiiil the lulal vhemicnl chanjccn taking
tiEac* in ibc linue; and we divido tb»)e change* iuEii thmw which either
uirrvlly or imlirLt'tly are c»n(.'L-ni(.-d in the building up (nDnbiilici, and thow
which an' in like inauiier cDiiocnied in the lirotkinjc down (kiilahnlic i of thu
lixing Hibetancv. We •hall exjiluiu im^tcnily wluit uu mean by iht- words.
"dinieUv"and " Judird-llv " u«sl in [hiN omuection. .\ud w'c may bore
mmM tne caution (|S .'10) ifial thotigh fur convcoicuca take we unv the phnun
" Bving BUbctaticv." what m rvully niouit by tite words u not a thing or body
of a particular clwniicul c<^ni|ioaition but matter ui>dei^Dg a aori«a of
chanift*.
J M3. irince llie eevcra] tiwuea originate through a ditTereiit'olion of tbe
unpler, primordial proloplosiD. ne may infer that we have a ri^hl to apeak
of a getMrnil plan of metaooUam common to all the tiBsnes, modified in vari-
oai pvliculars in various tiasuea. It ia more reasonable for instance to sup-
pose that there is such a general plan oommon to both muicle an<l glau<l,
than to i>ii|i|K«e iliat the metabolism of the one ditlera wholly from or only
atx'i^l (11 tally ri3>vuible« that, uf the oth»r. And we may pnifiiably take tlie
oatritioD lit' auscle as exemplifying, in the midst of the features apedal to
tbe nusclo. the general plan of vital meiabolbm. The muitcle in a uorual
Mala of thing) livcfl ultimately on tJie pruteids, fai«, carbobvdnttes, salts and
WMtt of Uw tboil, and on the uxygeu of the iiuplrvd air, but livtn directly
(Ml the blood which briugH theM things tu it. Taking thi- pruleids first wft
may ask tbei|unt)on. How d<)«» ihi' bliHxl >upuly thu luuacle with proisidaf
TIm' hliM-id cunlniiM three cla-ticH of prutvicin: 1, *erum-albuniin, '2. glo-
bultn (paroglobulia), ainl -1, librinogen — that 14 lii say, tlic IxHly or bodies
nnocwiMd in tbe clotting ofblnMl, nhwe nature we lull in ^ '2'i n» tun wholly
and clearly made out. With regard tu tbe function of ihese thrtv kimU of
pmlwKis in the uutritiou uJ' muMle arc only cuncluiion^i ul present are indi-
rect noes, based chielly 00 the mtults of cxpcrinienis as to the relative value
of llieae substancn in niainlaining or mtunng the irritnbilily of muscle. It
is fouwl that when tbe woshnl out frog's heart (!i 162) is fed with dvObri-
natol hliud, the reslxrntiou is as fjood as with whole bloud ; oixl that while
tbe etiectii uf j-lohuliii are univrtam, and while peptone and lUbumuw appear
looct in an iujurious m.inner, the rwiorative oflects of eeruro>albumin Am
morkcil. Kruiu tlieee result* we nuiy provlsioiuilly infer that the muscle in
iu (total} anabolic dianges take* up and m livve uptui the senini-ullHiniin of
I
Nl'TBlTIOS.
the blon<3. Bill ibU n>nrlnsbu must be ivKiinled ns pmriMonal only. aiuI
iii<lcr<l iiucenaiii. For we niuM renwuibcr ili«t the blixMl iuiM>liea not only
ttitt futiil (iiivluiliiifc oxyKCtij for the niiiscle. Wt »}»& tJie coonitioiu under
vbicli tbc iiiu*c!e can ifvc niul arail iUelf of tli« food nfftred to iL The
C»ni[)k'x ucri'>ii!i lhmii|,'li wliit-h n (.■ertiiiii udatitilir uf i<nilc!(I and other iaai«-
riiil i> built lui inln Itviiij; intitcular suWniiix- Dec^l lor th«ir execation a
fnvoralili' nit-iiitini. nerd (vrtuiii jibyvjful and rht-niinil condilinott ; aix) 11
luitv Im- ibnt llii^ tuviiiiibk' iiifliit-iice of wenim-iilbiimiii ix »itD]>ly iliic tu jti
]>r<-M'noi' in lonii! wiiy nfsi.-lirig ibr! iruiisfurniHltMi into living subclani'v cif
raw iiiHlcriiil dill miininitij: id the muvtiibir fibre* nod not to it« flUp[)l;iD];
nvv raw Rmtrrial.
Dcxirivc i«. w we hnvv r<-|>m telly Miid, nlwnjrs pncecnl in th« bU»d b
Mnnll quantity, mid appoire to l>c the iiiily cnrbohydrate can>ti(iiHni of bbnid-
pltuina. Exficriniciite curried nut on u liirge animal, such a« tlip borw or
cow, bare rhown that the venous blootl coming from a rauectc contaiDr Itat
dextrose Ibnn the nrterin! blood going to the muscle, nod ihat the dilfemi<*
is much iDcreflHcd by throwing tbc miiiwle iiilo eooiractioa. From ihL'vt
D>ay proi-isionally conclude tliat dextroie is an ewenUal part of the food «f
I be muecle.
The blood as we have seen aim coiitaiDH a certain amount of fat : and if
we push the analogy betweeu the whole body and the iiiuM'le, we may infer
that the muscle iaKe» up fat as food for ilaelf fri>m the bbiod. Uut we have
no ex|>eririienlitl evidence in iavor of this. Moreover, we have seen that fiu
uud earbiiliydrale arc in the animal body more or lea* imnxfi-rable. We
have distinel primf ibat the bi>dy can trausform cartmhyilraie ini»> fat ; and
it is very pmbnble that il can trausfonu fat into cnrliohydiaie. Seeing h>»«
niueb more easily a soluble (lilTunible carbobydrntti like Hugar isu Iw carried
fr<.>m place to niaoe by the llui<U nf the body thiin can iiiiiiiU<-ible bts, it
w<ent» rensouable to sujijioni- tliul vrhrn the IumIv hat> to ilriiw up-iD ita slort
of fut in ilx' cell" of iidiprwi' iii»ii<-, the fat. on lesvini; the fal-ccll is tran*-
fonucd into :<iignr, it* carbiin i"i to »poitlc being dealt out to the liMue« in lbs
form of dexinx.-. Imlccd, we may perliape, dwolltng on tbc fai.1 that a mus-
cle though itself c«»entiitlly of prolcjd buiUI. turns ovct I^ KT> in its daily
work w much more carbon than nitrog<?n, entertain the rieu- that what mm-
cle want.i as food is n corlnin aniounl of proteid plus iin additional quantity
of carbon in some fonn or otber. and that dcxtroae is a convenient t'oroi in
which the additional carbon ctin be supplied. And ne may hold this view
without prejudice to any opinion that the carbon m bntuKht, while betne
built up into the living substance, may be again arrungeil as fat, and in the
course of the melalwlism of the muscle may be later ou separated ftom the
living substance and deposited in the fibre its globules of fat. But our
knowledge is at prescni iusuflicieut to decide whether this view is true or na
The various sails bn>ugbt to the muscle by llie pbuma, though they itupply
ao eDersr are ss •rMeniiat to the life of tuu»cte as the eue(^y>holdin;$ proteid
or carbon cciui{>ound ; and exticrimeniA made with regard to Auroe of iliem.
oalcie salts for insliince. sho» ilinl thrir iireMUceorabsenoe materially aflect*
the maintenance or reeti^>ratti>n of irritability. Souti; of tfaetc probnblv plsj
the pan only of securing by their presence fitvorable oomlitioM lor liie dnv
metabolic proceceee, somewhat after the way in which the itnaeiMM of raick
phosphate ddermiuoe the curdling of milk; but Mmc we protMhlr ttugltt
to rqpird as at-tually entering into the procenoe tbeiuwlvw^ 6f these
mattent, however, ne know very little.
S 5M. The end'ppxlucu of mu.-b^ular mvluholixin are, us ire have soeo.
carbonic iii'id, lai'iic acid, and krealin or some other nil^)geilous bodies, aod
we have already (J til) cuid all we liave lo say concerning the foruiatioD of
I
I
tb«a prodactt. Wc mar, honever. bnflr comiiler here ibe <|Ueetiou. What
I* Out rvUUuii nf llit«e varioiiH imtHliolii.- pr'x.'OMe* to ihe einiclural el<eni«l>li)
«f ihe liMueT Wbiii ir« uiv iliat Itie niuiM;u)nr tibr« is coiitiuualiy timW-
gahtg tneUbolisai do we mean that ererj jot aiul tiiile of tlie fibre is umler-
goinp rhaiip^ ami thai ai the taiuc rnle^ We can bardir siipptibe tbia. It
•Htii utiliki-lr, fur iii»tauce, that the metalxilisiii ot' the librillar sulMtmice ia
irWntirMl wilb that of the iuterlibrillar atibdtaiiue, whatever be tlie vien n-e
lakr a* to ibe ]>r<>|>ertia or nieaiiinf; of tbe two »ul)fiaiicM. Furlber, if w«
•rrq)t tbe *uggt*tiooa made id ^ >*7 iiv to n roi) tract He substance, whicb.
llnMiffh liaviog peculiur iiiiulitio'. I>eiii||; twc^uliarh- ri-Iatcd ti> and haviii^c
ppfuTiar (■onmclioua v'nh tbe rtvt of the fibre, ranr in a bmad nnv be miu-
]mtr4 with lh« gl^Mgcn "( a lieputic rell, wt- cau coi:cdve that ib^n cuiiti-nc-
dlr •uhctaiioe may 1<« niaiiulueturcd vithout tb<' nholc of it at Iniiit bavins
bam at aiiv tiiue, aii integnil {lart of what uc niav in a Mricler h'Iim: cnll tbo
ml Uviog HlbMauoe of the Bbrc. Wv >bniild tbii* 1>c led to regarxl lhi>
BiMabotie emnta oceurrintr iii diuri'Ic n» lulling into two cIa«M* at Ien»t:
lb>w Inking )>Iacc in ibr living Rinri* {x^niiiini'nt fninicn'orlc. awl thoM bear-
ing uu ibr t'iniitiiion and dtvlruc-tioii »f itic ctinirni-lile Mibrlaiice lodind in
Utat living rrnniowork. Fnrlher. it" we Mipiiii»c tbal the n)elabuli»in by
wbich the tniiM-les nuppl}' «o mnt-h of thr beat of the ImhIv, and which, n;> ne
hav« aeen. may nixl do<« go oii indepcitdenlly of i»n tract ions, i« not n roe-
ubolicni of th« mine contraclik siilwlaiiec iliHering from tbe mi-tabolism ot
a ei>ntraction in bejitg au onl<-red that all ihc etier^'T ^<i08 tint as licAt. none
bcin^ Moploved to eJAct a chance of form, but >!> a inelaljolisnt of »ome other
"UMCinoftiiic" HbetaDce, we Miould bnvc to add a third ebi«« to tbe other
two. These, of «>iin«, are at present lualterv uf spectitation ; but «« tbe
whole, ihe evidence we can gather leudii. anil perbajM iucrea«inf>ly len<K to
(iuiw that in muscle there doe^ exist such a framework of wbitl we may call
morv dUtincllv living au))«i8iice which ralej the btstological features of Uie
Bbra, and wbuae mstabolum ibouji-h high in quality doea not give riie to
BMMTff discharges of energy, and thai the interatioea, io to ajMak, of tfala
frwnewurk are uccupteil by various kindii of material related in dillbreiit
dtfcree* to the framework and tliereloiv ileoerviug to he vjioken of as more
i>r lew living, the chief part of the energy tei frve by muKcle coming directly
frun ihv melabnliam of aome or other of thi» material. And tbe Hime vKV
■lay b« oxteiKlcd to other titwutv^ Both the framework and ihi; iiilcrcalalLxl
naicriaJ umlergo metjibolism, uiid havr, in ditlercnl degrei^. ihcir aimlHilic
and kalalxdic dmiigiv-. both are coiicrrmil in the life of the living Hulmliince,
bat 0D« raon- directly than the ulhi-r, and thi.-< is what was nH-ant by tlte
larnn " directly " and " indirectly." iimil iu S 642. >fuch n luotb) of axprea-
■ioo arems prH«rablc to the morv common one, hns»l on the aiialofii' of a
Gnarm, of tbe muscle fibre firing oH'ihe ctintrnclilc material; in the tirtvirm
there arc no such ooiiDeclioiix iK-tween ibo machine and the churgus as obtain
in tbe living iDcrbanism. We may perhapa further bo led by this to dis-
tinpitsh IwtwoMi growtli a* bearing on ll»c framework, and more icmimtary
DUtritioii as beiinng on tbe accumulation and cxpetidiliin' of tlie lodged
oMUcnal. We may add tliat since some of the material so bxlgetl in the
fruMWork will consiel of rahstances which have not yet undergone melabo-
KaiD, biit are eitlier about to Im worked up into the Iramework Itself, or are
about to he transfomMjiJ in a more direct way into some product of niflabo-
Unn, or are substauecs whose presence is in sume way oeooaary for tbe carry-
tBgoa of metabolic proeeeaei in which they tlwtmelves take no boility part,
«« must recuxnize a continuity without any sharp break between ihic mate-
rial ubieb we rct;ard as ]iari of the tiieue, niiil the lym|>b which simply
bathe* the liwue and llotrs through the interstices. Hen<« such pbnuea as
658
NTTRITION,
"tiKiiv [irotctd" auil " llouiinx |)ix)te!<!." $ 5'J3, nrc umledinible if iber mn
uii<le»ton<l t» iinpl}- n »linr[i tiiic of (l«niarc»tti>u belnd-n thv " tJntUQ" and
tli« blood or lyuipti, iliotigh lucful a* lodicutiiig two dilTcrent line* or Jigiew
of tneuibulinm.
S MS. Ttic jinMluou of nitifculMr niotabotijiiii [kum into tW ly(ii[i)i Imlliing
tile lUiru iini] mi, ncht-r by a ilirci't piiih iiilo tlin cnitilluriov or l>y u nxin
circitiitoii* cotiRW ihmitjih ihc gdicml iyniphntic f^-'Wn. intu the blowL Thm
fitlcol'thc citrboiik' nejd wu bnvo tolly trvutml of iii dcmlitig wilb rapintioD;
tb« littlo wo know concerning tli« iiitragcnous pmduci or pr»i)iicta du bean
minted in dvxiltng with iirva; the third rcL-ognizcd proiluct i» Itctic xdi).
Mtrcotnctic ncid. Did iin|- con »ide ruble timoiitit of oxid«lioii InIcv plflcv in
the blood Blrcam while tbc bloud is tloniiig alon;; the larger c'hjinti«lf, lub-
ject only to the influence of the vaiciilar walls, we might fairlr expcd that
the lactic acid dischar^^ed from the musclee irould bcnbj«cted to oxidUioc
influfinces while still within the blood stream of ibe laryer cbaoDele. Wc
have, however, no saiJBtactory evidence of any laaic acid betpg oxidiied is
thb vBjr. On the contrary, there i^ a i.'ertatii amount of experimeDlal and
other evidence that lactie and pr(.<Mnt in ib« blood i* touteliow or other ^
posed of by the liver: niid that if the liver fail to d<> ita duly lactic acij
may aiipear in the iiriov. It i« tcmpliu}!: to nuppceu that it mifcht then by
a iyntJtctic etl'urt tw converted into tclv'-''^S^''> '"^ Iivlt ibiin utilizing aotn*
of the mUNcubu- wiutlv prodiirt, 1>ut thi; uxperimflnia) and other vvidioicc i)
all a^ni>t thi» view. In fniilt of iictiial knowledge ire are led to iofir (hat
it it in the Iiv«T nxidixpil int» carbonic acid and water, thu« aiMing its v»i>-
Iribuliun to the mipply of heal, or prrpiin^ in mdm way for oxtdaiiun elMv
whore. Probably «iich n changrt i« not confined to the liver, but inkt* place
in other orgaof fiich nn the spleen. Thns the kind nf uclioii on which we
dwelt in Ireitlin); of iircn, namely, that the pnidiicls of the nictaboltsni of
one iHfi&u are larrie'l ti> other orgaiu for further elaboration am] poariblr
uiiliMiiioii Hpplieii to ibe unu-uttr'i^nous as well as to (he Ditnigeooua pro-
ducts of mo«cular inelabuliitiu : and if a oiuacle nWet t'tx to other noo-oitio-
^enout prutlncta than carbonie and litctic iici<l tbeae are pmbably dtcpoaed of
in some such way as the lactic acid. In xpeakinx of ijlyco^eu in the winter
frog ($ 4(il ) we xaid that puwibly the gtycofrcn *•> «turcd up might ari.to from
sugar brought to the liver IVom i>ther liiwuei. If that 1m> w>, we ulitnild fur-
ther expect that «ome nt K'usl of that »ugar, eitlivr ax Kucb or at wmv allied
Biibntance, would conie from the »kclclal inuttclcn which form m largv a part
of the body of the frog; aiid if bo, wc niuct conclude that uiidttr the Hpvcial
circnni Glance* obtaining in Ihc winter l'r»g thu muselot diNcharge into the
blood a non-nitrogenouR product not iti the form either of carlionic or lactic
acid. It is {K-rhaps, howe\-cr. more probable that the migar in ipiei^lion coniM
from a metabolism of the fat slorod up in the " fatty bodies " and clmwboiv.
^ 646. As far ns we can sec at prroent the plan of nutrition tlius briefly
sketched out lor muscle holds good for the other liMiiee af well, the chief
or at least the most oanspiflUOtls ditlerencee bearing on the nature and ttrop-
erties of and ibe changflB undergone by the material formed by and held by
the more distiDctly structural fVamework. Thus the mnda oT the Balirary
mucous cell linds ila analogue either in the coolraciile subsilance itself, or
mora prtdiably in some early nitrogenous product of the exptr^ion of the
ooiiiractile xilntance. Rucb as may correspond to the myosin uf rigid tnusclo.
Tbc mctaliciiiiim of the hepatic cell seems, as we have seeu, to be eapMiolljr
vharacterixed bv ii^ returniMK to the blood a body, viz., «igar, still couiain*
ing a connideradle umt>uni of enftrgy', available for use iu other part* of the
bMly. Aii'l tbi» -Migijiiit.i the 'juetition whether iu the normal mvtabolisni of
muKolar vulutancL' a similar itomething. .itill holding a considerably qDnnlllf
I
4
OS SCTBITION IN GKKKRAL. 667
,. natr^j. tome prototi) •ubsliiiiea for inttiincp, may not be rctiirDnl tu the
blood : tn iliiii llR^ mtliibolltn] of diuwIc i* iiapc rfc<-l1y dcvcnlwil in MTine M
tint lb« r»iilt» nro carbonic and Ikclk' uvuii' ni)'] mi Btilrc<.-<lcnt of uim. It' V
tbJB bv so, ihni n)uscl«s may bo uf otber iimt to the body at liirgc than m
own c<>nira(-lile nmchincn. jiiM ns ihc liver han other ums than the produc-
lioa of bile. And the Bonie coti side rat ions tuny )m- appltcd to the other
tknta Hi well.
$ 547. Whether the chief proiluot of the iucia1)oli»Ri of any tissue be a
IMMcmI tfibelance, or n fat, ur a carbohydrate, proteid substanco ia the pivot, ■
B to fpcak, of the melaboltsm, and nitrogenous bodies ainays appear an ibe I
(irodlieta of oielabolism. Tbi# ia striliingly seen in the outrition of plants
vbere. as far as oiert; bulk or weifiht is concerned, the active nietabolianK
tisoe ii insi];nifi<-aDl compare)! with the mua of products of loetabolum
liMped up in the form of starch or celluloM or some allied carbobydrnie.
Tb* protoplasm <>f a vep.-tahle cell soon becouiea a mere Sim bearing a heavy
Inrmn ot hi-aprd np mt^iabolic prvdncta and eventually diaappeare; and
of that lllu only a tiart eorrenpondii to what we tpoke of aoore aa iho
lirinic framework of the musde. Yet that scanty proteid-built framework ]»
nor* or Ins directly concerned in tiie imxliK-tioti of the carbohydrate mato
liU and the Tartuua cocavertiun* whicii that material underj;ot«. FnXeid.
nitrogen, clian^x are pntaugleil with the carbon diangea; and since the
product* of uictabolii^in iu the plant are nut as in the animal cant out of the
organunn, but for th<r mtvl pari hea[icd up within it, n'C find ibc plant storing
op in pan*, whvrt- if they serve no uwful pnqiiisc they at leant do no bann,
nitnigtoous pnidiicts of ni«laboli»m, such as those knunn ns vegetable alka-
l<Hili,inany»J'nhich by their amide nature betray tlieir kinship to the animal
■tUwtnoua pnxluct area.
I 548. The mtr nl which in the adult, leaving asid<^ f'lr the present the
rial nutrition of the voung. nutrition is carried on. and the characlers of
nutrition, are dvpendent on a variety of circnnisiances^ Knch tissue has
of course a line of nutrition of its onn which circumstaaces may favor or
hinder but cannot chan^,^ in nature : the nutrition of the hepatic cell cannot
be altered to Uiat of the niuBcular fibre. The tame tiemi^ moreover, has En
different races and diireretii individuals specilic and individual characters of
nutrition; the tiesh of a dog is not the same a« that of a man. the muscle of
eiw man livw dtSerently fi^u that of anniher, the metaboliiiui |>er unit of
budv weight is, ■• ire have >)ceu. greater iu the suialler urganiam. and so on.
Wilbin the limits and subjevi to (lie cunditiiins, however, thus llxed by
race and pcrsunality, general mtlueiiem produce general variatiuna in niitri-
tiun. The rale of uuiritiun uf a tiwiie for instance is de|>eDdent on the food.
OB iho amount and nature of the IWI material brought to the tissue by the
blood. W* have seen that pmteiil t\«»i, in cnutrast to carbon food markedly
ittcrMMt the mMabolisDi of the hiNly. 8ince this increase tells not only on
ibo siUogCBOUO but also on the carlxMi metabolism (§ fi24}. it cannot be tho
mult of a mere luxus consuinntion of the protud food itw-lf ; and unUw ne
nppoM that the prvfcnce of the cicow of pnMeid material either in the ali-
aMOtarv canal, or while pawing through the capilUries of some organ auoh
w the liver, ads as a siimulur to some reflex nervous meehinerj throvgb
wboae action the metabolism of certain or of all the ti«sue» is hurried on, w«
tniut conclude that it is the direct aeiTss of proteid material to the tissues
themselvee which stirs them up to inerciieoil metabolic activity. That pro-
teid food should do this, aivd not carlxihydrale or I'at. seem» to be connected
with the fact just dwelt on that proteid natorial is the pivot of metabolism.
I 549. In the preceding chapters of tbts work we have had abundant evi-
litoce that the metabolism of tlie tissues is subject to the government of the
I
668
Xl-TBITION.
central nervoH* ttj-nlcm : the coninieiiiui of n muscle, the tecnbtrr actlvitjr
of ft glMul. tlir imrwuwil or ilimiiiUhivl tinvlucti'xi of heart nil itlfon] in-
bUdccs of iiervotis inipulKv nllt-clini; nidalioliitni. In must of tlictv itwtaDD*
tluj cbnngcs induced tnll oiihiti tli<- itowiiunnl. knUibnIic, phniw ninl bnv« ■
duwnward ehnrnottr; (hue, uhon ii tuuHclr contracts. Ihf mult i* ■ ntnnr-
«on of more com)>lrx Wlies into t^iniptcr bodies; nn<l ihe ■amr. kn fiir ■• «•
(-nil MC. is true iif mosl other mses. But it is open fur ue to siip|>ue(< tbu
Dcrroiif inipiilficfl might affecl thv upnnnl. nnnboltc. phue mxi haw n cud-
Btnictive iiiMuencc. There nre no rcHsons for regonliug nieh an aetioD a^
inipoe^ibic; and indeed some nfaenotncna. eueh aa llioae (^ inhibitory oenrei
and the antagoniHin between thofe and augmenlor nerves, poinlclly suntU
some such view. Thus, we may eupfwi^e that an inhibitorr impulse prodoMK
Mldt obaogca in the cardiac mtiscular eubslance lliM tbu upward coa«triicti<ro
lUDCnMs areawisted and the downward dtaj-uptive proceceee checked, wberebj-
the aettins free of energy ia checked, and no the tieala hiodered or Mopjvd.
the inhibitory elfeci being followed by a period of rebound in which Hi*
ravings of the inhibited period are spent in increased action. Coiti'enMir «•
ruay suppose that an anginentor impuUe hinders the anabolic, and nwist* tht
kuiabulic cliiuijjes, and conversely aUo, when it hua done its «i>rk, Itaivm tlir
tiiMue with diuiiiiiithed ciii>ital manifested bv feebler bi-atit ur hy the »h*rDe<
of the |>nHer to beat. Ami .lirtiilarJy in th« ca>e nf the rcupimiory eenin
and of other ti^ueii. Whi-ii we have to xtudy the nrij^iiiiition of vtsua) (»•
pul»«» in the rutiua we ahull mine upttu a view ihni a wave of light eaajr
atfect u'hni we tihall call a vimial Hulwtancc dthcr bv promiitinjz unabolie
ooitatructivc I'ban^tB or bv incn»i.iiug katabolic deftrucltve cluingM accord
iiig to it* wave lenp[h. 't'hen; is then evideuoo, to a certain extent, for the
view oD which we an; dwelliug: but. without discuHing the matter nay fur-
ther, we may Kay that the cum-e pi iun, though •uggvstivc, haa not yet bc«n
demonMralrd, and to lar i-4in unly be ipokem of as probable.
§ fiSO. One value pcrbnps of »iich n view lies in the fact that it warns us
against awuming that n nervous impulse can only produce disnii>tive kata-
bulic cbaiigc< »iich as are seen in muscular contracliou or in secretion. The
elTecte of alimulntinp a uerve going to a muacle or a salivary gland arc Mrik-
ingand obvious, and the behavior of a musclenra ghuid as far an cuatrmclioa
and secretion are concerned u, within certain limits, under experimt-nlal lyw-
trol. But there arc certain phenomena, seen chiefly ia the connM; of dtwaar.
and lying, to a very small extent only, withiu the control of ex [H-rinicnt.
which seem to show that the central nervous ^steoi govcntu tlic tncliibolic
cbBDga, the nutrition, not only uf muscle and gland, but of various other
tinun in a deejier and more general way than that of smply pnimoting Inr
hiitdering) eontroctJoD or aecreiioo. Thus. m> we have aeen (§ J13), when tbe
connection betireeu a muscle and the central nervous mteni ii Mvered, tlie
iDusole eventually wastes and lose* it* vitality ; when all the nerves going to
the submaxillary gland arc severeil, the gland, innti-ad of Wing, lu in the
nonual condition, tnterinitlingly artivi.- and quieiteent, pours forth u continn-
oun " paralytic " secretion and vvcotual ly dcgent^rat«» and wnslcs. When in
a nibbit tbi: iifih nerve U divide<) in the Hkull the lorn of sttnsation in tboae
parta of the face of which it in the Hens«)r}- nerve is followed by nutritin
ehanms. Very so<m, wiihin twentv-four houns the come* beootnee doudr;
and tnis is the precursor of an inflammution wbicli luav iavolve the wbiue
eye and rnd in its total disorganixalion. At the aaine time the naaal cbam-
oen ol the aide o|>eriited on are iutlaiiied, and very frequently nieen make
their npprarnm-e on the Hjw and gums. Ami similar n^»ulls hare been seen
in other animals, including man. If the i;|HYaiiun he conducted in a vouug
animal, which stibsc<]Ueotly liveii to maiurity. the head may become bilate-
OV NCTRITION' IK OENKRAL. 059
nltv UHjruoiutrwnUy. u shoiro MprvMilljr by tbc «kull. Again, division or
botn vigoi ncrvci is verir apt cu ite liitlowed by iiiflnroiiwtion of both lungs,
b; buy d«M<MnUiiHi uf itu) heart, luii) m li/ ilentli.
In MVtttnl of tbeM inttaucca tbc elfod n a, niixot) miu, nnd the nroblciu
a>isplieat«d. Thui, in Um ea«c of <Iivtfloii of ibe fifth tierv«, aoeiiig how
Miait*i»tniclurv lh« sty* Miimd bow carofullv it >* proUicMd byUi* mei-hu-
of tbo ej'elids >n<l Uuira.it aeenu rcasonnW to suppow tliiu Um inSani-
I
auion ia qittstion tnigbt siianly be tb« roault of tbe irrittition cauawl by
Jul «od oootaciwitli fotvigii t>ixli«», lo which thecyo, no lon^r (guided nnd
iMXttecUd byMOMidons, tiictv b«ing destroyed by tlie wclion of the nerve,
OMUie «ub}«ct. In (li« same irny the ulcere od the llpe and fpiiaa nii;;hl be
•xpluDed aa injuries ioflielcd by tho teeth on thnec striicturee ia their iiiscii-
•ibve condition. And aome obaervora maintain that the indunaatwn of tlie
«yt may he greatly leneoed or altogeiber proveuted if the orgaa be earefully
«aftna up. aud in nil ponible ways protected ftom the irritsdnji inllueuoM
of fbrti^ bodiea. Other ohservers. bove\-er. have foiled to prevent the in-
AamiiMtMa in spite of ererv care. So, also, the iDflaniDuitioa of the lungs
Ihltovinft upon division of both vu^ua nerve* aeeina to be due, not to any
direct nuiriiive action of tlie puluiounry braiichea of the vsgud un the pul-
Btaaarv tia&oe, but to food accu inula tiujc in tbe pharynx, owing to tbe parol-
yM of the ceaopbsguB and larynx, and then piuuinj^ into tlie air paiaagat.
and en sMtJng up iuUammalion. Death ia ihttie ciunn is, uoreorer, often tbe
dnpla nmnlt of inanition caiJ4u(I by tite parulyia of llie ossophaguit allowing
no li>od to rv«oh (he •toinaeli. The phi-nomi;iiii of (he paralytii! aucretiou of
Mitva an abo of a coinj>li('jklcd nntiiru.
Bnt, even without iiiauling on xnch inatanuta a» the above, various other
phsDunwoa of disMae seem to iudicaic :iuch an influence of ibe nervous
■nii-ni on niilritiim lui we nr<' di^-iiwing. As i^iamplnf wi; might meotiou
lae rapid and, peculiar dcgcniTntimi of and lonof cvnlraclility in ihu ■kvletAl
■nuwlea in oertain nlfcctiuns of the apinnl cord, the changes in tho iniuclos
beiog nure rapid and prof^iuud ilian in the nervea; the phenomeua of bed-
auns^ especially the so-called acute hedaores of cet«bi«l apoplexy ; soue at
least of the caaea of veeicnl afll?ctioua attendant on spinal or oereMal diaeans
••r injuftw; the more rapid atrophy and lossof contractility in nosoles which
MIow upon contaaioas of nerves aa compared wllh the e&ecia of aimpte sec-
tion of nerves: the occurrence of certain eruptiona. such ai lichen, zona,
eethyino. etc., in various spinal or cerebral diseases, aud indeed the (renerat
pbenomena, and especially the toiMgraphy of (he eruption, of a lan^e number
of oulaoeous diseases. Lastly, nut not leaat, we might ()uote the general
proeeas of inflammation. These are eiamplat of disordered nutrition. To
than wo niigfal add as ioslanoea of altered but yet orderly nutrition tbe
msMtrkable oonnections observed between ohanges in tbe form of tbe fiogera
and gn'Wifa »f tlie uuls aud Iiaira, aud oertain interual malailiee, such, lor
iuatancf. as the " clubbed tiugers " of pbthisieal and otiter patients, and the
hke. Wc luijilil aliti) call attention to the influence of light on the uutitiou
uf animals. TlkC uxiwriooce of Mind people aud bliutf animals indicates
soma special coonection iMiwtwa visual seaiadons and tlie nutrition of tbe
skin ; and this can hardly In other than a nervous oounection. The elfeeta
of prolonged darkncM on nuiritiou tn geiK-rul and tlw exjieri mental results
wbMth abow tluil the total wclabollam or the liodv is influencod by light, also
aiigjast some nervous action. Tbe iufluonca of oold again in dolenniotng
the ipvwth of hair points in the an'nc direction.
Jinking nvorr allonrAuce for tlw inicf^-cntioo of bcl in tho praductiiHi of
the i>lieiiiKnoita'i]uot«d alM>v« of such coiuin>ia aotEoos of the nervous system
as are already well known to us. such as vasomotor changes, making ever;
660
-VUTHITION.
nllownme for tlic coii!»c(iiii?iipi''' of tliD fiiilai* or bltmino* of i«n>Mtron »t
tbf nliwncr nf ilioiw bcm-tidnl nflcr mails of musciiliir mtivily which dv
potnk<I out ill S Sii, n,xHiKiii/.iiig morcoTrr lh»t cliiiii|n« in one (>f)t)in rm;
aflii-t tht oinilitinn of (ithpr [)i»t)inL nrgnns bv (-tmngvi jndiiccil in iIjc com-
Scwition nr ijualities of the blond, tliirri.- »lill ipmnins ■ rcridnr m birh xtiM .
wtiiK-'lly to nuiot (o the conchi^inn tlinl ihc intlueiirc of ihv ncrvoii« crEtciii
ia not limited toeiiih ohnngtv of th<- nntEcli^ ni belong to the prodadion of
contrnctionK or the generation of heal, but benrs on iW whofe nutrition of
the muecle. i^imilnr eonfi deration 8 lend us nlsn to conclude tliat lb«> infiu-
pnco of the nervoiu ^vMetn benre (in the whole DUlrilioD of the glsudf, oftbe
bloodTeMcls, of the «^in sod of the conDeelire tiGCue in general, in ftctoF
nearly the whole body.
Such an iiifluenee of the nervous eyMem hae often Iwen spoken aa "Iro-
phtc;" and the term hae of^eD been u«cd aa if the growth and nourvhiiwH
of a liKue weie the result of nervous action, or, at all ereuta, could not b
complete niihout the in terveotioQ of nervous impulsn. Hence, in ihia »ie»,
the c-on»e([u«ucea follow iuf{ umn section of tbc KAb nerressregaided osdut
to the falling away of "iTopIiic" influcnceii. Such a view ha*. howe%'er, w
sound ba^9. All biological Mudie» tmch tiii that the )(Ti>«ih. repur end
reproduction of livinc subslaiice may go on (juite inilejiriidcutlr of any neh
vous Bj'Blcm. The while l>lo(.d-corjiui>cte> so tbruu^ch ihi-ir cydrs uumotMd
by nerroue inipult>c«, and the niilntion of the nervous nvstem ilM'lf caaiwt
bedepc^niliiit iiu thenclion <if that xjNieninn iliclf. All tKnt is n'ally iiewled
to explain there phononiciia Ex an acceptniKi; of tbc view thnt n nervoM
impulse may modify ihi* nielaholic cvimiIh of other lovun* than mu»cl<-* awl
glnniin. mid may modify ihcm in various viayv ; and further, (bat the nulrilioo
of i-ach tiMue i» in thu rumiilex animal body »o arranged to nfv^. the con-
stantly recurring influruocH Drought to bear on it by th« Dcrvous eyatcn.
that, when those infiueuccs are permanently withdrawn, it is ibrown out of
ctiuilibrium ; its molecular procesect>, so lorncak, then run loose, since the bit
hoN been removed from their mouths. And as our knowledf^ of metabolic
processes on the one hand and of the actions of tbe nervous systeni un tbe
other hand increases, ihne snppoeilions become more and wore reasoaable.
Oy Diet.
fftSl. An OTtlinary man living an otdinary life will need for tbe tnaio-
teuunce of vigorous health a ci^rtaiu amount of food of* certain kind ; this
we may take as a normal dier
Preaumiug that the experience of mun has led him to adopt what is cond
for biro, we may aRrertain apjiroximalely iho normal diet by mean* of the
statistical method, by cxamitiing the nature and amount of the daily food of
a very large number of individuals. The most valuable data for this ptirptce
are iIicm: gained by inqtiirin among pertons who choc«c their own food ; (be
mnll* gamed frem Ihc diels used in prisons or other ineiilutions, or uaong
bc(li4'» of men such as the army, though more leadily arrived si. are open to
the Direction thnt the diets in queslioti are determined in part by (be theo-
retical opinions of those whoso duly it is to fix the diet. Pnlting together
lh« various statistical reeults llius obtained, and selecting tbe quantities
which Fcim to be most commonly used rather than attempting to strike a
btrict average or take a strict mean, wo Bud that in an ortiinary diet for the
Inenlv-four boun the several food-itu^an-: —
I
Tin toul (itvtuUblo) pottntjnl owrgv or the lom-r n:iniit« u 2610, of the
h\^ttt3i0!» (kilogiuoinMMlcgrec) cnlorim, cnlciiUled, io rixmi] numbur*, on
Um i]«U <if ^-ViS, With «jch it aUtistiwl diet nc may compAn: mi ozperi*
fBtntnl A'lH. thill u to mj a tlivt arrivci) at ihrmigh n aonee of trials on nn
in«l!vi(fonl iimii whose body might be taken to bo an nvcrago one. that iliot
briu); fimitidvred a normal one in which the body, maiDUioio;; visorwu
btmltfa, ndtbcr ^inol nor lutt id wvight, and reouioed, moreover, in iiilro-
niioaav<i|iiilibniim wiihthi- nitrogen of the egwta equal to that of the ingcsla.
To make sure that under HUch n diet iho boily vas remainin;; of thi> same
composition, there itught to be evidence of a earboD ci(iiilibriu(n abo,
oUurwiw during the period of the exfieriiuent fat mi^hl ba roptaoet) by
wtUa- (mx f 'y£'2) ; but this is uulikely, aud we raay therefore aooept thie
DMihod as a fair one. It baa ffivea in the handa of two difTurent obeerren
the folltiwiut; tuoiewhat dilTi^rviU Ksulia, the diet A beioe that alreodv qiio(«d
A U
Prouidii . • lOOgramniM t18
K»U . . "lOO " 66
Cubohrdntci V40 " MX)
Sail* as " —
Water 2600 '■ —
Tli« total I available) pot«otia] enenry b reepectfully 2:Il<>aml 34)25 caloriea.
(In the whole, the dieU gained by tlie two method* iitfr«e very lur^Iy.
To put doKu a single column of figures as " Ok normal diet " would be to
uSxi » VMU and tulunve accuroeA-. Jf we deure, for thcoretleal purpuoea,
taarl«ci moio one aetof figurw rather than others, w«mi>;hi bu influenced by
th* coDsidentions that the lower amount of pn)t«ids In the ex [leri mental diet
wa* nearer tin- mark than the bij^h^r ojnount of some of the atatliKical dicla,
and farther that, wberv omI in not of moment the inih»iiiuii»ti of fat for on
•sens of cufaoltydnum In dowinible. Wc should he thun \cd t«i take the
txperimenial diet A a* «ii iImi whole the best or most " normal " one, and that
■I the uni: which nccmplo]<-d in the caloulationsoTS''''-^*- It will Ixiobncrvcd
that the potential enetuy of thU diet i* Um than that of any of the others,
am), as we said while then xpcaking of it, may bo con»ider«d low : but tbors
was no evidence thai it was insuHicicnt. Still, it miiat be rememboreil that
Dthber it DOT nnr uf tbo ixhc^ni is to be regarded iw divtinctty proved to be
the m>l normal diet. Against the eiperimi^ulal diet we mny ur^o that the
number of ex|>eriment3 have been ftw, and conducted on a few individuals
only at most, and that a larger number of etperimeais. with a variety of
eombinations of diflercnl amounts of the several food »tuflii. might lewd to a
di&rvnt f«snlt; tliat for iiislanoe, with certain ani'junta of fats and carbo-
Iiydratee, the atnouni of proteid need»I to mainiain hcalihy bodily e<|uili-
tiriuRi, including ntlrogenoUH e<|Uilibrium, might be reduced much below the
Inn grammea, especially if particular kinds of proteid^ fat or carbohydrates
were used, and Mitecial aiieniion (see $527) were paid U> the aalta. And,
indeed, a considernble number of oboervalwna haw been made tending to
rbriw thai a roan of average siae and weight may ctmtinue in nilrogenoiu
i
NV TUITION.
equilibrium bdcI in good ht-nltli wiih a dnilv mitriit nf miirh lew thnn
emnmcs protcid, wiili ii# littlo m 40 grnrixnco fi>r vximiplr. To tfii»
ihnll have to refer in cjirnkiiig of n vrgHiibJo dit-t. Apiinvl llu- prniiait
<licl, on the other hiind, we mity urge ihiil instind is nut nii unerring gut
and that the elioke of a diet is detcrmintrtl by maoy Other cimitneuince» thiin
the phveioloLnen I value of the food.
$ BSii!. TnKing, however, oonio «tich diet ab the above (o be tlie nppmsi-
mitel; Irue Dormal diet, v.-« miiy cidl iiliention lo the fnet that the uonital ^
diet it made up of eitch of the three great Ibotl-stu^, carbohydrates being ia,^
escefs. We may here remark incidcti tally thai l)i« diets of both the car~-.
iiivora niid herbivora ngree with that of omnivora in eoniainiDg all lhre«
food'»tul& : they difler froni each other ss to the relative pniporlions ouly.
Ai we have seen, the hoiiy may be mninlaiQed in ei)uilibriuin on proteid (mm/
aloue; but an eiclusively proieid diet is pot only bou){ht dearly in (In
ID ark el, but also [laid fur dearly within ihe economy; we are, of courw.
Duw speaking of man. To obtain the necesaary carbon oul of the carbni
nioiely of pruieiduniiecEi'sary labor is thrown on tbeeconomy, and the tntcm
tend* to bet-onie blocked wiih the aniiiles ami other uitroscnous waMe arittng
out of Ibe nitrogen moiety simply thrown olF U> secure the carbon.
Fata and carbohydrnie« arc mneh more akin to each other than b ■.•ither
to i>roteid ; and if. »n the one bauil. as (§ Hi'.i) leenia pdMibk- or even prob-
able, the fat of the tbod and of the l>ody i* L-onverled Iniu Migar either un
it« way to hei-imie bnilt up into the tisttue, or in the <H>urtt- uf tin- chnitgM
taking p1ac<- uutnide ihc reul living framework of the tiMue by ivhich it ti
rcdtieril t<> cirlionic acid, ami that, ou the other hand, enrhobydnitn <^D
fiirnioh the- fat whono prt^w-nw in the body ii nccMWry, we might ex|>cct that
enrbohydrnlo nWe without fat might with pmteid form a nonnnl diet. Itiit
on this point exiurience i.-> probably to be Inist^'d ; and wc niny iiilvr thai tn
every normal diet somu I'at at lewt munt be added ki ihu starches and the
mgara.
The advantage of this mixture is probably felt while ibe food ia as ret
within the alimentary eaiinl. What we hari; learned eonceniing digeotmn
leads UB to reganl it ns a coinpliented procew, and we cannot readily iiiiauiDe
that the proteolytic, aniylolvlic, and adipolytio changes nin their several
flOlinea, especial Iv in ibeaiuallaiid large inlcMtine, apart from and irrai{ie<-iire
of eacfa other. We are rather led to etippoee that the aceompaoimL^ni nf «»«
Bet of cliangetf, in some indirect manner, favors the others ; and it is for tiial
reason probably tliat we take our food mulb not separati^ly, but mixed in
tbe same meal, often on the swoe plate, and even m the aame miiuibful.
But apart fmm this the two food-stutl'e, faiH, and carbohydratea iuiim play
different i)art» in the ecunomy. ao that the one r-aunot be wholly subMilutetJ
for lite other; nml though, beyond the fact tluil the one soeins to be a sourm
of energy and iho other not, we do nut ati yet know the true phrviolngical
func-tJon of the hydrogen of the fat as conifiarcd with that of tn« ditll-renily
ditpfwed hydrogen of Ine carlmhydraie, we may pcrlui]» infer that the diHi'r*
tacD i>r use within the body of thu two kiiidi> of tiiod-Htuflii bean n<bt *>i mudi
on iboir ultimate couxumplion to supply envrg}', aa od llie vnriMia c>>nipliaitrd
prooesM* which they tindvrgo and nrrangrmcnis in which they tiike {uul
ueforc the end of their work in reach^-d. We have had a bint that (be
carhohydraic more rapidly Mipplies iho beat-giving mi-laholi«ni than does
the fat : and lhi» suggnts an ndvanlago to the economy in receiving daily B
certain pinion of the more tanly material, while at the sarao time il may
be taken lo mean that the tat before it is nM-^l to give rise lo energy has lir>l
lo be coiiverteil into tiugar. and so takes more time in itfi work.
Tire main carbubydrale of every <liet k starch, and as far aa we con leara
ox UIBT.
608
I
■t foarat. tli« »lnM) vhich is to lnrg« ii pnrt of tbu crrritU am) vegelnblcs
nuaiBCit by man is Ihv mrid boily in all nfthcm: tor the ii»c of lurh bixlif*
M iDnlin t* ra inwgniticont ilint it may \k nc-el«ct«!. Jlnii, howi^var, cou-
HimM DO i]>ctinsi<lera)>Ic ijiianiiiy of suesr, chiotly cftne sugar. Since tbe
•tarch of a meal does not become available for the economy until it has been
eonvtrtei) into sugar, we lui^hc be inelined to infer that it iras a nistler of
lixliAercDce ulictber Uie earbohy<lrBte at' b diet were supplied ns starcb or
asHii^r. But besides tJw fact that anr \artfe deAeit of starcb in a diet
might seriously interfere nith the general couree of diKe^tiou, eepeoially if aa
ur^d above the wreral digeslive proctnea are more ur leas oepeadent on
«adi otber. it muitt be remi-mbered Ihnt ihe au^ar into vrhich starch is
diBDsvd bj ditcestioi) i* muhotie, while cane suKar appears to he either
ibsoned >b cane »u(:Br or at moot only inverted. Miire>)ver. if our labora-
lorr experiments truly rejimeiit the dig««ti<)ii tnkin^ f>tac« iu the living
body, ooly part of the slarcb, S 198, la eliangod into maltitfe, irbile pari
bccoDwa mote variety <if dextrine or of starch. Our kiionledfce of HUfcare
aad of tbdr fate in the eoooomy I* too impi^rfoct for us to be able l» aiate
Ike eflecM on the Uxly of dignted Mnrcli as cnrnmrcd wilh tlimte of ease
•ugnr or millc sugar ; but tlint tlio»<> are or may bo aifforciit io shown br the
exfMirictMM of luedicnl prii4.-t)(v. In many <-!utct the total effect on tlielioilv
of a diet fnim which uini; sugar is as mueh n« po«<ibl? rliminiiK^l. ihough
atarch be allonvil, is rer^- diHerent from that ol <ino of which c<u»e sugar
JbmM an appreciable |>art.
Coaoeniitig cellnlii»e. which in ber4>ivora Bpi)«ars certAinlr to terv«asa
Mara of encrey and lo he a real foixl-stulf. otir knowledge will not allow Ui
to decide whether it has any !>peciiil iitan of its onn. or vfaelhcr the bo<ly is
eimply led to nlilize and make ihe best of what is a neccemry acci>m|iani-
ment <>{ the starch of vegetable food.
Cunoeniiiig tbe salt« prtv^i-ul in a diet, we need onlv re|»eiit irhat was said
io S 527, that tlieae. th<iuj;li iiifurcliug of ibemaelv<» little or do energy, are
aaegeeotial a part of a din n» tbe enervT-givinn food -stutb. inasmuch aa they
in sun>e way or other direct mrlab>>)Um and the diriiribution of energy.
And this is true not ouIt of llic innrjcimii- Milincf, Mich aa chlorides and
pbiapbatea. but aba of \bt so-calliil extrnctivcu. M ve have toea, the
pmence of ilxve bodin. both ihe oimplt-r inorganic and ilie more comnlex
organic snlu. in the blu»d or in the cztni-TBseular juim or lymph of titc
linuet i« owvntinl to or diivcls or nimlifiu the metabolic aetii-iir of the tvr-
oral ti«u«». The iM-ncHeiitl vtRs.'t>, as compoDents of »|xt'ial iVwtf, of such
ihinn as beef ten and intat extract, whkn consiit chii-Hy nf nails and ex-
IfaeuTA, with » very sninll Qiiiintitr of albumose or ulht'r forms nf priHetd,
and the rlTecIs either betietkiol or cfeleteriou* of drug:*, both turn in cominoD
Dpiiii their taking a pnrt of aome kind or other in. it may be upon, their in-
(•rfcrvnee wilh nieUiboIio procenea Tlie salts and cxttuctivc of a diet may
be looked iii»ii aa necessary daily medicioes. an<l a medicine as a mure or
lev exlnortf inary variittion in these elemeota of a diet.
Alcohol, to the use of nbicli aa a eompoocnt of an ordinary diet spedal
iatemt fur various reaaons attaches, (Monies in tliia class. For tbnngh ob-
■erraiiuiu show that the greater part of n nvnderate dose of alcohol i* oxi-
tlbcd within the body, iin<l mi lem-^ ai a 30^.irce of ener^v, man \im reeourve
to alcohol n»i far the ininuic qituuiity of i-i)«r)Ey whieh is supplied by itself,
but f»r its jHrniTfiil inlluence on the distriboiion of the ener^ fliruitbed by
other things. That inllucnee is a vcr%- complex iioc an<i cannot be fully di«>
mannd here. It is ■taii.il that imHleraia or amall doaca of aludiol diiuiiiiah
the consumption of oxyctn ami pnnluction of carbonic acid, that is to aay,
diminish tbe total rtsult of the nw-ulMilism of the boily, irbtle largvr biK
6S4
IfUTRITlOX.
Mill not intoxicating liwr* have n cxniimrr ellWit ukd inerWM tba
in«tnboli«iu. Bill Mich n »uit«iiiciu iitlbnis no nund bwit for Rn^eoDcli
lU li> th(> g«DoniI physiological etlvct of ulcohot, or lU Ui iU uwifulntM
pnrt nf All ordinsrv <tiet ; it doos not justilS' such a ci>nolii«ioi), for f Kiimfik,
U thnt ulcuholic driiike, tnkvu in nioiiermtioii. liy ilimini'hiii;; mvUtbiiliiiB
econoRiiz* lb« rcaaurcc^ of the body. Tile promiucnt i>hyi;i->Ii>giciil prohlvm
of dietetics is oot either to increiuc or diminish the tiM>lii6'>1tHin of llio body,
but to direct thnl nielnbolisni into proper chauoels: and whelbpr in each
particular case a ^i^^Q <l*'s^ of alcohol gives a n;;h( or a wrong turn lo tW
pbysMJogical procawcs of the body, depeiidi on the particubir circutDBtaooc^
of the caae. For the action of all these biidies of which ire are now speak-
ing, in contrast with the aclious of the food-slulTii proper, ia not only comptex
but variable; do cuniplex and variable that simple ex|)erience is at nreaear
a nion^ truHtworthy ftuide than spoculative physiology. We may odd tlitl
thu physiological action of alcoholic drinks is still further complicated bf
tbe hiut, lliat niinit ^iich driakit cuntain. liesiilea etbylic alcohol, various oUttr
allied HiibMt«iict«. whoie iictiou i^ ovt-n more potent than lliat of the cthyliv
alcohol itioir, anil wIiiik- pr^ciico very ninrke<lly dvierniiiiei the total eOcct
of tbi: drink, t^uch arl.icli^ of diet oh leu and oolfee txaiid upon mry mack
the same foiitiii); ELI ulcohid.
The qniintiLy of Moid which a man drinks or iihonld drink daily, or mar*
eorreotly the ([unnlily of water which be shmiUl daily add to the dry xjlidl
of hU diet, niuirt var^ widely acconliiig to clrcuiastttnoi!. It will differ ac-
cording w he is penpirin^ ijrcatly or not, according to the nature of the dry
solids of the diet, whether Iiir^cly ciirhi]hydnit« or not. and so on. A lower
limit, below which excretion is impelled, and a higher limit, nbore whicb
digestion and metaboHsin are injuriously atfoctod, probably exist; but we
have OS yet no adequate data which will enable us to Ox cither of then.
§ 65$. Ill the selection of articles of food to supply the food-etuH* and
other cousiituents of a normal diet, regard must, of course, he bail in the
first place to the amount of potential enertiy present in the maUrial. The
articled chowu f»r the daily lore must contain between them so much pro-
idd, fat, and carbobydraie repreunting so much av-ailable energy. But it
b no IcM iniporLant to secure tnal the energy potential in llie material should
be really available for the econoiny. Tht^ uniierial niu-it have such •lualilJM
that it in digested within the BUmt.-niary canal, and further that ila digMtJOB
and alworption do not give rise to iniuble either in thi; nlimeniary auaal or
in tluil socondarv digestion ciirrieil on bv means of the various metabolic
OWnta which we nave diiicu^'wd in prermiiinit seclioDN. A nuilly nutritious
(ubetanoo is one which not only coutaliii in itself an luleiiiiulp supply of
mvgy, but is of such a nature that its nui^rgy can W approDriatcd by the
•coBomr witb ease, or iit least with a» little Ironbto as |>os*i»le. We hare
approximate data lor (ktermlning how tisr an estimate of the relative tise-
fiiUMM of various articles of food must be corrected, bv allowing for tbe pro-
portioa of each which after au ordinary meal merely panes through the
alimentary canal, and the energy of which is not in aov way available for
the hodv's use. Thus, a niimoor of oluervations <»rrie<l out on healthy in-
dividuals gave, in (he cose of the following articles of food, the following
figures M the perceniagc. rockoncd in each case on dry material, wfaien
0oald be recovered from the leoee, and was, tlierefore, not digeeted and not
used by the body: Meat, 0 per cent; e^gs, 5 per cenL; milk, '■* per cent-:
bread (white). 4 per cent.: Mack brendr !•> per cent. ; rice, 4 per cenL;
moccaroni. 4 per oent.; maice, 7 pei' cent.: pens, !■ per ceul. ; potat*i«s, 11
per cent. It miisi, however, be remembci^il (hat the nclual correction to bt
made in any cose will depend on the mode ol' cooking of the lusiorial. ui
ox blKT.
Ill* dtknvUr of tbe iumI of which it forms jiiirt, tuul on the iitdiviiJiiiil
ofabiliUw of tho cvDiumcr, the lutlcr too v*rying under diffeTieat circuiii-
tfUIOM.
Thf nbovo nfen (d what nuty be atll«] roticb dij^eetibility, but bcniile* th»
\h«n »re other circuButaaoee tii be owwdefed. The mms food-fltud' iii two
mrticln of food, ihoogh aettiallr di)Nflted, (hat U to say taken up br ihn
Mlimeotan,- canal, loay, ev«0 while still within the alimeDtary canal, undcr^
cbang« in th« one case diBerinK from those in tbe other. A proteid may for
iiMM>c« in oi>e caae tend to be ontireljr converted into peptone, or to brvak
up into leucin. etc., or in other caaea to undej)[u other ohsn^ ; and a carbo-
bfdnite niay in one case be absorbed lu malltue, and in another give rise to
Uetie acid. Indeed, when we speak uf iIil- di^teHibility or the iudtj^tibility
of thb or that article of food, we do not in uuuij cum* so much mean tlie
(elatire amouul of the sulutauce taken up in some way or olher by tlw
lUoMBtary canal, a^ the chamcler* advantageous or otherwise of the cbaoj;^
whlsb it undeixoei in Wini; so taken up.
Hvace the purely fhomioal staiument of the amnunl of pDt^niiul euerKy
imaent in an article of food is DO snf<^ guide <>f ihr jihyiiioliigiciil vulue of the
MbsCanoc. A chunk of diecac viand* very high »n, ^norally at tbe top of,
• table uf the nutritive value of articUsi of food drawn up on exclusively
obtaUcal principUv, according; t>i the nnits of sne^y proscnt in n unit uf the
material: but it ■• verv low down in n corrMpondinf phyiiologicat table.
And similarly a dtsli ut* old p^nn ha« a rery diHcrent pnmologioal funi-tion
fr»ai a pUl« of frosli m«Bt, even whan both cootaio the aame amount of
oltrageii.
In thus romcting for dicestion the nutritive value of a <Ii«t it latist also
he boriM- in mind that the alimentary caonl, while chieHy a receptive organ,
is aim to aofDQ extent, § '265, an oscretory or;;au ; a free paaMjte through the
oueI is needed not anly for carry in^c o!l' u ndij^rdited mnlier but also for get-
tiiiit rid of excretexl matter : and the prexeui-e nf the former, up to certain
limits, asaista tli<? dindiiirKu of tbe latter. Were it pussihle to prepare a diet
every jot ooi) title of which oonld l>e dijjraiieil and ab!iorl>eil, llie itie of such
a (li«t would pmbubly bring about diiior<lor iu the economy, tbroUKb the
absence >•( a vuliicieutly rapid dischargu af tbe matters exorctod into the
alimentary canal. Hence cellulose and Tike anlMtaiices, even whm unuitUxed
thiou^ absonitloa, are not without tlioir use, and experienoc shows thai
dicastson may oc promoted by eating undigesiiblc tilings.
i 95i. Tbe several fbod^tufls of a diitt may be drawn from the aniiuitl or
from the \*«getabl« kin^om. Vc^table proteiils appear to undergo tbe
MOM clianees in the aliotentarv canal as do animal proteid*, and the main
•Ifecta on the body of protdds Irom the two sourrae eecm to bo the sane.
Oar knowledge at pnMeot, however, is too imperfect to enable us to decide
whether tbe fuDctioois of the two are exactly tli«8ame, whether the bod^
beharos exactly tlte aame upon a diet in which the proteids are exclusively
of T^fetable origin, as tip-in a did in which, otbcrwi*e tho same, tbe protmda
an paitl^ of animal origin also. Nor have we much b>iter knowledge of
ibe relative nutritive value of vegeuble and animal fats. And as we nara
aln-ady sai'l, we poseeas little or no exact knowledge as to (he part pbiyed br
Ibow extnctiTes to reipect to tbe aia[>uat and nature of which animni food
•triklngly diSbn from resetable food. In attempting, therefore, u Juditmunt
ftnn a pnrelr physioto^cal point of view as to the value of an exclusively
TCKMarian dfel compared witn a diet of both animal and vegetable orii;in. we
CM do little more at prosent than imjuire whether the former nuppli&i the
Mvenil food-stuiEi in adeauate i|uaulity. in proper ptuporlion, and in suab a
form as to be econuratcally ulilixed hy the body.
6A6
NL-TBITION.
Th« careful oxKiainiilioH during llirrc KDArair pi;nu«ls I'f Mvrral dan
each of ihv ineMtn nnH <^vtn of n tnnn. 'iS venr* olil. wri-i^hing AT kilok,
ivho line! for tnreo voir' livr<l on hd oxcIuKtvAlf vcgelitlile Oivt, vii., br««4.
fmil, nml uil, gnvo t))c fullowing mulu :
The (Inily did cunsisled on ihe »\trtige o( 719 grmi. eolid uwtlar aoil
1084 grms. water. It conlained
ntitmda
FntH
Carbdtiydratcn
(Ci-Iliilose) .
5i fCTamtnn ciuilainiiif S.4 N.
'2i ):niiiinit«.
057 g''i""i"n (alxKil } >ugsr and } ttUrcli).
16 gramines.
Tbe dnily fccct n«ighed, when fT«sh. 333 gnns. contnining 75 grtn*. »>lid
niniter, nixl were therefore both bulkr nnd watery. There werr ftrcMnl In
the tvctf, fnt 7 grnie., stnrch 17 grms., cellulose 9 grim., showing lIiBt 30
Iwr cent, of the tat. B per cent, of the starch, and 56 per cent, of I he cciluliM
md not boen utilized by ihe body. The eubject hnd really livetl on fm, 13
gnaa,, cnrbohydrntes ^40 grins, (and cclhilose T gmiB.). The fecea oonioimd
DO 1e«i Ibnii 3.4C nitrogen. If we reekon the whole «f (his hb prolcid. ibu
would giro 2'J ^tiiis. of undi^ealed proteid, so that thei'e ha* be«n ■ u'n»lc of
41 per cenl. of the proleids. lenviDt.' only 3'2 gnns. available lor real u«e is
tluibody; and indeed a very amnll (jorlion only of iht! nitrot;eu can be
rrgnrdnl as reallr discbargetl fmtrt the body itaelf. The loial stdid* of ibe
few* nntrt he reckoned a» portly excreta but cbn?Hy umliKealod food. If we
rt'i^rd tbe 75 anva. of »olid (eca as entirely undi^'atcd food. Ibe nbole aoUd
food available for the body niuH be reduced from 719 ^rins. to 6-t J crroa.
The urine of tbe day oontniDcd 5.38 gnnti. nitrogen ; this added to the -1.46
grms. nitrogcD iii tbe feces gives 8.79 grnn. ottrogen in tbe total egata as cmo-
pared with tbe 8,4 grms. nitmgenof the fiwd, indicating ••llghtTocs ofDiiro-
Sejiuun material from the body : but if wo «iip|i(wc that all the nitrogra in
le feoea was not in (he form of undignlcd fnixl we may neglvct this: and
Indeed the Nubjnct of the obMrvatioD woe in apparently ginxl bcnlth ami
stationary weight.
Com|iaird with either of the iinrnml diets given in ^ 651, the aboTO diet b
Striking for the low amount of proteids and of fat* and the relative escem of
airlmh yd rates. But thon^li such a diet may be fnken a» perhnji* fnirly
typical of the daily food of a rigid vegetarian, a much more richly iiroteiil
diet may be obtained from sources slill strictly vegetable. Thu« ihi: diei,
eotiroly vegetable in nature, nf an average 'lapaneae lalxirer of about ibc
BRmc weight as tbe individiud whose data we liare just given lias been esti-
mated to consist of proteids 102 grm.*., fat 17 grma.. earbonydmiea &7Jt frtat.
Anil the dii-t of a H'Umriiiiaii iK-uiant. liriug cbieflv on beans aud uoiie
with the addition of fat i^if i>oiiie kind, hnn been calculated to flirnitib no lea*
than protei4ls 182 grais.. fnt it3 grnix., en rhohyd rates 968 gnus. : btit the ml
nutritivo value of such u diet roust necil verv large correction indeed.
<y. 5 553.
The ctaminatinn of the diet of an indivitiual living with n fair nttmgviinus
equilibrium and nppareiitly good health on a moflitSed vegetable diet — that
la to aoy. oao whicb includwfmilk and cgg»— gave tbe following r I'nHetdt^
74 grms. ; fat. 58 grms. ; carbuhydrat'*, 4yi( grms. — a dkt which diHere from
lliu normal diet almost solely in the Icwer amount of proteide, one-third of
which, by the by, was supplied by the animal mHterial, ^gs and milk. In
another inslanoe, nitrogenous equilibrium and fairly good health were
secured, for some weeks at all ereitla, oa a vegetable diet yielding probrids,
•bout 100 grms.; fats. 70 grms,; curbohyd rates, 400 grms.; mit in tl'
nMrl^ th« whole of the Cut wiu furuiilied l>y the ntiimnt pruduci bailer, lud
IJebie'a extract wu (VmIjt uied.
Cuniloiitg ourvelvet, huwcver, to the more Mtrictty vcgelnrian diet, vre rany
toadod* in the fint place that, imlcM the ilnilv rcHxl be very large in amount,
the |woteid elemeot of nich n dift fnll> can»i<Wnihly boluw tli« lOO or more
gnm. pvcn in the normal diet. But n-e onniiot nuthoritativelr My that
Meb K radncLion is nccMMirily nn «ril ; tor, n# wc stated nbo^'e (s 651;, flur
knowledge will not nt prcwnl permit iis to make an autborilnti%'e exact
•lalement M to the extent to whicli the pri>lH<) cntir he trilneed without dls-
adrantsge to the bwly when nccompfliiicil by n<)e<|iiaie provision if (be other
eleiornt* i>f fond : nn^ this slatement holds (^>od whether the lio<ly bo nitder*
taking n rnittll or liirgc anionnt of labnr. A eecnrnl tVfltiire of »ueh n diet i*
the marked rv<luetion of the fat ntirl its re jiU cement hy ciirb«)hydnitee.
Although here aeain we ennnot make a dislinclly aullioriinlive stntemeiit,
Ike rvidenee wbien we posseea bears elenrtv in the direclion ihnt stieh ii
PtdiK-tino )» a marked diaadvantafce. A thiri^ and very charaeceristic feature
of the Blrictly vegetarian diet ia the relalivelv larRe aniounl of undi]>e«ted
Ihod lu«t to ilie body nud discharjjed na feces, feven when the diet is seamy,
M that the proietd element U low. tlie amount of fee» reliittrcly to Ihe total
feod it high; and when a more normal proteid contribution b secured by
smple HMftblhe fbce» liecomeM exceedin;;ly voluminous. Indeed when, lenv-
itt|> man, we compare the lierbivonma with the carniroronji mamma), we And
that the former a almittt its t-learly dbtiuguivhei) from the hitter by its fre-
quent and abundant feet-* ■■!■ by the nnniomical fcaturon of it^ nrjca nidation.
We have already nryet) that. *inee the fceei *ervea a* a meann ul excretion
of the real voMe nrmliietN of metabolism, a certain amount of vehicle to earry
tbew away U of n<lvantaK« or even neccfmr^-; but there ara no fndH at
pnaant known in iiit which show that the lai^r in[<«t.itjnl current of the
partly vtcelable diet etfeetf any such good ns csn eompcneslo for the obrious
waiM of labor iiieurred in its lmnsp>irl nnd mana^menl. to My nothing of
the opportnntties of mischief olfen.-d by n inan of material more subject in
the dominion of forci^m or^nntf^ms than even to thai "f the l>udy iiself,
ihnugh these op|>orl unities nre les^ thiin with a correspimding mass of aniimil
erigiiL With rN|iect to these three feaiuies, then, the Mrictly vcf^etnrian
diet aeems, on physiolo;:tcal j^nxind*. inferior lu one of a mixed nature.
There are, as we said, other aspeeis. «til) of a strictly physiolnjiieal kind, to
be considered, such as the relative dij^estibilitir of vegetable articles of food,
ibe relative metabolic value of the foo<l-«tufl!t of ve|[etable ori|nn, am) the
inAuence of aninuil extroctivee; but any fuller dbcunion of Uiese points
•ould be wit of pliice l»re,
S 566. We have treated the diet discnased nbnre as a norma) diet, suitable
Ibr tnan under ordinary or seneml cirounistnucet. Ought such a diet to
t>e modified for the rarioua exigendea of life, such as labor, age, climate, and
the like?
Vt'c •hall disctm (be influence of age in the concluding portlona of ihb
work.
Wo mnv be inrlittc), at fimt ^i^'bl, l» niMume that the toinl nmmiiit of the
diet *li»uh] xarv with tlH> wi-i^lii, that is. tbe >'no, of tin' individual: and.
indred, in discvMions nn nutrition, statonicnts concerning nH'lal">tiii>ii and
amount of food are often given in terms of |ht kili> of twly weijchl. In a
broad Mnse. it may he tnte that n •ninll man needs Ina food tliiin a Iarg«
OM : but it must be remeniberrNi that, ax wc i>nw in s|>caking of nninial heat.
Iho amaller or^nism, having the rrhitirely larger surface, cnrri<-s on ■ more
rapid melaI)oltBm per unit ol hoity ucight, and so ntmla rclalivelv more food.
And, moreover, the influence of site is prulaibly far lest Ihan tlie influenco
NUTBITIOS.
6Xert«d l>f Ibe iiiliDni imlividunl cliitniolers i>f tli« i>r({nQi«n, pivinf; riwln
what we may call the ponoiinl i>>]iiaiiou of tnet«1*o)ii>Ri. The saialler mi^u-
boliara ofwoiimn, Icailinir t» lli<- xik of a wontitT iliol. in n-iinpanHl with tbu
of n]au,isli>bon-^iiri)i.-'l ju l]ii.-> tight nilhnr than with rvrcn'nce to theavenitc
le^er wd^ht of woniitn. The n-hitivc nn^tiibolium of thi- twn s«xe( Diar be
illugtraioi) br tho I'luc i^f nil nclivr nmn iiiiH bU wifi:, iMhofabmit the turn
nse aii<l weight, tin- miin bcinir rather the lu-iivicr nn>) thu woman rntber tfai
"hicr. who. in carryiiiE oiil t'ljrrtbpr an oxpirimctil on iht rtlutivc valuM rf
vcgviablc Hill] niiimiil foorl. both livci) for fiiito little on tbv Mim<- kiii'l el
li'uit, uut] found thut iiiitriiivo «>.)iiiUbriuiii wa£, in thi; on« caee aixl iu thi
Other, Riaintnin«d when
Tlic man connumed dailr about
Thenrifr
■"rowtdk
Pan.
ntrtm^lMlM
]W
70
WO
m
87
SiU
Tlic nicMt strikiii;; difference b in the proteids.
g 656. With re);ard to climate, ibc chief cons ide ration' nttnch to ICfapori'
ture. When the body h eicpcKed to a low tentperalufv: lhef;cneral nMa*
boliam of the bodr is iocreaii^d owin^ to a regulative action of the ncrvotu
svRteiu ($ 535), We might infer from thU that more food is necAMarrii
euM cliiufttei; and, aince the increase in the metabalinm appeari to tnanifcM
il»elf chiefly iu ii j^ieater discharge of carbouic acid, and therefore to be
especially a carbim metaha1i»in, we uiig^ht infer that the carbon elemeata of
food abould be e»|ieciHlly increased. When tlie body ia ex[K>eied U> bi)A
temperatiirM, tJieiinnie retlex mccbaniam teud« to lower the mctabolbni: bat
the effec!t» in thin direction are much Ion clear than thue of cold, aiid aoon
reach their limits; the bodily tciuperalu re in muinlaimv) coiMtnnl under the
iaflueDce of surrounding warmth not mo much by dimini«heil production lu
by increMod lom. ' Wc may infer from this that in warm climatM not Imk
but if anything rather more, foad than in temperate climates is oec«aMry in
order to supply the perapiratiDn needed for the grvaier evaponUoa and di»-
obar]ge of heat by the skin.
In both cold and warm climates, however, man trusts mucli more la
rarJBtioDa in hit clothing and itamediate surroundiuK^ to protect hiiu againM
cold or to Eunrd b>m from heal than to any marked variatioiu iu his normal
<liet. Ill Hie former he may perhaps be expected to eat sotDewliai mure,
tiinue, in xpite of WRtppinjiH. hia okto still feels iu part die cold, and thus the
ncrvouA mcchanttiu for ibe inereote of iiietabolbm » to a certain i^xtenl mi
to work. And xinre the inelabolbim thut) inoreAitei] apjieani to atTect cape-
cinllr the ntrlion of the hoilv, he mnv further be expected to incrca«i: Ui«
fat* rather than the rarhdhvilmtni of bin food, neeiag that the fiirnii>r KUfiply
him with the inoyt energy for thur wd^^ht. But it is very doubtful whetner
what he might ihii* hi; cxpcctv<l to ^in over a corn«|Hinding incrmM in
carbohydmtiv in not more than coiinierlialanccd l>v the iiicreaMxl latior of
di;r<v|ion ; and ihi- habits of the dwelleni in arctic cliraaCcM cannot safely be
taken ns giitdcH in ihi» matter, for their reputed love of fiit is pn>baMv the
roiilt of that heinif their most available form of i-Jirbun. Indetvl. the evideaoe
that the iticrean- of rnelAholism provoked by cold bmrs excUiitively od carboo
ooostltuents is so uncertain that it may Ih' doubted whether any change ID
the normal diet, bevond some increase in the whole, should be made to meet
n colli climate. Similar reasooa would load one to infer that man in tbe
warmer climate would maiulain, on tbe whole, tlie same normal diet, the
only cbange being perhitiu to increoje it Eli|;;htly, puasibly thmwini; ibe
inerea-te chleOy on ibe carbvhydrales with llio special view of ftirlliering
|icni|>iratian.
I
on DIET.
I U7. A fpccwl diet for tlie purpoK nf (ati«Din^ — that b to Mtv, for '
■ctnnalnttoD uf sdi)Kii« tiutie out of prfifKiriioii to ilie mt of ili« t>nily — it
Mt MVtIvd in llie tnfv of man. The \m*i!t tu >t<>n.' ii|i fat in HtliiMNM; iLuuc
b micb more dc|>rmlcnt »it ci-rlttiti iuhoru i|t)aliiim of the umnwu) uliich
w* cunot at [irvMDt iHioc itiun mi llii^ kind of food ; of two Wlin living
Ml lh« aanii! dii-t, and under th*t simtv cirouinvtarnxv, oiiu will bcfonx- Int
■ hilv the olh«r will remain Iciin ; and il i» tin object of llic amoilluriKt 10
ikr«]o|i by lirwding nnd »rli-cCion a "coiirtiliiliiin" which wtH rtorc up ibo
neat (at on th« clM.-iip<«t diiM. in ftltrniog anitnalo tb« cJiirf can-, when the
Mlrction of iIm' kin<l uf nnimnl hn» Im-n oinde, i« to provide ndt'iiinie cnrbo-
hrdmte food, which. » w« have h-cd, is tlic chief tatlrner; and Uic object of
thv liarRivr in rrnrin^ stock for iho biilcfacr te nminly to convert cheap vege*
labia cnrbohydrntc into dcnr animal fnt Pimbcr aids in fntlcoing rony m)
found in providing rcpum for the body of snch u kind that, whil« sufficient
nergy is expended) to secure ade'juatc digestion and nbsorplion of food, all
caures leading to an increase of melabolistn. by which energy is set free and
Itaves the body, are avoided aa much a> possible.
Tu avoid fai miber than to inoreaM it ia often ui objed of human can.
This may be eticL-led by diminishing fats and carbobyd rates, but also, in a
very nwrkfd manner, by relatively inereaBing the proteida. I'roteid food, u
«C MVe wen, augmeiit« ibe whole metabolism of the body, hurrying <m the
dMtniction not only of proteid but of carbon food; and a tendency to cor-
pulency mair be couatenicte<l by a diet in which fata and car bohy<l rates are
much renncted, and proieidu erv largely iDcreaaed. When, aa in what is
known as the Jianling niethoil, the uiel is almost exclusively proteid, the
nitrogetiuus nverwork entails dangera on organinnis which do not possess the
power of ridding ihemaelres freely of the large amoiiui of nitrogenous waste
which such a diet produe>r*. A lea* Mvere methoil in which the &ta aod
cartmhydmtn are diminished only, not CBtirely done away with, and lite
Srottida only moderately increnspd. is lew open to objection: and such a
ict, BMotcd by other hygienic conditions, has proved successful.
Ad increase of daily lood, largely proteid in nature, given un<Ier circtim-
MaDCa. such as a large amount of iwssive exercise and skin stimulation,
known as " maaeago," which will not only favor digestion but also promot*
■nUbolitin in general, may be given with favorable results. In this wayan
caorBWas metabolism may be excited, and yet so carried on thot the body
B^m both in tiesh and in fht. Thus, in one case, the patient with an
mhial weiKbt of 46 kilos, and a daily nitrogenous metabolism calculateil
■a 28 pms. proteid, reachtKl in the course ot about fifty days a weight of
60 kiloa, the dailjr nitrogenous melabolbm being raised on one oocaston to
182 ^ms. proteid, witli an average on the whole |>eriod of 150 gnuB.
Doring the treatment no less titan 8420 gnus, of proteid were taken as food,
g 6w. With regard to labor, since as we have seeit tlie energ\' expended
as work doite is not taken unt of and away from the amount set tree as heat,
the two forms of energy being so related that an increase of work dona is
aoooinpanicid by a greater orlcHii increase of heat set free, it is obvious that
a nan who is doing a bard dav'n muscular work needii a larger income of
aurgj for the day itian dors nn idlii nuut. What ne have learnt concerning
muscular mvtabo1t«n further >hows um that the additional energy needed ia
not necessarily to be tiipplirtl bv nn incirnK- in the prnii-id com|>oncnU of
the diet: tl>e energy of musi'uinr contraction doon not pome an was once
thought from proteid metabolism |^§ MO). The fact ibnt it is the carbon
tDeiam>li*m which ia augmtated in muacular work may suggc^ tltal the
BXlra (bod for extra work should bo exclusively carbon oomfwumts : and if,
aa SteODS pn»bable, the carbohydrates are more readily and dircrlly avaiJablo
670
NCTRITION,
fertlieAinctionBl meLabolItmof luuicle than are (tie fntA, we laigbi beAutW
led In reoofBinend an iucruwe !ii oarbolivdniua to funn a di«( e9|>«ciallr niivj
for labor. But wrernl cni»i(l«ntioi»«litiul(I iiiakeuH liexitaie before w« con
to silcli u coDclu«ion. A miitclc ii nut u luiiahinu wiiliUi llie \iodv vliicli oa
be lowloci aud Bred wlf irriMpcctire of tbe nvi nf the budy. In tJie jwriunu
ance of muscular labor, tbo ooiiditioD of the miiK-lo, the amount of cotr^v
available !□ the muacle itMlf, i« of eourw of pritoK impiirtaiiee: hui,ijii!
tbis perbajK etpociallr hold* good iii wevorc labor, of ^mit imiionaiiM tin,
we might aldo «ay of no 1«>« itii|ii>rtuiKf, b na w« have tir^ed Ci 391 iln
|K<wer of llio body ii« a whoU- to iiTuil iuulf of tbe energy lau-nt in lh\i iuu»'
cl<d, The power of doing work bangK not <Mi the niUKle sIodo. bill on iht
heart, the lutigs, the nervouH system, und. indeed, on the irholv IkkIv- It u
very doubtful whether wo c^*cr, even in supreme etlbrt«, draw upan luier
Uian a portion of the capital of energy lodged in the muscle ilsvlf: fitli^
b for more a oervoue than a inusculnr cuodition, and even tlie diuiudlt
muBCular fatigue is as we have seen (j H^J partly at least the resitlt tf thir
accumulation of products and noi alone the using up of available <Mr|;<.
In choosing a diet Jbr muscular labor we must have in view not theniiint
itrelf but the whole organism. And though it is possible that future rswan^
maysuueeets lUinor obangea in the various oouiponenia of a normal ifiel neb
tu woufif leaseD the siraiu duriii;; labor on this or that ]karl of tbe body.w
the inuiclea aa well ns on other organs, our present knowledge would ratkn
lead US to eondude that what a good for tbe orsanism in comparative im
b good also for the orf^bm in arduous work, ihnt the diet. Dorroal tat tk
former condition, would need lor the hitter a liuiit«Hl total increoje but g»
•Iriking change in ita conipuailion. lu iireuiiriait the body tbr aome cuain;
arduous labor in '* Lraining" as it ii eulletl. an inureaie of protod food, (ot
th<i purpoMe of liurryintt on the ^uural nietuboli-tui of the body, and tluu
of making " new lle«h ' and renovating the bo<ly. «o to speak, to vm« o(
the strain lo be put upon, may jwrhape suggtat itself; but even ikuW
doubtful.
The priaci]>l<« of sucli a conclofion with rt^nl to muccular work aai
be applieil with still gnuilcr conlidenoe to nervous or menial work. TW
actual cx[>enditure of energy in nervous work is relatively small, bat tk*
indirect influence on tbe ecunonty is very great. The cloMUew and iiiin
cacies of the tics which bind all parts of the bmly together is very clenrti
shown by the well known leadenciee of so-called brain work to derange ibt
digestive and metabolic aciivitiea of the b<]dy ; and if there be any did
ee[)eciallv suited for inldlectual labor it is one directed not in oi
toward the brain, but euiii-ely toward li^dileuing the labors of and ainq
the way for such juins of tbe body as Uie siomacb aud liie liver.
BOOK III.
THK CDiTRAL UERVOUS SYSTEM AND fK IXbTKL-MBKl^.
CHAPTER I.
THE SPINAL OOBD,
Ov SoMK FiuTirRKB OP TUB 8ri!iAi, Xkrveb.
I 558. Wk liav« csllml tbe rnusculamnd nervous limuee lh« maclcr Iimum
vf (be iHiily : hut n epccinl part of the nervous systcin, thiit which wc kDOtr
M the i.'cfltnO ttervouB bvbUiiii, the bmin mii) spinal cord, is supreme aiuonj;
llie uerroiis tissues nn<) is iniuier of the skeletal iniiscles as well as of the
len itf the bo<ly. We Inive already (Book I., Cluipler HE.) touched on
•on>e of the (general leaturei of the Dervous syslcm, aud have now to study
ill d«inil Um votk'mg of the braiu and epinul conl. We have to inquire
iihci we know coui-eniinK the taws whicb reculate tbe diacbsfg« of eiferent
inpubeB froDi tbe braiu nr fruni ibe cord, ami to learu how that dtKbarge is
iMtrotiued on tbe uue hand by intrliiiiic chanKt'i uri^inntiug, apparently, in
the Mibstaoce of the brain or of tbe conl, and on (he other hand br the
naturf and aniouui of tl>e aiferaiil iiupulttce ubich reach theui along aOereot
attvt».
Aa we shall *ee the ttu^y of the spinal cord cannot be wholly aqwriiied
from lliat of tbe brain, (he two being very cliM^ly rcla(«il. NvvoftbeleiM, it
will be of advnnioge to deal with (he spinal curd by itwlf a* fur a* w« am.
TV medulla uhlotinu or spinal bulb' we *luill cun.iider an pan of tbe breb.
Bui belure wo speu of (he mmiibI cord itself, it will be dcnrablo to snr a few
wonb GOBcerning lh« spinal neri-n, that is to say. Ihe nerves wbicn issue
fVom (ho spinal cord.
We have alrcsdy soeo (S 96) thai each of the spinal nctree arises by two
roou. an anterior root aUached to the ventral or anterior aurfac«. and a pos-
terior rxio* atta^^'hcd to (be dorul or piwteriorjturlaei) of the oord. Wu have
further seen ibnl ihc latter bears a ganglion, a " ganglion of tbe posterior
r<>ot " of ** spinal ganglion," and we have (% 97) studied the structure of this
Ipiiialian.
We stated at tbe same lime (hat while tbe trunk of a spinal nerve con-
iaiu«(l both efieivnl and aflV'renl fibres, the etlVrent fibres were ^ihered up
inhi tbe anterior rout and ihe aRerent fibn-s inoi tlie jtoeleiiur r^ii>t ; hut we
gnvt no pnwf of this aiaiemenl.
j 000. Before we proceed to do so, it will be as well to aay a few words ou
• llwlunntdalkoMoMkiaUeaicailr lui)a.Ii<iiprtiwnwdUBt«ltli^ilnMtlu want mMhilla l
l>Dn* iwWaaidloaMnMiw wlial*ii'lna^nirdt»«dall>a(4iwlto)IMIIifMi«r«l>r iiMdIadmnMrl
■iMwcKllsrcaatAtaiMmabn.tlMirbliafatMsnMnr SelmuB. in ariit iMtaul tlw •ih4 ti* I
iiTtrair^iWTir.MflwINilSUiwclilHilefctrottonfimonwIih— ypQwtMndirf Iwlti. 'neo^koUre
b la wa Bueiinkoa usi^ lu uxb plifwa* ■■ " bultar |«>*ll**-'
«72
THE SPIN'AL CORD.
tli« Icnuit "efTereut" mid " nlTeivnt." By d&rctit iicm-u Rbras wv mtn
DftTVe lilirea which in llie 1>ody iiftuall}' nirry impubrt rn>in [lt« cvoinil im
TOUS lystcni tu perijihernl (irj^im. MomI elkrciil iifrvc fibnw cHrr\' impulai
to miMcleti, Ktriateil or pliiiu. und the inipulwcii jnLwin^ nloof; tliem givr riit
to moreDienU; Iicdcl- tnoy are frequcntlr t>pokr-D of ni> " moUir " fibre*. Rg)
all eBlbmit fiUnv do not end in or c«m' iiii[iul«cti lo niiii>ciil»r Rhr>^; wr hiit
•een for imtlHiicc thtkt pudp cfliirfiit fifirw urc ^ccwlory. Slorewvpr. all tit
nervfi fibm goin^ to mii^ciilnr fibrt-n do n»t mtrvc to produce lumciDCdl;
>oiii<i of th«ni, ns ID tho c«m of ccrtnin vnguv RUm going to iIm- hvAtl, ir
inbibitorv sod mny wrvv lo flop movcmrDi.
By ■■ nflbrvni " nerve tibrc" wc nienn norvc fibre* which in ibc Iwily itHiillT
cnm' impuIsM from ixriphnrnl orgHii» U> the cetilral nervous ivsteni. k
very coniiiKin eA'pct of the nrrivnl nl the (.'cnlnil nervous lytteni »f impulNi
pnsNnf; along atlcrent fibres it that cliniij^ io consciousneas which we csU •
■' «n8»tion '■ ; heoce atlerent fil>rra or iuipuUes are often C*U«d " seDnrjr "
fibm or impulses But m we have already in part seen, aod ■» ire uall
aboitly see in greater detail, the central uervoua system may be airecied by
afferent impulf^s, and that in several ways, quite apart frum the developrncat
of any ftuch change of consciousness as may be fairly called a aciuiatMB.
We shall see reason for thinking that oll^rent impubeii reaching the tpuial
cord, and, indeed, other pans of the central Denroui svMem, may modify
reflfix or automatic or other activity without neoewarily giving rue to a
"aeiuation." HeD<;c it ia advisable to reserve Die terinft "cfl«t«i)t" and
" afltreui " pa more genenil inodea uf cxprevion tlian " motor " nr "Mi»ory."
We Lave «eeii iu trciiting of niuscie and iier\'e, that th« cluingis prxluccd
in the muscle aervc an iiur butt guide for d«t«finining the chan^fc* taking
plave it) a motor uervi^ ; whrii n motor nerve is separated frum ii» mufde
(§ 72) the onir chungi.^ nhich we cuii appn-ciatc in it w on ckctrioil change.
Similarly in the cose of nn afleriMit nerve, the central fVBtem is our chief
teacher ; in a bundle of alTeretit librf« iwlntcd from the central nervous stv
tcm, in a posterior root of a iipinal nerve for iuflaneo, the only change wh *
we can spprccinte is an electrical change. To learn the charadera of af
ent impulses we must employ the central nervous system. Uui in this
meet with dilTlcuttieg. ]n studvitig the phenomena of motor nerves we art
greatly awiated by two facts, tirst, the muscular contraction by which wt
judge of what is going on in the nerre is a comparatively simple thing, un*
contraction ditlering from another only by such features as extent or amoont,
duratiiiu, frequency of repetition and the like, and all such diflbr«ncM arv
capable of exact measurement. Secondly, when we apply a stimulus diroctir
tu ilie nerve ilnelf, the Gtlecia differ in degree onlv frum thoM' which tfmU
when the nerve u *et in action by nalural stimuli, such as the will. When
wt come, on the other band, to inveetiKaie the uhenomeua of ofl^nt nerrn,
our labom iire for the time rendered heavier, out iu the end more fruitful,
by tbe following circutnilancet : Pint, when we judge of what ingoing on in
an affcrrnt nerve hy the etTeris which stimulation of the nerve priHhicca in
•onie central nervuu.i organ, in (he way of exciting or mollifying reflex
action, or modifying' nuiomatic action, or affi>ctlDZ conMaoutnoH, we an
met OD the very lhm>hhol(l of every inquiry bv tb« difficulty of clearly dia^
tinguiahing tho cvciita whicJi belong exclusively to the aAerait ner^e fmm
thoM wbiui belong li> the central organ. Secondly, the eflecta of aiiptying
a stimulua tu the neriphcrtd ciid-or^n of an aflcrcnt nerve are ver)- dilTerent
from lliiife of nmOyiiig the Ninie «timiilu« directly lo (he nerve-trunk. This
may he shown by the simple experience of comparing the senration caused
by oringing any sharp body into conlHct with a nerve laid bare in a wound
with ilial caufed by conlnct of an Intact skin with the same body. Tbew
(liOtrencea rvval to lu u ci)ru|>l«iitx of impulaeB, of whicb th«
a« of motor nervee «ve m liitnlly u hint.
loll further ee* iu dutail later un itist our conaciouNica mav be
•flnietl ill uisur (liflcrfiu wnj-« by alti-vfiii iiu[iuUeei we inuM (iMtinevbli
aaly wiuory from other uflvrvDl iinpiiliieit, but aJiiu dilfereut Ictiid* i>f
ry impulm from eui-lt utlier. C'erutiii «lliif«iit uerveti ure itjiiiketi of lu
oitfeaal wnae, kikI ilie nature of the affereut impuluea pftuing aloae
•peeial nerirea, logMber with the mwlifimliuiia of couaciouiaeu vuuwd
br*rrivmlortbeae irii)iulM-Bi>i ihei.i.-iilnil nervous ■yat«ia,0ODiUlut« by tlMBf
•elrai ocMnplez un) ilitliaili bnuich <>l''tu<lr. In •unu) of the problenu oon-
ncctnl with ilio cnitral ivi-n-ou* untviu w« tnitU hnw to itp[K«l to the rwulta
ufadmlvot' lli«»vK|ic<.'iiil •eiiMv; but, <>» lli«ot)ier hauil.a kniiwIiHlge of the
ftntrnl iwrvou* kyrtciu U uvctwsirv to n pro[HT undentt«n4liti|j of tlie mmk'ui\
MHta: and du iba iibolw it will be niurc convcniciit to Mludy tlut fonuur
btbra Ibf laitrr.
We may, biiwcvor. digrtn here to rornnrk thnt the 'jumtioa whether an
ftUvrvut iutpulsotliUcra in iteclf from au vlfvretit itupiilse a uiiv uf grcitt tliHi-
eullf. It w tniv tliut lh« clcctricul clinngw. whicb alunc, lu w« have sai<l,
waQaaap)irc4'tHlv iu aii iMlate<l piece of nerve, appear lobe thoaamo in both
Uoda ol' fibraa ; to eacA the electrieul chan|^^ it pn>piigaled id both tjiree-
liaaa, aod pciewM the same fnture». Btit it would be hazardous to iuaiM
IM niuob un Uih. Moreover, we niu«( reiiieniixT that wbal we call a nervou*
inpabe, eapecialty ooe provoked by artilicial uiniulacioo, cunstitutcfl a tira»i
ehange in the nerve fibre, uod timt obaogce of a fioer, more deliuace nature'.
MOb ar cannot be oliowu b^ the eunrse methods lued to ileteet u " uervoiis
ia^Be," way lake placv iti. tiuil be pro|Migaietl alon^, a uerve libre. We
" have occasiou iuiiii«tliutely to puiui nut that the cMudilitui of au altt'reiil
8bre along ita wbiile length is <lej>efideui on a nerve celliii tlie }i;uu^lii>D
the poetemr nxit ; lb« tibre wheti cut otrfraui the oerve-ctll tU:|;eu«nilM
and dica. TbU mttau llial in tlw intact libre certain inBiieucei are prupa-
ntctl aloDK tlM- fibrtfrom ibe ecll in the ganglion to tlic jwriphttml cothngv of
mftbre^tkat i» toaay inadircciiiMi th«opMNit«of that taken by the ordinary
iwrvaua ioipulica : and it mnr br tliai in like manner in oflerent fibrw tome
ioiluetlMB ore propagated c«ntripotally from the p^'riphcrul endings to the
etntnl oervous syatem. Uur knowleilge of ibew iuHucnoca is extremely
UntUid; but it is important to bear in niiiiil the powibility of their ocour-
nooe. And we bad tbta in view, when above, in speaking of ederenl awl
albrent fibre*, we use tlie pbraae " usually carr^' impulses."
$M1. Thepruof that the alTelvntaudetrerenLfibrvswhich are bocb present
in the trunk of u -j<iual nerve are parted at tbe routs, the elFerent fibm
rtutainK exclusively in ihe ventral or anterior root, and the atlereut libre*
exdunvely iu tbe dumal <>r ptoilerior rciot. is as follows :
Wb«n the anterior ruui it divided the rauaolee supplied by tbe nerve ctue
lubi- tbrunu into ci)ninii:ii<.in9, either by the will or by reMex action, while tbe
rirtictures to which the uerve is dintributol retain itieir woaibility. Daring
iIm section of ihc root. i>r whtrn tbe proximal stump that counected with tlw
»pianl cord in stimulated, no feuory etiects art> pruducwd. When tbe disttil
■tump is stimulated the musrle» supplivi] by tbe »vrv« are thrown into ooif
tractiotui. When the (HMierior rout i*div-t<led the miutdea supptiuil by tbe
uarvacantiniie to be thrown into action by an vxercisvuf (be will, or as part of
n nflex action, but the rtrucium to which the nerve is distribut^xl low the
■ettsibilily which they pivviousJy poMtwod. During thv svclivn of tbe root,
ttnd when the approniuiiil Munin is stimulated, tbe senaury eflecis ore pro.
dooed. When the distal stump is stimulaieil numovemenu are called forth
»
lapai
TBI SPIN'AL OORD.
Tlii-M- rni-M liurnoMHt rail! Oist »eu»>'rr tinj>u)»n |>iim I'xcltiHveIr Itr tlw jx
rior m<>l fmm thn ]x:rtiilii-ral !» iIm' r«iitrnl i>rxiin», nii'l itint molnr iin|iuk
|M« vxcJHxivttly by In^ niili-rior tiM fruni Ine wntrnl to tlir prrifik*
nrgaiis; ttntl lufttr an mir kuDwhtdgi- ^lut thcMiiit; linldii gund uo(l
wiwon' Mil) motor but nlsit fiirnOirrDi nii<l«-flrn'iil iminilMA.
An exvcfrtion niuot bi' inad^ to thv iiIkiv« f^itcnil nUKrmi'nt.nii tuvaaMtt
the Ro-cnllrd " recurrent M-iisibility " which n v.itntittvd \n oi>iiertiiu« ouui-
inals, uaAcT ivrUin ciministiincM. It imn>otiiii«» hnp|i«n)f tbitl wbi-D 1)m
(lialsl stump '^f lh« diiHdcvl anlenor ront to diniulntol, mbiii' o( ]tnin an oil-
llflfd Tb«i? aiv Dot caiis^ by the conciiTrenl inUBciitiir oiitrnclMitu <>r
enuilp which the ^tiniulnliou ocranioos, for they pcnist nl1«r the wIkiIc tniak
of tlie nerve faaii been divided anine little way below the union of tlie nvtt
above tlie uri)(inB of th« muwular branrh«e, m> thiil no comnirti<>n* Ink*
|>l«oe. They disappear when the )xiJteri<>r root is Bubseqitenily r( t
lh«y are not seen il' Ihe inixeil nerve-trunk be di\"ided cluee to il f
the toots. The phenofuetia are probably due tn tlie I'aet, tluit ixindla nf
•AOfwry fibre* of the jMxiteriAr root atter runuiDX a ahort •li^iarx'V ilimn ik'
niisn) tniuk, Uini bn<-k and run upiranl in the anlertor n>ot (Ix-ini! <' - i
utcd ]>ntl>nbly i» the pia iuni«r) and by this recurrent eoura^ give lu-. i
iIk' renirrcnt aeniiibititv.
$ W2. CWcfniiu); tfif i^iir^/infion Uie poUeriorro()t,we lnn)'•ll]■d(flldt«lr
tIlat vr« have no vvi<Vn<« that il can act Ma evntiv of r«Aox action nor h«vr
w« any evidence tJiai il cbu s|H>iitanv>u)ily (riv* origin to rKerent \n^~<-' ~
und tliuenct lu a» nulomatio centre, a* can the ceiilral nervous ^ysteii:
The bodies of tlwiwrvcocUe behave iturwvf bat dilTorontlyfrvim ihe axis-"* lin-
ders at Minte d)«t«ii<-c from the cllx. thntif^h, ii» wc have aMu. these ar« in
reality procMaea of the Dcr\'cc«IU: ihu« the n«rve-cctl8 in tl>e inuiglino
appear to be more #enattive to certain poi«>ns than are the nerve flbree of
th« nen-e-trunk. But bevond I his. our knowledge otnt^minf; the ftiiMlino
of the gaiiKlton is aln)u*t limited to the fact that it i» in flome way intiaialfiT
conaeeted «ilh the ntilrilion of the nerre. Aa we have already '(§ 8S) enui
when * mixed nerve-trunk is divided the peripheral portion uegenenttf
fVom llie poinl of seclinn downward toward the peiiplwrr. The eenlml |"'f-
tion docs not m desenerale, and if the lcnxtl> of nerve renmviil In* not <>•'
great, the tvnml jiortion may Rrow downward bIoiik the eounie »f ilnt df^a-
eraling peripheral |x>rlioii, and thus iv^nerale the nerve. Thi# div>'<>*n-
eraiion t* olu^rvcd when the mixed mink h dividol in anv (xart »f il4 niunr
fn>in Ihv iK-riphery to clnae up t<i t)i« ganglion. When iIm; niBitcri»r roM it
divided iMtwcen the ganglion and llie «pinnl vonl, the ptirtmn atlacheJ i"
thiMtpinal cord degeiMralea, hut that alutcltud b> the ganglion remnin* 'n-:!"-
When the interior root is divide*!, the pruxiinal portion in connei.-tio]!
th« xpinal cord remaimi intact, but the distal portion lirtwcwn the ^ni mi nnl
the junction with th<- other root dtg«D*mtM: and in tho inixeil n<'iv>' iru'l
many deeeiMraied (ibm arc »ceii. whidi, !f they bo cnrefully traoed i«ii, irr
fonnd tolw motor (e(R>ront>librci>. If the piiaterior nxit !>« divided nnfal'.'
botweni the ganf^lion and ihi) junction M-iih the anlerior roi>t, the oaall PO"
lEoti of the poilonor root left alUcbe<l to ihc j>eriphvrnl side of the jraagllM
whow the wction remains ininct. as Aon h!>o the rufi of the root ma tbi
Sanglion to the spinal conl. but in the mixed nervotritnk are aeou numffM*
^noratcd fibres, which whea eiamined are fuund to have the dinribnlw*
of Mnsory (afferent) fibre*. Lastly, if the posterior gaD|;lion bo exric*!.'**
whole posterior root dcgeneRit««, as d» atso the sensory (alTermi) libm'»
the mixed nerve-ttMnk. Putting alt tli««e facts tof^tlier, it would *eeai ibil
the growth of the eftorent and aUereni fibres takes niaoe in opniieiu 'tirw-
tiima. and »tarl from difterent nutritive or " trophic ' centnuv Tlie aAnn'
TUB ATBUCTrRE OP TUB 8PI.VAL CORD.
675
■way ftwRi tlie fninglion cither townnl Htn poritihery. or toward
the ifMiial Mird. Ttie «l1erenl libm grow oulwurd from the iMtinl con)
tovani ibe [leripbery. Tliio ilitTercocc in their Rivd« of nutrition ia ft«-
()oniilr of i-reat helpio investigBtio^' lh<- roliil.ivixlistrlbiilion of Hferenl aiul
ucreut 6brea. \Vh«u a poelerior root is ciit beyond the piDi;lioD. or the
j[ut||;U(in esvited, nil lite allbreut nerve dc-geaeratv. and in the mixed iicrve-
hrauche* ihete aRereni Hbres. by their altered condition, can reiblily be
Inml. CnuverMly. when the auterior roote are cnt, the edereDl fibrtti alone
ilqpDenUe, and can be •Liiuilarly reoopiizied in a mixed nerve-tract. Whoii
■W sntrrior root \» iltvicKii some few fibres io it do not, like the rent, dogen-
nnuc, and when the [xsteriur root U dirided, a few fibres iu the aoierior n>ot
atr M«o to iWcneraie like thotu of the posterior root ; these appear to bu lh«
fibrr* whlrh ^ve to tlie anterior root its " reirurrcnt sensibility.' In the one
(iTrcrtain •piual iiervM al nil elleeU, it hii» alixi been asoertainiM) that whra
the posterior rixrt w divided, uhili- most of the flbrta in llif part of tlie root
(hui< cut off from tbe gatiglimi but Ivft attached to the mrd dcgoncmle, some
few do uot. Then few appear to iiare thmr trophic centre not in the ean-
^lion, but ID some part of the »pinal cord itself; we shall rufer to tSese
later on.
This method of distingiiishing nerve fibre* by the features of their degen-
«mt*oa, called tbe "degcncrsiion mothixl." or somelinM* from the name of
Ibe phrsiologist irho inlroducwl it, tlio " Walterian method." has proved of
f;nrat utility. Tbn« iu Ibe vagus nerve which is composed not only of fibres
which spring from the real vagii» root, but also of Hbres proceedini; from the
Bpioal accessory root*, the two may bo distinguished by section of the vaguit
and a|)iuBl accosorv roots reftpectivcly. We shall presently see that this
iDv(bo>) may be apiilied to tbe aifibren nation of tracts of fibres iii the brain
and spinal cord.
TiiK STBtrcTOBS OF TRK SrntAt. Cord.
I S68. Lying within the vertebral caniil the epiiial conl is protected hy
ba ** Rwinbnines," the dura mnter, the arai-bii<>i<l motnhntne and the pia
■later. Tbe oonsidrratioa of the arraii^vnu-til of th»e membrance and of ibe
leun* of Ihc dura mater and tirochnoid we will leave until we cnme to
of tbe vascular and lymphatic supplies of the central nervous syatem ;
histology of tbe pia mater may more fitly come with that of the s]»iial
rord itself.
Alouff iu whole length from its junction with the bulb to iU termination
In t)>e jv/hih terutiiu'ilr the spinal cord, while poswetioK certain geuerni fea-
lurrs. i« eontinually changing as to apodal featuree. It will be convenient
to Mady tint the pi^enerai structure (if *ome particular pari, far instanoo tlie
mi'ldlc nftlie tbomcic (dorsal)' region, and afterward to point out the special
fealurcv which obtain in the aereral regions.
A traiumnc vertical section uf eitlier a fresh or n hardened and prc|>arol
' lal oord at the thoracic regiau pnaseww an oul3ine which ii, roughly
ting, circular. In the middit? of the anterior or ventral surface is a ver-
Satare, the vfnlntl or anlrrior finare (Fig. 174, A. F.) running some
lll*twiil«ib»Wal**«(lietsrini"'lMwr'ft«i1"v>nmI" Ijt th* ptitior tlWnnlini-iMiHl
*ii» ■Mso tig ni>inaT»lr naw (be 4Dral«iiwli i<ui. ■oil ike Tenml or botlrun of (b* boer,
laria*Mnn*po<*t«Mriiii4knt«r|qr: Inl It this it dua«. IW iw* u[ ilia witi liormi 10 iltftWS
I «r IM OMtl bMiraen ik* lambtr ani otrntal ncknii ii apt i'> Ia*i4 lo ooaPBiaii i bsoet
bMMarfniawart Uinnola, ttthunx nf dfimal •»] ircati*) tw ai1»tfB< bj^bab>«B»a
,. eBiesnntnlna mif r riTinliaitjtwi uMd la 'Iciuiia oawar ibv bMilsiid maiwUigmi (m
[ Mi(iwlIt«V : aiiMTlat aiul Matarlur but bIw b« awd In 10* isim wnia stM^i la lb* <ua
■rtnr aiul imartw BtHiw and bM>. whlob Mtoi* wan Im ■oak aaa«Mil br <1B* lo b« Ibnwn
ST6
THE SPINAL COHI).
Fio. 171.
A Tkam.it,»6e D'liiMi-vr.NTiiAL Stmiis OK THE Spinjl Cori> (Hi:mak) at Till Level or tkeSiw
TllOHACIC (l>fR»ALI Skbve. (Shebbisotos.iI
Ma^iiJIled 1,~> liniL'?. t ini? IniT^nil balf t>ii]y l^ hlioxvn, Tbu large »>i»pl<^uou> nerriMwlla idn"ii
from ik^IuhL »-iH.'t'jii]L'1jsi aru shaded blurk to rcndi^r their relLtivtf size, sbalicaiiil poalllotl monol*
vli>us: [licoiitllibv of iliegrBj' riiHIIrr lnnlpwn uihiIc ibluk Bnd r]»rk In order toremU■^ltc<mfIJcDu^«■
A. F.. fliitt'riiNr i.r^ur^.' i I', y., |^*ti'rior n^pll^o : r,*^.^ oeiilrsl catial ; f, ff. *-, centr»l getatLtioua tab-
Malice: A, r., ririlirldr rrjcit : 1'. r., Inli'tnl (or Inlennedtatel bundle : P. r'., medlnti bundle or poW-
rii.T root <i[HiliiKl iR-r*v : ;.', j." filin'?<if i isicri or n ml paning. p' indirectly through tlw fotaanwrf
Utilaiiilo, )'" ill net ly into itray iiiHit^r; rr, 'i, /•., anterior gny Gommtnure: p^ ff. c, potenor^nj
ci^iniui^^ure ; t, -■.. oiucriiir Hhilt coiiinki^snrv ; anK cof., aulerior column; fa/, eof.. lalen-L coiun"'-
pofl. (■"'„ i-i-lvil-pi i-.ilunm ; «. ■/.. ihu tulMance of Rolmido ; «,, wptum rD&rklQK oul the eitinul
Itofierlor L-ohiiiiii or ivjliinm oriluniacli, c.p., from Ibe m^iari ptielcrior colomn.or column cftfvl-
HI. ;j. 1, '"^11= of ihoiint^rior Loni : 3, pijsierior column or reaicular cylEnder. or oointnn of CTirke'
Ihtf BtfHof thoijlliiijtri'dellnedbj a dotted line: 1, cells of the intennedio-lateral met or Wn*"
trarl or lalefHl liorn; ii, cilL" ff Ihe |io-ierior horn: ", cells of Ibe anterior cerrli; y, a tracmf'i'''*
iv^liig l'ri.iL> ilie vesii'iiliin-jliiiik-T III tlie lateral culutun,
way Bcross llie llikkneBS of the cord from the ventral toward the dowJ f""
fece. t)|»pijsiie to it on the posterior or dorsal Htirface is a correBponaiogt
deeper but n&rrover, dorgal or posteritrr JUsure (Fig. 174, P. F.) which, bo^"
I Fur tlii^ and nninv -iHCniBllntr flgun?* I am deeply Indeblfd lomyfHendmnd itonner l*!^'' '*■
Slicmtnil"!!, who hu'- liUidly pre[jflroa llie figiirtu ft>rme from bis original drvwlngii
wr«t ibc mli^nn«' (if ihc cm) i* reilopet) lu ■ iinrrow uclimu!) imiliu^ lh«
Iwii Isi^nl liiilv'fs, ntiich in » nortiml oord are like eni'li other iti nvery
ntpttt. In iW miilillf uf \hv islhiiiiM Wee ilie section nf a dinuli caiKil. fA«
flnulni/ miMi/ (Fig. 174, r.r.), whicii is itll ibat reinainHof ilie rcliitivdy ivitk
nnimt autal of llio «inbrvo.
EsL'h UcemJ biilf oorutMs of an mitcr zone of wkUe mailer surri)iini1!ntr<
•ici'pi nt Ihe istliRiiiB, an inner more or leas cnaceulie, or cammn-i<hii|ie<)
■as of yr.iy mailer. Th« oonvoxitv of «oh crescent iti turned lowanl tlie
meiliKti )in« nf the coni, tite liro creacenta being pluoeil buck aui] buck ami
juuied lofceiher by the isthmus just ipoken of. The Minewhat bniiMli.T iiuie-
nor extremity of the orescent, or head of the comma, u called the arU«ru>r
■vmii or ftom: awl the narrower jMsterior cxtremilv of the crotuent, or tail
of ilie comma, in called the pMterior eornu or hora. ^he part by which cscb
hnra i» joined on to Ibe middle part of the cTMceiit ia calkd tbo errvir, ante-
rior and poaUnor respectively. The i»Lhinits joiitiog tbe back* of ihc two
croKvat*, likt the oreaceota theiuielves, coniisui, for th<! m<wi part, of {rntT
natter, ibc buod ruuoinf; potterior ordorval to the cciiiml rannl bcinfc cailea I
tbe jMifmor ^roy nmimunurf {P'ijt- 1"4. P- 7- ".). »nd ihc band riinning ante-l
rior or ventral to the canul being cnlleii tne atiltrwr vniy oommijuurt (Pig>t
174, a, g, p.), Tlw iKNtU-riLir li-uure toiiohM the pocUnor gray coiamiMun^ r
but the *nt«rior gray cotimiiwurr- it wparated from the bottom of the ante*
rior gray commiMura is soraintnl from tbe bottom nf tbe anterior liMoro by
a hood of while nuUter,callc4t the anterior lehiu emamuMtrt.or more -imply,
ibc trfiite etimmimurr, or somclim** llie •inlcrwmmmi4»urf. (Fig. 174, n. c).
If the Miction be taken at the level of the origin of a pair of spinal nervns,
il irill be MCD that the anterior or ventral root, piercinis the white matter
onpusitw tbe bead of the comma id several diaiincl butnltca { Fig. 17-1, A. r.),
tUDnges into the anterior comu. while the nonerior or doraat nait (Fig. 174,
P. r., P. T.I. having tlie appearance of a rinf[le undivided bundle, piL*M«, in |uirt
at loaM, into the fitnUrwr horo. Uotb roots are dbpened length niir« iilong the
<ord, the hinder nxita nfone nen'e being clnoe to tbe fbremOMt rootaof iho nerve
WInw. butil is uiilytlx- anterior rixit^ ntiich arv disserted sidcwayt. Tti>< com* I
pact bnndli' nf tbe poMCrinr ruoi dividiv. with tolerable sliorpDess, the white j
matter in vach laiernl lialf of the conl into a [xjvterior poriioiii lyine botweeB'
tbe pii*leri')r titnuro and the potterior root ( Fiti. 17.1, juitt. eol. ), which portion
Btnci*, •• <•'(- ■hall 'vv, it runs in the f-trm uf a mliimn ainug the Icngtli of
the coni, iscalltvl ihe/iiMterior eo/umn.and into a portion lying Iotheouldd4rJ
of ih*' pintoriur rnot between it oimI tW anterior tistmiru, called ihe unCenM
hlrriil ftlMMn. This Isilor may bo eooitdcred as fiinher divideil, by xhtn
enlrancfof the siilcriorroot;> into m iat^rat oatumii (Fig. 174, b/. iy>/.} IwtWMnl
tbe [Hwterior root and the moHl external bundle of the anterior root, ai»l int»]
an ualrrinr fJumH (Fig. 174, 'iir(. Co/.) between ll»0 anterior fiwiiro unit the
moet external kun<lte of tbe anterior root. The {Htrl trarerted by the bun-
dlea of the anterior root, as they make for tbe anterior liom. sooordin^ly
helonirs tu the anterior column ; but tome writers speak of the nntenor '
oohinin a* Iving hetwccii the anterior ItsNirc and the nearest bundle of
anterior ro'il, thus making ihe rei^ion of the anterior root belong to neither
auterinr ui>r lateral column. And indeed (he diolinction betwceD ihe ant
rior nud llic lateral cititimn U, to a great extcnl, an artilicial diatinclion.
1 064. Thr " while matter " ooiuaBU exduMvely of meduUaied fibres sai
ported (larilv by ci>un>H'tive tissue and partly by a peculwr tiMue known :
neMfwT/i'n, <Ywhich we ■hull presently st>eak. Tbe Bbres are of variotnsix
biUBHiny of ihviH are lurgv, iuhI m all of them tli« medulla is cuntpicnti
m
THE SriXAL CORD,
Tbey run for the nuMC purl luii](itt»linally, w> thai in (raimTene t*t%io
the cord dmHy i1i« wtKili; of tli« «faii« mailer a]>peiirH uiider the mW
lobacompOHtlof iniuulircircloi', tbo traiuivt-FMr tH.'<XKiDi> of lltp loiigilu<^i <
diapuocd Bbm, imln^dcil in tlio Mi|)jM)rliu]i !>[ructurt». The " gnr ti.^n i ■
also coDtaina metlulUtifl lH>m, Uit thi-M- nrtr for ibe ninat |>nrt 4.-s<r"li:ij;
line fibres pomemng a inedulla which n|ii><Mire lo diRir frtitn Ihat •■fu
ordinary m-rve'fibrp, «*□(« it dot« ik>1 tXnhi n-iidilv with oatnic acid, but »
rcmiered visible bv s)>ecial modea of pniparalioD each m tbai known u
Weigeri'e. Hcnci- ilivM fiiw Itbrea are not appnrtnt in ••rdinart- camuMm
other apeciinau, and indeed (heir presence wa* for n long time uverU-kpl
Beaidee tbeae fine mcdullated fibrw. if «l' ntny cull Ihen stich, the em
maUer caD(&in», vbal iW while matler does doI. nerve-oells irith bnuxniog
prooe««m, nak^d aiie-cvtiDdere, end delii'ale lilaoKDls ariaiug front tbt
division of axis-crlindera or Ironi the bmnchiiif* of nerve-cells, all thl«
viiritius atrudures being iinbeddett in iteuroelia. Uwing to the relatir*
nbuiMlanee of the ivhile refractive medulla, llw white matter pcaaiam in
(Veak apecimens a oharaolerisiii;. opaque white color ; hence the nanic Tbt
gray matter Irom the relative acantinees of medulla Itas no woh opaijiF
whitenew, is much mi>re transluoeni, and in flreah specimeDH hu a gny <«
rather pinkiitb-i^ray color, the mlUh (iut Wing due tti tbe pm-ciKv i«rlU' of
pigment and partly of hloud, Tor the bloodve«els are much mure abuDtlul
in (he gray matter iban in the white.
Tbe jtin inatcr which eluseJy invents the cord all anMind c«in»i*t' nf iiah
ncctive liicue, fairly rich in elasli*^ elenn-nls nnd nbiiiidanity sLipjilinl uilb
bloodvcHcIs; it in indeed vsMcnlially » vn»-iiUr mfmbram- nrnl (iiniisbte iW
nervous elvinuDlN of the curd nith their chi«f supply of bhxid. Il wnd* ia
m intervals parliuons or MpUt of lUr same nature as ilsclf mdinliiiK I'luanl
tJie oeotral gray matter. The luirron' ikmIcHot tusure is completely fillnl vf
by a large Mptuni of this kind, indeed as ue have said is in reality not a
fieaure but n inrm t«ptnm : but the nnterinr fissure ii too wide for sueh m
arrangement; the wbute membrane dips down into this fissutv. f!)U'iiiiai; lli*
surface of the cold and being reHocted at the bottom. FMni lhe«e pfi'tiur<
aepta, secondary 5ner sepln still compofcd of ordinary fibrillnted cutit
tissue, carrying bloodveHcIs, branch off; liut these are soou uierL'ol iiii
|ieculiar supporting tissue called, as we have said, ueurolgis, 'Ibii out"'-
in the first place of small branchiu]* cells, lying in veriowi plnnra. i^
branching in exeesuve. so that the budv »f ibe cell is reducrd to very twall
dimenniuns, indeed at times alnnst ul)liieraied, (he nucleun ili)Ui|ii]ieit(iB|t
w hilc the uumAous brancbett are continued ns loDg, fine rdantcni* or (!<>■**
|Miniiing a devious but for tlie moat pan a Itingiludinal eotir^^i. In lie
aecvud place (bfse cells and fibiw or filunicnls arv inilHiUicil in a lionw-
SOCOUS gnutui substance. Relativclv lo the librvn and ground siih«uaR
I bodies of the cells (which arc culled IX-iler's evils?, e«|N-ciHlly bri-liw s>t^
as bear obvious nuclei, are very scuuty ; hence in sH'tions. egpe-HsIlt la
transvenw sections, of tbi^ coni lite neuroglia has often a dotteil >>r piind*)'*!
Bupearance, the dots being the (ninevem Mclions of ihc Hue, lon>:iiu<liiMlir-
dupoaed fibres imbedded in the ground subMaiwe. Examined chentialf.*'-
the neuroKlia is found to be oompOBed not like oonnective tismie uf pUU'j
but of n substance which appears to l>e olosely allied lo keratin. ihctM^
r(>n»liiuei)t of horny epidermis, hairs and tlie like, ^ 43G, and nlikh
tliercfiire been called neurotrratiu i see also § OS), And indeed this dmir^
llitiugli like c^neclive tissue a siimxirting structure, is nut, like et'naHii'
ti»ne, of racaublnslic. but of epibliutie origin. The walls of ibv DM"
canal of the enibfyo which are Irauxformvil into (he spinal Ci>rd of th* *>l*|
cunsirt at finit of epilhelia), epibliuAM oelli ; nwl irhile some of ibxc i
'.sill
THE SPIXAl. CORD.
ttccome niTvoii* flcnwtii*, mbcra bocuiue iietiroglia. The cjiiihdiiil cclb
wbich are i)«»uii«il tu Ibnu iieiiroglui becutue exi-Milinglr linincbtil, vrbile
their origin ully prmtojilntimic cell »nb*taiKc become* truuMlunucd ton Inrgo
esftil iiitd nciirokvniiiii.
The Bciinwlim liltii up lb« ii|iiicM Wtwrru iho niiliAting Inrgcr iwptnl pro-
loogUMD* nribe jiiu niulcr nint llirtiiirr bniiic'hi:'il ncpla vrhicn (tntlitig frvta
th« larftvr nnn cnrry minuiv bl<i<>lvi-:«M.'l« iiiLu the iotcriur qf the wbiie
niAll^r. In ihtte *\mcv* it i* m> nrraiig>.-<l ii« to form rl«licn(« tubiilnr canals,
of nty vnriflbli- *iii-, ninning for lliv niuBt rinrt in ft loDeillidinnl diivclMMi,
GkH of Uu-M- iiibuUr ninaTs is neciipii-i) l>r and wbollr fillt't) up n-ith ft
■odallated nvrvi-libmor cnrmpnniling fir«. A medultatwl ni^rvc-librp "f
Uw wbiu nutUir of the Bpiiinl cvri) rc>feniblr» a ini^lullaUNl iK-is-^Hhre ut' h
Mrv« (( SK) in beini; coinpoM^ of sn usifl-iylimkT ami a niedulln ; but il
pCNBonM no ]>riRiitive *h«nlh or neurilenimn. This is absent and indeed is
aoc wnntMl ; the tubular »hcatb of neuroglia alTordF in the «pinal cord tand
as «« shall see in the cenlnil nervous sfstem gvoersUy) the 8U]>port which in
• iMrv« m afloHed by iIh neurileninia. Nodes are, according to tnost authors,
abeeni, but some say they are preeeuL
The while matter of liie curd consists Iheu of a more or less solid inaaMof
■Muroglia, hitviuy ilie stTueture just described, irbicb is pcrineated by minute
oaala, aotue exceed inj-ly Hue Hnd carrying verv Hoe 2^ fibres, other!) lur)(er
and rarryinjc l)bn.ii U|i to the ■lie of 15 ". Thu moss is further brukeii up
into areas by thi'sinaUvr and larger vavcubir conneclire-liwue M;])ta with the
cdgt* und ciidiugw uf whicli the ntHiro^lia in M)ntiiiui>u>. Most of tlw iK-rve>
flbna, ax >v hiivc said, run longiliiilinsllr and in a transverse section of the
conl nri' cut Iran.iversely ; but ns we shall aeo fibrts are cuntinually pCMiig
iniu aiMi nut lit" ibe while Rinltcri nnil in sij dtdng take a more or li^m traoa*
ver>« cunrH.'; ihcM', bonrver, arc few oompnred witli thi<ie nhirh run in a
lon^ludiiinl ilirection. On the oulaide of the curd below the pin maler the
neuroc'i' is developed into a layer of some thicknwi from whii^h nerve-fibres
■» kiMMit : this is often siHikcn of as an inner layer of the pia maler ; but
being netiruiglia and not connective tissue is of n difliircnt nature Ircim the
|iui maUT profier. A layer of this Mijwrlicia] neuroglia also accomjianies
ibtf larffer KiKa, and a considemhle cjuaQtiiy is present in the lar|^ se)Kuni
tallrd the pusieriur fissure.
The |>ia mater carries uot only bloodi.'eMela but also Iviupbaticti : of these,
homver, we lihall speak when we come i<> deal with tiie vateu la r arrange -
BMDU of the whole of the central nervous system.
( 566. In tbv m-ray matter ne may dBilinguiah the larger, more eouspiruiiiiB
aenrr-fvlli' and the rntt nf lh<- gniy maiter in nhirh ihcse eelU lie. We
bare alremly H dU) devoribed the general features uf theae lai]ger nerre-rells.
and sliall have presently to speak of their ;i|M.-cial characters ood groupiug.
>l)-nnwhile the mwl important jioint to remember about tbem boiMi.? tlH^
lJS4-t llial titey vary Inr^-ly in fonu and size t* that while one procna may or
doca become aii axi^eylindiT of a nerve-fibre, the ulbers rapidly hraiicli, and
brwkin^ up into fine nerve filaments are luft to view in the re^^t of tbc gmy
matter.
Tliesp lari^vr nerve-cells f<im), boircvcr, a part only, aud in moot rvj^on* of
v cord llui siiudlor part, of Ihe whole gray initllvr. In a Irantverse section
Dm the tlionieic rvginn ' Fig. 174 ) ■ few only of tbcsv larger n«rve-c«lli> nru
in the whole sex-tton. and thoUL'h they awiear m<>rr numerous in te*.--
I fVoni Ihe cervical and «<>()ccinlly from the lumbar region* (Figs. 167,
177), yet in all coses iliey occupy the smaller purl of the area of ine gmy
Miter. The lartrer \t»n ol the urny mailer consiato, bvM<les a neur>;f;lin
npirarting the nervous ekoMOta, of nuni'e filameiila running in various dirvo-
i
680
THE SPIX3
lionn nnii funnitijr, ivol k pkxuA projiwrly s-^mlk-il. but ni) iiiicrlvwmmi "t
exiretiii' <'i<iii|i]exily. Thf«e filamenlfl are, on llie oik' Iihih). llie liiw ntnlul-
Iwwl litiriK -ijHikMi I'f ab'ive lu beiiiB rvL-ngtiiMi) with tliffii-uliy, nnd. i>n tfa»
•Aher linuil, m in -nii-ilu listed filamuita niiiKiiiK Tr^ni rmrly nidc snH coD-
a|iiouoiin nakod axi>-cvliii(Iirn> iliiwn U> tihrib of cxtronto cvnuitjr, the UUer
nrisinc iippnrently Httii-r from [h« <livii>iiii) nf nKiit-cylltKlen' nf ncrre fibm
p»»inj{ inl'i iir tint of tliv ^Tay ninttvr or Inim the cviniiniii^l bra(i('liiii>: of
[tr\KV3sat of iiitrve-i'elljt Ity llir mniUv nf |)fv[K(riition i>ow nvMilnblc it hiw
Mi-n iihim'fi lliat tlii« ()ii« nH-dtjIliitt-i) tiliriv, «(i fur friiiti h^ng nir*'. an in
«-rt«in \mrlr nf tlip fcnv ninllcr nn Mlxintlnni im cvem Ui prtpoti<ii*n»lv ovrr
lli« iioii-in«'ilullnUi) fibni' or tibrilR. T^slly. bciqiln the conspicuous nvrrp-
<'i-H» jpnkrn of nhovc. which, though of mrinus nies, may nil pi-rhap be
i'pdkrii of lis Inrgv, a very Jar^' niinil)«r of other cell* of •mall riw. l^•in^^• of
which ni nil events mii«( be rcgurHpd as true nQrTe-c«ll^ nn> prvKnl in the
gray matter.
The neiiroglin in which all thee« Mniclum, nerves-ell-, fine tneiliillntpil
nerve' tibree. nnlccil axis-crlinders, and line KlnmeniB are imbi-dde<l, U iilcttticnl
ill iT» '.^eiieml chiircierti with thst of the while mutter, Ihii, iif^ iiaiiiriillv fol-
lows fVi)m the naiureof the uervoiis elements which it «up)Miri8. iit dilli-fviitly
arran^l, Inxlead of forming; it srtilem of tubular vhaiinels it tak« nu iIm>
fonu of a Biionj,'e-work willi larj,'*' spacee ll>r itie lar^r nen'e-eolU and fior
pa»a|ies for the nervoiia filaoienla At the jiint-tiiiii of ihe gny mutter vith
the while nmtter. the ne»ro)[lia of the one in continuous with thai of ih«
oiber, and the ei>imociive-ii«ue sejita of the latter mn riKht into tbefornxr:
the iniiltne of (he firny niatier i» ii<tt iiniiKiili and even, but l>mkei) hy loolfa-
like proccoKadue to the tiepta. .Since, m vre havejiut Mid.aome of th» IrtM
nerve-cella are very small, and nitice the ncrre-filainenut like ihi< nmm^lia
A\iTv* arft vn-y f!n« am) take lik« iJiom an irn-gular nwir»e. it ofiva )>eci>mM
wry difltcull in a Miction to determine exactly which ia neuroglia and whlcli
are ncrvou« clcTne»t«. The ncnroglic orlU may. hone\'cr. bedifitiD;;uish«d |wr-
hape from the smaller nervc-cell» bviheir nuclei not being eo conspicuoua or
»o relatively large ns in n ncrvo-celf, and by their siaiiiin;; diflerentlr.
The gray matter then may lie brriadly described as a beil of nearoglia,
cotriaimng n certain number of bnuiching nervc'celli). for tlw muM |MR
though not exclueivelr large and conspicuouti. but chiellv iKrupied by vrlinl
ia Dot so much a pfexu^ as an intricate inleriveaving nf nerve- li lament*
ninninji; apparently in all directions, ^me of llitae ftlamenta arefuirlvcm
xriicuoiin naked axi»f cylinders, and a few are easilr reco^roiied molu^la
Sbna of ordinary iiw?; but by far the greater numtier are either exceiidini'Ij
line medullated fibres, wh(«e medulla is ouly made evident lir ft[iccinl m>Kt«»'
of prrparalion, or delicate fibriU devoid of medulla. With the D«rvou« web
formed by ihc*e Blaructitji the brauohiog proonKS of the oerve-eelU, on the
one hand, nud the divitious of nerve-Hores pa»siiti( into or out of ibt* gmr
mutter, on tin- other hand, appear lo he conlinuoua. It may l>e «i)detl that
(he grny matter it well mipplied with IdooiIvcawU. tbeae }mn]i; in it, nt atnteil
nb).)ve. ndnlively much more numerous than in the while mnit^r.
r i S66. The cenlmt canal in lim-il by a Mn^le layer of columnar epithelial
eel Ik. which are generally dcccrilicd nu beariiit: cilia; but i[ lit not i.>3nain
that the proccMw which may Iw «H'n pnijccliug fmm the mrfaoM of tbe
rella are really cilia. Thew epithelial celln rvA not <>n a diMlincl boaemeni
membrane, but on n bed of neuroglia, free uppurenllv, or »<<nrly tn. frmi
nervoitB elemenie which Riirroiinds the cenlnil canal anif i» »omctimt» (iio-keiii
of n» the mhil'jitli'i 'iflntinota nrnlraiit (Fig. 174. r. t- ','<■ Tlio atlnrlteil'
basM of the epithelial o-ll» are bnincheil or taper to a HInmrnt, and 1>erome
CoutiuuouB with the brniieheil celU or hbrva of llie neuroglia lieloiv. As we
■nta
I
I
I
I
I
I
TIIK STKUCTUnE OP TItK SPIXAL CORll.
•kid ftbow, the nciiroglin I'liMncnbt iirr lrnn'<f<>nDC-il vpUhcltjil celti ; mk) tbe
notiDalir attim nib. which n-lninitii^ l)i*' cliiiriiitli^r* of uiiithdiBl oelb form
% Icniug t» lli« miih), trilti lliv i'i;lli> nhiuh liiivc becumo unmched xud loM
thrir *'pith<'lial chanicli-re, iii'licalM llxi i^pilhclinl origin of (hr litli^r.
Tb* cviilml nuinl nilh llir rurroiindiag amof nniroslis forma tlx! twotral
jttfl of ili« )*tliiiiU8 iiiiilin)^ (ho Iwo Inicral hnlv« of thi; conl. Fiwicrior
iiarm]) U> tUi* cenlrni tnltm lir« ths pn4erior ymy eommiimiiY^ (Ftg». 174,
liO. 177, p. g. (^>, ooiopoMiJ i.'hi«riy of fino fiUmenU ninniti}; trnnavcnelf,
amt uilvnor (v«ntnil) to it linn lirat Uie tliiniier anUruir gr-iii eummMar*
(FisL 174. 179, 177, a. 7. ir.) of a einiiUr naiure, tiiiil ihcii iht rrlnlivclv
lU» irAi/r rtiaunt^art (Fi^ 174. 176, 1 iT, 11. e.) irhicli ta formed by imwlul-
IMmI libm criisiing over froTii oae side uf ihe t.'onl to the other, and lliu»
OMHthiilu M dcci»iHil)oD of tibrce alon); ibe whok len^'th of the mnl. On
tMoh aide the central mass of neuro{>lin of which we are aponking enidually
DiMKW into the central yray otaiter of the eorresiioDdiDK Uterul lialf,
Tbe end or bead (caput), aa it in freijuently call«il of tbe pueieriur born in
•>ct^pie<l no< hy itnliuiiry tcray matter, but by a {wouliar liaaue, tbe tuiManlia
yrtMiinntn of /totiHilo, wliic-h forRU a aort of cap to the ruore ordioary grav
matter, but dilfi'ia id uu and shape in diflVrent rejjinns of the twrd. ( Cf.
Pisa. 1 74, 1 75, 1 7<t, ». y.) In carmine and Aoine other modes of prepamiiaii
it b fVvqueutly iiained more deeply than it the ordinary crar matter, and in
siich premraiionn i* very ootiKpicuous, It niav be deaeribed aa conxiatiiis of
B MHDewhnt )Kculiar ueiiro^Hn iraveracd bv liGre* of the poMerinr nxrt, aod
ctNitainin^ a larj^v number of eclU irhtcb, ^ur the moat part Nnall. llie cell-
biKliea betng small relatively to tlio itiiclei, are not all nlilte. unme lieing
pnbably nervoui and others not. It iak<« origin from tbt- cclU formiuji (he
immediate mlU of the embryonic mmlullnrv canal. In tbe embryo, (hii
cwtai b rolatirely uide. ihongli coniprvswcd rn>m Mdc to »iili-, and in tran»-
tvne wcUoni of ih«' miyliillary tube appears at a certain «tftce lu a narrow
oral ulit nlace<l vertically and reaching atmodt from the donallo the va'Dtrsl
turfkce. The donal pan of this lon^; »lit is later on ct(»cd up by the coming
tggetlMr of the iralls and tbe obliteration of the i-renter pari of tbr cavity,
baring the ventral [tori to furiii a circular canal, tvbicb bv tbe development
of the anterior columns assuraea Lli« central pOMtion. During tbia clo«ire
of tbe dorsal part of tbe canal a mass of the oelb liDinjt tbe canal is cut
flvm the r<~it on eacb ude, and during the aubseqaent Kruwth takes up a
fwsillon ai il»e end of the posterior horn. Hence, (hou^h it never appar-
ently oiuiains any cavity, (be gulistauoe of Kolandi) may be regarded as an
ianlnted ik-nion of (he wnlU of the n>eduUary canal, which ban undergone a
developmeni «nmewhnt diflereut from that of tbe |Kirti<)n which remains as
Hm lini»i( uf tbe central canal. Trace* of thia origin may he acen even in
tbe ailalt. Tbux, in the lower end of the oord, in what we »ball tpenk of
prvacntly n the ooittu writHUnnt, the central canal widen* otil out domally.
and in leclion (Fig. 17o, A) priwnt* on each itidi' 11 Iwy r xtrvtching out
towanl tht- piuiliim of ibc [HMtrrior born. At ihi* region of tlie oord, thMlgh
tmtb while and gray malti-r iircdcvcloitcd cm the ventral »iirfaoa.lh4 iWMerior
ooluunsdo not meet OD the dorml nirruce, hut lwivetl>o centml canal covon^
enly by ti»uo which perhaps may be called neaniglia. but u of jx'cuiiar
natun- aiht origin. In (he cidf, in a pari of the dorsal region llic «ubitaoc«
of Rolandii is not confined t<> the lip of ifao |>oitleri»r horn, but is oonttnood
tu mosit it« fcllofT in th<' iiiiddlo line. (Pig- 17-5. It.) If wc imagine the
doml portion of ibe caiud of .\ to be c«t off from tbe ventral portion, ita
eavltr la be oblilcntcx). and tbe lining epithelium with <ome of tbe enr-
niunding elements to utulergo a H|)ectai development, the condition in B is
ranched by tbe growth of the posterior calumiw. From B ibe transition to
482
TilB BPIHA1. C0H11.
Uw normni state or iblnei n !ii Pig. 1 7o. r, U b very slight une. The cxtrtiu
dofMl tip of the lioni, hoiDK of a more open texture ibMi (h« Fulnuor* a/
Rolttndo, in somi-limiH culled tbe imn tpon^wta.
no. I?*.
A 8 C
tnwiiua TO iLU-fiiiAii mti Naivki! or mt iKmimn •)• ltoti!>i<o.
nellinmftMTiiireli ^lamwBiuilonilatvn-iidmwii i« Uw auaa Kah. liiHll ihnviifiraiM
gnqi OMWr W (httdad aiUi Hln Unanikl Uw wbtlv loMMr villi ittn.
A. truwnnc wMloo of the Iuboc md or llie enaai ««)l«lbitt> Id bib: r.«plk*tliuBbMa«ir
Mtdollur ouuU ; r. IsMnl cx|niirino oT Un ouul ; B, uaMnn* n«ltani»niwa|4Ml«M4"riki
valf In Un hmor ihcmck K^«a: r,nbHU«eaf Motentoi cMMntViMl: <:, imbmdm mctm
thioush mld-fkBCHtlr rcfloa irfatrd la mui.
iSM7. The grouping of the uerw-etlU. The Derve-cella. at all ereautlit
nils wbich are larj^e coough to be eiisily mid vriihoiit doubt reco|{iuteil M Ik
Derve-oella, foru, tut vre have seen, only a pHri of the (^y matter, anil <<■
WHDe pans of lli« oord, in tlie (liomcic regioD for instauce, ara »o Hfmm thtX
in » section of tl)« spinal cord in llii» region thin eaougb to shnvc it* liitiu-
liVi<'Jtl ft^alUKft »ali»fiu-li>rtly, th« bodies of a few only of oucli celb an: vittiUt
(Pig. 174) ; tbe grailur jinrt of the gray matter comists itot of tbe Uolin i^
coDipicuoua Do-vc-a-llti. but of a inai« of fihrea and Rbrils pawiuft apiurtsilr
lu an dlrcctioUM. In the ccrvieal (Fig. 176!, and especially in the lunb^
(Fig. 177) rvgiona, tlw nervii-cella are bolh aliaidiitely and rvlmtvcly nwr
abundant: )>ut even in a »<xtioD Inlcfn from l)i« luiutiar region tbe Der%^
cdlK. all tHit logctbcr, form the uualli-r |Virt of the ubule area of v^r
matter. Moreover, in mpoct of th« number of ocll«. all LbeMcti»u« ''
the fame region of the cord are not aliko. Seeing ibat tbe cord niii> '
Htilencd a> growing ctit of tbo fusiiui of a aerie* of jNiirct ganglia, eiuli ft*-
elion carrapondtRg lo ■ nerve (<■/. $ 96}, w« may luirty cx|ii.-<rt to fiud !h>
niiion not complete, »> that the narve-celU would ap]>ear own ni: <
opposite a nerve than in ibe middle betwitMi two nerves. In sonti <
lou'or animal» ibis urrangcnicnt if woU ohvtou*. and lh«r» ar« liome n«««'
for tbiuking that even in man the nert'o^U aro mtumvrkally iucimunl ti
the level of eaob nerve.
Even ubeu caaUHlly observed, tt i» obvious that th« nerve-cells atv a**
acatured in a wholly irregular manner tbroughoul the gray malier. Ixuu:'
ibr hutAuco, much more conspicuous in tbe anterior born thou I'lievVn'
and nwro careful obser\-«tioa atlon-« us to arrange Utetn to a cvriuin rxtriit
in eri>U|M.
The etU* of the uutmor hvm are for the moat jwrt large and i:ciiupicuiiii*r
tiTA to V-i^f in diameter, branob out in various direetioiiH, ami pr^^rot m
irregular outline iu wciiona taken in dilfereiit pIaiK-«. Wi- biivv khv |"
ihinK that every one of tliein posaestea au axt»-cyliud«r pn>vcN>. vcbiAiu
llie caoe at all events of must of tlie cells, iiaaaiug otit of ibe firay aa,i>t
beoomee a fibre of the adjairenl anterior root. They are ubvi<tii;i mi-l
apioiiOUB in all regions of the coni, tbiwigh much iiuire uuniei»ut nml !■' _
vidually larger in the i.i.T\-ii.Ml aiul liinil>areiiliirgeinenu than in ilif ili>v»^
regKMi, We may lurtliur, uith greati.T or lev nmcoau. diviilu ihviii ">'"
teparale grutipa.
m
THE STRCCTUBE OF THE SPINAL CORD. 683
FlO. ITfi.
CEKIKMI. NKMVE. ll^llh:i!HlM.IlPN,i
This If drswn on (he unine 'cHit n^ Fli;. 17), ilml i- iiinBriilliii lin.i'n liiiu'i'. r./. I.. liiliTal rfliriiliir
'imatlon -,r/.p,, ptwleriiir reilciilHr fiiriii^itliiri ; ji', liiw iihivs •>i' laiiral liiimlk-'pf !)»■ iNi.itri.pr ri«.i :
'.p'^, fibres ormefLlan bundkporiri)giori4»rnH»l i-iiterjii^' LTny iiintlvr frjpin I'Tticriiiil i-nsit-riort'oluiini :
gr^y matter of pi«lerEor hum ; .^;j,'i., l>iLi]i1]4.'«nrhNr4.'s U'li'M^lh^ tip Ihu ^|<ii)iil tim'<-s-'ry inTxt- : in
W laUiBl rettcular formalinii thvv arf "ifii Mil IniTi'it-r-ily ; I,, i- a miiiiml -f|iiimi ••( [ijiuii-i'tln-
■sue mmrklng out the rerebtllftr inifi ('. T rn.iii ihi- (■r.L--i.1 |i;-niiii!iinl ihtii i:. 1', T. : ;. -.. /i.ii«
WD^on; 2 T , ,}, y^ iBteral <^tr]]« "f llie iiiittTl'ir ikpni : r), cf-Ll- in Tlit rt-L'Upii ipt Ehi; liiii^rHl celi* iilrir
pfniBtlon. The other letter" of re(Vftii''o i\ri Uk- i^piiipij ii'> in Kil'. 171.
684 THE SPIXAL COBD.
Ill the cervical and lumbar regions a fairly distinct group of celts la seen
lying on the median side of the gray matter close to the anterior coluniB
(Figs. 176, 177, 1). This may be calfed Ihe median group. It appears alsx
in the thoracic region (Fig. 174, 1) ; iudeed. the question arises whether sU
the celU of the anterior horn in this region do not belong to this group. The
other cells so conspicuous in the hirabar and cervical enlargements, and ihire-
fore probably in some way associated with the limbs, may be spoken of m
forming altogether a latent/ i/roitp ; but we may. though with some uacer-
t:(inty subdivide them into two or three groups. Thus in the lumbar regiim
a group of cells (Fig. 177. 2;) lying near the lateral margin of the more
dorsal part or base of the horn may be distinguished, as a lateral nub-^roup,
from the cells occupying the ventral lateral corner of the horn and forraing
a vriilrnl or iinterior siib-yroiip (Fig. 177, 2"); and the same distinction.
though with less success, may be made in the cervical region (Fig. 176).
Further, we may perhaps in lioth regions distinguish a group of cells ploced
more in the very middle of the horn aaa central etib-ifroup (Figs. 176, ITT.
'2 ,J). But, in all cases, the separation of these cells, whicn we have spokea
of a^ a whole as lateral cells, into minor groups, is far less distinct than the
separation of the median group from these lateral cells, especially if 't
admit that in the thoracic region the median group is alone clearly repn-
sented.
In the thoracic region a group of rather smaller cells is seen at the hue
of the anterior horu, near to the junction with the isthmus (Fig. IT-l.J)-
In the cervical and lumbar region these cells are very scanty (Figs. 176,
177. 7).
The relU of the posterior tioni contrast strongly with those of the anterior
horn in being few, and for the moat part small. They are branched ; snd
though we have reason to believe that, like the cells of the anterior hon,
they possess each an axis-cylinder process, this is not easily determined by
actual observation; the processes do not run out to join the posterior root,
as do Ihe corresponding processes in the anterior horn, and therefore are not
so readily seen. These cells occur in all regions of the cord, and appear to
bo arranged in two more groups. The lateral margin of the posterior hora,
at about the middle or neck of the horn, is along the whole length of the
cord, but especially in the cervical region, much broken up by bundles of
fibre.'' passing in various directions and forming an open network, called the
I'tti-m/ retieit/ar fonnatimi (Figs. 176, 177, r.f. (at.).- In all regions of the
cord a number of cells are found associated with this reticular fbrmation,
forming the group oj the lateral reticular formation (Figs. 176, 177, 5|. In
all regions of the cord, also, a group of cells (Figs. 174, 176, 177, H) k
fliund in that [wrt of the horn where, a little ventral to the substaoceof
Rolando, the uniform field of gray matter is broken up into a kind of net-
work by a number of bundles of white fibres running in various directinn*.
This network hu.-* also l>een called a reticular formation, and has received
the name nf poalerior reticular formation (Figs. 176,177, r. f. p.) to dn-
tintruish it from the lateral reticular formation just mentioned; the t«o.
however, in some re^itms (_see Fig. 174) join each other, and thus cut off >
vcDiral portion of llie iw»terior horn containing nerve-cells from a doww
poniim. .r in Figs. 17(3, 177, in which no obvious or conspicuous nervcHrelli
arc preseiK.
The grouja of cells jusi mentioned, with the restrictions and modificalio^
S|jokeu of, occur along t!ie whole length of the cord ; but the group of cdll
to which we must now call attention is almost confined to a special region oi
till' curd, or at least is hut feebly represented elsewhere. In the tnoiMW
rc;riiin, csjiecially In the lower thoracic region (we shall return to the lim't*
W groub lilcr on), at llii; bn*e o( t\to iK^terinr horu (Fig. 174. S). jitet
|bl lo tlw curve rurnivd by llio |K^tvrii>r tfrny conimbeure tm iliis btiiils
B-vuiniii. SKTioii or nil snuAi. comi iBnui) «i tbr Lmi. or tuc Tiiir»
I inUie wDw Kvleurip. ITI, ITl, kiiil in iM «in« njr. rimpt Unl Ike oiKlto* ol
rM Doi runcnird, If. mdlui : rr. IMCfBtiUMe ; IV. latanl bia>d>« «< tmoto'
nn niQi|iili>J mdft ■.!. t> tb* kmvImI •oO or Umbimi'i mm. TteMhcr IMlai*
I »• Ik* MB* M in nSK 1:4. ini. The iktt* •gunv. i;«. IM. 177. an InMwfad lo ItlWBU*
{■dlAraiiUkl ••urn* of itectfiloBl. ilKinwk. aad lumliK conL
|Ijr to joiD tli« p(W«rior bnra, b ■*«» 011 neh tide o( rbe mrd a con-
(oi groap of rcllt> kliiiwn lU t'/nrit't roJumn, or lli« porirrinr rt-fiailar
jjn or vfrimhr rytiiidtr. TUc celU cuimiotiiig ibu group, tliougb vary-
uie •! diffli^rciit Icvvl*. are rnihar lnrg« c«lU, nnJ nra for tli« oxat
666
THE SI'INAL CORD.
[Ktrt (uBiform. irilli tbejr loiifr uxi« ]>1it«xt leD);tbwBys hIoiik rliv
(bat in iranEri.'rM) sectiuu* thvy olti^ti n|iiN.-«ir to have a ralli«r «inall
body. Tbc.varc sDrrouixIctl by. Hnd as il were iinbeild«<l iu, n nuMif fai
fibres, the mv& of which is itii)i«it«d by a dott«d line in Fij*. 174.
Abo coiispicuoiJB in th« thorade n^^ron ia ai]Oth«r KToup of c«lls liiiij m
tJie outer Mde ot' ih« middle of tb« gny nialt«r st abutit ihe junction iif lU
Ulterior and pneterior boras, l^liia is Known as the intermtitia-Oaer^ trta.
and issomednies called the lateral horn (Ki^- 174, 4). The oelb comiwoi
it are loinewhat small Fpiodle-Bbaped i.'ells »'iib tbeir Ions &%» [lUuij iniu
rertel}'. The iiroup is couapicuous, as we hKve Mud. in the tboncir ncpnia.
it may be reco^iiixed id the lumbar reiiioii ( Fig- 1T<>. 4 >, bul tn tbv nnitti
ngion becoinea (Mkufused irilh tlie mutt <lor»atly placed or Utefnl (iil>-gr«ij>
of the iuiieri»r hum. We shall, bomever. have to roiiim tn tbts* gMup 4
0(11* when w« ooine tu apeak of the difl^mnctt becweco tl>e atvonl k(mii>
of the wnl.
S 568. The tract* of u^hite matter. At flnt flight tha irhiu> matter ft ib
cord nppcan to be «f unifunii nalun:. We can umi the oerve rooti u dt-
limitate the antt^rior, pooteriiir, and lateral columns, bul we appear lo hm
no crilcrin lo diftinguixb jmrts in each o»Iumn. In the cemcat and u)i{*r
tboncic regions of the curd, a septum ( Fig. 174, <>.) in tlie poeterior oiltimD.
somewhat mon- cotitpicuoux than the other H^jilti, has enabled anatMniia h
dislingtii^h an innc^r median iKirCi'ui. iho au-tiuin potlerior toluuin, couhdobIt
called the ponlrro-mrdi'iii column or tvluma of ffoll (Fig. 174, m. p.i,{Ka
an outer lateral portion, the •■j-irntal jtoettrior rolumtK, eoinutonly called ik
poflero-extwnat eotumii or coluiati of Burdaeh (Fi^, 174, «. /i. r, the luml
Ert of which, nearer the gray mnltcr, has, for reaooos which ire ihaB n
er on. been called the pimtrriar ronl-iting. liut beyond tlib ndlhtr tb*
irrej(uUr septa nor other features will enable us to dislinffuisb one {mr of
the white matter as diflereut in nature from another. Nor have wc hiu«t
lucow when with the ^cal|]el we attempt to unravel otit the irhito ouUer
into ■b])arut« strandi. Neverlhelesa we haire oouvincuig evidence tliat tbt
whiM matter i» arrmg^ed in atranda, or traote, or columns, which havs Af-
ferent connections at their reii>ecttv« ends, wbidi behavo difierDDtly uidfT
ditforent drcunislaiicu*. which wc have every reason to believe carry od
diflereot functions, but which catiiiut be seiiimtod by the m^alpcl, bectUK
each of them is more or Ins mixeil with bbret of a ditronni nature mi!
orifln, The e^Hdence for the exbtcnco of Uiom iracta is twofold.
On« kind of evidence i* embr^ological iu nature. When a nerve Abn it
b«ng Ibrmcil in the ombryu. either in the spinal cord or elsewhere, ibt
eawiitial axisc ylinder is f<>niie<l lint and the l«n caamtial medulla u furtntA '
later. Now when the devolupntenlal history of the spinal oord is sludieil it \
is found that, in the Mveral regions of thi^ con), all the fibres of the ahiir !
matter do not put on the mcilulla at the same lime. On the contrary, tn^
oeriain imcts, the medulla of the librcs makes its appeanince early. In otMr*
later. Uy this method it bcoomes pomihle lo diatinjcuiah certain tracts frota
others.
Another kiud of evidence is supplied by facts relatins to the degonerati<
of the fibres uf the while matter. We have seen (§ dS'Ji that the ddgctwn'
tioD of a nerve fibre is the result of the separation of the fibre Irani v
trophic centre, and that while the trophic centre of the afferent fibraa
ihv tfauglioD oil the posterior mot, that of the ellWent fibre* is in aome ,
of tlio spinal cord. In the ease of the eflereni libn^ the di^vneral
might be spoken of aa rfssMnrfm^ from the apinal cord to the muscle* n
other iwripheral orgiuts. In the caxcr of the alTcreiit filin*:! uf tbo trunk i
the nerve, llic degeneratiuu i« ahtu one deacetiding fruui the ganglion dua'
glion dua-^
TIIE STRL'CTL'RB OF THE SPINAL COKO.
6a7
I aktn or oth«r peripheral organ. When, bowcvor, tho mcUaii U enmod
tg\» lUi> {MMvriiir TiHiI iif II npitm) »«rv(>, ihr ilc^ni'ntlinii titkis [ilitM
Cn Iff iltf ncirTi* botUL'i-n the xtH'tiDii mid l)ie sjtiiuil coni, it ruiw uj)
MvliuD to nnil iul" tin? njiiunl conl, ninl iniiy, tliertrDrr. lie rnllM
vernding tlegeDCTBtioii. Tliuit viii may Miy tlinl ulir'n n nrrvr trunk or
b«a m tifvtv root m cut iijitipMely ihtpm, nil thti lilinw, which arc (herebj-
pMrminl ("mm iliWr tmphic wiitr<», <Ii'grniT»t«. When lh» oervr trunk is
livitlnl. nil the (ibrm \tVtv lh« M-i:li<>ii iimlcrgo dacvnding degeneration.
tbt anit'rior root hi.- nit Koriuw. nil tho fibril i^f tho r-iot b<-law iho Kctioti
(Incrndiii^ 'li-)Kii«mtiou. It' iho jKwlcHor mol be t-iit iicioae, iill
fibra of (Im: root bIrivv the M-eiion undi'rgn ascenditig dc^nermtion with
«xc»]ilio(i of 4wrt«ia fibres irhii^h do not dogenorBte at ull, nod of wbioh
r thall »i>ctik Inter oo. •
Wba-ii i)i« *|>tiinl i-ord is cut oerow, for iiwlnnoe in lh« dorsal rvf^on, nil
le libn» of the uhite matter do not degeiierau ^iher in liip {Mrt of the
' above tho section or in tho part below. Some flbtvf, and indeed some
_.. of fibres degenerate, and some da not. Puriber. Bonie tracts defren-
hnUe ill the cord above the eeclion, and thus under^i what has been called
••ceudio^ dejfeneratioD : other tracts def^nenilc in tlie eunl below the
n. and ihu^ uDder)|:u what has been called a deacendioi; de^'eneraliou.
term^ luust, hunever, be used with caution. When a nerve trunk in
i Bcrote. the dc^nerattuii actually defloeotb, in the sense thai the progress
tb* deeeoeraiive ehnu^eti tim)' be traced downward; they beKiu at the
.ioo aiia travel downuiml at a rate sullidently tlow to permit u diBerenoa
ing obwrvet) l>^twe>eu the progress of degeuenition at a M{>ut near the seo-
ftod that of one farther tm. Atler*ccii<mof or tujurjr to the spinal cord,
<<r«Ttr. it U Dol pntnbte l» Inice any sur\t ]>ro^;reas either upward ur <Iown-
ard: iu the tnici> both above and lii^low the nection or iiijurv, degenera-
ioD cither iMrgins «imultAiieou«ly along the whole length af the (It^Drrntiiig
tract, or projimw* along the tract w> rapidly that no diflervucns can be
cd, as Inr as the singe of di^grnermiion is concerned, bclwee-n |>art« near
tbi«e far from the section and injurr. When, for instance, tho cord
ivided in the ocrvioal rq;ioii, subsequent examination uf tlie trmcls of
Ecalltd dcaoendiug degenerattoii show's that tho (^generation is as fiir
rauced in the lumbar re«pou far away from the soction as in the cervical
[ion just Iwlow the section. Applied to the spinal oord, therefore, the
rm detc'endiiig dt^neratioa menu* simply degeneration below the seat of
jurv or disease, nseeDding degeiation means sioiply dtgeneiation above the
of injury or disrate. We way add that tJie nisLoIogjcal fealtiree uf the
ion iii tibrce in the spinal cord are noi wbollv identical uitb those
the deceDeraliou of fibres in a tierre trunk. Thus, the neurilemma with
nuclei oeinc absent from ilie libra of the curd, no pruliferatiun of nuclei
Enlaee ; tbe axis-cylinder and nteduUa simply break up, are nb^rbed,
d (iMppear.
Similar defceueratioDs, asceiiilin); or deaoemliu};. or both, are neen wheu
taectka i> not carri«<l right through the whole cord, but partimilnr iwrta
the i>>rd are eut through or deeply injured. .Knd similar degeueratioiu
feccur a» the c»n«et|iiencea of d!i«iut) net up in partn of the cord.
I In ihi* wav the runulta of Mctions of or of other iiijurie* to or of diseases
tf the aptnar cvn), have enabled us to mark out certain tractn of iIm; white
■alter as undergoing degeneration and others aa not. and. iiiorMiver iMrliun
(nu'la a* umlergoing duncciiding and othcni aji un<lcrgoing ascenduig d<^D-
ntion. Kiirtjier, the delimitation of tracts of while matter by llie inocns
if degeneration agroai so wnll with the mult* uf llie embrt'olitgicol method
mci, o
L
688
THE Srl.XAL COKIf.
m to IcATo no doubt ihnt (he white nislt^rdoescomiAt of tntcU wbka
(Void mcli <Abtr in uature an<l in function.
TIm Hcvt^nil iracl* iliiii itniimicO vnrv in diflmDt regnoa of ibt conL
Tboy auiy Ix^ t)^!^!!}' ilcMi-ritieil a^ t<>ll«w9;
I. HtK'tt'iing trafi*, lliat is t<t uty, trai-tB which uiHlergD & dMcendiBf
drgi'iicrittioi) in tliu »on«u nolcd nliovr,
Th«i nitwt ini)H)rt«nl mn<l cuiuiiicuouii U ii Urge met (,Vig. liR, cr.I'i
oi'ciipyini; the ]Kmlcrior i«art of ihe JAifrnl column, ci>Kiiu|i cl'-- ■'- " '•-
oiiltr timrgiii of th« pixtitrior born, iiixl for tltv mutt |iart ugt
surfiii-c of the (»>ixl. Wo oluill htivc lo rrliirn to thi« lni<-t miir< >>>.
mul ramy horc simplv »r thnt it Is nioNt <li»tini;ily tnarki-tl out Iit I>
enibryulot^it-at und tlie drgvntnilion mvlh'xlv, (liiit it ai»y he Irnnii \
Ibc whole length of the cord from thr top of tW i.'crvionl region l<i ih- < 1 1
of Ihe wicriil rcuion, nnd ihnt it cntpn the rortl from tbu bniin ihnmt:>< li"
structures called ibo pyramids '>f the Iviilh, uhirh w nbntl fliidt \uu t :
TImbs pyramids crws over or decui««te as tln-y nn- nlnmt to ]in«*> r
card, forming; what is known as ih« dtn'uamion of ihv nyrainid- n
tract of 6bre* in tjueslion abBres in this decusmioii. H<
cnlled t\M en4»ed pyratnidal tract or mar« ftiiopl}' the j'yrHxo
no. 17*.
IK
t.a,l
PiAiMAiiTa ti.u-niiATK TiiattRNuji. AmuMinuMTo^THt 3«mtAi. Tiut-n 09 WHire Ht
If ru* ft'iKii. CiMn, iMiukiKDraN,)
TbeimUiililtUlranMUH'WTcliirilicnnhr'tmfBlutiTv. Tti* nUllunt nC tlH> ttMW *> itll
nglOM at lb« iM«<l KM *n<nra In r<i|. IW.
Tha MandlnK tmdK.lniMi ur itw>-i»ini( dtcMMnUuii. ■/» tluJHl wUli iIih*, II« i
tmol>.tncUofdeinD<llii(4tKvn*nik>i>.Mv>b*dtrt Willi IIrm; liindwiillair InMClinwvw^
ildauf tliScORl. viilj^ltia tvDnimlDlMfthMliK |4i(0nl»» 1 hr iitlior tl-l«.
cr.l' c»f Ld ifTnmiiW mil. "t mm nbuilj. i>rrurliliil ir*n iLr. iIlriKi pMsmlM '>*'*>
ikiul«<l oil tliu ilitr u|i|->d4t to lh«l on Wtltrb rtP. k iliMlnl, iu ontef t<' ln>!li«lo tto dUMwNi •(
ilin t<K> *> (r. >.mnlDg: c.h, cvnbplUr mil ; t.lr. ■»<) «r. iiffiiipi ■■■■■>niii> iIm- mtdlan p^""
Imrl orinclof nlim ol the poMtflnc (iwls VI, rriimmlUiR. «• liei|>:< ^.illjr ID IM ML
lliBt<nlc*)*o4*.lr, UtcMeral. IwiBlwr. kixl iIo*mI nioit: aic.it.l. !>>•■' : •■■nni»iitti*-
■Jewl. ilionuttioliietmitw^-" ''- ^ ■ — ■■ r-,- -,•—., cwialMitHl, nwrki
inii-ioi nilipc |i*ti'ii mcniiiiii .- iin oxMln ndm
|ii>lcnuiou]<i>aii r* 'ri<v«ii<. I ' Ik ixaWHoT liiini. ■"'
final Miia of LlaHstr** H>Me.
A (Diiillor, lew <'<'n>piriuiUH dmcMxIinff irscl ori'iipit'ii tlu- iDi'dinn nani'*
at tb« nntcriDr ii>himn (Fig. 17^, d.P. Thin i* nul only much Mimltcr hM
alto nuch more vHrinblo than iIm- rnwwd pymmiilul tnict, in not prnwiin
lk4! lo««r wiinialti. being found in mun nnd the moiikvy only and U'ltii.' *''"^
devvlopi>d In man thap in tlie monkey, and nwclKS ■ n-rtjitn way ocl' '
ikc spinal cord, generally coming to an «iid in thi> thoracic n-)(tiiii.
THE itTBCCTCRB OF THE SPINAL COBD.
m
fViini tlic prntmiil, and it b contiiiuntiuii af Unit part of the
ioh, unlikf l}i« n»t, thxt not dcctiatittc in lh« bull); thiM the
tract which i-omiiiK dimii from tho li-ft dkIo of lh« lirnin nms in tlw left
pyrantid iti the hufb, [himmv* <1owh iiiin elic It-ft fliiierior coliimti of the oonl,
ilsnce thi* Ninitlli-r inict i» wiUcil the liirrrt f,i/i\imi<ial tract.
Thvm tiro titK iliv most conKjiiciioiiR nn<l imjiortAiit (lw>ccti(liog tntcta, but
DtinK-R hnvr Ih-cd j;ivci) to two otbrr ijcsccniling trnclH. Oii«, known aa tlie
anlfi-o-latifttl dof^niling Irarl, is fl l>rco Iruct |>l)icc(l iu the nnlero-lHteral
eolumii, nml (eeo in scctKiti <Fig. 178. cicec. t.) w an elongated aren Mr^tcb-
ig fnini ihc: {ivraniidal Inict lowanl ih« anterior column and reaching at
ilOM M f»T as the anterior lisiure. The area i* lar^, however, because the
IB very diiritse, that is to t*y, the librm witli descending deg«neralii>ii.
fibree which dq^enerate below the se«:lii>n or injury, are very l«r;,t-ly
ixe<l up with libre^ which do nut degenerate; iu this respect ihia tract con-
tnau with ibe pyramidal tract, whicb is to a much j;r«nler eslent compuaed
frf fibres with descending il^^generalion, though even in it there are a coutiid«r-
ble number of tlbtcd which do nut degenerate. Indeed, tb>a tuilero- lateral
deacending tract a ao diflW tJiat it hardiv deaerveo to be called a tract.
The other ia a ainall, namiw, comma itbaped tract (Fig. 178, x), iiitnntcd
In the middle of tlw external pi>st«ri<>r culuran nhich bus been oWrvcd in
the cervical aii<l upper thoracic region*, and ha« been called the " dtMcnd-
Ins ciinima tract." But the dt^jeueration iea«he« a short way only below
|h« aectioii or injury oikI the group of fibrM iIiuk degeix- rating can' Itnrdly
COUildiTvil as funning; u tract <!omparablc to llic other (mc[«. Th« ar«a
probably n-nn-Hcnl.-^ tibn-« of the poetcrior root which take a <lftceiniing c^iurtc
nl\<'r itii'ir <-ntriiniy iiilo the conl.
Atcrntiing irafit, that iit t<i wiy, tracts in which the dogetMration talcM
_ aboro the («ction or injury.
A eompicuoiis uocnding tract iif a curvtil »1inp<- 1 Fig. 178, C. b.'i »ccu-
iat liw oaivr donal part of the bitrral i\>luiiiii lying lo lliv oul>ide of [be
~ |»nuuidal tract, between it on>l the surface of tho cord. It iipncars
b«gin III ths up])er lumbar rvgion. being said to be alMenl from the lower
lumber and Mcral cord, and tnay be traced upward incrrasing in size through
' e thoracic and cervical oorti to the bulb. In the bulb it may bo traced
to the reelitorm body or inferior peduncle of the ceivbelluni, and so lo lite
rebellum ; for the reatiform body wrves, as we ahall tee, in each lateral
lalf of the brain, aa the main conitection of the cerebellum with the bulb
nd spinal cord. Hence this tract is called the crrelrrHur Irati.
A »ec<^ind important asc*.-ndiuK tract occupies the median portion of llie
DHterior (xiluronD (Fig. 178, i-r,, s.lr.), and »o far coincides with what we
eKnl>ed al>ove ao tlie median posterior column, in tlve upper rejfions of the
ord. that it may he ualled the umiian fituttrior Inift; it extends along tlie
■hole length of tl>e spinal cord, varying at dilterent levels in a manner
ibich ne rihall prtMnlly studv. and ending alK>ve in tJie l>ulb.
A ihinl iiM-vndintF tract, called the wu^nuUn-j antero-AtbrirV Inicl, or tract of
)ower«, occupii-n I Fig. 17H, aac. a. 1.) the outer ventral part uf the lateral
nlumn. It ban tnmcwhal the form of a comma, with the Iw^ul tilling up thf
le left liMwceu pniji-^'tint; porliotin of (he eerabdlar and pyramidal tracts,
tJ>e tail stretching anay vctitriilly along lite outer margin of the lateral
slumn fHilaido the ani<'rO'liitcriil 1 1 ivcit tiding column, tho end of tl>e tail otlen
iching to ibo anterior VKfi*. It niuy hv traced iLli>ng Ibu whole length of
eonl. bnt it m itot eo dii>iinct and c^iinpact a Iract ■■ iIk two aiioeudiug
jcts juat mentioned ; the Ahmt with a^riuding di.'gNMration, that is to oav,
.e fibres degenerating above the section or sent of injury, are vvcy largely
lixed with fibres of a dillerent nature and origin.
41
090
TRG EPINAL OORIt.
ing inti
nwU, li^l
Urtnn.
Wo ta\j Atrth«r reniftrk tliiit Ibcsc Mveral tncts differ (com eaeli «i.
in como corns mtrkoiJIjr, as to the dismelcr of their coostitueul flbra. Thi
the cowbcUiir Irncl is compoeetl aimom (exclusively of reoiarknblj e _
fibres. The median posterior tract, on the conlran-. is wade upof ffaefibm
of very mjuable sixe, while the fibres of the antenv-lnleral ssoeadiuft mtl
are of » sue iuleni)e<iiate beriteeo ihe other two. The jiyrauidal ir
the Other band, is mnde up of libre^ of almost all nua mixed together.
The tracts then uhich an? thua marked out are. aa deeceoding iraoU,
CioiBiid aitd the direct pyrftiuiditl tracts, with the len diatiiiclorim,
nulera-Ialeral dwcendiog tract : and.a^u^oeiidin^ tracU, the (jenbclUir traM.
the median poaterior tract, and the les^ diittiiict ai)ten>-laleral uoat&og
Iract. If we iiu|>pose all these truels taken uniiy theiv i» Mill leA a raiitiilK-
abli; area nf white matter, uamdv, uearlr the whole uf the external {xaleni*
rolumn, the t.xlcrnal anterivr coluiim, iucluiliiif; the rt^on trmTeraed bv tbt
bundles of tbi; anterior rMi^ and that |iurt of the lateral column wlucli Ija
between the ant(ro-latcral di'Wfndin); tract iind tli<^ crww-d pyramidal tns
on the r>iit»ide and the gray matter im iho iuitidi:. From thi» am of vUk
matter wc mar Dut on one side tit prvMvnl the cxtental poetcrior eoluao
because, «« wo shall see, thU cvlunin i» largely compOMid of the Gbra rf
the poaterior root which pasa through this column, especially through ik
lateral part of it near the gray matter, on their nay to tlrair ultimate dtni-
nation ; hence the alternative name of posterior root-ione. We may liuc-
larly leave for the present the small zone of white matter cumpused of veri
fine fibres known as l/it po»lfrior maryiHol fonr or Liaaucr's £une |'Ki|(. IT?.
1^)1 l/ing dorsal to the tip of the posterior horn and in the lower reguei
reocfaing to the outside of the cord ; fur this, too, belongs to the fibres u tbt
posterior root. Leavin;; these parts out of consideralMHi we may lay ■•
roiEardti the rmt of the white miittcr, that the present Male of onr knowltdn
will not allow uft ti) divide ii into iipeeial traeta. All thi« area i* larnly
cODipoAcd of dhn-f ivhich do not under^ either aMendiDK or 'iMCOnail^
di.-gtiiieriiUon un the n-nnll of ft'ctinn, injury, or diseate. It has bi-eil tui~
gcslcd that ihc^Hj fibre* either htivr tin trophic centre at all or have d«ih.
ones, one above and one below, on cither of which they can in ciuo of iiefj
lean : so that when the fibre is ilividod at any level, the upjx-r jwrtion b
nDaridied from some centre shove, and the lower fr>»m mme centre '
A( all events, whether this bo the tnio explanation or no, Ihe fibres in
part of the while matter cannot be diflereniiated into Inicts by a study
llieir degeneration. Fibres of this kind, which wo can apeait of neither
asoeoding nor as descending, also oeeur in the cjctemal pwtertor column
rolDgled with the fibres of the posterior rmit. And we may repeat the au-
lIod, that even in the several ascending and descending tracts just described,
tapecjally in those which we spoke of us leei distinct or as more diffuiie. nnav'
fibres are present which under;go neither ascending nor deecendinK de^» '
vratiun.
^ Md. It may be as well ])erhapa to insist here onoe mote, that when tlww^
several tracts or the 6brei running in the tracts are spoken of as ajicendia
or descending, what is meant b that the degeneration takes place above t
section or scat of injitrj or disease iu the one ease, and takes place below i
the other. It ho* been siippnted by many that the uervous impulses whic
these fibres severally carry, travel in the name direction tut that taken by ll::^^
degeneration, that the ascej>ditig tracts carry impulmw from below npwu-
tliat is to tay, carry impulses which aiiriog from peripheral orgam pas*
various parts of the spinal cord or of the brain, that tlioy ore, in oilier wo:
channels of alfcrenl impulcce. and that convereoly tlK dMcendtiig tracts cai — "I
vflereut impulses. Tu this view is often added as a oofollary, tnnt tlie tr^-^ni
M
r iir— -
"J
Dr«i^
TOE STBUCTI'RB OF THE SPINAL COBU.
691
wbiolt di> not de^eneralu at att cam* impuliteit both «&y», and lience caiiDot
• bo OMtsidered aa eitli<?r afTun-iit or ofTuretit oliaiiuuU, but xiniply hm cuiuiiitiui*
Wtiltg ohiUlDela. UpDii thin il may b<; rcmarkci) (hat imjiMliHitJ ilu not iicccii-
iinrilv travel in the aamu (lircjliinaK tbe (U^caeratioii : wlion a xfiinal nerve
Iruiik E* divided Ibe afloreiit librm w vrdl M tbe ctTon-ul fibril b'>l)i di-gcn-
ttcntn in K<lwoendiDgdir«ctii>D toward the periphery, thuu^h Ihi! fontirr carry
impnUra lo the othvr dirvotioii. Htuicc ine dinK^iioii uf dvgviif'raiioii w nu
proof of the direction in which impulma tmvel ; morcDVur, m wo havo nocn,
oegenonUion doo* not mrtmilty travel along the 6\>n% nf tlio i>|>inst ord in
the Mme wav that it doe* along tho librM of a norvv trunk. It nmy bo thnt
(h« dMocoding tracts do carry impiiUoe in n doer n ding directi>)ii. thnt is.
efferent imptiUm, and that the ascirndiDg tract* aorve lo carry afh?r«Dt im-
pulse; but th« proof that iher do thus respectively act must be supplied
from olher facta than those of deji:eneration. Moreover, ire Bhall have to
I return to Iheae ascending and deecendin^ tracts and to Mudy their behavior
alnofc the length of the oord before we can use the t'aclB concerning them as
n basis fur iiny discuasioo as to their functions.
S 570. The eonrtetlicHi of iht nervn rooU. If we regard tlie spinal cord,
nod apparently we have riftht to do so, aareeultiug from Ihefuaionof aseriee
»r aegmenlA or metaateres, each aegmeat, repreaeated by a pair of apioal
uervee, being a ganK'ionic ma-«. that id to aay, a ntaas Dontaining nerve-cells
with which iien'ti filirea are cuuii«cie<], ive ihould expect to find that the
filHW of a spinal nerve soon allcr entering in, or before latuiug from the
«piual cord arc eoiineot«d with nervfi-cetin lying in the neighborhood of the
attachitM-nt of tbe nerve to the cord. We nhould, we suy, expei;t to lind
tbJR : but owing tu the ditKcully of tracing individual nerve fibres through
the tJingled maw of the Rubrtance of the ord, our actual kn<iwledg« of ue
(ermiuaii'in of tbe fibre* of the jKwterior root, tuid origiu of the fibna of the
nutrrior nml is at pnwnt far from complete.
Witli regard to the anUtrior not, there can bo no doubt that a very large
pfoporlion of the libr«« in the root are continuation* of the axi«-cyliuder« of
• cells in the anterior horn. The fibres which can thus be traood are of large
dianteter and appear to he chiefiy if not CKclusively motor fibrw f)r the
skeletal muszlee. In the fro^ a laborious enumeration on the one hand of
the number of 6bree in tbe anterior roote, and on the other hand of the
namber of cells of (be anterior horn in the areas corresponding to the nerve
Foota has, it is true, shown a very remarkable agreement in number between
the two. We raig^it be inclinnl from this tu conclude that all the fibres of
mi anterior root start directly from cells in the anterior horn, and that all
the cells in tbe anterior horn end in libra of the nearest anterior root, fiut
tuvernl ootuideratious prevent us from truatin^ too much to this observation,
mpccially in ibe caae of the higher animals. The anterior root contains
•finer fihre» than motor fll>r«s for the skeletal muscles, vasomotor fibres for
gee, secretory Hhrei and others; and it is a priori unlikely that th«fle
illd have origin from the same eelb as the nurtor fibr^ of the tikeletal
niusot<T>. Mureover, as a matter of fact. Home of ilie fibrw have been traced
Ihrougb the anterior hum, on the one band toward lli« posterior hum and
oq the other bund toaurd the lateral column ; otlten again are found to poss
thmtigh tlte anterior horn of their own side to the bottom of the anterior
Wurt* vrh»re, crowing over to the other side and thus forming part of tJie
■nterior white cnmmia>ure,tl)ey ap}>ear to ascend to the anterior horn of tbe
l.oil)i>r side. We cannot at proKent make any pocitire iiaiement as to the real
in and exact naUire of tbcec fibres which thuM upon entering the oord
._ I by the oolla in the anterior born without joining them, though those
Uch cross by the anterior white commissure are supposed to take origin in
i^
m
TBK SFtNAL COKD.
latbcri
puqi— ■ bi niMiUr ibst «UW ■ krgt Dttmber of tho Ttbriv <
mat. yimmilily tkan tapfiyiBg tb« ckdMil muH-lrs. tftl[« ori^n in lU
ckDb at Aa aaivMr bnia, rliannr itfon they iftu« froin tho oir*). nhm
Imtc Piaa utfcg aan. And similftrl5 we bxvc nwK>n to think that nil il<
fldb ia tki KBlenar BOfa dnaoCMad oat kxi^-cylinder pmnwee lflj< I
utterior now of dw aMe «U«. W« mair, bair«ver, imrd a Urg« n
at all eraaM of tte cdh of the BBlrrior bora, at the level of af «>
Bnleklo* aad « little ahnvethe level of the exit of uir pnnictilar n--
root, tm cMMitvdof a son nf iwdcae of oti|iD for the Ur^er niimhri
ihne, and tbeae nuai {unbabiy tke dteleial motor fibn-*. <>f that 0:1
raol.
TIm pMlmoT rwol eaten ibe eonl not in several bundles laterally Kattcrv*!
m doea tbe aatennr tmt, but ia a mora compRct inaM. ThU ninw, fanwever,
eoodfia n( at \tmat tao dia(iii«t bundlf*. wbich upon ibdr etitnuior itun titi-
mtd, tBk« dUKnvM connn. One bumlle, the lurner one, ^yiog to tli
or ininiian ndcof tba other, cnttnallDEul' rrlntivelr coaneGbrrA, aixi ..;....
tJie McrfMii immHa <FIg. 176, Pr >, mum* ohli(]u<-tr into the litlcral part '>r
ihaextenal poaiviorooluBB, ahica, m«« bav<- mid.iH in mniH-(|uerir'
nolna of aa iha poeterior root-toiM. Here iht- fibm changing Uiru
turn nin longitndiiiallr for aooM diataan upward (tonio, bowo^-vr. vvnsiin'
in the upper crrri^al rt^on, and probably in o(hi>r regionv, run a ebort dw-
UiikcedcnrDwar<lt bni evrnliialtr dttirr )^l, ai we «hall m«, toforai tbcni<iimn
poeMrior tract or make their way back inti th<; j^r matter at the hate •<(
the poaterior bom aitd lhu« j»4n lh« vniciilar ryhmk-r, thwigh some an; tt)4
to becoutinued on throu):h the gray matter into the anterior horn, tit
other smaller bundle placed to the outtide of ihe liimier, and called U«
tafrrai bundle (FlR. ITS. Pr^. niay be again divided into an inlrrmtHiyt
friirii'/e (Kic- 17". IVi lyinp next to the median huiidle, and into a «till nioie
lateral bundle (Fij(- 1". I*r"). The fornK-r, (iMidisiing aleoof cxtune ISbrw,
|dun;;e8 directly through the suhatance of Kolnudo at the extremity uT, U'l
ito tniu the ^mv tnattcr uf tlie horn, Ttliere Ibe fibres eh an^ ion; their directi>'B
mo ill |>art atleaxi liinjcitudiuulty in the gray matter io bandlee kumta w
" tbelungitiidinal bumlWol'ihi'pwteriur liuro" (FiKH.17B, 1 77, r. /.;>.), Me"
uf tibidi Hpimir to mm ou t» the anterior burn. The small nii«t axIMiui
or lateral jHirtitiii »[ ibe lalernl buiulle, consiatinj; of fine fibres and MBf-
tinea apokeii of km (Ar liitcnil liuiulk', on entering (he ooni at udc« aMend^
for Mime dinlance, and thu» liinnn the ibiii layer nf fine fihfM, tbe txattricr
ntarginal mni- or I.ifMU<-r'x xone. indicated in Fig. 177 by ui, (., nfaidi lis
bMwcen the actaal rxtrvmity of the l«)ni and tin- iturHiee uf the ct)nl. anJ n
the upper ivgioiu of the cord (cf. Fig. ITii.p') ninn »i>nie wav H|i«-Bri 'O
Ihe laternl margin of the hoto heiwct'n the gray iiintter and the i-n*
|)yn*mtiliil tmct. As it nsc«Dds thin layer eontiniiidly gives off Hlin-* ti^ iH
gn»v matter of tbe pwtlerior bom in ihc cclla of which ihey ai>[>ear In mil-
TbuH. while part of tho median hundto dow ni>t join the grnr Qiattvii
all hut goes to form tlie median posterior triict. the t\-«t of thai ^imdli'^"
all Ihe other tibresof tbe root, sooner or later, join tlio gray matter eiik-ral
the posterior born or of eotne o(ber pitrl.
S 571. Tke nteial fentvrt* of the *nftral rtyiouf of Ihe tjnnal roriL Tw _
cord begins below in Ifae Blender (ilameot called the/f/i<wi imninuU, "h''
Iriog ill tbe venebral aoal, in the midst of ihe mn^ vt uene root* a)"
(Be raifrfu MNiiio, raj^dlv mlnrgca at about the level of tbe first lumbar <
tebrn into tne ronu* mfifullnria. This may be refjaided aa tbe beghiaiiig
the lower portion of a Aiilform rnlarfceinent of the cord knuwn as the'iw'
twrlliny, wliteh rrarhn na high ai about tbe attachment of the n>»ti of w^l
M
THE STRt'CTURE Of TlIK SPINAL CORli.
U9J
or ckvODtb thoracic Dcrt-v iit (hi> Icwl of itifi otghth thoritcic vcr-
,tb* brottdoMpanof tbo»u'ellinj; Ix-iag nbout i>n|)o«it« tbo third liimbnr
Abore the lumbar swelling. Ihrutigh tho I tioracir r^i^nn tho soni«-
■rbst Barrcmeal cord retaioR nboui the satiiic <linmi.'Uir unlil it mchvs liie
l«ml of Um 6ni or kcodiI tboracic itcrvo o[>iKisi[« llie scrcnlli ccrvicnl vet-
' Ubn wbor* a Koood fuAifbrm onlurgi?n)unl, tho arrvical twelliny, hroadur mid
', tOBMr than Uh lumbar swelling, hcttitii. Tiw brnndcsl jmrt of the ccrvimi
' fmullDg ia about oppuaite to the HlUi or sitth cervii-ol nerre ; fri)m thcnra
ihi* diameter of the cord becomes gradually Honiewhat lew until It beginfl to
rxpaod into the bulb, but even in the hi);heet part id greater tbaii in the
thoracic repoD. The seotiinsl area of the cord increases therefore from
below u|tnrard, but not rej:ul«rl_v, the irreKularitr being due to the lumbar
aD(l oervii-ul awelliugn. The extremity of tbe lilum termiosle is said to von-
MM eDtirely afueurufclia closely invtsied by the membntiiea, even tbe oeniral
caaal beiag abMut. A little higher up the central canal bei^us. and nerve
celb with n«rve fibre* make tbeir uftpearsnce in the iieuroi^lia; thus a kind
of gray muttt'r cowrvd by a ihiu flU|>erScial layer of irhite mnlter ia e«tab-
liabnl. We have ulruidy rfforred to the peculiar features of the lower end
of Um cddus(SM>6): hut higher up the canal bocniDes central and small, the
poiUrJor otdunin* are dcvulo[>«Hl, and the gray matter oont^na more uertrtnia
eluMOta and rvlatiwly Uaw neuroglia, licc»>m«a in faet ordinary gray matter.
Fttiin ihencc onward tn very nr-ar the junction with the hulh. where iraoM*
tional fralurei begin to come in, the Dpiual curd luuy l>c Mitl to have the
gmeral >4ruclurc prrrioiisly dmcritiMl,
Tb* »r«-tiotial area of the whtKt matter iiicrCMes in nhnolutv lixc and on
the wivde in n Btendy miinner from below ujiwafd. In other wonU. in n
•K4>on lit any Icrcl, the nimdicr of longitudinal libra< forming the white
matter it greater than the niimlicr tit a lower level, and lc» than the numlier
at a hiicher level ; for any dilference which may exbl in the diameter of the
individual fihrv^ ia tnsuflieieni to explain the diffhrenoes in tbe total sectional
area of the white matter. If we were to measure in man the aeetinnal area
fiFeacfa of tbe Hmnal nerves aa it jmns the cord, and to add them tocher,
pawung along the coni from IkIow upward the remits put in the form of a
mr^-e would give uiMomfniich llji;tiren4thnlahown in Kig. 17!); the area gained
rat. in.
V w •• * I V f* '[ Ti bu X awn ^Tv n a ii i ww it « lii ■ li I
\ MMvUHi TiK I^KiTOU jinmoMU. ARLia or rna SruuL Xmvri^ raacanmc fsoai Bauiw
lD«aKa*lata*<iHva>idlvacTO, l»^l«LIV,IU.tM.altar«hlebTatata m*n. ta* Ml toml
,'jMt MtHmaU tlw t>4l«D uf IW (wnlMiiil Uic >l(lil liauil tlulupoftlui rutd, tlu iMitiinilt latl>
(wltag mn 1 1— 1 1 tj ilw BcnU, lomtHt, ihunwir: «ii<t raidiMl niRTn. Trti smnl driiv* wv ai4
ilmwa la Uit midv m«1c
by adding logotbor the ve^liounl areas of the nerv«s iiioronM in a fairly
•uady manner (Vow bolow upnard. Tite cnrre of tba aectlooal urea of the
whit* Bkolter of tha cord taken from below Hpwar<l would be very nmilar,
but if anything noro mftulnr It must ho onderaioud, however, that the
idiawnnonaaf the aren4 would not b« the tame in ihetwovafct. The oeotional
of tha «hi(« tu:itler at the top of the cervical region, though greater
0S4
TOE SPINAL CURU.
thiiD niiTwlitrc lover dovn, i« fur Ires Itian tho nnihd Hcllonil imtf ill
lh«iionTe hi-liiwlliBt level, ThcTihitc mailer is nnl Jbniio) I'j^all ikrHInt
IWtm the nervci nhieli join the (pioiil eord conlinuing lu run aloDg tkenr>i
up to ihv brniii ; ns we hnve eccn, foaw at least of llie fibrea eud in (ktgnr
mailer. NcTcrthelew, the wliUe matter in piming up tbe cord appon f-i
receive a permaiicitt sdttition iit ihc cntrnnce of each iierve. We niiiT iaSti
tliat each Derve lins a representative of ilEelf starting from the left! tf ib
ettlraiiee and niDning up to some part of ibe braio. Whether ik flhm
thus roprefenlnlive of the ucrve are coDtiuuali<jnB of llie very fibre* ef ihr
nerve itself, or are new fibres starting from some relay of gray mailer. «ilh
which the fibres of the nerve are also connected, is aiiolher (ineslion,
m.ita.
/
V lY iH u I * (» <ii 11 I 111 11 » K I* « >n V f* !• " 1 in lAi -fl V w ■ n '
tnicaiv •wxiiini twR Virutuum im tub AimewAL Asa* or tilt flai* H*nM or iWMOu
CoaD IlUhiii tn LMutu.
5 672, The gmy mutter in contrnst to the white mailer showa great v»ri»-
tiuDs in nrcn along the tciigih of the cord (Fig, \IHl). From the enirauw
of the CiK-evgenl nerve upwurd the nren iiirrea»e# very rapidly, reaching*
maxininm at about the level of the fifth liiuibar nerve. It then rapidly •!(■
creoacB to about the level of the eleventh thoracic nerve. maioiuii» abuut ibe
ttinedimetuionsall through lliethonieie region, and be){iuH tt> incrcue aciio
at about tlie level of the second thoracif nerve. Its weond max imam ii
reached at about the level of the fifth or sixth cervical nerve, afUr whicb iht
area again becomee smaller, remaiuing honever at the iip|>er cerrical TtgMi
much larger than in Ihe thoracic region.
i>S,l»J,
Ducaui OMiBtMi nu BtijLiiva Sktiiwi. Aa«u vr thk 8nK«i> XKHtvi *• iimi >n% t«i .
Smju. (Van.
Tbe meaning of these variali{ms bceonies clear when we turn In Fig. 181,
whieh shows in a similar diagram mat ic manner the seclionnl ureas of ihti^
•everal spinal nerves. It will be ohcervcd that the increase and deorea:*' nf"^
the sectional area of tbe gray matter follow very closely the increase am)
decrease of the ijuanlity of nerve, that is lu say, neglecting dJHereoees in th»
diameter of the Bbres, in tbe number of nerv'e fibres putting inlu the iiin).
Tbe Hdioiniil areas of the first and second Ba(.'ral, fourth and fil^h liiiubar
Dcrvea are very lar^se, and 0|>p<i8ite In ibeee the sectional area of the grar
matter nf the curd is very large also ; tbe eularv'^ment of gray matter wTiirk
ia ibeeownlinl causeof the lumbar swelling is ciirrelated to the large number
of fibres nhich cuter and leave the curd at thU region to sup{>ly cbictly tW
rnS »TRUC-TL'RS OF THK SPINAL CORD.
69S
limrr limh*. Similarly Uiv rnlnrgenieiit of grny lowtk-r vrliioli in llie ctwn*
till! onUKC »r thi' coivtcnl »wdliii^ it orrvlatcil to lh« Inrgi- number tif flhns
wbirlt eut^r niiil lonvr ibii' ri-giun of lliv vorfl (o hu|)]>Iv cliii-tly l)ie u]>[ier
linib*. III the tbonini' n-giim, wlicrv^ Ihc number ol nl)i>'i vtilcriii^ iiud
Irnvin^ ijiccord h rvlntivcly liM.lhi: wt-lionnl oreiiof tbe gray niiiltcr i« bImi
ins. Sinco tliv miiK-liUKtitii uf the ^owntl »|>1iwl nvn'tv nr« not exiii;(lT
uidiiliint fmm nrh «tht-r along iIk- Irngtb of tbc i.-nr<), lliv wclioiiiil iirra
it *o Gjcnct mnsure of bulk ; iho toml bulk of gray mnlU'r. Ibr inrttiiice,
iging to tvo nerves which cnu-r tli<^ coixl clo«e tog«ltier is lees tlnin thnt
two nerree fpvin^ nte to (he *ame Kctional hkii of gray nmtlcr as the
iibrmer two but entering ihe cord far anarl from each other. Still the error
'~hicb nay l>e iiilrvHluL'e<l by taking seclional area to mean bulk ia, Ibr preaeut
rpotn at all events, so &iusll that no may perntit otirvelves to say ihst in
(the tHjccessire refpons of the sfttnal cord the bulk of gray malur in any sey;-
uent » sreatej* or lees according; to the size of the nerve (or pair of nerves,
tight ana left) belonginK to that sef^ent.
rVoin Ibis auatomicfll fad we a]>pear jnstilled in drawing the conclu«oD
thai at all eveula a great ileal of Ihe gray matter of the spinal cord luav be
eaoHiileret) tat funiialiing a nervoua mechanism, witli whieh the dKirenl G^)res
pf each »|>inttl uvrrH lUHt Iwford they leave th<? cord, an<l llie afferent Bbres
■oan alXcr thev join tli« cunl %Te uioru immediately connected. It may be
thill iIm; whole of the gmy matter in thua directly ooniiovtetl with and tbii»
rwMi ami fall* with the JihrcM of the niTvcx; or it may l>e thnt there u v iiort
of nort) of gray matter, which maintain:) a tiiiiform bulk along the whole icngtli
of Iho mrd and servt« as a basin uhieb i-" here more and there \<xf nwoneii
bv the addition of the eray nialttx more ininKxlialely connected with the
ibrtw of the nerves. Thie question tiie method which we are now using
noot settle.
§ S73. Owing to these diflerent ratot of increa«e of the gray and white
Iter respeelifely along the length of the cord, we find that in iections of
e cord taken at difierent levels the a{)|)earancea presented vary in a very
i«tinct manner. Tlii» is strikingly shovfti by comparing Figs. 174, 17(1 and
77. Ax the level of the third luinbar neri'e (Vie. 177) the gray matter is
ery large, rvachinfc. as we haw seen, its maximal sectional area at about
ie point, so that although the area of white nkalter is not very great the
hole area of the cord i* imnsitierabla
At llie level of the aixtli iboracic nerve (Fig. 174). ui spite of tbe wbiie
matter having ver^' decidedly increased, tbe gray tnut(«r has nhrunk to sitch
rcry small dimensioDM, that tne total aectionalurea of the coni hu* markeilly
limioished.
At Ihe level of the nixih cervical (Pig. 176) the gray matter bun again
iCreaaed, reaching here, an w« have seen, itn nerond maxiniiim - the while
wUtcr haa also further increaneil, ami thni in(lei.-<l very ranniilcrahly, so that
le total an-a <if the cord in much greater that in any of the lower regions.
Further driaitn of tbe varying ?wc of the white matter and of the gray
i«r HI diHt-rrnt levels are aUo thown in the series given in l-'ig. 1>!'2. In
couibiniil with the three tigurw jn»t referred to, it will be obserx'od
,a1 the iK'Hal iucrmsc and decfetuc of the gray matter does not street all
r)d of the gray matter alike, so that the outline of thejjray matter changes
ry markoliy in pawing from below upward. In thflooocygeal renoa each
teral half is a somevhal irregular oval, aiid iu the sacral region, r'ig. I$S{,
c, the diflerentialion into anterior and posterior horns is still verv indistinct.
ihtj lumbar region ibe two hurna are nharply marked out, tliough both
posterior and anterior hon» are broad and more or leM (pia^lrate. In
thoracic region the decnuise of gray matter ban aHectuI iKilh lioru*, m>
690
THE SFIKAL COBD.
Iliflt t)ath Ar« pointed and tlandar, while tli« junctMn tietv«>n tHeni liu i
uiKlerj^oiie hi luiicb itiadnutiOB, so tint what liua hceo calleil (iie lal^nil Imrn
is relatively cunspicunua. In tbvoerrioal reffioo tlieRtiirDini; i^^m<eltfl^
cnucb more on tlie anlerior hum which again becM>mn lur;f« and bruad, ihtn
on the poMerior biim which *lU\ remains *i en der and )i-tiDled. Taluajilir
form or the gray luatlcr in the thomdc rugimi na the lunn? lypicitl fnrm'f
the gray matter we may «uy ihni whik' the? increiui- nii ihe lumlnr twtWm:
bean equally oti the anterior and piNtcrior hnrnit, ihiit iu ih^ fcrs'icai iv^hd
bean chiedv on the ant<-rl»r horn*.
Now we have no rafon (o «iip)>inio that cither affcrcnl impulMS reat^iliF
luinl>iir npiuiil cord in grcaliT nuRibcni fn>tn the lower limbu, or al'iiii;>i>Trif
the iiervw joining tlii» part nf tlic conl, or that tlwwc which do mulm
■re of n more Icomplex iiatitn; tlian i» the cn>c with llw nflirrent impul»
Fui. tlCL
■7
y
Ct.
..Ur,
Cs.
dp
.^..
kl^
y|i':
c«.
U.
AH lUfSTBAnxc Kon. or rn* tujvaa or tub Srauii. uontt xt Dirrum Lanuk
I0imuximu>.)
OwMvtWMvitnini id (lalD. knit nivwant the eonl maciiMMl fmr iiiiml Tlier ■'■'>* "■■
H>«* M ■UlhnM WnrU In lbu>lmp« taatlaal tliDtaciI. la tha ouUlUe b< lbs Rtmy iimllet, aod
nlMlTa poriUoo of lb* uurinr Mid inawrlnr flavm. Mid ate tbivw U>t tahMlont al dUhnnt
Of Ow MTinl ' OMU " o( Ibc vhlw mkltet.
4ilMt**plofihaMiwiil«fftTlfBl Mrv*.(^af UMARImrvlnl. r.nrib* vlfMkoornlnil. I>i«f
•MiHMl tli<waclo.t>(nf Ihc tilth Iboraclc L|Of thannl limhir. I4 Wlbr BRh Intntiar. aad Mc.
I ivenflil mcnl Orttve,
>rtiadlasar tlwinra>wvih>ainifiiu In Mi ITKi bMin Uw nadlu partiHar aolnau of t>t
OBSf flbtnoHiUnsiViMnilicivnl Dtrra i. f-, mq IumIwj anrm li.tn JUantitAttA ftom
«B.d. r. ofiUiimbtlMicIng 10 ita iliurKlo iwrtm. Inl',i»4Utliiciiaa k nM<to bvlwcvn aiij'
<•«><* of llhm i Ib l.loDlf BliKiuf •■rntl iiniia iiii 11 |n ■■mil it ; to UDiIV (he •<•« dona)
|«ttliM> conopiiuili lu Mcnil ttirt* *i»l iIk aeil la lumbar, nr liuntar Uumcli' wrim.
hiBg the cervical oard alooR th« nerrw of th« apptr limbo. Th« incraaM
•jr nnlter iii the posterior bonw b therefore uot correlated tn any in*
le In ibc Dumber or compleiitr of Ihcaflcreiit impulses rMchiDR the con! :
we may imiviaioimny concluilo ihiu at leuat a larjie part of ihe |[rny mntttir
w pueterior honi it not ipedally conoeroed in any i^IatHimiiim or traof
alMO of aflkreDt impulsea immnliately upon their nnival ai tho ooid.
•d, wc liavo seen that trhili! there b ample eridence to oooneot the
B-{vlk and therefore pranimably the gmy maU«r in genoral of tlie
T«B SPIN&L OOBD.
kflfB wilk lb* rSrrtnt motor Akm of ihc onli^rior nvit. tli«fic '
Urn al tb« ftatmot mot villi tbr Derv«-c«tli>,
{ir iiiili«<l Ally iiLlirr rmn I
Um mj mallir of thu poMcrior boro. We niair odd (hat, km «<> uliall yA**
i«t Mttr oa, Ki MiiHial i* ibt canrarreiice nf iigiproiiriitu- iifTcn-ni inip")*n y
Uw duetxriyiiiEiMtof cmnplexeoaHtnmtL-RM>t(ir«rdfi-rviit ini]>ul«ft>. llu<*'
as tcvcrlr ezprrt Iti find sdt faicmuw in the ncrvou* mMhiiniiiiiu i]<v<mI
to tlw ponly BMtor fiiDcUeB of'wrying out nxitor imptdvo withnui ■ com-
spoodmg iaamm is tha armut mufhiianma tmlon^nji: u> llir atlirnvi
inpalm. by tneatis of whieb thnt notor iiD)xilsn> are guid«Ni) and eunrdinahii.
HflDCP, were tbe \aiur DcmMis Dwduuuflim raslriclcd to tb« posMrivr bonp,
«« aboald expect lo find ■ gnalcr p«ra1l«li>ra (ban dot« Bcttially «xb( bc«nm
tfaeiB aoil ih« anterior liom.
( V74, Tka ehaofa in the arm of gray mntlcr illiMmtnl by tho ttvt-
■tute ud llim ■Hill given above n-trr tu ibe gray matter ns n wbnlf— ibil
n, not only lo sene-celk, but alH< to slrands and uetworks of nerve lihrtt
and nerve tibrik, awl indeed include lo a certain extent neurt^lia. ^Vr
hare Men (| o67j that we an able to ditti»];uif.h certain large and e»
HMcaons nerve-cells in tbe gmy auuier, and to arrauge tbeae iitto KToafi.
The gray matter enatains loaoy other small iMrr^oelTs, which we ntewt
able at preaeat to nane or arrange, btit uhne existence must aJwan h
faarse la mind. CoofininK oorwlvt^ now, hnwe\'er, t» Uie grftupa of larev.
nore conipiruvus iKrve-<«1ls, «e find that, broadly speaking, the dlie^dt^c^
eoccB whi*^ can be obierved in tbe rells of tbe anterior tioru almii; ilx
length of the cord are that in the thoracic recnon tlie iier^-o-i--ella of tbe sau-
rior honi are few, ami relatively small, wbUe In tlw L>ervical and lurahu
region, eapeciaUx in tba latter, iltey arc numcroa* ami large. It n not ewy.
even if pn«iU«, to ^lin^txh inlbe thorade region tbe ncvcral gm[H rf
cells marked in Rga, 176 and 117 at 'ia, ,i, ; : On' rowlian group ( Fi|:- iTn
17'. 1), indeed, Menw tn ba iba only group pnvent in thi- mi'ltln n >' ,
regioo (Fig. 17-1. t). The group of tno |>oaterigr Ixtni (Fig«- 174. 17ii. I^<^
6) anrpaKn> Ui be about the same in all n^ion*. ^M
Vt ith two otbcTgioupf of nerve-cells nriking diflctvnon are weii in >ii/-
ferent rcgioiw. The vehicular c}-lii>der, for instance (Fig. 174, 3), i* m'*
conapicuous in the thoracic rr^on. It niav ))« eaiil to reach frira ih<-
MTCnth or eighth cervical nerve to the thirti lumbar nerve, being [n'lhsj^
Dioet developed in tbe lower thoracic and iipiier lumbar regioa- It ■
abaent in tbe cervical region above tbe seventh or eighth oervical aerv^«»l
in tbe lumbar region below tbe third lumbar nerve; but a itniiUr ^r" I' '
oelle ifl prtseut oppoalle the aecood and thinl cervical nerves; as">"i-".'
more doubtful likencn la aeen in the sacral region below, and the coIuub »
•aid to have a repiveentative in tlte bulb above the epiual cord proper. !■
■eenis natural to infer that the mU» forrtiiug thia vesicular cylinder an""-
necled neither with the ordinary somatic motor libres ^reming Iho sln^
muscles, nor m ith the ordinary alferent setisory, aomaito fibrei ootniag ft"*
the skin and elsewhere, hut in some way with some special twin of flbrwi ii
tbvpoint, however, no niitlKmtati\-e statement can as yet lie inndc.
Tne lateral bora or int<'rm<'ilii>-laieral tract (F'ig. Ii4, 4) i« also mod
Rpleuous tn the ihnracic region. Tn the lumliar region it h Icitt or
with great dilliculty. anil in the cervii-al ri-gion veenm tn W memd
miift donuilly plao-d divinion of the latrral group of celU of itui
burn. It in iKwitilo that (hie j^roup ivprMtiiH in thr liniblnw ilt<
region the cells which iitv devdoiHfi itito the gnat lateral group ol'tlm
rior boni in tbe regions of the limlie.
TII£ STRCCTt'KK Of TUB SPl.NAl. COBO.
mm
, t979. Tlif wliire mnilor, ae wo havo ee«ii, iiivreaMS in eeLtional aici
nth ctiitniilrniblf rrKulHrity fruin belnir upward. If, iDstMd ut' a disftmiu
Bf thn ini'mivr of Ihi; whol« nhile lualltr we construct io » similar way
dbnania of ihe anlerior, {looterior, aixl latenil culuiuDS rtspeciively, we
lii><I lh*t while the K^-licnnl nren of ihe lateral coluniH Fig. l83)iu(TeaM6
■ iih Mom cooiitlcimbic rt^'ulurily from Iteluw upward, ihough not tu revu-
larly aa ilom ili« wliole ar«a of while uiaitcr, Ixiih ibc anterior (Fig. \Si)
I
FM, m
/'
•1 ■ I mil t v wMi V V fv << k i v<4 w VI i >f ■ " '
XHMa»> nMm*o nia VoitnoK* w nic MmMrtL AaRi or the t-in>*t Cilpim* or tut
ariKAL Cms jiLMio nt Lnumt.
Fki. IM.
I MDVKWvavlVMHI IVIVIWUWi II
«t« •w.vnni Twi VA*t»Tiina nc tut Skumh Aria or twi Avnaina Votiatm or TWi"
^■«i Omui aumo m Ldvth.
Fut. ("ii.
i' 6 "i ~T ^ J J V N > •
I TWI TAKUTtMM IS THE SumiWlI. AaC4 Or THB I'O'miOa fVttTVM* or TWI
iVout DwD ALoiO) m LuwrtL
BDil tli« pntlerinr (Fig. 1)<5) coluinus aKrt« to a certain extrat with the cmr
UMder in «buwiug h dmded iiR-r«nt« in both lb« luinbiir and Lh« cetviral
awi'llii))^. We maj, prnvuiMnnily ul li-u.Ht, infer tmm titia that, while coa-
aJdembli.- [Mtrliuna of both the aiKcHur mid the pMilerior ooluniQs ar«, like
the ndjoiiiing Efwy nialter, iu toma wa/ or oilier cuiicem*^ io the exit aad
entrumv uf ctJen'nt mid atltirriit fibrrw, ihe larger jxtrtiuu ut the lateral
coluoin i« coat'criH->l in the cranainiMiioii of impiiUca Iv and fro, between the
local DMtcbuniymK hi-low, ininH-diiitclir coniwctcd with ihe sevvnl *]>iuul
nenrca. and thv brain nlmve. Thi* cunelueion areme inddunttUIr oonfiriiied
(though tfaew tliagnuns miift not he i>trniiicd to rarry tlotailed tnKnnw*) by
tba Midden inereaM of the lalenil eulumn above the lumbar Kwelll0)c, as if
tba largs maw of nervouB mrchanif>m for tbe lower limla cnuventrntud iu
Utia ivgion demanded n sudden increase in the number of Gbrw cuiinediii): it
witb the brain above.
Thia mora or IcM coniiouotiB increMe of tbe lateral oolunm piirtljr explain*
tbe ehange of form iu the geiteral outline of tbe tninsvenc Metioii uf the
aofd whkh is obierved in podHiDg upward from the loner in the higher
700
TBB SPINAL CUHU.
regiODt. In the cocergeal. Mcni]. tnd lumbar rogioDi tbc oatliai^ tbou^b
TBrylDg «<iiiivwliiit ehwfljr owing to tho dtspositioD ol' the grof nwiter, bag
tile wnolo dreulAr. In the t)ii)rticic region, Mpociall}- io the upper part, ik
loenMH of the lAt«rnl cuIuoiiib incrensce tho side-to-«idc dtaio»t«r M waA
(hai the section becomes oval, nuH in the cervical region ihi; incrrftN! of LSt
>i(l<.vtu-Ndo diameter out of proportion to tlie dorao'ventral diiinioliT ii leit
mnrked. The actual outline of the whole trmaarerve tectiou b,li9«nir.
(k-tcrniined alao to u certain exlviit by the ckiue^sof forraofthegn^iiiMttr.
The cord moreover unJergow along iiH length a change n liich ig nit wrj
clearly indicated in lie diagraiuH Figs. 184, ji^'>. By comparint; the (rtin
of traiiaverae «caion» giv<'n in Pil'. IK'J. it irill be »eeu that the rvtaiire |>c»
(ion of Uie cuntral canal Hhilti iiToiiz llic Ini^th of the cord. In ilia wall
and lumbar Tvgion% the cotitral auiAlit ncorl)' at (h« centre <>f the drdttf
outline, niid the posterior nml anterior fissures are ncarljr of e<|Ukl iltplli.
Kveu ID iho upper lumbar region, and still nioro in (he thoracic r^oa. ibr
position of the central canal is ehiiled nearer to the venlml surloos, S4 (Inl
the posterior tiuure becomes relatively longer, deeper, than the oauriw.
This ehiltiug goes on through the cervical region up to about tlie Icniof
the Moond cervical nerve, where it u arretted by the beginning of ibt
ohoneea through ivhioh the apinal cord ia traDsforiued into tbe &r bnr
oumplicated bulb.
This leugtkt^ning of the potteriorlliBure indicates aoiDoreajM^ in ihe'lop*^
ventral diameter of the piuiicrior coluninn. and thii, not bein>: ficcoiiijiuaol
by a compensnting diminutinn nf the iiide-bo-aide diameter, ahnuj in Inn
that the pOMlerinr ci'iliimnH uncl<Tg<i nn inureoite in posing upward. I'm
thi;i tve nmy add to thi- provixiona! conclusion juM arriveil at with regaidto
the lateral columiiB. the further conclusion that m»av (xirt of tlw pMlcmr
columns alao is concerned in transmitting impul*e>, in a more or leMdirM
manner, between the varloua regious of the cord belovr anil the brain iiboTt.
The anterior columuii do not iDcreOBe in the same marked manner, llioufk
over nud above the increase due to the lumbar and cervical swelliugi a mi-
tinuei) increase may be ob*erveil, espeoinlly in the upper rervicid rvgioc; il
is in this upjier repon that the direct pyfamidal tract la best deveiojied.
§576. Till- provUional coiiciii«iouii at which we have arrived ai* furthM.
to a Certain exiMit nl irntt. coaGrmed and eslendeil by a study of lli*
b^nvior at the several region* of the cord of the *jtecia! tract* of •fciw
nuuicr dcscrihcl in ^ M'.
The pyramidal tract, that is to say. the croMdl pyramidal tract eotcii^
the spinal cord above from the pyramid, is vr-ry larLru in the oorvical ni^
having the form and riiinilion sihown in Fig. i'<'i, C,C,C,. Fmm llwaM
doimwnrd it diminishes in sixe. tha diminution being i^pocially ropid la thi
lumbar twellint* (Fig. 19^. !<,). where the tract, being nn loneer covered i*
by the cerebellar tract, comes to the eurboe of the cord; out it maybe
tnoMl by the degeneration method down as far na the eoocyi^l re^ioo, nn^
indeed appeiirs to be coexistent with tJie entrance of spinal nervua into the
oord. Diminution of the trad means a JeeAening of the number of fibna;
and since we eannit mppuae that any of the tibrat come sitddealv to an tiA
in the tract itielf, we are led to infer that alon)t tli« oord, from aliove dom-
wanl, fihn-a ari* «uoc«Mive1y leaving the tract and Tnuain;; to some other p«it
of tlw oord. \Vv Mo-ni further ju^lifinl in oouvltidiiig that the librae whitt
tki» silcocwivHv Icavn the tract go to join (he series of local Dervoua medi-
anisiiu with which the spinal nervcn tximniuuicate, as we have seen komo
to believC:, upon their entrance into the cord. Indeed, as we aball ae« loUr
OD, wv have reason to think that the ni-rvoog mi.»oh an isnu which thi-(ibrotii
qneslion join are those bclon;;ing to the motor fibrei of the anterior rooU
TBK STRCCTCHB Of Til S SPlSAh CORU,
701
This pyraRiidal tract Aoea not bt|;in in tbe p}'niiuid, but may be traced
(hroufifa tbe lower pans of the brain rifjht up to special areas id the eort«x
nr surfaee of ibe certbrnl hemiejiberefl; and very etroDjj reaaoDs may he
brought forwanl Jo support of toe view that tli« fibres of ibis tract are
fibres wliich <-arry inipulsee from the cortex to succetixive portioos of ilitr
Kptnal eonl, am) there {(ive rise to efferent tmpul»es wbtch poM to appro-
»i>riai« skeletal Diu!«)es. 1'lie tract, therefore, ia uol only a dewendiug tnct
Djr virtuo of the m<Kle of defeneration, but may he »pokeD of in a brond
•CDM *« a trHct of rfrereiii inipiiUes deacending ttata tbe cerebral cort«x :
and indcMl it i* rnainliiiuvd thnt it in the channel <if the pHrliiolar kind of
efferent impulM'^ which we hIiuII >|ii-iik of n» voluiitnry or viililiniiiil impiilus,
Wv may mid that »* tbe tract jiuiu^ iiloiig ii |iiilh, which »<■ iilinll siihM-
Inueiitly <)4«cribe, from the cerfliral cnrtex through the loner |>iirt« of the
Drain to lh« pyramid, it givn off librca to mecbaniiimN coniwcted with
aeveral of tbt cranial Dervo. mucb in the aam« way that it gives off fibre*
to the apiuiil nervca.
We muy therefore picture to otinwivw tbie pyramidal tract ai< itarting iti
tb« form of a bnmd sbcnf of lihrcs frum n certain district on the surface of
one of the cerebrnl hemispheres. I'nltin^ aside lor the prMeut imy pouible
increaae of the number of fibrtv by dlTi»ioi) of libres I though we have reason
tu think that this docs to a ccrtitin extent occur), we may regar<! ihe trad as
being at it? maximum at its l>eginniiig in tbe cortex. As il descends to the
I decussation of the pyramids in the bulb it loses a oertaio Dumber of tibres.
which paaaotr to the cranial nerve:^. Having croased and entered into the
lateral column of the cord it cooLinues to give off libres to tbe spiual nerves,
probably to tbe anterior root of e«cb in succession, aiul so goes on its way
down ihe conl coiiiinually diroinisbing until the last remaming fibres ore
given off ii> the liul oicvygeal nerve.
When degeneration a set up along this tract, as may be dono, by injiiriex
to |Mrticular aieus of the cerebral cortex, die main raasa of degenerateil
fibrvK, iiAer croaaing over Ooni one «idi' of the oerebn)-.i{HnHl axist to the other
in tiK- dcciMtation of tlw pyrainidn at Ihe lower end uf the bulb, during ilt>
further pnigmw ilown the spinal eord, keep* to the i>iile to which it bn*
cruaaed right doirn to the end. Hetiee, aa we have »uiil, it Lt c-idl«^l tbe
aevmed pyramidal tract. Th<! main mavit of Rbru, thv dvgcncrntion of
which ban Ikxih ntnrtnl by injury to the left ulilc of tbe bruin, croKMat over to
Hgkt side of the Kpinid cnnl niid run* down ihi; Intrral column of the right
nua to the end cf th<^ cord. Nrvcrlhclem Kime fibni' appear to ci«w over
igsiu in the spinal cor^i and tWn to run iil<iTig tli«' »inte mic as tbe side of
tfao brain injured — along the left side in tbe caw just mentioned. Sucb fibres
atv upoken of as " ro-CTOfsed Bbrcs."
The direct pyramiilnl tract I Fig. I'^'i.dP), except tbiil it does uot cron at
the iltKrusGatidD of the pyriimidK. Is olh'.-rwi.<c <imll»r t'> I he crowed pyramidal
tract, and indeed is a part of the same i^triind to which the CTOHod tract
l*elongii. When degeneration in this trnut is slartixl by injury to particular
%fta» of the cerebral cortex, say on the left half of tbo brain, the deseocra-
tioo may In traced ihruugb the left antei^or pyramid, and so to the left
itiiKiiao anterior column of the smnal cord, 'rhe direct tract is never so
tsl«DBiv« or marked as Ihe cr'>«sed tract, does not reach so far down, is much
imrt^ Turiable both in tengih and in sectional area, and, as we have said, is
timoet confined to man. Diminishing as it deecemh, it ntay be said to oeaw
tho middle thoracie region (Fig. IS'i, DyDJ. Taking an average, we may
%y that, of the whole Mrnnd running in the pyramids above the decussation,
fcbout three- fourth.4 of the &hfta go to form tlie croaaed and about one-fourth
form ibo direct tract.
go
We xholl
see later on that the impulses coming
702
THE SriKAL CORH.
dova ftlong Iho iinitod tract [q tbe bnln may, bnit'lly iiji'^akin;. bo Mid I*
OKMB over wholly from one »\de fi the nilier 'l>i-r<in: tfaev roai-h lb* tkclttil
tDtiselcs. SO thftt lh« impulses pasinj; alonj^ fibre* In.uy, tho loA iirrisM,
reaoh the muscle ^ of the rit^ht limbs atid rivht nie of tlif bulr, wli<ib« ibr
flbrej croos over al the dei^nMntiou tn firm ibe crvHWrl or romn'm nn lk( luif
side to form tb« direct pyramidiil trael. Wu nre tlicrcforc Ird I'liofrrrUtf
the flbrea io the direct tract, tn ibey paia down the cord, croM ovor in iht
onrd itaetf before tbey make coDoectioDt with thu ii lin-4 nf tbe noierior roit*
t*nil>3bly the cruuiii); '\» efTdetod bjr mcam nf witne of l)i« doca<«stiB^ libnt
whi<'h form the anterior white commiMun^, A purt only, ioilCMl a roilt
part, of thy commiwiiMt ran ^rveihi« purpose; nuMt of lli« fibrtM nf tb»oiiD'
inimtirt-. and id the lower regions of tho cord, where the direct tract no loiigK
oxt*l», nil the fihroi must have some olhor functions. Some of th« Glmttf
thU great pyramidttl tract leiive the tract, >w we have mhI, to join tomtt
tbecnnint nerves boforv the nymraidx nf the bulb are readied; aad iht
iiDpulfcs pa»ing along ihcso tilirfa al»o cro«« over to the oppanle side befbrn
they issue aloii^ tbe cranial nervc:«. Hcnco wo infer that tneae 6bra) dow-
eate above tbe decussatioD of the i>yrnmid« ju«t aa thoeeof the direct tnd
deeuBsate below il So that of the whole strand as it leaves the oeitbnl
cortex, while the main maas of libree croues over at the deouasuJon of tbe
iiyramidi, the rest of the fibres croes the middle Hoe in auooenioQ fVom tte
Uvel of the third cranial nerve to the level of tbe lower limit of the ditMt
tract : below the decussation of the pyramids the eroaHfig takea place by nxini
of the anterior coniniisaure of the cord, above the decuaaation by raeaaiof
what we shall later on learn to epeak of as the rapbd of tlie bulb, orbr
structures correaponding to this higher up.
§577. The eerebeihr Irael (Fig. l>»i. Cb) is, as we have aeeu. a tract rf
ascending degeneration ; the degeneration in it makeo its appesraxioe ab:>vp
the section ortheaBfll of other injury of the cord. It begins somewhat *od-
denly at the level of the seoml lumbar nervp re^on, being abntnt at leaal
as a distinct tract Iwlow ; injury of the conl at tbe level of the latddlc and
lower lumbar ncn,'c» lends to no marked tract of degeneration (though yn-
sibly KfttUred single fibres mnv H^^nerale). while injury higher up does.
The tract lies, n« we have Knid,cfo«- lo thvurfaoeof thacord in the poeterior
pan of the lateral column Jiivt outcidc the crowed pyramidal tract, and while
varyiug snniewhat in tha shape of its oection from love) to level, remain)
thn>ugnout n somewhat narrow orescentic patch. .\t the lop of the epiaal
C'ltil. it passes, as we have Mid. from the lateral columns into tbe roitifurm
iKidiee of the bulb, and so to certain parts of the cerobellnm.
When the section or leuon is liniilcd to one side of tbe cord, tbe dejpeu-
erdtton is similarly limited to the sanie aide, am! that along il» whole course
up to the cerebsHuin ; there u no erideno.- of any of the fibres decunatia;
in the cord.
The area nf the tract increases (Vom below upward. This has been dtUr-
iuinutl by the emhryological method, by noting the appearance of tbe
mwlulla in the ilbres. as well as bycompanng the extent of the degftneraltoa
following upDn a section high up in the cord witb that followin;uc upau a
saciioii lower down. From this we infer that the fibres corapnsing the Intel
must start «uco(--iKiveIy from other parts of the cord along its length — that it
to say, the tract munI be fed by tibrea comin;; from other 9tnictur<ai in the
Cord. On the other hand, it is found that the de^nerated area following
upon a section or injury dimioishcs as it is traced upward : when, for instaooe.
u section is made in the mid-thnraeic re;;ion, the area of de^eneratiou in tbe
tract is greater immedialely above tbe aeetion than it U higher up, say io
the cervical region. F^im thi« we are led to infer that though tbe tract it
TUB STBL'CTfBE or THK SPINAL CORD.
708
ij r«d tlong it* ooarM by librw iximin^ from ollivr |urt# of tbo
•oiB* of ihe fibres cnUtring tho Irnd, though like (heir ounipxiiioiiB
rgoingan tuoeoding cWc«>onition, da nol lilcftbom cotKiauc in the tract
up to tb« c«n)b«lluin, Init p«M ofl lo other jnm of the eort) on their
ipmud. Tltis, houevttr, in c<|iiiralent to Mvio); thiU the tract is not a
or haraogwoous one, but eon«i«ts of st teitst tiro wts of Abres, only one
licit t> continued on to the cerebellum nrid atrietlv deserves the iinme
enboUar." It mny {>erhii|« here be mentioned tliat while the flbreo
ouag tiw tnct are as a n-bnie coDsfticuously coarse, larj;e fibres, wiUi
tber« are mingled, especially in the thoracic rejtion, a number of mucli
flbra ; but iboee apparently undergo a desuemlinK. not au a.Heeiidiag.
i«ration, and do not titerefore really beIon)( to the traot ; they may be
which have strayed from the pyramidal tract.
) have a« yet no rerv clear evidence a^ tu the origin of the fibres which
oae the tract. I'nIiVe the case of the median iioaterior tract of vrbicli
iv« next to speak, no degeneration, ui leaat in tlie lumbar luid tboraoic
at, appeant in the tract after sectinu merely uf tlie roots of tlie nerves;
xtuce the degeneration the cord itself miiai lie injuruil. Fmin thi» w
mfor that tli« tract is not fed directly by the fd>r»> nf the [Hwterior roou.
obaervera maintain that the tract i* f«l by libree comiuj; from the
llftr eyUnder and [wint oni'thal bulh the tract and the i-uluinii begin at
UM lavel sonMnbot luddenly ; but the want of pantHclism biilwvvn the
• of the tract and that of the cylinder along the knglh of the c^nl, the
being M we aaid cunspicnou* in the thoracic rcginn wbilo the tract
ilr incrcu«ca upward, i* distinctly op^raeed to such a view. From the
:l)at tho dageneratinn taking nlacc in it is au mcendiag one. it i:< sup-
thnt the tract i» the channel for RM^rnding, that is (o ny, in a broad
. aflerent impubos. And conaidernblc iiitereeit nitncbes to the fact thai
impulaea sfaniild be carried, not to the cerebrum but to the cerebellum.
kooiftedge on this point, however, is very imperfect, and what can be
II the matter liad better be said later on.
78. Tlie m^ioH jiorienor tnct is the other conspicuous tract of asceud*
Igeuenition : it also is supposed lo be a ehanucl fur ascoiidinK. afferent
Gi; and this view is rendered almuit ivrlatu by the iutimute relatioiu
• timet to the fibrea of the posterior roots.
dcidli^ so far with the trncU of degeneration in the siiiiuil cord we
■Iwavt spoken of the detrenemtion an being th« ra«u1t of le»toiii of the
I eon) itaelC Kxp<:rimeiilsoniuiim3l».how«v6r,and clinical cxjwrlcace
shown that division or injury- of the fd>rcs of the posterior root* is fob
I hv tracts of ttcgenenilion in the spinal cord, though no damage wbut-
may have beon done lo tlic subslanor of tlto cord ilaelf. Tb<»e tract*
their amMtruMM in the nie<liuo jio*tenor columns, the exa<;t putli and
< of the oanatration differing wuh ih<* dider«Dt ajHitnl nerval. The
a of th« divtsion of different group* of ncrrca are >o instructive ttial
ay dwell upr>n them in detail.
the posterior ruoti! of two or thrrc lumbar nervea (on one aide) be
■d. an examination of the conl. aflvr an interval long enough to allow
«ralion lo be well esublisliod. will bring to light the following feoturea :
Srided roote will be found to have degenerated right up to ibeir eatnoce
ibe oord. A aectioo of the cord oppiuiie the entrance of the lowest
»d root will show no de^neration of the cord beyond ihst of tlie bundles
ree |M«in|[ in. A little higher up deigeDeratiou will be obierved in the
mI poetenor column eloee to the poateriur h'jru : and oa we ascend wo
hat thin dt!;teneration 6rat spreads over a large pi^rtion of the extertutl
■iur column, and then invstled the median poiterior column; thede-
7M
THE SI'INAL CURl).
C ration does not alfect the whole of iho median paeterior colimui l>ii
» intact ft #ninll <)or«il portion, roughly triaogular in ^iii>c, at (V- u[.
betvcen the tiseiiro nnd the dorsal surface of th« cord, aa well as WHDeportun
of the niorv ventral part of the column nearut the craj coiDiiiiasure. Sliil
a litile higher tip we should tiiid that tlegeoenited ItbreA had dbipp«nil
from the external iwrtion of the external poelerior oolumn oloM to (m gn*
matter, ihongh still existing in the nrnre median part of that column, an ktIi
aa in the median [toaterior coluiuu to tlie extent juat indicated, i^llli a liitit
bi>,'her up the whole of the de};enerHliuu would have dtMppeaned fran Ik
exteraal posterior column, hut the tract of degc^neraliou in tfa« mediaaf»
t«rior oolumn would remain, the extent nf ili-ii^enerniiun Ivt'iug^ ilependuit on
the numtter «f nniid which had been divi<leii. Liutly, by carrving lbs wv
tiona nlill higher un the ci>r>l we :iJii)iiM he iilile to tnu-ii thi* tract btla
ni»liau poaterior volumu nght up lu tlie bulb, wberv it would oonwtuo
imd.
If we divided son)« of the thoracie nen-c* invtmd of tbc lumbar wo AnaU
ohiiiiii v«Ty ftimilar r<9ulti>: a dvgi.iivrati(>ii of the external pnatcrior oulvmM
a litile above the i-ntrar)ce of thi? roolN, itprcnding aeroffi the column toiruil
tiK- mvdiaii lint-, and wholly di»ii]i pea ring at n certain height above, acott-
|wnie<) liy a dcgrnciatiou iif ii pnrt of the ra^dinn poderior column, rnafMii,
fnim H little distance nbuvp Ihv cnlnuieo of lite divided iien,'(vroote right kp
lo the bulb. This IiUler tract of dcg^-ncrnlinn would, however, not (xeiff
tlir >«nie position us that conecijucnt upon division of tbe lumbar nrrvn; ai
piji'ition wouli] be morv ventral, iioarcr the gray romnii»ure, and rvilier tmn
Inleral. Com|iiin' Fig. l>^'i, L),, whcrv Ir indicates the degeneration ilae In
section of the lumbar nerves, and dr, that of the thoracic nerv«>f^ If «i
divided xomo of ihc cervicul poslorior rool;; we should gel similar rcanlli.
with the ditlbrcnce thnt the Intel of degeneniliou in the DMdiaa poMMW
eolunius would uecupv a {ifaition iltll more veulnil and Mill DMtrekttn!
(Fig. 182, C'l c.r,), wiiile if we divided the »acrul nervea tbo irm-t .f '■
g«iieration would be duraul and median lo the tract belonging to the luoi i
nerves, luid would oc«upj more or le« of Uie triangle left below thiit tmi
(Fig. 182, O, B.r.j. The degenemlioti it will Iw underaUHxl » in all raw
vonhuwl to tne nune ride of tbe cord a* that <if tbe divided roota. We aay
ndil, in order to complete the story of the eAcct* of diviiuoa of tlw yunw'wt
roots, thnt (he vcetion leads to dcgwcnitioo of tbe marginal zone <LuMucr'*
tract) but this degrnenition rencliw for a cerlnin disianc« only up the enfil
and thrn ilisapjiean'. It will lie rcmeiiibfirrd thnt this »»nc is feil by libr«*
(of fine calibre) belonging to tlie external or lateral bundle of the poatcrio'
roots.
These results may bo interpreted as follnna: The (jrrent ranjoriiy of the^
•Bbrea of the posterior root, cut olf from their ganglion by ibe diriduiS '
degenerate eeniripelnlly lowarri [lie spinal cord. We have previously Mtrs*
that many of the tibrea of the root paaa into the exicmal posterior eoluni^
and nin up in ihet culuuin for tome dtsianc-e. The d^neralKMt obseirad ic>
this column fur «ume distance aliove the enlranoe of the divided roula abinrtf
thnt ibc Bbrea run lengthways for some diaianee in ihia column, while ihtf
diaapitea ranee of the degeneration a little iijgber up similarly ahuw^ that the
fibres eventuully leave tbe column. The appearance of d^neration in the
median puHertor column sbowa that wnie of these 6breB have |>n>Bi-d iniM
thnt column from ibe exterual poateriur column, and lite eoniinuatiou of thai
degeneration right up lo ihe bulb indicalw that Ibese fibre* pursue aa
unoroken course in that column nionv ihe whole lengih of the curd. Tliv
area of degenerniion. or more exactly llic luinibi-r of degenerated fibres id
tlie continued tract of d('gi:ne ration m the median piMtmor column is much
TU£ STRUCTCBK Of THE SPIKaL CORD.
I
I
I
lew limn tlmt in (hn U-miJOrnrj' or short iracl of defpiutratHni in the *-x-
terDKl jiuricriur (-olumn. Thiti (hi>«« lluit oumo only »( llio tibrm jinwiti^
inlo lliP extcrnsl poeleriur eoliimn goon tu join thu median ponU'rior column
an<l eo rcKch the bitll> ; lh« mi obvioui>]y Uikv nnn(li«r ]Mth, nnd nu hnve
alreadjr seeo reiison to Ihink thnt amny of tbcoi- end in tho gntj mutter of
the eonl. Hence of all tlio libm joining th« cord in n posterior root, while
soiue, aud tlieee ne may mM are chiefly fine libra, eoIeriDg the gray mailer
directly or paaung into the posterior marginal looe, anon make such con-
nections that ihe degeneration due to the section of the root spreads no
flintier, a lar^ nauber, and these chielly conn* fibres, Iwfore they make any
aucli connection peas into and occupy for some length of ihe cord the exttrunl
posterior column. We may here remark that though these fibres are spreiid
over the gresier |ierl of this column, they do not form the whole of the
eolumn ; Iney are mixed up with fHut^ of a different nature and origin. Of
tbcw fibrrn of the (Hjsierior root which thus tun in the txli-nial ponterior
Column uhiie Mill dependent for [lit.'ir nutritive activity on the ganglion of
tbo root, Mime, iodeea the greatir jmrt, Imve the traot and make such con-
OMtioni) in the gray rontler, that tlieir dogt-ntTfltion ceiutcx : otlierH, forming
the sniMller p«rl. piiMi into the mniian |Hi)>torior column, and taking up >
definite povitinn in tluil c^ilunm puniie an uuhrokcn courM; to the hull).
All imt filiri-s. iherclore. "f tlie putterior rootn do n<it en<i in the gray
matter soon iiRer their entrance into th« oonl. A reprvM:ntniivoof each root
is carrical right up to th<^ bulb by means of the median posterior column ; of
the axis-cyhndcr* which leave the ganglion on the root, a certain relaiivelv
small number punne an unbroken coiirw for some litllc distance through
the external posterior column, mul for the rc»t of their way through iTie
ntedian poeterior column, along the whole lougth of the cord above Ihe
entrance of the root until ihey lind nn ending in the gray matter of the bulb.
Further, each spinal nerve has this repreeenlalive of its posterior root placed
in a delinile poeili<;n in ihe |Hj«lerior median column, the arrangement Imiog
NHcli aa shown in Fig. 14:^, ihal the lower (sacral) nerves lind tneir place in
the more dortal and median part of ihe column, while the nerves abore are
•nccCMively placed in pi«tlti)ii« more ai>d more ventral and external.
Aa ftr as our knowledge goes at present we are led to believe that thia
median [KMierior tract is very largely made up of fibres having thi^ origin.
It allunls a cliaunel by which atli^rent inapiil^ie^ are earned straight up the
cord from the nt^rve-trunk without making eonneclioni) on the way. Wc
may rrpi-at that the path in eoufiued t» the ftame tide of the cord along its
whole length ; [here \a no crosiiing over to thr oihur side.
In tlw above description we have spoken only of the rewiltit following ncc-
tJoo of itu) Dovtrrior riMjta oulfide the cord ; hut it will be undenttood that
Minilar muita follow upon jwdion or of injury to or disoBac of the cor^t itself
•flbcting tbo poeterior oiluniiix or the bundltw of tho roota n« they enter the
cord. When such ■ Ifltion occunt ihervi may 1k^ ii)»tcrTi-il in the region of the
cord above the lecion a degeneration »f the external p'Mlerior column, reach-
ing some little distance up, nod a more limited dngeneralioo of a part of the
median p^islciior ciliimn stretching right uii to the bulb. The pneitixn and
fdrin ul the trael of the degeneration in the inclinn prittcrior column will
(lv|i«od on the level of the lesion along the Ecnglh of the cord, according as
it inlcrruplfi iho ascending repTcsentAlives ol the sncral nerves only, or of
the Ininbar and Mcrnl nervei^. or of the dorsal amt cervical ncrveo as well.
A complete section or hemi'^et-lton of tli« eort will produce results corre-
eponding to the division ua both sides or ou one side of all the nervee below
the wctwu.
We may add that while, according lo some observers, the Mraai) of Rbm
<:.
708
lirl'iiiK<"R 'o a particular root or group of rtM>H hnviii;; otice lak«i liiTl
|Hjaitii>ii in the luedian p<:«terH>r coIuud reoiatos uiicluiii^t until it rCMlio
l)ie bulb; Mccording to otben it duninUKe iu ares, soine of ils tibran iMkiof
viiDiieciiuDs in the cord itself.
M79. Tlie anltTo-taUral lucrudinp Irael (Fig. IHL*. use. a. I.) I* liw ntll
kunwit than either of Ibe two (OeccdiDR : it a iilw> more difliiM, thnt i* tu
any. tlie filirM underpins degeoeraliuii are more InritvJv iniwd iriili tlbm
of II tllflereDt uature ana ori([ia. It HpfMraru to extend (lnwii tW rord ini
lower leri-1 than ilie cerebellar tract, but ii« loKer limit liMOnlTMbacD
aocural«lj ilttlennined. .Since the defcuiieralion Inking iilncc In it ii n
MoMHiing une, it hua been inferred lliat it tervt-s an thf ]i»th fur aflcrtot
and indeed for MMivory im|>ulMA. DeigennrHtion in it io mvu only ftHct
DVCtion or injury iif the t>uhxliui(-e uf tbe eonJ ItJoir. mil iilU-^r divisiun i»r ikr
pMterinr roat«. If. ihun, it it to Iw rvgardod lu ii channel of aftrtnl
ira|Mikn pawing into it fruni ihu jxwterior roolt, tboM ImpiiIsM inual paa
into it along th^iM^ filiris of the ]>(>«tcrioi' root which fin<1 mvoiidnrr UdjiIik
«-ntrw ill Bomi- purl <\l' ihi^ gray miiU«r: in this mjMx-t llii« trart reMOibla
ibo coreWbr tract, ami dilfen from tlir nuilian |>wtvrior trai-t. The luttf
i< tiw direct continimlion up the cord lo iliv hulb of siirb (ihn-» cu are tiil
Uiuting lor their iiulrilivr aclix-tty to (he cells of the ({■"■K'i"" >»> ihl
(XMtorior raol ; the Hhm of iMh the former lni»t for their nuirilive aciirilT
to some part of iIk gray mntlcr of the oord, and prwiiniiibly tn iht^ Dnrvt-
oella of that gray matter. A further rcMtmblance mtween tbe atiteru-laioiJ
ascending iind cerebellar tracts must be admitted, if future reaenrchoi n*
firm the opinion of those who bold that tbe former like tlie Uilter, nt tlie itf
of the ooni, paiffl along iIm rwtiform body to tbe cereliellum. IikIci^, uiiikr
Mich a view it would appear probable that the autero-laleral tract i» •iiiH>lt
n more ditHiae and outlying' part of tbe cerebellar tract.
^ MO. We may now brietly jiaeh in review, somewhat as fullowi, the rhie'
facta which we nave tearued couceming the structure df ihn spinal rant
aliraya keeping in view ibeir physiological meaning.
Tbe ini|Hinam leatiire of the spinal cord ja the pr»ence of what we bare
culled "gray matter," and all our Kmiwledse goealoaliou' that ibe ini|urtatil
|Hiwcn of tbe Hpinal <i>r<l, by which it dimra ftxMU a thick niutliplc ngnr.
and hy virtue of which ne ^(iL-ak of it ax a i>ervou* centre ur oorics of cMUm
are in iH>niv nay or nthi-r aiMicialed nilli thia gray matter.
Wilb this gray iiiallor the fibns of tbe K]>inal iivrvw nr« cobtioclttL Ha
graater part of the librea of tbe aiilorior root certainly end in or mthtf uIh
origin fnuii the gray matter cloac to the attachment of the root, aod tlw ml
awtl pniliulily join the grmv matter at no great dkuivco^ The fibres of tb*
poatcrior root nin, aa we nave sovn. Jbr sonrn little dUtAooe in tbewbiu
matter, but if wo except ibc «pcrial Imndlc which ruiu in tlie mediu p*
iM^or ir»<-l right up too cord to Ihc twlb witliout joining the sfintl giV
nutlirr nl all, wc may aay that (be Ghn» of tbe poelerior root auo join m
gray matter not far from tliu atiiichmcnt of the rout.
Morphological rcasont lead us, as ire have seen, to regard the spinal oni
ns a kHck of segments, each segment oomeponding to a |iair of oerra;
nml even in the spinal cord of i»un we may revcqpiite a sei^nwotal fvmoi-
Work, iibM-uivd though thia is by fusion and overiaid by the aeveral iiioun>*
sural tracts. Each segnwot of this groundwork we may concetTc ufiD*
central mass of gray matter, eonoectea on each side with an ui)l«Ti»T iM *
jMeterior root, thus constituting a segmental nervous meclianium npMf'f
carryiug out certain fuiietiuns.
8ueh a socinent ha^ bc^en compared to a ganglion, but it dillV'r> mrikinj:';
from a gaagliou, whether of the posterior root or of ilie tpbueliuic snMiai
TlIK STBL'CTIIIE OF THE 8P1XAI. lOBD.
707
Ui'turo iii>il in function. A ;|.'niij{IIi>ii uiiil the K>^ir innllur uf ■
Dent ImIIi cixilain niT^'e-i'c^llif, uiitt kh far riTK'iiiMu i-iicli ulher; but
jhlniicii fur llii
niuiil pari cikIf. Iiib gniiifluMi tbo cunxUtitcnt
liT»-<-rll i« a >lrvi-liii)m(.-iil of tin; nsiix-rlinilcr gf « filirv iiilo u iiiic1e«to(t
|ll4xMly which lid on ibu oourae of the librr, and miiT, lU in n »plniK-)inic
KagUno. Ix) plnoed jm^t w-Iirro one fibre ilivliliv into Iwo t>r morv. ^Vfi have
|nr ttrkiiiDO that tbp cell, ibac i> I<> mv, the dik-Icur wilb the ndjiicciil wll
Ifaitaiioc. Bxcrcisn rh it»i)»r(iiiii itiHucnco on tbo nutrition, mil) "i on th«
kactional M-livity of llu- nrrvc libra, it net*, ii» we Iikvo Mx-ti, iu> n " trophic
tnira." There iire «lso rcnwciB fur thinking tbitt the mil aubstHiice w mure
Kulive, more rendilv retpoiijive to chiiDgist iii lU circuuiitiuircs than is tlie
Ks^yliittler at ramo dut4iiii.i' fruni the cell. But wg hnve nu satiifactory
riiteno* that ibe cell can automat icallv nrii^iaale Dervous iiDpiiU^s in it«elf
t the itutcuRw of its ovru inlrinftie ohaneas. Nar have ne any evideaoe
tst the cell can exert any marked transJorminK [lower over ihe impulsefl
jHoiDt; nixdi! the fibre : the iupulan wfaieb travel away from the t-ell do not
ppaar lo diUVr markctlly from ttune which travel toward it. The teverul
Mteooen in which there aeewed (u be evidence that eplsiichiiii.- ganglia ooted
I c«ntr«i either of reflex or of automatic itctiuD. ha\'e, aa we have Been,
■vkco JowD ; aiid it a not even suggcated that the {{suglia of the posterior
Hl» powwt any such powers. Th« gray matter of the spinal cord, on tlie
Ihirr IiHiid. an nii \m\f alrendy »eeii, and a« we xbatl nee more in detail, in
?wcinlly charactcHwd by the [wMBiwiion of reflex ami automatic a* well as
other' poircn.
In tiructute, moreover, nuch a *|)iiial «e;|i:nieni diffiuv Htrikingly from a
knglion and cxbtbitx ft-alurci iinKn<twn !n icanglia. In a ganglion the
trve fibrvK may divide, and in n «mall [icripbcral )rjn£lion tbc diviHion may
ire ri«c to very dulicatv fibriU; but tbi; fibro or fibriu rtuulting from the
tviaion leave the ganglion to follow their appropriile coureea; the diriiioo
trirca for dimniuD only. In iho vpiaal mrd, on the olhor band, both
Bereul and alKrefit (ibra divide in such a way thai their diviaions are loot
) view in the gray mailer; division here eeems to serve the purpose of
nioo. I'he efft^rent fibres of the anterior rout may be traoed nack aa a
rocew of a cell in tlie anienor horn. That ci-ll givei utl' other procesMB,
Ut iio one of these juxmimms is contiiiued on as an asi^-cylioder proc«a5
Crcu-hing acron the gray tnatier until it beooues a tibre of the |)ofierior
bot,or ua anything like auch an axis-cylinder proeeas. Ou (be contrary,
tl the prucMHS, exc«pt the axiacylindcr prociMi, divide into branches, and
bpiiiar to eml in nervoun libriU lust t^i view in the gray mailer. Cunireraely,
Un^ our knonlislgc of ihc juncii'iu of the |K>tUirior Dbnu with the gray
EiUcr in much more imperfecl than that of the junction of the anterior
HM, what wo do kooir leadd tu to believr tliai the tibrc« of tlie puaierior
Dot, cithvr by the meilintion of cellit, or by direct division of thi- axiii-
jrlindvr wilhtxit lht< mciliation of cidlit, nimilarly break up into fibrils and
re nimilnriy \tml in the gray mattiT. All the evidence goes to (how that
he antrrior and {fMU-rior rooii arv fiinclionvlly continuous; this functional
piitinuiiy i^ howovcr, eflectwl not by a gruae cmntinuily of axi*-cylind«H,
Kit in a peculiar manner through tite diviiton of brancfae« of norro-cvlls or
f axi»-ryHndorf> into the nervoux lanicle which forms such a spoctnl fvatur«
t the gray matter nf the t <<i'<l. We may, perhaps, venture tn rvganl the
;ray matter of the Migmvnial gr< Mind work, of which wa an? noir nlonc speak-
Bjt, ai riinatiluliii;; a nervoii" nriiwiirk i.ir web. fonue^l certainly in part bj
be rapidly dividing branchi-« of niirvc-Lvll», and prob.tbly in p»rl by iha
tivkNOUH of directly dividing ni-rve fibres.
Iji any ordinary section of tlie spinal ouni the gray matMtr presents to
708
THB 8PINAT( CORP.
groundwork. To
(I.
..f'
view miicli raontbiui ilr
»lnicttir
nr*' cUimt'd nov In hr iicnro(,'li«. ni'w to lip nervou* in imUirr, Ihr en\
mntwr in every wi-tion ehuns minicroiis ili»liiict ncrvt &\>m rminingiiiii
various dirpctions ; of thttc fibres n Cvm arc ordiDBir modtillmK-d lil>nv. mm
are non-iDtdidlfileit fibrcc, ihnt is to fuy, nni niiKnl ■xw-rvlimlcn'. mhi,
Othen, and these the more numerous, nre lh« peculiur mcdiiilaird fibm >\f
email diameter spoken of in i; 'iSi. A Inrj^e Diimb«r of tlie»e lihm. in-iM
all the lar);«r ones, ltioii;;h tliey go to make up what ire call gray nalUr,
are not continuous with, and do not belong to, the i;roai)dworK or aerVMR'
web, at all evenis do not form part of the grounduork wen in the iw
aeclioD as iheni»elveG. Tliey are itimply fibreia iraversiu^,' the ^roundxisk, iu
spaces of the neuro(;lia bed, on their way up or down (he cenl. or acr<w ihf
cord from one part to another. It may be that some of the finer ncdaliiKd
fibres do really enter into the ^roun<lwork, and so eonirilKite to ibe nemoi
«eb; but our knowledge is too imperfect to afford a clear decision on lUi
[loint. Our inability to define its exacrl limits need not, bo«re\-er, pitroit
our reeoguiKJn^ the- existence of the fzioundwork.
The |iTonjini-nci.- in this groundwork of th<- hirgcr »cr^-«-cellt ha* ltd t*
the CQDreiitinn llint the powers of the njiinal lu-ginent are exerciaed by thew
Derrc-cellii l<> ihe exclusion of Ihv other elenieni* of th« nrrvuua neli. fhil
such a view ba» not linen adtsjuatcly prnvrd. What ue do know i* that llir
nuclei and fell - hud itii of the (.■elln of the nnlcrior horn cxcrctK* an itnpotuni
influeoco on lliv nutrition of the fihron nf the anterior rnoi which pravnl
from tbcjn, and poivibly »\m influt-jiw ibc uutrilton of the other Imatba
of the cell* forming part of the gnnindwnrk ; and tbcw cclli' are prnhiUr
«> eon«picuoii» n fniturr- of every secliun of the xpinal n>rd hecaiue of Iw
imporluut Uisk intrusted to tbeni of mainlaininf; in due order the nntrilNO
of the long atreleh of motor fibres reaching from them to the routcabt
fibres or other iieriphcral or^na. The Rlires of the posterior r'intutsM
so obviounly ennnci-ted with the conspleuous cells of the gray niador: in-
deed, as we have said, it may be douliied, though the view ta mainiaine4 ^
some, whether any cell intervenes lo secure the coiilinuily of a pwlcriM
fibre with the groundwork, a division of the axis-cylinder wrvini; iliii [M^
C; and this Ik-chuie^ inielli^ible when we Iiesir iu mind that Ihe pMl«n<f
« are governed as lar as tbeir nuiriiioa in concerned hy the uervtcA
itf tlic Kungliiin on the posterior root, which ought pmhahly to be KoBttdmd
as much a part of tlie spinal cuni as the cell- >d* the an(eri<ir bom.
ncrve-ccU of the gim^lion is adc'iuate to secure the due nuiriliounf
nerve fibre until it joins the groundwork, aiid probably hei\m la tnr'
the nutrition of the groumlwork iiwlf.
Hcnoc we may )X-rbapi>, until fresh GVidcnco- to ihe contrary Is
fbrwanl, incline to the view that the powers of th« gray matter do
depend on the consjiifiious eclls alone or even chieflv. bul on the peca'.— j
moli-culur ronniitution am) naluro of the whole grouDawork. The nuclei 'r
the cell* of ihe nutcrior hum with the cell substance ndjaccnt to each a *^
the cells of the ganj^lia on the jMirterior root probably govern tbe nutrilit^
and *a the functional activity of the groundwork as well as of the tasui. '
ami entering fibres ; bul there apjie'ars to be w yet no coiiTincing evldet^
of anyolh<r [H-inlinr powers conlintd to the cells and iilisent from ot^M
parts of the groiiodwork. We may add that, in accordance with this vi^^
the other celts of the gray mailer, such oa those of (be vefieular eylim^S
are to be regarded as of itnportnnee for governing the nulrilioo of fibr
comniiasiirnl and others, starling from the spinal segment, and of the
ol' ihe gruundKork from which by their mediation the fibres aun.
ItJCTtTRB or THE SPI.N'AL
709
uiniiii; th'- fiia«li«iu or lli« grouuilwnrk of itic •egmcnl or of
iviay: iiujt\il*c* from iU
I 581. Tlic acgmental gmiinilirnrk of grny malU-r Wlonirins to cat'h pair
of •iniinl nvrva* ia «o fuim] with that of nil tlic mhcr paira lu to form along
ill* whoir l«ngtb of tho cor<l n mnv of gmr matlor which iippeAr«, uniler
ia vircimMMDocaatnllcvonu. tobevoDtinuoui in the mum that impuIsM
paM in all iliroirtioiis along it. But tmch spiniil Mgmeot l« in tidJitioo
i>Ct«H by means of Irttrtf of irhit« matter wilh parts more or Ifaa diaUiDt.
Tb« oroued prramiilnl tract i« such ■ longitiidiDal comitiiuiiml tract, con-
uactiog apparently each epionl eegmeot In succewon with a certnin p<irt of
ibo (■ortex of tho oorebnnu. Wo have r«uon to think, as we ah»ll «ee later
in, tbat iinplliMB descending this or that libra or tfroitp uf fibres of this tract
i^T« rise to the iaaue of nii>lor impulses along this or that fibre or u^oup of
fibra of an anterior rooi. We do uot al preKut know what is toe exact
manner br wliieh the libr« iu the pyramidal tract is eonnected wilh the fibre
of iIm> anterior root. [| seems certwu, however, that the conneotioii is not
■u the l<>rni of a fibre isolated fVom the rest of (he gray uiatt«r, continuing,
Hu to speak, Uie pyraiiiiditl lihre into a cell »f Ihe anterior horn nhtnce the
fibr« of llie aut«riur mot isiiic*. Moat prubal>ly the pyramidal fibre makes
eoDiMclious with the secmental groundwork spoken of above, whether with
or without the intervention of a cell ire cannot at preaent tetL The direct
pyramlfhil tract is a Iik<: tract of lewt (extent downward, and thu Uw known
anbero-lnteral dcstwnding Iruct i* probably of a similar nature.
Tbtr ccrvltellar and aiilrro-]at«nil uicwnding trai-lH arc in like manner to
he niganl«xl a« longitudinal commiwures between the Kuocttwtvc spinal tta-
■MOt Ddow and sonii; [MrC of the brain nborc Wo have nM«on to think
tlut tbwe tract* coiivev upward impuLseaof a nature which may ho called
tflbrenl, and arv. tlicreAire, in 8»nie way probably conaecbed witb the pM-
terior roots^ We do uot know ns yet the exact nature of the conneotion ;
but pr»bably in those caMS also the couimisstirat fibres are umt«d not directly
(Olbe jMslerior fibres, but iudirei-lly by means of the segmental groundwork.
And since these iracls do nut de^nerate after section of the poaterior roots,
but only afWr section or other lesion of the cord itself, we may infer ibat
their junction with the grotmdwork is eltect«d bv menn^ of trophic cells, by
^fcns of some or other of the odis ipoken of a little while before.
^Bhe median posterior tract seems to be a commissural tract of a nature
dUl^rent from any of the above. Through it a cettaiu part of each posterior
not is brought into oonocolion. not with its own spinal segment, but witb Ihe
bulb above, and aa nJih the hrain. which tbiu receive! dircot repreaeotji lives
of cacb attereut spinal nerve. If, however, a* aomu maintain, the bundle in
this tract starting from a Hpinal nerve below dimiuixho* m> it proceab upward,
tbrowing off fibrciii tn ]>aw vl«rwbert), though always carrying soini; Kbrws
right Up to tlw bulb, wc must add to the nbiivc toe further view that this
tract connect* aU> each po«lcrior root) iwt witb its own segment, but with
other nioru or Ii^At ilUtanl twemcnts.
$ 66S. .\ll the cvidenoo which we posMss goes to sbow that each strand of
Mch of th«M tract* nin« itolnt^-d. that is to say, makes no connections with
adj<Hniii'_- •tructurv at any [Mirt of its course, from it« beginning or end in the
brain and Its end or beginning in its appropriate spinal segment, or in the case
uf the RUHlian poslmor tract from its bcginoiog la the gaaglioD of a poae>
rior rout and its end in the bulb or in some tUitailt spinal segQent. In the
eroesed pynuaidal tract, for instanoe, we have reason to think that one or
more fibres run a quite unbn>kei) and isolated course from the cortex of the
cerebrum thnkuj^h Tariuus paru of the brain, •]«■( the wb»le length of tho
laorrl until they reach lite lowermust spiual aecniHilal meohanijm. These
rra
TBK Srt^AL CUR0.
i
ii|hW I
Ineli mem n m waj tn oaaaect («e Ngmtninl nirRhinutn wllb anpthtr.
TW NfBcoul mBAmaaama a«c bowvrer, coanvcteil tn^ilifr ; umI Uiv n<i-
— ai»«» betwem ibeai «■» to b« of twn klwU In the firvt t*'"*^^- ■>-* *'
have alf^df miggntlM, tbe wfBtnital pitcai of gny uinllrr mro *i> fiuri
tngBtkcr w to fi.mi «lut apricar* Cu b« • eootinuitv of gray mntt'T fn<tn 'iix
MM «f ibe onni to tike pUkt. TboQgb «* ainnot ■rtuitllr truck vur ait
UHoloficailr UirM^b, mud mrc «UII Um »wu* of iIh' i>livi>t<>I(i^icul nklutr if
tfc* kbjrinu of BfiTr>oeik, ibn*, ftod fibiili «')ii<'h nm^r up what wn Imw
ealWd Um giwindwock. «« msj wilh eoMkkrulili- prolmlnlily iiwiiiuc thu
tbt |MMan of Bvrrotw inpabn tHoag it ia dMrniiiiiitl lu much by
tUtJMi of UM iBBWml M br iti ualMHieaJ dicp'v'ilioii : ibnl, for in
rMtncliow to Uw Bow 0/ sa iapabc u« brought nhwii much nun*
qncad; bjr tlw reftiMl of tk* sideeata of imrvnus m»tier to take m
nabeaJordiHariMtnce wfaidi iatlMMWflce of the impiilEc; ihiit ie to
Mttleeolar Tuimiicti iIms by utaal bmks of ooniinuiiy in tbtr
ntaiter. Indeed, wc ksre stme ruoim for thinkinf; iliai fn'tunl ^tnn-igi
cuntinaity of DemKn RMternl ii not cvmtial lu funi-tional coiiiiunity ; il
s Dcrre fibril, for initanee. nuij pndocv iu due et)«ft on another xtvt\r iiW
or OR a nerv«-ceU, if suffimntly ia contact Kith it, tltuugh th« mien.
fai)» l» drmoaMnte actaal runlinuity.
But bnidrs tbe pay tualter there are areas of white matter whirh il^
bcUiog rithiT to tW n«'nre roots aa tkeee are makini; their wnv into tha
Buttfr, or to anv of the Iracti vhieh we have meiilioiHM). Thn^t ouni,
the strand* of tihrr* whtrh do n»t undergo eitliur BHoc-ntling or ilmcvudini;
dtgeiH-falioD wbvn part* of tbi- *ptnal cord are injiin-il nr diH-jwiil, Tht am
of white mailer )«ft when nil the vnriouN tractn nf aorcndiiig iind dcM-rndiiif
dtgenerattoo detaiM alxirearv titkcn nut. neemi, at nil ovcnts in thr ['
parta of the cord (Fig. IH2), rdaiivcly i^mall, aiw) ftitiiro obaervaii'in-
cvinlinue tlill further to r«daco il ; but it mixt Ik- n-momlM-rrd lliiil noiw.l
tlie above-na«uiioD^I tracts are "pure:" they arc ill) more (ir \r» mixi^t up,
aid MMne largely mixed op, with Sbrcc nhich do not d('i,'i-iiiTati'. I'ut
Inowled^ ia at present too scanty to nilow us to make any ^tnieini-ot a>ih
confidence conceminK the flinction either of the Hbres forming tlienlwr
natter not yet marked out into tracts, or cf the Bbres scattered amixig t^
acknowledged tracla. I)ut we iniiy, at all events provtsiooally, aiuin* thai
IbtMi fibres serve in the main aa ooiomtMures ronnectio^ tbe auceemn^-
mental medianisuis nith eui:h other: we may eondnde that changes taklag
place in one vrgmental merhauiitm can by means of titew fibres produM ti*-
relnird chnuge* in M>mc ixher dtMant segmental mevhauisio, without cslint
into action any of ilir gmy mnttvr of the iuterveniugsegmenliil mn-hanMii^
11ie commi>«tin-9> wbii-U no may MipiKiw to be iIium furniKbi-il bv xUtr
Dinder ■»< JoiigiludiMsl comtnisnim ounnoctins: tbe M-^oH-ntnl nKcbaui^*
of ili« «tnie Intend hnlf of ih* aninal oonl with each other. A irantnM
cnnedi'in brtw<tn the l«o Inleral bii1vt9> U afliirded in »i.tiK! meiiHUfvtf 1^
anterior while i-Dmmi«urL'. We nhilll •«. howevi'r. Inter on. rrawot Ii*
Ihiukin); thnl many impulses besldw thfln pSMing idong the iini«i>
mifBUie cross from one udoof the eottl to ibooth'T: luid iheef, w'
pass nli'O},' dipiinct libre» or along ibe pi'n<'rnl t,'roun<lwork. mux tn
ttif gray inftller of iW irthmiis forming tlio nnlcrior and poettrior gray n«
mlMUrea.
Thus, as far as w« can see at present. tli« ^pinnl eord oonslsi^ of a teno
•rgBM-ntnl mefltnni»ma with tlieir respeclire nlTcretK ami efferent rnoUC
gray matter of Ihe several segnvenis l>eing continuous nlong the corn)i
encephalic tiee «f white matter iKtueen the several segments and lbs br
»
THE REFLKX ACTIONS or TUK SPINAl. fOBD. 711
of liingituduial oommiMura) Uncts ctuiuMting b)g«tJi«r ihvKvvrnI MKiKtilitl
nwctiauinnw. bikI of traiisverw ootuniuaiircs ninning iNrgvly in tao gmy
maltrr.
The nKPLKS Acriom op tub Spixal Com*.
i W3. Tn ihc jirecedii))' jmrliou of t)iia work we have repeatedly seeti
int thnii);h we chii leurn niiich cDn<«rniiig the workinf; of an orettn or tissue
or imrt i>i' il>e body hy sludyiDg itM lieliavtur when isolated fnmi ti>e nrat of ibe
l)>Mly, ftll the coDeliuioDs tbu> guiued have to be checked by a study ol' the
bvliKvkir iif the aune ofgan or part while it is eiill an iDteK^l [tan uf tbo
intact body. All tbe Mveml oreaDH nod liasue^ are ta bi)Liod to]*etber bv
vmriiHia licH that ilw actioiin ><f i.'iioli ile|ieu<l on lfa« aotiona of the rest ; and to
mty tliat tbe life of each pari in ii function of the life of tbe wlM>le, ia no len
true than to «ny thni thv lifu of the whole ia n function of the life of each
pan. This u enpedally borne in utmn lu nbeii we c»mo to itu<ly ihe actioiia
of tbe lYtitnil nervoua avMcni. We may, uu anutomicol grounds, Mepunite
tbe a^inal conl from the nrnln ; but when vie come to coiifider the re)|>eclive
AincUOtK (>f the two, we arc brought fac4^ to face with thi'. fact that in actual
life ■ lai-BO part of the work oftho bnuii iacurriisl otiL by meaiuof tin* apinal
ami, and CODveiwcly the npinal conl dm-s its work hiibitually under ihe lutlu-
•Bcc of. If i»l lit tin- diritt. litddiiig of. the brain. Wp may gain certain cou-
rtutioiM byttudring the Wbnvior of the f^piiinl mrd iKiIatdl frota the briiln,
or nl* part* of the »piiMil conl isolatcl from c^u-h otbiT; but wc must be c\'ea
lUorr cniitioUB titan when irc wcro •IntHitg with other |>ana of the boily. and
mii«t grmlly Imitate to lake it lor griinto) that the work which we can
make the spinal cord or a part of the spinal ifird do, when isolnled from the
brain, i* the work which u actually done in the intact liody when the brain
tufl spinal cord form an unbrok&u whole. Mun!ovi-r, tliin I'auli'iu l>eeoniee
■ncnannciy neeoaary when in our itudies we pass from the simpler nen-on§
•fUeni of one aDtmal to i)ie itiore t^nplex nerrous vratem of another ; fur it
i* by the complexity of their central nervoittt Ayalems, more than bynnvlliing
clae, lluu tbe "hi);he«t" anlmaJa are dilTercntiatL'd from those "below 'them.
When we coniptire ii rabbit, a dog, a monkvv. iind a man, the difleremxa iu
tbe viucular. digealivc, and nspiratory wyvtt;nis «f tbe four, utriking an they
nay aP[>vHr. rink into in»igniliciincc <viiD[>arci) with the ilillcrvnci^ exhibited
by tbcir mjirclivc rcntral m-rvons *y»tcnu. WV need caution wIh'u from
tna rwult* of cxiwrlmcntJt on diigx or rabbiln nc draw coDoluKion* at to ihi^
digMioo or circtilalioii of ninn, but wc nenl far gr«uler nnition wbtn from
tlu bthavior of the i«olalc<l spinal cor^l of one of thcw animals wc infer the
b*bavior of Htv- intact spinal cord of man.
A further difBculty meets us when an cxperinieDtal invntigalioii entaila
operative inierference with the cenlral nervons system. Rcmoral or section
of, or other injury to parts of the hniin or spinal cord is veiy apt to give riiM*
in vanitig degree to what is known as "shook." The entling or tearing or
other tcsiuu of any eon>^iderable mm* of Dervons substance aHccIs titc Hctivity,
not only of the htruclurcs imnieilintely injured, but of other, it mar be Ear
dialant slruetures. Tbe nature of " shock " ia mrt as yet thorou[;bly under-
ttood, but may |>erba|ie, iu pari at all events, be explained by re^-nrding the
lesion as a very iH>werful stimulus, which, |>anly by way of inbilulion but
■till more by way of exhaustion, deproees or auapends for a while nonual
ftinctions, ajid thia trive* riae to temporary diminution or low of ooiudous-
new, of Tolitiun, of reflex movenienla and other nervoua aotions^ Tliu» a
•o-tioM throuffh the A|iiiuil cord, ei,'en when made with tl>e abarpeM loslru-
tuent and niili tlie ntnui^t tkill, m> as to avoid all bmining as much m ponl-
TUB spircAT. eoftu.
tile, limy for ii «http inifiMtni) nil rvikjx tictiviU- «f tlto corj. t>r ii>il«Ml «W I
ulivioits activitim of Urn wliuk- central iwrvuun ayittcDi. We niBV aiM ik
micli a "ahuok" of tW contritl nvrvotw iiyiitvm may aIm> Ii<^ |)riiliir>:il
tniddeti lesions not bmring diroccly oa the mntrKl D«rv<Hu s^rMvia, u
iuKtaiicc by exteanvc lujiirr to » limb.
Moreover, in rniiny caaea in which ihc cITocbi of ox (Mr i mental iDtn^enm*
have bran natcheil for Bonw oonsidpniblp time, diirs, tnnnlha, or yeani ifiit
tfae operotjon, it hns been obKirved, on th« nne hand, that pheDom«na «hkJ>
an conapicaoua in the early jteriod m«V eventually itUappear, and, <m Ibt
other hand, that aciivi(i«» uliiob are at lir&t abaeot may later tm make tkttr
nftpearaoce ; rooremenU fur iD^tanL-e whiek are at first fr^qtient nfVer a whilt
Aw auay, ami oonveraely, movenwnts which at 6r«t aeeoied iiupoKible an
lalvr on easily achieved. We have to dUtiiivuish or to attemtil to diuin-
Kuiuli between the teiD|>UTary and the lasting cdaria of the <>]>er«tiiin, Incluil-
fng auonjE the former not only thoK of ordinary "shock," hut nthen ft
ilowfir d«vi>I(i|Hi>unt or loti^r duratiixi. In niwiy instanow whvrv a )«tt<f
tbft <!<!nlriil iii-rvouK syM«m In by wwtinn or othnrwbe Hudd«uly (oparaui
frwit) till! rmt, lint ]>l)(Mi»in«iia MiKgi«l tlial tltc «>'|iiirat<'<l tMrt i« lit flnl (ifi^
fuuniily influiint-vd ii.-> to iic iictivittos by tbi; withdrawal of rarivu* inlluawfi
which |ir«riouiily were hdue exerted u{ion it hv tiw KHt of the «yiitcni. but
later on Nc«i>iuraoiiut«H itself to the new conditions, anil Icnnw, so to xiwik,
to act without the help of lho« influonocA. And indrvd it u |»whla thai
some of tbo eflecU of even immedintc " shock " may be due, not its suggtateJ
bImvv, lo the actMMi of an inhibitory or cxhaUBling stitniituB, but Ut iht
sudden CMMtion of habitual influences.
8till. iu spile of all thes« ditiicuUies.ii is puwible notoaly tt ■seerliiin Ai
workinK of an isolated portion of tl>e MUtmt nerrous aVBteni, but e^-en u
infer from tli« results some conclusions as to the riinre taken by that pMl*M
in tJi« working of the entire and iutaot systeni. There oan be no doubt, tt
instance, that the Hpinal oird am. (|uite apart frtifli the brain, cany oai
variuua reliex n^^tions, and, m4ire<iver, it duee carry oiil nctionn of iliis kiad
wlien ill the intact orfconitm it i.t working in contiict with the brain, ladixd.
the carrying out of various nflex aeliona Mens i» bo <iiii' uf the nvwi ituj>v
tanl fiinctions of the spinal cord, m much •» lliat lh'>ii|>h the brain, "i >'
len»t, piirUi uf Ihc brain oin aUn anil do develop rx-llox acti'm*. Uie npiMl
(•ord ofli^m tbu beat ficUl for tlw uliidy of Ihwc actiiing. Wo linre alrwl*
'$ 101) touched on the general fcutiirt^ofreJlrx itclions. iind (.•iMwIiert htn
tncidenldlly dwelt on particular instances; we may tlterefon onofin* M^
selves now to certain points of special interest.
S 084. Keflex movements tire perbnpe bast studied in the frog and otbtr
cold-blooded animals, «Dce in these the actions of tfae cord are leaf drpendnil
on, and hence less obecured by tfae working of, the other parts <if the otutril
tier\'oti» syatero. They obtain, however, in tne warm-bloodwl itiaoinml tho, tat
In tJtCM special prejiaraliona are necessary to secure their full ilevelopmMi-
In tie JVog the alii>ck, which, as we have said, follows u|>on divinua nf tb*
spinal cord and for n while suaitends reflex activity, soon paasM nway ; witbis
a very sliort time ntler the bulb for instance has been divhled tlie tnoM MB*
plicated reflex niorenients can be carried on by tJie frojt'A spinal cnnl "bM
tlie Appropriate stimuli are npplK-d. With the mammal the caaei»vtf5_
difieretit. F(ir days even afker division of tlie spinal con) the Mrti of I
hiidv supplied by nerves spriu gin j( from the eord below tJi« sectlcm nurt
hibi't very feeble nactiona <Mi1y. In the do^, for iiulanoc, ancr
the spinal cord in the lower dorfial region, the hind limbw hang i.
motionless, and pinching iIh! himl foot evokw as a nsixHiM' (riiltcr m^
irreguhir movements or none nt all. Indeed, wur« our t^mervatioiis lii
TUB KKKI.BX AfTlOSS OP TIIK SPISAL lORD.
ns
laliiit |>i<irtoil irt mighl infor that ihf nsSex acllop* nf the tpiiinl eoni in Uw
aaiumiil were Imt fvclilv ami iu>i);iiificnnl, If. hnwcvcr, the miiiniil iw kept
■livo f.tr » l"iigrr imrifxi. f.ir wvck^, or lnHlw rtill for iDnaiho, Ihouicli i»i>
oniun »r rii!<iu'ritiii>it of tin* >)>inxl i-»rtl tiikw pliu>c, rvdox movmii'iii* af a
poworiul, vnrtii) hii>) L-i>mpI<'(o charact«r maailtRt ihemwlvct in tlic liiod
limbv Mid hinder pnrt^ of ihc budr; ft verj feeble slimuluM uppiivtl l<> the
•kin of theso rainuiu promptly eivct riao tu exlengire and yitt codnliau*
movoraflBta. Indeed, the more the matter ii sludifd, the stronger is the evi*
dance thnt the nAex move meota carried out by isolUed portiotwof the vpiiuil
ownl of the mammnl are hardir le» deKnite. mmplete, and nnrpuwttil, lliiin
witneaBed in th« frog. Il 'n wurtliy of altcntiun. a§ tiejiriDf; oot thai
rka made above oii the ^rent dit)(rtentiutii>n uf the central nervoiie ^ystvoi
in the hi|;hiT antnials. that the rellex phenomenu in RitiniiualB vhtv vory
touch not uulj iu difl^rent ■pet'ii'ti but iiUo in different individunis ami iii the
•Ktue individual under dilferenl dreiiuistancM. Kaoe, a^, and previous
iraiain):. wetu to have a ruurked etftK^t in deicrinUiiuft the ext«nt and char-
acter of Ihe nAex actions vrhii-h the Hp'tnal a)nl a capable uf ennjion out ;
ami ihi»« seem also to be largely influenoei] by paMln); oircum»tances, sudi
as whether fixid hait been recently taken or nu. It hiu been OMerted titat
the iaolnteat apinal oonl of the mhltit, nhich hax Iw«n the «ubject of so many
experini«nta,H,as oomimred nith thni of the dog ami many other nianroalt,
singularly deftd«ut in the power of carrying unt complex n'fli^x inovenients.
In ■(tidying n-tl<:x actions in man wn an- incl with the <liirii;nlly itiut ne
nerar have to deal with a pircion <•( thr >riinal n^rd xsparatod fri)i:i the rest
of ttw central ncrvoii* ■vdlnm niiiltT the tiivnrahlc circunutani'cn uf ciLl>cri-
mpittal invrslignltiin. In mnn, we inii^l be cxnivtil to examiiw reBex actiODt
«tth«-r whiU- the irhnle nervous systoni i.« iuiacl. or when a portion of the card
ha* b««D wholly or {uriinlly scprtratcd by «onie more or Icm diffiiH diMMO
<ir by some accident involving more or law crushing of the ncrvoas Blnictuna.
Hence, the cautiou already given, as to drawing inferenoea concerning man
from the reanlta of experiuiRnts on aniumU. acquire stJII greater force.
i 0S&. Couliuing oumelvfls at llr^i to the rutulu of ex|>erimeuts on animals
, wa may ny thai in both cold-bhioiled and w arm -blooded animab tlte salient
V0bMure of ordinary redex actions is their |>urpatiefiil liiaracter, ihouj^b everjr
jr variety of movement may be witnewed, from a simple npoam to a meal oom-
j»l«x mameuvre. And in all reilex nioveuienlik. both *im|>le and compiaz,
wv can rrcogniiee certjiin determining influeucu) which more or I«u directly
eiMilrihuio to iho shafting of tliid purposeful character.
Thus the fltaturm of any movement taking plaoe as ]iart of a reflex action
are in pari dettrrmiiMtl hy the oharactent of the Btfcrenl inipulwat. Kiniple
MTVOU* inipuisca genvratod by the direct Mtimuhitiiui of atTrrvnt uervv fibrea
gmarmlly rvnkc aa rcAcx movements mrrvly irregular i>)ia.<ms in a few
nmelM ; whereas the iDor« o>mpIicat«l difleraiitiat«l H-RHorv im|MilnB gw
•laud bv the application of the stimulus to the skin, readily eivo riaa loi
lanaand purp'iwful movcmonts. Itiscaj>tcr to produce a complex reflex
action by a >lighl ))mDun> on or other stimulation of the skin than by eYOll
slnwg iodoction -shucks applict diroctly to a nerre tmnk. If in a hraiDlaa|
IVog. tbc ana of skin ■•upplicd by one of the dorsal cutanaooa nerves be bsm*
rated br section from the rest of tlie skin of the hack, the nerve beini; felt
attached to the mum of skin and carefully protected from injury, it will be
fiMind that olignt stimuli npplioi lo the surface of ibe piece of skin cosilr
evoke reflex actions, whereas the trunk of the nerve may be stimulated witb
even strong curreais without producing anything more than irregular move-
nents. In ordinary meuhanical and chemical stimulation of ihe skin it is
not u single impuUe but a series of impulsw which [Mu*ei upward aloni; the
TnB si>i:«Ai. ronu.
MDSory nerre, tli« i-linngiv in wliirti may be r<ini|i«rMl tn tlti.' chunttM ia i
motor nerve thiriiig tctitniii^ In cv<tt' mfl«K notion, in fiirt, ibp o-nlnl
inrchBiiisrn m>v )>c Wkcd u]H>n o* hi-ing thrown iiilo notivity ilirouE^ t
Rumination of iW nfTvn-nt imj>iiWi> mncliin); it. Hcijw whil*- * n-tlux :•' -
U ntiiililr cnllnl f«rlh l)y cvitn fi-cbk- in(lucl»i>»-tih<it-k» ii|i|>lteil !■> l!'
if tlivv iHt repcnti.tl nifliriciilty na)>tdly, k dolttun' iDiluviion-shock ii in*)*
fi-cfuikl nnltM it lie Klrvrng rnoti^h ii> cniiM in iho (kin or norvn ehuievtf
Ml ckctroljrtic nature sutbcicnt to give rite of thtmMlvn Ut & aarios of in*
pulMi.
£586, When n mnscl? ii- thronu inln mnlrnotion in a rvfl«x aiTii '
pilch of the ifouiid which it )^\'ee lortb doee not varj- with th« stiniiii
IS coDAtiini, bein)* the snmc an tliHl ^-iveii forth by a muscle ihrow n iiii-> <.'*■
traction br the will. From which we infer, eveii bearinf; in mimi th» iH^
ciiJ^inn in ^ >fO rouoerDine tbeualiire of the niiiscuiar ^xind, that in a rcte
action the afferent imfHilMB do not !-im|ily jmss ihrouch ihi- iviiire In tkl
same way that they iiags aloii); nflereiit nerves, but are prufoundly modified.
And in accordance with thi!> we lln<l, aa we *)tnll see, tliat a reflex actnia
taken up an aniouDt nf time, the grcnier part of which Lt spent in the carrr-
fng out of the central chaugm, and whicli though variable iw alMura aioch
lon^r, aiw) may be very much l"n)^T, than that tnkm up by ike mn
nuBBfcG of a iiervotni iropulw along a cormpoiKling length nf iten'i~ ^*■'-
The term reflex action ■■ therefore nn untwitanb' one. The artcrrnl :
ia ni>t xtmpty reflectotl or turned aaidc into an cffi-n-tit channel ; <tO tta ■mut
nt the r<rntr)-, it ularlN change* of a different nulnro from ami mort cumplM
than il* own ; and the miw of effcn-nl inipnlw is the r<»iill of th<iM muff
complex chanec* not the merv continnalion of the simpler aHeri'iil tiapulf^
In other wonis, tbc inlorvat between the advent nt the central organ <'
nlh-rcnt, iind lh« exit from it of etfcreiit impulsn, is a busy linu> Cit lie
DftrTOlM (ubslnnce of that organ ; during it many proocMes, of whicli we hare
■t nment very little exact knouledKe. are Iwing carrie<l on.
§ d87. The character of the movement ffirming part of n retlex aetloait
abo iulluenced by the iiilenaity of the stimulus. A alipht alimuluB. ra<}iM
gentle contact of the akin with aome body, will proiiiice one kind of mni*-
ment ; and a Mroiig atimulua, aueh ma a almrp ]>rick applied to the aantr ipri
of akin, will call forth (juite a difTerent movement. Wlien a decit{iiutf<l
make or newt it Mi»j>ended and the >kiii of the tail ali^htly touchol *<tll> tlit
finger, the tail IwJidn toward the finicer; when the iikin ia pricketl or batX.
the tail w turned away frvim the offending oliject. And m> in manv "H"
inataiKiv. It mu«t be rememliercil. of i-ounw, thntadifterenoe in thei:i
of the MimuluM cniaila a difll^rcijce in the charai-ten of theeflereni {»(■»■-'.
gnille CODlact gi\m rite to what we call a nenmlion of touch, while a ^f
prick gives riw- to pnin, consdousnew bcitig diftVrently eflN't4ii in tlie w
caMf hecjiiiM the aflerent inipiikta nro dirfcn'nt. Hence the intlann* ■■
i|iK-Ktion are in reality fiiller illustrations of the dejH'iidence, l« whiefc w»
called allentton above, of the chitraclerR of a reflex movement uu the HM^
ncten of the aflerenl impulses.
Furtlier. ns we have already pointed otit ($ 101 ) while the motor i>i|w^
started by a weak atimutus applied to an atlereiU nen'oareirARsmillrH rW
a few, tlioee started by a strung ^limutiia rany apread to many eHerenl u«rv^
Oranting that any panicular dllereni nerve » more especially aaaneiattd »il*
cerlaiit efferent uervea than with any otliem. ao that the reflex impulMf ^
eraled by atferetit impulaea entering the «>nl by the liirnier i-aa* with tiw
least reaiatfl nee down the latter, wv miiat evidently admit further that rfh**
efferent nerrcn Ut ab(», though leva dinwtly, connected with the «ome if^
ent nerve, the paaMgc into the avootid efiereni nerve meeting with a ff«"
TOE REFLEX Ai'TIOSS OF THE SlMSAL lORD.
715
rrmtik
try.
tra
rcHUtiino^. When n Trog in |xiiMn<.il with ilni'cliiiinv,
m tlighl loiicti on nny {mtI oI' iIu' ■kin may ctiiiM> <v>nvulsions nf IIh' whuJc
butj I tltmt » to MV. the nflVrcnl impiiUcs |iii>«ii)|^> iilnng Any ninglo aScn-nt
MTTv m»j gtv« rttts to iho iliwhiir);i.- iif e-ti'eicul itii|niUM along any or nil ol'
tW rMrrrnt oema. This pr»r<-a thnt » phjYiolo^ical if ni>l un iiiint^ninil
continuity obUiiiw betwi<cn all the |>iirl« of llic »j>iiial coni wliich nir cun-
ntsrd in rcHex acli<<n, thai ihc nervous network iiilervcniiig between thu
■ftfvnt anij eKervni fibn-f I'orniB alung the uliule leiiKlh of the coril a fiinr-
liaDally continuous tielil. This coDtiiiuouit iielODrk, however, we must eu|>-
be market) out into irac-te praaeniing f^reuler or leM resiaiaiice to the
of the inipitlses into wliicii alTereitt impulMH, coiuiiiK slontt tbie «r
■fl«rent nerve, are trnDsfuriiieil on their advent nt the network; and
linjcly the path of any series uf ini]>iilitc-i in the uetwork wit) be deler-
.[d«iI larKely by the energy of the ullereiit itnpiilM-«. And the action of
c'hnine luny be in pari explained by 9>uppo^in|r ihut it n'[1u<-e3> aud eiiual-
tbe iioriiiul resintanci- of ihitf network, tut that cvrii weak iinpulM^ travel
r all its triicin with great ease,
§ 568. Further, the movement, fiirming |Mirl nf a reflex aclioii, t'uriefi in
.rooter u<ToriJin^ lu the particular part <>f tliv bndv to whicli the Mimulua
pptird. The retlex aetiunit develoixil by *timiilattnn of ihe inlenml vio-
are difTerenl from lh<iM: excited by ^liinulntTon of (heskin. We have
>n to think that ibc cmtnii-tiou of vr other ehaogw iu a skeletal miuelv
itosr imxluee, by rvllex action, eonlntctioDR of other nUKlw: and nich reflex
•aUQtit al»o <iifl(rr from lhc«e vtnrled by Htimiihitifin of the akin. Id reflex
•cHoiM Marled by applying a tfimnliif to the skin the movement* vary largely
aocording to ths pnrtirnhir area of the ikin wbieh ia oflectcd. Thni, pinch-
iM ihe tbUa of fkin iurrounding the anus of the frog produces dilfennl
cncU from ihow wiliicaKd ivhrn the tiank or toe is pinched; and, speaking
generally, the stimulation of a particular *p<)t calln forth particular nwve-
ments. In the caw of the simpler reflex movemeiiis, it appeara to be a gen-
eral nile that a movement started by Ihe siiiiuulalton of a aenitory surface or
region on one side of the body, U developi-<l »u the same Hide of the body,
and if it spreadH to the olher side, mill n'mninii inibt inten.-te on the ttame aide ;
ir movemeul on the other side moreuver in Hynmn-trical with that od tiie
tna tide. It hu» been nminlained ibnl " cro«Md " or diagonal reflex move-
nta, Bi where miundiiliun of <>ne fur<:-f<iot leads to movements of the oppo-
hind-limb, do aul occur unle» lumc [Hirtion of the bulb he led aliacbed
ibe tpiiial con). Seeing that loeomotion iu fonrfoolcd animals i* largely
ed by diagonal mnvenienlK of the limlm, nno WfMild rather hnve cx|)ecleil
find the spinal con) ilwlf iimvidiij with nirchitiii«ni* to assist in carrying
in cMl; and, tnih-cd, it h aJlirm'-d tbat in the msc of cold-blooded animala
of nuinv young nianinials.alUr division of ihc spinal cord below the bulb,
'^tla •timntation will provoke a dingimal niovenwnl, slight pmpurv on
ir»fo»t for example givin*; riw to nioveiuenu in the o|>pu8ile hind It^;
ig stiniuhif, honcver, will pniduce un ordinary one-aided movement.
_ when in a Aoff Ihe euni hiu tHwn dividml in the lower ilxiracic r^oQ
•<> tlinl the hind limbs depend on the lumbar cord alone, a rhylhmicnlly
Tvpeai<-d drawing up and letting down of the hind lirnb^ \» nitiieseed wlwn
Ihne areallowol to bang down; and these movemeniit, nhich appear to be
of a reflex nature exi-ited by the pendant lucitiuu of Ihe limbti. are often seen
lo aJtemate rej-ulBily in ihe two limb». the right leg beini; extended white
the left leg is l*eii)g diiiiin up and rirr iv-r^. It niav further be obwrveil
Ibai if the foot of <>iie pctidant limb lie piiiche>l nhile tite other limb i^ pna-
rifdjr flexed the flexion of ibe limb which i* pinched i> acvonipnnied by an
7](t
TnS SPIlfAL CORD.
vxtt'iietou of tbc<itiii-r linitt. In tlmi^ twiwicu. bowcTCr. dlfl^rvul ftmouW
M iilready urgwl. tlilTL-r from c«ch ithor.
§ SBB. Kroii) tbeiM- iiml similar (iliriii>nii-iia w<- mny inl't^r thnt th« nnrmii
■Ntwork spolcco of above U, m (4> »)M^k. iuA{)jk<(I out into norviMu nuctw
\ttm by to« eetablialinient of linos of ^raiiicr or 1cm rMutwiM, w UuU th
dwturlHmiMe in il {[enerated bycertiiiii Di>l«rcni ini|>iitms arc dirwt«d int*
oettain efferetU ehAniKU. It may Ixr adtied ihnt though fioiiD)>i<rui»i»lT jm
(wspful luoTemeiiU seein lo need the coDCurreut ncci'm of several aF^nii-aunf
ili« cord, and ta a rul«, the itrmter ibe teni{<h of lb<3 i-»rd involved the mnrr
complex and Ih* more disliin-tly ptirpfiTeful the raoveiueot, still the mot'-
nwnis evoked by evt^n a ^^iiit-iiL nf the curd may be purjHMeful in cbancM:
henee we munt co[iclu<lo thai nvery sef;awot "f the uervuus neiwaik ii
mapped i>ul iniu niL-chaiiiaiii.->. Itut the arranxenteat of thi>ae meebkunaik
oapecialty of llie more complex ones, a not a Kxed and ri>;id ontt. We ran-
nut alvray* prviliot vxaclly tlwi nature of the moreaaeiit ivbicb nil) na»i
frrtm the Htim illation of any pnrlii-ular •put, beratiM ibe reault will rut
acoonliiig to the cundtlion iif thi.- Hpinal catA, e«peeiallv in rolaiion to 1^
Btrenglh and ehiiriictcT of the I'limiiluit. Moreovor, under a chang* of fir
oumsUnowa movctiwDt quiti-dilTenfiil from the normal an** may dim let If
ap|icnranco. Thus wh«'n a drop of iicwl i]> placed on thn ri^Ht tlank oft
brainleea trng, the right foot is alnrnst tnviiriably tiwd lo rub oA' lti>- n-r-
in tliit there appean nothing more thitii n mere " mochanicnl " reflex
If, however, th<^' rif^ht leg he cut olT. or the right fwA be ollierwiaa bn :
from rubbing oil' the aeitl. the leA foot », under tbo exeeptkmal .
MAOoea, aaed for tlie purpose. This at fin^t sigbl looks like an iniclii^'Qi
choice:. A choice it evidently iH : and were lher« many instanoes nf dIma
and were there any evi^leooe of n variable automaUsm, like thai wbioli **
imII " rolitJoa," being manifeated bv the spiniil cord of the {n*ii, we tlv!!''
he juHli6e<l in i4u pinning that the choice wa^ dt'termined by an tiiU-l! .
But. ax we shull buvc! ix-cawon later on lo )>otnt out, a frag, deprive
brain so that the spinal (^i>rd only in left, mnkiu no stxintaneoiiii ni i-'
at all. Hucb an i-nltre aWnoe of spontaneity is wholly inconaistviit "■■'■■
imnhmIod of inttflligvnne. Then a^nin the above vxpenmcnt, if not the odIt
imtance, h at all evenu by far tliv miut striking inMaooe of ohoioi> ua tk
part of a braiiilw* frog. We are, Ihtrrefore. led to otmiitudx thnt ihi* \itua^
mena must be expIaiiMx) in sonMOlht-r way tlian br bt>i»i; ri^fcrrrd to lb
WM'king gf an inl4)II!gMici\ Morettver, t)ii« oonclumon ia sutiporuwl hy At
behavior of other animals. Thus simtlar vicarious reflex m >vviuiints ebit
be witneesed in mammals, though not perhaps to such a striking oitsni i
in firogn. In dogs, in trhich partial r«-m'>vnl of the oon-bral bemisp^' ' '
AptNumntly heigntenod the rcnei exdiabiticy of the apinal c'>rd. the -
able •onttching roovomenla of the hind log which are called ftrth l>i' iniH'
latinKa particular spot on ll>e loins or side of (be body, are executed briti*
l«g or tJie Oppooite side, if the ley; of the same side be gently lietd. In thi<
oaM the vicarious movemeuu are ineShutuI, the le^ not being, as la thtraff
of the ttojf, creased over so as to bear on the ^pot stimulated, and f*a iM ^
considered as betokening inUlligcnce. Again, the " mechanical " naiart*^
reflex aolioos is well illustnted by the behavior of a decApitaleil toAt'
When the body of the aoiinal in this condition is brought into otntsri u
several places at once with an arm or a Htick, complex roAcx movrmeni**"
excEt«il, the obvious purp^MO a* well as elTucI of which ia to tirini- tbr bjl.'
mund the obje<:l. A deoapttatail auake will, however, with equal ami ^'*'
rMdinusi twina iisclf round a rod-hol bar of iron, which is mtJe to 1n>^
ita skin In several i)1airc« at ihi- s-imo time.
^ 590. lu consiiUriog ihi- naturwuf ibff events in the spinal oofd «'><(''
Till: REKLEX ACTIONS OP TUB STINAL CORIK
ri7
krvraiiae the belinvwr of l)iv fmg in the iofilaDce jiiat mentioned ko must
btar ip roinil iliat (he mov«ntcnt* in qucelion ire " cuoitliniilcd ; " ihal b li>
HT, no4 only stv many distinct inn^clee hrou^fhl into ulny but certain re)ii-
tic'Dfl mm niainUined belTreeii the amounl, tluralion, and exact time of occur-
Eice of the conintdion nf each luiiiicle and ihoee of the conlracltutia of il«
low mwcln tbarin^ iu Ihe movciucuU In the ahdenc^ of sueh (-(Hirdiun-
lioa the niovemeol would become irregular and indlei-lnul. We »ball have
KnwkiD later on iu dealing with voluntary movemeuiH in puint out that the
fl>t>rr) illation and hence tbe due acn)iii)iluhment of a voluutary movemeot
a (IrtM'iidcut on cenaiii aRerent im|>uli4es pufsinj: np frvin tbe contraettug
BU'Ctrs t" iho w-iilnil norxiiud iiytitt'm. and t("idin^ the dischargL- of ihe
tfrtf nt imiMibv* which rail furih the c<>iilruirliiinii. Whi-n iht-ab ntlrrvnt ini-
nilan afltct cMweiouaDeM we anenk of Iheni a« comtituting a " mu>i-ul«r
tDM ; " it if. a> we liball we. by tnc " muteular trntu " that wc become aware
if and can ajiprcciate llie coiiilitinn nf nur TuuKcln. But we have reaxon tn
fciak that the afl«-ri-Hl impulw* which rnniitiliite thu l>a»iR i>( the miiiTtiliir
Itnfe. whatever Ik their ciacl nature, iu cirder to play their part iu hnnginf;
•bout Ibo coord i nation of a voluntary movement need not puio rijfht iip to
tbe brain and develop a diclinct muscular "KCnNe," but may produce thi'ir
tffttt by working on Ihe nervoua ineubanixnui of the npioal cord with which
Ibe tnotur fibtt* carniing out the noTCinenl are connectol. In other word^,
|kbe coordination of n volimtary movement tnke« place in tbe part of Ihe
tailiRl cord which carrier out the movement, and not in the brain. thun};h
the tatter may be cooBci'iua of ibe whole movtinenl inetuding its coord innticm.
But if the <niDBl coid po««f>w mechnnifms for carrying out co'Tdinaied
punrmevts, which in the cuk- of voluntary movement* are discharged by
perrons impulw« desecndiug from tbe brain, we may infer that in reflet
(iclioaa the fame m^chanirm^ ate bro(i]>ht into action tliuti)-h they arc dis-
tbarcrd by aflertiit impnltiit coming ulimg afTvreut iK-rven instead of bv
ta>pulM» ileiteeiidiug from llw brain. The niovementa of rellex origin, in afl
tbrir tca!ure« fxcept thfir eKcitiiiii; caure, n)>)H'ar identical with voluntanr
BMivtmrnla ; ihc twu can only he di^tingiii.ihn) from <>ncb other by a knowl-
tdge of the exciting i-aii*e. Anil it weeniH uuniuttonable to nnppoiH: that tha
•pinal rord fhnuld piiweMi twu wet* of mechaninni.i in all r(9|)i-i.'()i idi-ntical,
we ibat the une if diiirl)iirge<l liy volitii>iial impiilw-v Iroin the bntiii aii4l the
plhcr by nllprrnt inipulfiii fmm nllirrent uerveit.
iWr are lid therefore to the concluvion that in a reflex action two kindo nf
ffcrcnt impuln-a are concerned ; the ortlinary aflercnt impidiH^« which di^-
ebarge iIm- nervout mechanism within the conl bikI m provoke the move-
Vtcnt, and ibr aRerent impulHv which conntvt that nervou* mrclianitm with
Ibe miucles about to be calletl into play, nnd which take part in the coordi-
nation of tbe noTODenl provoked. Tbe natura of thcw latter afferent
[mpuleee is at prrwnt obacure; nu know iiit yet little more than the fac* of
illieir existence ; but if we admit, aa we ecein compellnl to do, that the char-
jBcUf of S ntex action is determined by them fla well aa by the afTercul
Aupabea whtcb actually discharge the mechaniam, it seema ponible thai a
Jbller knowledge of lhef« coiirilinating alTerrat impulses may aflbrd an
aitetjuale explanation of the fact that when, an in the case of tbe froff in
ii|U**lion, the uxual wt of muscles cannot be employed by tbe acr^'ous
Ettvdianlsin, recoune b bad to another set,
I We liavc avoided tbe introduction of tJie wonl "consciousness" as un-
[aectfsairilv complicuiiug tbe (|ueelioD ; and it would be out of place to ditmisa
[|iey«li«logK«) probleoia bere. We may remark, however, that since we have
DM objective prooft of consciousDeee outaiile ounelvea, and only infer by
■nakgy Ibat mch and tucli an act i* an outcomeuf cuntciouHtetaon account
THE 8PIXAL OOBD.
of ict likcii«w Ui Ktrlo nli:r)i nrv the ouknnw of our mro OOdtdlMtBNi «>
cODcluHe ihtil iht! braiulniK rmg |m>mmmm no fteUrc cunxdotUMM Iflwau
own. lK<cnii«! alMMttiM' of .■<}ic>iiuiim»ii* iiK>v('ciMmli> Mxnu u> Iw InrooQciUliW
witli Uw cxiKiciicr »f n» iictivv oupMcioiwiicw wImmd v«r}' n«M>»rt> 'umwnm<4
chaogM. (^niwuHii>tivi«. iiK vFc r<.-0(>giiiM it, tetmt to Ixi nMcwsrily bperatiB;
an. nr to lie iDcJiwolulily sMiicinlcd with tbe [ircMtm) of, »a latnmati'!
re|)enlfd iutenuil Hiiniultis ; iu»l we cnnnot raooeivo of thnt stimulus Ikiling
Id excite raochtniinis of ronvciumt which, as in (he ca«e of the btaiatai
fntt, are confoMcdly prmont. Wc may. hoirevor. (Ii»linguub betwen ■
nclive cootinuous coiMt-ioasium. iwh iw we iiBitnlly iiCKlcrataiMl hj tbe Itfa,
mill a jiast'mg or monieiitarr condition, wliicli iic may apeak of as emntririn'
ilGtt, but which is wholly dteooiilinuous fmni hh anlecedent or fnim m - -
qiwnt eioiilar momentary cooilitioD ; an^I iudeed we may eiipixne :
complete couMMXHUeas of oureelve», ai)*! tbe similarly comjibitc <-< i
■Mm wbtch wfl iofer to exist io roaoy aoimats, haa been (iruilually .
nut of Auch a rudimemary conscionnieBs. W« may, on Vi'm view, rappf
tbat every nervous actiop of a oerlain inteutiiiy or cbamcier i» artximpaoM
bv M1IUI- umiiimc of oouai-ioii^neHa. whicb vre mny, Id h way. coiuimrv loliu
light emitted when a t:oiiibuBlioi), previoutly fciviuK Hoc to invbaiblv hml, win*
fiercer. VVc may thus infer that when the brniitlcMt fn*^ i« stirr«l bv luw
etimulu* to a rcllex act, the apiual conl is lit up by a momeuury Aatbtf
eoneciouaiiow coraiufc out of darkneas and dving airny into darkncM tmnn
and w« muv porhapt further infer that Htich n piMiing consdonaoeA vAt
better develi>|><!>l the larger the portion nf t)i« oord involriKl in the radex
aet and tbe more complex lh« moTwnHit. But sueb a momentary flaA.
even if we lulmii tbi oxiMl^ice, iiiionwthiBgwrydifltfreDt from cunMiaosiw*
aa ordinarily und«r«lood, i« far nmovod fh»iii iiit«Uigenee, and canaot W
appealed to lu explaining tho"ohoiot"tpok«o of abore.
^ Ml. Lttslly, the chamcten of E Kflax movement are, aa we Deed liinil*
■ay, dependent on the intrinsic cotMlitJon of the cord. I'he action of UrTchMi
jutt alludc<l to is an ineUtnce of nil apfnrcnt augmentation nf raflox WlAa
"hml explained by suopoeing thai ih*.' rrwirtamxH in tbe cord MW \imill*i
There are probabljr, Imwever, casM in which the explosive tourgj of i^
iicrvoua mibstance la pautiv«ly increased above the normal. OonvenH'lv. \"
variouit influences of a defUMsing cbaraeter, aa by rarioita wiMth>
other iMiisana, reflex action may be lessened or preveated ; and thi> j.^^
may arise eitlier ftom au iucrewe of reaisuuice or (Who a (liminuLion in ite
artual di«charge of energy. So also xiurious diteaaei may «o nfli-ct llir simmI
mrtl an to prtxiuce on the one hand inoreaaed reflex «xdtal>ilitv-. m thai*
merv touch may prodooe a violent movement, and on the other bund iliauii'
iiihcil rcllex exeitabilily, so that it becaucedilTicullorimpOflMble to call fuRk
a n-Ilex aeiioii.
S SOS. When we oome to study tbe reflex actions of man we shauU M
fintl iM^rhajici be inclluetl to infer that, oinee in him the spinal con! i* *'
largc'v iiivil OH the tn.'ttrument of the bniin. tln' induprndeni rellex nciiro-
of the eord, at IcM audi aa aflect *kelcul mui>el.j(. arc in him of iimr^ ^'
im|ioriKnc<! than tJnoy apponr lo be in aniniaU. and cxirarienoe seems I" *if
pon (bix view. But It must be reoMnbMfd that in his case, aa we ln"
alrcndv i>uiled ($ AM), we luck the guIduHx- of experimental roalts: **
an iiblij^cd l» tnial to the <iiitan^lt)d phenomena of disnue or to a «til<tr*(
tbe liehaviiir of tlic oord while it is Mill a {mrt of an intact nenrouasyX*";
ami each of ihrte nti-tbod* pnaonts difBculties of it« own. Tbe nwveniM'*'
wliieh in tbe iniact human body we ran reongnize aa indubitable nll«i
RClions, are lu a rule itimple and unimportant. They are, in by Ur 1^
greater numlwr of iustanoe^ occasioned by stimulation of tbe akin or of tk
Tlir KEKLEX ACTIONS OX THK SI'INAL CORD.
nv
BUraus tni'iiiltniDi?. for llic muM. |iart inviilve a iVn' muitcln only, iiotl ntrely
hdioMc Kiiv vcrv coiniilirx tiHinlinatiiiii. Tliv llt-xiiiu. fullon'od u,V exundiiu,
f Um \og, nhtch i» i-nlUil forth \ty tickling (lit- note >if tlie fiiotr or tlic wink-
k^ of tbe cyv when ll>v cuhk^i <>r vniijuiictivii is I»iicli4<il, ninj perba|M be
Inrdcd as tlic Ivjn- i>f ihvm' ntitvemcalK. A vcrv ■■diiiiikiii fona of r<4ex
Itmt is lluU iu wliivli u iniitck- or ({roup of mu^oU-* i» tlir^wD into cod-
tkctton l>T Btiiniiliitiim of thv uvrrlyini.' or m-igliboriiig »kiti, lu when tlic
bdominal muAclca L-untmct ui>uti >trukin^ the 8kin of the ubduniLii or tliv
•tide is ntnctcd upon MroKing tlw io«i<i<? uf tho thigh. A rvHi-x niitve-
kot mar occur a» thr result of utimuUlion of nn organ nf Kpccitil wuso.
frti of ti>(> ceiilml nvnoue »ri<liMn other thttn l\w gpiDiJ vurd serving n» lliu
Intre. A fouDd or n floeli of light rrndily prwlucts a start, » bright light
talus the 03'e niiik and may oiuse n ptrnMO to niwze (the groalcr coordiiia-
pa manifeBt in thi« lu-t bi'ing duo to the fuct that the complex r«»pinilurT
pcchnniifn b hrouuht into plitv, $ ^'l^). and reflex movoments may rcfiiilt
(DO s laste or smeri. A special form of reflex nctioo, or at Icul an actioo
iwmblitig a reflex action, is called forth by sharply striking certain leii-
^His; for iualance, linking the tendon below the pat«lU givcti rise lo a
^(Idcu extension of the leg. known a» the " knee-jerk ;" hut it will be beet
B dim^uM these " teudmi reflexee," or " muscle reflexet," ua ihey are ealled,
birr oti in uitolher conneetion.
rOn tbo nhoti- tbe reflex mnveiuenU carried out by the inluct uervuus
^stmil of mau UK, ne repeal, Di'aut^ aud i-um pa ru lively simple: but nc
m Dot iustified iu inferriug from thia that the htimau ipinal conl. left to
ItMlf. b inca]»ble uf ditlug more; that owiui; lo thr predomiimnt activity
H thv brain it has loat the ]>uwcnt [lutseNMnl by the «pinnl cord in the lower
niiuaU. For it may be that the cord, ivhcn juim-d t« ihe bmin, u through
nrioiu iuflucni'CM pnti'^eding from the InUer in n dit)i.-reiit cunilitiuu from
luit in which it i» when aeixinttod from the brain; indi?ed, we have reason
D think that this in ■» : and we may her<' remark that in the lower animals,
la in tunu, tlio developnirnt of refli.-x m')vemGiita is diliicult snH uncerLaiu
p ihc presence of the hroiiu.
I, When M'e turn lo Ifae leaching of diieaee, however, we again lind that
jlAex uioveoients carried out by the cord or by paru of the curd are, on the
thoJie, seauty and simple.
r In sonie tUtgtm of oortain diseases of the spinal cord extensive retles
koYUiMat* are wilDCMed ; but these are not purposeful, ouiidiuated move-
MDlM. such as have been described ahore as occurring iu ttofc* and uiniu-
balaaiUr ex|)erimental inlerfereuiy>. but rather were ezagf^nilious iif ilic
(unpler reflex morcunenls iriinmeecl wIk-o the nerTOUS ayatcni ii intact. In
Hirr of paraplepa (such being ih« term generally uited when diiNHuc or
Injury has cut oHT the cord, generally tite lower juaii of the cord, ttoot tlie
brain, so that the will cauuot briug ubout niuv«au)ots in, ud the mind
Serires no senmiiJou from, the parti« belon th« hvion, the legs for iiuUiice),
it swnetimo happens that contact with ibe iM^dclolhcs or other external
objects sets up from time lo lime rhylhniieully re{M;at«) inorcmeiila, the logs
lM>iu>; aliernatelT driiwu up and llirust out it^iuii. And an oxaf^raiion of
(be " kim'jerk' i>r ol her " tendon n;(lexv«"is a very cummon symptom in
Wnain «pinul di»cu>i-9. Il tit nirvly if ever ihul reflex moroments of a reully
puiuplicBie'l choraoLcr arv oliwrvii]. Mureovcr clinical experienoe shows
tliat in mau, whm a portion of the I'onl is isolami, reflex actions carried
Mt by iDeMW of tfant ]H>rtioti. so far from l>oiug exaxgenilcl. are much m'>re
^MUluonly exceodiitg feehlc or absent altogether, in the cnseii iu which the
Bfayidolo(Hc*l ocintiuuity of the lower with the upper p<irt of tbe cord has
DMO brakeD by disease, by some gmnth invading ihc nervous ftruoturea or
732
TUE sriNAL CORD.
suniulxtion. that the lime interveiiutg Iwtwmn tbe aelioii of the aciJ m'
tue and the iritbdniwal of the foot U very much prolooged- That is lo nj?
ihe MimulatioD of the opiio lobe* ha? cauMs impuHee Lo deaceod to tW cori,
which have there m interfered with the nerroiis prooesws eni>Ji;fed in canr
iug out reflex acltoas as jcrestlr Ut retard the generatioa of cfTervnl in)pul«t.
Of, ill other wortb, has inhibited the reflex action of tlie cord. And sioulu
rtaulta may be obtained in uianimaU by aiiiuulatinj; certaia paru of lie
corpora '(uadriKeiuina. which bodies are homolui;ouk ti> th«! uptir iDba a(
tr*)g». From this it has br«u inferred tliai there b preM-nl in this put tf
the brain a npudal mecbauisiii fur inhibitin); the ri-llcx aetluiu of tlw ipla^
cord, lh<- impulM* dcMcndint; from this mei-hanisni to tltc varicitw rcDini of
ri'Hcx tictioa bdng of a specific inbihltorv iiaturv. Itiit, as we harr alraattr
tvvu, impnlm of an orditiarr kind, piiwiiij; along onlimiry Ewit»iry
nay inhibit reflex action. \Vc have <|iiol('d iiisuuicG« where a alight
ulua, M in the pendulous ni<ivciiii.>nta of the .-inake, and whare a sir
ttimuluK, a;> in the oim- of the inii'turiliou of ilie ili>g, luav produce an Is-
hihitorr rvsiilt; we may add tlial in the frog adeijuatciy stroi))* stiauG
app)ie<i to any alle rent nerve will inhibit, i. r., will retard or eveiinholK
prevent reflex action. If the toe« of one kg are dipped into dilute aiil|>hurr
acid at a lime vfhen the sciatic of the oIIkt teg is being powerfully sliuiu
lated with an interrnptod current, the period of incubation of the n^'llex aa
will be found to be much prolonged, and in soiue caws the reflex withilrawii
of the foot will not lake place at all. And this holds good, not wily tu ihr
comuleie absence of the optic lobes and bulb, but also when only a (xirUto
of tlie spinal cord, sutficieut lo carr}' out the reflex action in the uaiisl am,
U left. There can be no i^uestion here of any specific inliiliiii-rr ccuim.
atich as have been siipiiriM^ to exist in the optic lob<p. ButifUbclw
that iiihibitiiio of reflex ui-Iidu may he brought about by iaipubca whidi
are not In tbeiuMlvu* of a >]>eeitii' inhibitory nature, we may hesicate w
aooejrt the view that a ti|irciat inhibitory mechanism in the tenM of not
giving ri»e to ttothing but inhibil')r\- impulses a prewnt in tlie optic l->to
of n-oga. and after removal of tb<- brain that the exalutiou of reflex *c-
lioDM which is manifest is due lo tlie withdrawal of nch a spccitic inhibiwt
inccbanism.
Tha prtMiicc of the brain does obviously pnotuQi' an eilect which may I'
bnadly spoken of as inhibitory, and a specilic action of the brain, lo is
tlhn of the will, may stop or inhibit a s|>ecific retlcx action ; bat we noit
not iu thrac malterre be lod loo much away by the analogy of '
and limited cardiac inhibitory mti^nnism. There we hare ap|
deal with tibrM, whoee exclusive duly it is to convey inhibitory li ,
fixioi the bulb to the car<)tac muscle, and inhibition of the heart, at' bsM
through nervous iofluencci, it«xc)usivety carried out by them. Bui ainuij,
in Btudving the nervous tneehaBism of rMpiration, we have seen reasw lo
think that atrereiit impulwa pasring along the same nerres and prabik>?
along the same libreit uiay, according to circurostancee, now {nliihit.n(i*
augment the respiratory centre, and nave thuA been le<l lo speak «f ■>'
hibiutry impulsw. that is, impulses produoing an inhibitory vd^'l. Sf*'<
from specific inhibitory flbrea. In the complex working of lbs oMiUil
nervous system we may still more expect to come across similnr instaaoa^
the same channels serving as the path, either of iuhihitiou or of aatfM*'*'
tion. In all probability actions or proceau^. which wc may apoM <f "
inhibitory, do play, as indeed we ahall sec, an important pari in the f^i^
work of the central nervous system ; in all i>rohahility many uf tbr |<^'
nomena of nervous life arc the outoMno of a oontesl between whsl " "^1
call inhibitory and exciting or aiignienting forces: but in all pmbsbl^i,
bat we »w
riha^iy
ppan^^H
>ryuiM»l
TIIR RKFI.KX ACTIONS or THK SFI3IAI. L'DBl).
rsi
nj to ouea of |innipl«Riu railed ttv nvciilcdlul orrabio^or nreniDoeof tlic
corA. The Kiircity ol' wt-l 1-rtinrkei) reflex aciioiiB iii«Dliij|ied itbovc ns cbur-
atMriltic "f lurh mKi'. mtiy {lerltiipa be >)ue lit (be Ihct tbat the«« diaonleni
tfantiilUm |>n-v<til ihr iialieiit living; Umg eimugh for ibe wparal«d cord to
nivivrr ibi- liHiili')ii» u'lui-h piMjH-rlv bebnijj; Id il.
; 594. /jij'pi'.id.in '-/ ivyff^ arlioii. Tlie relU-x nctionn of the npinul cord,
likf i^Imt iit'i'v-iu* notimu, iiiuy be ditnlly or ]iurtiiilly iiihiliiUM), ibKt ii to
Mty, niity lie iirr<«i4.''l or hirkkml in iheir (kvclopmciit by iinpiiltKui reaching
th» OHitn- wtiili' It i> ntri-iKly iii vdion. Tliii*. it ibc body of a dvcnptMtM
I ite dlbiwdl i» bung ilonti, »lun- rhytbniic pfiidiilout^ mOTemoito. which
tr Id bi- Tvttvx in nnliirp, eo'in inakr ibeir ii|>pctimncc. nnd these miir
* while iirmt€<) by tlight siinuilntiou, tii< by ^nlly stroking the tail.
We h»vc nlready eceii thnt th« action of »iich norvous centres aa the resjit-
ratory iind VHsomotor c«DIrf«. which frcquetitly. ut all events, is of a retlex
nature, nwy be either inhibited or nugm^Dted by alferent impulses. The
uticUirition centre in the ntaninial, which is also InrKely a reHex oeotre, may
he easily inhibited by impiilaai [tassini^ downward to the lumbar cord from
the bruin, or upward along the sciatic nerve*. In the case of dogs, wbii*r
ipinal cord has be^n divide«l in the tboracii? regioo, micturition set up ax a
redcx act by Hmpl« preduire on the alidunieu or by «|MugtD^ the aims i# at
oooe atoiiiKil by ^baqdy pinching the skin of the kg. And it ia a iimtt«r
of Gonmou experience thai tn nmn miituritioii may lie suddenly checkod by
an MDOtton or other eer«bnd rvenl. The erection centre in the lumbu'r
mrd, also iu large ineiwure a rt-tlex ceulre, la MBiilarly ausoeptible of Iveinf;
iahibiied by impulns rvschinfc it fri>in various tourcefl. And, indeed,
many niniilar initanceti of the iiihilHlion nf reflex movtnicnt* might readily
be (lunteil.
tS;vcral nppan-ni instaiicos of tJir inhibition »f redes nets arc not really
•uch : iu these cn*cs all the nervous prijce««« of the act may take i>lace in
their entirely and yet fail to produce their eH'ecl un account of a failtim in
Ibe muKular part of the act. Thus, when we onrselvee by an effuit of iho
will (top the reflex moTemeiit^ which otherwise would be produocil by
ticklioR llie soles of the feel, we acliieve this to a large extent by throKing
viilunlanly into ndion c«rtain inuscke, the coutnictions of which antuj^onti*
the aclioD of the muscles eDf^aj^ in carrying out tiii? reflex inovemeDU^
Hut it may be doubted, fwn in ibtve niseis, whether inhibiiiou ■* alwai,-* ur
wbdlU 111 lie explained in ihix whv ; am) certainly in very many insiancrs
ot reflex inhibition no hucIi muscular autax:i>ni*ni t* prMi-ut, and the reflex
art '» cliccked at ii» nervoua centre.
When the bniin of a frog is removed, and the efftctn of nbock liave pttsfvd
away, reflex actioiu arc itcvrlnpi'd miii-h more readily and tn a much ^'reater
ilcgrce than in lliv eniin- anininl, ami in niamuiais also retleJC rxd tain lily
bas Iwco obitrved to In- increa^cil by rvinoval of the o-n-bral hemitpheree.
This euggwu the idea ibnt in the inlii<-l nervous system the braio is tiabilii-
ally exerting wme influcnci- un the spinal c^tril, lending to prcviiil the nor-
mal devi'lonment of the rijiiniil reflex actioitf. .\nd no kani by experiment
llut simulation of certain jinrts of the brain ha* » rrmnrknble cfled on
rettex action. If a fro^, front which the cerebral hemispliorrs have been
reniuve<l (the optic lobee, balb, and spinal cord beinc letl intact i, tie £u»-
pendetl by the jaw. and the toes of the pendent legs Be from time to tinie
dipped into very dilute Buljihuric acid, a crrtain average time will be found
lo elapse between the dipping of the toe and the resulting withdrawal rif tlie
■ Jbol- If, however, the optic lobes or o|>tic thalami be stimulated, as hv
putting a cryalal of sudium chloride on tbem, it will be found ou repeating
ihe experiment, while these structam are still under il<e influence of tl>(*
U
a
734
Tllb 81'I.VAb OOBU.
of Ibe body as tb«MirraeoMiniulal>on of which euuls lh« set. the liiMUk
up ia lc» tban when the movenKnt ts on tbe utlier side of the Kxly. mild
beio^ made for tbe Inif^th of central nervous mailer involved in iheivot
tlial u to wv. the ceiilral ujieratiiMU of a reflex act are proMji^ted mom i
idly aloii^ tlie cord tban ncn^s ibe cord. Tbe rajiidity of tbe act vari«.j
ootine, with Ibe coDditi<in tt( the xpiual cord, the act being grvaily pruloaj
wheii the ennl becumiw exhausled ; and a ^irnilur delay baa been •rl«efT
cwMS of dbe«»e. The time thus occupied by purely reflex acllon» muM D'4
Im ct>iifoumlei) with the iuicrval required when tli« changes taking place in
ihe ciriitral iktvouh nyxlcm arc »f a more cuniiilicatt^l nature, and uiorevr
lewrtiMinrlly involve inc«liilo]>eratioiin; of the lalti^r wcxhall !^>c»k
TbK AlTOMATIC ACrlOXS OK THK HpIKAt CoRIK
S M6. We Bpeak of an action of an organ or of a living bo<ly as
tpoulaneoua or automatic when it apjveare to be not iru mediately due I
enanf^n in the cireuniatances in ithich tbe ornn or body t^ placed, hut i"
be thiTCNult uf chanf^amtDfi; in ibeorj^n or body itseifand determiii'' *<-
cauK» oilier than the influences of the circomaiancee of tbe niomeiii. '
Hulomaiic actloua are of a i-uutinued character ; ulbers, like tlic beat iii \i>'
hi-itrt. »rc repealed in re^lar rhvthm : hut the mont Mrikiuji auiuntatic or
ti<>iii> of the tivinz bo<lr, lhuc« which we altrihiiU- tu ihu workinn of the iriC
and whirh we mil vi)luularv or vnlitional, are charucti-rixcl by itieir app'
rtot invgulurity and variaob-iiou'. Such vaniiblu automatic aclMioi WS
ibo rauat «lrikine feutiin-s of an iiiiaol iH'rvuui> ]i\i>li'(u, but arv roMpinwiuIl
nbKnl fioni II Kpitial cnnl when ihc hraiii has been removed. j
AhraiiiU-n> I'ro^ plncnl in a enndition of «Ku^etoci]iiilibriun) innhichl
(timuliiit is brought lo hear on it, prutwtnl, for lUnlaiuii, from ■iidikii t^p
chaog«# in tcmjHTiitiire, from a lua nipitl cva|>i>mti<iM by iIh^ -kin ajxt
lik«i remaiiM perfectly roolionlno uuiil it dt<-s. Such ap|Mirenily BiHiQiaiiMBt
nmremenle a» are occaniuiially wiliinMit arv m tiltw and itel(h>ni. ibal mt rti
liardly do othcruioo than attribtito Ihcm to eome rtimulu*. inicrnal <>r ru*
nal, which has cet-aped observation. In the mammiit idog) uA^r iliviiiuut
tJie npinal cord in tbe doraal regioD irgulur and appiin'ntly ^[NiDUnn
moveme Ills may l>e obwrvcd id the parta gov«riic<l by the lumbar curtl. VH'
ibe animal has thoroufihly recovered iVoni the oixralioti the hitiil liiutH n.K^
remain (]uiet for any lon^; period ; ibey tnov« rcslleasly in various wnyi ; ta
when Ibe animal » suspended hy llie upper ("art of the body, the |h-wI«
bind limfaa are cunliuually being drawn up and let donu aj^ain n itb a aito*-
onoua rhythmic reKUiarity. sugf^ive of autuiuulic rhythmic diicbnii.t»fn<ii
tbe c«)iral mechanisms of the conl. In the oewly-bom maiumul ti»>, »f
removal of the hraiii, movement apparently BpoDtaii«ouB in nature an* Tn
ijuenily observed. But all tbeats movements, even when moat hiiftily ilerrt'l
oped, are very ditferent from Ihe tuoTementv, irre};ular and variable ia i^'
Mcurrence though orderly and jniirpoM-ftil in tbeiri-baracter.Hhich uemvp
niie as dblinctly voluntary. Evcii admiltiuj>; ihat some of tbo moiTOHi"*
of tl» brainleaa mamnaal may reacaible voluntary nvuvtrineutii in w hi u
tbey are due tu rbangca leking \t\wo in the «|<inai i\>nl it.ielf indci>eivi<n> 'J
the immHialeinfluenceof any »limiilu>, we are not thereby juHlirictl iii<)>*^'
ins of the Kpinal cord ait developing a will in tbe mubo lluit wo uitriliiC,
wiTl to the brain.
i 567, in tiic COM of tJie beat of tJio beari. iIm automatic rbyUiaaf ^^
charge of energy apiM-ara to be exclusively the outcome of the nulK*^'
uulriiive cliHugcN taking place tn Ibe cardiac FUbattiDce. llie boat ■*,' '
THE BKPLKX ACTIONS OF TIIK STINAI. COUD.
72Z
I
I
I
I
»
mimt, wv outHit ntltor to Mck for t)i« cspJiLnaiion of how vofM iRi|Hil«M
inbibk Uie IMM of the lienrL Uj: refviviico tu t)ic inhibiiury phviiiomcDa of
Zbe ccDtrul ncxrou) ajtXcm, (han Id Ktivmpt to cxpluii ihc Inilcr by th«
JitU* wr kuoit nf ilii' fonuor. At prtw-nt, hciwcvvr, nv iiiii»t br ontctQl willi
%.ht &ct tlint rxpcnriM-nlit <>n niiimnls xlioir t.liiil thi? Itniiti, not onlv liv Eotue
au^itiu or othtr, iimy iuliibil piirticular vpitial mHes tnnvrnteiili, but aku
linbiiqallj cxc4viiH9 n rtviminini; iiifliii.-nc«< on tho rvAox nctiviu- nf tli«
i^liitlr rant, though w« nrr iiiinnk- to stale clearly bow tlii» inhibitioD U
ourivd out.
W« uy "experiment* mi MiimaJs." because though wo know, aa stated
nlhtw, by an appeal to our own conaciouaneM, that an action of the hraiu,
an iffrorl of the will, may stoo a parlii'ular r«tlox act, we have no evideuce
thai in tnaa aeparatioti of tne oonl from the brain leiid^ as in aniiuuls, to
lie^fateoetl reflex activity. In diaeases, or injuriea to the cord, retlex actiooit
M«, M vre have said, sometilBM eiagRCmted, but it is podsible, and indeed
pnliabte. that (he increaae ia dne to the morbid proceasee produutog a
crMtvr irritabiliiy of the cord ilaelf, and not to the withdrawn of any iii-
ntbilory influeooes. In many caaea, in perhaps the greater number, uo
exsggeraiiou but u diminuiiuu or even abaence of reflex activity is obaerved;
w muob BO ihai could we tru»t explicitly to cliaical experienoe, we ahouUl
ht locliaed to conclude thai tlie sountiueas of apinal relex action iu man
van ilu<- not to any preoccupation of tbe cord by influeocea praoevltiiL' from
a iloiniiianl brain, but tu an iuherenl paucity of ajMnAl reflex imvliuui»iua.
ilut «e have already Mid all we have at present to nay oat tim point.
t 595. 7%e limi rfriuirittl j'or rejtri aetiatu. When one eyelid iii stimulatud
with a sharp clectmitl iiiock. both L>yelidit blink. Hi;iiui?, if the length of
time iiitervvning b^lneen the siiniulatiim of the ri^ht eyelid ami ihe ntove-
nient of ih*- led eyelid Iw mi'murw), ihi> will give llie total lime rwjLiired
for tl>e various proccwm which make tip a reflex action. It has been found
to Iw from 0.0t'>li2 to 0.0578 aecond. Dediicliog from tbmc figuroe the (in>e
required for ilie passage of allcrcnt and cfTiTent impulses along the lifth auid
facial Derves tu and from the bulb, and for the latont period of the cun-
Iractioo of tbe orbicularis muscle, there would remain O.Oood to U.0471
■mnnd for tbe time oonsumod in the cenlnil opemUoiu of the redes act.
The odculations, however, necessary for this reduction, it ne«d not he said,
an opco lo aounMS of error ; moreover, the r-Hcx act in question is canied
nut by the bulb and not by the spinal cord proper. Blinkinf; thiu produced
is a reflex act of tl>e verr simplest kind ; but, as we have seen iu the pre*
eclin^ pap*, reflex act* differ very widely in nnlun? and character ; and we
aciHirilioglv find, as indeed we have incidenlutiy mentiuneil. that the time
inken itp l>y a twUex nnjveioent varies very largely. This, indeed, b »<*ii in
hlinkin^' itaelf Wlieu the blinkini; is caused not by au ek-ctric ahock
appli4-d lo the eyelid, but by n Mash of lijjhl falling on the r«uiia. in which
case complex vibual proouMes are involved, tbe tinte is didiinotly prolonged :
moreover the reMilta ia difilBrent experimenla ia whiob light aervea a« tbe
•timulua arc not nearly >o uniibrm as when tbe blinking ia oauNd by uinm-
laliun 'if ibe evelid.
In genenil ii mar l>e said that the time required for any reflex aot variea
vtrj Gouderably with tlic strenglb of the Mimulua employed, being le* for
the rtroi)g«r stimuli ; tliiii we should ex|Nii:t, seeing that the elldreni impubea
nf tbe rettex act are not nimply aflcreol impulMH truiiimitt«d through tbe
ceatral organ, but m*ult from internal chaugua in the central Oi]gan Harl«d
by tbe afllsrent im|MilM or impultMH; and lhe«e int«rniil changea will aatu-
mllv be more intrikie ami in>ire rapidly aflccteil whi-n the aflbrent impulsaa
arr strong. It i* statul lluil nbeii ilie movement induced i.t on theaame sidB
7SS
rns sri.vAL t-oRU.
to » qumiou nhii-h Iins Iweu tiiucb debnteil. anil nhicb ha* nu
ind iiD|H>rtiiiit bearJD^a, nani«ly. Ui« (jiietilHui Kh«lti«r llio *mr
cUeA au automatic fuDCttoii in iiiumiMiiiiiig » lonu ot' iIh' nki-lflml muKinv
TheuueMion ia noi one nhJch, ttki! ibii rue i>f BrK-riiil tunc. <'«ii be *tKM
off bana by a*iR)|ile cx)ienniufli. Moat obaerTem nm'c that ibr- (n,-ii'>Q <'i'<
motor nerre dot* i)»l )inH]iii.i> any clearly rvcogprxiiblt' imiu(sliai«> Iriir' -'-
iDg ttf a muscle nipplinl by the m-n-^.in ihniiHRKr way ibiil i>ti:1toii •■( ■
COnitriclAr nerve iiti<li>iibl<slly giriin riMi l» n rvlitxatittii of tbo miMcutitr i i<i"
Id tlw art<:ri4vi govcnu'tl by it ; anil it lia» Ixi-ii infermi from tbi» tlutt -l- - i'
lono don m>t I'xtwl. But thrrr iirv M-vi.Tal AicIm la bo takrn iniu cntKiden-
lioii hnftin- wi' (7IIU oome to a jiiK ilcciKion.
Tbc«k«lcial muicln bare l>n-ti ii(«rrib«4l a« living pIao«d"oR iIw*iTftrIi"
In iIm! living body. If a riiim-Io be out away Irom il« atUicbmcnie ai nA
rod, it »lu>rf<D» ; if it bo cut ucroee, it gapn. lu olhrr word*-, the mii»cl<- a
the living body p<«wwra a latent uadtacj to shorten, which ie continuk^T
baiog count era dfd by ile ditpmition and ntiacbmerKF. In Miidying »<»■
cular cootrHctioo vie hbw (S si) that the shorteoing •>!' n ountraction ir id-
lowed by a relaxation or return to tbe former length, both the coiitracii'o
and relaxation being tli« reault of moWuInr elMii)Re« in lite living; miiwulti
mibMance. We have now to extend oiir view and to ivco^ize that, apart
from ibeoccuirence of ordinary contraction*, molecular citan|[ea are by mnnt
of uulrilive procraaea continually voing on in lh« muscle iii such a way ital
lite muKle. ibougb continually on we atrrtdi, doea not pernianentlv leu^n.
but retains Ibe potior to Hhorten upon removal or Isaening uf ilie strMi^.
and coDvenely tlioi));h piwMeunj; tbia |)oner of shorteDiiig giermits jtwlf m
lengthen when tbe .ntrelch is increased. Id tbix way tlw mu»cle is able f
accommodate iinelf to variations in the amount of stretch to which it is fnai
lime to time subjected. When a tlexor muscle, for instance, contracts \U
autaguiiiMiecxIcuiHir niufrle is put on an increeMd stretch WMi is cortte]Miai-
ingly Iriigt iH'iinl ; when the* cimtrnctiun i)t' tbe flexor puma olF tlw nteDMr
rvlnrnii to ils pn-vioun leogtli ; and to in othor inylHiices. Thufi by riTtiiv 'i
cerluiii cban^rn within ilwlf a mutclr maiuluinn what luiir I"- called ll* auo
nl Itngth in the iKnly, alwuya reluming to that UHtuml Icn^h liolh >D^
Mug uiorienH nnd after being rtn-tcbed. In ihi* ihr muiH-lii d<n» mi wm
ihnn do lh« niher liiutKV of Ibo IhkIv which, wiibin liniiln. retain tfcatt
natuml form under the varied Mn-w and rtrain of life ; htit tbe pre|
eonamcuoiis in the miiide: and it» elfeols in ■kclclal mu«cJ« cor
clovelr to Ihow of arterinl tune, that we may venture to ■peak of it i
lal loite. Indeed, (be molecular changes at the bottom of both are pinb
the eamo,
These changes are an exprenion of the life of the muscle ; they dJMff**'
when the muscle dies and enters into rigor monia ; and, moreover, dur
life iliey vary in intensity so that tbe "tone" varies in amount acoonllngi
the nutritive cbaDgea goinj^ on. We have seen reamu to believe thst t*
BUlrilion of a mufcle, as of other tirauee, is govcrened in wine way hvii,
central nervous Bvstcui. We eaw, in treating of muscle and nerve iSft'JI
that tbe irritabiltly of a uiUM'te is markedly affected by ilte *«li(iB"(iB
nerve, i. "., by ocvcrani e jniin the <M;»tml nervous sysieni ; aixt again 1 5 *''^Jj
in sneaking of llie wt-callcil lni[ibic action of the nervous system, we rtff^'^
to cbaiigt-s in the nutrition of niuw)™ occasioned by diimws of tbe anvfail
M'stem. And vX|irricnoe, es]iecial]yctiuii-al experient.'c, showa ihni the vi''*''
live changes which dctcrmim- tone are \-cry oIom-Iv depeiident on a H"*
action of iTie cniinil wrvoui' nysii in. When we hamlle the limb of a bcal'"!'
man, we liml ihmt it oirt-r« a n-tlain amount of resistance to r"BaT<'H'^
ueuis. This n-sirtanw. w hieb is ijuite independent of. that is to lay, «»"
I
I
Tim Al'TOMATIO iCTIOXS Of THE SPISAI. rORn. 72S
nHidlRnl, m «» havt trta, hy D*rrvou« ininulm>t> n-nchiiig iW mrdiac sub-
MlBUni along certftin ti<-rvvs: btit the net it nl (■xitU-ticv of ilii- Iwit u n holly
inilvprmlctil of thifsc pxlrnnooiia inHurnt'cs ; the rhyllimic ilUchiirge cimtintiM
whvii ihfv arc piiiirelv sltwal. The aulonintie rhytlimir iliiichnrec of n*f»'
raiiiry ini|i«l»c« from ihp reBpiratorj" ccntrv \» nho AvpenAvni on inc iiilrintiu-
moWulHr i-lian;^ of the <viitre, iHmc bdo^'. ne wc have nxn. Inrgi-ly <!i>t<'f
otttieH by the chaniot«r of the bloixl stivHinin;; througb il ; but tti ibis cum
rxtrinsic nerrous impulses, rracbin^ ibe cMilre nluti); th« v&{;ii$ and iflb«r
iwrve*, |>iay a ruiiob more impiirtaiil jiart tban do Bimilar inipulF^^ in the caw
oTlbc b*arL Tliey )ict *o molinually ou the centre and enter bo Inr^^ly into
its workiui:. ibalwe «re cotnpelle^l lu reijard llie aclivity of the centre na fed,
if we may use lli« wnrtl, aot ouly by ibe iiitriiinic inoleculnr nutritive pro-
ctflMa of the centre ttAolf. but al»o by the extrinsic uerrrKtH iuHueucta whieb
liw inirt (he ivmre fmrn niihimi. Tlif aiiiomaliain of the »piimt card as u
whole rwifnibltsi, in this rft»peiH, ihnt of thi- reK()iratory centre rather than
tJiai of ib« Ill-art. It ban for ili> bn^ix duubiinu ibt.- iiiinunic niolcctilnr
(kMWW of the gniy mailer, on whi><e riMnarknble cniriilutiun v/c dwelt in a
fiKVlmu aectioD ; lh« metaholic evenl.* of lbi« oulHlniioP are so <>rden-il ili Xit
gire rtw to dtfichnr^tv of rni-rgv ; but (be diitchar^ ap^Mwrs to be alsu tnli-
nutcljr dependent on the inflow into the gray nuitier of afferent impuUes and
iDfluencfs. The iiorioiil discJiar^ ofdTerent inipuliH» ttota the cord ud<
rioubtadly lakc:> plai-c under the influeiices of ihnc incoming impubn; and
It may be donbteit whether the gray matter of the cord would be able, in the
■henicvofnll afferent impulK«. to gttDcrnle any «uttain«d eeriw ofdiscbarKn
•>ul of it« merely nulrilire iiitriocie cbangea. The niitomntic activity of ttie
conl u fe^l not only hy intrinsic nutritive events, but al»o by extrinsic tntlitenca.
In thi* feature we may, however, lind perhaps tbe resi-in why the auto-
matic cMvity of the spinal cord is so limited as compared with thai of the
brato. Ill «)>ite'if eerlain »trikiii;; but superficial chanelen of wbiuh we
thall jipeak later ou, the ^niy matter of the brain pn^K^nis no hii«tob>gical
feature* to diiri-ivtii fmm thuw of the gray matter of the cord, nii to jii*tily
lu in cooHudiiig that ib« one in capable and the other incapable of devvlop-
ing the impulwA, which w« call volitional, out of tbe molecular nutritive
cliMlgO of il« milMance. \W arc, therefore, led to the coochuion that ihr
Alllar nutiinuitic activitir of the brain indue to the intriitMC cbanpM of ita
wMaiKv lieing «i loucb more largely nmiiitcd by tbe influx of varimif nlfr-
iMt impulse* and iutliieucce, notably lb<i«e of the special Mnses. To this
•|iMetton, however, wc obnll have to retunt later on.
ji 008. In treating of (he vascular syntcm vro saw that llie central nervoua
•ystetn exercised ibriiugh the vasomotor nervM such an influeDcvon the miw-
eular coats of the bloodvessels as to maintain, what ire spoke of as " tone,"
section ol vaao consirict'ir fibres Icadinz to " loss of tone." We mw further,
thM arterial (one, though normally depemtenl on the general vasomotor
centre in the bulb, could be kept up hy ibc cor«l itself, that, for instance, a
tone uf tbe bloodraaselsof the bind-Iimbs could be maintained by the ianUted
ilorto-lumbar eord. This maintenance of nrlenul tune may be spoken of m
one of the " aoiomalic " functions of the spinal cord. We have ulao seen
that plain uiuricular fibres, other tban lhn«e of the arteries, notably the
flbrca forming sukincten, such as the cardiac and pyloric spbiuciem of the
stomach, the iipliinct«r of tbe bladder, and espevially tbe Hpbim-ier uf the
anu>, iilsii poMeas tone, and that tbe tone of thuK sphiiKten is alio de-
prmlent on the sfdnal cor-l, or on some part of the central nervoiu mtem.
W* Deed not repeat tl>e dinctiMious conceniitig tlit*e moehaDisms and other
£i>f the sjnaal ourd exercising an automatic inllaeiic* over rarioua
we Inve refirrred to them here, since they serve at an inliwluetioii
728
TlIK itl'IN'AL OOttH.
ilufiii); ilic ]MTi<Hl of Mliock nr» liDtp nad toaalns. In lil^ wnmi
nuiuml, a* «>' linvr •Atil. tin- i-flcirU uf sbonk nrc mucti more liutiiig
thi- cold-liLiocln) iiiiiniiil; iixl in tlir ilog the tone of tlip slfel<<tnl mti
r>-CiirnM tmicli more nl'iwlv ihun in iho Cru);. Iiii)i>ed. when tlt« •livisioa
l)ie (yinl tiM IbIci-ii fit.icr low lioarn ihi- «kf1eul lone returns very Blnwlj-.u
niuy \>e niniulifle<l very fmhly, or cvvii l>e nbwnt altof^Uter.' But iitii
t'avurnlile circumstHDees, when a suflicieDl Ien|i;th of coni bns hceii l«A. t
fnirly Honniil lone i) recabiblulied. In mAo, in aecordnune wttti tlw fi
|tr«viously lueDtioiKy) (i W£), skeletal loue. wbicli luu beeii Umt thrnugh
cuutiiiuity of tli« oonl bein^ brokeu Iij diseaae or acei<i«nt, aji|>r«n rarnljr
ever to reiuni fUlly in (Ik ref(i»ui below the l«rian.
We luav tl>ert>foTe o» (lie wliloe of the evidenw ouii«l>i<)p thai Um
Ifiiaiii-e <>r skeletal (<>ue U <>i>e of the fuDctious nt the cord ; but w« amy
Ttpeut tbat the cuiulttitxi i>f tlte cord, un which ditfM'ndii the iwue iVoin
c<>ri\ ailing rfl^mit iirrven of tbi^ iiiBiienoM, whntevor tbinr naiurr, whtdi
|>rodti<!r loiti' iu tiM' iniiaelp, tatty \m; nud imh'n) ik, in itH turn (lopondail oa
xflervnt im|iuL-H!». ht the cue of tlii' I'rog iiu<>|4.>d al)ove lIh< ton« of tbe pM-
denl liniW di»iip|K*r» "r i« irrcnlly U9«i-n«l whCD iho p<i»ilprior routs uf U»f
sciatic nervM nrc divi<l<,il, th<iiif!h llic HDtvrior route be [vIX intact. In iht
alaence of tbe usual tttn«in of nJfeTenl iinpnlK-? |)eMing into it, the cod
oeaMfl to send forth th« influences' whii-li inaintniu tho tone. Henoe Ihi
maintenance of tone jircwnts mitny imuli'gi^ with a reHrx ai-iiun, apecnitl;
wlieu we reineinlicr thai, ns ^laii^^l above, tone poMce itisemibly into coainu'
tion ; aud it may be seen a mere matter of w»rda whether vo apeak of lb*
maintenance of tone aa an atitontaiic or as a reflex action of the coril. W«
may. Iiowever, distini;uiBb ihe part nlaye*) by the afferent impulM« in n»i*l-
ing ibu c»rd to a condition in which it is capable of mainlainiujc 1(i«R' ftvni
ibe pATl played by an aflereni inipulae in cauaioK a reflex ai-tiou; in tb
ftinnertbe action of the aflVrent iiupubta seems anatojtmia loibat of a »upj
uf arttirial blixKl in mainiaining an adeiiuate irritability of the ncrvooii
Hiance. in the Utter the atfereut itn|MttMa lead directly to a divlmrs'
energy. Ami it in c<mvcnient Ui dtilii^Mh tlie twit things l>r ilifli-i
naiuiv,
<: 599. The cIoki coniwction between tiaie and reflex action !s illaitraiMi
by the ao-calletl " tciHlon-i)lHinonM>na." uhich.oii the one bnnd, arc cNwiilrrtd
aecaen of ordinary reflex action, niid.on the other hand, hnve bi^-n reKartltd
B£ exemplify iiig a «prcial inHiienpc of the npinal rairil tm ibv' irritabttilrt^
the muBcW U is well known that when tbe let; in )>tace«l in mi esity pw-
tion, realini; for initiinoo on the other leg, a sbarp blow im the ptttellar trMta
wilt csuae a sudden Jerk fnrwartl <d' tlu- log. brought uInxK bv a eonlraftlto
of the quadricepe femoris ; it is iieceeKirA' or at leaal deeirnble liir a pn^
development of tl>e jerk, that the twidoQ (and muscle t should be euaieohU
on the stretch. Similarly the muaclM of the calf may be thrown inloadiwi
by tapping the teodi> Aehillia, put somewhat on tbe etretch by flexion of A*
foot : and in some cases the same muacica may be mule to execute a seriM of
regular rhythmic con tractions, called " clonic " ouniractions, by miditcaly
preaung back the sole of the foot, so as to (Hit thetu on the strctoh. TIm*'-
and other inntanoea of a like kind, at firat sighl apiiear t» be, and Indeol sf*
by many obaerven) mainlaiuetl to be, oasett uf reflex aeiioti, due to slbfl
ini)>iitM« olurlc) in lliv tendon ; hence iher have been frequently spokM '^
a.1 " teiidon-n'tlcx." Other olwervcrt maintain (hat they are not rtnex. I>0<
duo to dircMi «limuliili<in of the muscle*, the vibraiiotu i*rt up in the nMt"
Ices tense t^'udon Ixing iran.'tmiltcd tothenniHcle*, and mi (bmning llie IsU*'
into contractions. The chief ar^um^^nt* apiinsi their being reflex arf >b*l
the interval Iwlwwu the tap and the iMnlni«tion in very short i dO^l »« tH"
rtvni
tllx I
rn» AfTOMATIC ACTIOXS OK THE SI'lNA'fc COBD. 729
Kinil ', ubiirtrr lliaii the i>nliiinrr iiiUTvnl "f n rctl"x iflioii <*$ 51^5), and
It ll>c niovt-roi-iit i>or*MlJ> nlliT Mvti<>ti of llw- Pfrvcs if the ttnAria. The
W •rgttDM'iil i* {H-rhnp* nut u vrn- vlroiij; on«. nn>l iW wmnil mil}- Iw met
\ mppodnft tlml. iii oudi n cbm- r1 If ivtt, if not nluiirs, ibo rcA<-ic a^rt rNilly
|ta> in tlw tniuch) Iwiug ntartinl !ii it by itic vibrations lTnn«iDiU«<) to it
tog tht iradon.
put •nn if wc mtmic that ilip iiiovcnicntii nrc purely muMalar, iinriM
il earrial out in the niii«cl<> wiili»ut tin- help of thu umal rcfl«i cliniu of
kmtt iu){Hil#es, spiiinl centra, and elTeivnt itupiilees, we niuM nt th** Mine
ba admil llwt they arc close dependent on the integrilY of the spilUll cord
d at the ronD«ctions betireeu the card and the muede. In the C«M of
liDflU they disappear when the spinal cord is dealroyed, or the nerm
Ing to tlte niUKleB are severed, or even when the posterior roota only are
ri<l<M). The nwaaure of iheir de\'el«pin«nt both in luiinmls and in man i»
ki closely depeadeat on the tondiiion of die spinal cord and of the central
l^'out system KeDerally. They may l>e increased or dimiaisbed, aH^nienled
inhibited by a roincideot voltintary efTurl directed toward some other end.
by the coincideol development of a ttnfficieiitly distinct sensation. lu
Deral it may be said thai whatever favors the lu-tivitr i<f the spinal cord
nrla to iuorea^e ibem, and whatever deprnwm the activity of the spinal cord
If is In itiminiMb tliem. They are diniini«hc<l or waniinit in certain diieases
the ipiniil iv>ni<,i;.i;., locomotor ataxia*) and exa|j:^i-niti:il in others: so much
tude«xl that thrr have become of practical clinimil impurlAiKv as a means of
iKnoaia. WI>otiii-r «c regard th<!m il> inilunciv nf onlinurv reflex action,
«ioiuMer tlinl th<--v are carried out by the mii^ole ibwlf and thai the cord
brreim otdy »o Cur a;" to increnac, raainUiin, or diminish the irritability of
I^JUiaciitar fnbatancc. il rrmaimt good that tbev arc prominvot whenei'er
■■BoditioDt iucrewe the rodcx or other oxcitability of the cord, and
ffiilh or disapjiear wbon tlw condilionH lower or abolish that excita*
Sit.
t 600. Disease in man re\'eals other actions of the Bpinal cord which bear
ituree diflereut frum thwe of an onlinary reflex uioveiDent, and yet have
fa dcKribed as retlex in nature. For iDitauco. certain affections of the
rd ftK diaracterixad by the le^a beeomtng rigid in L-xtrenie extension, the
;Mitr of the Btrai)ihtened limlia being often so j^reat thiit when a bystaDder
ta up one lej- fmni the he«l the other \tg i> raised at thi- raoic tiote. Ttie
>idiir is due to the exten.tor muscles being thrown into a ittate of contract
a. which in to uniform and long euntiauul that it may be »poken of as •
onic" ruulmction ; iiuch a tontc rigidity may, however, tie replaced by a
Pie* uf rhythmic, "clonic" contravlions. It fuut someiinHS l>een olHcr\'ed
K llie limbawhen flesiil un,'«oj>pli- nnd frrefrom rigidity, hut tltnt rigidity
ft In M won as they are brought into the pixtiliou of cKlcnKJon. the leg
tomiDc Midilenly fixed and Mrnight fomcwhat in the way that a rtiup-knile
ring! back when oftencd. It wcms clear that the peculiar contraction is
rriM nut by means of the spinal cord, but the whole action, though it i*
tfn spoken of as a " musclo-rcfles." is very unlike an ordinary n^dcx more-
int. In an ordinary morement an extensor is brought into action when a
)b is Aexed — not when it i* already extended; and if in a reflex act the
tditiuii of the ninscic about to be thrown into actlOli detemines in any way
I dischar};c of impulses from the reflex centre, we should expect that the
etching of an exieiuer muscle by flexion — not its relaxation by extension —
Uld determine the dischsr^e of exleusur impulses. In the ease of ibe dis-
irs in •jueMion just the oprHisiie seems to take place ; iIm poajtian which
pt-ar* to determine the development of the remarxable ctviilractinu it pre-
e'y that in nhich Ibe urain upon the exteoMTs it at it« wioimum. It may
730 THE SPINAL CURD.
be doubted, therefore, ivbether the word rellex Bhould be used to deimie suib
phenomeiia ; but the phenomena themselves deserve atteatiuu, e^peciuliv,
jierhaps, as showing how in the disorders of the gray matter of the cdfiI <Iup
to disease impuisea or influences which are latent only in health Ihihhk
actual and effective.
It remains for us to speak of the part plared by the spinal coni. a^ il>,'
instrument of the brain, in the execution of vofuutary movements ami IuiIk
development of conscious sensations; but it will be beet to con$i<li?nhtM
matters in connection with the brain itself, to the stiitly of witieh n m^\
now turn.
CHAPTER II.
THE BRAIN.
Os Some General Features of the Structure of the Bbaix.
§ 601. It would be out of place to attempt to give here a complete clescnp-
tion of the structure of the brain ; but certain features must be kept fresh
in the mind as a basis for physiological discussion ; and to these we must now
turn our attention, a general acquaintance with the topographical anatomy
of the brain being presupposed.'
Like the spinal cord, the brain consists of " white matter," in which the
nervous elements are almost exclusively niedullated fibres, and of "gray
matter," in which nerve-cells and other nervous elements are also present ;
but the gray matter of the brain is much more variable in structure than that
of the spinal cord, and possesses features peculiar to itself; these we shall
itudy later on.
For physiological purposes the brain may be conveniently divided into
parts correspon(line; to the divisions which appear in it in the embryo. At
BD earl^ stage in the life of the embryo, that part of the medullary tube
which IS about to become the brain differs from that which is about to
become the spinal cord, iu that the central canal, which in the latter is of
fkirly uniform bore along its whole length, is in the former alternately
widened and narrowed, so that the tube forms a series of vesicles, the cerebral
vesicles, succeeding each other lengthways. At first these vesicles are three
in number, called respectively fore-brain, midbrain, and hind-brain ; but the
fore-brain, after having developed on each side a lateral vesicle, the ojilic
vesicle, subsequently transformed into the retina and optic nerve, gives rise
ID front of itself to a pair of vesicles placed side by side, or rather to a sinf^le
vesicle with a deep median furrow, the vesicle of the cerebrum, containing a
cavity divided by a median partition into two cavities, lying side by side,
which open into the cavity of the original fore-brain by a \ -shaped opening.
This embryonic chain of vesicles is developed into the adult brain by uneiiual
KDWth of the walls and unequal expansion of the cavities, certain features
ing also impressed upon it by the ))end on the longitudinal axis, which
tak^ place in the region of the mid-briiin and is known as the cranial
flexure.
S 602. In the hind part of the hinder vesicle or hind-brain, the ventral,
basal portion or floor is thickened lo form the liiilb, while the greater part of
the dorsal portion or roof does not thicken at ail, is not transformed into
nervous element*, but remains as a single layer of epithelinui, adherent to
the pia mater overlying il. and so forms a thin covering to the lo/.enge-
shaped cavity of the vesicle, now known as thoj'juiih veiilrielr.
In the front part of the same hiiiit-brain, on (he contrary, the roof and
Bides are enormously developed into the conspicuous eertbel/tim overhanging
the front part of the fourtli ventricle, while the lloor h aho thickened ntto
tiie ponti Varolii.
^ Flgn. isr» lo^l. vhich will U' Itiiinil iii '■in'c^'t'iliit^' ^^i-Ildiis, ikuiv vi illi ii.lvjiiiTiigr U< i^on.'-ijiU'iE m
~i2 THE BHAIX,
This ihifkeiiiiig of tlie pons is Jurgely made up, on the one hand, of tori-
ziiiunl nerve fibri's, which run tninsvereely from each side of the cerelitlluni
inm the |miii». or from one side of the cerebellum to the other, and, on the
other hand, of longitudinnl fibres, which run forward from the bulb and vt
wmp|>ed round l)_v and interlaced with the others. At the front margin uf
tlie ]K)ns these longitudinal fibres, augmented in number, appear as two thi^
strands, the n-'ira crehri, forming the floor of the mid-brain, the rouf uf
which is thickened into the '■(•rporn ijuailriijeiiiina, and the activity of which
is reduced to a narrow tubular passage, the a'jarduet of Sylvint, oiiUrn
ti-TlIt) ml ijiiiirlam rentricnlinii.
At the level of the fore-brain the crura cerebri, diverging rapidly fhui
each otiier as (hey pass forward, leave the median portion of the floor of ibt
vesicle now known as the third efulriele very thin, but form, eepeciallv behind
and veutricalty, thick lateral walls, which are further increased in thickoai
by the development on each side of a mass largely compoeed of gray maittr,
known as the uplie t/talamus. The roof of the third ventricle, like that of
the fourth ventricle, is not developed into nervous elementa, but remust
extremely thin, and consists of nothing more than a single layer of epi-
thelium.
S 603. In front of the third ventricle each diverging cms cerebri spreii'
out in a sma^ j;^^ia1 tiishion into the corresponding half of the paired vesielt
of the cerebMuQn now developed into the preponderant cerebral hfmupkerti,
the two cavities of which are now known as the lateral venirida. Tbt
growth of the cerebral hemispheres is not only much greater than that of tb
rest of the brain, but also takes place in a special manner. At their fini
iipi>earance (he cerebral hemispheres lie wholly in front of the fare-braiB m
vesicle of the third ventricle, but in their subsequent growth, while expuid-
in<r in nearly all directions, they extcod especially backward. Thus, in the
adult brain, on the dorsal surface they not only completely cover up tbe
third ventricle but also overlap the raid-brain, reaching so far back u to
cover the front border of the cerebellum, while on the ventricle surfice.
though in the middle line thev leave exposed the floor or ventral portiouof
the walls of the third ventricle, at the sides they are seen to reach as far back-
wiird as on the dorsal surface. The median furrow on the dorsal surfiw*
which separates oacii hemisphere from its fellow is at first shallow, but
ra]ii(!ly deepens, so that lis the hemispheres grow they become separated from
each other hy a narrow, deep longitudinal fissure, into which, as we shill
see, a fold of the dura mater dip?. This fissure is not only deep vertically—
i. '■.. from the dorsal surface ventrally — but at the front of the brain nwa
backward in the middle line almost as far as the level of the third ventricle,
so as com[)]etcly to separate from each other the anterior parts of eifl<
hi'niis|)here. known as the anterior lobes ; at the back of the brain also it siou-
larlv runs forwanl in the mi<ldle line for a considerable distance, so m l*)
separate frouj each other the posterior lobes. Hence the two great maajesof
the cerebral hcmisjthere are united with each other, not along their whole
length, but lor about a third of that length, the isthmus or bridge thus cod-
necting them lying at some depth below the dorsal surface at the bottom of
the longitudinal lissure, in about the middle third of its length.
At its first appearance each lateral ventricle is of a more or less oval form,
its walls are of uniform thickness, and it lies in front of the third ventricle.
During llie growth of the hemispheres it acquires a peculiar shape tad
iK-comcs dividwl into an anterior cornu or horn stretching into the Mterior
jinrtion, a ]>(>3terior horn stretching into the posterior portion, and a dwceod-
iiig horn, which curves laterally and ventrally into the middle portion of ihe
ln-niisphere ; owing to the great backward extension of the hemijpbere! tie
TTrRE OP THE BRJ
I
lat«rat vAR8B eofuc !<> lie iioi uuly in frunt of hiit n\»n ut the ■!•!«■ of, luii)
iiid«nl, li> a ccfUin esttuit, «bov« or rlursnl !» llio tliinl vtuilrii^lc : hikI <Iuriiig
ihk gronth of ih« iNirls tlm urifriiially wide V'»)m])*H) •i|H.-ijiiig nhirh iiliu'rH
tbtkiod eixls of ttic two lateral vetilrirltv in oiiTiimuiiknltun nitli (liv fnmt
ut Ih* third Tenlncl« beeune* DarTuvtul into a uliiliki- fnuMfjr of niiiiUr
fen. th« foramen of Monro, wfiieb still <)|H-inu); inli> tlxr lV»nl ut llii- thini
rnitriclr, tiovt lru<l'> on i^ai'li »Ulf from ■ point rather in front of tbo middle
of thr lairral vi-iitricU'.
A« the bomwiihi-n- enliirgw, llie growth of the wall* of tin- vi^iclv i» luit
uniform in nil piirL'^ At niioirly ppriod tli«rc may be nb#erve<l in ihv vcninil
ytmll or floor of tlii; viviclr it ihickiiiing, whirh iLnumiiig a npi-cinl. niorc or
Ina MfniluDBr, form and pr<>ji:t;ling into tlic cnvity ttecomv* the binly known
a* tb« oorptu tiriatiim. As HfvrlopmenC procc%(t* the corpus striiitum on
«aFb ridv bccomtK attachtvl to Ihr npiic thnlnmi:*, lying l>chinil rimI to the
■nrdian fide of itwlf, tlm railiiUing librm of ihr cni« cerebri piming belireeii
tbfitwo. nodalwni' wcthall mv diviijing the corpux ttriatuni inio two hodin,
called the neufUu-' rauHatiin anil utn^r'U lentinilarit. A notnble rmult of ihu
rmwth and change of position of ihc hemispheres iind of the ciideseence of
Uie corpus ttriainm mtb the optic thnlnmtis is (hat the lutler btxly, ihouek
really lielonging to the third ventricle, come* to projcci somewhat into tT>e
lateral ventricle ; a strip of the HOjier surface of the opiic thalamus, alon^
its outer, lateral edge, forniB n portion of the tloor of the lateral renirii'le in
the ntedian re|;iou on each si^le of the third \'entricle. Besides ■hi-' Hiiecial
ileTelopiDeDt of the corpus striatum, the walls of each vesicle, witii the
txee[ition of the median part by which the two veeidee coalesce with each
Otber, become (we are now speaking of the higher luammala) tbiclccned much
io tbe isine way all over, the surface being folded so as to give rbc to ran*
volutions or .vyri He|ianiled by furrows or *u/c*; and the thickening taking
place in such u nav as to give the ventricle its peculiar sba|)e. Tbe mtdiau
coalewod pan underKoes a ditferent and peculiar chan);e. t'hi." |>urt, which
att fint lio« ill fnmt of tliu third ventricle, through the cbaiigm bn>ught
■bout by tlio growth of tlic hembpberei to Mttti its laitiitiim us to liv ininio-
Idialely over, dorsal m tlic third ventricle, very much ax if this part of tbe
c«n>bnU vsidts hnd lieen folded badt over tlw fore-brain. Id the junction
ttaalf we nav dlMingtiiidi a dnnml and a ventral ]M>rtiiin. Tbe dorenl portion
it developed into a srslera iif traiuvcne oonimiMinrul fibru* paanng acrom
ftooi one ncDii'phvrc m the other. In the median ref;ion them fibres form a
tlliek oompact hnml, i-alh-d the nirjm* talbmtm, whuh may be eiposed to
view at the bottom of the longitudinal firwure. while on each side they spread
away in all directions to nearly all |uirlji of tlw surface of the hcniispber«e.
pHiiog orer and helping to form the roof of the lateral ventricles. The
Mod M not Hat but cnrvctl vcnlrnlward ; hence in a longitudinal vertical
»ect)on of the brain taken in the middle lino it prcMOlta ft curved form with
the concavity directed vcntntlward. While tbb donal porti'^i of the
junctfoD is developed at the sides as welt as in the middle hoc, the ventral
portion is devdoped in the median region onlv, and that in a special way, co
lliat it fonns below, ventral to, the corpus callosum an arcbeu gilule, in the
ahajie of a triangle with the a^x directed forward, called the jomU, which
lie* tnunediately above the thin epithelial roof of tbe ihinl ventricle. In
tfhinl, the narrower apical |)ort)oa of the fornix lies at some little dittiince
below, ventral tu, the ei>rpus calloaum, and here the Juuction b<4neeu the
two vcMolea is rvduoed to a ibin sheet, tbe tcfilum laetJam ; but behind, the
bmuler baMi portion of tbe fornix b arche<l upsoas to lie tmmi-diately under
and loueb the cirpo* eullwitim. Henee the septum luddurn haa the furm »f
a mere w lc*> triangular vertical sheet, broad in front ilQil narrowing behind.
I
7-il'
r'i> . -■ iiiiiv l)f '■iiin'tivt-d ■'! ;i-
iiK'i ■! ■ . ■ .«! Li iiarrnw cl'i^il cavuv
■ ■iL- r ■ .^ i;-.;le lli<' hiUTiil v.'nirW-.
"rii:i;' ■ ■■-iricle, iiiiil th'- lliinl vrti-
ill- [■■. ivitli iht- fiiurtb vi-riiri-.k
.-ii';ii, ;■ .- -:>ina} cin"il. lln" wlii'ic- *rir-
utji.t! ■- -L.-:"ii caniil, tlir lit'th vfiiiriilr
N i>i' . i' ditieiviii iirigin.
''!'■"■■ ■ :ueli!>n iH-twoeii ihi- vi^ioiw
A: ■'. _ ..- viiole IviiiTtli, nil'! thu; I'TM?
i-.w\\ ■■ -. -. Lt'ilier ; llio uiiilille )inrii"i.
vi-i',!- I- -■■;n. tlmiigli i-iirviiiir over iht
ami I-.:.-... .. . ■i.-i'ili! pari* <)f,llR' Iiomia[>htre.
Iiy ill- li. ■.■ . J, '11 ^i>U' Ibrwnnl ami backward
kiMui; ,.- - ".;- '.Iimtijili llie c'<ir|ius (■sIIp'ub:
i!ii- i;'!;ii. . „ :;i;.'U'^ wiili that <if the yiW.
csli--- ... ■■;. i: i>I' the JHIIctiiill i;- verrilil-
tliiliiiii. -aLfsintotvco baitiU, kinjwn asiW
! 60:J .■ ur oi'iuiuuiuis ivilh, iiml irtkea
iiUL ill :i ■ ..-■■\^. Ill like nianiner heliimi.
!■(' III.' ■ .:::iun>iis vcilli the wall? "f ibc
till' :v.- . < mass of llie lieiiiL^phtrc^. Ifiiii'
j;ivm[i; - ■ .--.I allwl the crura i)f [III' i'lrnii-
ri-ii .■;■ -,-: -he attai'hiiit'iit." in t'ninl viA
ii|ijii'in:i:: -jrimieii iiiln llio .■iiil)>taiice "I'llie
vi>i, Ip- ,. :' niix iippearii 'Hi earli si>if '''
iii^' in If.' ' : -aalaniii:^, wjiicli htro f^iriu' lb.?
;iiliilt ':■ . -iitrii-U'; bi'twirii llio ii]inf [bl-
lliini ■,. ; ■ - ,-[us In be a iiainiiv <iii bv wliifb
i-iiv'f ;'■ aiis with jiiirlri niitsiilc it^elt' In
tb.'iijii ii- ■ -: ;■ c.mtimiity llumjtb iberv i* a
till' \.:i!'- , -i-'-iiieii uvoi' bv u biyer .if i.|"illn"
■LIS is iniiile i-i)iiti(iiioti> ttitii iht
«;ii'.i :i^
»liii'ji - ; ■ ■..; luMian wall of llif latiTiil vi-ii-
rii].i.|lv !• ■ .;::tUiiiii has tlielbllDHiri^- |iefiiliar
LM'-li •■■].■: I ■ .--„::
,-.■!■. ;; !'..i.. ■ ! - -, .-.uiriele. like that of the f.-iirili
i'. I.. :':..:i. :■ ' ■■ .■ I'lai ileviiid of liiTVoiis eleiiii'iilf-
li;ui%u;ii-.: _.. -ae hiiiil part nf the i.-orpu* w!-
<■. a- . ■ ,;.■.' ■ .. ...r. tiiiivl veiitrii'le, tbe free ii.i»i'l
lii'iiii-|.ii.'. ' ■■ . -e iMrpiir' callosuiii ab.ive ti.irmiui:
huh ...p.- I ■.■■ .. T bri'ljie betwetii tbe iwn henii-
i-e|iiir;ii. ii ■■. - - ; , rsal surface of the liraiii. |ia.-iii?
the I-. r.i.i:i, ■■ ■ , " -,„.* .vor t!ie ^)|^ofthL' third vonirii-'k
leiiilih, i-'i !■■■ ■ __-,-.jiijiiiti referred to, ami ihiis I'.inn:
iif'-iit.j iln'iii ■■. -: ■ ,,-, 11. which p^rvw !w tlie aomal r"'
llio |.>ii.;(iii.ini..: ■ - ., .,. ventral to, the l>)riii>; : ii eauD"i
\i ii.-ili-; ;i|.i. • _ :- ■drills. At the lateral iili.'0"ftlif
i!:- H!ill- :l!-.- ..I ■■•■ il...i: *b«t of pia mater ]iri'Jei'l.< ff'™
iiiiriii;^ Hi.. ji. .. -j.r-ole carryinft with ii the UviTfll
l«|..,'ij.-- .iisi<i. -: "'_. ^t rtlgo of t lie luriiis actually ''"'i'
|i .iri.ti. :: |"i>i.-p! ' ' ■ J" - ■■^- "-aicral veiilriele; the jtari ol ibr
i:i- i:-.irK "iiii'i ■ '_„ 1* tulire of thi! fornix when the Is'ewl
!:■ vi^-|ili.-r.' : M«':-' • - " , ,....,■ I'l.nw. Tj thi< i>eciiliar iutniJU'i
THE BKAIN.
7S5
i ili^ ptR nuiier, by whicli tbe nutrition of tbe bmiu ix awislod. w^ nhall
Niuni wbeii w« com« (u 'jieiik of the vanculnr arinn^mflnto of tbo brain.
JlcttawllUf Vf miiT piiiul out, thnt vrhilc Ibin viwrulnr itignivrlb wviat to
«uk» Ui« cavity of iIm> tbinl vencncli; ontiiiiiouii with that of the Iiit«ral
f«nlfid« on Mcb aUo, ud mil thrw wicli tlir ■■xtnrior of Iha brain, it rcally
doOT Dol. do Mil The mvttr of iIm- lliinl vcncridv k voiAq complete by th'«
jAjrar of vpithdium formiiii; if nuif, biuI tht; citvity nf the luteral rentriele
£• nuMl« oanip1«t« by th« lityer of cpithi.'liuni pawing fnm the lateral edge of
the fomit orar tba choroid plezui to ttiv oilivr pnrts of tbe wall of tbe
^rttUtriele. To pum alonz thii line from tbe nctiinl cavity of the Inteml intv
that of tin- third wiitride one mtiat firot pierce the opitheliuia ooverinj; tbe
«3horoid plexus, thus Kainiog acce« to tbe pia mater of the plexua and of the
^r«li)in, and then again pierce tbe miithelium coating tbe uuder aurface of
she veliitu and forming tbe roof of the tbird ventncl«. I( bonly by ibe
i'-iranien of Motiru thnt n r«nl com muni cat ion ex'ttxe between the eaviiy of
the lateral and that of the tbird veiilrii'le.
Thu» bv the Iar|>(> j^rowib and backward extension of the cerebral haini-
epberea, tfte third Teatride cornea to form as it were the front end of the
orBbi«>«pinal axis, the erara oei«bri expanding on each aide of the third
vasirirle into ibe cerebral hemiepherea wbieh cover up the ventricle on the
4araal !>urface. but leave it« walT exposed ou the ventral surfoce. Allached
to the donal aur&oe of the third ventricle at \ui bind end, ventral to and
tonevhai prtijecitiii^ beyond the Imae of tbe fornix, lie* tlie pineal gland with
itaattacho>enl*.iheremnantjiornun<.-»important nu-diao oi^an;KndatIaobed
In tbe \-vntml mrfiuv of tbe ventricle, at the a[n-x nf a funnel-abaped pro-
jvction. the infuitdibuluni, lies tbe pituiliiry body, alao a remnant of important
■acatral i>tructurw^
S 604. We may then divi<)e the whole brain iiiUi a tfcric* of ports corr«-
xndiog to tbe main division* of the enibryonir brain. At tbe front lie
oeretira] hemispheres, with ihc lateral venlriclcs. develupod out of the
'bral veaieles : and with thoss are asaaciatod tbe corpora striata, the term
rebral hemisphere bi^ng somelimea wed so as to include thene bmlie*. and
tM>iueliii>e6 so as to exclude tbcin. Next cume. correspond iiig to the original
't'orir'-liriiiti, iIk parl» forniini; the walU of the third ventricle, Ciinjpiouoitt
atmODi; which are the optic thaiami ; tor thoie bodies, though they appear to
intrude into the lait'ral ventricles, belong properly to tbe third ventriole.
la Um mid-brain which follows, the cavity, now the tubular iMssan of tbs
aquedoct. b roofed in by tbe two ]mir«, anterior and poatenor, of oorponi
•luadrigemina, the diineusioos of which are not very gnat; b«t a thick
tiour >• furnished by the crura cerebri. In each crua we muM diatingniah
httiraoo a dorsal portion called the Ugmenium, in which a Urg« (quantity of
gnty matter la present, and in which a great complexity in tbe arrangement
of fibres cxtsls, and a ventral puriion, the ftet or emtl't, which is a much
inon nnifiirm nius of lougiimlinally di.ipoW fibres. Ai the cnira pawing
Ibrward divenn into tbe cerebral humiupbere on ea«h aids, the t«gine«itam
OMuea at the hinder end and ventral |Mu-ta of the optic ibaJaidu*: it is tbe
pea which fuppliea ibe ma» of fibres radiaiiii;^ into each cerebral hemisphere.
In a view of tbe ventral surfincc of th* brain, tlw base of the brain as it is
fteqnantly called, the enirn may bo seen omergiog from the anterior border
<if the pon». Thii) we have ajMkcu of n« tbe thickened floor of th; front
part of the hind-bniin, hnt in reality it eueroacbes a lilUe on,tbe mid-brain,
the hind pnrt of the oirihini ijuadrigemiua haing in the earn* dorso-vootrat
plane a- tlie front p.trt of the poo*. (8>e Fi^. 180.) In the main, bowerer,
the poiia baliHies to the fore pnrt of the hind-brain, the roof and aides of
wblvn are deralopod, at wo have raid, into tbo cerebellum. This snperfieially
I
786
Tiie BRjLiy.
meitiMcs tho cvrebml hciuMphvros in iu Inrg*^ eizo, atx) in (he rpnijil
>li>v(!li>[inicnt of its eurrncv, whioli is formod ol' grny matter fnldcti in ■
n-mnrlcitblo mtinncr mikI oflrn ^txikcn of us cnrux. The wfebvllaia. th-iueli
tho Ititcnil |)Drlioos ohIIc^I iIk nemi»phorM, pr>:>jvct above the tnedUn fiJt-
ttoii, called the vcrmit, i«, unlike iho cerebriim. n single mus: euili latml
hair. Iiowercr. ^ends down v<>ntrnlly a maBi of libm whicli, ruiiDiDg tn»
Tcreelv, purely etid in the poDs mid portly Are continued acn«a ibe pMa iatn
the otiier lateral half; this nitiss of fibres, thus oonstilUlinjf, as wc hav« nid.
■ considerable part af the pons, fDrmit un each side, juH as it Itara tb'
cerebellum to enter the ptiDs, a thick i>[rand, called the miihiU ftr^nuAt -i
ibe cerebellum. From the cerebellum there alfio pruc^edi^ buckwafl ^ii'
the hulb on each side a thick strand of 6brea, the inj'rritir frdup'i'
cerebellum or resiilbrm body ; and a third strand, ihe taprrior jifi
the t'erebellum. pa^ce furwan) un each side into the regioo of thv onqm
iiuadrig«niina. As the latter converge toward each other behind ihv mr-
|)Oni (]ua()ri^u)iuft the ajijtie becwtwu tliem is iill«i) uii hy n ihiii dW >i
nervous matter, Ihe vnU-t of Viciuuenii. which thu* for a little di*l«iin- ba».
ward furms a ruof fur the fruiit rmrc of the fourth rem ride, just whrit'.U
luxengc-sha^ied CKvity is narrowing to Ixicome the a<)urcluct. itchiiid iw
cerebellum und puns cunim the bulb, which, a* we have Mtitl. u tlie lk>k
eneit fluur of the hiud [lan of the hind-brain, tiw mof uf the caviiy benf
ben jnolically waiitinfc.
Uftbcac KOveral iliviaions the fir«t division, that of (be ccfebral bmt
sphum, iuchidin^ ihi; corjioni Kiriiilu, Hliind* npnrt from the rvst by rtsM
Mlh of itH (iriKin "ud the chiLrnchrr of its dcvelopntciil. As ire sliall it.
this nnaloniicjiTdidiiiction corresponds to a physiolodcal diflerence.
Of tbt other parU of ihe brain the crura cerebri dcterve »|>ci.-iii] m'
We may repnrtl these as starting in the conl. but largely a»i((in>:wii
bulh; ihey traverse the pons, where ihey are Hill lurther imrin
paiuiuK beneath the corjmra •|uudrigeiiiiua, with which no well a^ "
cenelK-llum Lliey make connections, end partly in the region wJ' the t.jtr
tlialaoii and walla of (lie third ventricle, hut to a great extrul iu the rffr^l
hemispheres. We may ui a certain senae ojuniiler the n»t of tlie hnin f
buill upon and attached to tbu^ fundutiietital iiasal or ventral Mrand'.
^ 60S. CWuecied with the brain art a scrien of (laired iMrv<«, ihecniiw
nervui. The Ar^t an<l neeund pair, the olfactory nerves mikI the optic ntnv.
diffi-r in tlieir origin and mode of devehi|mient from all th<- resttofuad*-
mentidly as to cause regret that they ant inchi'lcd in ilu> mmv cate^rv. Wr
duill n>ii>i<)er these by thcinseive* iu ilue cour!>i\ The reiiminiug pair-. fr\«
tJie third nair to die twellUi, fi>ritiing a iuu<'h more homoj^'tieou!' nli^ri,
nc shall al»o consider in their proper place. We miirt now tiini t" muI*
in erMter detail fame of the «tnictiiral fcattin« of the brain, and «« an
v'lw iidvautagc begin wilh the bulb.
Tnit Bui-B.
$ 606. The spinal conJ. as it nsccnds to t)w> brain, beicuroes chut>, i
the more complex Imlb. partly by a shilling of the course of ihr :i
white fibres, partly by an opening op of the narrow central canal i:::
wide and superficial fourth ventricle, but chiefly by the dcvelupmeiit fi i''
gray ntatter.
When the anterior ventral aspect of llie bulh is examined (Pig. IH.CV
It will be seen thai the anterior columns of the eord are iutrrrupied i)t •"*"
iliiUnce in the meiltan line hy bundles of fibres (/^f. iltc.) which, ajipmri^F
TRB BRAiy,
to riM up IVom dMpv imrts. erne over Froia m\e tn u<le aui) ^t cnnltaK
line of liK anUrior flaturv. lliU is the iteeuMaliou of l/i« /jyi-xiniiit. ib
wbiH) lh« |tl«ee of the anurior colutuns of tlte Bpiual iiinl li utkou bv lai
Uri^r, man pn>iniiM.'iil <«)lumi», ihe jiyramids of the biilb (I'l/.t. vrliirl iv
coiitiniuil fornnnl to iIm! Iiind niniytn »f the imuh. Oil ih*- uuior -'
lateral to, each pyniinid Ihsm |ir<ii<H.'(in][ oral iurib, (A'* olimrji Ito'lii
ymiN' i/tn* {«/.) MiNintiiis: lli« ]ivniiuM ln>iu a <H>liiiiiti of wlitt<- iiiaii<!
rvH'/m'm boily (R), whicK. ocoupviiix Ihe luttiml region of tti« l>ull>. -i.-j
traced back wahl apiHtini to itmliuiic tli« liiK' of ttic Inlorml oulumn oftlM
cnrd. and wli<rn tmiixl furwurd in m«u to ruu U|> to tlK> cerabollum attbf
interior piilutiGlo of lliat or^ii. On the imtitrrior dorxil iwpecC no wtfli
dwiiMalion ia men. The two ixaurior coliinntii of iliv ii>rd divers* ^""^
eaob otl>er, leaviue Imiwv«d iIicri h (riausular «|>«in-, ihc mhmu^ ter^dom*,
wliicli i« the hiiM pitrl of ih<- limongo-Hbaped slinllow cuvity nf the l^itli
veiitrk-le. An the I'urd luiwov into th« bulb, ilia poKtrrior (.-»liin)H na ■ wlwk
l^rowb bnuider, nod tlw diriaioD uito a inc<li4iii jxwlorior and an extenal
jKieterior t-olumti becomes veiy obviou» am) distinct by the nppearEiine of )
ciuupictious furrow iM>|iiii-nling the two. At suoie dnCaiice, Itnwvrer, ia fnoi
of the point of divergence of the coliiiniM or apex of the calaiuua Krigrto-
riiu, (lie fijrrow bccumee leae marked, und it eveutimllr liidee away. In if
ii>urf« the furrow I«kee Mich n line that the mediau [nieterior ntltimti, forw
ing the immediate lateral boundary of the fourth ventricle, tuid the a]>pnr-
aiiceof aotraod broad behind but thiuuint; atray iti front, while the exterul
fMiUenor oolumii. aluo broadeniii); as ii advaDcm forwatd, eeeitui Ui ite wedpJ
It) betu'eeu Ihe median piMerior column on its Dtedian ed|;e aud tV- —•'■'—^
budv mi ita lateral edge; beiice iIm former is here called the / >?
jHHiruiiui) ^meili* (iw. p. ), aiwJ the latter the faacieatH»{<it /unimif •n"-;^*'
(e. u.). Furilier furuard bolh oolumna aeem Ui merge wiili mch otlm whI
who Sbrea which curre rouod to fono part of the mlifunn body ; tli' <<'i
lions, buvterer, of theae two columna In eadi other and to ihe other i>h -
the bulb, na well ua ilie nature of the other aeveral changiii hy wtii '
oord in trattaforoieil into the bulb, are diacloaed bj trauavene vtmii-al !
Tcolml) sedioiia. to the »tudy of which we ninxt now turn.
A wction <l'ig, 1>*7. I) tiikiTU at the hind ninrzin of the decuwilin). nl
which Icvtl the lirrt i-erviail n«vi« take* origin, when eomrarr<l wiili a "■
tioD of ihe cord ai the \vv*-] of the aecond cwvioni iier^'e (Cf Ktjj. l^J '
shon-ft that certain i-hnng*-' arv almdy tiikinj; platw in Ihe gray matti-r. !>'<
anterior horns arc not much altered, htil the poatarior lionu are, as it *'-^
pushed out laterally and dunsallv m thai the ])oaterior columns, wbirh a* ><■
nMain their previoua great <hi>t&, become rery much hninilcr liiaii Uic* *rr
lower down, eiusnuohmg. to to M)Mnk, on the lateral oalnniOR. At ilie auM
lime the autMlaneo of ItoUndn (<■■;;.)■ forming ihe hei^^or eH{>ut of ihr b"m,
hut enlarged into a more or ten globular form, and lice near the Mirf* t '
the curd thotigh aeparalcd from it by a eomiiaci tract of Ioii);iiudtnnl it-'"
( r. a.), which, aa we aliall sec, beloofp t<i tlie tidh cranial nerve. A '^f-
oiderable developmeut of the reticular formation (/ nrf.) at the »i<le of ''"
]{rar matter ventral to the poMerior horn ha» also taken iilai-e, ami ilw ''^
the ahiAiug of the poailton of the jMMlerior 1i<hii hiu <lriveti the Uietsl h'm
(t. h.) DtMTer to the anterior horo. l-"riim ihid laletal horn a nwi uf thf
eleventh apioal aooeGBory cranial nerve {XI. i may be seen taking oH^i><
Further, a gfoai increase of ijfny matter rnund the i-enlrwl c-aitnl lun.r >!*-'
be obMrveJ!
Thtae ohanges, however, are of decree only ; vrbat seeioi to' be aa ab**'
luU-lv new feature ia the preaeuce of bundtea of 6bres (/y •if<.\, whiebH"**
iiig from the anterior t^dumti of one aide oroai over to aud are appaTAj^
I
'sr'lliiJUKkoflhciuilcriar horn of Ui« other side; ill
enauug ibv Itlim) |>iii>h Mldt tlit bottom of tlie anterior finun. Wlieu
th« counw of tliiwv libn.^ i< iDVf«tignted. either bv 8im|>lp iDit^riMcopic obKr-
vktioa, or Mill better by the indnod of de^nprncioa. it is fuiind that Uiev
luav be Into-') fnxn thr iitiKtrjor coliimn of ooe »iile, acruw the itnterio'r
oiiDDui^ire, tlmMigh (hr ni.'vk of the anterior born to the lateral ooluutu of
tbe oppOMt* tide, and to that pnrt of the Inleral culiinio whivh we have pro-
vtouslr dcwribed w tlio crossed pTnimidiil tract.
lo a NctKiii a little higher up < Fig. lr^7. 2), these decussating llhred form
an each side a largv ttrand which starts from a part of the anterior colDmu,
BOW l)ecomiug distinctly ninrked off* as the pyramid </V). and U appnrenily
loH in tbe reticular formation, but in reality paasoa on to the oriKiM>d pyra-
midal tract of tlie lateral column. This strand, as it cro«es over, oi^nipletely
cuta off tbe head of tbe anterior born frxim the more central fcniy matter,
«nd fornu with its fellow a lari^ area of decuwattng 6bred between the
botOui of tbe anterior Gwnre and the central gray matter. When u •iirfMe
now of the bulb ia examined the decussaliou is oeen to be cnecUsi by alter*
Data bundles, pawnf[ now front riKbt to lefl, now from tell to riftht ; and in
timnaverw aectiona we find currei^puniliDftly that the anterior fiMure appears
boat now to tbe leA and now to the riKht, acconlinic tui the tcclioD cuta
through a bundle podciio): from left to nfUt or from ri^^h to lell.
In wctiou atill hi^^lier up (Fie. t^". -^ and 4) this convpicuotu ttrand of
fibna craiiDg oblionely from Ri<le to »ide, will be no lou^r mcb ; dwumMt*
ing fibroi are »ecn iIokbI to the anterior fi-iture. but tbiste, of which wc shall
tptaJt, praently, are of different nature and origin. The librc« whii'h in
•Mtiona bdow wen- *tKo in the act of I'roMiiii); un.- now giilhcnnd into ma«M
«( lon^udiiial librv^, the pyramiiU (Py.j one on tuurh «ide of the anterior
Cmuiv. each with a M-c-liotial ami of u roundnd triangular form clearly
narked out from the ourroiindiu); Htruutunst; tbe wction ia taken above tbe
deeoHUioD of ibe prramidc Or, tracing the chaagta from beloir upward
ve may My that ihe decussation is nuir complete : on each side the whole of
ibe croaaed pyramidal tract of the spinal con] has, in tbe region of the bulb
bdow tbe level of the present Hctioiu<, croascd over to the other aide, and
joining uiih the direct pyramidal iracr. of the anterior column of Ihe cord of
the Mme side boa )>ecome the pyramid of tbe bulb. In other words, tbe
daOMMlioD of the pyramids is, as we have alrea^ly hinted, the piissini; 00
AxMD «*ch i^ramid, and the crosnng over to tbe opposite side of tue ooM, of
Umm* (ibrta which are destined to become the orosiod pyramidal tract of tbe
•piaal curd of the opposite side, while the reet of the pyramid pursues ibs
coune uo the auae side as the direct pyramidal cracU
f, 607. In the npiual cord the bottom of the anterior Anure ia separated
from ihv central canal by nothiu^ more tbno the anterior white oommiHure
and a onmiw hand of gray matter. oi>mpii£«d of the anterior gnir comouMare
nod of part of ihe central Kelatinoua Bubitancj. During the diwusMtlon of
the pyramid«, the dciAuuaiin); lihrui nuah, as it were, the central canal with
ita surrounding gray mutter to dome dtitance from the bottom of the anterior
flnure. In aectiooK atravc the deoussatiou the bottom of tbe fisi-jre duo* not
•eaio npproocb the central canal, but eootinues to be removed to some dis-
tance from il, and, u we pM> upward, to an inoreafin^ distance, br the
inierpoNtion of ti«ua wliiuti connNU Ltrgelv of deeiusating RbrtM. Those,
however, though they seeni to cuntinite on tlie decussoltou of tltc pyramids,
are ^liown by the cmbryologtcail nnd degeneration methods to have no noo-
nectiun with'the prrami>h, but Iwloug to another syatem of decussalioo. Aa
we have seen (S ^'^^ > thi- imtirtor comminure along tbe whole Icarlh of tho
Okfd cuutaini dccuuiiling tilin-a. SoBM of Umi* In tnc Upper part of the cord
■ rv fibret oraaaing frnni lli« direct pyrnniMal IrsolDf one aide ft the K^ay
nintt«r of the ntber iJd«. aiid ai> iiiuy b^ rcirarded ns part of the wbule pyn*
itiiiln) tract: but aihont un-of dilTLT«nt uriuin; and even in llie region of the
nciunl <Ieniaiiitioii nf tbf jiyrHUii'U mxnr »(' thi^ libnn whtcb enaa nrer An nut
hdoDg lu the pyramiiial Inti-t. Tliix fptcin of tIeciiawitiiiK fibr<9 bei-iinii>»
iBcnuin^ly prutuiii<int iilHivr! t)it- di'i'iiwvilioii of the [lyminli].-', iiin) lhn>iij;b
it the reuinl Krvn of the bulb brtwi-vii tlic a-ntral canal itml tlic itnU-rior
Amuiv \m niurh in<'ri>iL-M.il. Tbr lilirv^ lui ihi^v rrnM foriii ii iiii<Mt<< linv of
Fttrtilion, tho rnpht (Fig. IS". 4. ■'>, r.). wliicli incniajiw in di-plh in lh« upper
fartri of the bulb, kod on fAch nd<> nf tlic rnphd hiOp to brcnk up ih<' i;riiy
innttvr (wbich nnriously fornxH) th<; niiUrrior horns) InU) whnt i« uilli'd tlM;
rHietilar JormoHon. Wc ohall return to thi^ prwpnlly, but mity here call
uuntlon to a special dcvolopnicnt »f tht'w (kousaaliui; libn^ wtiich is smo
jatt above ttio uecuaation of tht> pyramids. In a seotinn at Ihia level (Fig.
147, 3) II atrand of fibrw {supra Pij. •/««.) may be seen to sUirt chiefly frooi
the gradk nucleii* but also to some extent from the cunealv nucleuE. to
SW««[> round the central gray matter, and to decuasate ventral to this between
it Kiid the bittl«nn of tlie anterior finure. This is called the tuperior (i:WuMn-
(loij. or, for renrans which we abalt Bee later on, the »en*or<j dwfnition.
§ 606. We must now turn to the poilerior fissure and its relations to the
fourth venlricle. We saw that at the be);innin<j of the prrantiditl decussa-
ikta, tbe poaterior bora^ had been thrown backward and outward ao as to
iacnaae w» noeierior columni^ The [luateriur fis&ure is ftill of ureal depth,
■o iliai by tbe increaae of depth an<l uittinteuao<* of depth the porterior
column, the laierol limit of which ioalill iiharply marked out by the swollen
bead of lb« poMeriur horu as well lu by the hif;heiit posterior rootlet* of the
first eervieal nerre, acijuire^ ul this level In mtticimum of hutk.
Fri:>m thia |>uiDl fom-»rd the depth of the )>ot>ierii>r fiuiira and the dorwv
ventral diameter of Uie posterior column* dimintiheii. The head of the bom
iFig. ]8T, 2) ie thrown Mill further oiilwan) into the lateral regions; tbvel-
opmcnlA nf gray mailer at the biu>c and to iwme extent at the nock of tbe
horn iiif tbew we nhall tnmV prrewillv) cticmnch (Fig. 187, 3) donally OD
the while matter of the oduninK; sod the central gray matter uppcnnt to riae
doHHilly at the expvnec nf thr ponlcrior 68Siirc, in coiticidcnco wilh tbe
'levi-lopMipnl d«ficribed above nn taking place on the ventral *ide "f the canal.
Still a little loriher fornnrd. in a section lor instnnce (Fig. \^~. 4} n little
wnv Ifrhind tbe apex of tlie mUi>tt\u tcrijtlorius, the central gray matter,
tthtcli still Ibnns a rounded maw around tbe central canal, is brought yet
nearer to tlte pneterior fiaaore.
In A section yet a bitle further forward (Fig. 187, 5) carried through tlw
liinilcr narrow pari of tbe fourth ventricle itself, it is aaen that the central
canal hna opened out onto tbe doraal surface, and that ibe gray matter, which
nusniklamait litR*:/a (.asittnal aieiiBMinni: n.)aaldoa<iremb*ltor tnci: tt miUBtm
Milr or laftctor ptdniwl* of lb* ranbffluni ; t p pilamal pn*l(rinr Mlumn. lw.-lmi|ii» oaaMlaa :
o.p luaAMi toMlM coIUMD. AMculiu ■ncltli: r. npho. t.k. UUnl Iu>nii a.^in. nooleuaf
•ka B«dlaa ptMM lor oaluiDn ut iir»cjlr iiuulaiu. f.p. it. iincli'iii ul UMicslatakl poaMclur uatama Or
rWHMtaoilMa'.cy ■ in > ni<ill*n illtMan *iid f.y.n. (1.^ Ulvml illTHI»n nf (ho Mint i g< olMiy
bartj': ■<.«. ««4bia aMBHiry. anil el.t. Uutml acertotj ulln: fi.oL lliIer4B*wr Imyrt: M l.m.
laMiml iwu«D-l>Wi«n niiclmii; n. s. ueiwla iiui'tow; a r. mmiaoiuf uilarlar bom ; / m, i*ll«a-
Ur tonaOcB^ I g lutvnuice of Ralaula : a. r r I. ii»lcrtori<K>I.UMl p.t.t 1 |dM«iIoc niot nf Rnl
MnMalBM**: XL rwiorirtaatMHMctraem: X11-(iitUUiartinn((oiat<>arTc: ■■ XlLaurltwi
al Q»mm» la •; Ihv aurtM* inky In lflw*il. hOBDiw. ihi«u(h f.E'i.S.la oooMrtloa with Uiv
ntm ft llM MtT* ; ( X Hiiun' "f auln fat of 1W sbmo-|ihM j mwl-taso^otfirir iMclaaa : X.
«. ■atffUHlWUnribfi n«iKat nacl«U*nUt"u*: IX.ii.BM«*idllig>a>t«<ll>e(liw>'pbu)rDKsl
tiivl«a>i V.« OMMidlBC cm) or lb* aftb iicrro: Mb. &«rtb TWUMa: tbs opMlrai or hDlag I*
liBUcaHdbralhkb'tarbUnu; uidniAaDd«.UMia>tb-Ilk«N«U0Bbrik>|n4Ktliwol«it u*b»B«,
r42
THE BRAIX.
In pn-vioii» »ccitun» «iirr»im<lc(l it, is noir cxftottd to the Hirfitn* i>n tliv H—t
of the vculricle, the nitilioti posterior columns bcin;; ihraitt ii»m]i-. In n ui!l
more fonrnnl »0(-lii>n tFig- 187, 6) this gniy matter in i:orTinp«>n(l«'niv «ith
the ini-Tfiiring nii]tli of (lie ventricle octupie* a still iri<lrj- atvn, tlinittinj
Still ftirlher Rtido the Darrowiiig upjKT enil^ of llie two puttterior tiilimin*.
During tlieso ^ucceeeive chsnge* (he large, wide posterior (iHiih exienia)
piMierior nnd nicdian pnaterior) cctluiiiii^ of the cervical apiiiul i:<>nt aad
Itef^innitig bulh. arc reduced lo«Dall dirniTiiioiiB anil iu llie eud <lianii|>eaf ;
but licfore we fpeak of ilie coune and fule of ibi? traclii of fibres euustituiiiig
tbe^c columiia we must lurn lo ihe imporiaDt cliangee of ibe gray iimlier.
S 609. A trniisverte tection itirniigh llie loner eud of ibe decutMlioti < Fijt.
187, I) BboBB, ua me linve said, few difleretices sa rtfuards thi? grwy luaiter
from ODC taken al the level of tLe teuond oervii-al nerve. Tb<- cbotijtn
noticc«bl« are muinW Iliii <'liiiu)^s in potiitinn of ilio piuiic-rior bortia, the
increnM' of central gni^' mailiT around tlie central ennui, the nppnjQcbt^tbe
lateral horn, fVom which spring the root* of the Hpinal Bccvaaory nerve, to
tbe anlcrior born, iind an increnne of the reticular lorniiuiun in tbo bar
ventml to ihe no^lerior born.
In thi; inidale <)f ih« (Iccutantion (Fig. IftT, 2) tbe di-cuantiiig Sbrc* are
cutting the hcud of the aii(criur horn nway front the Imm of tbe horn anj
tbe cenirnl gray imhiilHncr. and the iiiolnteil hflid is diminitbcd in *iie, bnng
aeparaled from the surface uf the «>rd by an tncrciu>ing (hickiH-oa of white
matter. The lalcnil liotu iind origin of the ipinal acccm^r)- r>H>t do not Rbai^
in this isolntioi), but an.' driven back again liormlly towani ibc pufia-rior toal
to join ihe rcticuliir formsiion which w iiicr^iing it) nrvn, while the lairral
column of white matter is diminishing in hulk by (be uiilulrawul of die
pyramidal tract.
^ill a little further forward, the anterior bom aeeuu at firet sight lo bave
wholly iltaajipearcd ( Fig. 1S7, 3 and 4), hut its <)uappeaniuce is cnjuei^ni
with an increase of tbe reticular tbrmation in the poaiiion uf the lateral
columna, a& well tta with the growth of lieaue mentioned nbove l>eiucen ibe
anterior fixture and the centml ^ray matter. In fact, between the anterior
pyraintdi on the vebtnil dde and the liirgely increaaed and laternlly es[Min(M
gnj matter on the domil nde. a large area of peculiar timue now L'xten<l* os
MOi side for a ctmsidcrahle distance frrini the niid<ilc line of the raphe, en-
croaching on what was ihv lateral »>liinin o| white matter: and a com-
<>punding arvA of liniilar tiwiie miiy be traced from this level ibrongh the
higher parla of the bulb up into the jmns nnd crura cnrchri. 'flic timor con-
sists of nerve fibres running lrtin.->venu-ly, loiigituilinnlly, and in other dirv^
liona, so ns to furm a nrlnork. the Inm of which are often curved ; and with
llicae fibres are Ibund hmnchcd nerve-cclU in eousidrrahie number, "iiuv uf
tbem small, both librcs and cells bifing a* elK>where cnilwdde^i in neuroglia.
Though diHering from thn ordinary gray mailer of the cord by the more
open cbarncler of it* network, it may Be considered as a liimi of gray matter.
We may consider it as being in reality the giay niniter of the nppan-oUy
loHt anterior honi hmken up and dispersed by the pnaage of a large number
of tibieM and bundles of tibi'ca, cG{iecinlly of the decnssaiing fibres "poken tif
in g tiOti, which ^ince they curve through ibis area I'roni tlie middle line
laterally are called arcuaie or nrcifunn fibres, inttmat aifuatt fibrtr i Fi{.
187, (>,/. a. i'.) ti> diatiuguish ihem Irom the nrtenml arfuiile fibre* (J. n, e.)
of which we »hall apeak presently. Fragmenis of more compact gniy mai-
ler, also belonging probablv to the anterior horn are teen at iutervaU in (hit
Rrca. Fig. }f^'i, a, OB. and elaenhere. We have st«u that nearly all tbe way
along the cord the gray matter of the neck of tbe jiOEierior liom is similarly
broken up by iMindies of fibm into what we tbetv adied the reticular foruta-
THE Itt'LB.
748
Itun (Fig*. 176. 177. f-,i- p. iind r.f. /.): kikI thi* ftrvn iti tbv l>ulb though it
poan-wri rhunicltfiv nf iu, own w nit" rnlM the ntif^ilur formalwn. la tba
n\nn iKlcrnl portinn uf this fonaniioD. thv network i« man omMi nud irregtt-
t«r. IIm> Rbrm nrv liner, nod the nerre-c«lls nre tnorp nlxtntjiinc ihiin in the
intduin portioD where the nerve-oells. except id the immeiliale neighborhuod
•if the raphe, ire lew numerous or even absent, uid the libr«B are coarstr.
Thcae two p«TU ar« •ometime* dietiDgiiished ns the outer or laiornl, and the '
inner (It iDcditn formatioo. In ltt« middle line the Abrea dialiocllr interlace
ud tleciMWte in no obU<)ue nmoner. some running; nearlj vertically in ihe
doreo-ventral pluoe. thus i-oiiMituiiug an we have eaid a thick raphe, which,
however, nl ili> evlj^ea |,>nidually merges intu the more u|>en network.
i 810. Within ibc iirou. btiunded oy the pyramids veotndl)'. Ihe ex|>a»ded
tnjr mnUer ilorMlly. tbe raphe in tins niKldle line, and the it bite oiatlcr
]al«TKlly. cerliiiii dintiuct compact cnaMM of ){niy muiier make their appear-
ance. H> we p«M' upward lowunl iht- ])(>n*.
Onoof ihL- nxHl im|Mir(anl nt' thtw givea riwe H> itie olivary body, ar hiftriar
*iit< vhicb, an <><■ have M>t-n. pniji-cl" nit an oval mam (Fig. IM, o/.) un eech
slHr cif ihu |>yniniiili, n:3(-hiiig Irnni a level which is Kimewhut higher up
than Um-- lr>wL-r limit of the pyrainid:<, almost but not i|iiito to the jxin^. The
ulivary ImhIv, as a whole. (iiiiiiRl-i partly iif white nuilttr, that ii>, of Hhn^,
and partly af gray mailer, si>ii)Him<r« (■allt-^'l tbc nlivnry nuclni*. Thi» latler
M dispowd in tiiv form of a hollow I1u>k ar riirviMl howl, wilh dn'ply fohli-d
or plailed walls, having a wide open mnuth <lirt!Ot»'il iiiwanl towani ihv mid-
dle line, ami forward tonarxl llic jwng (.Fig- 1^' ■ ■*■ •'*• 6. "'■)• The Ila*k is
filk'<l within hy white matter, and covered ap ou it« onLdde with white mat-
ter as well as iraverwd by libm. The grey Riftller thus forming this flaak-
naped nucleus consiM of small rounded nervo-celU. lying in a bed of limie '
ahich is parllr ordinary iMuroglia. nud partly a fine nervous network.
Lying to the median side of the olivary bwly. immediately dorsal to the
anterior pyratiiiil is another small mass of gray matter, in the form of a disc,
appeariuK in iraiiHvenw sections as a thick bent nx), in some sections c»ns!sl-
of t»t) (larts (Fig. 187, 4, <//. a.). This is the afttttoni olirjtn/ »iif/riu. A
verj- similar h<Kly ties dor^l to the ulivarv niicleun. in the lateral reliculnr
fnimation ; this i* also called an acccwory of ivury nucleuji. Win^idiNlin^iiiibfd
(F%, IttT. (i. tl. f.) by the name outer ttpertturi/ niiclfiin from the nbovc-mcn-
tloued (n'lrr iirwjunry micletu. It will be olxvrveil in thou; iraiiKven** «e<y
Umu that the inner acemaurv nuuleusis W|)antle't from l)iv olivary nucleok
by a bundle of while fibrt* (Vig- Iff^.J.S, fl, XII.) which, ninning venlrally
from llic gray luatlvr in the donuil region, como to the wiriixcc between the
aoterior pyrnmidn and the olivary body. This t* the hypogtomal or twelllh
cranial nerve.
On the Burlace of the anterior pyramtti itself is iwen oo each side a small
mass of gray matter (l-'ig. 1A7, *■. (I, •■. a.), which since it appears to be cnn-
oeci(«l with a tyslcm of superlicial transverse fibres, which we nhalt descriiie
ilirectly as th* rjlrma! anuatt fibrt* I Fig. 187, 3. 4, A, tJ./ «. f.), is called
the arnwfs t»tete*t*. It seems to belong tu the same group as (he accessory
ulivca.
Ljutlv, a small fomewhal dilfuse collection of gray maiti^r i^ seen in *««•
tioBf OS a rnumlcil mat« of irregular furui placed lateral to the reticular
formation (Fig. 1^7. \, o. G. a. I. ».). Thb, which at its firat a|i[imnu)co
seem* to be hmlded ofl* from the general mat* of gray metier ' Fig. 1^7, :t,
a. i. Ill and which is probably a detached |K>rlii>n oi the base of Ihe anterior
liom or of the Uit-ral ri-gion of the gray mailer, is calle<) the tattral or
aiiUro-laifral nucleii*-
Htnce, bnides the diffuse reticular formation, tbi* rvntral pari of the bulb
TRB BftAIK.
cvintniDR mora nhiirfily <l<-fiuci] collH-tionf of j^rKV matter in the nlivarv
nack'ns, nml ihouiluT hiHliesjuH inciitioncd,
i 611. Wc niUKt niiw mm to the duraal pari of the hulh. Here in thefini
place TC6 mual <lwlingiji$h belircen the jxirlioos of gruy niait«r which are
more iinmediaiely coniivcteti with the cranial nerve* taking ori)(iii frrmi ihi*
fnrl of the bulb, and the poriioDB wliicb have no !>iich nhvinim iiinr>ei-ti»D.
n tiM spinal cord, the anterior honi» luippU-, (u ir« luru m-en. ihi- origin* iif
the Mivceasive anterior motor nerve»: but in the trni>)>l<irRiulii>n of thu oirit
into the htilb the anterior honis have been broken up or ilMplovwl ; nnH tbr
(lariK of the anterior honid, serving a.* tlie nucki of orisin for motor ntnta,
lave been translated fruiii the vi'iiiral Ki the more doml rt-gioiia. Heac*, il
is in th« more dorsal pari uf tin.- ^ray miilter that nv have to wek for the
nuclei of origin not only of afler<-iil hut alno of ni<?lur cmnial nervrjt. Il
will )>e conv«nienl to oonuider all ihisc nuclei nf origin if crnniBl ncrvM bj-
thenuclvr*. and we may here eontine oursclvw to the grny matter of Other
natDrc. We inuy, however, »av that thiw nucli-i from that of the third
tipr\-c backwanl aru more or ftsa clowly luwoi'luted with the griiy matter
immediiitdy nurroundirig the wntral cunnl, Thi» rrnlr'il omt/ mattrr, in tli«
namiw *cnec of the irrni, i* marked out Komcwhnt low down (Fig. IBi. 31
by tti« tihrcn of the Kmeory deciissntion winch sweep round it: it appenn in
aectioUE higher up a« a fairly iliHinct region (Pig. li^7, 4) ; and it is tais pan
of the gror matter which is exposed on the floor of the fourtli ventricle wheii
the central cnnnl { Fig. 187. o, li) opens out into that ap«i». We but ex-
puaed ; but in renlitv the irne grar matter is covered by a Mi|)erficial layer
of tiflnue of a pecnlinr nature < indicated in Fig. IS7, b.d. by a ibiek bUck
line) similar to tlial which is found at the hind end of the conus medullaris
in the spinal cord.
Wo saw that at the level of the lirat cervical nvrv« cnueEdcnt with tbt
horixontal flalteuing nut of the (losterior horuii the po^Urrior coluiana awumnl
verv large dimenwions. In this re|rion f Fij;. 1N7. I ) they eonairt entirely of
white mnlter — that is, <if louKitudimd ribn-s.
At a little higher level, however, at the level of llw middle of the deouMi-
tion f'lr example, aii inlet of gray matter (Fig. 1S7, i, m.p. n.f makw its
appenrancc in the niiMlian poMienor column. A little further forward. At ibe
level -if the wtulilish.d pyriiiuid^ it will he Ken (Fig. 187, 3) thai thi» islrt
il' the hind end of an invavion from th« more centraTlr placed grav matter,
•and that at the same time there ha* taken plttc*- a similar inroad of gmy
matter into the exteninl |>osti-rior column (Fig. ^>^~,'i.f.p.H.): indeed, a
tllxhl ext«Dsion of grav matter into the oxierniu ixwtcrior column runy be
SMdi even before this (Fig. IriT. 'i. f. p. n.\ It wilt furtlwr be observed Uiat
UieBe gray masses have so largely memncUod on the white matter that iMth
the ntedian posterior or liixciculus gracilis and the external posterior column
or fasciculus cuneatus. instead of bi'ing simply tmcU of while tibre«, as tbey
were in the hinder part of the bulb and in the cord, have now beoorae
i-oliiniUB Iif gray matter covered by a relatively thin layer of whito flbrea.
Tliw* columns of gray rontier are now (called reswctively the malian piutt'
rinr nurluiui or itur/fiu/<itciruli gKtcUi*, or. more snortly. the ymeiU nueirtta;
and the fxUrtial panttrior nufieiu. or HtuteuA ftueictt/i eunfid', or the runmf*
titie/'K*. From the ventral asj>ect of these nuclei a larger numl>er of Abna
]>nw ventrally, with a more or lea* curved course, to form, at we hnvv Mcn
r$ H06), the su]>erior deeuitMttiou and to pursue certain ]>aihs through the
reticular fornmliou, of which we shall s|)eak later on. It is at this level and
for some litile dlntauce above (Fig. 1>'i, 4. -i.t that thew nudri aoinire tlii^ir
gn-fltcst develiipmiiil. Further forward (Fig. 1H7, 6), when the fourth VMk>
tricle has o|)encd out antl the nuclei of the cranial nerru are becoming can*
THK Ht'LH.
rift
anil ibc iMwIerioi- coliimite have been lliriim asiUle laierally, both
llirw; iiiicli'i Imrir ilimiut^hed id sixe: Hill furt)i«r furivartl they hecome Btill
UDiillcr, moA loWKrd the pons they it'^'ui'lir (lUapfieiir.
Tha UMi of gclsiitioUB suhelDDce, fonaiDg m the level of the liral cervical
ntm the nrolkn caput of llie lioru citee lo the Hurfare h«l twparuted (Vom
it hv a haud of tibrea ( lo.) of fine calibre, to nhieb we have already reftrred
aa belofigiog to the S(\h cranial oerve, luereaMM In bulk tit a txiineivrhat
higher IpvcI ( Fig. 187. 'i, 3. *. .7.) and forms on the Kiirfai^o a nlight projec-
tion, railed the ItiAereU <tf Jiotando. It Aotiii. howi'vrr, bccnaiM ihruM v«d-
Irmlly bv ihediverniMeof the poateriorcolutnna, and rDoro and morerAvemd
op !«■ tbe tibrea whieh are jcMnj^ lo form th<' incrMwiuf nvliforni hmlv (Fig.
IsT, 4, o, 6. ID. Uetaiiuog this |KwitJoii ihe ulct of gclatinntis i^ubctniiw
dimitmbea in sixe further foruun] (Fig. 188, '. ^.),undev<.-nluallydi«op{>a)r».
rtu. m.
r>9
TO
Wm^
^~i
TUH,
VHl
CaJ.
■
■ Ta>u«uM nta tM-ta iiwr Bui;3:d iiik fio*. iRMSaaiKnTuii.i Takm It tlw Uiw t<a, FI*. IM.
V P$. HKinWt: Jt. trntUem bcdT; ('Am. r*rtballuu>. f. mWi: J.n.t. ritcraiL/.A 1. luMtiMl
I MOMM nio: L bunatoof niM nuu cdlre la tbc lenUoular ducIimis; / piHWIu ko^mdlual
' bwil**: ■./.(. aueleiu nf IIk Iho'U-iiIui lent; i if, iiiiirrlnr nll)«: ti r, i iiivlifut niitialU (lite
Ifta itHMa ll knanlloTMcif hiimllnnlAlim Iij'wMrti ll li irmrmti; t.t, tutaUiiMotnoUiKlo;
lr,a.aw«iidlaKniMoraRhnetvp: Villi, nuoieiuor Uu-Hnnifenom: Vlll.audliurr nerrcohlelljt
OaitofMla* aaclllwr taH: Vllt. u millan nueitua; VllL;t laU'rai niulnn^ VIII, j «n— or?
■Mtaw ll' HrillMy n*rTO. IX. Km* of rw>( nfulMb aonc fanliw llxniiKb •irtdpllntt mil of nafc
I
$fll9. The jthret t^ At Mb. It b» <ibvi«i». fmni what hax already been
said, thai the arrangenienl iulo puitenur, lateral, aud Bntnriiir colunin*, »o
clear and delfniic in the Hptnal van\, liecontea broken up in the bulb ; indeed
it will lie beat, iu treatini; of the bulb, not Ut atMmpi to tmoA out th«ae
colonof, but toMneak uf the coiirw nf the MToral tracts into whidi thaaa
oolnnM may be divided.
The direct and cm«i pyramidal traoU of tha pan] unite to fiinn, lu <ra
have H<-D. Ihe pyramid of the bulb, and so pa* (>n bt the pons. We nood
aar nothing mora nt prcx^nt i-»iiccrning this important pyramidal atrand,
exoepi that, tu we trace it down fn^m the ponn to the ipiiuil oird, il Kirea nfl
to thf bulb itself tihrra which iniik': ronneclions with the lontor libr« uf the
ertuiial nervM [ir<ic««dinK from thi* region.
Coftceruing ilie course taken by the other Ic8< con'picuous " dMoending "
tract, the autero.|al«ral descending tract, our kiiowlmlgc is very imperfect ;
nothing ilefinite can be aaid nt preaent.
The rerebeltar tract, occupying near to the surface a poaition wbid) in the
■erfaa nf acvtioiu ( Pig. 187, V6.) apjwara naw rather more eeotral, no* more
748
THE miAi.v.
(IofmI, t'TcntiiAlly jawa int» llio rc*liform b>iilj, af vluch it fortn> a iT^t
|>Hrt. and ihiti rcnch^s llie ccn?belluin. Thi- ■nlpm-liiCrnil •KyiHliug irad
|Ki»ibly III*') mkcs thp Mtnc coiirBc, btit tliin i» not w yet iTcrtnin.
The median pi:«IeTior Irni/t or column, becoming tlie fawiculu* ^tnifili*.
ends in the gracite Durleut ; snd in n similar maniter the eitrriial [■■(tiiiir
column, or fasciculiiH cuneatus, enils in the niedinii and lateral miiMM i>f ibr
moenle nuclcuf. As we fauve seen, the while mailer of tbiw? Liiltimn*
dtminiBhee as the nuclei iiiereaw; and the Diidei after alworbiag, ■<> lOfpnk,
ihe white maitc-r diuintbheH in turn: the Hfcendini; d<-){en«rBtioii itWrvwl
in thcDe eolunms Mo|ia at thene Dudci. Il h a BUj[f;cMive fact that as thw
nuclei diminish lurnnrd the restifurni liody increawa in hulk.
The remaining Rhies ol' tbe cord, belonging iwrtiy to Uic amemr colit^^
and partly lo the lulcml column, nol giiihen-n into any of the alxivr-ta^^H
tioiied tracln, amieur lo fud, chiorty iit nil events, in the reticular funuamV
»r ihe Itulb it«eir, though vnmL- an- oarriml on to the higher parts of (he
brain.
i 613. TlniM, «r th« vtirii)ii» tracU or stran<la of Ihe aftinal cord, two tath
are known dcfiuildy and cvrtoinlv to pun as oonspicuouH unbrukeu MraiKH
thniiigh thu bulli to or iVoni higher parts: namelv, the |>ynimidal liact m
th« cerebrum, luiil tlic corebelliir tract lo the oercbcllum. All, or nearlv all,
the reM or the longiludiiml fibres of the coni reiiching lh« bulb and. as ura*
ncknow iit jm-^ini, in some pnrt or other of the hulb; and we m»y infer that
romcor other ncrvi^cells of the bulb H-rvc as relays to oonncct Uieae fibres itf
the rord with other pnrle of llie braiiu
The gracilc and cuneale nuclei sttuid out conspicuously as rclap nf (hit
kind, nnd through them the posterior columns of the cord make Mcoiidarr
connections on the one hand with the cerebellum, and ou iIk- other luuiJ
with various regions of the cerebrum, We have »aid (^ 607) that 6hra
passing ventrnlly frtim the ifracile ami cuneale necH sweep in a cui
couiw tlirougb Ihe reticular formation ik.-> tlie internal arcuate fibres
187,/. i.t.). The hiudmoHt of these form the ^uperinr decusHatiuu all
referred to as seen in iiei-tioua of the fore-ptirl <>f und in front of the pyramidal
■lecuuation (Fig. 1K7, 3, mpr-i Pi/. •Ire.}. AAvt ducuwaliaR Teiiinil to the
ccutrnl canaf, thrai^ librcs fonn an area called the inter-tSirary la^r (F1||.
187, \, in. oL), lying dorsal lo the pyrHmids between two (lie <ili%'ury Dttclei.
This layer may bo regarded as the iiind end »r ht^^inninji: on each side of a
remarkable longilndinnl strand culled the- fiUft I FJNy. I'^U, B. F., 189, F.),
iii the ronnei'tions of which in the front part of the brain we shall speak
herenftcr. Thus these two nuclei are ihe source of librvs which cnw* lo ibe
other )iidr of (he bdlb. and rcncbing the inlcr-olivary layer ijonni to (he
pymniidM run up lo higher parls of the brain by the fillet. We may add
l^nl llic lomiation of the tillct i* also probably lumslcd by Hbres fn>iii n imct
which lies just dorsal to the inter-olivary layer, and is <leriv<-d from the nntr-
rior columns of the cor^l. Besides it< fibres of descending dr^nemiion thv
anterior column contains fibres of ascending degeneralion, and tbcM' riw
donally in tlie bulb in form llie tract iu question. Though the whole Iract
is of some leoxtb, the component fibres appear to be short.
The gracile and cuneale nuclei ^ive rise also to other fibres whii'h. though
also iweepiog reiilrally and croMiug to the other side, do not. when they
m^h tlie iulei^olivarr region, asaiime a longitudinal direction, as di< the
libm Innning (he fillet, but as external an-uate fibres (Fig. 1^7./ u.^.)
pursue n course nbtch is at fir«t vititral alone the side of ihe anterior fissun;
nnd linn lali-ral over the vcntnil surface nf tue ]>yraniid and olivary nucleus,
by which path they reach the hiUnd sutliiccof tbc bulb, aivd do the rmtilorm
body aiwl (.■rrebolUim. In ibis way the two nuclei iu (lueetion contrihutv ti>
I
I
i
opposite «de nf th« bulb. Th<»e cxtenuil nrcuntc
n, «hich as llVy sweep round ihe veulml siirfnce of ihe pvrniaid tmrcnfc
arcuate nucleue, ihoti^b lli«7 vary much in iudividiial brniiKt, fonn a
(■oDNiIcrable |M)rtiua of ibe while iiinller neto on Uie ventml and btlcrul tot-
Ciuvt of the bulb : it ta by tbeiu ihai the olivary nucleus is covered up.
The cuoeate aod Krocile nuclei, bMltlea (hb eroaaed and Kmivirbnl nnintl-
about cuDoectioD vrilb ibc rettliforiu body of the oujioute aide, are licliui.'ttl
10 hare more direct connection with (be rcsiliforni ImiIv nf the unie sido by
■MUU of fibres whtt^'h piua by a nicirc or Uta dirvrl lateral path IVom t htm to
It. Accepting tbiH view, ne may mv that the twn nuclei are cooiiectoil nith
the oppcnte aide of the rereWlluni by I'xlcrnal tin-tiate fibreti, and with tlic
Mue Bid« of the cerebullum by the other lihrt^ juvl uienliune^l. In any vote,
(he conne«tiun betwt«ii the two unclct and ilie c^rebollutn ia larj^ and
iniporlaot.
Tbu* ibe in)|H)rli)nt »(i«nd i>( fibrrv which w called in tlic bulb the re«ti-
fona body, and hi):her up thv interior i>edLinc]c of tho erri'lielluiu, is con-
iirctrd with the spinal conl in two chief ways: dirvctiv by ni««Dii of th«
cercbrllar tract, and iiidirecllv by means of the euneiUc awl gracile nuclei,
which, aa wc haro Mid, diminish in bulk forward ss the ru>tifiH-:n bmly
iorrtfiMs. By tbo relay of the gracile nucleus it is brought into couiiectioD
with ibe mcdtui poetvrior column alonj; the whole length of the cord, and m>
with Ihil diririon of Ibc {Kisterior roots wbich (S 578^ in each of IIm> aeveral
•|ni»l aervc* goes to form that column. Uy the relay of tbe cunente nueleua
K » brought into connection with such i>art8 of tbe exlernal posterior cohiian
■a cod in that nucleus, and thus probably with oilier &brea of the i)0*t*rior
nols of ibe upper spinal nerves. And if we admit tluit the cerebellar irael
itcoanecied. by the relay of the vesicular ej-linder or by other ocrve-celb,
with tlie rest of the pii*ierior rooia of tbe apinal nervcw, we may condnile
that tbe rtatifiinn biKly is, by ni<-an;> »f thnw relay*, a prominent contiuua-
lion of all tb<- xpiiial [»»li'rior root*.
Tbe rraliforni ImmIv and m thfl c«ndwllum is also specially cnnncoled with
the olivary Ividy of the oppovilc «idc ; for when in young animal* one ilde
of tbe ccreMlum is re^novet), the olivary body of tbe opponite pidc atrophica.
Till) (^Mirw of th(^ Rhres niainlaining tliis connection is not ua yet nocurately
known, but they nrolMibly na«H from the olivary nucleus of one nde tlirough
tbe inler-olivarv layer, niKi m> latomlly through the reticular formation of
the other side. I>iistly, we may add that a imct which is aonietjmes included
in the mtiform bodv as its median or inner division has quite a dilltTeiil
origin from any of the above; the Ubren wbiob compose it cume, as wc 'ball
•ev. from the auditory nerve.
The fnnher connections of the bulb with tbe cerebrum it will be beat to
leave ontil we come to deal with the structural arran;(ement of the rMt of
tbe brain.
Meanwhile eoouch haa been said to show that ilie bitib iliflera very mat«-
riallj in Mmcture from (he apinal cord. The gniy matter of the Iwlb is far
mure complex in hj iia nature tluui it tliat of any luu-l of tbe cunl ; anil tl*e
arrai>i;emeut of tl>e several Mnindx and irai-(a of hbrui i* far more inlticat*.
The Hiruciural feature* on the nhulL- (icrtiap* uigfcctt that the main functiow
of the bulb are twofuld ; ou the •me hand, it *cem* titttvl to Mrve an a bond
eentr« governing the Kpinal oiinl. tbo variitiiii rein* of which, with ibe ex-
eapthma rio(e<l. ii hi<ld>, »■ it n>-r<-, in il» bnnda: on iho other baml, it appare
no Ina BdH|)le<l to act una middleman between parts of the apioal c<^ liehiw
and varioua regiaiiK of the brain aboTb Aa we shall tev. ex|wrimeDt and
nhvrvntinn ^vc Rup|>ort (o tbcM aiiggcaUou.
TUB BWATX.
Tne DisroatTioN and (kkMKecTtojts op the Oray avu White
Matter of the Braiv.
THK (iBAV MaTTRB.
S 614. At we fuu» up fiMin the btilb t« the bigbcr prb nf the brain, tbe
iliflt-ri'ii lift ion tif Ihi- gniy timtror into niorv or Ton npnraU nuun*. whkh
nx- linvo Ki'ii Imgiii in tho bulb, bccooH* still inoro Mriking. W* htre lo
(listiDgtiiith n Urge number of ttmu or collection* of gray ninit«r, mntr or
low rvgulnr in form mifl morv or Icm shnrply HclinoaJ, Jrom (1m- giirmuu'liii;
white matter; to such colloctioiis the sercnii temii corpus locus, nvurlctu.
hik) tho liko, hiive from time to limo been given, llwse arcu or eollvctioDf
vnrv grciitlv in si7«, in Ibrm. nnd in histological chiirttct«ra; ibey dilTer frooi
eacK other in the form, eiz«. features, and arrnngemeDt of the ncrw-celb, b
tlie ehantctera of the nerroiis network of which the nerve-cells fortD a part,
and especislly pcrhapi' in the extent to which the more distinctly ertr inattff
is traversed ami broken iip by bundles of white fibres. tiuJocff by Uir
analog}' of the epiiial cord, us'well ns hj the r«suhe of ex|>eriineDtg and
obaervntjons direcled to the brain itself, we are led to believe that tht com-
plex functions of the bruin arc intiiiiatelv aseocialed with tht« gray matler:
and a full knowledge of the working of ite brain will carry with it a LdokI-
edge of the nature and meonin]; of the intricate arnuigenMnt of the cerebral
gray matter. At present, however, our ignorance as to tbtse things a grou.
and, though various theoretical cloniHcations of the several colleeUoiu of
fny matter have been proponed, it will perlmjia be wisest to content oat-
aelvea here with a very broad and nmple arrangement. We will divide tbt
whole gray matter of the brain into tour eatogori<« only: 1. Tlio oentnl
gray matter lining the neural canal ; and thin we may coiwdrr the nuclei «t
the I'raiiial nerves, some of which arv cliisclv ammeiated with iL '2. Hw
superlicial grity matter of the roof of some nf the main divifion« uf the bnin.
such ax ihnt of the cerebral hemispheres, and of the cerebellum. 3. Thr
tntermcdiftto gray matter more or Icm closely connected with llie crnrt
Gtrebri. 4, Other collections and areas of gray matter. Wn will, moiv-
over, confine ourselves at present for the most part to their sonemi fe«tur«i
nnd topography, reser\'ing what we have to aay oonceniing tltPir histnlog'
vlinractcrs for another oocai>ioR.
1. Tht Ointral Omj/ Matter and Ihg A'lielti tf Ihe OnmifU 2fant».
$619. The veiitrldea of the brain, like the oeiitral caual of lite sr
eerd. <>f which they are a continuation, are lined by an opithdiiim whic
in ^lerat a single layer of columnar oelb said to to ctliate<t througboalT
thotigh it U olten diHicnIt to deuionstrnte tlte eilia. Beneath this opiibelliim
Ii4v a layer of somewluit [wciiliar neun>ulia, fonuing with the epilhelium, ••
wi.> luivv said <$ RII). till' rpfiiilttma, wliicli, well devel(^>e<l in the tltMir of tiia
f;>iirlli vrntricli- nnd in the walls of the third ventricle, and of tlw atiMdl
i* ibin nnd scanty in the lateral vcntridtw. ncncatb, and more or Mn a
nectdl with the I'jwndyma in tlie eith^ nnd Hoor of tbu third veniriclo. b I
fnirly conspicuous layer of gray matter, which i* well developed in the jwut*
of the floor exposed on tlic ventral surlnce of the brain, and known an the
lamina terminalis, the anterior and posterior performed spaicK«, the tubv
cinereum, etc. This layer is not continued forward inin the laiernl v<-itlri>
elea of the cerebral heniiBphcres, but it ie w«ll developed iHickward along iht
TUE GRAV llATTBIt.
T4tf
I
I
<FttS. 191. lf)3>. and in lliu fitior t>r tbts fmirili vvnIriclF. luiil
lh'niu)[b tbvi l>iilb Ikvoidm, m we biivc Men ($611). rontinuuus tvidi ihv
oDDlrnl tny nwticr of tin- cord. Tbc itcrvo-fcllfl of tbb graj nmttcr are imi
Uw « h»ii' rniall iind in many plan^ scanl.
<i 616. Thr Fcvcrnl ronis of the cninial nerrps from thc< ihird ticrr<^ hnck-
wiril niKv be tnictHl within llie bnin Bul»tnnce to apccial miltH'tions of gray
nijitlrr, callfid the tttiriri <if Ihe cranuit nefif». »onie of which lie (ilnap u(>on
tbr ihe I'dilrnl gray nuitter, while others arc |tlace(I at eutno dislunce from
it. The optic utTTe and what ia w>m«Ulii«B called the olfactory- nerve, nsniely,
tli«- olfactory bulb and imcl, mar advaDta^>ua1r be dealt iviih ufiart. aiuce
dim.' two nerves are not, like ifie other cranial oerreB, Dimple outgrowths
ttotn the walls of die original nenral canal, but are in reality enlougaied
Tcaii'le*. budded off from the neural canal, the caviiitw of which have been
obliieraied. We may add tbat gmrt uf tbi^ rvtitm, and of the gray mailer
of tl>e olfacion.' tract, may |M^rlm|i9 bi- cc)UHt<kT(.-d ua corresponding to Llie
nuclei of which ne un* niwakirig. tin- rrtiiial and nrojier olfactory &\>n* I>dng
MUkeeted with them very tiiiK-h n« thv iibrca of tho n-maining cniniul nerves
an eoanected with llieir rci>)K'(-livt! nuclei. In thir brain tb« Mgmi.-ntnl regu-
larity of the nejve nHita :ici <:-'tii!-|)icut>u« in the iiiintil cunl w very greatly
obacored. We nball have aomcthing to «ny on i.tii* pi>int Intrr uii ; bni at
preaent we may be cunlrnt to trnt ihcM<veral nerve* in a»imple to[iiitimbical
naanner. Tli«-y may ba seen in a vvnlnd view of tbe brain i.Ftg. fS*}, C.),
leaving the bniin at rartoua level* by what is exiled ihdr "i>iiperficial
urigin ;" the third nerve iwiiing in front of ilic pons, and th« biat or hrj'W-
glwml itrriebing back to ihc hind end of the bulb. Part, indocd, ol tlie
rkvenlh nerve, tbc »ptnal accessary ncrrc nroperly so-callMl, maLea con-
DfCtiona wilfa th« Rjiinal con) Ivelow the bulb ii» far back as tJie rixtli or
wreoth ecrvicsl nerve, or even loner ; but this part may by ibcw- codihh:-
tiana be distingnisbed from the remaining part of the nerve, an well as from
all otber cranial oerrca. I'be nuclei to wbicb Uie nerve roots may l>e traced
witbia tlie brain anbataDce, sometimes spoken of as ibe " deep origin." range
in [KviiiaD Irom Ibe binder purl of the bulb lo the bind end of the third
ventricle. Tbe nurleus of the byp»i{liiH!<al nerve begins ut ilie bulb Just
above the decutaation of tbe pyramiiff. the nucleus of the third nerve ends
bennttb the hind end of tlie flour of the third ventricle; and all tbe rest of
(he nuclei may l>e broadly dtwcribed oa placed between th««e limits in i,-arious
|iarbt of tlte llixirof the ceuiml canal or in adjoining *lruciuR», though part
of one nucleua, iiamely.tbat of the fillb nerve, can lie tnu-ed. ad we iJiall we,
back Into tbe sfiinal cord as far a* tb« second ocr^'icnl ner^-e. and probably
eximda Mill farther. Fig. l!)-t ba diagnm showing in a rougbly aptiruxi-
oiftta manner tliv nuclei of tbe «eveml ncnta a* tbcy wouUI nM>enr in a binlV
rvlew of tbe floor of the miutildol and founh ventricle looked at »n tbe
■I atpcd.
S 617. Thf tu*/Jtli or htfpogipMol ncnv. The iincleus of this nerve, which
it will i» convenient lo take fiivt (Fig. 1I>:!. XIL), is a long column of gnj
niaittr lying in ihr bnlh parallel to, and very cl««e lo, the median line. It
rcaebeafrom tbe hinder pari of the fourth ventricle at hIhiuI the level of the
him) end of the auditory nucleus, as far bn<-k as beyoiid tbe bind end of tlte
olivary liody. Ai its extreme bind end or l>e^nning (.Fig. 161. 'ii it occu-
pica a venlral jrasilixn and is n jwn of tbe anterior born ; tnenoo il grmdually
risee doraolly (Fig. 1M7, -i. 4, 6), but so long a* the central caDal remaJiw
oloeed, oootinues to oecujiv a distinctly ventral p<isition in reference to ihe
oentml canal ; in iu front |>art it is. by the o^itucup of tbc fourth vrnlriclu.
Imxight into an ap|karenily more dornl poHtlon (Vig. 1^7, (t).
The nucleus cou--tieta fuainly of large nerve-cells with distinct axis^yliixler
760
THE BBAIX.
nracccMe, which tliuiigh [lurauing ■ •omewhnl irrcgiiltir cuurtp niiiy br tncti
into the fibrai of the ucn-e. These, xtarting rn>m the renlntl *ur^nc« of At
uuL'leuH %]oag its Icuglli. ruu vciitnilly thmiigh lli« ivtinilitr formAtlon. Bttd
making tlietr war in n n-t'uf at huixtli'v, 1x^1 necn thi^ oliviuy uuolriu on ibr
Iai«nil eUif «licl tlio pyramiil n»<l nxiltiin nccxMory oliro on ibi- mrHiaa itdc.
f^tan the Rirfacc nloii); tho groovo nhicb MpnniKs the pyrttuitl fnun (lie
olivBrv bodv.
5 6l8. JlSf iii'dM or ^lonnphari/ngtnl, UhUi or raz/w, ouil flrrevtA or ipkutl
(ir«MOrif nent-A. It will hv ndTKiilagoous to consider these thne owrc*
togeth''.r,
in the spiiinl ncctMOry nervM we miiEt dislinguiah, ■» we b«x*e aid. two
rarts : the " spinal iu!ce»ory " profipr. Ibrntod by tite roots which come mS
noiu the oervic«l suinnl coni, reaching as lar down as the sixth or seventli
cervical aerre, and the "bulbar acceBsory," whoM roota oome off from the
bulb Just below the vagus.
The spinal acce^ory proper takes ori);iu in the ({roup of oellti lyiQf; in the
extreme lateral mitrgin of the anterior horn, from whence thi: fibmt pruoeej
(llrectly outward through the lateral column, and bsuc from ilie curtl along
K line imraeilinie belnceii the anterior and jiosterior roota; the up[ier roa*
undergo, uiih the portiDu of tlie luientl horn from nbich they vpring, tht
shiftUtg spoken of in $ flOti.
The Dulbar aoceeaitry starts from an elouiated nucleus in (he bulb whidi
U common to it. to the vagus, and I'l the |i1o««i>-|>har%-iigcaI : hrnee we ban
taken ihc*e three uervea together. TbiH (Fig. I'J^i strviclKv further fiirwaH
Ihnn the hrpoglownlnudeua, reatdiiiig thi^ level of tbetmiuvvnw tibn^catM
Eirin; acuKticui (Mr.), hut doea Dot extend do far bi^hind.
In transvcnw nectiou* of the hoth. which jMm a lilth- Iwlnnr nn<l a liltir
above the point of the calamus scriplorins (Fig. 167, 4, ■'>), too uuulet or rol-
ledtooBof cells art- seen in the gray matter round the central caoal. The
more rentral one it the hypoglonal nucleus, the more dorsal one the hegiu-
ning or hind pan of the conibiDed accoesory-VHgo-gloMO-pbarii'utreal nucleiu.
When a tiltlc further forward the central cansropetiB out into the fnurtb
t-cntride *by which change the hypoglacaal Ducteua (Rg, 187, fi d. xii.) U
brought nearer to the dorwl surface in the floor of the fourth veiiiricle) ihb
combined uucleus, increasing in breadth, is ihrowu to (he aide aud H.-vumosa
more lateral pontion, lying now on the side of. but still nuniewhat donal to,
the hrpcigliissal nucleus, between it and the noir diuinishitig gtacUe tiuclMa.
In thtii pDtiilion the nucleus appeum to conun of two part*, a tnedi
laleml, the ujedian part having ouiisjiicuuui oervfr'^jelLi of modt^rate luse,
]at*.-Rt] iKtrt having hut few cd^ls, una thute of small «»:. From this i
the niicli-U£ runs lurward. nainiaiiiing nearly the same pinilion iu the floor
of tbo fourth ventricle, but gradually becoming thlaa«r, and ends, oa w« hare
c«id, at about the level ot' the siruc acuaticfe on the donal aiirfiMO oorrespood-
iDg on the ventral surface to a level a little behind the hind margin of the
pom;
From lhi» combined nucleu:*, biil chielly from the median |mn. tibrea
fiwcep in a ventral and lateral dinx-tion through the donal part of the reiie^
illar fbroiation, paw ventral to. or in Mute caM« through the gelatinous Mb-
atancc and the Ntraml of fibres connected with the jiflh nerve (Fig. 187, V.
a.), and rcudi the onrfuce of the hnlh on its lateral aspect iu a line bHWMa
th« olivary and rcsliform bo<lici ( Fig. 186, C). Along this line may bo
wna.
I (Pig. 180, C,\B««-ricaof roots; of tlieM the lowest, the acoetaoryroota,cpriag
^n-nin the hind part, the hi^^hesl. the g I oero- pharyngeal roots, from lh« front
part i.nnd it u thcee ee|>e<-iallr which pierce tlie gelalinoiu stibstaao^^E^.
ilX. a), and the intermetftaie, tlie vagus roots, from the middle part iif
TIIK ORAV MATTER.
751
bined nuckuB, Hence we timr speak of the bind part of (li« wlmlc
Il« VI
I
I
I
beiDft iht acve»or>' nueleua, Uie middle [mrt u tli« vtupu nucleuii
frviit part of ihe Kl<B*u-[>hnr_vni;eat nufleuA.
Ail tbr litim, boH«ver, uf tlu- fiuis of ilu-iae tbreo oerves do nut takeorigio
tran the noc-leus in ({uettion ; *i>iiii^ »( the fitiiva iitart in a diffcivDt irnr. In
M«tiMl» of tlie bulb above the decur^alion of the. |iyranii(I a patch of ffrav
(ukIUt i* MCfi lying iii the lali-ral part uf thi-- n-tii'iilur fonnatum (.Fig. If^i.
\. III. >, about niidnay bKvii-n ventral aod ilunuil mirface)^ What i» tbw
diMlra**! by Kctionii 'n a cditnin of ]|Tay matur, ihi^ " nuckuN anibiguns"
(Fig. \'JZ, ii'i.),*In1cliiiigHhoiil a* farlurwanl anil backward at ihr combined
MVtwtry-vago-glomo-phnryngvnl iincli-un, bnt pUcM) (ll<inctlv more ven-
tral ty and KinMjwhat more rnlcrallr. (In' l-'ig. 1!:f3, it anil tfie combined
iiiicleu» lira rrprrm-utcd •m diHcrcnt ■Jd» of the diamiiR to iit-oid confusion
thrmiirb th« overlapping of thr shading ) From it nbn* curve round (Fiic-
irtT, X.iD.). tojoiD the accnsory vago-glos8o-pharyiigv«I roole, but e»pecialR'
tfae vagin naU. It mnv therefore Tie consider^ u a wcond nucleus of the
fwui (ftod powblr of t)io other) roole.
But tliwe i« yet n third MuroeofsomeDf the fibres of the nerves of whidi
we are speaking. In eections through the bulb there may be seen jus) ven-
tral to and a lillle lateral to the combined nucteurt t Fig. llil, 4, 5. l>, 1\. a.),
ihe circular feclion of a longitudinal bundle of tibres, In the binder sectioiu
I Fig. 187, 4) the bundle in a very ihin one, and -ilill fiinlier back it is lost to
viaw, tbough there are reasons for thinking tlini dome of the flbree are con-
dniisd back into the cervical cord, as far as tlie orlgiu uf Ihe fourth cervical
tMrre or even beyond : in the more forward sections ( KIg. 1^7, 6 mid 6), ii
iuu'ama in diameter and may be traced forward U> tliu rmiil end of thccom-
bilMd oucleiu into which il metres. It id a buudic of lilin^a whicJi starting
tuccMHvely in the laleral ^ray matter of th« cervical cord unci higher up in
the rvticular Kirmation of thv bulb, run longitudinally forward : tbe bundle
ai fint iocrcaan in sixc by the luhlition of fresh fibres at each step ; but fur-
ther forward the fihroi lc«vo the bundle to pass into the root» of the nerves
of wbicb we arc Kpenking.n'prciBllyof tbo glo»o-phnryngenl,and the bundle
evcDtaaUy ends in front by pauing into the gl(M«o-]>haTyng«al roots. Tbe
graf nwUer from which tbvec fibres take orij^n do«s not form a defined com-
pact area, is not llrerefore ii nucleus in the sense in wbicb we are now using
tbe term, but is dilfiised among tltc rest of the gray matter along a consider-
tble length. The Hbm are nevertheless 6bres of n<?irve mots, and the bundle
is called l\w ateetuliHy roirfof (he ghMW>-|AarTiij^>al. (belerm nsoending being
used UDce it is customary to tracv such ttructun^ from below upward, that
it fton behind fbrwmrd: though »in<.'e the Hbrc4 in question are probably af-
fhnalflbre* carrying impulses backward from tlie nerves t<i the gray matter,
" d<actn<l!ng " would he the more appropriate woid. The bundle has also
Iwm called thtfiurieiiliu /ntHaritu; and, since ila [xisitinn has been supposed
|o corrapond to iliat uf the area marked out ex peri m«u tally as the respira-
torr centre ($ 362), it ha* be«-ii spoken of as the renpirrtimy hunHit.
'hM- root* of ihttc throe nerves then, the bulltar accattory, the vagus, aiid
lh«> gliNvo pharyngeal, all learing tbe surface of the brain iiluiig tlte linu
betwwii the olive and the reatltbrm body, and all do far nltku that it is
imiia«b)c upon mere ionnvction to say ntierc in the wrics the fibrea of tlie
middle nerve, thr vagtm, lie^in and end, spring from three sotirce*, the com*
bine) nnclvos, the nudt'UH ainbiguu.i, and iW ascending nxit.
$ 619. Thr fiifhih or auditory nenv. This nerve iriflerv from the Other
ner^'e# which we are now conndcring in boiog n nerve of sncdal MUse; ita
arnugements atr complicated. In a view of the bote of the 1>rain (Fig. 146.
C), tbe nerve is seen to leave the surface of the hrain from the ventral sur-
752
Am'of tlic Ion' pun of the n^iif'nrni Ixxlr lit tliu hiiiil nmi^in of the panii
two nniidfl (ir nxits. uiw ^>f whkh wimU round liit- rcelifnnn liodj mm t*
Tviti-h \t» doml Hirriicc whilo the »ther nnpi-ur' fi eink iitln the MMtUHcdT
the hull) to lh« median sid« of iIm- rMtirorm hocly ; mod in n tnosnK Mo-
tion of the bulb {Vig. lt*tt) just behind tli« imns ihr two nxrta mar hcMCo
embracing th« rcstiforni bocfy, one pominE on it« dorsal and tbe otber co ill
vcnlrnl side. The former is called ihe aormtl root ((''iK- If^), ornnKtian
tfae lal«ral root, or since it readies badi or tow«r down than ibc tilliff. the
poMerior or inferior root : tl»e latter is called lli« renlrai root (Fi^- H^'f, or
someciiues the median rw>t, or aince it reaches further forward or liigtuir op
than the other, the anterior or superior hmjC. When we cmue tn »tu'ly thr
ear ve aJmil iiati thai oa« diviaioili of lh« audiiory nerve in iliitribuud to tbt
eodil«a alone, and isoalled the nervus i-uclilcarid, the retiloflbo ntrrv hcin;-
diotrihuteil to the utricle, Mccule and aeniidrcular cauain an tlM> nrrvm v»-
iJlHilaris. As we Hhall aee. there are mauiid fur thiiikiu); that thi; vn>nhti>tr
nerve <uirri«i up to the brain fVnm the Minioircular caiiab impul-
than thorn: or h4;Hid4» th<w< whtoh give rise to aensatiouit of sound. ' <
lh« oochlcar nerve anjieimi to ht; exclutiiraly conoeniei) in bMuine ; nii'i m
tuune Btructural de4aiU ihtvn two divi*iotia of ttie auditorj nerve difler (nm
ABch other. Hence it is im|M>rtHnt to note tluit according to oarcful ini-r>U
^tion* the cochlear nvrve i* ihr cnniiuiiailoaof iheiloMU root and ihr vn
uhulor nerve the c>»itiiiiiatioo of the ventral root. Witli thu>e mots of tbt
auditor)' nerve projier iiIki iwii» little in front of tki: ventrml root, the #>mU
nerve callwi the fmrtin inlirrmftialf Wri^terfi, whieb goce to join the Ruial
i»erve.
Tli« iiuditory nucleus, lu a wbok. is a hn«d iiias]!, having in tnmveni
BeclioQS of the bulb a fan>cwhat (nanguliir form, lyiu); in the lateral piirt>
of the floor of the fourth ventricle, rvnchitis in front aomewhut l*ey»"ii t^'
level of the slriie seu)>tiua\ and overlapping Mi ind the front paru of tlir rni-
eleiis ambigiuDB and the combined acoeSBory-vago-glo«BO-pharyuKe«l aa<itut,
it extends iaieniUy some distance outside ihe former nucleus.
The nucleus, liowever. consists of two distinct parts, a mcufiiM or laav
mieieiit [,I'"'K- l^''- VIU., «!.), characleriied by the presence of iraall »«lk
and K lateral or outer nucleus (.Fig. VXi. VIII.. /.). tlie cells of whidi Ut
much larger, aofiie of ibeoi being ^-ery large. The lateral nucK-^is t* yln-t<\
aomewhal deeper than, ventral to. the median nucleus: it alM> i^xh-iids lunhn
forward (Flip. 188 and H'J. Vlll..,i), so that tlie front end of ihe nboli-
nucleua b furnished l>y the lateral nucleus alone which at its front eml <«vU'
p(ea a more durwl |)OHilii>ii than at He. hiiul end.
Moreover tlit* audilorv uucleuH thus ptaual in the floor of ibr founb
veutricte is iH)t the whole of the nucleua of the aiidiliiry iterve. At i^r
coaver^nce of the dorsal and ventral roots on the ventral Kurlitir of l^r
reMiform IxkIv it> placed a group of ivlb, forming it Nwvlliiig which In i»
genermi apuearanoe and in the cliarnutcni of it.-) criln in nut unlike i pa-
glion on the poaterior roiit of a iipinnl iiervo. Thii in calhv) the ntwmf}
nucfetM.
When we trai<e lfa« fibreii of the nerve centntlwanl into lJi« bimin, nr (in^
that a lar|;e numln'r el leaat ot lli« fibres of tlie dorwil r<Mit, cuvhim Utrvr
(Kig. 188). cud. iicoinling to rkhiI obaorven'. in Iht- cetle of the oecaMitj
nucwuN, or in iH-rvr cell" lying dorail to the accvsMiry uuvleu! iitid e»pcdtl?
in a gmup »f idln giving Hue to (he luttrrt-alum ••ru/dicnm. which, xiuall it
man, a coitSj>i<'«»iix in Mime KiiinmU. Ilencc ihv farllu-r piirl of ttisXltt*!
root, as It U'lndM rouiKl ihc IiiK^rnl iind dorvnl ^urTacu of tlie rwlifonu haif,
coiuiiiUi lar^gely, if not wliolly, of 6bre8 which are ilcrived nut diiwily At*
the trunk of tlie iverve, hut indirectly ihrtMigh ihe relay nf the aciawr?
■MHW orHTwitT i'ellrt. Kviii;)iiii)c tbi- ilnrul surfaoe cif (he ratUbrm bodyi
lliCM Abrvt u|i[ir«r on tliu tlunr or Uie futirtli reuUricIf! m tbe ttrim antluet
I Pig. I^ti, «lr. t, Hiiil vad pnnly in tiiir iii«^)t»ii uucleuv, |iurtly in otbrr rcgiotw
•li (br hiilb. Thi* vsiu-t iJtU'rniiiiiilioii, )ll>n(.'Vl^^, ol' ibv vniliiifjs nf (iiU root
I' It.. lUl
4M<
VII q
./
le^Jtv*.**
>«(i
I
I
IWUCOH Tim iitMMr fiar or nii rm'iiTH VumMij'. 'SutgaMimvx.)
TMkvn In ibc Uiic III. Hc- lU
ftMMninU>Wnbi*irulmn»niv))-. fr, />.. IbP niiMnlaliil mnlnll liBuvKn llbiB M (he |uiM.
TktilM'lMl |iM(»rilM |iin*'i>'. rilndtoiCcBiiajaMlw mlnclalwIUi Iho itecper Itaintene nbmi.
''..U* ni)r«. 7)>.. Iliu intciltHa: c X., Uw i«iUII>ni bodf orUArlw i«diing;»<tf ihv oMvWItiUB,
•ul ••HI* iiUhurlir i K ^.. lk« •U|«rlui |i*ilunr1r«->riliDMrabutl«in : r., rapbi;; i. a.mtwriacoUw.
I* il.cMfaiil*Miil(iinafth*»Rlwllnni. ^.a^ihc nucloui of Ibe (aaf : a. |i^ (nberchoflRoIuida ;
f. X,.*MHau thjaimti nlciia in ih* TennUMi|«rlor of ilw (anbrilnm : l. Iniiulhi fttna ilwi>llr*U
ItottMlnlar iiucln*: Vlll..iii>(4fbUiatMHllt«yDm«.ltirmivl-i>imill>nlMraui. pniMciMnK
AOB Vlll ,i. Uic(iwii[iui-iribv liteial BOiIlkirr Dotlcai : Vtt. n . Uw duoIbiu «r (ba MitMllilor
(MUl iivm , VI,. lb* MMkaiaf tbo ititli oerw ; VII u., Illina of lb* wrvuih nnnv cut VRMa*
ito|r ta«i-'"<"<-llbr audgDn^lhttlilbbvlbivlHuliMf (mm Uw|nw«* VII,^ Mb. Utr knirth reb'
nWk. b«l« rooM in !•) Ihc rvrateltam : Ihv iluutliiK nt ih* CHirnl fta^ M«IMf ilniD«>tUl*>r <«r-
tiMMatluf IkcivntrlrV [m. »<r ib« Mktuf iIiutJipKy, •imlRid.
>• « niMter uf coDHideroble dtllioully : »ura« obttrvtn rv^rd tlie aceOKwy
Dudout Ha l»(>iu«li<f^iuf, nut witb tlit: G«B«riwi and iriih ihe tpiual ]||^nglin,
but with tbe (itlitT. im«, vrantal nucid ; anil lu uiy cum we must probiibly
coDuder thv luwliwu iliviiiion ul' llii- auditory nucieui, not u a nucleii* io
tbe wtBM! ill wbich we aiv now iisiiiy il, liul rather u m MOOBdAiy oonnectioii
within tbe )>ulb.
VVIh-u <m tnuv tlio ventral n>oi, vealibuUr B*rv« (Fig. 189), inward, w*
-IB
7ol
TRE BRAtir.
lind ibnt RUikiiii;. acordEog to inoel nWrvere. nu nmueclldos bI dII nith
llie aceeworr nucleus, it pggaes (Fi;;. I'^l*. viii.) lo ihe intdUn Nr)« of ih*
rcMifbnn bmh-. belween it and tlie asoendin;; root of t)i« fifth u«rt*, sod w
resobee Die lateral division ot' llie nucleus, in the Inrj^e i-elltt of wfairb niaM
at least of its Bbn» arv eaid to eud, and which, ilierefnrv. niav be regarded
ax the nucleus of the ventral roiU. Un this (loint, hou'ever. all autl»ur» an
not neretil. Tlte lateral auditory nucleuM, with the Bbre» proc«ei]iu(: to and
tn>m It, lying tut they d<i lo the median or inner tit<b of the niitifonn biMiy
|>roi>cr, are aumeiiineti spolcen of ax the iniMlinn or inner diviiiiin of tbr
irslifDrm hmly ; and fnmi the nijclciiii a. consideriilite numhcr cif fibnv \imm
lip with Ihv rextifiirm hiHly into Um cerebellum n* a conlinuatiou of tliia
'* median division of llir rrstUorm Iwdy," Some author* maintain thai thiM
8bres aro continued •imiglu on frtmi the nerve to tht- •.vrclwlliim: but thr
more r..-c«nt invrntintiom M«ni to show that ihcy nil make conon-tions with
tlie nervocelU nf the lateral niicIruK oti their way. TheBC fibres oon'litute
B connection botwc>cn the auditory 'vt^libultir) iien-e and the ccrubelluai.
the phyeiolugical *igTiilicKnce of which we 'hall f«e Inter on; we inny. per-
haps, coDipare it to the connection between the poeterior mots of the §pinal
nerves and the cerebellum through (thfl vewcutar cylioder andj the cere-
bellar tract.
Tlie other central connectiuna of the tateral nueletts are. like tboae of the
■cceasory and of the median nucleus, complicated aitd ohHciirv. Hut we
may call atteutiuu to a tet of fibres which, uartioK apnarentlv in the nc<«i-
Rory nucleuK, run directly Iranaverae in tlte ventral nseion (if the tegnientuai
}ntt doiwil bi the iranvenc Uhreti of the [mott, furmmg what u allied the
lr»i.'tiiim (Fig. 189. Tp.).
Lottly, we may add that the fibrea of the peculiar p«rti<t inlermrdia appear
to lake origin from the iico«Mory nucltiH.
5 680. Thf ttvrttA or fitiaf ngfoe. The nucleus (Fig. W-i. VII.. and
Fig*. IJSS. 1W». VII , n.), of this nerve (it being borne in mind that the
mi>tor libres for the orbital region (the corbicular muscle, etc.). though they
run in the trunk of tbb nerve, really belong to tli« thini nerve and lake
»rit;ii) fri)iu the hind part of the nucleus of the third ni-rvc) narroirer Ea
front ihnn hehinil. reaches from the level of the btri^t acUHlicie toroe dlatancc
int'i the regiuii of the pons, and occu|nc9 in the midst of the reticular forma-
tii>», a little doraal of the patch of gray mutter culled tlie upper olive, a
pnfilion cinrres[i'tndiug cIo«ely to that of the nudetin amhi)^ii«. Tlie cell*
of the nucleiin are large, and ixBtiecu well-marked axii^cylintlcr pmcettr*,
which are gathered up at the ilnrsal nurfara of the niicl»iis to form the root.
This, rising up diirenlly. d<.-»cribe« a loop (Fig. Iftfl, VII., y.) n>und the
nucleati of tlie nixth or abdunnit nerve, running forwmrd fur tame little dis-
tance dumi to that nucleuit, and then descends again vcnirally. lousing to
Ibc lau-ral eiile of it^ own niidfUi', between it mid the ascending nwt of the
tifth ( In.) ; it thus gains ilir niirface of the brain at the hinder margin of
the |H>iis. lateral to the uhdurens. ophite the front end nf tl>e groove Ite-
twei-n the olivarv hiKly and the resliform body. As ii thus encircles the
nucleus of the nliducens, it looks as if it were receiving fibres from tliai
body ; but the evidenoe gow lo show that thmo fibres limply pus through
the nucletts, and do aot lue urigin from any of its cells.
$ 621. The tirtliorabtiueeiu tirree. This nerve starts from n compact oval
nucleus (Fig- 193, VI.). lying at the level of tlie hinder jiart of th» pun*.
and therefore of the front pert of the fourth ventricle, in the oentral gray
matter of the floor >if the ventricle, or rather juet between it and the reticalor
formation, a little on one aide of the median line (Fig. 181), VI.). A flight
~ tlic llnor uf the fourth ventricle, emineuiia tcroa, mark* its posi-
THE QRAV MATTKK.
TS8
I Fin- 11)3. «>.(.)■ Til* nucleus «oni«iiw fairly lar^e nervo-oolU, with
„nct axU-cyliiiJer proceMM. TheM are gathered at the nieiliati tUh of
nurli-uN to' form the tliin ruot. whi<.-h piL»ii))c ventrally aii<t litlvriilly. at
■UDT little ilisiaui-e ffoin tiie nioilinii niphc, llir»U){h ihe retictilar forniaiion.
niiM baiilcKnnl above the [lyrnitii'lil l)iindl«» of the pou*, and tiiinllv roniM
to tkp BurrnM.' at thd bliiikT edje of tlic poiu, upposlto the front em) of ihu
pynuiil (Fig. I8B, C).
nu. M.
via.
*•
>th
V*
Va
THiwiHiii ni( ivm «T na Xttt or tiik rimi Naarc (HiiRaauKina |
lnilicnn>' l)£ ric. tn.
C. UnltKtImttntUlona lodr . K K. •aiMrtflriwliiM-iaurilwouBIitlluiD. f. dl. niBtlsn-. f. I.
Uln«l MM, r. R.. MsmwiUl nilrular *L«mitli>M. %i. e„ raptaOcM imimnaabi*! uTtlui poai^
l_ iDilenui k«NCtni4l«Ml bundla; V. *.. auiortai Trriiili iKUQcmt of lliiw Mte aiv ihOWM. one
twtna dslarb^ : bcoinai ihria uic iiibrtfriiinf tiilel Ul>l o^rn r>r ibr •Mikn u* mn) : VW^ nlic
nl VliBiiMU « itMerlur nlum^ i.. niplit: ly.. i-rnniliUI alina; tr. P. aiar —Mai o4 lbs |»ni
*.a,«i|Mta«atlTaL t, itewil on am Irft •■■I* liidMalot Uw |i«iltloii of* taodle of lonftiwliiul
atom vkUli rutr l» UM«4 tomfd tiiu> ite •uMhaluBlo *a(k«wL V. w., aout imeteai V •..•«
wry nadMH', •*> V,. notaoriacnnhiwcvc. Iih. iMnli raiMck : ilwdliutefnuiMlin) nuiwi
oBlual M IB nc- ">■
$ 628. TheMh or rri^tuniiia/ nenw. Tbia narro, aa it oooMa to tlw aurfaoe
oo Uw vanUvT aspect of th« poii* (Fig. 186, C), near tho IVoqi odgv, at
aona diauoee from the nMliaii litm. cuniuta of two purta, a anuller motor
not UmI a lw!K«r Mdtaorv ruot, the Ull«r boaring Ihe lar);^ [^ii);lio<i of
Gaawr; and tfaa orlglu of (he uerv« U in ntan^ way« cnrnplex. Uijih roou
nutj bo traced la ao »hlti)(M diraction (Fig. 100, V.^ innanJ and toward the
doiMl urfaoa. through the |Miu to the raiicular tornuliou beosaih the door
73ft
THS VBATN.
of the froDt part of ihr foimli vcntrk-le, tlie •maltcr mouir root ukiii^ u|< a
po«iitioii median to tbe larger tfcnsory root.
Here the motor root conice into cunnertion with n ixillwiion nf iwrtiMxJb
(Fi|p». 1{I3 and 190. V. m.), whidi may ho rmrdrd ns iu nudrut; liut (bit
is not tlie wliole nucleus of tbe motor root. From tliv level of tbo oaiitm
there atreicbes forward as far as tbe level of Ih? ■n(i*rior cwrpon wwki-
gemtiitt u hiiudle of lonicitudinal libres wbicb. »inoe it is usually tnwM ftoni
the front biickwartl iiutil it nasepa iolo the root of the Derve. b fpnkni of
H Ihf dr"-rniHn<j mot of tb* tifth nerve.
Tbi» di'^ci'iidinjit (xMit bfj^ns ua a few scattered bundles of filirw u ik
Icvi'I of llii- anterior cnrji'ira i(uadrigemiua. in tbe perifibcrul lattnl pui
of the ci-iiti'Hl gmy miilli-r ttiirroiiDainjc tbe a(|iiei|iiet, dnrwil, and bunl
(Fig. 19'i. V, (f 1. io \\\c nucleus of the third nerve (Fi((. 19'.?. IN,,*,
From Ibcncv the fibri-s |>ims liarkuai-d, aujcmentiDfc in number, am]
' ' " iMeralu
increMinf b
form n compact bundle, fcmiliiniir in Ininxverwe lectioti, Ifini; Interal totb
fourth nerve na this i» rising dontatly I Fig. 19!, V. d.) ; atilT ii
Fk. 191.
■»:■
■^c?-
.^^^ii&SS'
Tiinuihii n« F'oNB.pARior Tiiii l<am, (MtRKDfarMi.i In ilMlliiPtO. Pif. iim.
IV. pTTBDiMal llhmt f- ''■ Obrw Ihim tlM Itaailil ohms: X I*, nipcffiir |vil>iiic1e .if Uw nip
IVilumi rm. median pratian. f lalenl pcinloa «( tkc lllrt : '- piwwrfor bmciliirUiinl >iwP> ,
P. C </■ lawtior ooiiiuis iiaNlrtNanliw ; is fibiw wlilrb lnwoiw dplwhri ftum lh0 Blltt. niid riiRlH'
'omrd tiitin iiht tnrwnmali jan ol ilm pm nf ilw nn» i. r. tooai acnilc'w . •■■ P. Q. iiut*>in «t
ItiopoMUlot CDTp(iai|Mdi1SEinlDB-ihvcidillB*Uui«4CMniiIiitni; IV. biiii-lloor Ibr tourtli imvt
dMiMMttnir. n'. ■. lunuclMiii V.'i, ricH«aillni iwH or itM «nhi»n»; ^v, ihr oiiiwObri : i-.f im
rnSon of n-ii(nl cnjr tuaim.
number Jn thtir CKirie tnckwanl, (her (^dually aasume ft more ventral
poeitioD ii> ibo ttaunluot oprn* into tlie fourth ventricle. All mlong iit
I'oune thii (ItacruniDg root ha« altacbed to it lari^e (70 •• or more in duuu-
eter), spuie apberMdal nerrc-celle of etrikinr appearance; these, Uomtrtti,
•eonloo few to givu origin to nt least all tne fibres, nitd there mn> wqm
nsMOS for conuccliDg this root with the collection of f^ray matter itdM
"locw cwnileuE." (Fig. 191. (. *■)
We mar probably regard this deac^-nding root aa bdoneiug to tbo niotoi
division of the nerve; but it ia Mated thai nianv of the nhree of tlib ru»l
TUB ORAV MATTBB.
inlo ths motorj Tvnt, 4-v(iiititiilly flniliiig their wi\}-, MCOrdInx to wiiie
ohMrvcn>. into iht oplilhftlinic kraiich,
Tlw »ri«ory rool mar be timiUrly irjiciil into it uiiclciiii, the sonitory
DUclciM I Fi^ lO.^ aifl iW, V. t.) lying laU-Titl Ui [bo motor iiucli^u*, niiil
CoaiMoU*) with this U ihv slrikiii); tract ol' Hbre*. to which nlrniiilv wt- hnve
' lOftcqtwittlTnltiKied, and wliich u ralloiJ the 'Meriii/ifitf roof of thu I^IVh nerve.
This m»o.-n<)iii(,' root be|;iiiB as n buDcllc or bittiHlu of few fibres which
■ar be tnicctl Iwckwan) as far as at tcadt th« level of tlie Mcoiid cervical
aervt, and it cooa oouapjouous iu Irangveme aections (Fig- 147 «t t^., V. a.)
m ftMmiliiuiir patch or white luatler furioia^ a sort of cap on the oulsidft of
the Bwollcii caput of the posteiiur Iiorii, between thia etnicturc and the lon-
gitudinal fibres which are beiicinnins to form the rcetifonn body on the
Fia.tSt.
\
TiUHiiva ma Can am AiTKiiinB ciiM«vi«t iji'uiwamD't. (SniSRaHnew.)
OBv-hniroiilr Ixbotiii.kiitbsUuall*. 1^. IM,
^ UN pfmKtdal innlcm nr Ibc |m. » Ihv nclu'i »< 111* t«oc«u|4Hl bf libra (rnoi ibeffDiiUl
fatitmt aCihttvntx. fr o. the kkiuo uoea|4cil bj nlma mmluit (rem itoocclplial lotUonoT iba
MHU ; r Mm* cKunlna trim Uit lUHt -. 'tr. tba opuc im« ; IF. tbe IIIlM, I. U» lu«m pMlMi. m. tb*
i<1lsa lunti : I. Uw iiatcrto* lungHiidliiBl btiadic : it> a. (be bmolilQtn nf iiic uiierlar CDrpoi
i|a»ili1a—lliiiw ; t. lllmi (ram Ite imlcrlur aiinaiMU* of Ibe cvNtmiu . r. n)0>* x ^ •hWuhiIa
■Itn ^ jr. a. nd (iiifImu ; C p, t Lilmil. kih) C y. <l nwdbia wrpiu (nilculMam . fn. pulTlnu of
iifrili Ihalaniu: A. V ■- iiu«li>ui'>r Kn; niallertrf knlivtur Coriiiu ■)■*'■ ilStBiluuin : 111. n, Burloiu
ui III. HUM acrra: III.' luoiMi rmiii ilisdifHiI lanotlll. ■..tb« nocItuiorilM thin) nan*. wUcll
era* lb* acdUa Una la miBivr witb mMleli ilerlrttl Ihm Uie nurlcia of Ibe opialie ilda : i. ml
nqartdBl lar« oT abfa of lb* ■nLmri'. quail.: d.*i.4ee|t layers Ji%^'aa&atA tumniiidvl by
«»bnl fnr awnu.
mrftce. pMuog upward, and oonllnuBlIy nugownling in bulk, the root
elingi, M it were, to \Xw g^laiinouN mbstnaco of tbe canut of the posterior
horn, and nnk* nilh it inwnrilly iind vcDtmliy as tliis becoinca eovered up
fint by the nstiforni timly mid Kub«o|ii(-(>(ly br the iauiiag trunk nf tbe
EMt nighth or auditory 'ocrTc (Pig». 19^, l(5d>. Panins still fi>rt<ard,
jrond the dtMpiN-nriDg freUtinous xubatance, tbe root, ■ttll growiuj; larger
ium) divid<id into wwral distinct biindle». runs into the reticular funnation of
tlie {Hitii aiKl, It-aching (he level of the sensory uudetis, suddenly bend*
rouod nod joini> the vrnsory root.
Tbi* iMoeDdiiitt root dillors from the deK-endinj; root in not having ooii-
•picuouvly ntiached tn it any collection of Derve-cvlla: iii thia nvpool it
rmecDbles the nscctitling root of the gtoeso-pharyiij^iil. dihI »e tii«y add purl
760
THE 8RAIX.
w* ^lall »ee later on, ibit> uerve b now exdu«iv«l>' efthmit, wluiivvrr U nii
have been iu imire jiriiMitive beingtt. W«!<linll alaoMie lnr>>r mi, ihfti im;>'>''
iliartiug rr»in llie4.-<!ret)nnii(if <io«i<ii)e pttm to ltieo<Tve<>f ihruthor >i<l- i
ii to Mty d«Gtuwiis; and ihin UbImo tli« auw u-tth th« tiihor rflV-mii >> < - >
iicrvuH. Tht Sbrei irhioh appear to take nrigin frum tlie hrrvv-tvlU <it ;'
niideui do not crow over afur emerging frurii tlM,- iiucku", but kwp t<- ili>
MUM lida : thore U no dwtnnt il^^iuwatinn w in the oiiM> jui>t notM of lir
fittirth norvo. Thcr*- an;, Imwcvcr, lilnv* :Fig. Ifl'i. III.') whicli laaviag tin
uuL'liHi* crmu tbc nKfliim raphe from oih- side In Hir "thvr, un<l thetf pcMibli
are iIkt imths for the <IccuMiilioii of ibc imputare : hut thf>r may ur fibm
paasinc Irom thv crtiK acrow th« raphe to the iiiK'leos. 'this iiervr ku
■peci«r relations irilh the optic trad, but of tlnete nv slinll sfMult wli«a at
OODC to deal with the fuitcttotiJ^ of tbe nerv«s.
$ 82ft. In attempting to uixlen^lnnd the nal(ir« and relations of these craoU
nervee, it must be borne iu mind that, while niorpbolofjical studiea lead uilD
beliere that, as the vertebrate body baa been developed out of an iiiTenebntk
ancestry, so the brain of tbe vert«brat« has arisen by a series of luodirmtiin
from Ine n«rvouB Btnicturea placed at the head and arouiut tli4! iiiuatli uf vi
Invertebrate, tbe same atudiee teach U6 that auch an evuluiioo liu bMi
a<N.-omp) talked by uiettiu of profound changes. We haw. for innianre, na«>ii
to thiitk that the mouth of the vertebrate does not correspond to tha mouih
of tbe iiiverlebrale, but ii a new sLructure, wbi»c aj>pearanoe luu been acoooi-
panted by a conHblenible dulocation of parts. ^V« niual a«-oiirdinflT expert
ti> fmil tht! iiidicali'>na of a neKnienlal urraii^-rovnl grMtly otwuml iw I^
one haix) by traiixiMMittiin, aiHt iHt tbe otiwr by fu>ioD.
The twdnh iir hy|»i>|^lnMul ucrvu i* one vh<i*v nalure vritms fairly *impl*-
It ii in function t-xchisiri'ly iiu rlfi^rvnt iMirvr. TIm- Inrgi? ■'•^llx, with iva-
Kpicuous axbicylindiT proccwtTi, which characttrriM ita nui'leun. an* i-xacilj
hko tliiiw- af thf anterior hi)ni of tlw spinal aird which give origin to ihr
tibrvn of an niitt-rior root. Tli^ niirlciM, worsorcr, in its iMisilion curre^pco)^
tu juiri "f iIm' anterior horn of tlie uhiuiI cord, if wv laVe intu iM-ciRint Ibr
shilling involved iu the decumation of the pyramids, and in the new der»l(^
uiflnts of the Itulb, If wo conipara Pig. 1^7 with any Eoction of the oord, at
aan that tin- liyjMigliMHil nervo cnrraipoods to an anterior ruot of ibe t^ioal
eon\. but that the fibtt», atlcr leaving lti« celt* from whidi they lalie llwir
oritpii. IraviTW in the former a large tract, aud iu the latter case a unall Uae*
of ti<aue. Whether tbs whole nerve corresponds to the tibrm of seventl ftf *
vanati- fusMl lugether, or to tboae of one eegmeoi spread out longUurtinally, a
for our present purpoaes of secondary importaiice.
Itecoipiizing the bypoglosaal nerve aa toe hooioloKiie of a spinal uiteri»r
root, we may go on to claim ilie nuclei of the third and fourth uerviv f
similar groups of cells of the auterior horn, giving rise to atiietijij n**"
Tlie ]>oflilion of (he nuclei, the character of tbe cola, the fiincti<ia <•( thr
fibres, all support this vieir. The caw is iwrha|ie no4 ao clear as tliai V il/r
hypogloesal uerve. since there are reasons for thinking thai thaw ncrv« \it*<
unoergoDeiu the course of evolution greater changes tlian ban the hypn|leMl
nerve; still llieM reaMinii do not opfmae lite abo\'e cnrjunion.
The Ducteuv iif tbe exclusively motor sixth nerve does not exactly i^in*-
sjMind to thua« of the third and fourth in iMsitlon ; but wc may probsblr
place it in till- »ame serieti with iht-m. Thus vre Itave in mics-vmiiun. tbittkinl
lourtJi. sixth, and tweltlU nerves, with their respective nui'li^i. as the aalcr»r
rootA of iicrveo of tlicir sevdrs) segments.
Ill the filYh ixrvc, tbe dislocmtion and fiuioD spoken of above has latr>-
duced ditHi-ulticM. The motor nucleus, with the fibres of ibc motor rout i"
whi<Jj it givw origin, han by some be*u oiMuadorcd n* humohifivius u il't
UV MATTKB"
rtxit >kf nil onlinarv spiiial nerve, the fibres of which, as we
iiilii the amy miiiler tviihaiit IteUtg obviouBlv connecteil with
Iti ilE loner ivart nt Icael iL cuiihIhis orextreiuefv fiue flbret, atid
-• vcrjr much like n aiiiliuuHiiuii io ihe bulb of the margiOBl
-i> t *) xonc uf ibe f'piiiul curd.
3, Thg/ourUi or froM/f'ii- wtriy. The duc1«us of thin uerve (Fik. 1&3,
• ■ wliiiim uf aoiiiewhat Inrce luulliiHilar cells ou riiuh *iile <>f ihu
lln« lielow lh« ui(iieilLirt (ti^'. IIU, IV. ».), renc-hiii); t'roai the level
inction i>r ilic anierior nnit [lutiterior corpora quntirigeiuiDa to the
(v«l 111" the iBltcr body,
I, einrtiii)^ IWim the hiteral niirfttct- uf the nucleus, <lo(« not titkc lit
airnl iliroctiini, but sweep* luierullv and dontiLllj: in the outer Inycrv
ntral gray niiillcr (Pig. liH I, aDil »> curving round to tlie dorwl
Itraclm th« valve of V'icuMcn*. where in the mctlinn line it tteCU»aic«
tfrlhtw in Ihe ttilisliincc of the valve ; ttuch a (1ocii»»i!od at a ilietitDci;
> nucleus ofori^^n is except ioiinl in iho cranial uci\t». I.c«viug the
see of the brnin in ihe valve, il takes n snperticiid coureo curving ( Fig.
it) hilmlly and ventrally, and ninkc's ils appearance in a ventral view
lie brain al ll>e frciQl ed^c of the pous, on ibc Inlerol edge of iIm.' crus
J AM. /'Ar Ibird or oculx^nuUir urrvf.. The Ducleua of this nerve (n|j.
^'1. III., \^'J., III. II. ) is a culiimu of, for the oiwii part, fairly large ntulli-
nlar calls Iiinj^ «u each sid« clime tu the luedlnn line, iu the gray matter of
'' irrairal ruial, just durwil to a bundle of fibres nliicli wi- sJiall t|>eak of aa
l<Migitudinal puot^rvor hiinillv; it reaches from ihe level of the pcnterKM-
nmiabwre tn iIk- third TcnlricJn to the luv«l of the junciion of the anterior
■wl posterior corpora i^uftdrigeiniiia. In a wcliCMi laken ihroui^h its middle
I Fig. VJi) tli« nucleus t* wrn Io give iitT fibres whirJi run verlicitlly Uiwaitl
iha ttatnl nirlacc, Iravcning the tegmentiini and n body ( fin.) irhich vrv
•lull pfMCiitly i>|>e«k uf as tho " red nucleus," bul appnrvDtly nuking do
■nections with these stnictiires, and pierce the median edge of the pes,
trging (Fig. 186, L') on the surface to the median aide of each crus. As
nchi 4i1«, tb* tiirii'n MrlMam ■Dd optio itMkiaiu Hai* brai cut swajr h(n4»nlall)r lovnnvBUh)
■IflSh IU ot>li'r Eu tbnw lliMt lalnml iljveniiw. L. lalenil. E. P. rxitnutX I'mlritm. ur>l II, t: nH"
4lui ivMonor mAddio or Uw ear<l : /. /*. Infnrtoc lodiincli. S. P. ni|itrlur iitduriule. osd P. BiMilte
rarfuacif «r ilrtcnvUltiini.nllcBiaonM. Tho doiMd cnrrM tln«>.iip)«iwidloi(«r, unibv Hibt
bairaf [hi>aKni«ini>iii<'i> tliBifctied Bner.t'. outddcUirnRnreiuisu^asft Ikseppsr siul liivtr
hiaiMlmisiif Iheioaaua Ikstntimi )»|im. ThvoiilUua iirdinlhimli nauf^ltabown tifa liwM
iU«k IIb*. Ill la* IIm* et Ihe nnltlclc ore tit/wu. on llii! nsTil half: ffi. Kit** poMv>li''r; n.
rH^nnwa byi<e(lt«ai : r or. irtcoBuni aoiiUi'iuB : 1. 1. •mlorail* MRa : i. •■. ilflB aelMlsiw on
4nMt(iB : /. o. tmm sainlia: i.r. \nc\a iimiltiwi t g. nlr* of VIbuhkbi; Q^xmuit\ot unl <]n.
■Bi*ili,T(iirvu«i|iM<lit(*aunniD: /U- |4u(al(l*nili AV. UMuntUncof tbare4iiucloi»: i, Uiu UiM
naliMa, In whli-h i' \o»irMi.v (he iindilk iii koR MUUOilNnn: flp.o. llM |>Ulai« of Uiv fofalt.
teblu>l wWr)il>In-tl«tpl In IhcnutiTDrihcllilnl it«aM*tallisl»llo«a(lb«lnfiioiUbal«im (' Cf.
IksfMniof ikrnriuaeallouni. IwtHMn wbioh and ibo 6>mu IbcntKraAcnalMihsnAli tcd.
MWtltlatflated: r. partliinolrarmliill'npr ftonislhtailqAanouiMraak On Ois 1*fl iMs an
ii« ■ II . C&nflui unaliun : O- r.o|i(trihalani» : rr.ivlirinu: 7.«LRibm«IiiiaanMtliK: rt.j.
HiraaMal wiinu miking lh« plan of neiviliit, of lbs cliuiiailul iJins. On Uit itKbl (lilasivaiiiairi :
S. C. Iuad«f. at, end of tall of nuikOf nuilnlu> : 0|>'. (^p~ihclini|art*af tbcgMHiaiallldar. snl
n. Biitswin f iti* niii'kiw ImilnUrli -, .v. a. onieriui biipIhu . .V wed. intdiui niKlaia. -V bii. bustsl
•«(lfaa.aa4 l\'. |iul<lMr>>r ibeoi«li>tlwiMn<i>: Cia, ftmit llnib, <V- <">** '■>' paOi f>p.biad tinb
at Mamal raiaole: O. evlonsl oa|*>la: f- elauMnis Ttt DunMrab III. b> XII. Indtcal* Ills
Mtlu «( Ux iBp»un onnlsl mrwt. alt tfaowa nn ibo Wfl tble wlUi itw cteapdea of Uw aciiipiy
i^aih^i ibiriiriiil tX., X.. XI„ nhich «> atotil nahulnn la plarvd lai lb* rifbi tblB. I'. ii Uw
■MM nadaiw at Ik* nna avrn otib ilig dGacoHllng no. r. a. tbe wtiKXT anvlsat of iW •smv
alt'- ■>— >—-.■ -•-■.-'II' - "vK, VIM n, nivllaa nui'ltu*. Vlll. I. latoal andnis of Ikt ■IkIUoIt
n,' iiiaL TM atlvnilliiH nvt oftba ntnOi asm la nvn si llwWiHttndal'
■■ .\.X.XI.
■60
TirK BRAIS.
ire shall see later on, ihis nerve is uovr exolusiTeljT efferent, nluitev^ il mm
Imvebeen in ntvre primilive beiii^. Wesball also see later ixi, Itial iin|iul<r>
Marling flxiiD the cerebrum uf one side pose to (lie nerre ur ilie ntlier Aiilr. ihni
ia to say decussate ; and litis is alsi> the ou»e with the other eflen-nt cniniu
nen'cs. Tlie fibres which B»j>e»r in iiikf origin from Ihe nerre-celUor tb
nuoleuD ilii not croe over after enierj^iuK from Lh« nui?l«u«, but keep t' ■!»
f»mc ^litle ; tlien^ is no distant deens»ation iw in the cane just nuUn oT ihi
fourth niTvc. There aw, however, libmi ^Fig. lil'2, 111.') which Icnviiii; il^
nuckui' crnM the tni'ilisn ru)))if from oiii^ side to Lliv uthe-r, and ihiw (Msibli
aro the iiiithi> for thf d<'[:tiiw:itii(ii rif (lie impuliiesi : b«it llwv ninv tir likm
pasring irom the critK ncmm tho niplio to Um' iiuckus. I'liw nam Iw
gpecinl rplnlion* with the 'iptii- tmct, but of tli<»e we shall «penlc nhm ■>
come to dcjil with tht' function" of the iicrvM,
§6Sfi. In nttoinpling t<^> iindcrvtnnd ihenntureudrvlaliansof tlicwtnaiil
nerves, it musi bo borne in mind llinl, while Riorpholn^cal Mudtos lead lu ii
believe that. Zf the vcrtohratc t>iiily hiis been developed out of iin invcrtebnu
ancestrVi so the brain of the vertebralc has arisen by a series of niixli&imiiani
tcom the oervoue structures placed nl the head and around the muulh uf u
inverlebrnto, the fame studies teueli Uf that such an evolution has bwi
ac(.'oiopliahed by idi.>hdb of profound ehauKes. We have, for instance, nturn
to think lliai the mouth of the vertebrate does not correspond lo the ntoodi
of the invertebrate, but ia a new structure, whow^ ajipearaiice has been aomn-
panied by a considerable dislucnlion of parts. ^Ve must accordingly espn
to lln<l tlie indications >tf a eeKmenlul arranKenicot greatly nbavund ootlif
«ne hand tiy transpusitiou, and on llie other by Alston.
The iweltlh or hy)>ogluMal nerve ia one whoaa natun; oecju* fairly Minplt
It 19 in function exclusively a)i elferejit nerve. Tlie large oclhi. with an)-
nucuous axis-rylindor prooCMC*, which charaeterin' ii« nucleus, areenctl*
Ilka tbow of the anterior horn of the i>piiial cnrd which give origin tollv
fibres of an anterior rout. The nucleus, moreover, in its iia«iliun corropMiib
to part of the anterior horn of the Mpinal cord, if we tnlic into ai'oounl ite
shifting involved in the decussation <>l the pyrumide. and in the new develop!
ments of (he bulb. If we compare Fig. 1>S7 with anysMtion of the cnid.vt
sec that the hyiwglnssal nerve correspi>nds to an anterior root of the spioal
oonl, but that the Kbree, afler leaving the celU from whidi they takeibtn
nngin. traverse in the former a large tract, and in the latl«r case a small ttatl
oftissue. Whether the whole nerve corresponds lo the fibres of several scg-
meniF fused together, or to those of one segment spread out loajptudinally, «
for our present purposes of secondary importance.
Recognizing the hypogloenl nerve aa the bomolofrue ot a spinal antenor
root, WL- may go on to claim the nudd of the third ao<l fourth nerna at
^niilar ki^'1>|'> "f cells of the anterior horn, giving rise to anterior root*
The {luaitiun ot the nuclei. Ihe character of the cells, the fnnctiun of llw
fibres, all aupport thia view. The cade b perhaps not t»i> clear as that of tlir
bypogloanl nerve, since there ore reasons for thinkiuK thai these ner\-cF iMn
untlergone in the eourae of evolution greater changea Utan has the hypogUMal
nerve; siill these reaiiona dn not oppose the above conclusion.
The nucleus of the t-.xcliixively motor sixth nerve doea not exactly ooitt-
vpviid to thcMe of the thini and fourth in pwitioo; hut we niity pndialilr
place it in the tnmc «rri(.i> with them. Thus wo luivc iii auoccwtnn. the thin
itllirtb, sixth, and twcltlh nrrv», with their rutpcclive nuclei, a* lliv antorior
root* of nerviw of their acveral aegmonU.
In tlic fifth nerve, the di*locaUon ami fueioD spoken of above has iatit^
duom) diSiculti'v. The motor nucleus, with the fihn* of the motor rout i'>
which it giveaurigiu, has by some lieen conndered m homologviiis to
OltAV UATTBB.
rBi
jii»l <lo»PtibMl ; bill it is at onc« AbvioUB that we cuniiut look u[)od tliU
lil\ti nerve as correitpouding to one H(>i»al uerve, niili kit ulterior ami
t>u«teri<<r nol, mat m tlie Buper6ci)il resembluwe Beenu to be. The fenUirts
• •kf the TeliMirksme oteeading root forbiij this. The Rhrcaof tins root taav be
tnoed hnok, w> we bav-e Mid. to the very beKiiiuin^ of the bulb, and. iiKleeil.
Into (he ipinal cunl Iwyood ; as (tt as can lie aHL-oriaiued, ibey are not iu ild
ubviiiua iu»l direct inanDer L-oiiiieoled with ii«rv«-eells alou^ their couru;
but the bundle »r fibre* diu^pt. as wl- have seieu, to the fcelatinoii!! aiilMtauoe
uf the iKMterior horn of the npinal cunl and to the cnntiuuation nf thin along
lb* b«ilb, mad Ihe Ghm are lost in thi« sinirture. Tli« rout, tlicreiori^ a« «re
hiTO Midi, oonTHMiKls very cliiwly to |wrt at leuvt of the pwlcrior root of a
ipiaal nerre. and, Iboiigh ine matter h«* not yet bven experinivatnllr proved,
>ro may infrr that the tniphir cnntrtv 'if tliew Gbrcv w lo Iw found in tli«
pdk of'^ the Qanerian ganglion.
B«it if iheascMiding root b« of tlw naitirv of a pu)l«rior root (and we may
facMeotilly remark tnnt the term asoendiDZ Uiu> been unhappily chmwtn.
phioe, if it be an alfetvni mot, the direction of the impiiUt:* whiL-h it '-arriw
irill be a <)<WM>n(ling <>mc, iinniely, from the entrance in the pons unriiftl the
hin<liT partiif, we ran hardly siippi.ei' that it btiloii^ to a single i>egroriil, or
tthe L-ompleaiNtt of the motor root alone; in it, most probably, the poi^-
rior libf«6 of several fie);meul# are blended together. Further, we may
perbiipe inter that the other libree of (he Ben«ory root which end directly in
iVhat ire have culled the Eensory nucleus, are in nature quite distinct from
the tibree of ilte ascending root : and if so, diffiotiltiea arise oa to the nature
«ik1 l>omolo|[i«i of the nucleus in question. These, however, we must not
IdlaouM here, Dor can we enter into the question of the nature of the deecend-
ting root, conc«ruing the fibres of which, as we have said, auth»riti« dill^r as
|lo whether they jaw into the niolof or wtuory root. We have said enough
llo ilinTr that tlii* lifth nerve in extremely complex, ami that ilv apparent con*
formity tn a «implc spinal nerve \* in reality mialeading.
The tibrea of the vagu?, glouD-plinryngeal, anil hiilhar acotwtory, mkeii
logMbcr. arc partly cftVrcni. partly alli^iroot. The combinr.fl nucleus of these
Ithree ihtvoi, the orllx of which an- small and dcvuid of conspicuous axi§-
icylinder {irocessee. is usually rcgan)<.->l a» n M>ns'>ry nucleus, and in the dia*
Igram, Fig. 193, is riiaded aocordingly. It may jwrhaps be oumparrd to th«
fiiniry nucleus of Ibe fifth. Thu»,ttie oscrndiiig root, or fasciculus soHtjiriua.
presents many analogie- with the ascending root of Ihe fifth, and we are ltd
bo r^anl this as, like it, a gathering of tvriain allcrent Gbrea of the poetenor
boots of several segments ; in its case also the lemi ascendiDg is niflleadtng.
Bill there are many difficulties in ooiiiieetion with this nucleus, as with tne
fifth- We must not enter into a detailed discuwion conoeming them, but
nay rennark that we have here perhaps to deal with complexitiea due to the
Ihct that certainly many vagus and glosao- pharyngeal durcs, and probably
lotoe of those of the fifth, an- apianehnic in funotJoo.
The nucleus ainbiguus (vntatns largu cons|>tcuoua cells and we may proba-
bly regard it as u motor nui-lctu, i3i[M<oially of the vagus fibre*. We may
•no perluiP) place it and the nucleus of the m^venth nerve in the same cate-
gory, and lurlher i-laiv with them the motor nncleua of (he liftli, looking upiia
■U three ok -v) many detaehed [MrtJoni of gray matter, corranondiiig to some
Dart nf ibc anterior horn of the- siiinal tv)r.l. Whelhor tney are exactly
■onologous to the hy]iogl-j««d nncl<-ii«. and their fibres to simple anterior
roots, is not so clear.
Lastly, the auditory iH-rre. both from its character as a nervo of siKcial
trnoe and froai the remarkable IVaturas of its niieisi, bevtn more difficult.
■62
THE BRAIX.
M(wt i>rubiitblir it rcsiiltii from ihe fttkion <■>' niore roola lltsn tim; Iwi itn
iiupoauble ni pivMiti to obtniii « ulor owception of the nnUirc of iba ■! '
nerve.
3, The Sup^rfifial fimg JAiW<r.
§636. Tbowboleof ibfi suriiiccofvnch cerrhml h4-nii>irher« for funis Uilr
depib inwiinl consiflf i>f prav mnlter, |>i>«w»ing ■[lecuil ebnm' ' :•
nllc<) tb« uorlicnl pny mailer, or ihe rortr^ trrrbri. or nhortl;. it
tlw eorttx. As wr »IihII *ce, bv il« tiistolomiMil nnd still nmrc bv lu {•Ihm-
bifiionl frnliirrf. it MiuicIh apnn from nil oilier kinih of gmv iuiiEIlt.
Tbc irhrlc of [he surtiii'e of ih<? et rebel In ni is nlwi eov-crni wilh jtrai luait't.
whi<'b. wbile {>o»«;f sint; lenluree of il» own, so far rawiiibic the cvrvbrnl oir'.(i
ill ils hislulofiical clmraoter^ ibai it too htis bei'ii spoken of u ooriex, m itr
cortex cerebclli. Itv ilH fimclioDal niauirratalioD^. however, it iliflvK *Mi
tnttu the cerebral corlex ; and mik* ihereare muny advantages rn Wiiix^li
la use the ffbrd cortex in eoniiecliun with the c«rcbriini only, it ti ilmriliii
not to Hpeak nf a cerebellar eoriex but to employ the lemi "superficial fm
mailer of the oerebelluro."
The third ventricle and the biuder purt of the fourth wntride an fi
moled in by nervotia uiaierial, and potirSfl no superfieial gt:ay raullei it ' "
In the corpora i(uadngenitna. « hich funi) th« rtM>f of the a<|iieduci "^ <
nf the niid'brain, j;!nir niutli-r !-■> pretfent and poMeMM. in the cum: "i u<-
anterior corpora 'pindriici-iuiuii at feiut. churadersi to a certain cxinit uil>
^OUB tu lho»e of the eurti-x and to the ccn-bt-llHr AujM'rfleial tfray matUc ltd
It will he best t(i consider the gray matter of Uivw bt>lie» » "^ '
another categurr.
3. JTtf InUrmedittt«Gnif Mnttero/lheCrttntl^tlm.
S fl87. W« hiire seeti ($ WM) that Ihe cnira cerebri form llw
pan of A svEtem of loDgicuiiiDal tibrefl MrelcbinK from each ceret
Rpltere to the bulb and to the spinal cont, Tbt« system of libre», upoo mM
we may consider the various pans of the hrain to be as it were fouD'M, ■'
may speak of ae the crural aysleui. It is. it is true, not one eontioun*
strand, but a number of diRerenl »traiids, having diderenl be^inniii{p id1
endings; tint these all contribute In the crura ami are so far alikeaslojotlili
us iu considering them «« a sydteui. The cortical gnty matter of (■chbnal-
qibere is, as we iliall m-c. omnecud with vurtouapiinaof ihit>sysleia.*tuliD
oufl sense we may regard ihi* nMrtn a« heginniug in ihe cortex ofwi
hemisphere, and emliug in the npinal ciml. But certain mniw* •■( ^^
matter in (be bewi:<pher« not »trictly cuTticul, and wretal ini|>i-rtniii m.-i*H>
and arena of gray nuitlrr lying brineen the hrminpherv and 'I
ronueded «ith ltie*y*ten> : nnd th^^tr n<> may .■'pi-ak of a* the ' n
gray niatier of the mif^t nyxtcm."
(.^fptif Kiriiilum and </ftlie ihaUmv*. Of nil the«c novtrnl collectinR^ -'
gray mailer, the hirgct. moat coDipiciinHi, and perhapn ibc immt iin
nre the two nniMHv in the front part of the systom known as ibe c»r|"
turn and optic ihulamup. The former is, as ne have mn (§ 603), a ■■'
m«nt of the wall of the cerebral vesicle, iho latter a dcvelofunent n( i
of the ^'eaicle of the third tentricle. They are. tberelore, of diflereai
allboiigh ill lh« oniirM of the growth of the brain they become eloselv au.
to each other, they are at the outset rjiiite separate and distinct. Mort
an we slinll see, they differ front each other bo eaeeutially, in their naiurri
TUK GRAY MATTER.
rus
nlatiana, ihut ihry rniinoc be coitfiHercd as hutnulogouB bodies ; bdA llie lert»
" bHil |riiii)i;lia " ••Adi «]i|ili«d lo ihrm i*. therefore, unrortunale. Neveitlie-
k«8, it will render ilu' iltxcnjition of tb^'ir luiirgrnpliicBl relations eaakr, it'
for n little uliile ii« coi»i<l<'r ihi-iti ingi-lhtT.
When the latent) vnilHcK^ is liiiil o[>tn from above, iiart of the corpus
■Irialiini it m*ii pr<ijnrtii)}i: into the cJivily of (ho vcntncli-. I» fn>iit the
prnJMtlDg part i« lironit, Innnitie thn lateral wall niiil iiait <il ihe ll<M>r of tbe
reniriotv, and to it> nn-ilinn nitli- lies ilw ravily of im- vi-nlricle, i«|iarated
tmm it* fellow by tbi' crplum Ineiduni. FiirlhrV Iwck tbe proji-i'iing part,
beoomtna ^adtinll/ tiarrowcr, nminHs a mori? lateral pwitioii and pawn
bito th« dtMending hem. In (his rnrt of ii« couno there lie« on itk lunlian
tide, wpamte'l fn^m it hy a narrow band callod the tienia seniieircularb or
stria tcnninali*, the optic thalnmuB. a oarniw Anp of the siirfiice of which
It aven projccling out»idc th<> edge of (he choroid niextia. If now, nitt only
htAlt lattTitl Tcniriclet be laid open by reinoral of the corpus calloxuin and
Ibe fornix with the velum int^rpneiliini and choroid plexus be taken away,
>o a* hilly lo expose the third rentricle. but also, in order to obtain a better
view, ibe orbolc of the hinder Dart of [he cerebrum coDCainiiig the poi'teriur
borna of the lateral ventricle, tie completely cut away, it ja seen (Fig, 1!^>^)
that (Im- two optic ibalanii {O. T.) preeent ihemaelva as two large oval
bixUe*, placed oblitjuely athwart the diverRinjc crura cerebri and convfirgiiig
in front to form the imiutdinie walls of the third ventricle. In fri>ni and In
Ibfl Mdf* of the optic thulaiui are seen the cor|^>ra iilriaia ( f. -V) foroiiiis
■ntvrtnrlr the lateral waJ In of the I wo lateral ventricle*, and dirergiiuc behind
lo allow (if the iDterjiosilion of the optic ihalnini. Ou each >ide of the brain
then ihcK two bodies, tbe i-or[uis Mriatiini tiiid upti<- thalaaiUK, np|ieur as two
maiwes of ^y matter placed on tlie crus cerebri ai tbi», diverging from its
fellow, befina to spread ou( into the cervbrei] hcmisph«r«.tlic corpus alrtaluiu
beinc placed fonicwhat in front of the optic thalamus. Th« relationit of the
two hodirs, moreover, arr such that while the optic thalntnus alone forms the
wall of the third ventricle to which it properly beloDfjs, and th« oftq>U' Mria-
tutn forma jvrt of the wall of tbe lalcrat ventricle to which it in tuni pro|>-
eriy beloon, the optic tlvalamus also projects into and seems to form |>nr( of
the wall <^ tba lateral vciilrtcle, though at its origin it had nothing to do
with tka eor«bra] rceicic.
We apoko just now of these bodies as l>eing placed on tbe crurw cerebri,
IniI though their dorsal surfaces ihiis nri>ject from the donal surlace of tbv
dlrerging crura, a larf:e portion of each Itody is, io lo sneak, imbedded in ths
BubaUuice of the divorKi'iv crus, and what ia aeen in the above surihce view
is only a part of each body, and, indeed, in the case of the coqwa Krialuni,
only A small part. In order lo understand tbe nature and reUtions of lh«w
two important budira we must Miidy aeetionH taken through a cerebral hemi-
SpherM in varimis planes (Figs. 1!H-201).
Each cms is made up as we have seen of a doiani portion or tegmentum
consisting largely of gray matter, and n wntnil jiortion or pes eonsistiBg
exclusively of lougit^idinally ditposod tlbm. The tcgmmtMm end* partly
ID structures lying ventral (o the thalnmits, pwilly in the thalninitn itself;
kJHJ we ntay for the present Iraw this part of the cms out of ctiii»i deration,
Tbe fibres of the {ic», while conliniiing their oblique course forwnni and out-
ward, soon rite don«IIy by the lude of lli« thalamus and hence, in a trana-
verse dorwweotral section at the level of (he hind part of (he thalamti* ' Fig.
194), are aeen Icavinj! (lu-ir previous position veniml lo the substatilia nigra
iSn) and |«ning (Cip) hy tb* rid« of tlw ihaliimiie on their way (n the cen-
tral while matter I'f the hcmisnbere. In this part of their coutse they fonn
n thick strand reparating the ihalanius (In) from k large maa ofgray matter
Moei finilwbly it resiiUa from lh« (Visinn of inons rwrts iImvd oil*; but ii ii
i»i|K«0ible al presetil i<)i^>Uuh a vtear oonc«|)Uoa of llw tiKlure uf ibt »)' '
uerre.
■2. The Superficial (imy Matter.
{ 6S6. Tbe wbi)l«(iriW *urf»c«of Mch cen-bral hemisphere far •uine ItUir
cailcil the coitinil mv nm(l«vr, or the rorltx eetrhri, ur hHotIIv iiti>i
Uk cortex. Am wo diflU fe*s, liy iu hintolwiinl mxl »till mora by ilo )ih} mv
logE<-Bl fi-HUirc». it »tiimlii aimii from all mher kiii^I* "f grav inKi'ler.
Tb* whtjlcof theeurfiirctirihi'rcrrlM-Uiim itnilwi covrrr^I *ilb ([Ti) '
nliurb, wbil«! jxKWMing Ifnlun-n of irn own, w fur rii*-nitilc the cerrhn
in it* biKluUieical rhBritclen Ihm it loo lii»> Ix-i-n .-i><>k<'ii cif an i
cortex crrabcili. Hy ils functional inanir<>(iiiii>iii>, boni'vor. il li"
from the c>.Ti'brKl corI«x ; nwl ^imr ihcmnrr nmuy ii<1viio(nc*« in t^
to um the vroxl c>i1«x in coniieditjn with tbc nn-bnim ou^. it in •!' '
iiot to tfi^V of « r^rcbellnr cortos but to rniploj tbc lAnn "Nuporfidtl ;r*T
nintt«r of tbc ccn-belluin,"
Tbe third vfiiirick aitd the bin(l«-r part of ihe Ibtirth vp»ir>L-l« nn' dm
routed in br nervous nuilerial, and pMtves no superficial gmy nioltrt ft 'i
In th« rorfmru qiindrigcniina. which forsD th« roof of the mquediiLi f\
of the mid-bniin, ^nty malter b preeent and pomeses, in ih« ciu<- "i i"
iinterior corpora (jUAdrigemina at koM. characters to a wrtain <?x(«tit aiioi'"
gotiB to ihoee of tlie cortex and to (be cereiiellar euperlicial gny mallvr : b»l
it will be best (o concider cite gray ntatler of thcee bodtee as beUwi^tt
another caUnory.
•1. The Intenwdiale '>/»y Matter i^'Ae VkKrtU Sgdem,
S 6S7. We have ifeii (S (WO that the crura crrehri form llw iimmttwni
part of ■ ijrMMB of lon^iludiiinl librm ■■(ri-tvbin); fnvm tiich ccnrbrni '■
spheraiA the bulb and tn tbi- ipiiinl conl. Tbi* «yiitein of Hlirea. a\m'.
•nt: mny convidrr \hi: vnrioui^ jinrls of tbc brain to bo M it ititv fmiti:'
may i<|icak of a* lh« critnil (yaU'iu. It iji.il in tni(% not <>»■- < <
Mrand, bm a nuinlK-r i>f difl^rtnl Mrandii, hnving ilillt-mit \*;\l
endings; bnt tluvenll voiUrilHile tn ifae cnira am) an-ti»far nlikt'.j- . -
ui in coDcideriog Ihrni no a syHtcni. Tbo oirlical gray inntX'r of cai-h bivi-
•phare is, as wo shall ee«. cnnnoctcd with various yunt of this sreh-ui, inil '»
WMWfwewc may n-eard this syMcm as begini>ing in iha conex of iMch
hrtniiiphcrT^ and ending in tin- spinal cord, Hot nrrtain masea ofgn^
mailer in Iho hvmispbere not slrit-tly cortiral, and Mrt-ral iiniMiirtaBt IMMR
and areas nf gray matter lying between the bemiepben- and Oi
fonnccted niili tbeByslem : and lbet« we may speak of as the " .n
]fniy matter of ibe crural »y^tem."
(VrpHf Mrintvfn and optic, tkatamu*. Of all ihese seTenil O'Def^i"!)* -^
gray matter, the l«r:g6M, moM oouspicuous, and perhaps the m"-:
are the two niassts jn the IVoni pari of the syalem known a» ih'r ' .
lum aod opiic llialaniu*. Ihf linmer ia, as ne have seen (§ fiOI) i, a <i'
menlof tlie vail of iJie cerelinil veniuie. tbo Ult<<r a development nl'il
of the venicle of the lliint ventricle, Tlii-y an-, llit!ri--ffin>. >>f diirvrtm i .
■UhodKh iu the counte of tlie growth of ibu brain tbvyb4>c»iiM>cl<>«ely«iir
to eacn other, tbey are ul the tmtMt ({uitc tcpanito and diitincL Mi <
as we shall see, tbey diffbr fnun nxth olbcr so CMCOttially, in tbcir naluu ~-
THK GRAY MATTER.
ioD», ihnt tli4-)r cnnii<>l )>e coit»i<lcr«d as liatnologau* bodin ; and tbe 1
^ hMil Knnglin " ••i\ta applied (o ihcm i«, th«n>forr. unronanalc. ynuilfci
Iwi, ii will rctiilor the description of iLdr topogniplticB] nlatioM ««Hr. S
Jnr a little while we coD*ider them lo^elfacr.
Whm the lueral vcniricle ia laid open from nbovi;. f«rt of the («^»
■iritttutn ia »eeii ymjvciiug into the fuvity of the rcntnelr. In flMn «W
prDJrnin^ purt n linwd. foriiiiitK the tuleral Hall and Mtt i>f tb« i
vrntriclr, aiid lo it* nK-diau side lit-A tbe cavity of iW vnttiidt^t
Inaa it* fellow hv ihv trptuni lucidum. Further back tbt faijaataf ^■'t-
bteMnins gradually nnrroiner, bmuiik* a luore lalervl pMliBB md fHM»
Into the dMCMHliDg Imrn. In ihiii ran of io counte thnc fia M iSa mBm
••(Ic; w-|iaralc<l fh>in it hv a oarmw hand ealM the iKsia MBMomlaB ^
airia tvrtDiDnlt*. the a]>lic tbaliiniii*. a narmw ^np of ihe wuhm 4f vteA
b ttfn projodinj; uulwth; tbe vd^ nf ihc- choroid plexBL If :
both lnli.'nil rmiriclea be laid i>iH.'n by reniovnl or tbe «oipa*
lite fortiix with tbo velum iiiterpiwiltiiti and choroid plotaa kr
»■• p* lulty to expose the third TODiriclc, but nlao, in eadar •» at
ri«w. Ibe'whotv of Ibc bitidi-r pari of tbe orvbrum onotnniar ^ '^
of the lateral venlrich-. be COtnplMely cot avar, it m «n: Ts
the two opiic ibnianii {(>. T.) prrvi-iii tbeoatlMi a»
», placed •pblir|ticly athwart ihc divir^ine eran mrfK
fVont to form tbe tinniedialc walU of the third VoMddft. Ifc l
the tidn of the optic thalami arc «cen lh« <«rpwi ani» 'C £-i
■ntrTinrly the lateral walls of the two lateral reDtrictnci
|(> allow of tin ioterpotiilioii of the optic ihalaisi. Ott i
then these two bodictt, the corpua flnaluni and oyUr
muHra of gnf toaller placed on the cru* rrrwbri ■■ Urn
fellow, be^na to aprvad out inlu the cerebral * - • - -
't^nc plared anruewhat in fh)nt of the ofAic t
two Wlit*, DMiTTOTer, un iinch that while tbe i
wall of ih" thini reniricJe to which It prripBl^
tnm fiirnia imrt of ihr wall of the lalvral r^Ot
crly Iwlonva, tbe optic ihahtnioa aUo jirwicfaa MOt'^t-
thr wnlt of tbu lateral ventricle, thougli ai b* •
with thr crrehml vr»i.'lc.
\V>. •[liihi- jimt itiiw iif tbe«r Ii>v)m« i
t ■ ■ ■ ■ '■>ir (Inrul Buit -
' > A tiiian [art.
4
7W
TUE BRAIN.
which, niughly Iriunifuliir in »evtioo, is divided by iiiirtiti'm* if wliiu iuit«i
into tiiTve jiarM ( tip', <ip", PI \ and of wliicli we snail i^jteiik direcilv ■* ibt
niicleUB leiitii'iiliiH^.
PM.IM.
\
.IV.
•r
\
V.3
-M4
'v
'v'^"^:;
>
ty.d
y
tNMia-uoi'TicOiiUMetir i Tiit!«vcnut i>oii9D<vK<itiii- )-u:ii<» maut-nn the Iiiu4n 111
iMii^ti, tl Lkvil PimiMDii iiirtiE KxElur t>ilM(a-vii. r,ir>|iLi <ii»uii.'uJTa>.|
He. iiui.-lo<n aititatiu; In (bo uppBr pKrtoT iheflnM. ihaimlloaarilicniulein h ikiMikl
MMiOUVt M**!"!' *t>Ich (uec<i«di Ihu wHur fruitl vi><1 or huiHl: in Ih* lowvt pui n( tlM%iitl
WMlda p«i«ai> Itironch (he lall i>r iric iiui'lcui iimr Il> mil. una llil* |snlnii of It ka*ftirll»«Ui
«lmnie» beta •UQ-U)(«d fmoi Uio trmj imtivr of .Vu. ducIciu unfv^U*, man lUilla«il| UMl
NalUf i> Ihr •^*H. ^', dp". hIoUiu |■lUl<Ill^ 1H-U Uunlu Iku xvmtob.axl K pUtMnni *t taiMi
toaltcubrt* ; 'IT. lilt' aiitmnr: (tr. itii< iiiiii^i. nii<l la, I lie lalvnil niicloai u< llw opHc iIkIiobi*
«t<»liihi.-* iMIlesl la)vr" Ijinit duii <» (.v. Uie inBUrlor liDitiof Uic lu«cre*lcii|aiilcMl«»
Mlalnc uiMin iiruidi »( itlim u]ili-li iiiliiKlt wlIU It In llir IIimttmiH balwvrn tbf anlMMK^
lotrnikl niirlrioii Diiviii- IiHti>1 nii-l llv liilrnil niivleu mi tlir-tlhcrlta Ujrr •lia'lnl lea >W1) O
%ht flpipc r«i<«entlntf ibt liiti^ma] iiiolu].lur)' Uiuiiib nf iJii* iliaUpuw^ cuiuLi41ii( IvntctfOi *t*
nUtMT, iflhvrcullKiiviu of nblM mttlcr wllliiii tlu- llialaiuiu aiv llvUiv buadltril \10| 4'iiR.
HOd f, I III' Innmrii-I of Ih« aiite'riir iiUbirnr Die funilt : /' Ihtf upfa)r*o-l of tbcuiHlM|(U'^
tIM tvBti. Ulm If. (III! cotinu ckllwiun ; tub, cut\mt lubibaUuileUK. runnJui- •> f-i->ii niuluia*
BUM «rul> tbt Uultimu; .^ . >utM*tiiln iiign: rL <>i»i'i[iuu ; ct. oiMnwl '
■•onioa of AiitafotccuimlHim : h. Itir IiifiiU or lulniiil o( ICfll ; tt.Mnairen' i.< .< ■ ■^■
lnt^utnvt iMItni ntmilvle; I'.S. Im ihu rulUou ufiliv Uilnl renuicle; ItKiHiUiiiaiil lUWiaa
•TV iiwAc ■UifninmulMlIf •IMloc* b} Ihlrk liWk lliiu*: 0^. ot'lc tr*c<- f>. P. pUMll <
r. uinp-mi Me.
THE ORAT MATTER.
If iuiileiid of (flkiDg a traiifvoreo Be tuliP a lon)»ilutlmaI doreO'VC'nlnil
Kor as it » called !a|>iital ! amotion (Fif;. 2U(I> wc lind ihsi tbe librae loruiiij;
|fhe Mranc) in c|ua(i(>n ilo not coniiniie pnriillel to each other as they riw dor-
■bIIv bill diverge in a rsdiatini; ninnner, Inriuiii); the eo-calied corona radiata.
5f a^ain we take hDriutnial seitiona at i>ro|)«r levtU (Figa. 19S, 199), we
tid that thb Blraod or rather thick haiid -if doraally dineoted radiating flbm
^ot only Btretcfaee l,f Vp) between the thalamus stid the firay iush juU kpoken
pf, but reachinK farther forward panea { da ) Ix^iwueu {\v- »ain« gniy maw on
the bieral «ide and anutlier cray man (J\» on tin- mtdiiin tide, the lalti'r fVum
kji |H<Hiiti)n being evidently the {lart of the curfius etriatuui which |)n<im-t)>
Inlo the lateral ventricle. The uiiue hoiiiontal «eetii>i» ftirtbcr («ach ii» i hut
|he front j^urlof the hand (Cia) ia bent at iin angle upon the hind |>iiri I dp).
1l upiK-urs then from these tectionti that the fibre* nf the pe» a* ihey rite
hp dorauly into the- heimaphere uprend ont in iho form of a fan hr-nt u|Hin
Itoelf. ThU run-like cxpaiuion of the pen is vall(-<l the ini'-rn-il caiauir, tho
%ngle funned by the bend bein;; rallcil iia gnni or kntv, {C'iji i\w juirl in
!(r<int of the knto thu/r'^nf limh, und lli<r pnrt iH-hinil tlie knee the IuhJ limb.
'And horizontal M-ni«nH iit lirvelit more domal than thiw given in Fig». 193-
,19'.> would »hnn that the lihmt componine ihb fan-like intvrnal oa|«iil«, as
'ifaey rofe doiNtlly, curved auiiy in variouH dirctctiooH U> reach nearly all pans
tf the furlaoc uf the hi'iui»|>ben-. We nuiy add that though the internal
lapeule is mnibly cnaiprHW*! of librei! whieh tliu» strt^tch all the way from the
Cerebral cortex to the pea uf the cru», it also cuntainR other fibres of which
we »hall *peak later or.
< 1; 028. The gray mniw leparnted from the thalamus bjr the hind limb of
ibe internal capeiile is called hh an hole itm nueleuf feiiticiilaru,aace in hori-
■ontal Hdion it presents a cerliun ihoiifth distant resemblance to a len». Of
the three divisionH inlo which it is xplit up by the partitions nf while raatter,
two median ones Up'. Up", are spoken of together as the g/of>ui pnlliilnt.
Dame being mea to them on account of their |>a)er eolor. Tlte third,
leral diTtwoD f/. tft called the putaimtii. The use of these tno nanie^ for
e two difi^rent parts of the one bo<ty, ap|)eani to lieju^tiRed by the different
nections and leaturf« of the two parts.
The gray nisie which in a horiiunial section (Fig. 193, Xe) 19 separated
ftnm the nucleus lenitcularis hy the j'nmt Hnib of the ext^rtml ca))sii1e, oimI
|irhic)i projects into the lateral vemriclt-, in called the tmrle"* rtiiulaliM. The
ucleus caudatUH and the nucleun It-ntunilaris form together the corpiin ^iria-
01 ; the farmer, nince it projects into the lattTid ventricle, being the part of
: cor]>iifi »1rtatum tern when the lateral vtuiriolr in laid oi)eu, is fumetlnief
ikenof as the intTa-veniricnlarporiinn of thi; whole body, while the nucleus
irularis, which is whujlr hJcld<-n in the hennHplK'ni and in no part prfjeclif
to tht- talrrai ventricle, is callH the extra- ventricular portion.
But only a part, indeed only n relatively r>roall jiarl. iil' the nucleus cauda-
|U» it difcloeed in such a horixnnial wctinn ; to h-arn the ximewhat peculiar
D and relations of ihe whole nucleus a number of Mrlion* of * hemls-
taken in diHerctit planes mu>t Im- gtudicd : and then' will nt the «arae
I explain why the nurlcus is uiDcd " laudiitus.'* Thete t<«ch u« tliiit the
has somewhat the f"rm of aooniniu (Fig. litii. The thick, rounded
i fbnu the lateral wall of the front pnrt of t\w Inlcnil ventricle ; tbeoct
body passes backward, narrowing rapidly and diverging somewhat later-
ily ; iu its course it archee over the nucleus Icnticiilsni. curving oo much
\il the end of the tail, sweef ing round the hinder border of ibni biidy and
iMiging its direction, runs eventually ventral to it. In a horiMnlul i^ction
ken Hi a certain depth, such as that represented in Fig. ll);t, only a portinn
the bead or hiuly (,Vc) in the front part of the figure, and a truwvene
7W)
THE BRAiy.
•cc-tioii of the end of tlio luil (.V<-) in ihf Iilml )>art »r tlie fikiTe are Men ; lU
Ihv interfiling mrtion of tltc iiuctcii* lii» iiUive the plaue of the •Mios.
Ill n irniitvcrao, ili.< no ventral wi'tiim tiik«n »'iiui-whnl niil^riorly ihroufli ihr
frout iimb of the cnptiilo (Fig. I9')), thc! hrJiil or Ixily i>f the u'ucleui »u4i-
(lis t.Vr), which ha.* not yet n-nchtil iu gn-nK^M >litnpa«i<>u». u aecB ]jiog
Fta. Ml
ee
eia
M
UuaiLuuuTir optukc or * Tujnna« mMMO-VBCisiL Aicidw tiimi.'om thi. IIaud
rauc IMtnj jir « LnBL AmKwoK to no. i>i. lAniiiaiNaiDs.) Kntuul itj*.
Jtf. iluclcu(««litiu: Up'. 'Jf>". (lotiut imUMoi, via bent IB two Hsmoun. •ml II. irimmT-t
ItMltlU ttuUcnUrb: OT. oiulc (linlBinua. ulUi (w. uiMTlur uumiuwun> ■■ cliim nIMI
kiturtor Haiti fit iBicmA) (niatiV, 4V. dtcntikl cajiial*; 4ip- npUl Inict : c<. oiffHn i
tmix : Ir. a (pMD ihti III III u|>|i*c i«rt brloaci U> tbt iMenI T«iiitet«. In In inm i
nw Rdd of nlKiaDliMU UmmmM |d« aMn. Ibe •U* Mdik of wMth,(D«>««*a*llfe4
taraaOttibanU ftam: tUt OM vnM OMti^aa In Uw nuMniar itoMaUoauitmBr
At. ik« tenia: /'.ftoaoklhtoi /'.■■rlMBllolir: r-UrapsnlkilK. RKKmicreiauiMnibtc
PdimiMHr. wUckbdutdodlBPIc. t*l>. Ii in ihb flciuc kfl uiuluulod.
(lonol lo the nucleus lenticularis, »«paral«d frum it by die whita oiaM ot tbi
fruut liiub(ri<i)of the capsule, lhou;{h this a Bumewlml broken U|» hv»ir
ot'^roy iU4tler puoaiuji from one nucleus to the other.
Tejilnil KctioD, Mkeu Mill more Mileriorly, Ibrougl
lu a tranavcrtt, <1
the frodtol lob* I
TUE GKAV MATTKtt.
76;
> bwl n( llip nucl«'UK miiiliitut i* aeen at nbr>iit iu ((rMtttl tixo, nnd
Jhhilig iiiirktiii lontN'ulnriii i XI). n'pmiciili.-d Uv (lie putamcii ftloii«,
ihg (utH'il wilh it. llir Ino tHifloi lH-iii;|; >M;|inrAtoii by n *iaall <|uiitility
vkh* tnntU-r of ihi' iiiUimal i-a|i«iilo,iinr| t\v.u Inrtcoly lirokvii up l>r bridlet
rnr nuutor, giving ri»o In ii vtrinlc-il niijiniMiiM'. In ii similar tvctiuD
lurlhrr r»r«»r<l. tin- iiuotcii^ Iriiticiiliiri* n-uiilil he aWnl. the hciul of
iiueli-iiscnuilBtti- iii>iK!«iriii^ l>v ilin-lf. lU^Ciitiiiii^ In l>i<- liiii<li-r juirt ur
lM.'iHUj>lii.'rc, w« 6nA in ii ilon>u-vci]trnl wctioii Uki-ii thruugh lli« himl
Pto-UB.
CC
-■-CI
iKuniTir fMTUJii or i TBAirrrEMH; DuNro-vtsnui. iacttox i» Rmflr Uuin>iik«ii iViW)
, b«ad cf iHicttiw OMMIMiB. sBil .VI, iIh Inuil unit of ilta (MUunia irf tlw uuolciu luuUfulAN*
Blii# fuad «llk II; «•. «or|w> taBcanin, oat tli(««Ii at lU hml h«Dd Or rMiDm, *a tfau bMk
tl tat nM/nl ivithiiu Ira dwHii-. bowWD IbaH it tana thv tnhnairlol* <vcmniTlu lb*
tn tUL-Minn »t ; (r. kMcal nniflcla ; O, clu«nua i f, miniul tota. COcUial gnijr muMr, •» l«
1*0. left uiwtuulwl.
b of the c«ijeule (Pi>:. 11^4) tbml while ibe iiucleux IcniiculAriii U horu at
treaUrdt airs, (he tieit^ of the uud«iu caudaiu* (.V<*^ lyioK dorvial to the
leal lentionlarib uod wjiianXfil fhmi it by a ci>iui<lcriibTi.< ihickn(«> of
irnal caiiwile. biu lourh diniiiiuhiM); the aamis Kclion laorvor^r Hhiara,
iral to the iiiiL-leuH Uiiiiruluru nm) clinging to the dcsoendtng honi of the
ml rentricle i^l.r.ii.), the cxlrenw tip ofth^ tail of the nucleut catidnlui
?»ioo« about tu fb»o with Ihritnuill miu« ofgraymnuer called thsriKctu,*
ftfl'tlir (.Vrr). A UgEttal (lOD^ilmliiial cliimo-vt-ntnit i wdioii taken at
ic ilidiance from lb« median Inif (Fig. I!*7 ^ nh<m» the curved c»uree of
largtr imriiiKi of the iitu'lni:* rAii<liiiu>, the cxtrvma head, aa well as the
tr |>art of ibi; tail lyiii^ >•«■[ ■<!* (lie pUiiir i>f llw Mctian; and a aimUar
THE VltAIX.
MCtion Ukeii nonrcr ilic niidiJK'^ tine (Fig. 200) sIiowb how the nudeui b
tbo middle portion is l)n>keii up by bantu of fibres of the iDiema] oepnlt
trnveraipg it, nnd thuR coiitrihiitiiig ut tlio Mtriiilcd anpeantnoe; tlie »mt
fcction nlM> sliutrs that tho globiii lulliilus, u well M tne puiwuen, beiicaw
con tin II 01 1« with lb« iiiielcu« cautUlus.
Fill. I9T.
-^
.O
vN)
'^'
o
DucouimiTic Onuira or a Sujittil aaamoi takci raMtca n» Riciit MDinriiDi >Xi*
sun mcoi TH* HoMi. ^TRrjtiK. 'itnuuuiBTTOE'-i Uolr nuiuni (i«.
Tb« pltiiv or the nctlim lannllnilriBCiKBl. but <1Ik>i>I]' If I'ltnct. .Vi-, tbr oaudatr iiveliui a>Mal
lolfaclcnaf ttic lvIlen.VriDni«il]r1liiciitlmuiMrlure>lcDi, i«(lBblori>ieI*lun In hr
|«nof llii«*Utim<i(*i>I, II tonmui arch ntcnrBWIUriiTrr IIiir|nm] niUKt om, tlwpiu
■nd Op, the (lolRW i«llldui or ihc lontlciilniniwleitfi Xn. Iliv itnnilBlcild nueleiw; t1. (L <t,l
Inunnl («[*ulc ; Oi. ibcaDtorlurcuniiiiliviLn ; i r llm Iilrirtm tliiitl nrWiiii nf lliii hiImiIiiwi
Oftlkal -. I: ihi' iKiluIat iaW; T. Iliu tviiiinrvl.
Thus, u'hvn wc sjniik of tlic curpus slrintuii) its » wholo wo i»v«n a luge
iiiiuH of gray mutter lyine liil«nil to the optic iliiilnmiis, rcacbtng nesrlrM
far back us that body nnJ strccohiiig iiiucii furtbcr Ibrwanl, as far forwaiilin
fact as does tho latcnl ventricle; but it is iiop<)rt«nl to rvn>riiibt>r that Uil
divided into livo massea or tiucln. which ari> fused togi-thrr. and that in-
perfcctly at the very front only. These two nuclei are. ihe one the camna-
sliaped nucleus cauaatus, the bulk of which is plaeed Ibmard projecting int*
the lateral ventricle, and whteti on the whole is the more dorsal portion of
tlie whole body, the other tbe irregularly shaped nuckus leuliciitariti. At
bulk of which is pliiceil further hack than the lateral ventricle, by the aiilt
of tbe optic thalaniii£, and which on the whole is the moie ventral ixirtioo of
the whole bodv. It is no lex^ imjxirlaut lu remember that tbe radiatiiif; l)hre^
which we calf the tuternal capsule, pats in tbe hinder revioa of the wbala
body between the Uutlaiuuii and (he nudeuH lentieularia, rorming the hind
limb of the capsule, and in the fhial region between the uucleiis caudatu*
ntid tbe uucleuit laiticularu, formiog the iVonl limb of the caneule. the froat
and liind limbi being heut na each other so au to form an angle, the so-oalltd
koce.
TUB QKAV MATTKK.
rin. IW.
7mt
<*
ff."-
itf^
i
'f?^
Of-
ai
r**l
^"5^
*.
I
Vm* or Muht Htir OF BuiAiii or Uisi *> n»ciMMi ar * Lwnim'DDUL sicncni ik iiic Uuman
LnatiOKfraii mi LoHom-inHAL Fiw-ku. {AiiKiiaixiiTMi.) lUI/ iiaiiiiml iln,
TlNto*,KtBMtna|liiiilIiu)icccloDM 0. lUK* lata Uiciiuv J>, uul IdIo Itie rrw ontrf. nbkh
llilbCMlablk|B«l)rwnM>Mtld1ir«rB« tnlo Uio hcnil>|ih«i« •nd )«■«* oiil of Iba notion. A port
«f UN Tcntml Mirfluic of ttn vrm li tfaown In Ui» iiiiiM ton matkad VX. Ai 0/. tliv anirni ranni
•>r iht iftMl coM fa n*n oivnlnR udi lnu> Um /ohiU oatnclf (41b). ararbnng by tho taicbetlnin
lUafHol Id lb* mUUIv tiiiiri. kuil innlas oD bf tb* afwrfvcr tOHUb tb* poAHw, 9 P, and duenor.
</■*. r«rp«ni tptadriariittita Into the flUnl rmMelr (1). Tbo potttlor osTpOi qllBdl1(UIilMlill ii «aif
(IvDOB bdilnd miiii Uia rairr i)f I ir-ttim, aiiiwiitil to UM «ip«rfor podaBola Of Uw e*rrMlnin, Mid
■vcniDa Umclntillaal McUon oTprbauclna ilie finnt parlor Iho Ibunh TanUlola Tta rorpoia^Ua<-
(KHUaa tt'"" ralallnlr imall brcauw Uio mcU&u iiaiaw Id Uic latdlaTi line in ibo dDina^im
banrMffllkartibCanillvn bodln'if Uif cmiialni uid [ininBlUiliiljr In tnrntiJ tboin la Uiw •ocaon
uf Ihanoriallr iiIbcbI i>(iiniliili»»f f, wbich ororbaii^ tbc opculng of ibe itqiwduet lato Um tblM
twpfct*, anil UiB ilulil aim ol wlilch niiinliin In ttit bitaml wall of Ibr tUM vtowlolo la allows by
aa nnakated iiBoL Thanor of Ih* tbIM rtniclotc It hcd lo b* (iratabad bjr Uw «rc*i «f tht JWbM
f, afcB»a naaliaded In lon*ltniUnal acvuon. r«i«iDrlr ibe tody oTIto fetnts paaai )>!» lte4lT«K-
bi( rtfbi ^rtrtrn-jWItiir. whina ^liUuoM. and laluuu vl» uudn tha onrtiBngliii roundadUod
rod or ipimmm\$inottheearji»trmlloiu'n. Inlhmi rbv bodj of Iba Ibnili l« Man |iaalii| Jbat behind
Uia ttafimn* •canion uf Ibc onlrrMr fammlHun .1 Inlo tbudlTaistuKlfsl>liM'<rtiirv<W>r/. whloh la
tMtintto««atltl>«(clif*1nilio lnEn«linl1i:^lh*<xniHolplnvaidUia<wpaaia«>iHilIlkv*e*«MMiM
M. Tht BMll whllo i-rma Ituincillatrlv taliliid / liidiau* Uw imlUoa of Ite /liiaami qf ITava.
Tb* bii%tat malliin lurftas of Uw ogiUf rftiiUiiiiu. » r. it xen fbnninf lb* laMial wall of iba bInMt
(and, oatlBii la Uw eiaalal Aami^ Uw nuni doml i ikh of Uw ihlrl vauUida. and on tbla. tulow
llteaM)iarilwFla(al(VUid,l>a«tCOn*tiadod. UxatcUonafUwaitllorwfiMlfenMilHnd BM««Fn
thr* tOiBBl (land i^aiid Um *[iltfiluni (J^jkwMi Uwhinilandorjndrtaarof Iholbaluiuui |i[«jm<.
tof InMlbaaMAllad imunne dwunof Ihcbialn. thonn ahadad In Uw Biui*. bf which (bo |iu
nwWi. laaJiiiiilianiaUi lliii >i»laiiliM cil iifll Mliiiiwaiin ilniii llii iiiii<*llinn. |alaaaiiaw
to (ba UiUd taoblria. Ui» iwWoii of lb* nhw b*ti«*n<ni bjr lb* iblii bUak Una -^f-""! (Kb
thaii|il«iliBn iDtlw Ibrnli. Tkr franl laad moH! ivalrali |ian of (ho Ihird Tcnltlvlo ta aeni tt>«ad
Ui Um li^mUlHtlmm, aiiulidd lu nUcli U (lie iMMItry btrlg II, «u La wcllun *i t In remit of ilio
IslandlbdtnDittnnitbaopnfwmnrt aonwat UiaiyAcrfMawallnt Oi>,MraIablaf Ran whkit hi
iM aoiWrtM WNnmtamn U Uw hMiaa vrmiaofti. MrMtMof bM««MUw«atp«<*li»iiMosiic<n
la langllndtawl anUon irUb a NriaM appaatanw. and auUnt In Omt ai Uw nMMM Jl and babind
■I llil»lrtWitnw ;>i ilniMltj aiiil lliii ftiiiili iiailiaflj la aarn inriitwdtall thii afptaaa birtilaw Wf . bw
iftap^4af panof iMt baabnnuuiavay laetdulodlKlsaaUwrtfbt laleiml ranMtla. tn Uw wall
ct labldi la aaao Ihv btilgtnc aadtw tmittaltu .VC Abnte tb* Ooti'iu Callgenm I* aam Uw nM*bU
•vAm ot tbe dibt banila(b«re fentfnc iba rttthl launl vail of Uw lonxtiuillnal Ownn. On (hia
aiailal aofftM appna tminodklalj ald>r« iba ceriwa Mllsiuni Uw anbnl (ijirM AnalMtai (;,/*.,
dallMdBbonbgrUMoJItav-aMrtKaaJjlHiiw/^w. TbawboleollbcniMbcrwan In Uw fMoial F«0an
49
r68
TBB BRAIN.
MctioD ukcn n«rer llie middle lino (Pig. 20O) sbovra how ihc mclnu ii
ilie iDtildle puTlMKi in l>n<ken up b^ buiu« of Glinis of tb« io(«nia) optV
inivening il, nnd ilius <x>i)tributing ta th« ilrlnted anpcanueo; ih--
aectioD nim ahows that the globus pullidua. u wdl u tb« putanMi, ^.
contiDUMiB wilb l1i« iiucleua caadattie.
m.tf9.
0
^Of
O
m
MviiUMaATir otmutM or i Ajuiimi. Sncnn'i itiiiui rnncita ihk Hiuin ItuKwimn <)i
•OM nam tiik UnOAL tii-ari's. ^MMuaixniw.) llnlr iui>iml tlw.
Th«|<l«i>eo'llH.-Malu»l>ne«Uul)ra^lu>I Ihm tttKhUrliirllari. .Vp.lhxaiidMetiUclMic
laUlOliAiK UwlMI(nA>lnaMtlrlU*'lt>t*alil«t)or»tviil,toriRfciarilMI(>It«'* i» -
[wltirUipaMOfMrviiani, It ftmuuiuohnf (n;iM»«Bt«r il»tm] Batu
Mddp.lbetli'ba* laluduiofUie taalkutojiivkiMj AXiliii MnjrsdBic*! nil - 'til
MlM: J>.ilwiarl(Mlobe: r.ibeumpanl.
ThiM, wlwii wo spMik of tl>e corpus HtrijUiim iw it wli»lu we Hi«iia » Urfs
liiMB of gnir m«IUr tying lateral to tlii< ojitic thiilniiin''. r\-«cliiat: ticarly u
fkr hack na that bod}- and atretdiiiig much furthrr lirni iin), »« Ihr furwaKl in
fact an Aim* tbe lateral v«ntricle : but it is impunant to remember thai it »
divided into two masea or nuclei, which aro ftieed together, aw) tbit in-
perfectly nt (ho very (Wtot only. The*e two nuclei uro, the one tliv coiitin*
shaped nuclcii* cauoatus. t)i« hulk of which is placed forward proitrcliiie tnl"
the lateral ventricle, and nbid) un the whole a tbe taore doiml {Hirlion nf
the whole body, ibe other the irregularly shaped niicleua leulicuUro, du
bulk of which is placed further hiiek clmn the lateral veolricU, hy the wl*
of the optic thalaniua, aud which on the whole is the more vent nil [NirtiMiof
tb« ifboK bodv. It iano lene im)>ortant to remember that the rs'listiui; libt«^
which we call tbe inleninl capwile, |iaaa in the hinder region uf ibi- ' '~
IkmIv between the tbalumu* aw) the nucleus lenticularU, li-rnuTr.' thr
limff of the cii]iaulr, nnd in ibe front regioD Iwiween Ihi' v.'
iiud thf Mucleua k-nticularif, tonoiiig tbe ttoal limb of the cui ,
iwd hind limU Wing bonl od each oilier ao a» to form an angle. tli<< ti-eaS»
knee.
no
THK BRAIK.
i 626. The uptic tlialamua lu a wbnle U s soniewliat ovul nuus of ^^
inittter lyiii);, aa wc liav« said, nthwftrl d»e iltrer^itix crus. in wliicb tl b
|uinl_v iiubeijded. ItH curved iue<Uuii Mile ■■•ivereil with » thm Inyer ofna-
tml gray matler foriuH the IniemI wall of tho (htnl \-«ntncle ( Fi^ 193, IH
199), mkI in n loiifcituOiiml verliciil seclion of thu liraiii taken id Ote lltw 'i
tho midille af the t^ln) ventricle (Fi^. 1!)><, O.T.) » hvii iH'cupying tbe m»
belnccii the formx aiiil hiii'l imk) (vple-iiiiim) <if tho cin>u* culoMn mqit
■tnd t)ir iliveri;itii; itiim below. It* man- ur !>»• utraiglit lateral border tbur
on the internal chimiiIo (Pi>n. 19.S, 194, 19^). Il» ilmviil Hurboe. M «• hxu
alreadv nvn, iiliut furinj" jmrt of thf wall of tin; thinl vcntriolv aiwl bt fnr:
bnl ibcri- liw clow ubnvi' il the |iri)|i>iii^t(iou of the piii mnu;r. fomiiRg tkr
velum iiHerpiwiliim with iia cliorotd plexii» (5 fi()S). which crr«|» in owr ii
bcnontb ihc nrojvctine liind eiid of tho coriius cnllosuiB aiid tile fbnii
IFifi- 11*8). It» ventnil aur&oe is fused with Inocnis; indew), the t«^timiit
or <jor«aI portion of tho crus ntajr be said to end in it and in cortaiD HnK-
tures IviiiK vciitnti lo tbe thalamus, in what is called the " mbtlialunic
retail '^ (Fig. 194), while the 6bnw of the pes jnas firet veutral and tlua
lateral lu it to form the intornal ca|>Bulc.
Tbe gray inatt«r of tbe whole bodv is more or leas dislinetly divided bt
>heet« of white matter, as seen botli id bonMiital aiid in vertical lentiw
iFi^. li*'-i. 194, 1!>9), into three parta which have received the name a(
nuclei, namely, the 'pWum or ianer nuofeiu ( Fix- 194.ii(),i*hivh with the thk
layer af central ^ray matter forms the side wall of the third rentricle; lb
larjcer /'if«raf riWoiM (/7i\ which abui« u|M>n the internal capaule; aixltki
nmnll anitrior nwnlotia (Rti), which lies on the dorul aurfiioe nf tl>e fnti
jian of the body, and which thus at iu tntnt end appean to proje^i into tW
lateral veiitrtule.
ThcH- three nuclei form, however, not the whole of the optic thalamus, tral
only ibe birger front portion ; licbiiKl ibem 11'^ the important |>orti(iii calb^
the pu/riiiiir. into which the hind fxirt ol tlu- meilian nucleus merpM ; (his ii
partly imlxvidod in the crue vontniUy, and in tlic hemisphere laterally, aal
IS )>artly free, coming to Uic surface beneath tlie hind end nf the corpui <al-
loeum. Id a median longitudinal section of the brain (Fig. 19S) it is tht
pulvinar which Inrms the cushion-like (hence the name) end of the thalann
beneath the overhanging eplenium of the L'orpus callosum, by the nde Df tit
Cineal gland: and in the horizontal view (Fig. 193. /Vr). in which lb
emlspheres are euppueed to have been removed, tbe same pulviuar it mm
frojectiuit over the cms by the side of the anterior corpus quadrimninMi.
be buried portion of tbe puKnnar is eipiiaed in a tnuMrerw aecltoa tatM
through the anterior c<>rpu«< qiiadrigemiuuni (Fig. 192) ; the extreoie endtf
tb'iK pan of ibe pulvinar ( I'vt) is here 6txn lyiuK dorwil and lateral lo thr
l>et of the cms, immediately above two inoaMS of f^rajr matter, ihe crrfx-n
getiiciilata ( Q/l. C^ot.). of which we oball speak later on. Oni- uf tbeae. thf
Inlerat corpuii geuiculatum C.g.L), is especially ooaneoted with the oplit
Uuei (o/t)' oi'd, lis wesliall me hereafter, tbe pulvinar iUelf i-< aUo ctiniteolsd
with the optic trad, and 'u an iui))ortant |iart nf the oootrat appitratui uf
vision.
S 030. The mibnUtiUUi nit/nt, Uv re<l nufUiu. nwi vthtr p^^f iimtter of &>
ttgtiuMtum. Nerve-cells and groups of Dtrvo-cdls, or snu of gnj tnalier,
■IcHMlB
In innt ft ttw miiM>(M:ita»l itani*, irmiah iJIrtiM bjr n«tim^ )• calM Ihs margli
inlhtmlMlepuiBtMlnaion»tior>ioCtbt cetabml MlalaDcr bu bna nmovrt Inordarlo Am
(tM tMlHnn of III* BrsinW jbmt* or jiMUH qf Ililait4<i,/t..»nd IninunUstrlr Mow IhlthMnaiM*
01 PA.C.tbi^ punbvninii i(i4i>I«. Iii ihaoanpltal nvl<m. J>K.0.1t lli«f»i»n*A>iirtwaAiHtM«lr.MH
C Ilwfww, irhllB tl O.L timn ■ p4rl or lUettntwU Uibalt. Tl. ka |«n of ih« AV^Nf I
aedpOal eantbiiUn. Uie (tMior p«n of whlob 1* lildilcin to vlai* by ibe pM» ■■■d cnn
THE RRAV UATTER.
771
lo daicm spwtial nunm, »rc scnttored throU);hutil itie legfiuMitiim
' il* coiiTW. But, beaides tfaf«o ami tb« niicl«<i or lli^ltiinl am) fuurtli
tl atrvtt, of which no have already spoken, certain lar;;«r ci^Ut^dioaa
_ Kv BtattM' doBcrve atlention, A conapicunus maas of gny niatler, n\r-
ilftr in iruntvcne s«clion, placeiH in die ruidat of the tegmentum on each
de but fiomenhal near the middle line, and atretcbing frAiD the hinder
;in uf th« third venirii'le heneaih the anterior c»rpus qinidriKeiniQiim
lf>2. I<J3), is, from tlie red tint il poawma, called the red nuoletui,
itt, ur locu4 ruber. It id trttveraed by fibrea of the thin) nerve as thew
ik« their way ventrally from tlie nucleua to tlie surface.
We mu»t ooutider alao oa belonging to the leicnientuni a largo are* of
ay matter, aumewbat lcD»-aha|)ed in nerliim (Fik- 192, Sn). which lie*
twcen the pea oiid t(.-gmentu>n. xharplj: marking olT the one fr»m Iho olher.
Da it< dark Kp|itMtrani,v. duv U) the ahtiiidaiMe of black pigment, it is
thtr tutulanti'i nigra or lorxu ni^r-r. It acquire* iu Ikmet dimonsioni
It till- mtddk- iif the li^ngth of ttif cruH. cuming to no 004 in front (P<g>
«) mid Jailing nwny bt-hind (Fig. 191), w the crus pa«MS beneath the
rior corpora tjuadrigeminii. Th<«c twn. the red nucleus und thcxub-
nigra, are porttapa the niuel imporlnnt wlk'ctions of gray matter in
Qontum, bul wo may nild tbnt at the frant of the crua h tbo aiib-
. nigra comes to an end there ia aeen in a somcnhnt rimilar potition
itral to the liind part of ihe optic thnlnmua a colloctioo of gray matter
Jl«d thie nrpiu ntlAfuiiamietim (Fig. 194, t'.ib).
At the hinder part of the crtia, ae it ia about lo plunge into the pona, while
ba pee, now decreasing relativelv in aize, still continues to be ordinary white
■composed uf longitudinal bundles of medulUted fibrea, the tegmentum
iktt no more and more the structure which in apeak! D;f of the bulb we
illed reticular formation, and which, »:• wo aan, d«wrvei to be coiuiderei) as
. kiod of gray matt«r.
Jftc yray mattfr 0/ Ow poiu. When tbo conjoinod crura as wo trace them
vani plungn iM-iiojitli the ponw, the longitudinal fibrvs of thr i)os of each
in, nf wr^ have said, aooii split up into bundlw niid i>cntt«.-rcxl among the
verM fibres bciouging to tho poiks itaelf. Dorcal lo this splcni of irana-
I and longitudinal fibre* formuig the poos proper, botween it on tbo vea-
jrfiwe abd the centra) gray matter with tbo pnst«rior corpora quadri-
CD the doml mriiM«, is n region which may be called u^^cntal.
t it is a cnntimialion of the tegmentum of the cniA. In the front part of
I poo* (Fig. 191), where the posterior corpora numlrigemina «till form the
tina] roof of tlie section, this tegmental area, wnidi ia much broken u]> by
I atranda of longitudinal ftbros, of which we shall apeak later on, oontaios
* nerve-celb, and ii largely composed of reticular furiaation. In this
I on each side agnnipof nerre-cells, tii« lotvt raritleiulVig. lill.JLc),
thivh we have alniadv referred (§ 622) aa probably serving in part aa
[thaori^n of the dcncendiiig root of the fifth nerre (V.<f.). iust ventral to
rhidt it linL Thin nojuirea laiver dimvosioiu further back, in the front
, of ibi' fourth vcnlrtdc (Fig. 193, Le.) lietwc^u the levels repreiented in
190 and 191, and ix a collection of large spiudle-nhapcd nerve-cetla; il
bluish tint when itj> bhiclc pgnicnt is Mon shLniag through the sur-
Bg more or lew tmn^iKircnt material, hence the name.
Um hinder pnrf of ih« pnnx (Figa. 189, 190), where the cerebellum is
^orerhanging the open fourth ventridei the reticular fonaatioo of the
Dtal area is still more conspicaoos. The only special collection of
^aoatler in this region to which we need call attention i> one which, con*
f, like ibeolivary body of the bulb (or inferior olive), of a wall of gray
772
THE U1tAl>'.
niBltcr Eurroundiiig and Burraunded bv while mailer, is rallei) thtl^fpiri
(F^ IKll, l»U. *.«.)■
Toe veDtraJ fiarl of the pone, nrllie pons pr<i):er.niilike ilte pesortbecni:
coiiiaioH, niix«(I with the flbree, a verv cuDsidtrable quantity ofgnr mtiifr
Tliia is fairlv almmlaui in the from part of ihe jxina i Vig. 191 ) Wo« li,
corpora quailrigemiiin. hut increafefl even more behind this * Firb. 1«I>, JMh
Heucc. UKiugh Ihr ponn proper is largely built up of traiiiiverec and iMuiiu
diiinl fibre*, niid iboiifih it coiilainH no comijact H;^ri'^Blioi» of ^raj autltr
rc(t'iving ii|jei;iHl DBiues, it due* contaiu Hialtered ihriMigbout it t very lirp
i|iiaiitity (it'gruy mailer, far more, indeed, lliuii w pr<-«rnt in ibe t^jTaenul
|Hirtion : the gniy matter of the pon* — thul U, of tlie jioiu proper — ibuii k
n'gardeil an fiirintnu: ii very iinixirlniil part nf the gray matter of tbi- crnni
mtrni, ami nf no litlle phy«<jlpgicul wgjiificancc.
Dchiiid the pons the ernral vycleni is continued into the bulb, «ith 'hen
etructuro no h&vc ulrcndy dealt.
4. Otfirr CttlUttiont of (!nty .Vatler.
$ 631. Of tbese, three descn-e cliiel'utt«Dtion,aDd may lie cltuMtl ta|{«ll)n,
tboiiKli they differ in nature.
7'fte gray mattfr of the airpoui ijtiadrigrmina. On cidi M«le of anil Mn*
what dorval to tbe eenlral gray matter of the aqueduct which, a> we limr
•ceil, is well developed, etpecially on the ventral «ide, cnlleciionfi nf ^i
BUtller form ilie chief part of tbe corpora ijuodrigemina, both anteric>i Mil
pcaterior.
Th« gimy matter of Ihe nnierior corpora quBdricen><'>* (I^'Jt- '^i •*■ V- ••
» more ditlinclly marked nff friim. and MpnmtwTby, a uider tract cif wfciif
mailer from tbe central giBy mailer of llie utfueduct llian is ibal ofilir p>
lerior corpora quudrigeminn i Fie. 192, "J''/) : it it. moreover, of a <)ilfmii
nature. Indeed the tuo pairs ot bodies have quite diHereai relalioDf, art iJ
diflerent nature, and perform dillerent functiona.
fiwyjdryi ffenicvhilfi. The two oplif nervea, an we shall aee in detail l»t«
on. give riae. ihmugh the optic decuMiulion, to the two optic trnela. ([arit
optic tract (Fikul IS'J, lir*. 0;i), wiiida ruunda the criia cerebri on ii- Teainl
nurfece to reach the vubxlame uf the beiniaphere in the region Mxatk
opiic thulamua, and »■ it doea au ia described aa dividing into n Inleral in'
metlian portion. The lateral portiou juat a* it «nei'|)<i round the far nip,
that ia ine initer or laternl edge, of tbe crua l9rar* a mun<led awcUlnir ' P.-
146, R and C, C.ffl.} the lateral or onttr eorjiii* t/mifvlalmK, (he lui ■■
wfaicb confiNlH largely of gray matter (Fig. I!>'2, Cgf)- Tbe median i
fimilnrly beam unottier like twelling orr-upying a more median piwi-
mrdi'in or innrr eorpu* grnirnlalum i^^Fig. 18*1, A and K, Cm), iJie initr>c
of which (Fig. 11*2, Cytn) ii1h> concists of gray ninticr. It ■* to hengntBal
that tbece two bodies should bear the nme nume, for tbcv nrv diffimii a
tbeir origin, in their conneclinns, and in their InnctKHM. Yhe luli-ral l>*N
is nid to be derived from the fore-braiD. that is from the vesicle of lli« ikini
ventricle, hae definite conDcctioita with tbe reiiniil optic tibre«, and i* dii
tlnetly concerned in vision -, ihc median body is derivol trom ihe miiMifvi'
ia Dot definitely cDnnecUd «ilh the retinal fibres, and ap|>enrs to bo ta oi
way ccncemed in TiaioD. We »ball, howtter. return later on to the conixe-
lions and probable functiona of llie«e bodies.
f'nrjrtio dnita(vm if ihf etrtinltuw. Id tbe midal of llie »■» of ■1*
mailer which ia formed in lite interior of the cerebellum by the coDtMU*
of llie three piduncle*, is ioiind iFii;. \^9, CD) an area of cray MliO'
iirraDgei), like the olivary budy of the bulb, an a sharply (bided ur |il*il(^
THg GItAV MATTEK.
rio. tm
rT8
OVTUn or DeiMUSTiL ScmuH of liku^ v> now tub Ifrnumn, <'tr*i'u. MunlMM.
I te Mkan M a Inol mni« Tnutnl lliUi iliowiiin >^ m Tliii«ntr uiaUwof (baracUs
■mUlanaiukailei.biit ibMOf itao)[|iiu*irlBiuuiaa'l«|iU«ib>Uiiiiii li ihaded ; or,
«itila<l«IIMiM.>ti0irtiitUi« axUao. LOBBl.AUd aauilar iiuctul ; .ViL iiucleiw knUcularlt, akvir-
H« t*cpatiBaataJ|CkaB4»irln<i«r4lTMei>Of (IU(labut fvllldin rtrr •mall : XC. nocteiB «»u-
.Ikakifvbaad tafroalof. auillliaitliDtalihliiitall tMhIa<l.tIi«lhalaiuu>; O, ih* knee uT Ifet
. mini"<|>("ta-'JNf."nutf (tafodUoaaftfeipgronil'tBltnetaaaiiilMtaiMd
Ita «m»»l bun* iBdIcale tmadlr th* niaun pmMmu dT ik« Mtwal coiMdMaiUa <rf ih« una,
iMMd ae«nrdl«c lo Ihs wavuiuMi wllb wWcb UOTal* awwniBd : Uiui Ar. iDo^aMcMb o( Iki
*T«t. JM. <4 ibc bead ; Ik.oritwlonco*: ••tt.uriWnuulb.SW.arilwAaalilar; A,«#tlM«)!bnri
P^ at itw hauil : .tM.uriboalilaa«n: Hi^vltiir hip: jTa. of iba hate: Mftof lkafca(i4llto
iBai<0ni-n<vl|>tal imt : vc, Staw to lb* iiralrlHl tota : <%■■ vfU" nllatioa Ai iMi lavcl (h* Hint
a th^ltvaui Mt^laUianiW'Uinbnr tiMMpMtolMhiMlof ttenrauiUal tneu nw aloM* katl-
MBMltl'. ^Wllrl irtth Uig iiUm <ir lbs koUiM. CC Tlf. M. /Vaa. <«. tlu nMniH of U« rarpiw
(•ltaNn,'MllWif«nli*'<'<h«winr both «IU MtO* barlBMMallr. Th* tlOtkdark lint liiillciaM*
i>a WwiiilaiJ nr ibiiiaiHI ( Un •itMioranl 4«n«iiUiii kcrw o4 ibo bu«tal nuiral and or im
Ife* tttM nnUlotab (lia lw« T«alil«l«* Mac U4 Dtwi lilo onn br tiM rvmotal or (ho nliim aiiil cliu-
WM H"*** ***- 'n«DtaloiMlinal>lb(B>FB'pai1a<tbUniirUTl»>llaam tkcfonili. Ul*nl lotlic
nadaii* l»lUlraUi1> l> ■«« is omMim Ikt clauiLram. Itw mrwi sf ilio liUa4 <rf MM ao-I Ihr npot-
fala«atM«««lilUanontUn''<t*MW*n1 0(11*11 fli ln»rl>4 lutteir irkloh !• lbahlii<l-(«rl
MtteMMo*.
^
774
THB BRAIN.
band in (he shape ul' a Ratit or boit\. An In tbv HiniilMrr uliTanr tiodt tbt
gniv tualler of itie flank i* cuveml up by and il* iDi«riur nllcd up with *liil«
niaUcr; the uoutli of tli« Haik ii on each niilf, dim!t<<<) lownrt) Ihe mf^lui
line ; the fibres paw cbii-ftv to tbe ttiporior ]H-i)iiDctt'.
Thent are ul»n othut itilWlioiu ofsniy miiirer in ihv ci-4)tnil uliitv nalttr
(jf the ctTtbclliini, Din^ iif which, ciillcu ificniiclciunf tlir njof." iiCooaecierl
with ibi- two iiifi-rior |xtliinFl<».
THE AMUmEXEMT OF THE rUIRES OF TDK URAIS-
{ 63S. The systeiDB. Iracte, and btiodlce of fibres m H'hich the white imtttt
of the brain is arran^d, may be distiDKtiiahed from each other, partly ibniugii
mere inechauieal separation by Dteaos of llie n:B)|>el, partly by l*eiD^ incri
out with the help of the microacupe, but, as m the 9j>inHl i-nrd, miu'li nion
fblly and completely by diil^rences of (fev«lopo>eiil, aitd by thci nieihM '^
(legrneralion.
')C
UCTUVS or 1 9imnAt Sbcvmoi tii»ol'iih tak Ummnimm—ittx. itstaaiKiHH >
Tlwustilnii lj uktu not far tollwrlchlor (Iid moillui pluMMnd lnanc-lMlf UlMarur infuwl —
TheKiai mitilvc o(lIie uurpiu ■Irlaluiu HiirJ Italamu* U tlMd*! .V<% ^. Uw caodM* nmt^K. ^
ltepu(Mn(n,»iul i^.tlw|lai'i»i»iil'liuor(lieleiiilca1ttfaiKil«u«: or. iliiiu|iUc iliMluna: C3.iw
InHmal oianlt wltli a ilraatol ■I'l'snuutn nnallnf aiiprajilmMd)! Ibu illfvrUon lmt*n kf "*'
biwtH* pHUot tfiM It from ihu i»nlon nt ttitmit ntdlua orar U. la >hMn wn nf hun*!* ■■> ^
bfcaitlTill**laCIilil>edabwiMtij-slna,yroa,ap]r'*'><l4*llJM«in, P rOMllattTlMht* tola mitel n**- >
tn<lil*> ttraaU, 4un»liiinl>t> iihn. (uh >, tnil Imatenaoiil ila*. mc.) mn*. ami ■ '>«|—>
dfMal ifaMni. •m- : lb* tlimiliin ^r thr tmn of tba intMwit o4|niil* l« IndleaM br f - (X. <■*
«Tiuetrtbt1:0b llici»<allad-ipuv rw1Uiic>iu]MHliizInlotti«(i«clH>*l ■■>t»i rv. tbt >9faiil*.~m)>'
■ha C0Tt*i> talluuiu; r, >. r. llH-liilafal mUrlQlt cut acfnai iii l>in« iliITurani t4a.--
Hiiliaiilin II f'li Hill inHliilnliT III iii« aiilliiii nutofltietvivl-'lUiiBltami i . .
rtfbi.
We linve »eeii that a o>arlcei) feature of (lie hniiii i^ Difttendil \>\ "
crura cerebri wbicb, iiiniiiii^ forward from ibe hiud-parix of tii
spread out into eacb r^rohral hemiaphere. We bare alau ;«i-n tliat i >' '
in the wiiU: twti»c of tb« nonl conaiaia of lvi> parlK, a dunMil part, i>.
Dieiiluin, and u v<:Dlral part, ihit iim or rruKln, ami that tlimv two iMri*
very stiikiitgly fWtin ea«h <rlhor in HlriicliirL* and in rvlations. Titr |*-
mt£ cxctuMvHy of bundlts of longiimtiiinl libm, and wv may Ini '
from the cerebral hemiupbenv iiilu tli« tioiii< and aoinv of ihem hvyi
pons into tbe bulb and Kpiiml oortt, Tnv Itftnii-iiiiin) in more niini'
Btrvciure: it cviiaula of gray matter, and of tibnn and biiodleB "(
d<&r
I.ON'ClTL'tHSAL FIBRES OP TBK PIUAL 8TSTKU. 775
luivini; various relntiouB, bolh wilh die oilcoiiom of gray mutter Ivitii; uitliin
iuelf and wilh aurroundio^ ittnictur». Il too has connections with the parta
lyipg in front of it, and with the parts lyinj; behind it; we may trace it too
^M^kwan) throufcb the pons into the bulband forward to the optic thslamus.
If we allow ourselves to oonoeive of the optic thaliunua aa constituting the
froui ending of the tegmentum, we ma}' arrange a large part of the brain
Into (wo main regions, into h tegmental re):ion stretching from the optic
Uialaniu* through the donial nortion of the pons to the donnl portion of the
bulh, ami into a re;;i<>ii. which we ma)- <-all the |>edal regKMi, stretching from
ibe intrrnitl c>intiu)» through tht^ venlnti |nriioD of the pona to (be ventral
jKirtion iifthe oiilb.
Tlir Hhn» of thp hmin. iis ii whoK'. iniiy Ih.- hmodly rlii^itu'd into longl-
liidinal tmcts mnnertinf;; piirt.i of the hriiin with )iijcyH-i:din){ |Birt.-< nml Into
ITOIM or oummiwurnl Iriu-tii hi-twiTn ••"v lutt^nil hulf Mnd the olhi^r, bikI
iio tncta vonnecU'<l with the ^^vrnil [-ranial nrrvi-t>. Talcing the loiigitu*
inal fihriw fiml wtr may in BcciirdiuKv wilh the division junl explained into
,E petal and a tegmental region, coiiiiidcr ihiM^ iw forming on the one band
pednl. and on the other hnod n tegmental K)-»tcm.
Both Nystcms begin, us we xhall we. in the cortex of the eerrhrnl htmU
ihena, Wc shall linve to deal with the tv^cmphy of the cortex later on.
t nay hero say thnt the tin>t broad diri»ion of the whole eiirfiice uf a
bemtaphen ie into four mnin regions; frontal, parietal. occipital, and temporal
(Piga. 194. 195, m»>-
i«x<irrt'i>ii>Ai. FitiRB) or tuk pbdal Hvsrreu.
S 633. 7'A<- piiraiHidal fr<ai. We bare already (fi S7fi) said that ihe
atdal tract of the spinal eiird may he traced to a particular >«£'<■» of
be cerebral cortex, ^^e Hhall study the d<rtaiU of this region, it'll ich ia
often spoken of &.i tite " motnr nrra " later on. but miiy here nay tluit liniadly
apeakinc il is ixirii-tat in jinaition and cor res {Hinds Ut the partu of tbe cortex
giathered rixitid the tiwiurt- uf Hultiud". Fibrm piu»ing from the gray matter
iOf the (^>rtex of ihU n^gion to the white mattrj- Iwlow. and »i eontrihuting
lieir share to thu central wliiti^ matter of iho henii.iphere, convert (^Ig*-
~l), '201) to form part of the internal cxponile, namely, thnt part which bi a
nn»iiilnl aection {Pig- 109, Eyr to Ply) oceupic* the kne«- and strvtchea fur
acre than half, or itcarly two-thirds, along the hind-liuib of the cnpnilv,
tftten the optic ihalaniiis on the inside niid the nuchrus IcniictilariM on the
^Utaidc. From Ihe knee and liind-limb of the capaiilc they paa« by the side
if and ventral to the optic ihntiimiis <; Pig. I'M, 'JOl ), and so contribute to
'fenn the U-(;;inniiig of the cms cerebri. In thus copverging to take up their
Hitiud in the ra]«iilc and in (heir further panaga to tae cnu the fibna
h>w a course of somewhat complicated CTirvatnre, Aa we trace the cup-
nilr from more dorwil to more ventral levels, wc liiid it CODtiuuttllv changing
form 1 the exact shape of the rfl|«ule shown in Fig. 100 only holds good
Kir the level at which the section wa^ taken: it difl'ers somewhat from thai
'II in Fig. llftl taken at a sliKhtiy different level, and sections still more
il or still more ventral wixild present still greater differences. When
examine a series of Iiorizonl.-il sections, taken in sticeeasion from the
dorsal to the ventral regions, we lind that the knee shiAs ita poeittOQ and
bangea in the width of its angle, that the two linihs vary in direction iu
! and in sliape, and that at la^t the bent, flattened capsule pafaee into the
re or leea nwndetl erus hy the rapid diMppearauce of tl>e fore-liiub, and
oonaequent extinction of tl>e angle; so that in one m-owe it ia tlie hind*
OvruKI m * TK^ttvian Uohm-tdctkii, Sbcdon or iuk Ridht U4Lr or tuk Diatx.
(KklanU dai) (SonjUMania.i
TI>aaNQoniihlaliUlak*n*t Uir ltr«l of Iti* kna* or ilw ratwilw. ami 1* tlwntat* hik
bMWOTn Pki«d aiKiini 111 Fl^ IH knd i^*!. U luuvdnaed is iliiuimis iiw rouiw <if iim«im«Iium»^
the pynmlilitl tnU, 0 T, Ojrtiv llulnniiu . St. Baoleiu iitu<l>tiu— Ui* bout tatly mpfttn IB tm
•crtlon: n. imlniDi:)! : tip". (^'. Divtio ram of llii.'(liibnt palUrliwoT Iha luettil* iMMImUria <'
llw clauflr.au. i~£, ibu uimtuI i:a|aiilB: Iii, llio lilaiwl of IMI: ca. ilia aiiUrbv'i
•bwM l» midpt II dlUlnol ■iid Hip nbi<<i fincn Uio l«m|>Dn>4pbi>niilJal Inbr wlikk («■ |gV> It Mn
Illdk*k4 br brakes llnca: Op. lbs ofiilc Inct: Inl. the radot Ihs dMruiidliiit bum ottlM tatn<
niilrlct*'. F, Ibt Ibmli: /*. ib« nid uf ibu aniHlnr iilllsr of iha foriiii in ibr (■••of ilwiMlwHi
( f, oupw oallotum ; (l r. anivrlor [«it of iha o-K<i'iial 1-itw /<■ !• ibt rvntml BwK or ikaan c
RoluidiX Tin fouoa of the Dbiwof ihc vynmuiliil imi-i ivmnopt*! ninniRl)- irllb ■••- ■-.■■> i<t.
add ami and bnuMvIlh t|<Inal iiemmanituf tliiH4tDniii>rCivl with tUs (iMali4 hrn' ".tl
Mrnn, I* tnuiirn br brokoD Uncfc TtiHvanallnMMiiTcrfiiitt ibiothrlaicmalaai- tidi
lllimitbouM In rDipK* uili* Dy.inaot ibn* DhnalneaiiiiiarBl wHb ibc burtuiiial •wiiii« iluwa
In tit- IWiDd Ui«acltlal finurt ibnim In flE. Vn. fflBdlOUtillia ttumt nt 'Ibt iiuni aatethit ut
4aml |an ot Uio wmfioKhocdiiliBl Racl Tlw fliu diMMt llDoi «aiiTpr|iaR In ilw cTWfM* '•
r r, Indlnta lb« roUiM nf [h« csUoial llhiT^
14 gbvioui that the flUroi of Ui« i>yranI<Jiil tract, lik« tliv othvr Sbrw uf lli«
capaule^ are eoniinuHlIf eh«agiii){ ibcnr directioD u lli«r pan ihmngl) itie
I
I
I
Moreover. « bile ibe fibres tnm diflereot purld of the " motor ares "
. defloite |MiHitti>ur in relatioo to each irtber %s they [xua inio the
apanle, their rebUire pomtioiM are nol cooalant, but varr soroewbat. To
ihn point, however, we shall retura wheii we dime to apeak of the fuuciiofl
of thia tnet.
In the crut thcM fibre* rua excluDively In the |ks aiid fonn a compact
■mnd (Fig. 192, I'll), (iron I lying tlie c«ntral ami Inrgpr portioD of the pest
betWMO a HQuill DittliaD )M>rtton ou the Uii«t<le aiid a lateral portion on the
OUUMc Maiulaiiiiiig l.hix jXMiiliun aUmg tliv cru* th«y entrr tlic p<iiu, Iwl
Iwn tb« (mviouiily coiiipiurl Ntrnti<l i* K]>Tit iili, by tlie iiitorlneing traii»A-«no
ibrm of tho pon«, into a immbcr of McatIon.-il luiiKllm, wliii;)i, huvtcv-cr. n» a
wbok. Mill kec^ th«ir cviitntl (weilioii. Th«y f»rin thn greater part of. but
not all. the butidlM aocii cut tntievenely in tmoavonw aooliofw wf the pons
(Fin. ID<i, 11^1). Further barkwani thev bcci)m«' the pyramid of the bulb,
ukTm) give riM in the spinal conl lo the dirccl ami onwscil pyramidal tract*.
TbtM tibm &OID ihe motor area of tht; cortex of the oorebrnm are thus tbi-
■oorM of the pyramidal tracts of tho spinal cord, and hence the wliole
straiMl of (ibm frum ibe cortei dowDvard has been called the pyramidal
tract. We have said (§ 5713) that ne have reafions for thinkioK that the
pyramidal tract in the spinal coni make* oonnectious through the gnj
inatier of the anterior born with the anterior roots of all the Spinal nervei
in succeadon; and nmilarly ve have reason to think ihiit along ita oouree in
the crus. in tbe pons, and in the bulb, before it reaches the oord. the tract
also niakea connectionti with the nuclei of thoae cranial nervea which are
motor in fbiietJon. During tbe ]>a«Hige of the tract thmiigh llie iuteroal
capeule the DhreH dentineit for cranial nuclei occupy the knee, while thoae
belaogiBg to the npiuul eun) run in the hiiid-limh. Some authora limit the
tam pymmidal Iract to the spinal moiety. Mni-e thiv alone r<irins the pyra-
mid ; fnit thii* w undesirable.
Tlib> tract t* well marked out hv the drgoneration method, and thr dcg«n-
eralioii in it is a deMooding one, tlie trophic ccnim of the fibres lieing cells
in iIm- gray matter of the cortex. Removal of or injury to the cortex of
Ihe aholc motor ana gives rise to a degeneration along tho whole tract, and
reoMval of or iojnrj lo part of the area givei; riw to degeneration of some
of the strands. The tract is also well marked out br (he embryo logical
method: the lihrett belonjjing to it aci^uirc their medulla at times diSifrent
from tboee of other fibres.
Anltrior 9r frontal eortieaL Fibrea from the gray matter of the cortei
in front of the motor arvn also pass to the intetnal cajHule, bnl occupy the
fare-limb {Viic. 200, fron). Thence they psK to the crus, of which they
form Ihe ^uall inner, mediao portion of the |>es (Fig. 192, /->.). and fron
the crus |iato into tbe pons ; in transverse sections of the pons tbey are seen
as scattered bundles (r'ig. 191, F. C.) to the median tide of tl>e pyramidal
SbresL But here tlwiy seem to end ; the deseneratiou of the tract is a de-
Mending one, lutd <vuMt here. Most probably the fibru> end in tbe uerve-
oelk of the gray malter, which, a» we nave wen, b abuitdant io the jMMia.
It ia also }irot)atile that tlimugh these nervo-cellii the libros of this tract are
connected with lniQ»Ti,-npc libres pasrinff along the middle orebellar oeduncie
into the cvrebclhim of the opposite nide; but thin hu.1 nut lieeii detinilely
proved.
l^dtrior or Irmporo-ottipital nrtital. Fibrvw fivm the gray mattw of
parts of the cortex behind tbe motor area also converge to the iDtrrnnl cap-
sule, forming the hinder end of the hind-limb behind the pyromiital tract
rFig. IWt, A'). Tbeee fibres also contribute lo form the crus ocrchri, paaiiii
iolu Ihe pes, of which they occupy the outer lateral portion (Fig. If
ns
T1IK ItRAlS.
iV. 0.)- From the cms lliej- pass inlo the woiib. where, like lh« llbni
the preoedinR tract, tlwy np[)ear to end. and probably in » Vikn lowiiil
ThU InicI haa lievii <]«fiLTil>ec) lun one of ofvodin]' define rat inn, Init in all
probability like Ihe precciliiiti ia one oC de^t^nAlug deceDeralion.
The nbove tlirw tracbi n( fibres may. therefore, all Be reganl«d ■• rtartiiy
iVom or liavin^ tlietr trophic reiitres in the <.y>rtical ifrny matter of the htm-
H>here>i, as all helpiux to fonn, 6r«t, tlio inti-'nial ca[uule uid then the |NSif
the cniA cerebri. Hut while the pyrnmidiil tract mnes, in part. In the •)<iiiil
cont, the other two ocaae nt the jhhih. and pmhablr throuKU tho ^ray niautr
of thti poiin iiinkc c<inii(.i.-ti»iin with the cerebellum. Purtber, while tW
pyraniidul tnu-t citming: from the middk' rt-giun of the cortex oocii]qeia
niidille [Mniltiin in the cap*iilc and a niiddli^ |Hviiti<ifi ju tlte cru», the «ystcB
from the I'ronl jinrt of thf cnrlcx uiTiijiir* a front pnnilion iti the capnulc
and an inncT or TDcillan iKwitimi in llic crii# and the sy»ieai from (he Iiia4
part if [he curlex, n hind pi>»itton in the capsule ami an outer »r latml
jtoiiitian in the crux. A> llic thrt« syMWDi paw from the cortex throuf^ tb»
caiNfuIr tu fonn the jh* of the cru», their mrition)> in relation to ench olher
are nhiiUil fnun "oc plane inlo another. .\» the fibre* "prcad out iVom thr
pea through the capxulo to all parl4 «f the cortex, or, put in another way, u
they converge from the corlox through the capmlc to the pv*. they form *
fan, the ciirnna radiatn. which u not only ctirved, but the consUtuvnt part*
of which cm« each other.
Betides these three systems nil paaeing from rarimit rVK)OD« of the mnci
to the crud. there is yet a fourth strand conlributod to t£e pta by tba Mt^
bnl hcmiiphere, though not stftrting in the corteic. fnn thf nHfhu*
caudattui fibres pas' down to the cms. and take ud a pa«itian in tbe pa
dorsal to the tracts just mentioned, occupying a lana-uiapea area imm«diu«lj
ventral to the auhstantia uifrra, and prolubly passins into the suhMantu
nigra ilaelf. Tht^w; uiunut be traced (hrtber down tnao the poos, when
they ap|ieiir to end, tlmugh jKWBibly aome terminate higher up in Ihe mI>-
stantin nigra. Thi.t traet hiw a (lciu?endinE de;i:eneratK>n, and maybe re-
ffantv<l M a tnurt a:ialu)piu.-< to the front and hind oortical tracts though il
ntrgiiis not in the eorlex but in the Hucleuv catidattia; it is not, however,
a very pure tract, many fibre* of the pyraniithtl tract pu«anj{ iotu il in
thejK*.
Tlicae are the main tracts of the pocUl nyMcni. For. though tbr nneUv*
lentSculaii* girc« ofl' tibrea tn the internal caprale, our kli(>wh'<lge of the
Ibrther course of thew ■« at prcn-nt imperfect, and though there wem lo W
lun^tiidinal Rhrew connecting the biilh, the pons, and the [M* at Tariou*
]€«««■. thcmi are not nunicrous, and at nil evente do not form eumtpieuQw
atnindp.
I.0MU1TI;DIXA1, rillltl» UF the TICoHEIITAL ifT)*TKU,
$ 6M. 0>rlietU JUrt*. Although the fibres of the pedal syaleni form, a»
we have tven. tbo greater pari of, they do not form the whole of, the internal
oapaule. Fibrce OOmiOK from all or nearly all [larti- of the eurtex |Jk'UkI>
they help to li>nn the internal cajieule. rlit nol go on to form the wm, but
paw to the optic thalamus (Fi):. 1!J4, './.) and appear to end iu inv gray
matlcr of that body. In Iheir pai»age through the cap«ule the flbrea of tbii
iiAtare from the frontal and jiarietal n^ons of the ci^rtcx. ncciipy the extremr
front end of the front limb m front of the frontal straml of the tibrcs of tlie
of the pedal syuem (Fig. VJ'J, Tb.). The fibre* from Ihe on'ipiUil aiid tern-
-nral rc);innit — tho*e fiirin the oodpital nigbuu being the moat iiiiRM-rou*.
'-lUecil being vwy cwupkuouc — occupy the extreme hiixl end of the
t
LON'OITCDIltAI. riBRKS OP Tilth TBOME.VTAL SYBTGH. 719
hfod-limb of the capsule. Kebind the teroporoHicGipttal division of the pedal
■mem (f'*iK. \iH>, Op}- ^-iuc^, as uc shall lee, w« have reaiton to aaeodau
the (x.-eiDiial region of the oonex uiih vitiioo. thv fibrv* ihuii nidititiag to for
from) tne thabiroui through tho exlri-tne btiKl-liinti of tti<- ('ii|^<i>ulc Irom (or
to) the occipital MiMex hnve Wen fiilleil ihi- u/ilir railoition.
All the aoovi' traci.i of librr*. Uioujfh joininji; tlii; llialnniun aii<l not potting
on to the f)M, (nkv purt in ihu forinaiiou uf liit! internal ai|Mul«t. Btit ■ con-
•idenble mimberur rihrc* I'dmiufj; from Ihr ti-inporo-occipital rveiiui nod
etpecially (rum the letniionU region iiiiw to tbr thaUinn* without joining the
capMile: thejr ban rentnil to kikI U-hind tlio pm m thii pluugCN tnto the
hembphere tii Wcoine llic cni«uli.', urid s<> r«icli th>.- thiilikniii)>.
We tniiy Iwrw p-rhapii liivcrgc for a monH'tit to point out the contnut
betwreii the optic thnUuiiiv and the corpus strialum, or at lcn»t the oucleua
caudniiuL Thi> fonncr don not contrioule lo the pedal tyxtcm, lh« latter
nupplic* a marked contribution, The former receives Ubres from alt purta of
the cortex : there are no luch special oontributions frotn the cortex to the
latter. .\nd lhi« differenoe accords with the experitrncc that when partit of
the cortex are removed, or are eongenitalty absent, no defeneration or want
of developntenl is ohserred in the corpus striatum, while degeneration or
want of development is obwired in the optic thalamus as well as in parts of
ihr pedal and le^'mental syatCBW. llenoe, while we may regard the opiic
thalaiiiu* as nn intrrnieiliale mass of Riay matter receiTing tibres fn>ra the
cortex, and cnuneeling the cortex with lower pans of the tegmental region,
tbtf corpus strialom, Appears mthcr to be analogouH to the cortex itself, (o
be a special modilicatioR of the cortex, sending fibres down into the pedal
l^Vtcn, but itself receiiing no special tract* of Rbrcs from the cortex. Indeed
wcmay probably regard the corpus atriaiuiu as tlw oldem part of the super-
ficial i;ray matter of tbe heiuiiipliere, the more onlinary cortex being a lattr
develupuem.
Tbe tef;mentum pniper, lying vcutivl to the hind end of. and behind the
thalamu«. in which reui"'!, a> we bare sct^n. the conspicumis rcil nucleus is
•iluated, t* thus, by riuinon of its connection with the ihalamuH, indirectly
connected with the cortex. But Ixvidcs this, it has direct connection* of It*
own with the cortex. Some of th« fibres of the optic radiation, as well a«
fibm from Uw temporal and occipital regions described above u sweeping
round the baae of the iiitenml cansuh-, are saiil to pass not to the thalainiis,
bvt lo the tt^mcntuni. Other fibres fmm the frontal and parietal rt^ioaw
liavfiting the lenticular nucleus in ihe sheets of while matter tnlittiDg tlM
Bod*lM into parts ar« also said to reneh tlie legmeotuni either by piercing
thiwigh or byswaepjng round the interual capenle. On tlieir path iltcM
fibres are, according xa sonw obMrverv, joined by fibres coming from the Icn-
lictdsr niteletis ilself, and possibly firou the caudate nucleus, wbtcfa fibres, on
the Tiew that these nuclei arc modified ooriex, may also be ooosiderHl a»
conical. Thus the fore-part of tbe (egmenlal resion is brought into ample
connection with tl>e cerebral hcmiipheri- partlv bv libnr* joining llie thala-
mus, partly by libres |m»ing directly to tin- tt-giuentum pniiwr.
Tbe mode of degeneration of tbeee cortimi hbnv of the tcgraeiilal system
is at present a matter of ditjiute. Nor i> the general oatiin' (if tlie tibrvs
concluKivety detenuinetl, though it in eem-nlly Mip[K<s"d that tltey cany
impulMd from the thalamus and parts oTthe ICEmentum to the cortex.
I 63S. I» the tegmentum from tbe subtlinlmic region to the bulb the
reticular furmatiuit u>, as we have scrn, more or less abundant ; this, and the
occurron'r of variouB bundloi of fibre*, gives the region great complexity ;
and «e must confine ourselves here lo touching on one or two important
looglludinsl Mrand* nhich iravcm it-
11k tufterior peduHcit af tlu eerebeUma in on« of the mosl impoflant ofl
llieae. TkU mi each ude nf n bundle nf fibres which, Mkinc origia chiefly
from the gray muli^r »r i\\r iiucli'ii* dviiiutii*, und the tnuiUer noigbliorinit
ci>lleciii>n of ft'iiT >nalU-r, liiil iilii» in lutrl Trom th« tuperficiKl zr%j nuOcr,
leuveo (be ccrubulluin in fruot of, itnJ l« iho niudian side of tae rotifortii
IhiiIv hdiI ]<«i!M(v forwnnl lawurd the ciirjxirii (jitudriffemiita to converge vith
ii.-> telliiw. At RrM ih« two pmluncle* am viipcrficiB) Kiid doraal in pontioa
(Figa. 189. liiO, .S'. f.) iind the spnco bclwocn lh«m i* bridged over by tbt
vnlve of VteiiSA'n* (Fig. li"), Vln); but, still conv«rgiog, thej mmu tiak
vwiirally Ih-im^jUIi the ["wtwnur corporn qundrigrniinitaiid at I be level of the
juticli'xi Ix-rwix-n the itnterior itiid pintt^rior corpora ijuiulrigvimoii meet t»l
dccu»iai« vcntml lu thtiee bodies iii the VL-nlrnl region of tbu t«gnieiiluai (Fi^
li^I, iS. P.). Beyond the decueaation (hey arc coDtiiiued forwanl in the tcf;-
nitntiun ventral to ihe an(erior corpora qiiadrigeminn u* (vo straurla, oik
on vaeh side, which appear to end in (he red nncloi.
In ihU nny ihe peaunclea connect tert&in parta of the cntv ina(X«r of lb*
n<r«bcllnni with the tegmental region, and more particularly with the nd
inicleiu, and thus indirectly with the simctures with which that region u
itMilf connected.
ThetiiUt. This, at ve have iiecu (J, GIZ), takea origiii in the bulb, b tbf
intcr-olivary layer bulweeo the inferior olivw, frum fibres which arc derived
thivugb the nupra- pyramidal or nejuury decuieatton iVoni the Knu-ile ud
cuncate nuclei. Fnmi thin origin it patwa forward on each udt iw a Hal
band into the lejfmiMitnl region of thr pon*, receiving scceflBtODs from tlw
superior olive and ')thcr collecticin.i of grav matter, and dividing them into
two straiidn. the mtflian (FigH. 1!)0, 191. Fin) and latorat (Figs. l!tl>. 191. fl
and Fig. Ififi, HF) filial. Tlif lunral itiviKinn end* i>artly in the gray
matter of (he posterior corpus f|Uadrigotninuni. and partly in the while
muter nndi^rlying (Fig, 11';!. itm) the luitcrior corpus <|uailrigi.<minum ; the
median division puseing further forward appears partly to eiid in tlie gny
matter of the anterior corpus ()uadrigeniiuiim, but [lartly to be condnnca on
lo the subthaluDiic region of the tegiueiitutu ventral tn the tbalamiM.
to the tholaiuui, and so to the cortex-
Tke toiufitudinrU pontunor bui>dlt4. In a transverse wction ihmagh '
funvpart of the pona at the level of the posterior corpora quadrigi-reinnl
rather conspicuiua bundle of longitudinal Gbree (lulled the loiigitndini
poaterior bundle) in hkh <in each aide, cut transvereely, in the dorsal n-gifl
of the tegnientum junt ventral to (he nucleus of the fourth nerve I Fig.. \'J\ '
Traced backward fmm (he Hijiii'duct l>encatli the fourth ventricle, it bcconi
len eanapicuouii (Fig. 19U, I) though niaintuiuing ii^ position dorwl to ibt
reticular furmalion.and at llic hind end of the hulb up pen re to be a continn-
ation fomard of thnl•l.^ libnv, "ground fibnw," tif (he anterior column of tbt
cord which probably f^rve n» HuccnMive thurt luUKitudinal coraniiamna
between the si^aieats of the cord. \^'hilc the vomewhat aualogoue lillei runs
ventral to the reticular formation, this |KP«t«rior longitudinal buudle ruoa
alway:> donwl to llitit ctriictuiv. It may be traced fomunl as far as tbe
nudeui of the lliird tiervc, as is seen in traniveiac aectloiu lyiug immediately
ventral to that group of cclU i Fig. 192./). but it» further counet-tious f>rwanl
have not as yet been determined. It is relatively more pniuiueui in the
\nytfr than in the higher animals, and its fibre* acquire their roealulla rela-
tively early. It is supftoeed to be connected with the nuclei of the nerves
t^reniing tlw niusolce of the eye, and so lo be eoncenied in the marements
\m that organ.
TrttrUfrnm thf ixtrpon <iHailri^mu\a. From eoch corpus ((uadragcminu
tliere puMw obli<piel}- forward and downward on each aide a band of filir
TBANSTKBSK OR SO-CALLKU COHMlSSlBAb FIBBBS. 781
ineded wilh tbe Ktar matter af lli« coq>ua oaA kwtua ta iu hrafhitm.
« anterior bracfaium (Vig. I'J'J, 7Ai), as *re shall t>e« in iltaliug witli Ihv
^»tic uerve. joins ilie latcnil ii<r|iii!S geniculaium aiid )ivI|ih t<> I'oriu itie nptio
tract, Irtit »boi«orit£i^let'|'er Iviiif; libree proceed lo ibe octipitat oirlpx tltnu-
ing part of ili« tibr<-!i nhich *» liuve (§ (>34) described a* riiiHiiiji from the
ooci[Htal cnrtex to uml jwHt ibe thalaniUH. The tMHlerior brui-liiuiu pBnac*
ta tue Rwdiau cvr^ma ({i-tiiculttliim ; biivinjr receiTed tibri-n tVotn nud nromblv
E'vrn fibnt* ujitotbat l>i>dv.ii ia i-nuiiuuM onto the li-i;iiiciiiiiiii,iiiid accord-
g to tome ■iithnm through the legm^nlum by th« bind part uf tbi^ bind-
linb of ilw iiiienml cii|wulv t» iho (fnipHrnl n^nn i>l' ih<> cortex, mingiint;
tn hs couiM Willi libroi rrom ibe thalamus.
TIUXSVEIISR OR MOCAU.KD (t>MMt880RAL ftBIIDf.
S 636. Tbe two chief mrmn nr« thow on the oni: hand bolongiDg to U»e
cerebrum, and those on (be other htind bclongiDK !'■ the cercbrtltim.
/n the ftTthrum thi> mott imposing mius of irumverw tihr«s from the a»T>iu
taOottim. Starting fram the cortex in Dearly all parts of ibe hemispbere.
ibo Rbrefl convert loirard the thick body of the corpu! callosum placed in
the middle line, aud llicnce diverge to nearly all parts of tbe cortex of tite
haniipbem on the oilier side, interlacing in tbeir course with the cortical
tbrea of the pedal aud le^iiienlal systems. It is Buppowd that by lueaOB
of theae fibrea. each part of ibc curies of one hemiepliere w brought into
'i>n with the curreeitondin^ part of the other bemisphere.
s thcae callcael nbrea trom ooe lieiiiixpherc to another, the vbite
)er of each bemispliere contains fibres cbIImI " asttiiciaiion Rbrea," {Mannj^
fh>Di one cnnvoluiion lo aootlier of the same heTiiis|>here.
The iinnll antrrinr tvhil< mmtniMuiy though it in placed in the front i«Tt
of the ihinl ventricle (Pig. 198, A ) and, in jxirt of it« counc, lies along the
ibalaiDiiB I Fig. ll>'>, (])if i» really a coinniiwiirc of particulnr parl« of the
cerebral hrniiFpbt're*. .\ [rurtion, very Hoall in man, boEongn li> the olfac-
tory tract : the reat takes origin on each side in a limited jxirtion of the
cortex (Fig. 194. Cc). which we shall Inter on speak of as the tempom-
vpbetXHdal convoliitioa and in which calto<«l tibm are deficient, whence it
arobee forward throng tbe globus pal lidns. past the thalamus (Fig*. 201. ra,
1&&, (bl to the front nan of the third ventricle. It may be remarked that
ibit ootnmifisure is still tbnnd in those lower animal!: which do not poeaoro an
obvious corpus callosum.
The ainall poeterior commiMure niay be regarded ns mainly a commiNRire
between the two tliolami, but It alto belp* to unite the tegmentum of the two
aidea and some fibres are Bai4l to paas on each nde inM the hemisphere. The
middle or soft commissure of the third ventricle ( Fig. 193. c). thuugh it con-
luiiu IraDiTerse flbres, ia in the main a collection of gray matter, indeeil. a
part of the central gray matter.
Tlw/orMLT, to^ber n tth. al all events, part of tbe septum lucidum which
join* it wilh the corpus cullosuiu, must also be regarded sa a comminwral
structure. But its relations are peeulinr; for while, l>ebind, tbe direrxing
poatsrior pillars begin in tlie cerebral liemttpberes, uamelr. >n the «alu of
tba dtacendiiig Ikoni uf the lateral ventricle on each tide, in /Vont the anterior
pniail or eolumn*. Icnviug the cerebral hemispheres, pus nloug the lateral
walls of the thir«l ventricli.* (Fig. 19M, / ), and apparenttr eml in the gray
matter of the corpora ulhii-antia. WhetlKr tbe band of fibn*. known as
Vic<id'Asyr'8 bundle (Fig. UU. I'fr), whieb running in the lalcral wall of
the tnird ventricle leads dorsully from lach corpnii alliiciuu up to tbe anterior
TBI BB&IX.
nuclMUof the llixUiuu*. li reatir lu t>« ronxiilfrtMl lu a ounliiiunli'in af iki
fornix it dbptitorl ; it mar more prolmbly bi; ivgartlc^ w n pari uf tlie tjtum
spoken of itnnvc a* oiinecting tnv vurtcx ifilh tliv (hnliimas.
h tlte eercbrllam true onimisaiiral 61>re«, «rv supplied hj the auiUlr
jwJuhoIm ; but by nv menne itll lh<: (ibrea uf thcev p«duDcl« w of Hm
nature. The Hbrat of the tnidillv poduncle, in oootrnci to those iirtbi up»-
rior peduncio which Klnrt cbieflv from the nucleus dentatui, or other iotcral
jtntv iDRttor, nnd U> th<i«e of tite inferior peduncle which start chicDr fraa
the superficial gray mnlter of the vermis, appctir to start from tho HiperfieU
gny niatt«r of the wrliote anrfnoe. from that of the median rermts at well u
fnim thai of the lateral hemispheres; ihej lhu» form the greater part oflltt
ceDtral while matter. Sweeping down into the pong, they form the trau-
verae fibmi of that body, interlaeiiig with the longitiidioal libres of the crural
^ytiem and iotecniDgliDK with the abundant gray matter.
Of theae tnuisvene fibio ot' the pom, a ctrtaio number are trulr ocmiiw-
fural : they mako no coDiieelions with eells in the poiu. but coutitine tluir
way uiibrohen across it ; they »larl in die superficial gray mutter of uoe ndf
of till- c-efebelluiu and end in the superficial )cruv raatlvr ai the other lida.
tho parts or the gray matter thus united being probably Liirrtupondiug put*.
The niDBt vetiLrally place<l trannvente Obrca of the potu which form a Mpcr>
ficial layer of white matter, free from gray matter {Fig. 189, tr. P,) an
i>robably of this nature, as arc alao the tninsrerac fibres plued mou dorialty,
just veutral to the tegmeutal rcfpon.
A lar^ number of the traii:>vcr»e fibres arc not of tbb uaturo. 'They
orow from one aide »f the ocrcbullum to the onpoeita ndt of tba pooi. b«'t
end in tbc ponM apiAreiitly iu the ner^'o-celU of the grajr mattar ; and tl b
soppowl, that by thcM nDrv«-e»il» iboy arc briHigbt int*) counoctinn with th«
longitudinal fihrm of tbo pedal syflem and thus with tlw oerv^rum. They
are Iransveree a|ipcndag<>* of the [ledal system, not true commtaMiral fibns
though thev do eroe* the median line.
It N further supposed tlial other tibr« of the middle peduncle reaahiag
the pons do not croffi the median line, but k«ei>iug tu the same aide and
chaapng their direction, tnke a loni^itudinal upward course either wUh or
wlthocit the iaierveniion of nerve-cells, and so make their wajr to the leg*
awntuui. llul this is not certain.
We mual ubu consider as oommissaral stnictnrea the numerous ftbres
crowMug, or serving to fonu the raediaa ra|kl>e in the bulb. Thin raphe:, with
similar commissural fibres, is present in the tegmental portion of llic puu,
and, indeed, in the tcgmenttiui itself.
Fibres also oross from one Aide to the other in coDDOction with the cranial
nervnt, but theae •» well at all the ti^acta specially ooonected with the cranial
nerves, ineluding tlte ul&otory and optic uervet, had better b« cooBidered
bv ihcmselvvt.
SirUMARY.
^ 837. It muv perhaps apficar from the foregoing that the brain coansts
of a number of isolated maasos of gray matter, and some large, some small,
OOQDected tccethor by a multitude of ties of white matter arranged in per-
plflXlDg iotncacy ; aixl the nd'Iition of numerous ooUectiuns of gray matter
aod strands of white matter of whirh we have made no mentioa would »iill
Airtber iorrrasi- the perplexity. Nevertheless, a systematic arrangenent
mav be rec<3gni]:e<l, at least to a certain extent.
The least conspicuous, but perhaps iu point of origiu the oMnt part of the
brain, teems to he what we have called the central gray matter. Thia soems
SL'UltAUY.
783
iy t» m bci] fur the development of tin nuclei of tli« cnuid
I Ui« oTJitrnl gnif ranttor wul more or leu a«odi»tmI wiLh it ramea
irtiAl we liuve called the togmuitiil region, of which tbe micular tunnatjon,
eomiog into prominence in iht bulti end continued oo to the »ubtbalamic
nigiaii, Ibno* oe it were the core. Belonging to the l«Kmeatnl Bfsleui are
nniDdnias mMtee of gray natter from the conitpicuipiis optic ihnluiuui and
th« red nndetti in front to the MVcral nuclei of the bulb buhiud. Tbia com-
plex tvgnientnl ay»tem, which may perha|M be regnrdvii tu a more or leu
eontiniioiis column of K^ay iimlter, comparable to the gray niatler of the
B(Hiial cord, serves nit a sort of bnek booe to ibe re«t of the central iiervoua
■ntetn. With tlie s[hiui1 oord it ia connected by various liai, lieiii<leit being
aa it vere a cootinuatiou of ihe spiDal grey nutter, and around it are built
H|> tbe Kr«at mius of the cerebrum, and the Hmaller but still larj^cr maM uf
ihe oeivbellum: the leas important corpora (|uadri)[einina we may for sim-
plicity's Mtlte neglect.
At the tiiud ejid we find varioua jmrta of tbe iipinal con! becoming coti-
uected with ibijt tegmental system, either panio); into it and becoming, a* far
as our preoent knowledge goett, luet in it, or iu^iplyingntrundsurtlbna which
paiiing into it Income through it connected with other |mrU. Thai the au-
lerior column of the oord exoluslre of the direct pyramidal tract, the Utcml
ctilumu exdutive of (he criMwd pyramid:!] nnd (xirvbHllur trtuttd (aiidpiiMibly
the an lero- lateral avcciiding truci), tu;;i^iher with jiarl >if thi- jiiwtrriiir column
rsar to join the teemeiilal NynlL-m, while piirt of the jwist^rior cnliimii, after
rday uf the grncile umi curi<-alu nuclei, |ubu«> through tht- nyntcm lu tlit-
fillet dnrtiiKil for various ittriii;tiires.
At tin- front end we lind iill parln of the c«rchral corlvs (,thvugh some
ngiooa. namely, the rnnipDrD-nocipiliil, lo it gr>.iit<ir extent than othemi, con-
nvCied with tbe thalamus aud other |Mirl» ol the tegmental ayslem ; and. at
a* ire have seen, tbe corpus etriatum rany postibly pooseta like connectiotu.
The relaliona of the cerebellum to tlit» spritem are notable. On the one
hand the cerebellum ia directly connected with the system, partly by fibrce
which paas from the bulb to jwn the reatiforra body or inferior peduncle,
partly oy the superior peduncles which, as we have seen, are in n measure
lust iu the tegmentum, and partly probably by fibres of the middle peduncles
also making connections with the tegmentum. On the other baud tbe cere-
bellum form* around the te);mcutal ityslem a great junction between the
H|iinal curd and tht- cerebrum. Ti> the Hpiual cxtrd it ii joined in a direct
manner by iliv oerehr^lbir tract and |>oa>ibly by ibe a ntero- lateral asoendiog
tract, and in an indirect manner by the reluy of the gnieile and cunnte
nuclei. To all partn of the cerebral cortex, it up|i«ani to be joined by tboae
ooMpicuoua Mtraud* of tbe [ledal nytitem, which. a.i ire have seen, end in Ihe
pooa. and tlicre miikii ii>nuectiuiii< with the fibroi of llw middle jiedunijc.
And we may iK'n; jN.-rha|ici tviuark that while tliu connection bclw(.'en the
etnbrum and oereWlam in whidly a erUMetl one, each cerebral heininjihere
being jojned with thvoppiMlu half of the cL-rel>ellum, tbe conuL-ctioiu between
tbe kpinal coni and the o<Te)>elIum are hirxelv uncnewl one*, that by tbe
ocrvbeltar tract being nholly uncronooil, and that with tlic jioatcrior column
by the relay of the gracilc itud i-oDi»tv nuclei l>cing in [<«rt uncroaied.
Thus the cerebral oorlcx Ium a double hottt, so to s|>vak, on the real of iha
cmtnl nervous «}-«t«m, fir*l through the the Kwmautal systom, and soeoudly
through the oerebcllar junction. But in addition to this there is another tie
betweeti the cerebral cortcs and the wliolw length of tbe oersbro-epinal nsia,
or at leut between it and tin- whole scries of motor mechanisms in sueoosston
from the nucleus of the third ner\-e to the nucleus, if we may so call it, oi'
7M
THE 8RAi:r.
the nbierior rixji of the coccmwl n«r%-e, iiura«ly, ihc fcrt»t |iyntmUUI Inn,
which thiiM upiwura m a miuMiiEd}; *iipera<iil«d lo all tba rat uf lb« «enuid
iKrvoiis 8}'8tem.
Wh«n the cen'.bral hemuphrres tire reraorecl thit prntmidal trart &Bi
airny aa ilrN» nlsii the peilnl syslein Iciuling rn>in ihc ccrchrum tu lht> [uru,
but there rtill remninfi ihr legiui-jital :>_r»lcm with \t* ifrcbellar and otbff
iidjuncls iind this, n^ we shnll see, eoni'tiiiiU'a n nervoiiK lunchioor;, capaMt
of ciirryitig luit excealinglr compHcntcd acts.
Ok the Phknomisa RxiiiiiiTKu UY AN AxiUAL DKritivKd or na
Cbrkbral Uemhpiieres.
{ 038. The cerebral heniiHpheree, as we bare more than once iiisistciL
•eetD lu Btnuii apart t'nini the reel uf tlie brain. Id the ciue of aomv antnuib
it is pnwible lu remove the eerebral bemiapherea and to ke«p the aiiinuJ doI
ooly alive, but io good health liir ii loii(c t»ii« — days, weelcH, or eveu nicalh«
all«r the operation. In xui'li raae we are able to Mudy the Itehavior of u
anininl posM«iiinK no uerebrat be in sphere*, and M> iiimpnre it with thai of
an intact animal. Sueh un cx|H^rimetil is best i-arned nut on a frog. In
this niiinitti it is i^ompiinilivelr vn»y t*> reoMVO ibc cftrcbral beinbphmK
irichiding the pnrls ciirn^p'iiiiliTig In thf rorpora striata, leaviog bchinil
intact itnd uninjured the ojiiir thntnnii with thtr opiii' nt'rvcs. th« oniic lohf*
(or repre»<^ni«tivi-s of tlic ooqicrn iiuiidneeiiiiiin), the small eerrUiflura, aiid
the bulb. If the animal Ik- cnrrfnlly ft-d and attended to, it may be tw|it
alive for a very hing time — for nu^re than n year, for invlanM'.
The ealicnl foot nboiK a fm^ Inciting the eenibral liemisphercs is that. ■*
iu the case of a frojj deprived of il« whole brain, the siftns of the workinirur
an inl«lli(cent voliiion are either wholly absent or extremely rare. The
presence of the bulb and the middle parts of the brain (for «o we may toe-
venieutJy call the cerebral stnictuive 'ying between tl>e verebral benmpher»
and the bulb] insures the healthy action of the rascubir, retpimtory, ais)
otli«r iiurritive systems; food placed in the mouth is rMdilyanil easily swsl-
lowed; the auimni when stimulated executes vnrinus movements: but if It
Im )eA entirely to itavlf, auil care Iw taken to shield it from advenritiuv
stimuli, either it reinainN |>crf<'ctl^- imd pennaDciitly ljuieacent, or thr appar-
ently spouluneous nMvom(!nt:> which It carries out an so fijw aotl ■<( Hmitel
as to make it very doubtful whether they ran fnirty )w called vuIitiunaJ.
Such a IVtig, for inxtjincv, nfl«r heing kept alive for some time and made lo
exhibit the phenomena of whicli we an- aliout Ui s[M«k, has been placed on a
table with a line drawn in clinik aroiiti<l the urua covered bv iU body, and
lef^ to ittclf. has KnW']iH'Dtly been found dead without havTng stirrM oul-
sidc the chnlkitl circles
Wc must here, howrvcr, rejicttl Iho caution laid down in § S83. as lo the
ultimate eflects of an upcnition on the <^vnlral nervouH syvtem. The longer
the frog is kept alive and in gotx) health afler the removal of the cerebral
beiDtSpnera, the greater is the tendency for np{mrently spontane^Mis more-
meots to show themselves. For days, or even weeks! auer the operation
there may be do agon whatever of the working of any volition ; but after
ibe lapse of months, movements, previouely absent, of such a cbaraeter as to
auggat that tbey ought to be called voluntary, may moke their ap|>eanuK«.
To this point we shall return, but may, in the meanwhile, slate that eveu in
their most complete ilevelopment such movomenU do not negslire the view
that Ihe troz in the alMence of ihe cerebral livnii^pbcres b wanting tu (rhat
, we onlinari^ call a "' will."
WITIIOCT CEREBRAL HEMISPHERES.
786
I
Wr have t*xa (httt a froji from wliiuh ibp nhotc brnin lin» t>«;n
Bvdi. KiMi ihp spirul cor<i only kfi, npponn ^iniilnrlr 'tcvoiti "f » " will ;"
^tlt iht? I'hcnompnii pmraUii by n frog iiQwcwin^' Uic mi<J<)lv |>ortioni) of the
bmin fliiliT wii]ely Irum th-.«« iirvfciiieti br n fm^ yn^nt.Mig n tpinml cord
oaiy. We m«y, pcrhnp*. Iirondiv ilowribc tlic bclmviorof n iVog from which
ibe cerebral hcinW|>ht>iTfl oiity hiivc been remorcd, bj* Miviac ihnt such an
aDinial, tbuugli exhibit in)* no epoiitaneoiis movements, cao by the ttpp lien t ion
of appr»|jriate slimuti be iiiduceil to perform all, or nearly all, toe luove-
meats which an eu(ir« fmt; is capable of ex«ciititi);. It <.-aD b« inad« toawim,
la leap, awl to crawl. IJetl Lo itwlf, it asiunioa wltot may be called the
natural posture of a lro|{, with the fore-limbs erect and thchiri(t-linibedexed,
•u that tlie line of lh« uxly makes nu angl« with ibe surface on which it is
rtviing. When placed on il« back, it im met timidly regains this natural f>as-
tnre. When placed od a board, it does not fall frum the board whtii the
]mUtr a lilteu up so aa to dbplace llie auiinal's centre of iiravily ; il cniwla
up the board until il gains a new poution in which iia otntre of gravity is
Ratw^ 10 ila proper place. Ita moveiuenta are exactly ihiwe of an entire
tng, except that tiiev uev:d an cxienial stimulus to call rhem furth. They
dillvr. moreover, fuii<{umeniully from thoae of an entire froj; iu the fullowing
iropocuuit feature : ther inevitably follow when the lUmulus is applied ; they
eOBM lo an end when iheitimiiluB ce«ae« lo act. Br continually varyiug ttic
inclinaiiou of a buunl on which it tii pluceil, the frog niav be uiaile tu OOD-
linuv crawling almost inde6uitely ; but directly the board i< nmile t» assume
■ucb B poaitiofi that the body of tlie fniir i* in equihbrium, the crawlinjf
ecaiaa: aiid if the {KNiiioH l>e n^it diaturlwd the animal will remain impassive
Hul quiet for an almmtt indefinite time. When thrown into water, ihu
rivalure liegins at onre to «nim ntioiit in the mo>t regular ninnnrr, and will
foniiouv to swim until it is exhausted, if llicre be Doihiug pn-Mciil on which
it caii come to mt. If a small piece of wood be placed on the water the
fh]|> will, when it comes in contact with the wood, crawl upon i(,and»u
flone to rest. If disturbed from its natural posture, ns by bcin^ placed ou
Ua back, it immediately slruf^gles ta ngain llial p^ture ; only by the nppli-
calioa of continued force can it be kept lying on its back. ^Sucb a fro|(, if
its flanks be gently stroked, will croak ; aud the croaks follow so regularly
and surely upon tbe sUokcs lluU the animal mav aliiioet be played upon like
a musical, or at least an MMMMic, iustruuieut. Moreover, oruvided that the
ofillc nerves and their artatwemeDls have nut been ii)jure<l by tlte operation,
tne ■noTementaortbeanitualappear to beinflnenceil byli^'hi; ifilbeurf^
to moTS in any particular direction, it iceuis in its pro^rvos to avoid obata-
cio*. at teost sucu as cast a stixiuj; aliaduw ; it tunui its course to the riifht or
leA,orsoiDetimea leajis over the obsiaiele. In tiict. even to a careful observer,
the (lilferenees between such a frvi)c and an eutire frog which was simply very
stujiid or very inert, would upfieur nliKbt and uniui{>urtam, except iu tlii^i.
that tbe animal without iLh cerebral heuiiapheres in obedient to ever)' stimulus,
aud that each ttimulua evukw on ajipropnate movemeut ; whereas, with tbe
ODtiro animal, it is imfMaeible to predict whi-ther aiiy result at all, aud if si>
wbat roult, will follow the applieatioii of tlits or that Htimuluo. Both may
bo rvganleil as maohineH; but the oiic u a mnchiiie and nothing niurv, tbe
otiwr is a machine governcil and checked by a duminunl volition.
Now such ronvcmeni* as crawling, leaping', swimming, and, indeed,
a* va lasTO almdy urgi-d, to a grtaier or Win extent, all bodily movv-
nieiits, are cnrrie<l out by ni«iii»* of cni)rdinntu nervous motor impulses,
inOueaced, arranged, an<l p>v)Tnc<l brcoineidcni iriisory or aflerort im pulses.
MuBculor movements are iletermincd by nHerenl inllueiKOS procoadiug from
ifca noacU-s and couttituting the foundiitiou of iIk muMular seusa ; tiey arc
Ut
780
THE BRAIX.
iil«o ilifocled by means of nOVrenl ioipukes piMiiij centripelally aloOf tlu
fCDMiry nerves of Lhei^lciii. the eye, lite ear, and other ur^iM. Inij<>p«idiii[l|
of the pnrticulnr afferent ini|itilM«, whii'li uetiuft a-i n Mimalus call f»nk lu
muvi-iueot, very miiny other afrnretil im{iul»eit nre couct-riM-il in t\» pom-
liun and oniirdinntii)n of the renullani iiuMor impulto. Kv^ry ImhIiIv iann>
tuenl Biirli aa thoee of wliirh we are H|>eakinx i» the nork of ■ nK>r* or In
comiilicated Dervoiiit nieclianixni. in whii-h iliere arc not ouly central iiiJ
eAerent, but alsi> ntren-nt ftietont. And, fnilting nuidr the ijuivtion of nn-
■riouBiieM, frith whk-h we liuvv here nn occiisiixi lo deal, it i» evidvni ihsi b
the frog ijeprired of its <«re1irai lii-Tiiii'|ihi-mi all thon; lactnn air |
the ufliinent no lew ihnii the ci-utnd nnd tlif clfurenl. TIte nuiehiort' i
the Uf^eMKry and uiunl lintlily nutvcnH-ntu in {ircwnt in nil it* cimiili'iitirw.
We may rei^rd the thnrv. ihercfnr>'. which the cen^liral h«mi<>iiher<.-<> uit a
cx<.-uultne the movvmciil^ of which the entire onimid i» c«|ittt>le, a* lhai4
putting uii» miichinery into action or of limiting its prcviuuii activity. TU
reUilinii which [he higher nervous choiigv* concerne<l in volitioo bcAr to lUl
mnchinery nmr be compared to that of a EtiniiiliM; uliniys bearing in fninl
ibat llie ('fleet of ft slimulus on a nervoiie c>.iitre may be either ta Uan
activity, or to inemM, or to eurb, or to »top iicliviiy alremly prewnL Wi
might almost spmk of the will aa nn intritieic fCimtilue. Il» operati-iot on
limitc<l by the ninchinerv at itn eommanil. We niuy injur that iu ibe in^
tho action of the eerebrai heniisplieres in giving shi)|M> tu a bodily laoveranit
is that of thraiving into activity pnrLleular piiri« of ihe nervous nuieblncry
Htuated in the lower pnrt« of the braiu and in the spinal cord ; preciMlr iIn
same moveraeiic may be initialed in the abaencv of the cerebral bemiipDmi
by applying $iie)i aliniuli ua nball tbrow precisely the oatue parts a ikai
luachincry iiilo the oaine activity.
Vi-ry niurked la tbe conirust ticLweea the behavior of auch n frog wbk^
though deprived of iut <^e^L'brnl heinupbftrei, Btill retains the iittu>r partai^
the brain, and that of a fmg whicli poaseam ■ tipinal cord iinly. Tb« UUii
when plooeii on ita back maki'!< no attempt to regain it^ mirinnl jxMluni; ia
fact, it may be Mitd to have eonipletely h»t itA normal |HMtiire, for una tthai
pineol OB it* belly it diK'.i not Nland with il^ foTe*fvcl erect. 0* dots the othft
nnimal, but lie* Bnt on the gnxjnd. When thrown into water, instnwl if
twimming it nnkM like a lnnii> of lead. When pinehnl, or olhcrwim- »liii»
lated. it doM not crawl or leap liiraard; it iiini])ly tbruw* out it# limlab
various wayf^ When il< tlank* are xirokeil it doei not crnalc ; nnd niiraa
board on which it is plac4.'d i* inclined Kulliciently tn difptacs it* centre of
gravity it makes no ctbn to regain iis balance, but tnWu oQ' tlie boanl likca
IifekM mast. Thmigh, as we have m«i), the \-nri«us part* of ibc spinal cord
of the frog contain a large amoimt of ctmrdinaiing nuicbinery. to that tbi
bniinlen frog may, by appropriate stimuli, l>c mailc to execute various par
jKveful coordinate movoiiienl^. yet th<'»e arc very limited compared aitb
those which can lie similarly carried out by a frog posMwing the middle oaJ
lover purls of the brain in addition to the sjunaTcnrd. It is evident ibu •
great deal of tbe more complex machinery of thi< kind, specially all tit*
which bos lu deal with the body aa a whole, and nil that which is entttiuriMil
WttJ) e<inilibrium and i^ 6[»ecially govonutl by the higher senses, ti iraird
Dot ID the Hpinal eord. but in the brain. Wo do not wish now to dtM;ua> iJic
details of this maeliinery ; all we desire to innst apun at praami b tliol. ia
the &og the nervous uiacbinery required for tho execution, as distinifutthrd
ftiini the origination, of bodily movements even of the tnosl cyMnplicnlcil kind,
is preaent after complete removal of ibo cerebral hemispheres, though tiim
mnveoieuta are such oa to retjuire the cooperation of highly diffiuvatiaivd
aflbrent iinpulses.
WITHOUT CBHKBKAL II £UI3Plt ERES.
7S7
\
{ 640. In wmrm-liloodfil aiiiiniiU tlic rrniovnl of the cca'1>ra] homUplicraa
■ttcuditl «ii(h much grciUr 'littiruttiw ihnii in ihv ciuw of tlio fro^. Nov«r-
' I, in tli« bml ihi' (i[)cr>ti in iiiity be mrrietl out with ftp|iroxtinntc huc-
Pisenus for iualfliice huvc l)C(iD kept alive for five ur six wraks KlWr
cniDpleta ivnxwul of the (.■ervhral hiWMWpbenw. with the cxccpli')!! of portioui
of tne cnira and corjtoni strima imfliMiately surrounding (he optic toitlanii;
tbeM |MiU were lelt in order to eiuurv the iotacl voodition of the lattcr
IkkIw!*.
Wheu tb« iiunifdiiile «flbcle of the operntinu htiv<! pnscd olT, and fur uaw
Uiu« alWrniird. (he apmaranoe and behavior of the bird Br« alrikiii^lj
•iiuiUr to tbuM of u bird exc«e'liui;Iy el«epy and etupid. It U able to itiain*
tain what up|n:ar» to be a completely normal jxMlure, and c«n balnuLi! ilaelf
wn ouc IcK. ftfler the fo^hiuii of a bird which has in a natural way ^nue to
»)««]>. Urh aWe iu iierfed (piiet, it will remain iiiipassive aiid mAlioulew
Jbr B loD); tiini;. When >ttirn.il it tuuvoD, abifis its [Muitioo ; and then, »ii
ling IcA tlitne, returns tu a uutnral, easy posture. Placed i>ii iiA siib- ur iia
ick U will rcfpun it» feel ; Ihrnwn into the air, it flie* with conKiilerub'*
Ervciitii'n for *oRi« diotance before it returns to re^t. It fri'ijuently tuck* il«
rad uDiter its wiDgH, and at liitie* mav lie m<vu to clean iui feather* ; whuu
ii» beak ia pIiiRK<.''t into corn, it euln. It luny l>c induced to move not iinly
l>y ufiliiiiiry stimuli applied to the akin, but almi by ^tulden hiud sounds, i>r
bj ttMbca of li((hl: in iu> (light it will, though imj^ierfecllr, avoid obttuclos,
MhI it* Tsriout roovementv iip|H^r to bo to a curtain extent gnided nol only
t by touch but ix\»y by visujil iuiprMnoii*.
V III a certain number ut'coMt ibi« slMpy, droway condition poMoioffand is
laacoMxlcd by a phase in which tlie bird, niiparently «{K>iit«neously. without
■ the intervention •>( any obvioitH iiliniulu», nixvcs rapidly about. It dixa not
I At, that is to say, it does not raise itself from the ground in IliKhl. but walks
" a&oui inottaantiy for a long while at a time, periods of activity alternating
Kith periods of rejKMe. It iceniB, fruni time i" time, to wake up and inove
■hiut. mid then to g« to sleep afiaiu : and it has been observed that during
tJie ni^-ht it ap|>eani to be always asleep. It is obvious, therefore, that the
•liTpy, •(uieaeeut cunditioii is nul due tinply to the absence of the carebntl
bviutfjibrra, but U a tem|><>rary effect of the operation, and that s|iontanootu
BOTCaBCUta, that is to say, movements not started by any obvious rtimuliu,
may ovenr aAer removal of ibe cerebral hemispheres. But the luovemcnta
sn witnR«<^l 'lilTer from those of an intact bird. Thev' are. it i» true, varied :
aiul Ibe varialii'hM are in part dependent on external circum.<taitc«i, iho bird
b«ii>g vultled by tjii-lile. and sd we have said, viuial seotttiMUs. ur. to be mora
exact, Dv imprctiiiiUM niud<: upon the sensory uurvesi of the skin and on the
In-tina; out they d^i not shuw the wide variations of vo'.unlury m'lvnmiriiU.
The bin) never llies up from tbi; (cround. never spootJtn eon sly piclu up oora,
and ilanimliiui, monolunouii, rmtldj walks, rcnumblinz the continued sn-im-
tning of tJic frug thrown into tlte water after being deprived of its c«n.>bral
hvmispboras, forcibly «uggv«t that the aoiiviiy is the outcome of some intrinite
itupulae generatvl in the nervous machinery in iu>me way or other, but not
bv tbe working of a conscious inielligence as in the impulse which we call
I tbo will.
I 8till we mutt not shut »ur eyes to the foct that Mpjnlonoous movements,
whatever iheir exact nature, arc manifMtexl by a bird in the iibwnce of tlie
cerebral hcnii^jihcres, and )>eci>me the nnre olnking the more complete the
recovery from the ptiutng elfcol* of the more o|>er*tiiin. Ciuld s.ich birds
be kept alive fur any considerable lime. (luMihty further dqvel')|inicnta oiigbl
Ite witneawd, and iudee<l cusw are on r.icird whore birils have bsen k-ipt
ftlivc for moDths after llie operation, and lisvc shown spontanoous move-
78S
THB BRAIK.
mentsof a still more varied charncter than ihoae jiut described; twl itttnA
CBses lli« reinovat of tbe bemiiptieres has not been coniplele, [tonioos of ihr
veolrftl reeiniti bein;; leti behind ; and. ihoiij;)) h iiioTe ivtiinaul left ■nanj
ihe optio thalami ean hardly be legnrded as a sufScieut rause for tbe tpie-
tSDcity uf irhirli ue are ^iienkiiig. a larger maM, still more or lav ralwa^
its Do'mat etruclure. might have a markeii eJlecL Aod we may her»m>
bii[ia remark that all these fhets seem to puiiil to the conclimion that nii
may be called meuhuiiicnl Bponianetty, sometimes spokeo of as "auiitmntlmi,'
dillm from Ihe &p(>iitaueily of the " will " in degree mtlicf ilian in UtA.
Lookiiif; at the mailer from a purely physiological poioi of view (thttnlj
one vihich has a ri;;hl to be employed in ibeae pag»), ibo r««) aiflhftoct
belwceu an automaiio act and a voluiitanF- itei in ibui tbc rhiiin of iihttio-
logical evenis hetwec-n lliv acl and ils |diyaiid(>giud niune in iii (be ow or
fliorl and Him]>le. in the ulhcr long and complux. Wa bnvc men that a fn^
bickin); ila cen-bral heminfilieres, viewed fr»m one stand p<-irtt, apjioiri in tkt
light iif II roechaoicat npfiarntns, on wh{<'h c«ch change of eirrumilanon
producn a direct, nnvarying, iiievitablo efli^d. Ami yet it is ou rKonI tkai
such n frog, if kept alive long eiiongh for the mort complete dianpnorawt
of Uie direct cHl-cU of the oiN'-nilion, will Imrj- itself in the cmrtli at lb(
approach »f winter, and t» able to calch uml swallow flic* and olh«r hoi
coming in ils ncighborhofxl, allhougb in olher respects it shows no Rgntof
an iiilclti(,'rnt vuTtliou, and mtswrrv with unerring mecbniiical oenaiiUjrto
the play of ilimnli. Wc mnv add that in some firhi^ the removal of Ihi^
cerebral heniii<j>bcrrs, Hhieh m these animals f'irm n relatively small lort of
the uhoIcbrniD. produces exeetdtDglylitllu change in their general iMtnivior.
These. however. arc nol thecunsiaerationE on which we wi»l) here to dwell;
we have qiioled ihe bchavinr of the bin) depnvi-d of its cerebml bemisphcR
mainly to show ibat in tbtf warm-blooded auintal, n^ in tite more IdwIv cuU-
blooded frog, the parts of the brain below or behind the ocrclirnT bemi-
Bphem conMilute a nervous machinery by uhicfa all the ordinary bodili
movemeuta may be carried out. The bird, like the fbig. sulfiT^ uo poralya
wbeD the cerebral hemispheres are removed ; on llie cuuirary, though ib
movemeDla hare uoL been (tudied » clooely as those of ihv Irog. the binl
without its cerebral bemis|)lieres aeems capable of execuliDg at nil v^'enia aQ
the onlinary bodtly movemeuts of a bird. And in tlie hini nr io the frup.
the afferent impubea passing into tke central nervous system, whHherlkt
give rise to consciouwess or do, plajr an iiii|)ortKiit part not only in arigi-
uaiing but in guiding and coiinlinating ibe i-ircr«ut im]>»lKii nhicb stir itv
muscles to couiriirt. the cin'rili nation bemg elfeete^l parliv in the spinal cori,
but largely and in<leL-d ehiclly iu ihe |)art* of the brain lying tichind thr
cerebral hem in ph ere". It \* further worthy of notice that spuntnueily u(
movement of the kind which we have docriliid, is much more nromtneut iti
the more highly developed bird, than in tbe more biwly fmg. The cerebral
lieniiephcres are not the only part of the central nerAouK nystcm which hu
iiDdcrgonc a grcuter development in the bird; the other parts of tbe brain
have also acigiiired ii fiir greater complexity than in Ihe fmg.
^641. Id the mommiil ihe removnl of ihe cembml hcmispbcrce ■ still
more difficult than in the bird; the animal cannot bu kept alive fbrmnrr
than a few hours; but in some nuimmalu it is pvmibia to obserre dtiriue
those few hours pbeuomcnn kindred to those nitntased in the bin
aod in the frog. The rabbit or rat, from which the whole of both hemi-
spheres has heen removed with iho execption of tbe parts iiumcdialely
Bumiunding the optic ihalami, van stand, rvD, and leap. I'iaced od its tide
or back it at once regains its fcM. LeA alone it generally remains as
niutioulera and impnesive as n statue, sure now and tbeu wheii a [>anii)g
TTITaODT CBRXBRAI. REMISrHBRES.
ipulw BMiM (o tilir it to H iiidili-u but britif mnvement ; but *oractiniot it
Mcms mbtMt to n morv ountimicd iiii|m'iii; hi niuvo. iu vrliiirh caM dnith
WmtanjfommTery^fttAilf. Budia rabbit will remnin W ininiiln u«vtli«r
niUrlT hveillMs of k mrrot or cabbHf:«F-Imr pliuv^l ju*i befurv il» mwe. ihiHigb
tr« tnnrMl Iw placci) irilliin iu moiitli it nt 4iicc bl^gin* to cat. Wlwn atirn^l
it «ill with omm nni) KimlinvM run or leap roraiiH; nml olwUcIc* in its
coun* an varj fmgurntly, with mure or Ivn taccan, avutilrH. In »i>mi.- cium
tbr aninul (ni) ba* bmi described m follonin); br inn\Tm<'nt» of the h«wl ■
brijflit \lgln )iH<l in front of it (urovided tlint tbc o[>liL- ncrrw and tracts
hna jHt hw) itijnrcl during tbo operation \. m nlnnini: when it «bril1 nnd
HNd nciN ll naik' ncnr it, and ii« erring vrbcn pinclKil. often witb ii too^ nnd
ncemiogl; pinintivo scfMii. So pUiniivc U tbo cry which it thus girca forth
Itai to •n^'gisl to the ohae<ri-er tb? cxiitcncc of {n«i<>n. this, hovrover, ii jirob-
■ My K wnmj; inleqtretation of a tocuI uclion -, ihc cry nppeKrs plaintive
limplr bccaiiw. in eoiueqaenre of the coiuplctcncne of ihe reRoiE nervous
Iaiarbincrr and ihe ubsetioe of the usual netraintJ, it is proloni>od.
Without instsling too much on sucb results as theec, and alloirini; full
wpigbl to tbo ohjecliiin which may be uived. that in (onie of ihew coses
part^ of tbc rercbral hciuispborM uirrouniiiil]; the optic thalaroi wrv left,
llwre still remains adequate evidence to show that a mamnial such a* a
rabbit, in Ihe same way iks a fnig '"'d a bird, may in the cuniplele or all but
roplete absence of the cerebral bemiaphem maintain a nnlural posture,
frooi all aigna of dttturtiaiice of e<)uilibriuni, and U able tu carry out
ilh auccaag. at all eventet all the ususal and commao bo-lity niaveiiienis.
And as in tbe bird and fr»K. the evidence also showd that theae nmvetueuU
Dot only Diiiy )>e started by, but in ibeir varryinji; out are guided by aud
rdinated by afferent iiupuliNSi aliinj; aflenml nervea, iueludin{( tlMMeoflh*
' MiiK*. Rut in the cjise <tf tho rabbit it Ik even ittill cleuor than in
am at th« bird ihnt tbc oflcda of lhi.i«c atrvn-nt inipulgw are ditfomit
Jron thoK which rtwilt wbon tbe impiilxr* gain aci-cm to an intact brain.
BjDovemcni^ «f (he animal seem guided by iraprcssion* mndc on its retina,
III as on 'ilhcr scuuiry nerves ; irc luiy perhaps speak of the aoimnl as
Milijnct of spnsniioiis; but ihcrv is no satisfactory cvi<lcncit that it |>os-
■ either visual or other perceptions, or that tlic fcnrations which it ex-
pwtema gives riw to idcti». \u avoidanou of objvols depends not so much
on tlM (om of thmc as on their iutcrfereoeo with light. No image, whe<tber
pleasant or terrible, whether of food or of an enemy, produces an effect on
U, other thnn that of an object reflecting more or less linht. And we may
infer that it locks the possession of an intelligent will. Uul it must always
be renwrnlKTcd that some of tlie phenomena are due to tlic operation pr\>>
dudoR other results than the mere absence of tbe part removeu. We ratut
bear in mind that in all theaboveexpenntenla while the positive phenomena,
the things which Ihe animal c^mlinues able to do, are of great value, lh«
pegativc pWoomeoa. the things which the animal can no lunger do, are of
much lew. iwieed of doubtful value. The utorv carefully and sUOceoBfully
|li« uperinwDls are i-arrie>I out. tbe narrower become what ne may call tbe
*d«fiaCDcy pbenotnciia," the pheDomena which are alune and directly due
to aooiething having been taken away. Were it puasible to keep iliu mbbit
alive long enough lor the mere elfccts of tbe operation to poas e»m|dH"Iy
amy, we atiould not only probably wilnew, as iu the case of thi; tiini, »
greater scope of movt^ment and niort frequent •[lontancity, but |)iM>ildy lind
~ dillicullv in describing the exact condition of the animal.
i 642. tiilliertu uttempu to witiioa similar phenomeaia Id mom highly
inimi mammals stich ns the dug have faile4l ; iheao animnN <lo not
frora-the operation of removing tlie whole of billi ibdr humispbcro*
rfto
TIIG BBAi:e,
Miffi<ri<^tl7 to €Diili1e Hi to jiiHge Kbclber (bey, like tht freg, Ae hM. tot
the rabhit. mil cam- mil i-oonlinalc Ixxlily luovenietiti is ibe •iMMS^ihc
beii>i»|>hrro, cr ulteilicr in tlieiti lliw jwri of ih« lirun. so lufrlj A i i^f 14,
hm utiirpid fiinctii.nii nliuh in lb« lower inimala bdon^ tsMfer (wn
Our kDonltdffe is lHT),t'lv criiit>ne<l ii> the px|i^ricnr« ihu «wa li kdaf At
ceicbnil convolulioiig nre rrnidvei) lierenml it MTtfml a) aniwi, tk
aoiiDul may be kept altve Rod iD kocmI li«attli for e Iouk (Jne, wmMf mrtlkt
Bt lenvi, e^'^n aDer ih<-ie pans «r ibe brain havr bmn rnjund t4> nfjoHll
diinenriuna,ai)d tbat under i1ie«e i-trctini»tBmt« ibr animsl '»w*i^Mi
to carry out ntth wme limiliilinni hb txditinrir bodily mntTtoFOti^ Ml ife
exbtbita a tiponiaii«iiy vbvkiuily betukfning tbt^ ptDwikioi) not -
couwiuiu Tulitii'D hut of a cvViain amount of iuldliccocc^ I
KJIIinfc to Wlkve ihnt a nirrc fnigoM'nt, ki to Mfnik, uf thv iMtDnpcwrn ni
take on n>o«>t cxlnidrd iiontr*, lucb nn oxprrirncr ■mn* to tlxnr llm iail*
dog It* in lb<- riibbit and in the bin), ibt^ dmlopawnl of M>«alM Uf^
fonclioDs i* not limitt^d to iW crn-brgl hrmiapbnT*, tbat ti»andAmi
\ovctr portion* of lh« bmin in thr bighrr animRlt bk co«Bimrt4 «kk iki
lower do not iocmisc in bulk mvroly n» ttie ii)ftnim<nts of ibr htaitfk»m,
bul like the hcmiiplicrv* ncgnirc more nnd mure oomplpx fuixtioMi V|
niny perhnjis go so t'nr a» to atk lliv '|U(«ti^i vbcthtr the vvlitv* md fakt
li];t^nce nliirb sudi a dog «sliibilB is nt>l as luuch the product of ibtfam
lyiiiv' behind the bcniif^plicref as of the !lunip left in ibe fniBU
II ve csn thus say little about the condition of a d'j^ nitb-i<tl iW canbrij
beiDi«pber», Ke can say still leva about the toonkey. ithicb ia all aaum
toui-biuR the cerebral nervooia syelein aervtt as our hm, tadt«4 MV«*^
fpiide fur dra«iug inference coucerninf; man; but in all profaafatlity 4i
innnkcy in ihio reaped bearv nunieiA liai ili« Mine reluiimi tu lac dog lul ik
doK btani to (be bird.
In sliort, the more wc study the ph«Domena vxliibited by aDinak pcan-
iiig n part only of their brntn. the cliwr uc are pii*lM><l to ibe or '
tbat mi shant lino can be dniwii l>clwc<'n volition nnd ibc Ui-k of '
or between liie [wufsfwioii nnd nVisence of ink-lligcnce. Bclwi^n (ht ttox '■
ftene prepnralJKU at the nne limit, and mir n.'ne(-inuf willing ttlmailbf
oibcr, there is a continuous gradation nithoul a break ; we aoaat (i *■
any linear Itarner iu the bruin or in the general n^-rrnnt trft»m,ni»}
" ''fy^")*' <'''" tben- h volition and intclli;^>iicc, but up to thb ibvrr ii •«&*
Ink, however, is not llie (|ueflii<in with which we are now dealing. Vht
we want to point out h that in the higher atiimals, iocltiding at ImM am
miimninb. a* in the frf>j(. uller tbe renioviil <if the cerebral heinispb*rM fnt
tliougb roDMloiia voliliim and )nlelli;;en(-e appear to be Iiir^ely, If i»>l relin','.
lo»1, ibo boily b Kill tujiable uf exeeuiiug all the onliuary luovemm!* ak^l
the animal in it* mttunil life is wont to jHrf >rm. in i]tite of ibene murtmrti'.'
Docoaulating tin; ''i>rij)erati';n of vnriotu affertul impulses : at>il i^>i il-rr
foro tlio iK-rvMi» Riacbiiicr>- fur lli« csecution of thfse movii
•nnif part of the brain other thnn the rttrebral hcmiif^ere». W . .1
•ons ^ir ibinking that it it liiiian.-d in tli« ttrudurea foriiiiu)E thi
nnd himi brain ; lu wc iiball rec, interfen-ncc with tbne parts pruiutT- •■
once remarkable diiNtnlcre of movement.
The Mactii!iert op 06BDi».vTEt» MovmnKiti.
§ 643. We may now direct our attention for a while m s.ime «m»»ili- >
ninc«nitng the nature uf tlitt complex nerrou^ muchiner^' fur (be O'
Mon of bodily moveinentB, and m|m'tally concerning lh« part playtil "!
TUB JIACHINKHV ar COORDINATKI) MOVKUKSTfi. 701
impulara. MoBt of <nir Icnoirleil^e on this pMOt bu bccti ga!rieil
[by N ttudjr of uiiouilft not ili'|<rivei) of. bui f.il\\ pttetming, their c^i^bnl
0«n)M[ilivr«A, or by dnluvtiont from the data vf uur own exprrinice ; Imt it
p pcwsible [n nuat cum to eliiniuntc from Ihe total rojutu the ptietinmefla
<Mliii-li arc due (<.i ibe morkirig nf u oimiciriiis inielliKenoe. Buine ul' the ntoet
■thkiiif; lactn lieuriiig tm ibi* niiiltcr bave been tcained by studyint; tlte dfecU
' ofM-ntttvf tiitcrfeicncv nilh ccrliiiii |>urt9 ufthe rtileninl ear. known v tli«
(dtrular Cttnal*. Tbe dctniU ui' llic ntruiturr o( ilie^ |Hirts nc nliall
ibo Inter on. wlieD vr« txtaie l<i deiil ultb bcariuji^. but we riiay hen aay
iluil Mch internal eiir pouMMtes tlireo membruouuii oemii-iri-iilar canaLt, dit-
ned in tlie ihrcL- pluDiv nf i>|iaM! (»ne bitrizoutnl, and one in each of the
,wo vertical plani^, fun nud utl and Hide to tide), eavb meinliranoiu mual
iMine •umiuiiili'd by it bonv ciuiid of niairlj th« Mnie flinnc. nod being ex-
||mndrd ol one <'nd inii> wlint is vnllnl an nnipulla, on irbich fibm of tba
■udilorT iHTvr end. Kui-h nieniliranmis canal, in cvminon vilb the envity
•f tbo intiTniil i-sr of nhi^h it is a pr >In illation, coatiuna a Hniit ullini to
|yRi|ih, callol rn'lolyinph. and the »|>aor tH'twii-n each menibranoii* canal
mm] tt»ci>rra{Kin<line bony canal is in renlity n Ivmphnpnec contuininj; a
Ruid whieb is virlually lymph, lh(iOf;h it ie called by the «DCcial nnnie of
Jn-rilyniph. In birds iolerfc'rence with Ihe •emicireular cnnals (iroducus tho
ullonin^ reiTiiirknblc remits :
I WlH^t in a |>i^'««m iho linritontal membnuuHis Mmieircular cnnal i» cut
|tbru«i);h. tlw bir«) is »b*erveil to be coDtinuallv moving Us head from side lu
ikitle. If une of the vertical canals be cut through, the ni-ivenientf! nn-: uu
Ed duUH. Tlie peculiar movements may not be wiincnfcd when the bird is
rfoctty (piict, but they make their sppenriincc whenever it is disturbed or
kmpt<i iu any way to stir. W1il-u the injury is eonlinoil to one cnnal only,
|«r even to the canali of one side of the bead only, the condition lUler a
Iftthile |ttMes away; when tbe eunalt of both tides have been dividetl, it
becomea much cxaegenite<). iatts tiiuch longer, an<l in some ca»<s » nuid to
rrunain |)ermaiiently. AlW such injuries it iti found that these jievuiisr
[movetnctits of the htriui are asHieiattHi wtlli what ai)|]Cars to he ii great wnitt
Itif ctntnlinuiiiin of botlily niovemttnlfi. If the bin! be tbmnn into the iiir,
lit llulteni and lalls down in a hclplvn itnd coufuHw) niimner; it np|H-iin> to
hi\« lost tile [Kinrur of orderly (light. If plii(-«<il in a liaUni.'ed {Haiiinii, it
nay remsin for «»■>■' time iiuiet, grncritlly with il« licnd in a peculiiir jnm-
|UI«: buLdir4.<ct]vit i* dtitiirlHnt, the niov<.-rnfnt--< which it alteTngita tocxocuia
sra imfCiiUr imd full xhurt of their [Hiquw. It hiw ({real ditEciilty in |MCk<
Ii^ ui> food and in <trinkinc : and in gciicnil iis Itebaviar very niuob rescm-
mm that of a penon nho is «xocedingly diiixy.
It can bear perfectly well, and therefi>rt^' the »ympt'inii^ cannot bo rvgurdcd
•a the remit of any abnormiil anrlilory sen»ntiun». such a« " a njnriog " in
Iba «an. BesKlcs, any such stimulatiun of tho auditory nerva sa the le^tilt
•r Ibaseetion irould f])ecdily die away, whereas thcMi pbcftomaiui nay lost
for at least a very consiilerable tima.
\ The nmvemeuLs are not occasioned by any partial ]«^Blrsi^ by any Hant
of |Miier iu particular luuacles or group of muscles ; though removal of the
canabvf one si<I« has be«n dateribed as leading to dimiuisl»ed muscular
force DO tbe same side of the Iwdy. the mere diminutioa of force b insuffl-
I ciciit tt> explain the plwuoniFua. Nor »u Ihe other hand are the movements
idue to any uncoiitnillabic irajmlsei a very gentle preosure of the band
■ufficca to ttMi tbe inoveiuenia nf the bead, and the hand in doing so experi-
COcaa noatraiB. The awixintut! of a very slitiht sujipurt enables movements
etiMmEBe impombleor mutt iliHieult to lieeosilv exevuied. Thu«. llioogh nhen
laft aU«e tbe bird has gn»t ditliculty in drinking ur picking up corn, it will
792
THE BBAIIF.
rnniinue Ui <M willi r«M if ita bosk bv plunged inO trnlrr or into ■ bMp rf
biirli'v ; thi; iiliijhl stipjuirt of th« vtntcr or of the grain tf*fM MlSehni h
MfBilr ila niDvcnicnrs. In tbc miDO vnr il can. «ven wi[h<>ol H»iUiD(t.
ciL-na it» fenthcre and ecrnich tix hmd, its beak and foot beisj; in that
operation* giiidrd bv contiict wiib it»o«n body.
The amount of diaordvr thu« induced diflbre in different birds, and hw
mownxwU ara mon affiwted thnn otbon. As a g«^npral rule, it mar be mH
that the more complex and intricate a niovement. llie IViller and aon
doiicuie the coordination needed to carry it out succeMfuIly, ll)« mm
markedly u it disordered by the opcmtion ; tbu». after injurv to the oaU,
while a pigeon caniiot fly, a eooee ja tflill able to awioi.
In Duunmals 'rabbits) lectioDs of tlie canals aboproduoea a c«nniii amoiA
oT lonofooordinalion, but muoh leaa than tliat vitnesMd in birib; aaJ lb*
inovenieiits of the head ar« not m niDrkeil, peculiar DarillHtiiig monfocall of
the eyeballs, dilfi^rinK ■■> direciiuu and character aoconlinit to the canal or
rfltiaUojieraled upon, beeomiti); however prominenL In tho (nrgaaini^
titina of the head are seen, but there '» »omeI<«sof coiirdinaiioii in tbeiMT»
mentsof the body. In fishes uoefiect ul all i^ pr»duc<.-d.
Injury to the buny canals aloao is iudufCrieiit to prciduce the arnptaw.
thf iiiembniU'iUK raniilii themselves inu.it he divided nr injunvl. Thr- clisnc
tcriirtio movements uf the hand may hnivever hi' hniught about in a biiil
vfithoitt opening the bony canal, by stiddunly hrating or ooolinjj a caoal,
citjHvially Its ninpullur temiinationii, ur by tht> making or brealliitg of a oun-
6lant I'urrcnt directed through the runat.
Then? can bv no doubt that the«e chantctt.'iriHtic movements of the head ai»
tlte result of alTerciit impulsns started in the nervous eiulingc of the andiliiij
aervfl over the ampulla of the canal, and conveyed to the brnin ahmg thsl
oervB. And thai injury to or other stimululion of each of the three canah
abould prtxluce in each cnse n dillcrent niovcnient of ihe head, the rlirertioa
of the moveuienl bein^ diHereui accordiu); to Ihe plane in which the i!BJmI
lii.'M.shriws that these impulses are of a peculiar niiture. Tliis is futiher illiu-
Irair^l by the fullowinfc experiment. If the horizontal canal )>e rarefully
laid bare, and the membraDoua canal opened so as to ex[H»>e the eiid'>lynipb,
b'owioz giMitiy over the opened canal wiih a 6»e >;laM cauula Kill prmluo
n definite movement of the beoil. whii^h i& turned lo the one side »r to iW
other, ac«or<line: as tlie cutretit of air drives the etidolyniph tuwaitl or awiT
from the nm|)uTlu. From this it is inferred thai a movement of t)»e endo-
lymph over, or an increased prtssure of the endotymph on, the nervous end-
ings in the ampulla gives rise to aflcrenl impulses which in sonic way ddcr
mine the issue of efferent impulses lc«iliug to tJie inuvenient of tlio hroil. It
is further suggested tlint since the phun-s of the three canal* Itc in tlie tlirse
axes of space, anv change in Uic position of tlie head miiM lead ti> obangta
in the prcwnre ot the cndolym|>h on the walls of the ampulla' or to more*
DKOts of endulvQiph over those walls, and mi must give rise to irainilpn pav-
ing up Ihc auditory nerve ; and thai sinoi; cvory change of jvHition will ufTvct
the three cannU difliirently 'nhemw the change* of proMure of lb* rtulo-
lymph involved in a " witvc ofxiund " will afloct all throe ampulb< equallr)
those impulses will diHl'r according' to thi^ direction of the chaoee. A Still
further extension of this view gu|ij><>«e:> thnt sinoe in any one position of the
bead the pressure of the eixlolymph will differ in the three ampulhe, m«rs
position of llie head, ai^ <li«tinKui>ihcd from cbstigv of position, is ade<{Uat« la
getierate afferent impoloes diHi-ring in the different pntitions.
Let tis now for a while luni aside to ourwivos and exnmino tl>0 CoHfdid
tioii of the niovemenls of our own bodies. When wc appeal to our ui
sciotuoiess we find ibat our movecnents are govorawl ud guidod 1:
THK HACIII.VKRr OP COdRDIKATEU MOVEMENTS. TfiS
«* niaj mil satDW of eqiiilibniini.W nn nfipnvinlion of Uii' [HMilW of our
Wly and ris rektioDs to spnce. Wlicn tliU mrw of oqiiilihrinni i* dixtiirbiHl
wr mr wi> are ilizzy, and w«> then tttigger and reel. Wing no longtT able to
RHinlinatp Ihe nHivcntfiitH of our bodicfl or to adapt lh«ni to the p<mti->ii of
things around ub. What is the origin of this iense of rquilibriiini* By
what nicans are we able to appreciate the poHilion of our body ? ThcfV can
U> no duiibt ibat this appreciation is in Ut;r« nieaauK the product of ¥■*»»]
nod tnrlile Hmiitttiuni ; ire recognize the relatiuns of our bndy to the l-hin^
around u« in jtreat measure br sifcht and totieh : we alito Icarn inueh by our
muHuUr ceoee. But there b«or»«thint; b«»id«a theee. Neither siKht nor
tou<-h nor nutoular kom can help us uhen, placet! perfectly Hat and nl rest
»D ■ lutrisontal rotating table, witJi the ey«s f>lmt and not a muscle stirring,
«« atiMi)|4 to determine whether or nn the table and we with it are being
tmiTexl, or to ascertain hnw much it and we are turned to the right or to iho
left. Vet under auch drcuniii(aiu-«« we are cnnscioiu> of a chanj^ in our
pneittoD, and some ab«erveni have b«en even able to pose a tolerable sucoeaa-
All jud|[nieiit aa to the angle tlirougfa which they have been moved. Tliere
can be no doubt that meh n judgment 'a biuted upon the interpretation by
ooaadoaaneMof siRirent impulswwhioh nrt- dupeiideiit on the p(«ition of the
body, tint which are not alU'rcnl inipuln^ Ix-limgintr to Mcnuilion^ of touch or
wht, or lakinjf pari in the mnitciilur M'no:. And it in uiyed with ureat
plauitbilitr that the atrm-ni impulwx in •{uwlinn arv th(4ito which we have
ju«t rtfi-mvl to a* ftartLiI in the f^miciix-ular I'nnaU.
If ni- ailiiiit ihf i-xiM(-u<v of Kuch a:n{ni]Inr impiilw'V.iru'v may renttire bo
to call ih«ni. and nii>):niEr thc'm iiai'ontribiiting largely not only to uxr direct
ttcrceplton (if lli« pofjiinn of the lu'ad and thn» of the bixly, but alvo in ii more
indirect way to what we linvc called tliv iV'niw of tijtiilibriiim, we »liottld
•iperl to find tbut when tltcy nrv abnormal the acnte of etjuililirium ii< ditt-
turbfd, and that in coosequeDce a failure of coordination in our movemeata
mulls. And the Io«e of coordination which we described above a* rt!>idling
from injury to the aemicircular canals has accorrlin}>Iy been attributed W> a
dsficieocy of disorder of normal am|Millar impulses.
But vc must here disiint^uisb between two thin(i:B. It seems clear that
wh*n the membranous ciinala are injured or otherwise ttimulated nlfereiit
tnpulsea are generated which, on the one band, may produce [veciiliar more-
menti of the head, and on the uiher hand, teem able when the injury is largo
lo eauaea loeaof coiirdinatiun of buddy movemeuta. But it do«« not necea-
•arily follow ftnin thu that in a normal omdition of thiniT' alftreni iminilan
are continually pasting up lu the bmiu from the semicttctilur canals, am)
that (be low of codnlination which tiillowt upon injury to ilic canals b due
lo theae nonnal impulxs henij; deficient or altertnl. It may he that such
normal impulses do not exist, and thnl the lotn of ciiiirdiaaiion in the result
of the central raacliiner)- fiir cudrdiiuuion tieini; inti'rferisl with by '(uite netr
imptilaea gencraled by the injury to the canal with the conNCquttot Im of
endolymph acting as a ttimulu* tu llio endings of the nerve. For llie expe-
rience <|uoli-d iilmve, though it provM that afferent impuW* other than those
of MgUl. touch and the muKcitlur tcn«c do n'ftcli the brain and aflim) a biiM*
fur a iudgmenl lut to the puiiitiun of the boily, don> nut by ilavlf prove that
thoM im|)ul*ni come from the lemicircular canal*; the arrangement nf the
canals is undiiubtc>lly *ugi[c»iive ; but it it (luiio jKitoiihlv that the afleroot
impulara in itur^tiun mny Ix- generated by one or other of various obangaa,
Tommotor and Otberw, of the tnaucs of tito IxHty which arc involvMl in a
change of poailion. And if it bo trv« aa affirmed by tunic olwrvcr* that
both andilory nerves' may lie completely nnd permanently Mevvm). wiihutii
as/ aflbct ou the coordination of moTcmcnla, it is obvioua that thu incufinli'
TBE hhais-.
nation wl)k'h follow* upon icrtion of (be Mtnicimilitr canRb m dw f ■•m
sjircinl irrilnttoti Ml ii{i hr tlio nporalioii, Htiil nut lo tite incr* abariuv nf u;
normal aiii|Hillnr impulses. On the other hnnd, if lhe«ffi>ct« w ihoM nf
irritution, it is dillicuit to nndereltind hoir thcj mi. us fl<.-c<anlini; tn emiii
obwrvcre lh«y corliiinir do, become p«rmaneni. It )iiu>, bimrrr, ka
mmngly urged tlmt io hicIi cwm nf pcrmmnent inontirdiniition. llir (i(«rai)Di
hit» Fct up wcondnrr mischief in the brain, in the cMvhoIlum for iMlaan^
wiib uhitrh, » we jiuvo »eeii (§ 0I!>>, tlifl vcMibalar auditorf neno tMbs
itpcvtiil connections, and ihut the permuaeot eSteta are re*]!; dnt to iht di»
eaw f^in;; on tiera; aivl we have reaton. lis vre »hall w«, to ihiiik thit iW
cerebellum is concerned in the (;»5i<ti nation of movements. Ii oannii<,ihrf»
forv l>erepirded Ha««ttle«l that (be mnnl# iire tin- rH>iirce of normiil iin|wlMi,
or that our coiiscloua appreciation nf the i>rtMiii>o of the buul ainl (•> »( Ik
body iu «pace is baaed oa aiich impnUw. But mieb a view b nut diapmtd;
and in any caae it remains trui^ thai injury to the rana1> tlo«a In fniw nj
or other, eitht^r by piieraling n^w impiilw.-* or by allcriu); prerxiMintt uui,
so modify tlie flow nf aHercnt imiiulMa into tbe machinery of co urdiaMM
as to throw tlial machinery out «f gear.
$ 644. We have dwelt on the»« pltcnomcna of the •emicircidar (auah
l)ecauM> they illunralv in a alrikinj; manner ihc important part played bf
nH^rent impnlM-s in tho couritinatinn i>f movemouii. We taw rea.*oti t)
ibink (S ■'>9l>) that oven in an nnliuarr rrllei mnvemeiit carriMl out by th*
tpiiial ennl or by n porlinn of lliv curd atli-rent in)pulii«)i, otlM-r than ihoM
nhti'b cxrtte the movement, arc al work, dvli'miiiiine fruch coilSnU nation at
in pnvcnt In each a cfl»e tlic cionlinating ajfi-n-nt impulm,^ ar« rclaiinlf
Ktiupli- in charoclcr and rtnrt chictly, al all event* in the muiwlca cunKni«<L
In an aiiinml [low.'ninf; the lower parlsiif the bruia, thtiuirh di-priv-d uf ih»
eerehral heniiH|i)ieree, the coonliuatin;; aficreiil impitlicp. in ncoiilimee «tU
thu greater diversity aud complexity of the nuivements which the Biiimal it
able to execute, are far more |>olent and varied, Ilct'iiles aHi^rent inipubw
fVom the mutiele«, f'trmiii); the basts of what we have called the muneulu
seMe, atlcrent impidsen fnim the akin, forming the lia'is of tbovnge •tft'Kirli
in the wide ni4-nniu^ of that nun], niher afferent impulsw of ol^wure clla^
acter from ibo viiu^-ra and varioim tiHue^. and tl>e peculiar alTerent umpullir
im|Hilw« (if nbieh wt^ have ji»t epukcii, iaijiortanl s|)ecial afterent impulM
borne alnng th« nerviw of tight and haarinj; come into play. Tbe fr"g, tfe
bini, and even the nmtiininl, deprived of the wrebral hemijipberen, tbinim;'' ■<
may *}iow little itigtut i>r none at all of having a distinct voliliou, », aa wt
have ur^^l, indubitably afre<-lcd by vifual and auditory ini|>reisi»M, aail
whether we admit or no that nucb' an aniraul can riichtly be spoken of ai
being cimnciuu^ we eaiiuol rr*i»l the oincbitii'iu that affierent iin|niliin* ttarted
in ill retina or iniernni ear priMliieo in iti> central nervous ajaieni chanitca
Mmiliir lo those which in n c'iin<Wini» animal form llic basis of visual and
auiiilory sensatioDs. and we niit«t either call ihwc <^hang«d scnattioos or find
for Ihcrii some new wonl. Wh.ilcver we call them, and wbethw eoiiMliNis-
IMW i" ilirtinetly involved tn them or no. Ibey o1ivi>Hi:>ly nlav an ini)H>rtaat
pari iu> fuetore of tbe cuonlinniion of iDOvenieiit«L Imk-id, when w<< apfiea)
tA ihv experience of ourselves in |ioeeca»ioR of consciouHHMS, wo find tliat
though various sensations clearly enter into the eo&rdinatioa of our movr
m«<nte, we carry out movcmcnis thus coSrdinateil without Ih-iu^ distindly
aware of these co5rdinatin;; factors. In every movement which we make
tbe coordination of the movement i« dependent on the impulsnw or inllueams
which form tbe basis of the muscular sense, yet we are not distinctly coa-
sciotis of these inipulE«9 ; it is only, as we shall see, by special analyMs that
we come to tbe codcIunod that ne do posseas what we shitii call ■ mUMuW
tNTS. 795
So agniti. tiikiiiif the iniilter ffoin ii jiotiiculiat 'liB'ereiit (vkint of vi«w,
if o«r ttmvrm-tiis. iimrktHJIy. ■» we xliall 3««,tboM of the eyel»ill«. >re
n<rinliDaln) hy viMial gentiiitonn. nm) wlicn vr« tJntt or wlien we tlunra to
nitiMc our movewmta iirc CHinlitiBieil by ih«livIpAr*eDBatlnu3of muihI. In
th«M MM* dittiiMt ■eiuMtiana in the ordinary i^nwofllie woH iiitorvene; if
wt ciuinot M« or cnnoot hear, ibc movcmetit fHila or in imiierfect : y«t even
{n ihrM cnw* wo »i* not dirrctly cimciou> of ilic tcftntlonii u pofirdinniiojt
fiictoni ; It DrMis nn'fiil ■luilpf* to |)ni70 ihal the Mccna of the aM)veinent
in rmlly iJrprmlrtit nn ihr wuinl or on the *ighl. TheMHiMl nthcr fiictii oiit^
mt lli« virw tbiit iht- fHiint nt whii-h the vnrioii* nlTerent ira|Milw> whioh
form lb* Ixint" of the »eitfUlioii» nf n conwinuit indiviiltml rntcr into tHc 00<(r-
dinfttiD^ mwbnni^in is or mnv lif w)ni« wtty *Ih>t1 of (hr »iiigc nt itbicli the
coni|kl(<I« oonrereion iif tlw imjHilw into n pcriVcl iwnwitiou mki^ jtlnm.
TliPfvc-nts which fonjiiuitpwhiit wp inny call vi»UHHni[Hilm^, n» thiiM; Ipnvo
thp retina to »«wp along the optic nerve, lire we tno«t nilmil very (Hflercnt
fmm thiffc which in ihe npproprisic jmrls of ihc bniin ooiMiliitc whiil we
Bmy ciill consCToiw vision ; mud pmhiibty Itrtncon iIkt beginning nnd (lt« end
there tre pragreMive ohangm. It in probnble.wttMy, that these ritusl eveuis
mnyslR^t lh« coSrdinating mechniiisin at eoniertageof iheir prmrrMs before
iher reach ihcir final nnd perfect form. If this be so we mny fitrlher con-
cluae that lhoU);h. when ihc whole ncrvotis mnchincry isprcaeut in ii^ entirely,
the aflerent impulsef which lake pnrt in cooniinalion miti>1 iaevitablr at the
nine lime ^ive rise lo eoDM'ioua gcntoitionB. iltey nii^ht Mill eflect Iheir coAr-
■linatinj; work, when, owing.to thrir imperftciion or lack of Ihe lorminal
part of tbeoervouH mneliiuery.ibe impulses failed lo receive their Bual trana-
formniion. and coosciuus feiisations were nhaent. In other word*, the coi'>r-
diniit int; influences (if »en*ory or HlTereiit impulses are not eMenliolly dependent
OD che cxixu-nce of a diminct (viniciou^cit^qi.
{ 645. We have rntncd this piiint piirtly for tht^ Mike of illiMtmlinir the
working of ihe coordination rnnchincry in ihir nlMcnc^ of the eercbrnl ho'id-
gi>h(?fe». hilt alfn in onlcr to aid in thr inti-qirftulinn of the ml^Vclive mtidi-
tion which we ■|>Fiik of w ^•idrliiirivi nr ili/v.iniTui or verligo. We coni(iarei|
the omdilion of the pigeon after iin injury to (he temicircular cannU to that
of a |)er«on who n giddy or di/f.y. nnd, in<le<-d, verlign is ihe iubjeclite vx-
pnrHion of a dirnrmiigmncnt nf ihe ei>ordination rnnchincry. eitpeeiatly of
thai C'wemcd in (he mainlennnor of IxKlily oqiiilihrium. It may oe brought
about in manr wiits. When a constani current of adoipiate slrength is stmt
lliroiich the bead froni cur lo ear, we experience a pence of vertigo; otir
ntoTemenla llivn ap|M-ar lo a bystander to fail in coordination, in fact lo
reeemble three of u pigeon whoee semicircular t^nals have been injured;
and. indeed, ibe effects are probohlv produced in (be same way in |I»b two
oaaea. In whni a culled Meniiyre'a disense atlacks of vertigo H*m to he hsmi-
eiated with disease in the ear, being atlrilmted by ninny lo disorder of llto
feiuteirculnr canals, and cases have been recorded of ciddineai as well as
deafnen reoiiltinK from disease of the auditory nerve. Visual teoaattoiis are
very potent in producing I't-rtigo. Many perwHia feel giddy when tbey look
at n nalerfall ; and this is a case in which both ibe aenae of giddiness auil
the dtumngemeiiit of coordination ia the nuult of the action of a pure aen-
aailon and nothiti^ else. In the well-known ioteitse vertigo whieli w enuseil
by rapid rotntiou of Ibe body vbual lenaaiiun ptnya n part nhen the rotnlion
U carrieil on with the eyes open, but only a |ian : for vertigo may be indure<l,
though not *o readily, by roliilion with the eye* eompleiely shut, hi the
lath-r rate it ha* l>rcn HUgg4-*li-iI ihat the vertigo is cauieil by ahuornuil am-
pullar inipulfr*, hut thi-t« enn only conlrilnite lo the rexull which ia (n the
min rautol by direct disturtionceof the brain. When the rotation i« c«r-
rnc BRAIN.
ri*^ oat with Ihi? evM open, lh« vfiriii^ wbich is felt when ili« raUlisn i
\» pnrUy rauseH by the vtstinl ««iiMtii>ns. nn accwiDt of the hrhavliir t
CjtcImIIb. coostn^ to b« in hnrmony with the reft of ihc MOKntinnt nnd •fitnt
impuism which iiel[t to make up the coordination. The roUUioo «eta up
jwculiar oecillnliu); mnrementa of the cyebiills, which continue for •obh timt
«ftcr tite rotation lias ceased ; owinK to thwe movements of th« eyf^k llit
visual sensations excited are BUoh as would be excited if cicteniJil abJMtl«m
rapidly innvinK^ whereoa all tbe other senntioaa ami impulsw whidi in
alMctinK the central nerToinajruAmirewob as an> excited brobJMt* at rat
Jn a nonual slate nf thinKi* the visual and the other seusalioDS and ini|wlMi.
which go ta make up ilie cui'irdiiiatici;; niachiuerv, are in at'cnrd vitii Mti
other in ^e^«^l>u<^e to the event* to the external world which are gWim nm
to tliem : alter rolatiiMi (Imj- are for a time ia dida«coriI, and tlie ooordiiiaiiag
maehinery i<i in <!0ikae(|uaiicc diMrrnugwI.
When w« inUrrogat* tmr own ofHUciouaneM, we Hod (hat we an nBtdii-
tincllv cv)uiicio4i» iif ilii« dioiuicord; the vitual senialions are ii> pnfiotBiit a
cnrntrJoufin'M, that we really think the vsternal world b ranidly whiriiag
n^uiid, all ihnt we arc furlhor eondou* «f i« thv foelinj; of |{id<lineai ai>4 nor
iuabilitv to make our Wlily moTmoent harnioniw with onr viiiial i
S.I that even in i1wciim« where lhi> I'm of co5rdlliation is hrr>Ui(hl I
wnsntioiis what wo raally appreciate by nuauu of our convinuMiaa 1
dixirnin^mcnt of th« c-Mnlin.iting machiiiory. It b th^ appncialiun tJ
this disorder which consiituUw the fMling or vertigo; both tue (Ssding
ghldiiHOB and the diforderod movements ar« tbo outoonw. one «iibj«Mii« u
tba other objective, of tbe tame ihiii};. It is not booauae w« ttti ffjiAii ihM^
wo KtAgger and reel : mir movements nre wrong baoauM the madiinerv ii u
fanlt. and it u ibe faulty action of th« machinery which nUo makei lu M
giddv.
We may h«r« perhaps remark that it is an actually dtsonlei*d otaditbw
of tbe coordinating meehanisn) which ^ivee rise to the aAoctiou of ooMttMf
neas which we call Kiildinee». nut n mere curiaitin; of thi> mMihanism >ir M
failure on its |)art (o make itself etreclive. Cimpleic hIiiidoeM limiU L'
niue of activity of tbe maohinery hut leavee the remaiiular intact, aad i
gidataea* is felL Ho ai;iiin in oeruin diMaaas of tbe nervoos syaUa
muscular iMiAe ia enterlered with over c^naidarable regions of the Uidr, i
fa these regions coord i nation fails or is imperfect, but the central maeliiMt^
b not therebv afliboted, thoujch it4 area of UMfulneas u limited, and no gMJi'
neas ta exp«ri«nced ; aud M'> in other inslaneesi.
S MS. Fomid mortmfnU. So far as we have direU on disurden of iW
coonlinaiiiif; machioery brougltt abMit by tliu aeiiun of various aSmai
iiupulM^. We have now to call attention to some peculiar pbenonieDa whiA
retiutt from oitcrative interference with jiarta of the brail), and whieh in SMW
inxlanrui nt leiwt may 1>v taken to illuninte bow this complex naoluSHT
workH when some of ili inner wliwU ar«! broken.
AH invcM^tori) w)i<i hav» ptirl'nnni-d experiments on the hmin bare ob
eerrcd, as the rmtdl of injury to varioux |urt8 of it, remarkuhle iniivcnw
whidi have the apiH^rnncc of Wioji irraistihle, compulsory, foroix), ~"
Vary much in the eslCfll to which they arw driveloped : some are so i **
hanlly to d<;wrve the name, while otiK-n hk i>lnkin|ily intenae. On
nuat common t<>rins b that in which tbe nnioul roll* ineeMantly niund
lon^tndinsl axis of iia own body. This b es{)wially common aft«r tvriiq
of otic of the crura corahri, or of the middle ami inTrrior pedundaa nf I
oereMlum. or after unilateral SMtioa of the pon*. hut hu also basa '
nessed after injury to tlio b:ilh and o-irpam <|iiadfigeniina. tJomKimiu iW
animal rotates towiird and loniotinica away fntni tbn skis operated na.
»
^
lAhrr fi>rni b that in whieti t)io animal exectilea "circuB mnveiiieats," i. e.,
coniiiiiinllj inovra rmiiiil nnil ruuml iu a circle of lua|(vr or iih'>rt«r ruiiiua,
•iiinclinMB toward hik) HomclimM away from the injuiwl oido. Tliu nuiy lie
MTU Bft«r Mveml nf lli« above-raentluned operations, antl in one fiirni or an-
otlirr ia Dot iincminioM aflt-r vnriouit uniliitcrnl iiijtirim tu ibe bniiii. Thvre
it a vnrirtv uf ihc circiut inovcmeiil, lliv "cKx'klianil ranTctneiii," himI ta
ocnir frTi)UHiltv after InnonMoflbe piMcrivr curjinni iiuaJriguniiiin, in wliioh
tbi' Buinial oii>v4ts in n nrcle, wttb tbe lungitmlinat nxin of lis body oa a
ndiuM, anil the end of il« luil fur a mnln!^ And llii* form again may carily
{MM into a simpler rolling movement. In jet anollier fiirro (he Animal roUUM
ov«r lli« trwwvfnw nxi« of itn ImmIv, tiinililw head ovw heidit in a >ori«« of
MMiH'taanllR ; or it may run tnnauiDtly in a niniighc liui- liackwan] or for-
ward until it is atojipcd by aoiM obMnclo. The*i.^ lallcr form» nf forced
wov<^mc»l» are somelinM* aecn »1\tr injury- to itir eorpiiK mrintiiui i-vrn whun
a v^rr limited portion of tho gray mniK^r j* iiHK-led. And many of thcM
furc«(i movcmrnU niny rtMilt from injurka which npfvear to be cuolincd to
th« c«rvl>ral ciiricx.
When tbe phenomma are well davelopcd, vvery cflbrt of the iinimni bringa
on a movement of this forced character. I^ll lo iieotf an<l at n.-«l tii« ani*
nial may ptment nitthing iibnormal, il« ()o»Tiin! and ulliKidu may be quite
naluml ; but when il in excited to mow or when it attempts of ItMlf to nwve,
it exeeutce not a nataral movement hot a forced one. turning round or roll-
iug over a« ihe case may be. In xevere cases tbe movement is conttnuctl
until tlie Bntmnl is exhausted ; nhcn tbe esbaustioD passes oM' tbe iioinial
niAT remain for some little time tiniet, tint fome stimulus, iniriaaic or ex-
trinsic, soon inauijuratcs a fresh outbreak, to be a^in followed by exbauatioD.
Jn some of lb« milder fonns, that lor instance of the circus movement
with a lon^ radius, the curved character of tbe pro^resxioD appear* aimply
due lo tbe tact that in the clTort of lucuwolinu vuhtiitual inipulata do not
gain oucb ready acKxsa to one nde of tbe birtly a* tu the other, die injury
navibg eauaecl aome obtiacle or other. Heuoe th« contractiomi of tbe mu*-
clca of one aide (the left for insianoe) of the body are niortt jKinerful than
Um Other, and in conwequcnc* tbe bcxly i* conlinuiilly thrunt tonanl tbe other
(the right) fide. A* ts «<'ll known we ouRfulvrf, wbeu our walk ia not
guitled by visual wmuitioii*, lend to dencrilio a circle of fontt-nbat wide
radiut, iW (k-vialinn being due lii a want of bilateral *ymm«try in our liinba ;
and the above circii* movement i» only an exnggt'ratiun of tbut.
But tbe other mure itilense fomu of f'jrecil movemcutM arc morn compIE-
caird ill their nature. No im-rc blocking of volitional imi>ultcs will explain
nhy an animal whenever it elteniptc lo move rolls rapidly over or ruihca
irreaialibly forward or biickwanl. It ia not poaiibic with our prevent knuwl-
fdf^ l4> explain how each particular kiiKl of moveoieni i» brought about:
and. iodcvd, the (cvernl kinds are probably brought about in ditlereol wars,
for they dilfcr ao greatly from cncb oibcr that we only clan them together
because it is diflicult to know wlicro lo draw the lino between them. Uut
we may regard the more intense lurme as illiisltatiug the complex nature of
what we have cvlte^l the coordinating nwchinery, the capabilitica of which
are, so to speak, di«closed by its being damagol. •'fucb groM injurica as an
involved in dividing cerebral stcncttirea or in injecting corrosive mfastanoe*
imo this or that pun of the brain, must, of necenity, portly by blocking' tbe
way to tbe impulses which in a normal state of things an oootinually [m«b-
Id{; from one i«rl of the brain to another, partly by ({eneiating new unUMial
impuWa, »eriou«ly nUed tlK> due working of llie general coordinating machi-
uerv. The fact that an animal <--aa, ai any moment, by an etfort of its own
w)l(, rotate vu its axi« or run slrnijjht forwaixl, shows that tbe nervoiu
79S
TRK BBAIK.
i-nntintio bi mt iritl) mm if ili bt-nk bo |>luDg«<d intn water or iitln ■ hrafl
loirli'v ; the iili|;lil aiipporl of tltc wat«r or of the imin seema siifllvical M
fl««<l}^ it« mnvcminits. In the mme itny it can, even wilhriiji kiwfatuei,
cimn itN fcnthors nnd scrnich iix bend, its Iwak and foot b«uiK U 1^
u|irnilUin» guided by cimliict vtilb its oKn body.
The nniount of disorrlor thus induced dil)i>ra io difTeKot bird*, uid tarn
inov«iiHnt« nro toorv tMeclvd thnn othora. As a g«ii«rHl rule, it mar ha hU
that the mnrc oamplcx nnd intricate a niovenient, the fnll^T and nuiC
driicnie the onnrdiiinlioo needed to carry it out HiocevHlullv, the nun
iuark«dly ia it disnrdered by the operation: thuR, aAer injury to t)te aaab.
while n pigeon cannot tty. n fjooac is etill abl« to swim.
Id mammals i rabbits i serltunii of tlie canals aisu pmduoca a Cf-rtjuu aDinaU
of Inas of coordination. bn( luuoh leas than that viueaned in binU ; and ikl
moTemenis of iW head are not m niarke<l. peculiar oacillnting RKtvaaMMH rf
the evebatU, dilfi-rinK; iu dir«eiion and charaotcr aooonlin^ lo tho ctnl tf
nuiafs operated upnn, becomitiK lii^wever protulDciit. In the frog no (l«rW
tiuna of the bead are Men, but tltere it ixHneliMDrooJSnliQBtinii io tb«BM*>
inents of the body. In fiihea ho ell^t at all b produoM).
Injury to the bony canals aloiw u intufficient ta prmltiMt th« sritiptaai;
the tnembrannus canaU llieniaelTei Diu*t be diviiki) or ttijurvd. Tm elMno>
tcriuio niovementa of tli« head may ho<irov«r b« bmuglit about tn a M
without ojWDing tlie bony canal, by widdmly boating or ooolinj^ a caul
tttpecially itii ampullar t^rminatiiMui, or by the making or breaking of a «a>
atnnt eurrcnl directed through the eniinl.
There can l>c no doiilit that tbiwA cltarnotertXic ntoremeDia »f tbe haul W
tlifl nsull of aflitritit impulMv •Uirtecl in the nervouii rndinf^ of the auiliwy
nerve over the nnipuUa of the cnnal, and conveyed to th« brain along thil
nerve. And Ihnl injury to or other stimulation of each of the ibni- iaoiIi
should produeu in cadi caw a difl^ront movement of (he bead, tlie ilin-i-iiaa
of tl»e moTemant being dtflbmt according to the plane in which iK<- <-:iii«l
lies, ehons that these impulsoa are of n peculiar nature. This ia furtli' :
treted by the following experiment. If ilie borizonlal caual be m!
laid bare, and the merobraDous canal opened to as toexpoiw the endi'Iunjih,
b!nwina> gently over tbe opened canal with • fine glass canula will pn«laa
a <lefinrte movement of the bead, which is turned to the one aide or to lb»
other, according as the current of air drives the endolyinph toward i>r »nj
from the amputta. From this it is inferred lliat a movement of the «tii>
lymph over, or an iocrensed pmsure of the endotyiDph on, the nervou* rait
IngM in tlic ampulla givM rise to aflerenl impubee which in wmx' way lUut-
mine the ia«ue of eflvrenl impulses leading Ui the movement of th*' liciul. Et
E« further niggMed that since tbe planes of the three canaU lie i» th« itine
Bx<s of H))aoe, anv change in the position of the head miial lead tn ebans*
in tbi^ prtrwure of the eii()o1yni|>li ou the walls of the ampulln or to no**-
meniK of eiidoljrmph over those walls, and »o roust ^ve rise to impubas ]m»
ittg uii th» auditorv nerve ; and that t>\nct every ohanae of position will aAM
the ibrec canals <lilfureotly (whereas the changes of pre*urv of lb i"
Ivmjih involvetl in a "wave of sound" will affile* all ihrae ain|Hillnf<ju.<!it
tlioeO impulsM will difli-r aeeordin^c to tlie dirvelion of the cfanngi^ A itill
further extensioo uf this view supjio^nu ihni miMv in any oiio pxsilJun nf til*
liend tbe pressure of tliv cndolymph will ilitlbr in the lbnx> ampulbe, iwrt
position of the head, bi> ilitiliiiinitfJied from i-)iungc uf poaition. is Bd<<<|iMt« to
generate afferent impulM-a <litt«rin|; in the dilTi'rcnt ]>««itionfi.
X^ us now- for n whib^ turn Buidn to oiirM>lvni and examine the ooOntins-
lion of the movcmentN of our own b(Mliw. When wi: ap|>cal to our iian r-nt-
sciuiisaesB we find that our movumuiits arv governed nod guided by "hsl
TnE HACniXEBY OP COUBDINATBD N0VKXEKT3. 793
a' tatty mII ■wnM:<if equilibrium, bv an H|>prEciiiLioil «f tlw pwilion of Mir
Kxly nnd i\» rvliUions to R|mcv. Whvn ihio Mtnsc of equilibrium in <li*turb«(l
Mr wo nrc <)iixr, nnd we then elitg)^-f nnd reel, boin^ no loiigtT nblc lo
icvirdinitlv tbo movcmrtils of our bodiw or to lulnpc lh«m to lb<! jMHilinn nf
tliiriK" nrimitd iii^ Wliat is tbe origin of ilii* «i>»p of equilibrium V By
bat mMui<> are irc able to np)ireciiit« the pneitiOD of our body t Tbim' cnn
B no diMibt that this appreciitlion is in Inrgfl meMUr« the prodncl of visual
nd laciiUi nenntiona: we reoot^izo ihc relntiuns of our body to tbr tbin^
Ikkiih) us in great measure by siKbt nud touch ; we also lenm much by our
UM-ulnr HUM. But there ismmetbini; bfl^ide0 tbeae. Neither sichi nor
luch nor musculftr ien»e can help ua wben.plaoed perfectJy flat and at rent
11 h<>rij:iinia] rvilatin;; table, nilh the eyes «but anil not a miiBcle stirring,
'«■ Rtii'Uipl U< delertuine uheilier or no the table and we nith it. are being
vod. or to ascertain how much it and we are turned In the rif-bf or to th«
pft, Vcl under such oireuiii8lano« we are cinaciuu? of a clian}»e in our
loflitioD. and eome obiervers have been even able lo pam a tolerable suooeM-
al judgment as lo the anitle through which lh«j liave been tnored. There
BO be DO doubt that such a jmlxment is baaed upon the interpretation by
DaaeioaeaeMofalTereDi impulnoewltich are dependent on the position of the
Kiily, but which are itot affeKnt impulaea belonging tottenmttont ortou<;h or
iRht, or lakint; part in the muscular »e»*e. And it in urned with Kreat
Hausibilily that the uiTervnt iwpulntvi in qiit-Jilion an; ibue which we liave
Di»l referred to a* itart«d in the Mniiciniulnr ranuln.
If we admit the exinlcnoe ofsuch ampullur impulwi*, if we may venture to
call them, and recn^iniie lti<-nt lu iiinlributing Inr^ly not only lo our direct
rceptiun of the [Kuilion of ihc Intiil and thuii of th<! botiv, but a1»o in a more
direct way to what we have oalliHl the scnou of equilibrium, we should
(lect to find that wbou they an- abnormal ihv vrnw of rfiuililirium i» dia-
rfoed, anil that id conwquenc" a failun- of coTirriimilioii in our movi'incuiM
Ita. And tbe loM of coord i mil i»n which nc drvcrilHfl above n-i nsnllin;;
iiu injury to ihf Mmicirenlnr caiinlii h»:> acconlingly been uttribuied lo n
ticirncy of dinordcr of normni ampiilUr impiitmi'.
Btll wo inurt here di»lin|^iii«h Ijclween two th)ng». It wcm" cWr (hat
bta Ibe membranous ciinnle arc injured or othcrwiM' vtimulatcd alTcmit
Dpulam are Kcoemted which, on the one ban<l, may produce [Ntciiliar moro-
IMilB of tbe bead, and on the other hand, ecem able when the injury lit lar^
< oiiiK! a Ion of coordination of bodily movcmeuta. But it dom not iH)c«a>
rily follow from thiti that in a normal condition of ihini^ atrvrenl )ni)>t)lMs
re cniitinually pa^siu); up to tbe brain from tbe semicircular canal", nnd
It tJ)o loM of on6n)ii)ation which follows upon injury to the canals is due
these noriDa) impulses being doHcient or altered. It may be that such
Drma] impulsea do not exiai, ami that tbe loea of coordination is the reeult
f tbe central naehinery for ooordinaiion beiun interfered with by '(uite new
ipulsd fTcnerate'l by the injury to the canal with tbe coasequeut lose of
laolynipli acting' ae a stimulus to the eudini^ of tbe nerve. For the exf»-
>nce quolol mIiovo, lhotif(b it provcti that atferent impulaw other than tbrM
' aitlbt, touch and tbe muscular sense do reach the brain and afford a baais
r M judgment aa to the pi^iiion of the body, doee not by ilaelf prove that
idse impubn cone from tbe eemiciroulnr canals ; the arraDKemeut of the
tnnls is undoubtedly sugMstire ; hut it in quite |)o»>ihle that the adWrent
i)mUea in queation may be (generated by one or other of various chanxea,
laotuofir and others, of the tbauee of the body which are involved in a
fbaniri- of |M»ilH>n. And if it be true aa affirmed by some observers that
^tb auiliiory nervca mar \*e completely and [wrmanently severed. with<mt
~ ly eflect on tbe co6rdiuatiou of rooTements, it is obvious that tbe incoSrtli-
TnB BRAI!9.
BRtioti which ro11<iw*B upon eoctioD or iho semifiniilnr caimN n iIiik (OMim
sprrinl irrilnlinn set up by th« nperaUou, nnil not to tho mfre nb»rtioc of Itf
normn! nmpiillitr impulse. Oa the other hand, if the eflecta nn- (1i»hi at
irritHlinn, it is ditBi^ult lo utidereUiKl how ihey can, ns arcordio); In in-ruil
oWrv«rB thev certniiily do, beooms perniniMDl. It bus, b»wpwf, tn^a
Mroni;ly urjt^d ihat iu such ciwes of p«riiiaii«nt ioooSnIinaliou. ih« u\'<-—< t-
hne set up wvoudin- tnisdilef in lli« bntin, in the cerebelluiu fur << -
with ti'birh, m w« have aet-a (§ CItJ), ih« veatibular auditory nerve m^m
Hpet-ial coiioe<rlii>i», aiid that th« pennanoDt effbcts are really due to thi- ■!»
mae Roint; <>ii h«r<-; niid ne bav« rv«Kin, u we shall sc«, in think iliai iht
cerebellum is coticvnm) in th« nr)0itlinali«o of nunminenii). It ckiinM.tlwn-
fore heKicnrded iw iwtl1<-*l tbiit tho caaab are thi^ «'>urre of nnrmiil tm|>ul«^
or that our contciou* apprcnstioa of the iwmiion of lb« head ami »•> i'( iW
Iwdy in fpacc W baMd on *ii<-h ipnpulHo*. But tiiich a view is not dtiipr'vnl;
and in any oho it rmiaiDR triK- that injury lo tbr oauaU d»«>« in vrnf war
or otlwT. *'ithfr by ^ntmting iww im[HiW« or by altfirini; prri'xiaiii
HO moiiify th<' flow of aHorent impulmi into ibe roachincry of co orWn
tm to throw that niovhiiwry oiil of gt*r,
i 644. Wo have dwell on tbow phennnwia of the aoniiHrcular ram!-
b)x-atif« thry illustrnio in adrikin^ niann<>r tliv important [wrl pt.-.
affcmit impiil««B in th« coordination of niov<inK-iit». W<> knn r>u
think (S '^W)) that ev«n in an ontinary retk-x Riov4-nu>nt carrio'l out br tlo
»{i«Dal cord or by a portion of the nirti alK'n-nt iinpiitMM. other than ihiw
whit'h «xcile Ibe ntovemenl, aro at work, itvtrnuining Mich coCnlination ai
it pnwnt. In wch a nse tho eo&ntinaling aA«r<-nl inipuhxM an n' '
ilmple in fihaneter and start chiefly, at all eventa in the iDitaclca wi
Tn an animal poase^iuK the lower parts of the brain. thoui:h dt-privnl >'i tr>F
cerebral bemispherf^. the co^nlinntint; afli^rent imputsee. in ncc»nlaii(« uii^
lh« (jrenter diversilv ntiil <.t>niplexity of the movemenls nliifh the an'
ab'e In execute, an* far mure |>otenl and varied. He* i- 1 v* n tie rent '.n
from the niii«r]c«. forniin!; the basis of what we havi^ cnlled the it
sense, niri-rriit ini]<utM-B> from the skin, forniinK the l^ai't* of the v-nf ni i< i'
in the wMk rm-aiiiiij( of that word, other aflerent imptitsM of i
acter fmni the vi«*ra and various tisftues, and the peculiar aU'er- n
impul»e« of nhieh wo liave just s|ioketi. important specitd afTeivnt imi'iii"
home alonji the norv«« of xjght and Iwarinj^ ooino into piny. The ft'L-. !'>•
hjnl, an<l oven the mammal, depHveil of the cerebral liomispberea. tli
may show little *igm or iMwe at all of bavin); a distiuut voUtion, t<. ^
have tir):eil, imhihitubly afloe(«>l hy visua] and auditory imprawinnt, Mri
whethi^r we admit or no that Mich an animal can rightly be ^■"■^.— >'■'
Iwint; eoimciouK we cannot rvillM tho conohision that afliirent imj'
in it* retina or internal ear prodiieo in ita central nerroua ayntem mu':---
similar lo thcwe which in a ccKiieiouit animal form the baaiii of vfamlw
auiiilory w^niiatiimf. and ne murt either rail tbeae obaneu aenaMloU M lal
for tlii-in »»niv new won). WhaCevoj- we call them, anil whaiher cuaudM*-
nMK i* diiAinelly involved in them or no, ihey obviounly play an imp<ftwl
part iu> factiioi of the ci»jnlinulion of movvnienlH. Intlvi-<1, when Wo ■tpT'*'
lo the eipvrieoou i>f ourMlvi-M in pooKwiion of o(Mi«riou»n«M(, wa Dml thii
though varioiH seiuntions clearly enter into the eodrdinatioo of nur nuin
mcnlM, we carry out raovomenla thu» cnordinaitKl wiilmut l^-injj dWintttf
aware of these ctKinlinating factors. In everj- nioverm-nt wliicli w* m»l«
tho co6r*linntion of the movomont is de|>enilcnt iin the impulst-s or inflortW
which form the baa'^fi of tlw mtiaoular sense, yet we are not disum-llv e^*
sciwi* of these impulse:*: it is only, as weahall m'«, by speetal aniitrui tlul
w« come to the eoncbiaiou that we do puescn wliat wo snnll imll a mUMmlv
I
»
I
BSBN. 60 xgain. lakiiiK iW nintlcr fmm n ttointtwliitt fllflV^rGiit point of view,
manir of utir rooremfiuti, marknlly, u we fthnll mo, ihooe of lh« eyelwIU, are
oMinliiialcd hy vlMial aetmili'ms. and vbo) we xin^ nr when we dance to
muaieour movt^nH'TiU arPivH^rtliimtrd f>T(li<! hi.-l|iorM>iiMition!i nraotind. In
llieee cawa dinliacl .■x-iiniKioiin in [ho urdiimn- m-im^ of thi? wonl intervene ; if
we nuinot hm ur );niiiiot lit-nr, ihr mcivinni-nl luilii nr i» ini|>i-rf(-(rl : ¥■-'( fven
in thrw cnsn we iiro i>»t directly (-oiiin'ioiiii uf ihr *cnMilii>:u> iin ni^inliimiinii
fartnni ; it nro<)K ciircrul iiriiilv<i:> l'> pmvc (hut lh<- *nc-(T!B< «f tli«^ mi>vi>m('Rt
ia ti-tdly drgieiultiit oti the Mnitiil "r <>n iho *i[!ht. Thr^r and other fnci* hik-
!;PM (he vH-o' (liiit ill-' |H>inl lU whk'h tii<^ vnrioiiit nfffrrfnt impiilw^ which
firm lUc liiwif uflhv ffttfatu^nnof » ci>ii«pioui! itidivi<luiil witcrinWi Ihe co5r-
(iiftating mct^banisDi is or mar be »»me way Khort of the stage at which (he
cnniplric cnnversion of the impuliw into a p(^rfcct sctiHition taken place.
Thr i-vrols which conflilute what we may call vi»iial iiiipiilees.M theae teatre
the n-liiia to sweep along the oi>lic nerve, are we mii»t admit very diRereat
Ituri iIkwc which in the appropriate parts of the brain conotiliile what we
tnay call conscious vision ; and nrubablv between the JM-cinninf; and ibe curl
there are pmgrcasive changes. It is probable, we say, that iheae visual eveiile
niay alTcri ibe niordinaling niechnnisin at some slni^ of their prodTesa before
they reach their linal aiiit perfect form. If this lie so we may further con-
clude that though, when ihe whole nervous machinery ia present in its entirety,
the nifercut impuism which take part in ciWirdioation must inevitably at tbe
aanie lime give rise to conscious sensations, iliey migbt alill effect their eirfir-
diiinting work, when, owing. to their im|*ifrelion or laok of the terminal
fiarl of t lie nervous oiacbinery, tlie im|>ii)^a failed to receive their final trnns-
forinntioii. ami vonseioils senaations wi^re aWnt. In other wonl.*, thi> <iH'ir-
dinat ing influences of arniKiry or afferent impit Ilea ate not ai«wnitnllyde|>en<lent
»D the ex'otenee of a dialinct coniti-iou^ueas.
$ 64S. We have raised ibia jioint partly for iht^ Mike of jlluatmtinir the
wnrkin^ of (he coa'nliiiation innt-hinery in thi- almcnce of the cerebral b(nni>
ai>here», but ala» In onler to aid in the inlerjiretation of the sutiieclive eondl-
tii>n whieh we spenk «f a* ^iddin<«D> or di/7.init«> or vertigo. We ci>m]Mre'i
the eondition of the pigi^m after nii injury (o the M-micircuIar canaU to that
of a iH'nwn who in gidily or dixty. and, indeed, vertipi is ihe subjecliro ex-
irecsiim uf a di:>»rriing<'rnent of the coiirdinalion machinery, especially of
eoiicernisl in the niarnlcnance of bivlily <xiiiilihrium. It may be hriught
„jl in manv wayn. When a constant current »f ade<|uatv strength is aeot
trough tbo hcail fr-n. . n . ■ ;ir u, exjicriencc a *en»e of wriigo; our
BtovemeDta then api" < .n : r to rail in oiirdination. in fact to
rei^mlile llioao of a pigeoti ullo^« xmicircular canals have been injured;
and, ii)de«il, lh« eAciia arc prohnhlv proditi'cd in the same way in the two
eiiaea. In what is callfl Meniere's disease attacks of vertigo seem to be asso-
dalcd wiib diM-ase in th« car, l>cing atirihulcd by many to disorder of the
eeininrcuhtr canals, and c.-ims have been recorded of eiddineaa ea welt aa
dcafiiem resulting from disease of the auditory nerve. Visual senaations are
very potent in pnxliicing vertigo. >Iany persons feel giddy wlien they look
ftt R waterfall : and ibis is a ciise in which both tbe sense of giddiness ai»d
the diaarrangement of cixirdinution is the rMult of the action of n pure aen-
Mlioii and nothing else. In the well-known intense vertigo which ie <?aused
bj rapid rotation of the body visual sensation plays a part when the mlutioa
b rarried on w ilb Ihe eyea open, but only a part ; for vertigo may be in<luce>1.
tliouKh not to readily, by rotation with the e>'es eomplelely shut, tn ibe
latter ease it lias been suggested that the vertigo is mused by abnormal am-
ptillar impulse*, but these ran only contribute to the result which is in tbe
niaiu cauiwil bv direct disturbance of tli« brain. When the rolaliuu i* car-
jj
802
THE BRAI.V.
Bi<lv« into the nuclear Injer : or. »ut in anollwr wsy, ui«duiUu-<I film*
ing out of the uudear layer at all puinU converse tii funn ilie ccGttol ohlK
iBRlter. Some of these nbres, ss we have seen, beeiii ur end iti ihc relUrf
Purkinjj. None at them appear to join the cellti •>( tlie uucliMr layrr, inJ
ve linve no evHleDce that any of them end or bi^^to in any way in tlin nadai
layer. A certain number, Iiowe\'er, mav be setui to paw tlm>ugh the naciw
layer sni) Wtoet^n the cells of Putkinj/- into the molecular layer, wlwn Initee
their medulla they divide and apparently iwintribute In tlw numnoM Unik
of tlie molecular layer. The pTWUinjXiun, therefon, b that all iJie Ohmrf
the while matter bi^n or end atlier in tlie cells of Purkiujfi or tlM Ibribgf
the molecular layer.
The uipcrtic'ial gray matter of the cerebellum then rcMmUn the gny
nuHtTr iif lli<! tpiiial cord in to fnr as it oonniKl* of hmncbiDg nemnlU
ncrvv lihrwi, and nerve (ibriU imk-ddcd in neuroglia; but the di»f»«Ji"i
BDd f<.«turc« of the ncvcral furlorH urt- p««uliar. We may take-, jt-riiiijn, m
the key of the slnivturc the fihril* »t the molecular layer ; thii> layer i* nU
lively very thick, about 4(Kl f. murh thicker than the nuclear wlii''h. bi*-
ever, vnric» in ihioknca. Wing gewrnlly thickest at iho top of tlie UU;
hence the number of libriU in it may be spoken of ns enoratuUB. T^n*
fibrils •eem certainly to bo conooded on the one hand with the call* ■•( it'-
Duclear layer and on the otber hand with the scnitercd small cells u( timi
own layer; but we have no eridenco that cIkm two sets of fibrils an cna-
tinut>u« with each other : on the contrary, it aeem< more probable that iW
two eels of cells repnseut two independent syelems. We can hardW dmbl
that these fibrils arc in functional oounediou with the mednllated ^bra''
the central while matter; but we have no clear evidence that tlio fyiUiD
of Mattered cells is continuous either with the cells uf Turkinjf. and so aiik
lh« medullated fibres beloufpng to those cells, or with the iiiMlullaled 6bni
which end iodependenUy in the molecular layer: and we have no evidna
at all that the system of the c«lls of the nucktar layer is nKinoetcJ wA
either. We can hardly think otherwise than that Itie nulecdlsr efanga
which sweep to and fro along the tangle of these fibrils (whose nntiithah
frobably gorerned and hence whose functional svlivitv i» probably renUtsI
rlhe nuelear and scattered cells respectively) are intlut-uvud by or an|;tatfi
toe nervous imjMibea paning along the medullauxl fibnw of the nhiir malK:
and hence we must conclude that either a ounlinuily exintii which has m J*
esra|)ed detection or, what is quite iMsible if nut probable, that one SM
can act u|iod another hy nimple contact or even at a distmnc«. Paitbtf.
while the cell of Purkinjf-, niili its large celMiody and nucleus, its (MBi|ridi'
ous Hxi«-cylinder pr<ictM and its other branched proc»M>« prvaenls wmkJ
analogies with a motor cell, such us ihcate of the aiilcrior hum of tli« ijaMl
cord, and ra>M« tbo pmumntton that the im]>uttN» which move aloog i>
axis-cylinder proctM, proem outward from ihc rrll w motor or at leota
eHeiviit impulses, welmro no dirrvt proof that thi« is Ko, And thnui^itil
templing to suppoM that lh« other mmlullai<Hl fibres, which like the bbm
of a posterior not are lust in ihe grmy matler, nilhoul tW iolerveutlon «f >
coDspicuouB cell, carry atlcrcnt impiilani. we have n» yet do proof nf |1m>
All we can say is that tlie gray mailer is ciiiiiircled in two dilferent wsf)
with at least two sets of 6br««, which probably, thcrvfore, have diStnU
functiuDB.
We may here add (he remark ibat tlie large body of the cell of Purkisj'
lies, a* inileed do tlio other nervous eleiMDts, in an appropriate span «
the bed of neuroglia. Between the surfaoe of iIk cell ao<l tbe wall of phi'
roglia is a apace, generally eo narrow as to be potential rather than aciult
bill which may sunictimes be considerable. >\ hetber small nr large it r"-
SOMK HISTOLOQICAI. FEATCKKS OF THE DRAIN. 803
I
I
I
Itiiw lymfili, anil tbe cnvUy in whicih tbc oell IJet ii ui oonneetion nkh lite
lympluUHii of tbc bruiu. Ktich (till then Vit* in a lymph xpave ; but ve
tncroly mention tiii! pMrt now ; nc ihiill havi> i» riMurii In tbv niHtt«r wh«n
«• e(Hn« to iliuil witli ibc tymphalio ukI vuculsr uTmngemenU of Uie i>nuu
uid qdnul ounl.
TA« Cerehni CorUix.
^ 6ft0. While the euperfiml gray matter of the cerebelhim ioa not differ
ttrikin^ly m to its hiatu logical f<paUiK« in diflerent re(;ioi)s, very considerable
ditfert^ncea are obserTe^l in diller^ni ivgiooa of tbo nrebral cort«S. A )ten-
eral |>laD of structure may perhajw be reco^i^ed, but tts ne paas from one
|i«ri of the cerebra! surface to another *ie find otodilicatioDS continually
lakin)^ place. We uiuat content ourselvea here wilh attenipUni; a iJeacrtirtiou
uftlu! f:ei>erHl plan ((illoiDeil by an iitdicatiuu oflbe more striking cbantcUr-
utiai of certain rei^iunii.
Hie eortical gray matter, having fui average Uiickneaa of about 3 min., 1>ut
vmrying coundtradly in diflerent regiuns ttom 1.8 mm. in Mime parin of the
occlmtal lo)>« to -1.2 mm. at the dnmal aiimmit of the (irecentral convolution.
w, like other ^rmy matter, comiMmed of m-rve-cellA, and of nerve fibn.'n and
fibrib lupiKiried by nvuro^lia. The nerve-cclU, at l«iutt the uonxpicuoiig and
Muily raooKniit^l norve-s.'eltii. an- Hcatlfrcnl, and appear, in nectionii, to l>v
inibewlcd in, and »e(<oraleiI fnitn eat'h other by u not inc(inKi<Ifrabli' but
variable c|Uan(ily of itnnittwhat {ii'Ciiliar ground milmtanci-, not unliki.' that
which form* w InrK* a i>«rt nf tfi* mcilf'ulur layer uf the iNTi.7l>clliim. I'nrl
of tlib i^und nuhMtance, irbieh nnjiarcntly i« not cvmtiiuit tn any piirtifiilnr
layer, but vlrvtchon Ihrciiighniit tlie thicknt.'wi of ihfi (^irt^'X i« iindiiiilitcdly
omrtigliBl in nature, hut jmrt, und priibnblr the grpjiter |utrl, is nervous in
Dalurc; it iw largely com|t<iM-d nf fine tibrifi! tr«V(!r»ing it in various dirtc-
tioHH, tlie irangvvrw mvii^nin of thi^- lihriU giving it n characteristic doited
or "molectilar" appvaranci" ; nud ih<* ninjnriiy of ihwe fine tibrilaara prob-
ably the TOiilinuatiiin* of branthing ncrvr-cvlls or dividing nerve fibres, the
remain<ler Ix'ing neuroglia) Hhrils. Tn this r««i)cct it memblits llic molec-
ular layer of tlie orrehelluni, hut it !», to a mtich greater vxtent than is that
layer, tnvervcd by niediitlaled nerve lihro>, especially by fine medulUted
fibres like thoee wen in th« gray rantter of the spinal cord. § 564.
The iKrve-«etla imbcKldcd in this ground siibdance is more or leas distinot
layere an? of various kinds. The raoM conspicuous, abandant. and character-
istic nerve-cells found in tlifl cortex of all redona of the cerebellum, are tboae
which from their shape are called pijritmiiUu etUs. These vary very much in
utJi and have been dMin^uislied as "small pyramidal " cells averagiux 12 ^
in length by 8 ^ in breaitth, and " large pyramidal " celU, nmfllimee calleil
" iranglionio cells," of which the mcliiim size is about 40 /■ iu lenittb by 20 u
in br«dlli, Some of the tatter, occurring iu s|)ecial regions are of rery ]arg«
liie. 120 ;> by 50 A and have been calleil "" jjianl cells.'
The features of a " targe pyramidal " cell are very characteristic. Suoh a
edl nitpears in a well prepared vertical section <>f the cortex sa an elongated
itoaceles triangle placed vertii-uMy. with the \imv. lookiu^ t'lward the under-
lying white subdtance and the taperiug a|tex jioiniiug t>i the •iiirluc«. The
oell HubfUnce u finely grauulnteil or titirilluled. tli« fihrilliL- aweeiiiuji round
in vnriooa directions ; il not uafrcuuvnllj' cuntnins pignu-ut. In the nii<hit of
lhi> cell lubatance ratber near the base lit-n a largi-, ciciir. cDiispicuous round
ar oviti nucleolatiHi nucleus. At tlic haHc tbv <vll »uh«t«iK-e ih pmlongcd
illlw a nURib«r of proceMuai. One of thiiK', generally Hinrliitg from alwut the
midtlio of the ba«e, runs for Morae diataiice without dividing, and soon acquir-
B04
THE BBAIir.
ipea ni«diilla may be noognieed u an axia-eylinder proooe; ihelbrv
which it ijivM arigin sweeps with a noiv or l»8 curved couree iiit»tb«B
jnvenl nhile matter. Iii some instances llie Bxis-cylinJ^r prooav, kj
T-division like tliat seen in a f^nt;''^'' of^ * poetertor root (S 91), gliti
to two libT«8, one of which mav take a horixonlal direction ; in lotiie Rc._
of tite cortex, the occipital I'or instance, tlie axia-cirltn<ier proems a tuA b
^ve rise hv division to sevemi tibrea. llie other pmoesses JVom the but.
especiallv tfiose fhiin tbe aogla nf the triangle, rapidljr branch inio fin
fibril» u')ii<!h are soon loat to view in tbe jprotiiKl suMtanee. Tb« aiiei<f
the Iriungle is also prolemgetl 10(0 a pn>cew, wbirh, jc'vin^ ofl* fine lat«nl
braiiclKs, niHk««, as it were, i>trait(l>t for the s^irfnce. but ultimately bnac^
inj; into niii- librils la loat to view at some dislnnee fn>rii tbe tiody of tbec«ll.
Tbc cell Vu-r. in n cavltr of the ennind ^tilutance, which it apMan nonMUr
to fill, but from ibc wallj* of wbicb it Homeiinics abrluks. di'velopinr hHntn
itself and (be wall nfihe cavitr as[i«cc which may ooDUin not only ixnfh.
hut oct-nsiimidly WuoocytCH. (n preftarod tpcciaMnH the rMractioD «illiin ai
cavity of ibo artiricially shrunken cell mav tie oft«a obacrred.
The "small pynkniidal" cells have inucfi the Mint Mtum: that i«inMf,
IIk cells are charactfriscd by ihvir prrnmiilal fum. though this is nsianiJ/1
not w distincl, by ilieir verlioal |KMiino, awl by the poasvMton of bnartiii^
Erootcwe which are bwt in the moUi'idar ground subvlnDtx' ; thcprfMiK*.
Dwerer, of a niidbnsal axiscyliudtr process has not been clt«rtj iIvuh-
•I rated.
Otber nerve-cells are more like tbc ordinary nerrtscells of the ipiMl mnl
and of tbe internal cervbnil gray matter ; they are branched cells, of Imft*
lar, not of pyramidal, form, and for the niuet part small, 18<> by 10^ I'M'
may be eliarsoterixed by Ibe relative large siic (7 »i) of tbe nucleus, uddl
not poaacss au axidc-ylinder process ; ai least, such a prooan has not yet '
detnoDslralei). They are freijuently spoken of as "anpilar" cells.
Another kind of cell, tbe " liiulbrm cell." which is found in all repMSol
the cortex, baa a characteristic spindle-sbape. the cell subetanco hetoit pn*
longed at the opnosite poles into taperin);, ultimately hrandwd pmesn
The long axia of tlie cell is generally placed borixontally, following tbe vu-
vatiire of the cortex, and being Llius at the sJd€* of the sulci venial to Um
surface of the brain ; it is, however, at times inclined at various angti
Siill unoth«T kind of cell, the " granule cell," or " nuelear cell." b ana
which ihe nucleus n surrounded by a relalively small iiuantity nf cell
Ktunce, £1 ^ by 7 /i, more or Imt sphcn<ul in form in onlinary preparatii
but probably brenking up into dclirate branched procwMw. Cells nf thW
kind nrespamlyscatlvn-d throughout the (xirtax genomlly.but in [larticulM
n-gii(ii» — e. </., the occipital — are crowded togatfaer into a layer, which in
many rmficcts ra^:mMi!)i ihe nudcstr layer of the oervbetlum, and has h«
cnlli'd the "granular" or "nurlcar" layer.
Lantlv, itimughout the cunex are found, Ixsidm indubitable aerve-cdk
and inifuliilalilc Rfiirogliid eel In, numerous Nmatl, ennicwhal irregular cslh.
ccnwniiiig which it may be debated whether thej are rcsilly iiervmu or
Mmply iii!ur<>Kb"l )'■ nature. Honover, in using the namea given above fot
rite vuriou:! kintU of Dorva-calli, it must be remembered that many tratui-
tionnl fiirm.i an- ohaerved ; eells. for instance, may be scon interiuediulai^J
form Wlnt'cn pynimidnl <.<clls and " fiisifonn " or " angular " cella. ^H
The iurdu]lai<-<l nerve lihrra which take part in tbc cortex may be enP^
sidered provtsinnally ns forming two categoriee. In tbe first place, flbns
sweep up verlieally into iho cortex from the subjacent " centml while 1
niatt«r," taking at first 11 curved conrse as they enter into the gray matter,
and iheu np|M'4iniig to run straight toward the surface. Tlieae ore orrai^td
m
soys HISTOLOGICAL rBATCRSS OF THK BBAlX.
in tha dMpcr levels in hiindlf*. leuvine verticnl oolumnH at Iho grnr mauer
b«lwrcn tnoni: but at more siiperficiiiltevt^ia ibe biinHlt«BpK9id omniid Kre
cnuluallr lost to view. Beeidee tti<«e diMiocrt vert4cal litirra nod bundlen of
Gbm, of the (l^iiIla^v modullAted kiiii), which we have resMti to think are
the vimIm (or Ix^nnings) ou the onp hand of Kbra of ihe pedal and tegDiental
(yttcnu^ and on the other hand of libree of tJie corpus (.'allnatiui, or tlie other.
wmmWtirnl fibres spoken of as " afaociaiioa " 6br«a ',i. CHrij, an esceedinijly
Inrge mimber of fibres of the peculiar fine raedullftted kind run in rarioua
directions, forming a <ten»e network in the jrround subilanec of the (tray
mnlh^'r hetw<«u the eeita. We may add thai this system of fine uiedullatM
librfc is of late growth, and la not fully ileveloped iu man until two or three
ycare alter hirlb. Many of the niedtillateil llhree, coaree as well as liue, take
a horiioDtal direction parallel to ih« surfiii«, aud in certain regions an
specially <l6vek^>ed iii(<> a lajrer or into two layera so oa to form a horizontal
streak or Mreaka.
The raacular |>ta mater umwla cimidv, u ire hav« saiil, the whole Kurfaoe
of the cortex, dipping down into Ihetuloi; and from \l, a* in the case of the
spinal curd, proecMM oarryinj; h!nudve«eU un>l Ixnriuk' lymph ><[>ui:ei pa-in
inwant to Hipply tho fcmy inaCtor with bloiul. Ilia wliilc, aa wi: shall w-u
later on, the supply of hlooilvea«ela to the gray nintti.T i* roDsiderablo, tlio
truljr euuneotive-twuc vlnmeiit* of tho iiia nialer pron-xam an- fmn method
into neumg'lit. Imnudiatclv hi-ni.'«tli tlie nia miitrr furmin^ tliv iiniumluito
■urftee of tl»e cortex » a thin layt^r coiuisling of neuroglia only.
§ 6U. Tli<- nvr%'0-cclU of tho abov« nm'enil kind^ arc arrangml more or
k» dislinrtly iu l»yer< parallel to the aurfiicv, so that the whale th!cknn«« of
tlw cortex may by nicniu of them bo, more or less successfully, divitUxl into
« Mntn of lonos, one above the other : atid wc may, as we haw said, n'vng-
nbt on ibe one lumd a general arrniigenienl oomtnon to the whole stirfacD,
Ukd ma the other hand mod ill eat ions existing in theseverHl region*. The
generul arrMigoiiMint miiy be said to bo one of five layers or lones, usually
counted from the surface inward.
The fifth layer, lying next to the (-enlral whitfl matter, fairly uniform in
chamctera and thickness (about 1 mm.) over lh« greater part of the brain,
is cbaracteriied by the presence of eomewhat sparwlv scattered " fusiform "
cells, though other branched cells are preaent. It is broken up into vertical
columns by the bundles of vertical fibres, and its demarcation from the white
matter below ia somewhat indistinct owing to the fact that in the brain tlte
white matler, eapeeiolly that lying beneath the cortex, coniaina cells and
small gruupa of celb Wing between the bundles of flbree to a much greater
extent ihau don the wuite matter of the spinal cord.
The fourth layer, lyiug abure tlie preceding, varies much more both in
tltickiMW (0.&& mm. to 0.13 mm.) ami iu its cliaracterB. The constituent
celb are on tbe one hand lar^ pyramidal cells, and on the other hand
"granule" or "nuclear" cells. In ■'ime n-giona It may he aubilivided into
twu layen, the araall " nuclear" cuUh tii-iiig »> iiliunituut as to form in the
up|i«r {lart of the layer a M'iiarnt« Inyi-r cnlUil the "gmnule" or "nuclear"
layer. Thin fourth layer, Itkn [he pri-ctnliiig filVli layer beneath it, is split
up into veriic-al columns by the bundUu of vrriii-al fibril, but to a less degree.
li is marki^l in ibi loner part by a horiitonliil xlrvAk <luc to numerous, mostly
liDe. niedtillnled fibnn running horiximlalty. Iu ihi; c>irttoc of the island of
Rial this horixoiilal larc^r i« ilcvrlo|H'<l into a cotispiciioiis iihcet of medullatetl
fibrea, separating the rourih mid lirih layen hy ii iliHiincl interval of obvious
while matter. This fifth laver of fuiiifonu celU. thus ik-taclied fri>m tbi.; rest
of the cortex, is what is called the nhutfrum (FigH. IDS. I'M, r/.).
In Ihe third hiyur, the constituent c«lts are the ctiarocterislio pyTamidol
t'l till, bill froiu the usili-
itself mid the ^viill ufili.
lint occasiDiiiilly ti'iicoi'
ciivity of the nrlifii'ijili'.
The "cniiill ]iyriiini
the tells are ehiir:ii';r ■
iii'L 60 liistiiK't. I>v r.
liMfe^ws wliii'li in.
imwever, of ii lui-'
wmteil.
Other iicrvi-i- '
ami of tile iii[< . ^
liir, not of \n t :•
may l« eli!ii:ir
not pnee(.'!i> ill! '
(leinoiistniti <l
Anothi'v '
the ciiiiiv
)iill)reil al
The h.M-
vatuiT "1
surthi-i' .
Siil!
whifli ■
etiiiii'i
hill I'
kiii'i
n';:i
mil-'
t'ii' ■■
RATrniS OP THE BBAIN.
nam and In tlie monkey, tbuugli iml in some
nlilni) ill very definite clusUra in tbe motor
• ■•■■l region. In thit oovipita) TVgian
.. -Murth laver 13 very conipjouumt, miid
iiivJullAteil iibrea preBenl, tbrmn 11 while
: ibtt motor region, the nrnui)fi.'iui^iit it
ihtv<> i)cHcril>u(i H-i till! gi'iicriil [iTiui. Tbe
i.iii k''<:, iiH in aliiu tlu' ftmrtk liiynr; )>iit the
iiH'ti thinner ihuti in tlii; moinr rv'gioii, as
I'liirlh hiyvr. whilir thr Rl'ih layer, thiit o(
I'Tu, Smiiil '■ iiiiclfJtr " wlU arc iK-rbn]*
^'hijut nil l*y<T» than in tht? mi>U>r region,
111 ill thp ocopital rcgiiin.
lie iraniiitiiin iii «triK-turc from one region to
'i:tr]> nixl ili»liii<7t, dnil in pcrhn|« mpwially
ilor region Imrkwanl to tliv occipitnl rcgioD.
jiiiie htRtoTogiolly the limit, for inatAoce, of tbe
I fx peri mentally.
I ' Iiritin, lor innUincc in tho olladory bulb, of which
. I'fv grenl modihcationH of the general plan may be
,_. \Vo cannot enter upon those, but nnjiy just refer to
I iir bififtomnipUB. At the ventral end of the ternporat
a^wanipi, the atruclure of uhoie cortex follows the tceneral
Jrtiril »o ua lo project into the cavity of tlie desceodins horn
Evrlcle, fiiriuing the rid|{e-like prominence known hv the
liubstance of the cornu ammonis is therefore cortical 8ub<
the side uf the ventricle by a thin prolongation <if the
J*r, which i* in (urn covered by tbe ei>t^ndynia linina the
ticnl w-i-tion uf this substance &hows that while the tiHh
I nrv rcdiicMl tu ^mall dimensions, the thini layer, that of
Fcclls, in w«ll devoluped, though narrow. The celU are large
*]«np, Biitl the tajwriiig proi-eeaa are arranged »o n^gnlarly
t;uillr in iitatned pregmratioaa, to a marked radiate apjH^r*
■t of tlH' second layer there occura a large lievelopment of
ttb and a sair(!en««s of oells, giving riae lo a " lacunar"
^dihI ili« lir^t (ir m<i!ecular layer is uf some coDHtilei«l>le thick*
I Ihe prominence i>f the jiymiuiilat cells in thiii region, tho third
enemi plan of llie cortex ha« Hom«tinie8 been spoken of as the
' the cornu uninioniit."
he prvsent Htat« of knowledge it t* imixjiviible to conw t^) any
onclusion cnnovrning tho meaning uf thn vnriuty and arrunge-
FcellsaHd other con^ttitiicnte of the CJirtex. Tbo cells with their
nerve fibres and the nerve lihrils form a network of gray
we may compare with tbe gray matter uf the epinal cord
(btch is obviously, iw wc might qxpocl. far more complex than
We mnv conclLido, and experimental oMorvation oonfirnis the con-
, tlie large pyramidal oolls irilh recogniinble axi»-cytinder pro*
■B trophic centres for the fibrm which appear to slnrt from thein.
t may, though with Ices confidence, explain the large vize of thcN
kibe motor ref^on, by the fact that they give ri^fe to fibres of the
tract Mrett-hin;; a long way from their origin in the cell, and
demandiuji; gr«at nutritive aelivily on the part of the cell. We
ape ttlX) conclude that these fibres are efferent, motor flbra, ilee-
tnj impulttes from the cortex to the peripheral, or at least dtataot
806
THE DHAIX.
celU. TlicK are for ibe mrnt p«irl iur\cv, (luxi^h dimiiiUliini; in m
lieliiu' ii|>warrl. ami the layer Iiua I)c«ii rutleil i)i« " layer uf larjte jiyr
calls," lli(iii)(li in cenain ivt.'iuii!i tlw largM pymmldnl nlla, ana notaUyl
giaot-celli' an^ fotiml in (Ix: [irecedine. founb, layer. Tho celb are. on
wImIc, acattenil M)tnim')iat ii)iar><rl}r, ImKi^h Irwiutnlly f^utliureil into
ffr»up«, auA unionB tWm orcur iiinall "nuclrar" aiit) nthcr vvlU.
DuiMlliHt of Vfrliral filir!.-* r>{)rMi<l out rapidly id tbi* layer, ao ibat
culumnar arrniigcmi-iit 1wg"iui'« liift, unci many of lliv fibrni umlnublHilJ
Ik-odioc uxi»'Cyliii<K-r jin>cr«H'» of the fiyramidal cclU. Thoiigb llw lay
varitit in thioltm-iw (1 mm. toU.4 mm.), and in w^mv of ilx ft^tiirx-H in diffirnOI'^
re^on*. tlio chamctcrifttic pvramidal cvliK arc prcHiit over th« wIkiIc (urDuir
of the hcmifphert-. In tbc loner pari of the layer a MMnd hurixooial aimk
of cliMfcly interirovoD horizontal tibrr* freijncDtIr make* ita appaamwa.
Tile Bcconil layer, {•cuernlly a thin one, tlnni^h vnTvinij; from C- "' i ' >
0.7ij mm. in thicknew, is also formed by pyramlilitl cclli^ Ixil ittdi-
fruni (lie layer below by the nbwnce of largo and medium-siMti cclU ouU l</
(lie ptMonce of nunien>uH small <«ll8 cluaely packed together; it hw btia
called " ibe layer of »mall pyranmlitl celK" A« we have said. ihcM- tninllR
Iiyramidal cells dilfer tonii^wiiiil from the larger oella; and tliv ccU» in lb>
ayer are Bomolin)e« described ae ''an^fular."
The 6rsl ami mi«l uiijwrfiiiBl layer is i-hflracterieed by tlie pr«duainaMt
of Uie noleeular Kf^mud Bub»taiice. tite vella bein|[ few, far lietwoeD, Mull,
and irrefcular. The k""'""!!! subsinuce ilaelf seems to be iin)re Urgcly w"-
roglial in nature than in the other layera, and. as we Mtd above, its eiimur
surface appears to l>e furnislii-d by neuroijlia atone. The layer i« i^-iwnll'
spi>ken of as the " jM>rl|ihern] " or "auperflcial layer," or aometimia ae thr
"molecular" layer. The tapcriiix venlca) p«i>cest*«e of the pytaiuiilal crIU
may be traced into thin lever, wliicli indeei] varies in thickneas aceurdin^ lu
tbaabunduiice of pyriimiilal ccIIh in the subjacent layers ; num«raiu su*>-
what 6nir me<liillut«Hl tibriM atxi traverse it iu a honEuuial direction.
$ 662. Till- ^neml arrangenKiit juM dencrtbed variot, as we have said, in
dilTercJit r<-gi<>n* of the cerebral nirCue. We must eonieat ounelvca bnr
wilh pointing out the eluinict«rialk« of two or three ini|Mirlaut regions.
The region which w« have (§ tfSS) callcil Um' '" niotiir an'a "" or *" rppM,'
is charactcri/«<d on ihc one hand by the gnvt ihlckuuM (1 mm.) of ihr thini
layer, thai of Inr^ pyramidal cells, as acll as by I bo number aiiij siceoftlK
cells contained in it, and on the other lumd, and ctpcdaltv, by the prMni-
ncnce in the fourth Inver of remarkable clustere of very large pyraniiibl
cells, of the kind which are ntferr«d to above (§ OftO) as being friM|otuitlr
called "ganglionic;" ii is in this region tlial "giant celts" ura rmindin ik*
fonrih layer, namely, in the up|>er part of the proceutral and at the Hmniil
of the poetceninti convolution, and in iho |)anicc»tnil lohule, aoC|ulriflg thfst
grtUatt eixe at the lop nf the precontral convolution.
The occipital rej^ion a characterixed by the prominence of the " gfasalt"
or "nuclear" cells. These not only (brm a dMinct rlivbion of the foat^
layer, but are stiio conspicuous in otiier layers, their arrangement beiai; tart
tliat some anlhon> have been led tn divide the cortex of this region iutow""
or even ei^ht lay«-rti. In ihe present state of our knowledge we awv bo oiu-
Icni wici) in.iintiiijt that thii great mark of tht» uccipilal n^ion is ue abun-
dance of thi-w! nmall " nutdtrar " cells, toother with Other small " anguUt"
cvlk, whnn^liy l)<c |iynirriidul celU sieem to Iw made less con^pieuiin*. It i*
worthy of iiotii-e, however, that in the third, but more eapecialty in tlw
lourtb layer, n t'l^w er-U* of verv lar;ie um are met with, which by thrir
large branched c<-ll luhiitunce luxl conapicooun axtveylindcr pmotM rweaibir
the large cells in the motor rrgiim; hut it Uiuuld be notvd llinl whilMkos
SOME IIIfiTOLOaiCAL rKATITBSe Of THB BRAIfT.
„ cells occur (itl least in man Mod in ihv ni'^nker, (l)i>ut;Ii not iu s'>in«
ipf tlw lower KiiimnlH, sa the nbbit) in very definite clusters iu tiie iikHot
region, ibey oceur ain^lv ia the occipitfll r^ion. Id this occi|nl«l rvKiun
Ibo layer of hoi-iifintai Abres iu the fourth larer is \'ery oonajiicuoaiB, ati<l
jtwiar to the number of ordinwir medullmted nbr«e present, forma u while
pumk Tuible eveo tn the naked eye.
t In tlw firoBtal region, in front of llie motor reigion, the arrancreuedt b
inore ia aceordauce with wlial we have described aa the ^neral plan. The
|wo pyramidal layers are well marked, aa is also the fourth layer; but the
layer of large pyramidal cella in much thiiiui-r Ihaii in Iht: motor re^jpon, as
|b also, though to a Um extent, the fourth Inyer. while the lltlh layer, tJiut of
jfbaifonn celU, it tluekerthanuWwhere. Stuull " nuelear" celli art: prrhapa
■nore abuudanl in ihia rcgi»ii thn)ii;|!;h()Ut nil layer* than in the motor region,
but an (kr le» oau^ivuouH than in the? ocrlpitnl reifiori.
I We may here remark that llxr truiiaitiim in viniotnre from one re^cion to
■nother U very [^niiliiHl, nut Khurp and di»tuii;t, and i» p);rhnn» avprcially
Cradaal in piutiiii^' from the motor region bm-kwanl to the oc.'ei[M[jil resinn,
Pt ia not [HinibU- t^) reco|rDize hiMoTog^irAlly the limit, for inxtanec, of the
r*>r n-jpiin lu dvUTminea eipcrinifnlally.
In ais^tial rreiona of the brain, for inittanoc in the olfactory bulb, of which
fire ahnll «|K:nk uUor on, vorv grcnc aiodilicutioiM of the general plan niiiy l)e
Iphiierruil in the cort«x. We cannot enter uimn these, but mny just refer to
the comu ammunis or hippocampus. At the vontrol end of the temporal
lobe iKe eyrus hippocampi, the structure of whow cortex follows the general
plan, ie thrust inward so ne to project into the cavity of (be dowending bom
Af th« Inicnil ventricle, formint; the Hdgeltke prominence known bv the
Bbove Itame. The substance of the ooniti ainiuoni* is therefore cortioal siib-
Manco covered on the side of the rcntrii-!e by a thin prolonj^alion of the
^entral white matter, which is in turn covered by the cp<:iidynia lining tlie
IveDiricle. A vertical section of this substance shows that while the- lifth
id ftxirth layers are reduced to emnll dimensions, the third layer, that of
lUrKo pyramiilal cells, U well developed, though narrow. The cells are larM
' r«narkably lon^;. and the tapeting proeenee an arraOKed n regularly
to give tifie, es|>ecially in staiuM prepantioDa.toamaHted radiate appear^
loe. At the lerel of the second layer there occurs a large development of
pillary bloodveaidB and a ecarceiMw of cells, KivioR rise Ia a " tacanar "
.p[«aniuoe; and the finrt or molecular layer i* of some considerable ihick-
bcM. From tiie promioenee of tJie pyramidal celU in thia region, the third
^yer in the general plan of the cortex liaa Bometimes been spoken nf as the
" formation of the cornu ammonis."
§ 663. Iu tlw preaent state of knowledge it is impoastble (o come to any
MJtfacUirv eoucluHon concerning the meaning of the variety and armiige-
Deiit of tlte colli and other couatituenla of the cortex. The oelU with their
brancbc*, the nerve 6bres and the nerve fibrils form a network of gray
matter which ne may oumjMre with the gray watiex of the vpinni oortl
i% MO), but whi<;h ia obviously, m we might exnect, far mom complex than
llhat i*. We mav eoncludc, nml (■xperimental omervntion confirms the con*
Iclusion. thill the lar^c pyramidal cull* with recognisable axi^i-eylinder pro-
' asee Mrvc iw trophic cculrux for the hlim> which appear tii atart from llieiD,
nd wo may, though with hw cutifidi-nce, cxpiaiu the lai^r »jtt> >if tlwiw
b in the motor region, by the fact thai ihcv give rinft to fibrvn nf tliu
ramidal tract slretcliiii<; a long way fmm tdcir origin in the ci'll. and
ereforv demanding great nutritive activity on the |)itrl of the cniH. We
lay iwriiaps also conclude that iheae fibres are oAbrcnt. motor tilire*. >i(*-
' to csury impolsM irom the ooitex to tbe peripheral, or at Icuat distant
TnB BltllN^.
mechan!am Ibr the executiirn of thow mivementd ia rcadv «t hanl in itie
brain, wnilin^ only Ui hi d'txharfied ; ami it u euy tu ooouelve how »ii<rh ■
diNfliiirKe nii|clit oe iilfecled either liy Uw uibKitutiun for tliv urill of wmc
])(>tvtit iniriiiiic niTviviil irtipulM: or l>_v «uiii« niifdirnclioD ul* ToltUonal iia-
)>iiIm«. PenciiiA wbo ban- experienccl iimilar forcci] iuovenivDt« u llin
retull of dWttw repoM tliat they nrc freiiuciilly MicompAniod, niwl aoaai la
be CBUwd, b<r liiMurbLtl vUual or other sciualiotu ; thu* llwy ituribuU th^r
KiiililL-nly falling furwunl li> ihi^ (lociirrtucc of tiw KimtiaD ihdC thr Ki^mnd
in friinl of ibuin i» MiiMcnly linking away bonoUh their (mU ^^ ilbuut
IruDlin^: too cluHoly to tbn intvrprHuliniui thu xubjvcU of thaw dUanlvn fpwt
of ihcir own fi.*cliiig*, and rcmcmlxTing what wa5 saii] above OoncvrniDg Ttr-
ti)fo, «« may at Ii-wt coni:liiil«^ tbni th<> nnuaitnl nvivvmrntit art- in aiaaj
cues due tu n ■limirikr of the cucird inn ting nK-clii>iiUni. bmugbl abocii hy
stTBDge or disordered statmry im\m\»vs- Ami lliin vit-ir i» Miiipurtnd b_r Uis
Jkct ihnt many of thvne forcvd ni<jveincnt» aro aco(im]>anicd by n [icruliu
■ii't wholly abnormal piwilioii of the ryes, whii-h alonu might peramfm tx^Ham
manv of the phrnomcii:i.
§ 647. The phetiomrnn presented by animals d«prircd of ihoir ccrabnl
hemiAphiMTV show that llii# niuchinfrv of coordination is supplied by cerebral
structures lying l>^lwccri tha cerebral hemisphere above und th<; U>p of tbt
spinal ourd bi-loiv. But when ivc ask the further qucstinn, how ts thit
luacliinery related to the various elemeDts which go to make up this pan
of tJie brain? the only answers whieJi we receive are of tiie moat imiieriiwi
kind.
Jd the case of the fro^ we can, afier removal of the eorehral heniiitphi^rc*,
make an experimental dutiuction in the {lartJt left bvtwci^ii the optic llialoaii
witJi the optic nerves and tracts, the optic Inltee, and the bulb with the ruili-
tucntary ccrcbi-lluni. Wtien the optic thiilami ari> removol, as might bs
cx|H'4-lL-d, tbu evidence of visual imprca*ioiM muilifying thu tnorcmenta of
tlie animal diMpjieant ; and it is .stated that apparently 3iponUi»<--jus mow-
tuents are mudi more rare than irbcn llit- ihalunii arc intact, Whi-ti tbe
optic \chtt B» mil lu the cerebral bemi»i>hcw-« are retiiuvc.il, Uio power of
udanciog is iMt ; wlien itiich a fro^ is Ihmwn otT its biihineo by incliniag
the plane on whirh it in phut'd, it slips bock or fnll.t down ; the special o>-
ordinaling nifchuDism for tuaUncing must, therefore, in this iiuimal havst
efwcial connection with the Ojttic lomM. liui nflcr removal of thcw oritaiM
the animal is still cupalile «f a great variety of cuontinate niovecnvnu;
unlike a fmg retaining its spinal coni only, it c«u swim and Icnp, it miiB-
tains n normal puMur*.-. and when iilaccl ou its baek immediately rcgaini
the normal posture. The cercUclliim of the frog is so small, and in n-
moving it injury is ki likely to he done to the underlying parts, tiiat it
becomes ditlicult to ny how much of the coonlinatiov apparent in a fny
noasctning ccrchclluni and bulb is to be attributed to th« former uriotlic
latter ; probably, however, the part playol bv the former is small.
Ill thu case neither of the bird nor of the mammal hare vc any rxMl
iulorniation as to the behavior of the animal after removal of iIm) poitt
behind the hemispheres, in addition to the hemispheres ttieiuselru. Our
knowledge ia continetl to the results of the ablation or of the Btiiuutallun t^
Earts, tbe cerebellum for iustance, in animida in which tbe rest of the brain
U been letl intact. Ubeervaliims of this kind have di^eluKd many int»t>
eating ftcts, besides the forced movemeuts juAt referred to. but Ihcy liiive uiil
led to, and indeed could hardly be ex|>ecled to lead to, any clear views as to
the point which wo are now ilisL-ussin){. It doa not folluw that every ]«rt,
injury, or stimulation of which interferes with coordinated muvemt^nts, or
gives riae to de6nite, forc>ed, or otiter niorementa, ii to be cunsidored as pan
I
I
I
of thr Diacbinery uniler coiuiderntioii. Tlie cnrponi stristft and cerebral
hrniu|>b«rt* ruriu, lu vr« ha\Q aceii. u<> pan of ih« luaobinerv. yet injury tu
llim) mny (li.i'trtk-r ibi' mAcliint-rv ; tiiic) lb« fact lliul reniaval of ur injury
tu tbe ctrcbc-Uum •lisonk-ra the nmoliinerv a do pruof by iUelf (bat tli«
co/rlii'llutu i* lui uaeutia) |iart til' itid luucbiuery.
] f nv may Irum t» iloluclionii fruni alructurul arniiijteinenU, ira mi|{lit be
incliiiril to infer llint the luiatoRiioal relatioutt nf what ve Iiave milvd the
Ici^M-iitnl rcKtuo from tli« bulb unniirt) poiiil to ita scrviiij; at tlw fouudiuioD
ol llii' iDiicliiDeiv ID (lUMtioil. twhiud it hiu full conncctiun^ with vaHitua
|mi(n ui ih* rani, while iu iVunl by mtaiu t>f Iho otitic ibulumi and atiierior
Nrfxira <iua<lrigoioiiia. if not by other wuyi a« k«1I, it tt itu far luuoL-iatutl
aillt tliu optic lu-rve* that tbc pricb »i«nii njieu fur visual im[>HlM» Ut fniin
si-cv-M tu iu Tu thia luuiitiatiQii. Iidwcvit, vr« muNt ad<l thu nrcbdluui, nn
aivouiil »r i(* rvliiliont to it, to Ihu iNird, and ti> the lulb through th« nnti*
form If xlicf. inoludinz ila tin with ihv iiudilory nervt:. And it' we add the
cvrpln-lltini WK miiat also probiihlv add th« |>i>ii:<. W« may sxcluda tli« pea
of thi- cTiii. Finer this ic compoM-d cxclu»ivcly nf Gl>n« bringia^ tbe cerebral
I>dni<[ih<'rc8, including iIm oonxirii ctrialK. into ci>it»<?ct)on with thi? |>i<n<i,
bulli, ami conl, nnd m with tbc cuonliiintiiig niiiL-hiiii.Ty itaclf, n* well n*
with i<lli«r {wTis of iho nervous mtcni. And ob<crralk>ii. im fat i\» it gov*,
auii|Miri8 this iJtduclioit from nnaiomical relationship*. We will, however,
iitivT tihiit eiee wc have to say on this point until alt«r Kfi hav« dUcuncd
tile carrying out of VoluDUry niovemcnis.
Ox SouE HnroLooicai. Fkatureb of the Bkaisc.
I
I
$ 648 TIk while matter of tbe brain, ns we have niremly said, like lliat
of Ihv >[>iiial cord, comi*taof medullatcd fihrca of varlout tize* imbcildctl
in ni'uriigiia niii) Hui>|H>rl(M! by iM;]tta of eonnvclive timue (k-rivud fnini the
pia oialrr. Savo loat cvll>, or even gronjin or ron* of (.-i-IU, fur the IDOK
part vmnil e«lb, about mmtv of which it may be dcbalctl wlu:tlivr tbey U»
aervfe-colb or neuroglia ei,-ll*. arc fiwjUiMitly nccn between the filma ami
bniidlca of fibres, the white loallcr of the brain M«ni tncoiially iileiitic«l
vith that of tbc Rpinat or^.
The gray matter of the brnin in etDeral also cormponds to tbe gny
Rtattcr of the cord in coiiMfliiig of brniiching ne^vc-ccll^ fine inr<liitlau>il
fibm of |>rciilinr nature, noiMiirdullatcd Kbm and lihHls, with a Ivw unli-
narv nicdulhiteil tibm, all eupjMirled in neuroglia.
I'hc " ocotral " gray mnlter u cxtrvmely like that of the ootd except ibal
the ncrrou* elemeno arc imbedded in a relatively larger ({uantiir of m-u-
roglia. Immedtatvlr undenicath tbe epithelium lining ihceei'eral vciitriclca
and tbe aqueduct, tbe neuroglia ia especially develo|>ed, forming a distinct
layer whicb may be rcgiinU.^ at a continttatiuu of tbe central g«hitiuous
aulMiuncc of the spinal cord, and which, with the epilheliura overlying it,
forms what is known as tliv tpemtyiMa Tbe " nuclei ' of (be cranial iiervw
are, as we liave seen, companible to tbe groups of nerve cells in tlic si^iutl
cord.
A K'^*' <'^ '^^ ^^^ ff"'T matter of tJie brain may be apokeo of as more
"ditliivc" or " st-attered." more broken up by bundles of fibres than is the
case in the spinal cord. Tbe "reticular lonnatioD" of the b.ilb and of tbe
iegai«tital rei;ioii is an eilriiue furin of this dilfusc t;ray matter And even
in web collediotia of indubitable gray maiivr us tl»e enrjMi« sirtatiiin, uptio
thalamus, and the like, the pure gray matter, if we may um tlie term, ia
much wore interrupted aiM broken u|i b/ conspieuoua bundhi* of white
THK DRAI^.
flbrefl than is the case in any region of thp sfiinal cord. In tUc oorpon
quiidrigcniiDB to Uw gray iitait«r is broken u|i by »l»e«(a or bundles of mhkt
matter.
The oi^rve-oent of the aeveral ciitlectiooe of (;ray matter are not all dika;
ihey present in dld'ereut re({ioni( ilitTt^rences in si<e, furm, and in other tftu-
sclera. The cells ot' the Ducleiix caudatus, for instance, arc rather small ti>d
oDeu round or spindle-Hliaped. whil« ihoee of tb« oulk- ihslamits an larp^
broDchcd, and rich in pi^'iiicnt. The cells of the sulwlnutia uir,tn ui
■pindle-aba|ied. of inod«r»te xuc. mid mi luadtd with blaek pij;nient (iu iiiu)
as to jtuiily ti>e name; thoiw of the hioui ai^ruleiiit are very lar^e and
siiberi<-al, iviih juet so much {%iiiGnt as tn give a bluioh tint, liut nui
knowledge of the lioer hiMulogioi) details ut ihe rarioua mosaee of k^J
matter is at present too impvrlect to aflbn) auy bsfiia whatever fur jJir^
logical deductions; and it will be hanlly [imfiinhte to dwell u|wiu tlicK
Two regions of gray matter alone e«II for ipeciul CMcrifitiou, the conn
ccnhrl and the superficial gray matter of the cerebellum.
The mperfieiai yrajf matltr 0/ lAe cenMiutn.
g 649. The ifurfacc of the cercbolluni is increased by bdnc fgldsd «
plaited into Icnfliku folds, and c4tch of ihcw primary Iblds U tinllulT
folded into a number of secondary, nlso leaf-like, fbl<l* or lamttUn. Eua
of ihcM lamella) consiets uf n central core of while nmltvr. the fibre* vT
which pn«8 inward to, and cuiilrihiile to form the ocairnl whice niatlrr of
the (i'n.'hclluin,aDd of a supcrlieial layer of grar omtlcr. A M-ctinn thpLOjti
a laniclin perpendicular to the »iirliice shows that the gray matter codmu
essentially of two layers: a layer Iving next to the white matter formed !«■
denevly crowded small cells. ealW the nuehnr layrr, and l*tween this sad
the tiu|ierficial pin mater a much thicker layer of peculiar nature, aUledlke
mottcular laiicr. Detncen these two layers, and connected, as we sball M^
with both (j1 them, lice a row of ver}- larure and remarkable cells, called dii
k/I* of Fvrkiriji, the bodies of which abut 011 the nuclear layer, and lbs
loiig branches of which traveise the molecular layer; these cells so ptiesd
may be Mid ta constitute a third layer. Before pmcce<liu]r further, we maj
b*re remark that a section uf the lamella, that is. one of the seeoudary, bM
one of the pHmarv, folds, while Htill remaining a vertical section (tbsl it
perprndicular to tlie surliice) may be carried through the lantella in dillmat
planet, and ibai of these several plation, the Mictions Ukeu in two uf ihcia
am e«[H'('ially inittructive, onmely, the one taken in what we may Mil tk
longiinilinal plane, pasfJng fKim'tbe lop «f the lamella to i|s baae, anil tht
one taken at right angle* to the farmer, in what nc may call tl»e tnuunroae
plane. The nuclear layi-r and the molecular layer pn-eent lb« snina brwd
IVntumt in both Imigituilinal imd Irnnsverse n^-iiunn, but the long bmncbcd
processes of the celli* of I'lirkinj^- >.ince tbcy run in the tniitsveree nlaue an
iidciiiialcly fven in iransvctw: sections only; longitudinal seclioiu soow only
their proh'tcs.
The molecular layer is of a peculiar mlurc. lu manv m<Nlci> of prenn-
lien and in ntauy sections it appears chietly composed ol'a granular or avttel
groutKi Euhslance; hence the name molecular, na if it were an aggre|[Klioa
of molecules. The dots, however, are sections of fine fibrils, some of wbtcb
are neuroglia fibrils hut otbera are undoubtedly nervous. The layer eoOMlt
in fiict partly of nervous elements, and here perhaps even more than elie-
where it is extremely dilBeuIt to say with regard to many of the eli-tueati
wbrtlicr they are neuroglial or nervous iu imiurw. A considerable {Mrtioo
fAL rSATCBES OF THK BBAIX. 801
of ibe wholt ana of the iiifili.-culiir litvur U lakeii up l>r the cniispicuoua
bnntchtd prwCMiM of iW wUa i>{ Purkiuje ; «u>] KcatlcnHJ about lie numer-
aa* iinntl cell*, tame of irhidi mi* oeuroEli* oella, but >om« of wl)ic)i are
iit)d<>iil>le>llj Derv«-celb. The moit conaMCtiouj feature of the \&ycr, boir-
KTor. M tlw pmeon in lar)^ ntimliera of tli« linu fibrils ; but before we *p«ik
of thcw it will be diwntble to turn to tbc cclU of PurklujA nud the uuclcar
Isrer.
'The »11 of Purkinj* ponxwn n Urge (40 ■ by SO »<) flatk-^haped I»odj-,
nirroiiMding a large, cum imciioiml, elm r, rounded iiucI«uh; it ha* oiucb ibo
apprnrance of n larzc giinglioD evil. The biMv nf the fliwk nal* on the nuclear
laTrr. ni>d frtiRi it ihcrc proceeds af ingle nxiv'CjliiKU-r praorai which pwaing
throurit the nackar layrr foioewbal •ibli<)urljr, nii<l in il« piiatagi* nnguiring
a BMOaila,3oJiia the central uhiir Hihoinnco ns u mi-ilnltatoil tilrn?. The cell»,
■a wa have aatd, form a ainglr laver only, hiii riavr ihin vovcn the nttclcar
layer over the whole of the lamolln, a cunvidcnible iiumlx^r of the (ibru of
tM white central matter, tltotigh imlv a vpn- (mnll fracliuii of iho whole, ara
Uina ilenved from lh«ae cells of I'lirkinji^. Tbe niirn)»r<l nt-rk of the flaik
niDiiiD^' outward in the nioleculur layer divides in an arboraceni fitehion into
a lar^c niiiuber nf bmnclieB nhicb. fepreadiug out laterally in iIk; tniii»verie
itlane and iitretchini^ a* far as the MUrface, rnniifv through (bo tnolocular
aver, aiid are eventually lost to view aa exceedingly line fibrils. Some
ooterTers maintain ihu aoiDe of the line proceosee are continuous nilh pro-
e»[B of the small nerve-eelltof the innleeular layer, but this is not a^liottted
by all. In any case llie fibrillar terminations of these cells of Purltinjt^ oon-
tribute to the iine jjbrils of the molecular layer.
The nuclear layer in ordinary stained specimens has the appearance »f n
nan of nuclei clueelv crowded totfetJier in a bed of reticular nature ; aud
•inco tlie nuclei ttnially stain deei>Tv. the layer Htands out in stmng conlraat
to.tha nnch leas di-eply nluiootl mulcciilur layer. Careful emniuaiivn with
•|»cSal modci of prcpamtiun >hi>wit. bowcTer, tlmi while wnM of the nuclei
an nuclei bdoagiog to nL-tiroglia and bloodvcnwU, ilw majorilv belong to
iinall iwrve-fialb of a pi-culiiir nature. In tboc cells the nucteuM is sur-
rounded by cell solMluncr mhich fiirniing n thin layer iinmedialvly anmnd
the nurleiii<.fschteIlydix|Hi*f<las thin r]>r«uiting bmnchc«,woi« of which end
in a i>ci-uliiir nrborceccnix- not untiken mii»>?le rn<l-|>liilc: thcee prooevoe oon-
iributc nilb the neuroglia t" fnrni tlu- rcticu la r-lo« Icing bed spokm of above,
Noprucewctin be Iruecd inwiinl to the central while mntler; but one of the
pnoetees gives olf a branch, which piissing vertically outward lakes on tbe
appearance of a delicate axi> cylinder prooeea and runs, without dividing;,
into ibe molecular layer for a Ttiriablo distance, aometimes reaching clo«e to
llw mrfacc, but nt last divides at right angles into two Hbrils, which mn in
the hutgitudinnl plane in opposite directions for a considerable distance, auil
are ultimately loet to vtow. ^incc theae cells in the nuclear layer are very
numerous nod each gives rise in the above manner to longitudinal 6brils, tl»e
molecular layer is traversed by a multitude of librils, visible as such in lon)n-
tudioal sections but appearing as dota io tronsvene eeotioDS, in which &e
cells of l'urkinJ6 are bc*l displayed.
BeaJdea theae longitudinnl librils proceeding fmni the cells of the nucleur
layar, epeeial modes uf pn.'|iaruti'>n HimiUrly diH'Kiee numeruuB traiMvenM aji
well as more or lees obli<|ue librils. Mitny of l))»c »p|>ear to result from the
branching of the small nen'e-cells of the roiili-mliir layer, and soow of ihoae
ao ariviiig descend ti> tbe layer of the cella of I'urkiiij<!- and end around the
bodiiw ol ibwe cellii in n-nnirkablc neat* of fibriU, williout, however, actually
making connwlioiu nilh them.
The meduUatvd fibre* uf Uw ovutnd nliito matter of a lamella pas on all
TirS BRAIX.
sidee iata the nuclear layer ; or, nut in uiather wsy, mctlullatt^ ftbrf* pai*
iiig out of th« nuclear layer at alt poials converKC to rorm the ctnuml •hiic
matter. Some of titeae fibres, us ne have M«n, bi^ti or i!!i>d in the cWltor
Purkinj^. l^oue uf them aji|icar to join thv ccIIh of Ihe nuckar lajM*. and
we have no evidence that any of them end or begin in any way iii thr nuelav
layer. A certain number, hdwcver. Hinr l)C *«vn to paw ibmngh tb* noctrai
layer and between tlie cells u('Fiirkiiij<'-:n to die mulcrular tayt-r. triicrt Unnc
ihdr medulla tbey divide and a))purcutt)- cunlribulc to thv numcroa* fibrih
of ibe molecular layer. The prcnuroption, iliercforc, i« that all iho libn> of
Ihe white matter by^in or end eiilier in tlie cells of Purkinji ur tlw fibrili of
the molecular layer.
The KUperGcial gray matter of the cenrbclium then naeniblca the pmj
matlt-r of the »))inal cord in so far of it canRtila of braocbiiif; nem rcll^
nenc tihnai. and nrrvc iibrilii inibi'ildrd in ncuroclis: but the diapoaitiM
and fi-alurcM of the »vvtTal foctora nn- peculiar. We may take, |)erlMfa.M
th<ikcy of the ilnirturc the Bbrilsof the mdleciilar layer; this layer is reU-
Uvcly verjr tliick. about 400 >, much thicker than the nuclear which, bw-
ever, vmrica in thickncH, bung piutrally tbicknt at the top »f the Ml;
hence Ihe number of fibrils in it may be epoken of as cuormoua. Hioe
librilti acim certainly to be ci>nn«ct«d on the one hand witli the cells uf ike
nuclear layer and on the other hand with the scattered ainall c«11b of tfatir
own layer ; but we bavc no evidence that these two sets of fibrils arw ooa-
tinUQUS with each other; oa the contrary, it seetin more probable thai tbt
two seta of cells represeut two indepeuHoot syMema. We can banlly doult
that these librils are in functional ouniiection with tbe me«lullaled iibre*'/
tlie central while matter: but we have uo clear evidence that lite iiT*t*m
of scattered cells is continuous either with ilie cells of Purkinji-, nud *o witk
tlio medullaied fibm belonging to thoie celU, or with the luL-^lulhitt^ tibrw
which end ii>d^DdeDlly in tho uiolucular layer; and wc have no evidtao
at all tliai tfae srMcni of the cells of the nuclear laver u connected >itb
eitlwr. Wo Cflu tiardly think othemiie than thai llie molecular chants
which sweep to and fro along the laiiglu nf thtvc fibrils (whoae nutriuua i*
EitlMibly governed and hrncc whoao Atnotional activitv » probably r^uUied
Ihe nuclear and scattered colls r«»pMtJvcly) arc inducnccd by or originate
) nervous impulses pauing along the nH)dulla(4.-d tihrcs of tbo white matter;
and henc« we must conclude thai cither a c<Miliniiity exists which has as vei
««rai>e<i detection or, what is nuile possible if not probsblc, that ooe SGvi)
mn net npon another by simple contact or even at a disunce. Fuitbcr,
while the cell of I'urkinj'^, with its large cell-body oud nucleus, its oaiia|ilra'
OUS Kiiaoylinilcr procew and its other branchml proceeds prceeota nuuiy
analogies with a imitor cell, auch as thura uf ihe anterior bom of tbefptsal
cortl. and raises the preaumption that the impulsn which move along in
axi* cylinder procesa, proceed outward from the c«ll as motor or at lea*t ai
•Akivtit impulse*, we have no direct jiroof iliai this is io. And though it it
Umptiiig 10 luppoac that the other medullatcd libres, which tike thr libra
of a noatcrior root an loat in the gray mutter, witlioul tlie intervcniran of a
r'>iH|>iouoiu coll. earry aflVrcuc impulse*, we have as yet no proof of ihia
All we can say is that lh« gray niniier is eouuected in two otflenut trays
with at least two sets of Bnm, which probably, therefore, bave diOknuA
funi'iioiu.
We may here add the remark that the large body of the celt of Pnrkio^
lira, ns inlle<-d du the other nervous elements, in an appniprialo spacv in
ft >.: I :,f neuro^'lia. Itetween the surface of the cell and the wall uf nea-
II siwco, generally so narrow as to b« [Hitential ralhcr than actual,
loll uiiK'h may sometimes be consKleTablv. \N betber small ur lai^ge i| ena-
SOME ntaTOLOGICAL ?EATDRKS OF THK BRAIN. 803
imino Irmpli, awl tlic cavity in which the cell li«t ia la connection wjlb ibe
Itmiiliatica uf llio liniin. KhcIi cell then li«» in s lymph-»pac«: Itut we
merely menti&n tho fnrt uow ; ive nball have to nrltirii to the matter when
w» eomo to <Ich) with ihv lirmiiluklti; luul vascular urmDgcnieiitn uf the bruiu
■ltd splaal flord.
TA« ftrrrfm/ Orfar.
$ 660. While (he soperiioiHl gray matter of th« ccr«belliim does not differ
atHkinKly us to i in bislolofjiml fcalurcs in i.lifR.-rent rv^ions. very considcfnblc
difleffncn are observed id <liircr«iil rcgiooe of tltv cerebral cortex. A gpn-
ernl plan of Htructurv inny perhaj^ lie i«ro|;nixed, but as we pns§ from one
purt of the cerebral surface to another we Biid n)o<liticstionB continually
lakinj; place. We niuflt content ounelvea here with atleinptiit^ a deflcripiion
of ibe funeral plan followed by en indicatiiHi of tli« more otnking charucter-
latit* ot certain regions.
Tbe cortical grav matter, hating an average thieltneee of about 3 mm., hut
rar^'ing Considerably in different regions from l,» mm. in «>me ]Hirl« of tbe
occipital lobe to A.'i mm, at tbe dorsal summit uf the preoenlral convolutiuoi,
t», like »th«r K^ay matter, coni)ir«e<l of nerve-cells, and uf nerve fibre* and
flbrilt supjHirtci] by neuroglia. Tbe nerve-cells, at leasi the oonspicuous and
«a*ily rtco^lxed nervo-ceiU, are scattered, and appear, in aectiona, to be
tmbwided in, and scjiaraled fWrni each other by a not inconaiderable but
variflbW iiuantity of somewhat peculiar ground suWtauce, not unlike thnc
which furiiio «n large a jiart of llic moleciilar layer of the rereliellum. Part
of this ground «nb>ilnnce, which n|){iart.-cilly i< nut confined to any particular
layer. 1x1 ( xtrftcli'w thrnngluHit tne tbiekncn of tbe ciriex U undou)>tcdly
neuroglial in nature, hut jiRri. and probably th« gn^tto* p^rt, i« nervous in
nature: it is largely c<ini|><F»cd »f fine fibril* lmTen>ing it in variouM dirtw*
tioRs, the transvcr«e socliona of thtvc libnU giving it a characteristic dotted
or " molecular" apiK-nnince : nnd thtt majority of these line librils arc prob-
ably the continuations of braiiching nerv»ceJle or dividing nerve librce, the
remoiwler being neuroglial fibrils. In this Ttamct it resembles the moleo
nlnr layer of the cerebellum, but it is, to a inuon gmter extent than is that
layer, tnivcned by niediiltutcd nerve fibres, especially by tine medullated
flbree tike Iboec seen in the gray matter of tbe sjMnnl cord, £ o64.
The nerve-celts iiubeddcd in this ground substance is more or leas distinct
layers are of varioas kin<U. Tbe most conspicuous, abundant, and character*
istic nerve-cells found in tbe cortex of all regions of the cerel>eUuni. are tliose
wbieh from their shape are called pymmi't'U ceti*. These vary very nmcb in
•iiM and have been distinguished as " small pyramidal " cells averaging 12 ^
in lentrlh by 8 r> in breailth, and " large pyramidal " cells, sometimes called
" ganglionic cclb," of whieb (be medium sixe is about 40 ii in length by 20 m
io breadth. Some of the latter, ucciirring in s)>ecial regions are or very larjfe
rise, 120 III by 50 *> and have been called "eiAnt cellii.'
Tbo features of a "targe pyramidal " cell are very characteristic. Such a
t*\\ nnpeara in a wet! jin-'imml vertical section of tli« cortex as an clongatod
niiMN-liii trianglu plaeeil vertically, niih tbu Ivasi- looking toward tlM under-
lying white tubiluncv and tlie ta|ieriiig a|ies |H)inting to thi^ Mfliica^ Tbe
cell subataace b finely gnnutated or filiriltaiol. lh<; fiTirilb- nwecping round
in various dirfCtioiu ; it not unfreuuentlv c^nlainn pigmrnt. In the midiit of
this cell siibntann- rattier near the nose Ik» h large, cli-ar, conspicuotu round
or oval nucl«ilatr<l nucleus. ,\i ibig base thr veil iiulwtaAC* ia prolonged
into n number uf proccsM^ One of tb(.-ae. g«nenil)v ulnrting from about tho
middle uf the base, runs for somo distance without dividing, and soon aequir-
804
TUB BRA1.V.
ing a roedulln may b« recngniud » an axis-cylinder prooew; tba fibrp lo
uliich it |iv«s origin sneepA with a moir or I«m curved onuree into Uw hI>-
jm-eiit white matter. In suiue instancee the axu-cylimlcr prooen, bj a
'T-(livi»iuD like that eecii in a ganglion of a pDet(>rior root (^ 97), givwriK
lo two fibres, one of which may take a horizontal ilirodion ; in sotoe renoiu
of the cortex, the o(.-ei[>ital for inAiance. tho axbcytinder prncea baaiilla
give rise hr divisiuii to eeveral librfs. The other praoeeBee from the bat,
njiecially inow from the angles <>f the triangle, rapidly branch tab) flat
libril* which are soon lost lo view in the |i;raund Nibstance. TheaoexW
tlic triangle in also prolonged into a procem, which, Kiviox olT fine latonl
branc-he>, nnilce^. as it were, strait{ht for the surfaoe. bul ultimately bniteb-
ing into fine lltirilti la lost lo view at aooie distance from the body of the ail
The cell lit-* ii) a cavity of the (;r<vund aubiiluDoe, which it appears nomallT
Id fill, hilt from the wallH of which it sometimes shrinks, developing l>etiien
ilMlf and the wall nf the cavity a «)»cc which mxy contain not only Irmpb.
bnl occmiioiiaHy Irucocytce. In prcjiareil apeciinens the reiracliun vitliin lO
cavity of tli« artificially shrunken ctlll may be oflen observed.
The "small pyramidal " cells have much the same features ; that (■ to ist,
tlic cells are characterized bv their pyramidal form, though this U milunlljt
not to ilif'linrl, by their vertical piMilion, and by tlic pnwmion of braDoUlf
proccfBCf which are hot in the molecular f^round «ub«lancc; (h« ptfWDcc,
flowever, of n midluiMLl axi-i cylinder pn>i.>M has not b(«n clearly ilomoS'
et rated.
Other ncrvo-cclls arc moro like tho orrlinary nerve-cells of the apliial »nl
and of the inlcronl wnihral grav matter; they ar*- branched cells, of im^-
tar, not of pyramidul, form, and for the mo»t jinrt itmnll, 1H>' by IOf>. Tbct
ntftj be dujacterixcd by Ihv relative Inrgo sine (T n) of tbo micleua, aarf w
not po8MM an axis-cylinder process ; at least, »ich n praCMi bas tkot fet bto
demonstrated. They are fmiDently spoken of as " angtilar " cells.
Another kind of cell, the " liinfoTTD cell." which is found tu all n^omt^
the oonex, has a eharscieristic spiDdle-ehape, the cell substance being |>r»-
Iniiged at the oprioeite |Kiles into tapering, ulliniately bmncbed pfoeeso.
The long axis of tlie cell is generally placed horizouially, following the cut-
vuturo of the cortex, and being thus at the sides of the sulci vertical Ui Ibt
surface of the brain ; it is, however, at linws inclined at vuriotia angles.
filill another kind of cell, the "granule cell," or " nuclear cell," isowis
which ihc nucleus is siirmunded by a relatively small ouanlily of cell nib-
BUuioa. 9 !• hy 7 ^, more or Imb spherical in form in ordinary prenatatioDi.
bat probably breaking tin into delicate branched procaaea. Cells of ihii
kind are sparsely scaticnil throughout the cortex generally, but in particuhir
rvjiioiif^r. y., the ncctpital — are cniwiletl together into a layer, which ia
many rcKpccIs rovmbl(« the nuclear layer ul the cenbcilum, and hsN licea
calkVl the "granular" or"nuchjar" layer.
Lastly, ihrou^diout the cortex are found, bcddde* indulHtable iMmr*«sili
ami indubitable neiirriglini C4-lti>, numerous small, aomowbai irregular cclk
conceniing which it may l»e dchntc<l irhelher they are really ncrvota q»
simply neuroglial in nature. Moreover, in umag tlie name* given above for
tite various kimU of ncn'ccclls, it must be rvmemborcd that many tninri-
tional forms an observed ; collf. tor instaoco. may be seen interniediatt) in
form between pyramidal cells and "fusiform" or "angular" cells.
The medullsied nerve lihm which take part in the cortex may be coti-
eideivd provisionally a» forming two calegorica. In the finti place, fibres
SKcep up vertically into the cortex from the subjacent "central white
mnttcr," taking at fim a curved roune as they enter into the gray matter.
and then ap|)earii}g to run straight toward the surface. These are armngctl
AOUE HISTOLOGICAL FKATUBES OF THK BRAI>*. 805
the dcep^ levels in iMiDdlfv. leavinj; verti<.'n1 columna of tlie |>nir miitler
belwcou iDCtn ; but at more Eiijierra-iul k>vela ihe biin<lleB s|iren<l out aud ura
cniiliiKDy loM to view. Uraidee (hue dixtiiict veitical fibrea uid buodla ot
nbm, »f tlie ordinar;- roeduDnted kiod, which we have reason to think an
Ui*«ads(or bef;inniiij^) on I he ooe band of fibres of the pedal and teKuieiilul
•yaMmai and on ih« uiher huud of Bbna of the corpus calhwiiin. or the otbci^
comtniawrftl fibrea spoken of as "asocial ion" fibres t^ GW). an exveedingljr
\nTgf number of fibres uf the peculiar fine iiii.-iliilhuod kind run in Tarious
dircctiona, forming a di-ti«« network iu the jiriiuud substance of the gray
tnaiicr between llie celU. Wo itiuy add tlial tItU svuem uf fine medullated
fibres » of late growth, iind is not l\i]lv developed xa ninu until tiro or three
jreare after birlh. Many nf tho wedullaled fibns, coarse as well as fine, Inko
• horiionial dii«ction porullel to the surface, and in certain regioin are
sfiecially developed into a la^r or into two laj^en so n to fonn a horizontal
•treak or Mreuka.
The vaicular pia mater in\-rets clnwlv, mu «i> have xnid, the whole tiirfnoA
of the cortex, dipping down into ihr *ulci : nnd fmni it, iw i» thocitMof the
spinal cord, [irnirwn carryhig IiIoimIvc-'Wi'Ih nnd hi-jiriug lymph-^MOM pan
inward to supply tlic gray matii^r with bUxul. Rut whilr, as wo sfaall •••
later on, the nupply of oloodvcMcIs to the grny matter is considcrmble, lh«
Iruljr connL-ciivoiixHue elemmta of the »in miiier proccmta are soon momd
inio tteunjg'ia. ImmrdiatHr )>rni.-«th the j>ia miUvr farming ihu Immedtiite
MlHkeo of thp cortex io n thm layer oimsting of neuroglia only.
§ 651. Tile nervp-«II» "f ihi- above several kinds arc nmiiigiil more or
IcH distinctly iu Inycn piirRlkl lo the surface, so that the whnk- iliickncwof
tlM eoiUx may b; meom of thoni be, more or Ivs succ«Mfullr. dividt-d into
• MliSB of lones, one above the other ; and we may, ns we have uiid, recog-
niu on the one hand a general arrangement common to the whole surface,
and on the other hand uiodificatioos existing in the several regions. The
geoeral arnuigenMUt mair be said to be one of five layers or xones, untally
CDUnted from ilie surface inward.
The fiAb layer, lying next to iIm "eniral white matter, fairly uoiforni in
dwracters and thickness (ahuut I mm.) over the greater pun nf the brain,
is characteHied by the presence of sotucwhnt sparselr scattered " futiform "
eelli), though other branclted cells arc pn^tenL It is broken uu into vertli-al
c<>luinns by the bundles of vcrlinil fibrui, luid tU dciuaroatioo from the whltu
matter helon i* somewhat iudistiucl owing to the fact that in live hrnin tlte
while mntli-r, uS[H:('iaUy tlinl lying hnnc-ath the cortex, cuatuim celU and
small gr<iu|« of cells Iring betwocn the bundl>« of librva to a much greater
extent ih4H does ibe wnite matter of the spinal cord.
The fourth layer, lying above the proccding, varies much more both iu
thickness (O.i-'t mm. to O.l.'i mm.) nti^l in its charncterSL The constituent
cells ore on the on« hand large pvrnmidul ctrlU, and on iIh; othrr ham]
"granule" or "nncle-ar" cells. In wnie rvgi^U" it may Ito subiiividvil into
two layen. Ibe small " nuclear" colls being oo abniidani as to fumi in tlic
Uf^r iwrt of the layer aiepiirate layer called the "granule" or "nuclear"
layer. This fourth layer, like Ibe prcooliiig fitlh layer beneath it, t* split
Hpintii vertical column* by tbo bundles of vertical fibres, but to a lestdsgrwo.
It is marked in its lower pnri by n horiioutal streak due to numerous, mostly
fine, niedullatcd fibres running huriionlally. hi the oufiex of iha bland of
Kiel this horiioutal layer la developed into a conspicuous shaat of nwdullated
fibres, eeparatiiii* the fourth and fifth layeifi by a distinct interval of obvious
while matter, i'his tifth laver of ftisiforni celli. thus detached from the rest
of the cortex, is whiU i« called the el'iiutrum (Klgs. 193, im, e/.).
In the third layer, the ooujttitueut oelb are the characteristic pyramidal
806
cells. These nre for tlir mimt pnrl litr^c, tlioiigh (liminnhintr in mu fW>in
below upwarti. ami ihc layer hns l»wn cnlUtl (Ik- " layi-r i>f UtJt; pynniiiUI
oellK," tl)iHi);h ill certain regions the Inr^gcnt [lyramidiil oelll. Aiiil noulilv llw
{(iant-cclls are luiinil in (tie pnnM-ding, luiirlti, Uvcr. Tlie celU «nr, oii ilw
wbole, scatlcreil Bomewhnt sptirffiljr, though frvijiietilty gnthervd inlii mail
ETuups, and among them oeair small "nuclear" and other iiulU. Tbr
buodltfl of vertical fibres tprcad out rapidly in litis layer, so tliftt lin-
ealumnar arrangement becomes lost, and many of the Rbm undnublully
become axis-cyliniJcr proceBsoa of lh« pyramidal cells. Though tlra Ujw
Tories in thickness (1 mm. to 0.4 ram.), and in some of ils features in diffeniA
tenatti, llie characlerialic pvramtdal cells are present over the whole surfoA
oftlK' bemisptiere. In Llie lower part uf the layer a second horiuintal dreftk
ol'cliisely iDterwoveQ bori^onlal flhres fretgueiitly makeii iu appearance.
The second layer, geneniUy a thin one. though varvius from O.i'i mm. |«
0.T& mm. ill tliickneM, is alai> funned hy pyramidal cells, but ii^distingul'lieJ
from ihe layer b«li)w l>y the nb>euce of lurgt nuil medium-sued cells and Uf
lh« proieiiee of ouiiit-Miiti amall cvUf dosirly packed lu^tlier; it has ben
cnll«<i "the lavcr i>f *mnll pyramiflal celU" As we have said, tlteee unaUer
Iiyramidal c.a\\i dilfer niitii-what fr<ini llie larger cells ; and the oella in tin
ayer art souieiimai dwcrilii-il on " ituj^ular."
The lin^t ami miwt «ii[)t-rti(-iul layer is cbaractoritMl by tlie predominance
nf tliv molecular ground MihiUincc, l.Iie i.-clU being j'cw. lur between, naall,
ami irrt'icular. The ground wilxitantH! itself Mccmn to W more largely tte«-
roglini in nHlnrc than in ihc other layer*, and, us i*C Aaid above, its extrviu
•urface ap|MMtr« to Ix- fiimi»he»l hy iiuiinigtia ali>iic. The layer m gmeraliy
spoken of as the " peripheral " or " sujHTliml layer," or sontclimiaia* ik«
" molecular " layer. The tapering reriical proccsMs of the pyramidal eel!»
iBfty be traced into this layer, which indeed varies in thicknesa according lu
the abundance of pyrntuidal cells in ihe subjacent layers; numeroos euBtr-
vhat line medullatcd fibres also traverse it in a horiiontal direction.
^ 652. The general arrangement just described varicH, a» we luiee aaMl. in
diirereiil regions ul' the cerebral surface. We must content ouraelree hut
with pudnting out the eharaoteriHtieg of two or three important regions.
The regiuu which we hnve (^ O^l.'Ij cidled the " motor area " or " rc^iaB,"
I* t-hnrncteriud on the one hiiml by the grenl thickocae (1 mm.) of the thin)
Inver, that of lar^ pyramidal cells, as well us by tlte number and siee of the
cAU containeil in it, nnil on the other hand, and iwpei^iallv, by the promi-
nence in the lonrlh lavcr of remarkable cliinter^ or ver)- large pyramidal
celN, of the kind which are n^fnrred to nlmre <.S G'^Oi as being IKfjuently
called "gangliunic :" it i» in tlii« rvgion that "giant celU" are lixittd in tlit
fourth layer, namely, in the upjicr part of the precentral and at t)K summit
of the (toetcontml convolution, an<t in the immreutral lobuU, aix(uini>g tlidr
gnmlCEt tsite at the top of the preconlml cunvolnliuiL.
The occipital region is chonivteriwd hy the prominence of the " Bmnula"
or "nuolear" cells. These not only form a disiincl division of tbn fourtb
layer, but are also conspicuous in other layers, their amngeroent being surh
that some authors have been led to divide the cortex of tbu region iittu seven
or even eiglit layert). In the present state uf our knowledgo ire may be con-
tent with inaijLing that the great mark of this occipital ra^on is tna abun-
dance nf tbese small "nuclear" cells, together with other ^mall "nngiilar"
cells, irheivby the pyramidal cells seem to be mode l«n coti«piciioUK. Il is
worthy 'if notice, however, thai in the third, but more especially in the
fourth layer, a few cells of verv large siie are met nith, which hy their
largo branched cell subtttauce atid eontiptctious axis-cylinder prooeas rceemble
tbo \»rge cells in the motor region ; but it should be noted thai vbile ll»eae
flOMK HISTOLOGICAL rEATURES OF THE BBAIN. S07
iatfit orlU occtir (at Irast in idbii and in tliP i»on)»!y, though not in goate
of ikf lover aiiimalH, as llie ralibit) in rrry iJHinito cluMrre in the motor
n^on, ilicy occur Niiglv in lh« occipiliil nxton. |p Uua occipital rc^oa
tbr layer of liorifrinuil ^brcs ia the Tourth layvr is verr COiupicuouf, and
owIdc to (h« number of ordinary medullaled ohm prpsent. foriua a white
alf«aK vwhie even to tbo nalc«d eye.
Ill tbi- fr-inlal rC|;ioii. in front ul' lhi> motor region, the arrftni^eiuent is
more in ac4'ordiuioe with whiil we have described &« the ;;enerui jilun, The
two prminidal layen are we'l marked, as ia alao the fuurth layer ; but the
layer of larf^ pyramidal cells ia much thinner titan in the motor re^'inu, u
is aJao, though to a lew extent, the fuurth layer, while the tif\h layer, that of
fusUbnn oell«, ia thicker than elsewhere, i^mall " nuclear " celU arc perhu|ia
more abuu(lau( in tliia region throughout all layers than in the motor region,
but are for leo oouspicuous than in the occipital region.
We may here remark that the transition iu structure from one region to
anniher u very gradual, doI tihurp and dtdlinel, and i^ perhaja e^pucialty
Knidual in pwuing from the motor i«Kion backward to the occipital refciou.
It ia not uowible to recogoiie bistologioally the limit, for iustauc«, of the
tmitor repon na determined ex pari men tally.
Id sneuial iwion* of the bniin, for inntaui'e in the olfactory bulb, of which
m shall nicak later on, very great nioilirnutii.iu.'' of tht; ^ncral jilan may bo
obMTved n tbo cortex. We eaiinol onl«r unon thcM«. but may juKl refi?r lo
Um ooraa aminoDis or hippocampus. At tnc ventral end of the temp'>rul
loba ttw cyrus bippocamiii, the structure of whose vurl«x follows tbo gvnoral
plan, b Utrust inward so as to project into the eavity of tho dcsoeoding horn
of llw lataral vetilricle, forminir iIm ridge-like proininence known hi- the
ahoi* oamc Tbe substance of lh« oorau ammoDis is therefore corticnl sub-
ttmuot eoverwl od the side of tbo ventricle by a thin prol»nj^tion of the
eentml whito matter, which is in turn covered by the ependyma lining the
ventricle, A vertical Kcliou of this substance ehows that while tlie fifth
and fourth layers are reduced lo xmall dimeuaioiis, the third layer, that of
large pyramidal oella, is well developed, though narrow. The cells are large
and rumarkalily lon^, and tlie tapering procewn are arranged so reijularly
aa to give rt>«. t-«pecially in siaiueil preparations, to a marked radiate appear-
aiHieL At tho level of thv aeeund layer there uccun a large deTelopment of
capillary bloudvesMhi and a seareenuM of cells, giving rise to a "lacuDar"
■ppearaDoe; and tho Tirwt or mulocular layer is uf some couiiderahle (hiok-
una. From tlw promiuence of tlie pyramidal cell& in tbL- region, tlie third
layrr in the gVDcrml plan of the cortex has sometimes been iipukeu of u the
" iiirniiitiim of tbe coniu aininoni*."
(663. In the present state uf knowledge it is im|Hituihli^ to oomt" to any
Httsfiutory conclusion conceniing the meaning of thi- variety aod arrange*
moiit of the celUnnd olhrr comtitiient* of iho curtpx. The celln nith Ihrir
braacbee, the ncr\-o tibres and tho nerve tibrils form a network of gray
matter which ne may oompar* with thir gray matter of the s]>inal cord
(6 MO), but which is obviously, as we might exiHict, far more complex tlinn
liiat il. We mav conclude, and experimi-mtal olm-r^-niion confirms the ooa-
cJuaioo, tbal the large pyramidal cells with rpcognirahlc axi:M:ylindcr pro-
oaana serve as trophic centres for the Kbrcs which appear to start from them.
And vre may, though with less confidence, explain the large siu of thoeo
cells in the motor region, by the fact that thcv ^irc nno to fibres of tbe
px-rainiilal tract stretching a long way from their ori^n in the call, and
ibcrcl'ore demanding great nutritive activity on the part of the cell. W«
may perhaps also ooncla<le that these fibres are olferent, motor fibres, dea-
liued to cmrry impulaea from the curies u> the peripheral, or at leuit distant
80& TFIE BRAIX.
pnrts. AimI ir« may fitnher. with bonover ilUtinotty !>'« confitlviict.MHM
tlinl l)i« n/x of tl>e ci'll iit ctirn-lateil t" (lie tm-r^T nliirb haa to bo toiBaHU
in llif ilii>cliiirgu nl' t-ilVmil. iimtor im[tui»c:). ll' wi- nc(i'p( i)h«« cooaarinni
wc muxt nlw) Iwjir in mind llimt mich ii:ll:i, irilli nxin-c^'liuiliT iirocaHBooi-
linnecl on nii filirii>, tirv iint liinilc^d tii, tliougli most ■)Hin<liint in tbtt moUt
ngion, hut nrc fuunH in nil regions of Umi cnrivs ; uid wv muat litnc* cm*
elude Uiit iinpulMS, which vre mtiat vnll pRi^rrDt, proceed from oil pan* iif
tho cArtex.
Il M ohviouf, howcrrr, thnt the conncctioD of the conical networit of gn^
matter with lh« fihrt-s of iho whitv mntlor is <!Abclcd in mrt only, muI iMt
n fRiall (inrt, hr thr mpthrK] nf axis-i-ylindcr |irace««M definitelr pnilniiiKd
from tlip cell »iiWimcc of cclU. A pnrt, nnd probably a greater pari uf Un
fibree sweeping up from thp subjacent whil« tiinUer, vhcihcr they be fibiv
of the pednl iind Icgmciitnl »y»toDiE or c-iilloeal or "aMocialion " fibres, col
io the gray matter in eome other war than bv bodily bein^ continue ip lli
cell BiilMtance of cells; they plunge into ai)d brenli op witliia the network,
of which fibriU no less than ccIIb lonn a conspicuous jtart : and we may hai
rejieat the remark which we made in speaking of the cerebellum coocerauj
llie uctual continuity of the element* of the network. Moreover. bcMte
the vertical fihrei obviously coming from the subjacent while matter, we
have io this gray mailer to deal with the lihfes of homotiisl nitd otber
dirc<-lt'>u», which may come from while nuiUer not far <tff, hut which nmr
coniv from some Dei|;hl)oring ^ray matter; our praaent kDoirleclge will M
enable ua to settle this |>oinL
In the spinal cord we were able to divide all the fibres tnlo affi>tcut at»d
cfferrnt reiipeotively ; though even here we met wttJi some difficulty. ))nl-
ing with the cerebral cortex, which, a> we have already aeon, is rrnniidjr
espeaally conccrneil tii voluntary moTcineiil« and in ibe dcTclopnieDt of (ull
eensaticnE. we mar he teniptctl to con*idrr the fibres connected with lb* ptj
matter as similarly ilivisiblc int^) motor and urn^ory ; and no may go on IS
suppose thnt the tihrn joining (he cnrlcx a* axis-cylinder processes of ntag-
niiahle cells are moiur lilircs. ntid ihnl all the other fibres joining (be ffij
matter in some other wiiy are sensory tihn?s. But io doing so we are goio^
beyiiml our lether ; in all prohahiliiy thp nervous proecsSM fpHtij! on in tbs
cortex are far too complex to permit such a simple claasifioalion of the fatw-
liuna of fibres as that mto motor and sensory; and any atteoifH lo arnuin
either fibres or regions of the cortex as simply motor or sensory h iirohahly
misleading. But we shall have to return to these nuitteis when we deal viu
the functions of the cortex.
Ok VoLCJiTARY MorRURHTs.
r i»ov» I
. of tW I
$664. When we examine ourselvea wc rccngnixc certain of our
roenta aa "voluntary;" we say that wc carry thcro out by an cflbrt
" wilt." And when wc nitn<«s the rocivcmenlR of other people or of animab
we regard as also voluntnry xuch of those moxpmeniK as by their characters
and by the circumstaiii'cs of their occurrence ecom to be carried out in the
same way as our own voluntary movemoDts. Kven in tho caw of some of
our own movement]^ we are not always clear whether they are m]ly Tolun-
tary or no; end in the case of other people and of animals it is atill inore
diliicult to decide the question. It would be out of place to attempt to dis>
ciiiv here how vulunlary moreoienta really differ from involuntary niov»
mcntF, or, in other wonls. what is the nature of the will ; we muat be muimt
to lake a somewhat rough use of the words " voluntary," '* voHiioiuil," and
609
" will " M « Intuit for phyii"logiritl HiscuBRion. Wc may. however, remark
lliat iia fur v llir muftcuUr side of the act. iT »i> may »*>' *\ii;h an vx|)red«ioa,
ia concerned, a roluntary movement dues not dilfi-r in kind front an involun*
laiy nioventont. It i* perfectly tnie that a fkilled mnn may by pmetice
Icam In exonito miiKiiliir manceuvrce which he irnuld not have Ininit ta
txtvvie hud not nn intelli)^Dt volition bcco operative within him; hut our
awn eX|)«rienoec teaches ua that many more or lea intricate movenipnl*
which have nndoubtetlly been leaml by help of the will may be carrioil out
under ctrcuntstancee of such a IcimI ihnl we feel comiielled to refE'i'*! thom
U, at ihe time, involuntary ; ami it may at leuat he debated whether rvcry
iDoveiDeiit which we can carry "Ul by an efliiri of the will may not nppcar
«nd«r appropriate circumatancai ii* part of an iovoluDtary act. In the «ue
of the lower aniroalB. in the fm^ deprived of its cerebral hemispherM, for
itMaDee. we have Hceti that viiluntary diffrr from involuntary mivomonte.
not bj their eMuiitial iiuliirr, hitt by iho rvlation which their occurrence
bouv to clreumtilnnceH. We have lher*'fiiri' to m-ek for the distinction
between voluntjiry and involuntary, iii>t in Ihr cim Ordination of the miucular
Ktid iiervouti com[iiinenU of a movcmciil, but ia the nature of the procMS
which *tart« the whole act.
The ht*toriai, related in a preootlins section, of variouH animals di-prived
of their cerebral hemiiphcnr, while they have further *hciwii tli<! dilfii'ully of
ilniwini,' a «hnrn lint' lietwc'ii tlio pmenoc and aliwiicc of volition, aucb aa
when *c apjmi) m <nir own conticiousncM we room able to draw, have tauf^ht
a* that ill a broad een»o the iirmrncc of volition is, in the hinhnr vertebrata,
^priulent nn tlie pomewion of the cerebral hemiephen?« ; and wo have now
tn inquire what we know oonceming the way in which the cerebral cortex,
(or tbi>, a« we have wen, is the important part of the cerebral hemiitphero,
by the help of other pana of the ner\-ou3 syxtem carries out a vuluntary
movemcnt.
$ 6U. Wiib this view we inav at once turn to the reanlla of ex[>erimental
Interference with the cortex. When the aurAice of the brain ia laid bare by
removal of tl»e skull and dura mater, mechanical atlmuhition of the cortex
pnxlucca little or ii» efted, lhu> afTortling a contrant with the nwulia of
nievhanii-ally alimulatin^ other portion* of the brain, or other ner^'ous
mmclurt-a. And for a long time the cortex war upokvn of •« InaeoMble to
Htitnulation. When, howt-ver, the eleclric current t* einjtloycd, either the
make and break of i1h* constant currmt or the more miinat;mhle inlcrrupled
current, vtry markiil reiulii follow, II is found that certain movementa
follow upon eWtric otimiilntion of certain regions or areas. The rcsulta,
inomiviT, dilTer in dilTi-rcnl animals. It will be convenient to begin with the
dov. on irhich animal tht^ obtcrvaliow of this kind were tint oonductett.
When the surface of the dog's brain ia viewed from the dorml surface •
short hm deep soleu* is smn toward the front, running outward almost al
riebt anzica from the great loDjptiidinal lueure; this is called ihf irueial
aiwM (Fig. 2<''2), the syras or convolution in front and behind it. and
■weepinx around its end being called the aiffmoid gyrtu. It will hardly bo
proJilable to discuss here cither the homology of tbia sulcus or the names of
iIh' otl>er sulci and convoltilioits of the dog's brain. We mention thi<» sulcus
tiecause it is founil that (^liinulaiion of the cortex in a region which luuy lie
broadly d«scribeid as that of the neighborhood of this crucial sulcus givca
riae to movements of varioui parts of the body, whereas no sucli noventeots
result from stimulation of the extreme frontal resioii in front of the area
around the crucial sulcus, or from siimuUtiou of the occipital rt^ion behind
this area, i'ertain exceptions may be ma«le to this brood ttoteraent, but
TOE BRAIK.
tlwse it will bo b«st to <li«cu« la reAtnnoo to tli« more highljr <lfTtlii|iNl
monktr.
The region of the corl«x in the iwighborbwtd of tb« crucial wIm* dwjr
theu bo t«rmrd nil " exdtablv " or " motor " ragion, iiiadmach u ilimiililwa
of thi<i roRion leads to luovcmente cnrriiMl out hjr akelotal muwin, wbik
etim Illation of other re^i'ins <loe* not. Further, Klimulatioti of Mrttcnlar
dislriciti or arvm of tlie re;ii<in leads to particulur movvmeois earned out bf
partkular musolte. Fur iuitiaDoe, stiniiiUti«n of the more nwduD parti «
tbe errtifl beliind ihe k-rucial aulcus ( l^xji. 'iO'J, C) l«mla to movenetitB of tha
1iin<I-ttmb. nhereaa atimtilalton of tbe lateral part or oul«r «nd of tba tarn*
gjixn leads to inoT«neiit« of th« fore-liiab, aod im najr h«t« dtatia|[ilU
no. m
^
Tiia Arm op tbi CuniiAL Oa»voumM« or n> Vwt. teooann* to liirao tin tmnn.
(1) B. Tbcuoitbi iht muMilaiof ilMMck. (31 A,Tlia ana ftir Um extenili« »oi aMBahaW
Ih* itaf*.limlv fllK nauwkvtktOmoaaailraMUaBaf UwftnlUBii. 'liCTlHuaM IM
blMd-Unb. Konnlaa lnuin«fwl)r Uwwd *m» tfunUam I *nd 3 Itma bkihI i I* ibm ■Mav*"
•aim*. W D, Thir bKtBl UM.
between nn areA atimultilion of wbich (Fig. 202. E) ImiU to flexi'>r,
fore limb, and an area (Fi^. 202. A) etimulntion of which leads tor^'
of tlte saiae liinl>. lit a atoiilnr way stioiulalioD of ollMir ureas wi:
"motor" region leadg i» muven»eaia of this kind or of thiu kind of ;i<
of tlw ejet, at the muulh, of oilier p«rta of the face, of the tont;n(<. *«'' "'
on. Obviougly iit tlic <ii>g this ref[i<in of tlie oorteix hua conn«cti<)tu •iU
tbe skeletal muscles wliieh ild ni>t ot>uin betw«eea other re);ion« of thocnrtan
and thi«e miMcloe; and rurih«r, the reunion tn questioo ta lopoi^itr^''^'''
dillereiiiiutod. ao that certain area* or dislriets of tliu re^ou are -
connected with certain »kel<Ttal niuiwlee or >,'r<>M)a of niu»clw. U'^ >»•'
■peak of ft '* lucHlt«uiii}ii of function " in Ihid rt-uion as oonipftred with nibfi
regions of the cortex, and in tiie aevenl areas within the rogioa as eomptn^
with each other.
Tlte muHoloa which arc ihuN thrown into oonlractiou are the mutd*''
the np|HMitv side of tbo body. When " tlie lure-liinb areb," a» we niay <all
it, of the riuhl hemiepbcre ia elimulntnl, it in (he left f»n>-limb wbicb i*
Riovet) ; iinif so with the otlier arena ; it i> only in exceptional cum*, tt io
certain niovementa of tlie eyti. that tbe eflttd is bilateral : m Diov«iii«it oxi*
fiuni to the aam« aide h Uist •Umulateil la never wituewMl.
tir TOLUNTART XOVBHENTt
811
I
The KitilU nre most ctear wheii llic current «!iii|)] ()>■«< I u a Mimulus u nit
*tmi|;er iliau ujusleufficieflt tujinxlucetbeapimiprmitf m'iveni«Dt (roufcbl^
apeskini;. n ctJirenl jtiat perceptible to the limuve uf the-'i|)ersti>r U in ordl-
lutr* ciuen a u*eAil i>De\ ami when (he cnrlex in in ipiwi i)ii(ritiv« condition.
In luy ox)ierin)i^iit ihc r««iilt« ohuTriMl )>y tliceurlicr rtimulntiom. W)on after
Um cortex bu IxH-n expowcl, aro lh« brat ; nlU-r n>)>i-«l<i) attuiulatiiMM the
Mirfiior n apt to become bypencniic. iind it iw thra rrv<|ucntlv obaerverl tlwt
tbw ntOTCiuentM nwultiiix from the Rliiiiiilalktii uf h [uiriicular area are not
OOnSlKtl to the appmnriulo miivclc*, but «prvad to the (^Trnpniidiiig niUNclea
of the opporito aide, (nen to musclfl* oouiio'letl irilh other i»fLical amu, aiMl
at iMt to Ifat iDUKclffi of the body generally ; itt the Mini« time the move-
BMDta loH tbcir diMinclire piirpiwfut chiintctT mid (be niiinial i» ihmirD
into convulsions ol' an e|>ilepiif>irm kind. It not urifrrqiinuly Imppeit* that
an pj[|*finwnt hn* to br Mopped in «>iwr»]iieiicc of tbo onset of the*c epilep-
tifoftii c»nviili>i(inH. The n»poii»o of movement to sttmulation niiir bo
obecrvnl white the animat is under the moderate inHiicnoo of an nnresthvlie,
ImjI a lo<> profound niiir^thesia leMona or annuh tbo eflccts.
In order to rarry out a cinder annlvi-ia of the phonomcRn it is dcsimble- to
«atch or record the coniraelion of n pnrticulnr group of niufelee, or porhniM
better still a ]>anicular muecle, p. p.. (be nreti for extension of the hind limb
■nay be studied by help of the eKC«nsor dif^itorum communis of ibe limb.
Wh«n this is done the fullowini^ important facts may bo obvrved : The anm
of cortex having been found which jfivea the beet niovemenls. and the
Kimulus haag no stronf^r than is neoeaHTy, isolation of the area from )t«
lateral Burrouodintp by a circular indsion carried to gome little depth n-ill
But prevent (he development of conlractiitns in the muscle; but tlieM do
ceaae, even without the circular incision, if by a horizontal section th« eray
cortex is nparated from tbo fubjncent white nuitier. Alter removal of the
corlMC. Mimulatiiin of ilie irbitn matter underlying tbo area producea the
appropriale conlraction ; not only however i« a stmngitr ittimnliiit Deocautry,
but aUii ihr latent ptrii<d, that is ih<! time intcrvtmin^ Iwin-i-cn lh<- beicrnning
of the n|>)ilicatioa of the Ntimulntini! current and the: lH-(;inning of the mut*
rular c»ulractii>n lit appni;inbly nhorteued. The approprialiT cHiulnictions
Dot only apjiuar when the. whit<' matter immcdiiitcly ln-Kiw the I'ortox ia
slituulalrd, but hy niukingndccewiveh'irisonlul aoelionii and stimutnlingi-Hch
in (urn, the *-lli-i-t m«v, so to #i>r-jik, be traced through the eititral white mailer
of ihu hemisphere "(own to the internal capoule. \V<t nmy cnnelud" from
thcM- rc>ult«. that when thv eiirrcni is ap|>linl to ihv surface of the cortex,
oenuin parts of o^rtain struetum in the gray matter are stimulatod, tli«
pruMas naving a nuirkea) la(cn( period, and that as the outcome of tho
ebuigea induerti in the gray matter, iniptilsva paa along the fibrea Icadinic
iloirn from (he gray matter to (he internal capsule and m by tho pwlal
kyslem uf fibn* to ue apinal oord and motor s)>inal roots. The anatomical
consiile rat ions advanced in a previous section hiad us to «ippo<e that tlic
fibres in cjiiestion belong to the great pyramidal tract, on which we have bo
much insMed ; and as we sliall see, all our knowlcdi^ conlimn this view.
It must Dot, however, be Fup|>oeed that the several areaa stimulation of
vhieb produoes each its di<linetive movenieot.are in the doK sharply defined
lVo<n each utber ; when the term area for exteonon of the hind-limb is lued,
it mu«t not be BupjKxied that the area can he defineil by an otii'int!. within
which Htimulation prodiioM niilhtiig but extension of the hind-limb, and
ouUade which iximulntion never pruduee* extension of (he hind-limb. All
tlial is mesnt i* that rxteonion ••]' thv hind-limb is the salient and ftlrikinf;
Nwlt ofMlmulatinit the art-a. Wbr'n we study the various raovetnenia, and
wptdalljr perfia]a wlien we ntudy, bg l>«lp of a raiihic record, the oon-
812
TUE BRAiy.
tmctSoM nf vnriotit injiviiliinl mu«oI«s Ksnltlng from th« ftimnbtioe of
Turioua paru of the motor region, w« (tixl mA nnlr thnt Hk areai (bf fw^
ticuliir muvcinctiU or pnrticiiliir miKclm nra very <liflu««, but tkAl tbetevpitl
arau largely iivcrlnp «ich olhi-r. Il' for iiiftaiin^ «t> were l» map out oa lb
tuns dia|;riini the several nms belonging (o four nr fiv« muntes of diKfM
nnrls of the botly, biicIi m the cxtpnwn of Ihr Aigitf nf Uie fora wii] of tb
nind-litab, lii« flexors of the ^ame. nod ihe orbiculitr muaclc of tb» fftfiJ,
that u toaay, tbeBevernl Hrena irilhin which in turn Mimtilntioii of tbeeDlln
produced coDtmclioii of the jiarticuliir miifote, the ovcrlappiug would br»
mat that the whole li^ure vould appear bichly o^nliifd. In a iiimiUmT
tno excitable motor rej^ioa as a whole would };nidually inerg« inin, W hriiL«ti
up into, the uiiexcitable tVontal. occipital, and temporal regiona, io frini.
behind, and below. In other words, Lh« localuation in tlie coriax of itx
doe is to n marked de^'ree iniperfeci.
In this respect the do^, corittitondin^ to ils position in the animal biennkf,
is iDterniediate between aiich nuimaU as the rabbit, the bini. and the ln^,iia
the one hatid, and the more highly developed monkey on the other : asd thu
is oDO rcnsou why we have taken ihe do([ tint aitd dwelt w long anon it. In
the rabbit, a umilar localization may be oluierved, but far (a» oefiaiU, la
more diffuse; it Itevomea Mill Ii9« in the bird, and ii hardir racoKoiaUcia
the fro^'. It will not be profitable to dwell ou the detaiu of ui«M Inn
animals ; but the plieiiomcna of iha monkey, lending up oa tbey do lo UiM
of man, call for ai>eeial notice.
§ 656. Whitn in a monkey, in no individual for inttanee belonging to ibt
jfeaoi Maennu, the fiirfan-* of the oerebnim it expton-d with refenMel*
tlie tfftct* of electric «liitiiilntii)n, it is found that when the eiirrvnt ia appM
to the precciilrul or iis<i<iidins frontal nnd the pxrt-centnil or oacondiaf
parietal convolutions which lie respectively in front of and l»hind ibt
iniporliint central lissore or flsanre of Kolando (r^. Fig. '2t)^t, movwiMlitl tl
theforr limb follow. The " motor area for the fore-limb "thiu diaeufmdii
more circiin)scril>ed and dciinile than is (he oorrospondiog area in iha ivf,
lu outline (Fig. ^iHj is roujrhly that of a tnincaled trtanele biaocled by IM
rentral li»ure, with the broad bone at some distance from tne m«»ial liar.ul
the triiDoated ajtex reaching on the lateral surface of the hemisphere t»i
well-niarketl bend in the loner part of the central flsaure. Behind, it mi-l«
as far as the inira-piirieiAl ftMure which somewhat eharply defines itf bind
bonier, aud in frunt it cciMea no leaa definitely at aome little dtmincv IwhioJ
ihepreceniral tia&iire. Further examination ihows that the whole areab
divided into areas correoponditi); lo movementa of imrticular parU on)*
forearm, and that Ih^M- nn' nrmnged iu a defihile relation to each olhrr.
In tbe more dorml jiart nf llic area, nl (he liaiv of the triangle, atiiuulmtm
produceM movement* of Ihohouldcr (Fig. '20J <; if the electrodes be vhifitJ
veDtraliy, movemciil« of the elbow make their uppoarunce; if atiU mm
veulrally, movomenle of the wriM come in, and tiMWC are in turn MioeeeM
\-entrallv by movement* of the digits generally, of the f<>n.'finij:i-r, and Laitlf
of tlic tliumb, A very striking experimont may b« made by applying s
cunent of suitable streiigtli, lirst at the lower ventral border of tli« am,
and then gradually advancing iipwurd toward the mmial Hne; the thumb >>
moved lintt, then the foretiuger, then the r«^t of the digits, then llie wrwt,
next the elbow, and lastly tbe shoulder. Further, in certain parts of tbt
am tbe resulting movement is flexion of the appropriate segnieot of tba
limb, in other pairtti extension, in certain porta abduction, in other paM
adduction, and so on.
Similar eiploratiuo show* that the " area for Ihe hinddimb," ties on tbt
rocdinn side of (lie area for the fore-limb, stretching besides on to the mc^
OSr VOLPFTJ
VOTBMBXTS.
nrflwe al'ififr ilio margiual eonTolution whicli fanm l)ii- dortnl pi>rlioii of the
■nil of lli« gTvM ltiii];ilud)Dal IWiire ; it rvaclioM lut fur liiu-lc m tlic intn-
■ivtal lulcus. utd is aueocetJed ui rnml t>^ tbn " uvn for tho iruiilc " ( Tig.
)), WilliiQ liiis general Hrcn fur the hiixl-linib wc mmy xiniilnrlj distiiu
lU) •[weial urans Tor i)m> lup (Fii»i 204, 205) in th« rroiil porlton, for the
tnc* UMi ank)« behind thia, and Ibr the diglu atill furtlx-r backward, the
•n* for th« greu lv« being, however, Id front of ihe nrai for the other
I digit*.
Fib. SB.
I
i
littftr. frvitla/M
V
J^r
xi'*
>.
»MpTi9r frantal
X
^
t
->'J
X
•et/fittm/.
I
Orrun or aatto or MmKET (JAioinui ro euoii rnutupAL tvM ifiwiiKt*} «ra otu (Oovto-
UTTKUUI. r^wliiMvroiialUiHonuiYUidSnitnuLI NaiorKl «lw.
Tin bnln Ofiml li Ui* aimu u ibnt In Fit. IM. «nd ili« i>u Otim ihoiibl k« cunnilM UgtOm.
Ont M«b mIcw^ lariiowljr i-rliiwtl v«rT Ihlck. tb> mum I* WritMil 111 imM mpUali. om «Mk BTHM
la mNm > tndkaw* Uw «n>LU •l«t>nMiau. lurdlr %o In wll«4 a nlous vblob I* loppcNad lo t«
buMWliBim vltb Uw wpfrior tanHl nlnw oT man : aiwl Oi r. ^ iiwlarljr ladlaaia tuM whow
liiwnlrnlw an ntn nirlain rorioimiriionri'uaalVi'SOT.sm.
In fVont of the areas for the limbs and trunk, on the inediaD dorsal sur-
fncv, dippiiig down into the mesial surface along the marginal eonvolution
(Pig. 2(u) and reaching laterally on ihc lateral d'>r>ai1 aiirface to the dorml
MlremltTof the precenlral 8iilcus(KiK. 204), i« lhe"nrvn for the head,*' that
ia la aa^,' for ibe raovemenia uf Itead Drought about bj conlrvctions of llie
nuaclea nf the Deck.
Vcntfal to ihii again, in front of the prec«Dtml iiik-us, is (h« "area fur
Ibe eyes," that i* to aur, fur cciuractiona uf the ocular mueclee ; and behiuii
814 TilK BRAiy.
ll>c nreoentnl Nulcuf, venlml to the arm area lies m snuUl *n» (or
of Uio eycliilii, l)ri>iight nbuut by cualractiiHM of ihe orbioularit ntoirh.
Ventnl to lliis aguin u ihtt " nr«a fur the fuoe," in irbicb we may diatinguU
no nmi fi>r the mflulb, tliiil in an un-a atiaiulation of tihirh |>roduL-at cWpf
in lh«i biici'al tirificv, vjicning, filuittitij;. <lruvrii);( to unc tide, (He., uiil mt un
for n>uvenM-nl» 'if the. limguv. Thnw two mms rvticb dunnnRnl to tlis faauit
of !?ylviiin atxl iHii'kwnru tn the line of the iHlra-|)«ric<al >u)vua. In fnoii/
thcni, occiipyinf; «ll thu vi-ntnil pnri of tlw proctntrml convoluliua iiU
rvuL-liiiig furnHrd lu for lu tb« precuiUnil Mulcua, wli«ra it nioi<ta iba un to
•ITJ
*«..■ - ■ "^
« ION
lip "."'t -*iji.u!i: , oiiiTs
VISION
t^T
Lett Uuamau or rat Ciiuai*Lum or Mokxkt yttaraaut. Viav^ raox n> Lm >m «•
rtOM AtovE. (SiiiuiNnroH mHit llnOMii and HkkvoIi.) VbIuivI ttm.
Thv flvum ihosi iha i<aiIUonior ihepurtinnioT luc cortex nxMnncd wKb mnTfawntrf wni
101 ri«. kiid Willi lliowrw.vof ittlit, uDcli. niii1 IimRIik> TIm torUrml KNa ronneettd wM Ha ■»•
■mrDUuf tlw lifU iliiuIfO mrtlcall; lurruB. Ihai w)Ih Ihc mintmenta t4 llic WB tiod*mil>II;.ial
Ida) «iUi Uw uiUTOUunU of ib* trunk jii a ilautlnit <llm!lloii: Uio ana (OUiicinl alDithram
■MliNoT (III liiail iiwckj. tk<x, 4Iid eja u dollpl Tlicc«i|i>« oT Uw OhlaTllBUia b ladbuM )f
tfucle llnix,
the e^, lies an area (tinmlation nf which pmducen ntovcumts ot tb*
jiburyDx ur laryus, at w«ll u tbo motiih or face, itud which maj be diTJM
iuio areas for inantii-aiiou, for ■vrallo<ring, and for tbc prodocUoa uf Lh*
voice.
We luiglit s|>ealt of tbeK Eeveral arciu iu aiiotli«r nny by referring to tk
OK TOI.USTART MOTEMEST8.
eio
ncrr« cooconird incnrniiifioul tlwMvcrol miivcmentit, though itnloingwi we
mtut ntii«nilMr tli«l lh«re t* Dot nn exoct corrtopoudcncc between tlio tvlntive
pantiii) nf m mufclc nlonft; rli« nxia nl* the body or aloDg the axu of » limb
Mid Um raUtivo potiliuQ along tlio oervbro i[>iiinl nxi* of lh« norvc or nerves
govaraina lb« mutdai We roar, however, wtopting this method, nott? ibst
the MuraT aod lumbar oervtt are rcprvMnted \>y (he moat racaial portUm of
tta.X6.
)Ua*l Unet ortNX I.nr lUi' or tin Ktitv i^r Ui'icn, l>Mrt4V«i> hy Sutmv v nrR
HnlJlV SUfTTlL PLAKX and RDtOTXL OT THK CUtMll.ltU. IVUUUIHUTOH ■Acf HOMLtV kod
7«M hMclMd and MlppM tiMU «# UK tnrtlee (bow ih* rvglcdu or Uw mtcs coDiwMid wUh »aT«L
mmatolih»fitl.hin.Up.tafI,trwnJt,tadtitrtiweaYtlj. TiMHvaalpaMtlDaiaril»«nuo(aci(Mi
IHliXtfIrt ■!«> ttMim and iwrf ■mi •tiOi niaiimu tmalitm t« Indtratgl tr Iha ■ffnupif au wortl.
tt^HM^o^—ttontMimBJiliwiBrt thetorpnifHamai, <f.. ff .tf..«na thioiitlithaaiitwior
caaBAanr*. «. qHtiif iha lal) pillar o( Uw fiitnli. K: baUiid ii ii»i UinM iw anwtUir fart of
ihBf«wla]PlvopaUMB«iMdiui. il4. nor* ivfUa ad quanniDTimliViiluin); itot; tto IvR hal/
of tka foaalu ftfiaMMMooiCy,tteapileeeaiiUau»cut acioc^ in. UwimtcJ ihvtlilnlnaRte)
turn: rM. Om bontal tola; Oe, Uw onlrlial fula: 0>. dia cuscw: JV^ Ih* mnncw: 0>,
0^ 0/a. tba iTTv* fc«>l(ati» : iho tinlciioivd DMirv •««« U> (um lb* a|i|cr boundaf} ut ■»■■?
fynia Id IU npis-CBlknl t^n 1' <1h «all«ci-iiiamlnal ; ivy. ibc |«ileli>wri(atal Oman.
the whole motor area and br the hind diviiion of this mntal portion ; that
the lumbar and thoradc nerves are repn^culed by the fVonl divt»IoD of the
mma neilal portion; that the upper thorncic with the lower cervical nerires
beloDg to a region iytnf; lalemi lu, nod the upper cervical nervM to one lyinff
in fmnt of llie prectnliu^ area; mid. lastly. iliHl thu rt^muiuing lateral and*
vrtilml portions of tlie whule uiolor region afijierluin (u tliv cninial nerves.
Rut tliv Ui]»ij;riiphic-ul ditferenlistion does not come out im> clearly by this
inrlhoil na by thut of iiikiti); fur our guide distiuettve niovemeuu of the
teveral imrt» of tbe biniy.
It will be ubicrved that all tlieae areai laVen together, RprMented by the
portion of Figi. 204, 205 aliaded in one wav ur auoiher, occupy chiefly tite
parietal irdoo of the evn^hral surface, thoufih they alao reach into tbe frunlul
region, fiumulntinn of tbv fnmuil region in fnnit of this motor area or of
the occipital region behind, nhethrr on the lateral or on the meaial surfaee,
or of the temporal region, wheilier abo on the lateral or on the mniol sur*
fkcK, or of the gynis fornicotii* (Fig. 206) coADecling the frontal and occipi-
tal npons on the mnlnl furfiice, and running ventral to the nuirginul ^rut,
doM not give riec to movcntcnte ; or, to be more exact, doea not give ruw to
816
TBIC BRAIN.
tDovemenU coropantble to (o those just dMcribed M resullitiK ''^'" MiiiHli^
tion of vaiioUB purte of llie motor reginn. MovemeDls do take j>liuc «faA
ceilaiu parts of the occipital or of the (empoml reinon are BtimuUioJ. bat
ibeee an nut only feeble oud expcriuienlaltv uiicerlaiD, but appear to btti
a diSereni nature from thoae r«ftitl[iii)f fruiu stiiutilntjoii of the uiolorragtM,
it will be cuDveiiwDt to apeak of the iialun> and meaning of thn kiJul af
Riovenieut when wc vonie to diecuaa the dovelopuent of aeDsatJon*.
§ 6ji7. It i» obvious fruu the foreguiog thai the inecbaniBro for tfaederelt^
ment of these moveiueiils uf cerebral origiu are far more bijfbljrdifleroitiittil
in the monkey tltou in the dojf. ilul even in die monkey (ifaraeiu ud
allied forniaj ihe ditlbreoliatiou ia atlll very iucouiplcie. If we ezulore, fct
inatance, the area fur the nrisl, we l!nd that )t« hmiia ate illddiixd. In
some parts of Ihe area we obtain utovenicula of the wrift only, but in othtt
Eiirts of the area sliimilation iirodut-eB not ouly moreinL-nts of tUo iirul,
ul al»o of the aboulUer or of^ibe digits, or of the nvck; mk) M> witk tlw
oiber areas.
If, hiinever, not a Maeaciu or other ordinary monkey, but th« more higUf
derelojiMl vurangoiilanc betaken aa theaiihjcctof exiierimenif.thedilwo-
tiatioii ill found to bedislindly udviiiic^d; the fcvcral arena are m<irc olinrplj
dvfmtil, and what ia iiupurlaiil to iioli-, the rcflfx.'clive an-an (riid to be Mf»
raitd Iriini ouch by putllon* of cortex, ntiniulation of which givtv rtw to aa
moveniMit at all.
The o[ipiirtmiiiiLit of »timnlnt!ng the cortex of man hinurlf have been tn
and far lietwei-n, and bnve for tlic mofl piirt been conducted under unfu\-iir-
able circumntBUCi-*; but an far as the rcvulH m> obtiiineil go, lh«y sliow ihil
the lopoKtaphiciil dirinbution of Bn;n« for the acveral movements is carrM
out on the mnic plan as in Ihe monkey (wc are purposely cunlinios ouiwItm
now to the results of arlilicial etimulation) ; and, moreover, Justily the cq>-
clnaoD, which li priori reasons would lead ua to adupi, thai in man the dif-
ferenUaiiou is advanced still further than in the monkey.
Thus, when we survey a series uf brains in sucoeasiun, tVooi the nior« lovlf
frog, through (he biid, the rabbit, the dug, ami other lower mammals up la
the monkey, the aiuhmpuid ape, and do to man himself, we liud lui incrn»
ing diU'ereniiaiiou of the cerehral curlei. by which certain areas of the oortsi
are bruiighl iutu sjimal couiitctinn wiib veiluio skeletal or other mtiacln ia
such a way that slinmlatiuu uf a uarlicuiar portion of the gny nwtUr jptm
rise to a particular nioveuieut, and to that atone.
§ 6fi9. In treating uf Ihe structure of the brain we u>ok« ($ 633} of tie
pyramidal tract ua utsriing frum the motor region of the cortex : anditii
obvious that the librca of Ihin tract miixt be conevmcd in llio development vT
the movemenU which ne huve junt described. When the mavDnieniSBn
biuujjhl about by nliuiulntiuu of the Rbrai in some {latt of their cuurw, in tbr
inieruiil ciiniiule, fur in»[anc(r, [here can be no doubt that (he stiniuloliaa
starts inipuUi-n which, iruvclliiig down Uiu tract to ll»C origins of oertsin
crauinl or itpinul nerves, in (oinc way give rise to co>>rdtnaie motor iinpuiw
along the laulor fibres of ihc ner\'t9 ; un<l tie mny with rcn«un speak uf the
iinpuUc* ihcD passing along the tract u motor or cSl-rent in nature. Whni
the stimulus is applied direct to the cortex, wc may assume that procwu.
started in tlie gray maticr, cvcntualo in similarly eilerent impulsca aloo{
the fibres of the tract. All the evidence lends us to regard tbia traet aaati
eRercnt tract.
Whet) i))« spinal cord is divided in ibe lower dorsal region and tbe tieo-
trodes of an electrometer on brought into connectivn with the troneverte cot
surface and with some point of the Kingiludinal surface above, the electro-
nieler giv<s evidence of currents of actiuo (.uanilested aa negative variatiuiu
ON VOLL'MTABT H0VBMBXT8.
817
^11 •InnirctttuincuTnnlornirrcDlnf n.M,§ 67) wtirnerer the motor sre« of
the hitiiMliiili i« HtiimiUt«0, but not when othiT pitrtttof tbucorl«x nrastiiaa-
tnu^. Wc- hnv« iilr«adj Mid tlinl «limulnti<>n of aiiy nnrt of tlit> motor
p'lD may uiMlcr Bboonnal c'in<Hlinn# givo ri»c to general epilcptilbrtn coii-
iont: vhvti tbtMoecur <liiriii): ciicl)an<>xpcHmpiil)u< the above, ctirrcnta
' Mlion nianif«t thunMlvce in lh« lower dorMil lord, whether the stimula-
tion givitij^ riw) to the convulsMtis bv ajiplieH to tlie area for th« hin^-limb
or 111 any fiart of tius motor rrginn. It Km bMin further obeerTe<l that the
carrcnis of action d«vel<i}H!>d irithin tJio >{pi>i>l <^'^ lallv in a very exact
manner with the muBculnr movonicnta. Th« eonvulgioDB begin with n bus-
lain«d " tonic " contraction of tbo mueclee, and the electrometer eIiowb a
(tmilar auMained current of avtiun ; this is followed by rhythmic moTemeiite
of the mwelci; accomjitinied by correBpondiag rhythmic niovcmCDls of lh«
awrrury of tli« «lectroiaet«r. Without in^isling too much on the exact
interprelatioD of tli«e results, we may take them as at teaat shonin); that,
«hi-n the lootor region of the cortex is excited, nervous irniiulses acconi-
|iaiii«d by " currents of action " paaa downward aloog the fibres of the pyra-
mi<Iiil IrHcE.
Thv [ittults of Btimulating the fibres of the tract !□ tlieir cmirae throu^
theeoraDaradiatauodilieiiilenial oapaule, and tbereeultHubmined hy#tudy
mg the de)(eDe rations fdllowiug upon injury to or remnvul of the seveni] parts
of the cortical motor region, agree in marking out the pullia taken by the
several couilittienta of the tract throu^'h the central uiittt^ matter of the
faemisuberc, the coruiia rudiata aii'l the cajwule. Comparing ^'fP- -*>^f -'*'*
with Fijp. litO, 200, and 201, it will be seen that the purtton i^f the tract
daiincij for the cranial nerves, and so l4>r the movements of the eyes, the
nouth, fact-, tunitue, pharynx, and larynx, starting fVom tlie ventral jwrts at
the morf frunlnl di«trirl of Hie motor ivgioo, take up their jHMJtion ai the
knw uf the internal opBule; and the portion destincid lor tho«o upper
cervical iM-rvts which carry out movements of the head through the mu«cl«a
of the nei-k. siaHin^ from the extreme fnintnl and dorsal |>arta of the area,
la nlw) appareiilly direclcd to the knpc of the cap>ulo. The rest of the tract,
tuning from t^ie part of the area lying at once behind and mcvial to the
above, occupica in the catwiile a petition posterior to them id the hind-limb
of tbe capaiue ; and it will be obwrved that the inict for the lore-limb which
beipna on tlie surface lateral of the tracts for tbe trunk and hind-limb, shifts
its course in rclntion to llieirs. so that in the cajMule it is in frurit uf them,
not lateral to titeiu. It mav further be observed that while in the tmcts for
the trunk and bind-limb the same tore and aft order which obtaiiiecHi ibe
surfuc-e is reproduced in the capeule, even apparently to the Strange prece-
dence of the ankle over the knee, the order of the aeTenI elements in the
fbre-linib tract which is lateral on tbe surlnoe becooiea regularly fore and aA
in tbe capsule. In tbe capsule the several elements are arranged in a linear
order, corresponding brou<lly i» that of tlie diatribution »f the musdee along
the lonKitudiual axis of the body ; on the cortex thev are disposed in an
order the cause uf which is at proient nut very clear, nut which i* probably
determined by the r»(>e«iive rctatiuus of tlie several parts uf the motor region
to the functional activity of the i>ther paris of the cortex. In the tibi(ting
ftoni the one order to the other, tlw several oMutitueiit fibres, as nv liave
nid. deai-rlbe a *uiUL>what iiccultar ciMirwo; and when we remi-ml>er, a? atatcd
la i 6tU{, ibnt the onlcr slioun in Fig. Uiit it only the order <ili(ainin^ at one
particular level uf the capMilc, and that from the doranl beginuutgs of tbe
i-aptulc in the corona nidiatit to its ventral end >D tlie pes the caiMiM is con-
tinually cliaagiBg in famx. and iia fibm therefore continually shifting tlteir
relatione tueach ollker, tl^wltole course of tbo etfvaral fibrm of tliv tnu;t ffvin
U
I
TITE BBJlIjr.
their origin in tlie corteic uotil tbey are e"t'>^'^i^ >*P >■>■<> ^ eutnl
or the ]>e« (Fig. lU'J, /y) idubI be a very complicalcM) one.
When the area nf one heraieph^re ta etiiiiulated, ilie movemnit miiA
reaulu ia in most caa^A seen on the other tide of the body, ami on that odiir
tide alime. Thus when the area for the fore-limb b stimulatMl on the ML
bemiti|iliere it is the ri^bl fure-liiub which ie nutveil- TU'a b in sconnluet
with uhac we have learned ol' the pyramidal tract and ils ultiuiate cniin
decusration befure it reacliee the motor iiervee, the decuamtiou either oetat-
ring n>a»tively as in the case of the crwKil pyraniiiUl tract, or in a mtn
scattered tiiauiier along ttie upper pan of ihe spinal euni in t)w cue of ikt
direct pyramidal tract; and. an w« have aeen, there is a nmilar decuMatiio
for Bucli pari of the i)yraiuiiial Iraa as in connected with the cranial ucrra
above the ilecuatation of the p]rraiuitU. Kxce|>t in the case of certain arot
for U'lveiiienis naturally bilalentl, of which we stiall speak pmeoUf, Ik*
ragveiueni ia normally on the cniHiwil fide, and on lh« eroaiid tide nly.
Uu'ler abnormal oonditiouit. howevtr. the liuib of tbc other sill* — lba( ii^sf
the BiUiie tide at the hemisphere sliumlalcil— may move abo. But Mchia
ahuormal ni<ivenient of the same side biut not tli« nroe character* a* ib«
projtff luoveuwfil of the crowed limb. lunlcnd of being an orderly oa>ili>
uate inovenieiit, it is a more aimpic, cither tvlanic or perfaaia touie, *t
rtiythmic. clunic, uontructiun of the muscle*. Obviously iia uwduuisa b tf
a dilleruiil nature from llint by whit-h the proper movement of tbe croaal
limb is effecteil ; but it ih important to bear in mind that a nKivcmaatlf
ihe uncriwwetl limb riiay tiike place; and further that, tbc abnormal caodi-
tiuiis continuing, rqmilar mnvcmeuls of an uncoordinated character a»t
iipread to the hind-limb anil other jiartsof the cn»Kd aido,th(Miffh llwitiaia-
laiion bo alill contiuod t<> thv arm arc*, llirn t<i Klbcr jmrls of iImi uiKtraal
side, until, ti* we have said, the whole body if thrown into epileptil^im mi-
Tulamw. Thi* feiiturc muHt not be forgotten. In fact, it may be Uiri;
instfWd upiin thai while we may speak of a (nrlicular cu(~>niinate oMnatM
us being the normal outcome of an ordinary careful stimulation of a par
ticular arm in a normal condition, it is no Icea true thai did'use uucu»rdia>t«i
movements, cilloiioating in general epilcpliruroi coovulsiuns. are tb« nstunl
outcome of tbe Btimiilaiion of any area in an abnormal cyindition. Aud ia
attempting lo t(>nn any opinion of the nature vf tlte 6nl Act, we nnal beat
the second in mind.
As ve Mid above, the movements iveulting from cortical siimulaii<>ii aa
most conveniently described in terms of parts of the body — of the arm, of ik
thumb, of the tongue, elc. Tbe moveinunla of the same pan uay ht
further distinguished by means of Ihe n»menolMure usually nd>ipt«d ia
speaking ('( muscular movements, such as flexion, eslensioii, ftbductMO.
adduction, etc. ; so that, within tbe area bearing tbe name of aome parlicnlu
part — such as the wrist, for i^slall(^e— ne have lo distinguish an area for iW
fleiion, and another for the extcimon of that joint : and in like mnniwr ia
refervnco to other parts. But it will be rMdifv umieratooil that it ts easin
to map out tbe area for a {xinicular {mrt than to dtminjj'iiab the an-as e»r-
responding to the aeveral movements »r ibut part. Huiice ilir uMmcnclaUirt
UHually 8du|>ted in speaking of the m>>l<if n-^ion U one biuol on tbe (kartaof
ti>e biidy moved, rather than on the character of tlw movements. Tli« ntort
cloeely. however, the ui»vemcnt» in ijucation nre studied, the mora proliabb
it appenrb tliat tbe liicalixaliou nbich obtains in the cortex is easentially a
localixation currea{>uuiling not to tiart« of tlie body, or to nervw, or to inaa-
cics, but to movcmcnio. In coniiilcritig ihi* point it must be fcmenibeieil
bun' rude and barbannii* a mcthixl of Mimuintion is th.it of applying electMdes
tu tlie surfacv of iliv gray matter compared with the natural ttimuliuioo nbtufa
819
taiiea p^Att during oerebnil action ; ilie one prrtbably ii about as mudi like
>tt is rtrikiof; llie keys of ■ pisiio nt a distance with a bnviiiMiii'k
to Uie exeriition of u skilled musidnn. Wert it in our power la stimulate
Um rortes iii any way at all approadimf; th« natural method, wo should iu
■II iiniliability arrire at two results ; ou the one hand, we should be able lo
produce at will a ramly of muremeiits of ditfereot de|;reea of vuinptexiiy,
■one very simple, otbcra very complex, aud for these we should have to use
mtBtt sugKested by the characters apd piirpofe of each movemeat, au<l Ity
thcH khwe: on the other baud, we should tind very decided limits to the
Dtimber uti kind of movemeiiU which wo couhl evoke, limiu fixed in the
CBM of each subject (Minly by tuherit«d or|{iiiii£utii>D, partly by (be iruoiu^
of the iadividual.
Some Buoh results of relln«d ex pvri mentation are, indeed, already fore*
■)ind<iw«d by ibe rude ratultoofour prfisent r»uKh methods. The uiovcmetitjt
which uauullr follow ttimuUliciu <if the inutor region, an<l which *ie have
dnMTtt>e<l a* llcxi'iu. utc. an-, no t'l ununk, the elementary fuctori of ordinary
budily moVL'uivnif, thu di-tathtiil and iuiperfeul chords uf n iiiuitical piece;
and iu the fotluuiu); fnclii rfliiliuj; to llicir priHluotiou tiv can recMgiiize llw
iuliuvotitit of iirjtaniiatiLin and hnliit. As we have Mtid, stimiilution «i' the
ruulur area of one beruinphi^rc pnxluow movementn, at n riik', wbicli arc
limited to uue hUU' nf thv Imdy, and thai the opfiiMtlc side Now b<)t)i in
unnolve* and in the bigbiir animals a large number of bmlilv oi<ivonic-u!«L,
capeeUlly of the tinilHt, arc habitunlly undntcral: and. puKiii^ luidc ihi^
4)il«alk>n why there nhould be two halves vf the bntiii. ainl why llie <in<> half
of tbo brain »h»uld be OMociniwI with the crow half of the tKNtily, wc may
rMOgnixe in the unilnlcml cr>»!cd niovcncnt raulLing from Htimulniion of
the cortex an accfirdancu with natural habits. But some movements of the
body ire ordinarily bilateral; 'he two eyes, for iiutnuce, are ordinarily
niuveil tii^lher. and the two side* of the trunk move together very louch
mure Ire'^uenily than <lo the two fure-linibs ur the two hind-limlw. And in
acioordance with this we lind that Dtitiiulaliun of the motor area for the eyes
on eitlicr hcniisphcrw pnxluceit moveuii-ntd uf hi>tb eyo», and atirnulation uf
the trunk an^a uf one hemisphere is aUo very apt to pruiuce bilateral at'tion
of the trunk musclei ; iu such instances the movemenu on both aidm are
cjuiie normal movements. We may iociileiually remark that remoral of the
trunk area leada to a jcoiid deal of bilateral de^'i^nerntiim, that is, to d«)[ener-
aiiun ufftiramlit in iheiiyruniidal tracts <iftiuih*i<le«, whereuMch a bilateral
deiceneratiun is L-»wparatively scanty after removal uf the itg ur arm area.
That it is the mtivemciit and not the iiart moved which ts. so to afieak,
repcvMnled on the cortex t* further ihowii W the relative m.i^jnitudia uf thL>
several cortical areas when (Ii«y are map|ie(l out accordiii); tu parts of the
iMxJy. Tlte are* for the arm, fur instance, cf. Fig*. 'Jll4, 205, is, so to speak,
raormoua compared ti> that of (be trunk when (he relative bulk* of theK two
partt uf the boily are oon^idercd ; and within thv arm area ilMilP the spncc
ooeit|ne>t by the thumb and lun-linger and diyit* i», bulk fur bulk, out of
proportion to the (pace allnited to the vibouhler ; so aUo the area fur the cyM
or (ur tlie mouth is out of prupvrliun to thu stxv of thoM organs. But thiM
niklive uies of the rc*pectivo nreoa becomo inielliglbla when wc bear in
reUlirc mobility, nimblenoM, and delicacy of exeuuliun; in themi
the shoulder is far behind tli« thumb, while the eye* and muulfa
moot other parts of (ho body,
^c are broui;hl yet a step furtht-r when we conpdu«, Id rMpMl of the
rorlieal motor iv^ion. anintals uf ditlert'nt i^rodee of vrgonitallon ; and the
results thus obtained lead us to the conclusion that (be motor region is orrc-
laied not to movemeuts in general, but to mowuietila of a jMnicular kind.
8S0
TQK BBAIN.
Tnking in wriee tko nbbit, tli« doe. tbe nonkcr, Mid man, w« find io pwring
from ono to tho otlwr, lui incmso m proniincuioe Knii in dirrvrrntUlion oftW
motor rvgion Bccompnnicd liy nn incrviiKiiti tho balk of tlii: prntmiilAl met;
nnioDf: tile mmt}- itrikiiig itifttrciK-rs betwBcn the bnin* of llwM Hvcnl anl-
mnl«, ihcpc l«o fcnitii'PK. thp jncrcasing complpxilj of tbr motor n{(lon. asil
lli« incrtnsiuj^ i^ize uf ihc pyrnniidiil tnicl, nr« among tbr tncet rlriking. Tbi
»ii« of llie nrramidnl tract is ilwlf corrclnlcci to Ibo comploxily of lb« motor
region, find, beinf; the marc rBsily detcrniined, may be uwd as iodintiog
butb : the difl'^ienee in tbe sine of the jiyniniidal tract in tbeao a&iuaU ii
eeen tUI along tbe whole length of tbe cord (I-'ig. 20tt). }4dw aa nguil
rio.ao>.
rvi
MAN
HOHKWY
OQS
DiAfliuH IO liuvncATi mi Rkutit* bus or the funAMWAi, Taimr la nu Doo, Hovui, u>
Mt.S. ItaSmMITOH.)
Til* flgnM ibDin In oailioe Uic laicnl hilf of iheciirt. Mlka loiol of HioaitblbonelowH
tn A, nwD ; B. monkvy : C, dot . A It ■ rriirodncUcin uf (* iii rii. Iiq ; b uid C tn ilniwn c> IM
Mm* dw M A. JV- iludod oblliilKlr. the p;runi>l>J tncl ; Ihe depth of iihtdlDff lndl<at« IMO U<
Inol ll Dorn rrovdnl « Itl) tnic jirninWiii nbmm wrll m larfcr In A (lun In a, ■ndin atkanliC,
In 8. jyiiui oallilnc ponlnna(ll)^I7nmlll■l inoi Kfanldl ftoinUiCNM bf the «*tfIj(1Iu ImM
JV.il. Uw direct tTTWnldal incl. pnwnl in maii oulr. Tbv (Imr ■MWrotat rMIvsIt luvtHC
t«nawll)«wciluii XHiolen rrdiunTeijjbuiiif |iup|>]r,
mera ouanllty of raovenient, if we may use such an exprcaion, the diflir-
eiicca DdwiL'H ihte« aninmla are of uo great moniiiit. If wc were (u take
tbe nniount of energy txiicnded aa luovi-iiivni in tvrfuty-four boutf per
grunmiG of iiiiinrle prenirnt in tbe bodv Eu t-iich of the four caan, we alxwtil
ccrlniiily not find imy i:urrta|iondpDce Dctwcun that und the fiw of the pyr»
midnl tmct. If, however, wc take n particiihir kipd nf inoveinviit, what «a
niDf pcrhapa cull (killed inuvnnrnt, that la, miivcnicnt ratrii-d out by meaiu
of iiilriciilc chniigt-a in llic cciiirul ntrvmia ayalcni, wc do find a rcninrkabk
p<iralldi«ni in Ibe above ctim-» hvlwccii thv iimotint of viich akillnl tnovcmtui
vnlciinp into tbe daily life of the individual and iW cia; of tho pyramidal
tract. In thcae two rrviwcla ninn is much nhovc ibc inoukcy. and the monkey
far above the dog. ^^ v may concUidc then Ibat the conicid niutur region »
in Fonio nay »]>i'cinlly couctmcd with tb« kind of movement which we ban
i-nllcd "skilled.^'
$ QS6. These skilled inovciiienlB are to a Itirge extent, though not exda-
aively, voluntary luovcmeuiB. We hnve in a prcvii>u< section eeen reaaon to
ibdiere tbal the cerebral cortex it in some vny especially awocialed with llie
deretopmenl of voluntary movemeDts. Putting logelhvr this conclusion and
tlw oouelusiousjuat arrived at we are iiaiuntlly led lo the further concIuaioD
tlial tJi« cortical tuotor region, uilli the )iyraiuidal tract belonging to il.
pliiya an inipurinut part iu carrying out voluntary movemeata. Di> tuber
&cl» support tbJH view, und if ao, nlmt light do ibe^ throw on llie c|Uce4ioD
at to what purl and what kind of purl the luolur region thus plays?
ON VOLUNTAKr MU VKU KNTS.
esi
1u ihb eoaneclion we nntumllir dnira in kwiw what are lite results ofS
rvniovi&R frotD aii mfaerwiHo Iniact luiimul the vrbole motor r«Kia<i. nod more |
flqweiftlly ihu ur iliat iwrtictiUr (M>rLiuD of iL Before prm.'eediuif further,
bmnver, we may mioe mure call attenlion to the uiiitiuii f(iveii in $ oOA, aad
npMtad in S 641 ; imleeil, when we eoiisi<ler ihe hiftli organixMiou ami com-
plex AiBctiooa which obviotuir b«li>nK (» the cortux. when we bear ia mJaii
llutl it ai>i>eara to ffnwrn. am) miim thcn'fore W hound l>jr clone Ihw to almost
all ilie Kttt of the L-viiinil iit-rvnuH *y«teiu, nc inuat be prc|>iire>) to tliiil after
rWDoviuK A portion of ci>rifx that the pure "<iefineacy" pheuomeiia, iIiocM)
«hich naull fntm the inere itlixcacc of a pieco of llie oortex, are larfcely
obtcureil l>v the other cfli«t« of ihv opemtton.
Ill the rabhit the rcniili* luivv liii'ii nlmiwt pureir neeAtivuL When in thi*
aniual the (nirl of the cortex which may Iw coiixiilereil M the nurtor regiuii
U rerooved, nolhiiiK roninrknblv it obverved in [he uovoiiietilJi of ih« unimiil.
We can hnnllj xii{ipi»(! that (ho 0|>eralion« of th(< contml ni-rvuii* Hvntcin nro
the «ame iu an o|M-rnleil iif in an intJict tmimni, nnd tliv ditlrrvncv* imlucwl
ought to be brlntTcil by the movuniunts uf tlic body; but nl prtnont ifaey
hare <wca]>ed olwcrvation.
In the do); the reiDoval of bd area i* followed by a loM or dimitiutiou of
voluntary movement in the cvrratpooding part of the Icidv. Wb^-n. for in-
alHn<x-. tiie area for the fore-limb is removed froin tli« lull bemi'pbere. tho
ri)£ht fore-limb ia eumoletely or pAriially "paralyied." In carryint; out it*
o^innrv movementa the opernle^l nuimnt makes little or no u«c uf ila riy;ht
fure-liinK Rut this stntv of thin^ is temporary only. After « whi la the
animal renins power oi-er the limb, and in HueeeMful ooaea reooTl^ry is so
complete that it is impoasible to point out in the limb any iippreciablo devia-
tion f^>^l the norniiil uw. And eareftil examination after death has shown
not only that the aren had been wholly removed, but also that there was do
Rgeneratitm of the hot imrld ; the removal uf the cortex leads in such caaea,
M uaaal, to de;^neralioa of the cornspoinlio;; »lrand iu the pymmi<tal tract
right awuy l'n>m ih^- rerehral mirface, tu the endiuK* of iho strand in the oer-
Tjoal and donal spinal curd. Nor raiii it be ur^l in such awei tliat diOused
remiiaJita of the arm an« hnd Iitvd lofl in tlui raiuunin)i; |iatl.-> of the motor
refiioH ; for the wholt- motor rc)pon has been nniOTcd. an<l y>-t the aotmal
haa r«coTereil to sueh an ext«Dt that a casual ohaerrer could detect no ditler-
cnoea between the movementa of the (wo >i<les of the body. CluMr examiua-
tiou did difi^loiie certain im per feet ion* of movcmonl; but the Ofiention had
involved injury t« or priMluced c)isn);ce in structures other than the motor
rtgioa.aw) the impcrlectionf niij^ht have lice n duo to tlir additional daniaan.
Morcan it Iw nr^ed that, in such a ca«-. where one oide i» removed, tne
reiuuning hrmispbere takes on double functions : for the ^rvutvr part of the
motor an-u have been reuMved on both sides, and yrt the animal's m»v«-
rornto have iMvn so far apparently complete that a casual olnervcr would see
nothing si ran}M in them. Ag^in, the whole motor region bos been removoi)
frcicn one hemisphere in a yooni; puppy, nnd sometime later whvn llw (nixrc-
ments seemed to have rcoivered iboir ncM-nal condition, (he removal of the
mot4>r rej^ioiii of the other hemisphere has produced merely a paraltvis of the
criMsed side of the body, an<t that as bef>>re only of a temporary chanu'ier.
Two things have to he noted here. Iti the first place, the removal of an
area does siTiH^t the iDOveroeuts which are brought about by stimulating that
area, it leads to their dinppearanee or at leant lu i;reat diminution of tlwm :
and this aHiirda ati niliittional arcumeut that the ooiuieciion between the araa
and the movement ia a real and important one. In the aeoond iilace, thtt>l
physiuloffical effect ia lemporar}' only, thouj^h the analomical results of the
operation are permanent, for tlie cort£x is never renewed, and the pyrmmidnl
823
TBB BRAIN.
Iraet degenenitea nlnDK its « hole Irogth, never to be n»tar*>l : th'w •b'tui
thai we have to dest bcre with events nf m wry complex cbnnictvr. Wliiv
a pHrliciilar iDiiveineiil multf from Blimnliitioii of the tip{iro|irinl« eartii'sl
nrea, we may be sure that whatever takes place in the mrtox anil nloog Uit
pTrunii<)al Imi'i, inouir imrul^^es, duly co'Winated. pass alun^ certain >&!«•
rior roola lo certiuii rhim-'U-s; and we know ihm if we reniDVol a sulSnfM
length of Mfh (if thi'U^ anterior ruoia that fianicrulnr movemeut mikiM ht
li*l for the rt-rf "f the life of thr incliviiliial. We may. iherefuf*. infer ikai
the cvvutN whi<'h, nhait!vi-r be ilii-ir eincl nature. lukinK plucv in the rona
and along tlio pyramiilnl tract Ictiit iiltiiiiutrly to iIh: imuc of molor Impute
ulong the anlcrior nmti. ililter ■■RH'nitMlly fWjni the evcDta attending ik
trnnHmiMion uf oriliimry motor impulKii.
In thi: cn«c of the- monkey, the mult* of reuioriof; parla of the tntlhil
raotnr rejci^o hn\'e nni iH-rn no nc(?onliini n* in thi^ I'luw nf the 'log. Tht
two animnU a^i> iirrfvclly in ■» fnr that tho roniovnl of a jwrlicnilar an*
leniln, a* an immnliale rrsiilt, to the him of tlM Oormiwin'lini; miiveiiMil ;
hut wliilo in *nmp inxtance* nrcovery of the moroionil Iiiif in (he ninnkeyu
in the dog aftvr a nhile taken place, in oilier instaiH^cs llx' " iMralran" las
appeared to be pcrmanmC As a rule the pamlvHS Miiml by a lar^ IrtM
is Dot only niorv extensive, but also of lonf^r dunitioii than that cniiinl hr
a small one; and natural bilateral moTemcnts. as of ihe erv*. rv«pp«r
earlier than iinilatL-ral movements. The facts, however, witltin otir kimwl-
edf>e relnlin); to the permanence of the effeet arc neitlter nuincrimit nomad
enough to justify at prevent a definite crincluBioii. On tbe one hand, th*
paeitive csma where recovery has takeu plm-e are of more value than ths
iM^tive ones, since in the latter the recovery may have been hiodend hj
concomitant evenis of a natun^ which we may cull atciiltiilal : and It if at
k'li^l It priori mtBt unlikely that thi; pyramidal Iriict nici'haninu, if we niajr
nsO tho i>x|>ri'iuiiin. I hough il mnv dilb-r in (bo moiikrvam) the dog in ckfTt*
of develop mi -lit, ditl'i-m m> eawntmlly in kind ihat damagi? of it lend* in lb*
one case to pcrmaoont, and in the other to mere (cm{iorAry, lij«« of ftinedoa.
Wo may adil that «<■ .nhould further oxpect to meet in the monkey with
more prominciit and more lasting complicnlions due to the suluidmry cdrdi
of tbe operntioii, and il may be lionbled wlMrthcr in any of th» rvcurdid
evperimcnis tiK- animal ha* been allowed tn live n sutBeicnt time for ibw
subsidiary eveniK In hnvn clearvil nway, having only ohnt ve have nilbd
Ihe "delieieney" phenomena, due to the loss of the (sorties) area ainnc. On
the other hand, it muit be rememben-d that iho mnventenls of the u>onk«^
are more iulricute in origin, more "skilW than those of the d'lf;; and A
may be lliat diHerences in the characlcre of movenieuta determine the pdw-
bility of ibeir recovery. In illu''tr,«tiou of this we may fguoie the expe-
rience thai, after the removal of the arm area in the monkey, a oenaia
awkwardness in the movements of (he (humb is one of the last eBVcta oftW
operation.
§ 680. Bcibro wa pmoeed, however, any Ainher tn Iho discussion, it *i|^^
be of adviintnge to turn aside to wliat is ktionn concerning ibe c«>nl^^H
motor region in iiiRn, As we have already said. the<in.'tiL*al eoiujiIeratJai^^^
lead us to bcliere that the cortical motor region lu man u disfiuaed tu nccunl-
ance with the plan of the anibrapoid aiie a* aacertaiuod es peri men tJilly. bvl
with (he diflt'rentiation carried siill lurthcr; and llw few caaat of OJtper*-
mental stimulation of (be human cortex support ihi* view. Our chi«f
knowledge in ibis matter is derived from the atudy of diacase; and In tbit
the advantages of dealing with one of ourM-lvos arc lar|i«ly connterbttlaneol
hv tbe disadvantages due to disease being m olV-n anatomically dillbsa and
|tli}-ei<iIogically vbangeful and progrcMii'V.
OS VOLDSTAUr MOVEMBXT8.
828
"We Mi'I almrn (hat iluring «s]>rrint«nt« on niiimali* atimulnlitm of any
pATt of iltu iTKilor rrfpon may iiiii)<t nbiivnDnl nKiihlimiii Ii-ml in i;Mi)!mI
vnilvpUform ounvulawxin. Now cliiiicnl ftudy liiia showii llijit id man certniit
klildl of vitilvptic nllni-k* iirr of Miiiiliir cnrlicnl nri^m. In tlione ciim!S it
his bcon ooHrrcH that thr mtark ln-^in» in n ptttliciiliir tiravcnieat, by con-
tfaetkmt of (Mrticiilnr miiKcU* or of ilir imiK'k'* >*{ it luiriicular rrffi'in nf
Die tHiiJy, «f iho haix). foot, tor, ihiimb. <>ir., and ihrii xpmiilii in a liofiniic
ordrr or " miirch " over iho miiK-lc* of other n-giorw iiiilil the vrhok IhnIv is
iavolvrd. When in nn fxporimrnt un iin miimnl cpilqiliform conrulsions
Mipfrrfnp, thry jimilarly »i"rl fn-m ihe region of the l«>iiv. the motor area
. of whii'h u Iieiieaih the electrtxlrx at the time. nnH »nnii]iirly spread by a
i]e6nire " march" over (be whole body. Hence, in ibe htiman e|iik{>tiforiD
attack* of which «e ar« cpenkini;, it bns been inferred) thHt tho immediate
«x«iting oaiiM of the Bltsck )■ to be nought in events Hiking pliice in that
part of tbe eorlex whieh cervee as the urea for the movement trhich ushers
intbenllnrk. Further in4|uiry hns Dot only coofirmed this view, but baa
aboBhomi tbnt the tonography of the eorti<«l areas in tiuiti. u» thus deler-
tnined, very cloeely follows that of the niookey.
Other diseases of (Ik eartei have been ninrked. among oth«r sytnptoins,
hy Ion or impnirment of pnriicular niovenieiils. In must of siieh co^es the
cortical krioD bns been of such sn extent ns to involve a number of s|>eetal
areas at tlie same time, nod «o to lend to luna or impairment of inoTeraent
over relatively cunaiderable rej^ioiifi of tbe body, such as the wliota of OUB
arm ; and ii> general the teaching of these cases of disease, while eoafirminit
ilia deductioDS from the monkey, and givin;; us some general idea of tite
topof(raphy of the human motor cortical region, has at present given u>
approximate n-sulla only. Figs. 209 and 210 ahow in brund diagrararaatio
maniwr tlw im^iuud and relative extent of tbe motor areas for the leg, arm,
and fae« in mnn, ni< far ax bai yet beun awcrtaincd. To aiwist the reader
we givi! ul tbe ninie lime dingrams, Fign. *2))7 and 208, illuatraiiog iha do*
ncDclalurc of the surfaiT of the hunmn brnin.
One ana is of special and in«troclive interest. Sp«*ch is an eminently
"skilW movemctil. We havo »wn that in the monkey tbe arxia for the
mouth and tongue lies nt the ventral end of the eeiitml timiire or fissurv of
Rolnndo, ventral to the arm area, and that the extrrrae ventral and front
p<irt of the motor region just above the Hwurc of Rylviuo supplies nn area
which we marked as that of pbonation < Fig. 'i04). In the monkey the area
of pbonation is determine<l by experimental stimubition; in mnn, in a
similar position, on (he third or lowest frontal convidmion, R>metimes eallct
Broea's convolution, ventral to and in frvmt of, and probably overlapping
backward, tbe area which in Fig. 'JU!> it marked " tnc«," and which incluwi
the mouth and tongue, clinical study has dtschwetl the exiatenee of an area
which may l>e spoken of ns tbe area of "speech." Lesions nf the cortex in
this area cause a Icm of or inttrlerence with speech, i
known as aphatia; to this we shall presently return.
■8 shown in an approximate manner.
The movemenls of epeeeh are eaaentially bilateral movements. In tbe
dog awl monkey various bilateral movemenia may be excited by stimulation
of tbe apiiro^riale area in either hemisphere ; and analogy would lead us lo
Biipjiotie that lu man the movenK'ut« of siieech would be connected with (ho
ff)e^-h area in b<>ih one and the other ncmispbere. The results of leniomi,
liowever, show tbnt it is in moat caaca eapeoiallr tbe left bciniaphere which
Is connected with i<|ieeeh; it b a leaioa in tbe third frxmlnl convotutinn of
Ika left hemisplK're, olten nsaoeiated with other Iwions of ilie same b«Rii-
•pban leading to paralysia of the right aide of the Iwdy and facv, which
'h, the condition being
In Fig. :209 thia area
89$
THE BKAi:r.
c«ue«» nphuia, it Wing nnlr En cxeoftfioiMl etut» thai Ihc oonditJOD
from & IraioD of llio comspf>ii<ling nrca of curlcx oo tl>e right bemiapbore. '
Id maD, thcD, clinii-al study corroboratcc tlxt oMicliudoot dcdtiCM turn
R 0
.0*)
I'?-.:
nr-O.!
^
^■«A
-*'
;^>r5
lO B C
^
't>.
'■^^
T E M Po BUL
-OB'
UluMT iUwu-HDi or Uni'. «to*tu)
Ftc-W,
0..B, ,E
.^;^«»i«.
^*/.
TM
I."
UntjJ
.N
■Xo
ot
The ume oa mx Uboal Scwao. (OomM.)
tn botti llfiixM ttia uiipl an luiliioMd tr <Mle >nd ll» oMiTolaUoni I>t roniKii tni*. TIm IhUaa*
lliBlUto(wnici)>iioi]}miiiiiT)>crhB|Bl)coI UN to omDMIlcti Willi Uwrc Ikllla >Dd tboMcf tt*
tnln or Um luonicy iFigi OK uul afl).
Oyrf «r nnnvlurriHit, rm«ntni1 nr anUrlar rantml — unndlns ftonul. fodccMnt or ir^ilw
tmn«l*>*«siidIni[|iiuteDiI. SupMorMmpanl-lDtM-aaigliMlvilnKcnironi. Triuvular loboto
aninau*, C»nti»J tobt-MUwl nf Rsfl. frntiionitnl int>Ul*~lli« tnoaUl Ikta of Uh- raprfMr
(raoMl. >HIilD lliv nuflRiil trnu. (.IhsuIdtii — Uie |«/i of tin qtih toniliialiB wblch ul)Daa>tl>B
eorpo* oalknun. Ojtut hljipaiamft " aaiilotUi lym*. UhmikIi Uu laltn nuDO li KiinrllnM* i*-
iMoivl Io ibc ftwii t«tt of tliB lil|i|NMiuii|«l grtvt : iha iin> mir ta caitUviwl oi a aoarUniMUiin i4
UnsTTua fhrnloMiu. aod ilw Uim lnc«lher. flitDi[i«B Brtn, turn btoa tallnl "Iha ^isl linUo
aiM«r/bnira. OmUmI — HfiUndIc or of Rolando. nqViuUcnlBr— farlOI»<]eel|4l«L t^nsul
— tain'infleMJorwiBfUmeiUiWr'puMBL TVmionwpbencMalloto— tempomllobo.
rOLUN-TABY MOVBUEXTS.
825
rimenliil iDTefltiKiitioD of the dog and of iho moiikcT, but (till
FM ill iinoeriaintv oa Ui tbe (|ue8tioa what noil whiit nloiio arc tlto
lutelj permwieni eSeeta of the loos of a oirucal arra nn<l mithiag eUo.
tfa* OHO hand, io Um cam* in which reoovery of a moviMnenl Tullow* upon
rio. nti
f~i.'^ t
lft<-
-J»
Oe'
M EARING^
Smal 9emr»ea or nm Rioirr CMtaau. HEHnmmt or u«m ih OvTuya. to iLuvnun
T«i Cmtiul Aaau. (KcdoMtf (Torn natniv.l
TM pnlUon of llw anw of Ui* conoi couocraid whb noTamant* of ItM Am. arm. &nd les. •x'l
■1B> Uie wn— of dibt mJ twriin «» *ppwjlm«t«lr ilwuni, TlwimlUMartlM «>«« eooiuwMit
«iUi itiMfk (BfOM** omifTlHilao ihnn fbr iba Mike Of dMttMlmi of II wlib (bo tadtlnncf IM
Ml tb*r«i>«>cnutiuii i>r*imui»itwranajce(miil U■^ bowvrer, In ibc ^ bomUvbcn
Oe. L, wclplul lobt. Ft. L. frooUl lube: ft- 1. MuiponJ lot*: ^, /. itw •*■«« of
1i*: C./. ibei»DUmlitMim(Ki>Uiv1lc]; Ctn./iliidlralM ib* putUon or IbepMicrtorcailcf (lie
■flia«l HMire.
m. m.
Fr-Ll
lOii>
»A
Tf L
TBS Hboal SumTAOi of nn Rnun Cuekkii. ducnriiKiii or His dk Ovtum. ni iLuwraiis
Tat CUKTICIL Amku.
TbavBualwwuuviIiowoonticclnl with the iDomnmliof Ik* IcC ftod with thcKMia fif righl
a*d hhU. n. L. am IkMiM polo ofUie boioliplioni ; Oe t, ibaocM(«ul polv; ft L. ilic wot.
pMkI id(c: Ch /, iIm aUkM»«i4iitiiBl taam nfantlni Iba musln>l sjnu •Idt« D'Om tiw tcynu
fcwiWWt l*law : Cf. naifet lb* iltiiallnD of Ibe omukI llBar*. ili« Omits li>«lf not Wag aiiiMRiii
ealfeaBMtlaiicMaf lb«bml>iAian. Tb«niiiiiBfaIlamiauida«*Blerto(eoanl«nii«««Maii
taatmrnMau.
?2* THE BRAIN'.
its I-r* T ImpairTB*EiL H w •■■vn for us to niopoae that the lerioo itoelf ««>
r«ir rorsr*. »c-i '■fis.t »i'e tfc* «ire of the m&Iady the cortical area r^inai
hi n-«Ea! ocii'-.o. *~*a cEw other hand, where the diseftf« contiuupB, the
ivrmioencT . f :s^ \.m -4 anr novemeflt may be attributed to the disax
d'icz T.ire t'ras rr-rr^r H»r«fHi the function of the cortical area. Aphasia,
tst'T'.ial'r ia T'«:dj r^mc*. ha? been followed by recovery, but in such
ra**^ i; fca* befo "rrr.-e*'! tbat ibe donnant area on the ri(fht side ha* heen
awaken*-! M ntiv;:T m ibe !•■*» of the left area ; and in support of this
Titw oa^ee have ^**n fW»->pi<d in which a firet aphasia, due to a lesion on
the \ih ^de. hs^ hreri f'-illow^nl bv a second aphasia due to a sequent legion
fptxurrinz ■■n tbe ri^bt si.le. On the whole, perhaps, the evidence of clinia!
»ludv tecd< '■-' fb>''w chat in man the Ion of movement due tn the destniclioo
bv -lijeafe <->f an area is a permanent one, though actual demonstration of
lhi# i* waufin^.
^ S6L We mav n->w retom lo the discussion of the qnestinn. What ia lli«
rart plaveil bv a m.»tnr area, and bv the contribution from that area to tHe
piran^idal tract in carryinz out the movements with which the area isaao-
ciaieil?
We mar premL>e ibai ihe evidence points very distinctly to the roncluoinn
that Khatever he the nature lifihe whole chain of events of which the cortical
area -'eem^ to Iv a s^n of I'enlre. the fibres of the pyramidal tract serveM
the ''Iiannel of nrivewe^ which we must regard as eSerent id nature. Ii i)
perfectly irue that in many case!> at least the removal of a cortical ares hu
le<l io diminl-'heil vendibility of the part iu which movements are excited br
stimulation of the ar^a: and there are many facts, of which we shall pre*-
enilv qu'Me a very ^rikintr one, which go to show that the cortex of tlie
m-ifir Tvjiivn i* larselv intluenccil bv sensory impulses from various parts "f
ihe K»lv; but we i-annot suppose that the pyramidal tract is the chsnM!
liv «hioh siii'h *en*-'ry impuli=es reach the cortex. As we have previotislr
a -'>6:l ur^e<l. the fact that the defieneration of the fibres in the tract iii
des^vnitin^ one. cannot be trusted bv itself to prove that the direction in
which the tibrwj carrv impulses is only that from the cortex downward; but
this adiieil i.i the tact that when the fibres of the tract are stimulated at mi
psrt 111' thfir iiiur*e. movements, the siffns of the occurrence of eSerent eva-
tril'u!:al impttlfv^, are produced, leaves no doubt that the tract is one nf
efferent libre*. Hence we may infer that whatever be the nature of the
events tskini; place in a motor area during the carrying out of a movemenl.
the ]wrt (ilaveil by the fibres of the pyramidal tract is that of carrying
eflen'nt inipiilj*^ from the area to the muscles concerned.
Let us con'^iiler tirsi the movements of speech in man, the evidence tonch-
ing the connection of which with an area nn the third frontal convotutinn
a|i|>ear; lui verv clenr. ^^|>eech is eminently a "skilled" movement; it
involves the ni<>st delicate coi'rdinatioD of several muscular con tractiMU. anil
we m:iv ivrtainlv sav of it that it has to be " learned." The whole chain of
oo<inlin:Ued events bv which the utterance of a sentence, a word, or any
yncal sisrn is aixiimplishei! consists of many links, the breaking of any of
whiili nrll lead to failure of one kind or another in the act. Something maT
go wrouir in the glossal or other muscles, in the nerve endings in th(»e
nuiiscles. or in the fibres of the nerves, hypoglossal and others, between the
centml nervous system nud ibe muscles, or something may go wrong in that
part of the eenlral nervous system, the bulb to wit, iu which a certain amount
of coiirilinatiun is carried out just previous to the issue of the motor impulse.
Paniiige done to any of these parts of the mechanism may lead to duiubnes
or ii> imperfect sjieech. In the latter case the imperfections have a certain
character ; if we are at all able to gather the wish of the speaker, we recog-
627
ittf thnt Ke » Bll^mpliiic m inter r)i« rii:)il norHn in t)i« rifiht R«<|iiciiro, hut
tliitt his rflbrts are frtwf ral<^i) hy imporfcct cniirdiitfltioii "r imtwrrcd nni^ciiliir
■etiao : hit flpech ii " rbirk," the B^llablM ik hlnrred mtd the like. On-
w of the bulb at times WaAa lo iinpert^ apecx'h of thia kind in whirh thr
p«ri«clinn may ho r^cofiiiiEeiJ as due tn Ow luck of proper co^^rtli nation of
Enninr imptilwa. The alfeetton of speech known aa"apliaBia," which is ca need
Ihr Iwioni oflhe cf-rtex. is ofa difli-reot Hiaracler, ami the furmH of imperft^ct
'^Speech caused hy bulbar diwaite huve juMlv been di»i intuit lied f^>n) (rue
aphakia by the uoe of oilier ternii. Csm* uf complete aphaiis in which all
power of speech in li>«t. do little more thRO help us lo ar(<erlain the to)x>-
graphical pnsiimn in the corlex of ihe "speech" an-H, hut cosee of fmrlial
■pbsrla are e^|)ec-iiillr iuMruHive. Without ntiempliiii; to ^> into the details
of the Biihjrri and inin the ninny c<iii:<idrr«ti<in>< which have to be had in
I mind ill (Wlinfi niih it. for tberu are difrcnnl kind* <if aphoiin. we may ren-
lure to Niy that the *irikii>jt friiturv of jiarlinl aphaxia in thi> failure lo sar
certain «oril>i or orllnhlr*. and the tendency to riilwtiliitc *omo wrnnu woril
or *yllnl>h- for ih*- ri{;hl une. The words nr sylhihli>» nhich are iiilered are
Hchilv fninnuMCi'd iviihoiit deftrt nf articulation; and iu many raw*.
ttiou|fh the Hf;ht woril rnnnnt be pnHluccd nr i> direct cRorl of the nill, it
Inay he utlrrt-il under thi- inHiicncf of nn cnioiion, or indcc<l M>melimc« aa
ibc n'»ull of »"nic jwrcbiml procrwun more cnniplcx than lh(i*n involvcrl in
lbs nif-rc volitional i-tToM to i<ftv ihr uord, An inslruclive cii*e i» rcfvirtled
of m mau nuflriintr fmrn vli^'ht nphaxin. who after Mvcra) failum to any llie
word " no" by ilwlf. nl laxt Hitd. " I cnn'l «iy no. rir,"
From the phrnnmeoa of jinnial npha^in we may draw the deduction that
the corlirat upcech area does not carry out the whole nf the coftrdinalion of
Ihe iinjtiiW* iiivoh-cd iti nrltntlalion. Thnt co'^rdiDation is exceo<lini;ly com*
ptex. kikI ue oupht (ifrhaps to recof^iiie in it more than one decree or kind
of ciinrdinalion. The laiture of articulation in disease of the buln show^ that
• certain amount of cmi'rdinaliou lakes place ibere; for the affection* of
qiccch due to biiltmr dieea»e are not ibe »ame as those reetdlinir frr>ni ihe
intf* loss of this or that muscle or nerve. We munt. of rouree. admit that
■nme, poanbly a creat deal, of co'kdination of n certain kind take« place in
the cortex, for the bulb cannot by itielf be made to fiiteak ; exactly how
inuch. the knowlcdj-o at |>r(«ent at our disptwal leaves a matter of threat
VDcertainty ; but it in itiijiicienL for our present purpoae to reco^iao that
whatever may be ibe nature of the eventH takiuK place in tlie cortical arvm
durioff tbe act of Bix^ech. those events make u^ of ibe machinery already
proviiled in the hulb. ^^le word iipokeii doe* not stort. mi lo speak, ready
Itmide in the oiriex ; it is not that a f;roup of impulMs atart fKini ibe cortex
with their coordination fully achieved, and pass aloiitc certain ncn'e fibres M
certain muscles, making their way without rhunire ihrtiuffh the lantcl^ of the
bulb, as if this were merely n bundle of linen oflerin); pathn for. but exercis-
ioK no influence over the impulaeH. We miiM rather anppoae ibal Bome*biDfc
lakes place ia the cortex of the third fttm.lal convolutiua, aa the result of
which eflerent impiiUea paaa alo»j{ the appropriate fibm of tbe pyramidal
tract lo the bulb, ami there »tart a series of evenla leading to the i«>>ue of the
Ordinatod inipidi>ra hy wliicb the word in *p»kc]i.
(668. We have uu n-ajHin ulialL-ver to think that llic cortical area for
speech dillbrx in il* fuuiltitiii-nlal ohamcterw from other didikwa of the motor
legion, and an- Jij>lifii.-<l in carrrinR on to other ari-aM tbe deduction we have
JnH dnwD in ninncc-tion with the (pt-rch iin-a. With that end In view ne
nav now turn back to the expcrinwnlal multn olitaiucil on the do|>, and tt
will niakr our dltciitninu iiiniplcT if we lake aa an illustration Mine large area
such a» the lore limb aren.
8S8
THB BBAIS.
We have wen that Mimulntion of lliij) urtA pruduoe* what ire msf. to Btart
with. »|)eiik of •implj' im roovemrnt* of the fore-limb: ami gtiMed by
nnalutcv of Mf>eech in mtiD we mnv confiHvntlr conclixlc ihni wlien the i\of
volunliirilx mnrvw iho foro-limb, tlic net u cnrriMi out ttv ni«Hiu u( event*
tiikint! pliicc ill ihc fon-linil> cortical iir«&. The fimjilicily "f the ekctri-
cal pfaenomeiin i-fsiillins from cartiual slim illation, which vrr dti«(-rib«d in
9 658, Diigbt at lirst si^nt lend ui to coiirliiilc thai the wlmk- lumber wv
fiiirly mmple ; anti tndcciJ sum« wrilcre ni>prar to vntortain lliP oooorptiM
that in a voluntary niovcmriil Kuch as that of the foro-lirab, nil that
Inki^ ptacu is that the "will" itimiilatM corlnin cclU in thn oortieol mm
cnii*inglhe<]ischar^of nxilor impiileea along the pyramidal fihraa onDMVtn]
with ibufir ccIIb, and that th^se molnr impulses tnvel strsif;ht d'>irn llit
|tyrnmidnl Irnct to the motor libras of the appropriate nerves, undrrcoiag
powihiy »nmc cban^K at the place in the cord where the pyramidal tibn
make* iimction with ihe fibre of tlie anterior rout, but deriving their chi>^f if
not their whole co'^rdioation from the cortex itself, that is to say, bdng eoOr-
ilinated at their very startin;;-poiut. That such a view is nnlenablv, nsd
thai the simplicity of the electrical pheDomena is miilendiu^; is shown by llu
(■illowint; two oonsideratioiu amonj; others: Ou the one band, as was sbnoa
in a prei-iods eeclion. the co4'>rdi nation of movemeate may be carrtMl out ajiatl
from the cortex, namely, in the absence of the lieniisplieTes ; and we caa
hardly suppose that there should be two (|iiite disliocl systenu of oo-'krdin-
tion to carry out the aaroe moveoieut, one employed when volition was the
tnovinR cause, and the other when aometliin^ else led to the nM>venient. Un
the other hand, the analogy of speech justilitM us in conoludinf; that the cor-
tical prnoeme« <lo take advantage of a ooonlinatiuo effected by the aclwia of
other parta of the nervmi* nyslern.
Hearing thid in niiiid, we riiav recall attention to the remarkable effv^l*
which result from the rcniovnl of iho area. Thefc arc twofohl. In the Hni
plave, there Is more or 1cm cixnpli'tti paralysis of the limb; all the mor*-
menu of tbv limb are f jr n time ineHcciive. It » not that purely voluntary
movemenis are alone, so to speak, cut out. the relli-x and other moveMeau
arc a\m impair<'4 or lemporarily abolished, and, as wo havu almidy sai<t, ia
manv cases at lni»t the sensations of the limb are interfcrxvl with. Thev
tro(ible« are of cmirso in part tho oflects of the more ooerttive interferroa
belonging to what we spoke of in $ h6'i. as being of the nature of shook.
But. even giving full weight to thL< consideration, there remains the llsi-t tbil
the cortical area u aatociated with the various coordinating and other am-
Tous inwhani^ins balooging 10 the limb by such dote tie* tfaat these art
thrown into diiorder when it is injured. And side by side with this we may
put the remarkable fact previoniily sialyl, thut during an abnormal oonditiuD
of the cortical area Bti mutation of the area, instead of producing the appn^
prifite movementa confined to the limb, may give rise to niovcmeuta of oibtf
parts ciilmtnaliii); in epileptiform GonvulMuiU.
Ill the Hrcoud place, ihit paralysis ia temporary only, tlie voluntary mcft-
menu are after a while regaiiied, ami that in spite of the fore-liub moiety of
the pyramidal traei pcniiauentlv degenerating alon^ its whole lenf[th, oaxket
it nor the cortical area ever being itgeneratiM. This shows that whatever
be the chain of eventa in the intact animal, it ia jMesible for the " will " of
the animal to get at ilie musclca and motor meehaniams of the fere-limb br
some other path ihaii ihnt provided by the appropriate cortical area and
oorreapoiiding |mtliof the pyramidal tract; and ibeMcts previous! v recufdvd
(S 659) ahow that that other part ia nut the correip mdin/ part nf the pyn*
midal aystem belouging to thti other half of the heini«|iiiur«, an<l inilM-d it
not any part at all of the whole pyramidal system. The '• will." whatwi'T
I
ibe proccMM hy which it tiikt^ origin, am) nlxervver b« the pl<w vrhcrr
ih^TBrv carried vii.U nlik in thcitWncvof the pyramidal i^stem, to {iruduoe
il* •flbet on tho inotvr Gbm of the brnchinl nerves !>/ working on oLh«r |mh1b
of th« CBDtrml itcnroiis FVEtcni.
Ht-iict' whilo fldmiliin^, m wc miiH do, thnt in tli« intnct nnimni the cor-
licul nrcu and |>ymmidnl tract piny tbcir pnrt in cxrrviiig oni volunlnry
noTemfiitB, th^-ir action i« »oi of that »iinp)c chnmck-r fiippuvcd by lh« view
rrfprml to nbov«. Od tlio conirnn', we mro drivQO to regard iheni ralher ib
linke. in)|N)rlnnt link», it is true, but flill links in a complex chain. A» n-v
have already orged, we may prolwbly speak of ihe changes taking place in
th« pyraiDidal fibres n« being in the whole of the natiirv of efferent itDpulsea;
but we thould be going beyond (he evidence if we concluded that ihey
Here idetilical with the ordinary efferent impulses of motor nerves. And
above all it must not be left unnoliced that the corlicnl area has close, if not
direct, oonnecliom of a sensory nature nilh the part in whose movements it
b concerned. This is (hown by the foMowiog remarkable results whieh may
Drake iheir appearauce when Mimulalion of llie cortex is carried nhile the
aniioal (ihrgr is in a particular stage of the intlueiiL'e of morphine. If a aiib-
luiiiinial blimulus be found, that in, a current of such iulciisity that applied
to a mulor area it n ill produce no movement, but if increased ever so slijchlly
will give a feeble contraction of the appropriate miipclcs. it may he observed
that a dU^hl stiiuolue, sucb as gently strokine the skin over the muscles in
(fiwfltinD. uill reiKJer the previous submiuimul slimtilua elleelive, and so call
liirth a ntovemenl. Thii^ if the area experimented on he that connected
with the liDiug of the forepaw, and the euhminiaial stimulus be applied to
the area at interval*, after several applications followed by no moTumeuta, a
fentle stroke or two over the skiu of the paw wilt lead to the paw being
d1»l the iwxl lime the mininlnf U a|iplic<] to the area. A umilar rt«uli,
hut l«ai sure and vtrikin);, may follow ujion the Mimulntion of pnrlM of the
body other ibsn Ihv |mrt ■'ornvpfiiiding to thi- arm stimulated. Then, nirain,
it haf Iwcn ol>>crviH) ihnt in M-rtain other xlagcs of the influence of mnrpninc,
ihr cimei anil the r<»l of thn ncrvoun system are in »uch a condition l)uit
th« npplicniion of even a momentary ttimuliM to au am leads nut to a
•tniplt; movement, hut to a long -continued (ooic contraction of the appixi-
priaii* miMclea. Under Ihcec nr<-unift«ni-n a gentle stiroultiiF, fucU ns strok*
ing the skin or blowing on the face, applied immedintelynller the application
»f i1m< electric stimulus to the area, snddrnly cuis nhort the contraction, and
brings the muscles at onco to rest and normal Haccidiiy.
Tbcfc expcriractitfl abow that the development of the procc8>«s in the
conrs kiiiling to the mne of whut we have agreed to chII efferent impulses
along the pyramidal l)bn.-8 i« markedly nH<?cled by sensory impulses, and
specialty by sensory impulses starleii in liie tkin overlying anil correspnnding
to tliv mosaes put into movement. How those sensory impulses reach the
cortex we do not exactly know; but ne have no evidence In show Uiat
aflerent, centripetal impulses can travel backward, to to speak, along the
pyramidal fibres ; and it is more reasonable to suppocie thai the senior}' im-
IioIms in question reach the cortex by the ordinary paths of sensory impulses,
which we sliall presently discusa. We may therefore take tbe results of llie
ex(>enmenls as sliowing how close is the c^jnneciion of the motor area with
Uie sensory inechaDisDM of the spinal cord and lower parts of the brain, and
as illustraliDS the oooiplexity of the chain of cvenis by which tbe motor
area brinirs about voluntary uoTementa.
ji 663. We hii\ e alKive used the fenentl phrase " movenienia of the limb,"
since in the ilnji it h not ttwy In pick >Hit cerlain movements a* lieing pnrlicu*
Inrly skilled movoiin-nln. In the monkey such a divtinctiou i* easier. In this
auimal, aswe luiw mid, recovery of voluntary mvvetueot also ukrs place
I
830
TnS BRAIX.
nlier reoiuvul of a corticnl area, or at least lias dune so in many laisei ; aud
white tbe j>heuuui«uii iu)m«iliaiely lulluwiu)* removal uti iti« whulv twembl*
tbuw iriin<*ie<l in tbe dog, a certain order oC recovery may be observed : lii)
more akill«<l iiKiveiiieiiin are ibe btit u> return. When, fiir iuataaoe, tbe aru
»tt» in remuved. tlie delicate movemeiits of tbe hand, uf tbe tbuub uai
liu^r, are the lust to be reratablitilied : and u uuudiiioo of Ibin)^ nuy k
met with io wbidi tbe atiiiiml aAer removal, say ul' tlie arm are* id ilie Itlk
Itembpfaere, Uf«B by preference the lelt biuul at a time wbeu, if [mwaiiJ
tViMU ueiing tbut baud, he m able to ane llie ri^lit; tlmt U to !»y, the raouvgry
iu tbe riichl limb alter the removal of tbe area on tbo ivii aide it ituarly tna
nut (juiie coiupleie; tbe " will " can j^ain aooan l» tbe ri)(bl baud, but lui
ao caaily an to ilic Ml band, nud tbitt latter i« UNcd, tliou^li under ordinary
cireuiu*iauo» it would nut be uMcd.
When we turn to niiiii, in wliuni the great (luvclojtinvut of the iiyraraiiU
ayileui and dillereotiultnii nC tlie curiiciil ami is |i«irallelwl by tlio (irvnt-
notice of tkilJcd and trainol movements, thfl MiBlaffy of the i>UpnoDMU<>r
•pe«vh, if it be true. ii» cliiii<'iil bioiories accid to abow, that di«iructioa bf
tliaeaie of the *|ire(rh area of both «id«« causes [>«rniHne4)t aphakia, woulil
lead ua to oiuii'luile tliul n( IrJint highly ekilM vuluntAry moveoMnU ai*
carried uut by the pyrnuiidiil vysLi-m and by that alone. But in nfervnm (u
thia it must he romombervd iliut xiich a perranuMit aitliaeta may ba dtia. w*
to mere lou of the pynimidal chiiniivt. not to llie will b«ii>g merely nnaU*
to gain aoc«M to lower coiinlinatiug itii-cliauUinB, but lo the abMDce of tlw
diHJwtditiaicd cortical gray matter, by rnuon uf which absence the will cnno'it
initiate the fintt procvsMa «( the ncl of it|<eech ; it may Iw that were it abit
to do M), the pru(W»es to started niijiht, in I Ite nlwcncv of the pymtaJdal tntd,
lind some other way to the bullmr nu-chnnism, ns in ihc caau of the ittiskilM
moTements of the dog. This [loinl, however, clinical biuorieft have not
definitely tettted. Moreover, io dc^in^ with the phenomeoa of the nervoet
system uf oiau, as revealed by diAea^e, we meet in relereiice to tbe oanlnal
cortex the same dilHi.-uUy ilial we dwelt upon iu dealing with the dpinal eonl
(§o!f2). Lesions of the pyruuiidal sysiem. of the InlemaU-ajKiule for JustaiWR
leail ti) Ihe loiis not only of skilled, but of all volutilary movements ; aceort-
lag to ihe character and jMaitiuQ of the Imioti tbi> or that jMirt of tbe bod;
is wholly withdrawn from the iutlui^nc-e of ihe will. Ami it t* pOaaiUe Iv
inaintaiu the thetiid that muo lius l>eco(iie no <leveIop>:d ua to his nvrvoo* tjt-
turn and the motor oiirtex, so aci-ii>tomcd Io make tMe vxcUiiivcly of ik
pyramidal system, that the will haa l-nt ilie [H>w«r, mill |»iiuvwpi| by Iumt
iiniraals, to gain access by sunie paih ulbur than the pyniniitlnl om< to tlv
immediate nervous mechanism* iif movement. Tbe dnla liir loriuing a tMit
IkcUiry euuelusion as tu tbt.^ pi>iat are so few nod uni'crtaiu thnt it would b
nuprufitnble tg discuw the (giK^tion licre; hut we may v«ntnn; to ptiut iiui
that, griuit as is the devcluptiient of the cerebral cortex and th<? pyr^uii'lsl
aysteoi in man, that development is aocompaaicd by ■ hardly lew slrikis;
ezpauuoD of other parts of thu bnin not directly OKinoctisd with tbe pyra-
midal ty*tcui nhich we have previously seen rauoil to a««iic!ale with iba
ejiirdiniilion of iiiovumuntj). for ciiimpio ihe oerebelluin. Ami. indeed, it ii
clear that, niliuitting the pyramidnl tract to bo the onliiiary chsuoel by
which volitional impui»c« ftnta lo, or by which the will gains accees tn, ili<
motor mwhanisiiiK immcdiiiU-ly associated with the anterior tuols of tJiU «
that »pinnl nerve, we must also admit tlial ibom volilioiinl impulsed paaiiaj
along the pyrminidai tract, or at leiist iome of tbe prooeatea ooasiiiutiuj; tlw
will, are in connection with, and thus are influeooed by the oonditiun i>f.
olher parts of the braiu. When, fur instance, a gymnast executos a sklUeil
VuluoUiry uioveuieul iu which all bis four liuibs and other jHirta as well pnr-
ON" VOLl'KTARy HOVKUENTS.
881
I
I
I
liialxiJj' nri- involvol. it u pnilMtlily tlw nM llmt chnngn of tin
iMtiirv or i-HVrtiit iMi|HiU>« fvitvp ilowii hii pyntniiilnl inid, and timt tiMH
j»i|HilM-a. Hlurtiiig ill o (U-liiiiUi unJcr from hit corUix, ihiil in Id my. having
uitik-r>,'<iiiv a n-niiiii uuKiuiit of initial outiiinnlion m lli^ir very nngiii. iiR-vt
Mitli liirthur oonJiiikiioB in tbttpiiial grtiy Rivtlvr. wbii-li Mrvn u n m4 uf
iturlui ul' (irigii) tur ili« inolor nervM conoerned in ilie ntovcnwot botbrv Ihvy
iMtiu BR oniiiiary tuotur impulses aloii); the niuerii)r nxita. lint tlii;' t« nnt
ftU. Sbuuhl ihe ;;yniiiiu>l'H seoiicircular ctintilH happen tu be injurvl anil hi<
cerebeiliim tbereby bo Iroutiletl, or mischief fall on some oth«r part uf lh«
drain wliioh like ibia baa no direct cunDeciioa witb cither Ihe pyramidal
Inct or tii« luolur curlpx, Ihe movemenl foils thrungb lack of coordinnlion,
thuagh bulh the cortex, the pyramidal tract, and the spinal motor mechan-
iaraa ranaio as tbey uere belure. Obviously, the carryiuK out of a voluntary
muvement b a very complex proceedin;;, and the muiiir ciirlex with llw pfrft-
loidiil tract is only one part i>f the whole meclmuiam ; k> tar from the whol«
bu>-iitea4 beiujf L-»nliiu.-d lo tliefie, it is perliajM no esa^^raiiuD to say lluil in
eai.-)i iiiuv<^u(i:iit uf the kirxt mu^t paria of the whole hrsin bave a greater or
lea* abare.
The exact luUure of the part playe^I br t)ie cortex and the pyramidal tract
iu voltiuDiry moveiDcoia uiir present ko<iwle<l;,'f n iniKk'i^iinte to detini!.
Wh'ii We iMua in review a wriett uf braiua Irom llie luuur Iu tite hi};her and
•cf bow ibe pyruiiitilal nyiil<.-iii is, »i> to npenk, grafUd un to Uie rtst uf the
Itmiti. wWn nf ob-i^rve huit the iiit-rea*iQg ditferentiatioD of the motor cur-
t«X niiu parallel In the iucrciuiog pOMemon uf skillctl educated muV6m«Dl»,
wc way ]ierha|« flUppnet.- lliiil '* A alrart cut " fnnu thi; ourles to the urtfjpria
of tJw aeveral mulur ui^rvcs. vucli hh in aHunlcd by the pynirnidal tibn-s, Irutn
tbo advantagea it uiren t» lh« umrc primitive [wth fr^ni i>e}(dieut tu M^ncnieni
aluo;; iIm CVNbro-spinal axi« baa by natural selcctiun bcien developed intu
bcin^ in man the cbivf and trn^t important instrument for carrying out
voluiKury movements ; but. we rcjicnt. il remains even in itn higheat dsvolop*
laent a link in a chnin, and a knuKledga of buw th« vhola chain worka ia at
preeviil bidden fi\jm ua.
We must Dot here wander into psychological prob!e>Ri«, but may ropnt
that in the above diocusiion we have used the word " will " iu a genural acoM
unlj. A man may be broufiht into a condition, for instance in certain byp>
nolle phasee, in which he can carry out all the various skilled niuvomcuU
which Ite has inherited or which be bu tearneil ; and yet, ncording tu some
(letiuitions of the word " will," thuM ntOVeiDeals could not lie said to bo initi-
ated by his will. It can hanlly be doubted that in sucb cases the motor
cortex and pyramidal Itact play their usual part. lint we may pass from
>uoh caaea aa these ihrou^'h others, uniil we come to casei where a skilled
niuTemeut which baa t>een learnetl and praciiwd by the wurkinj; of an iaielU-
^leul will, may continue lo be carrit'd out under cirLntoutances ifhich seem to
preclude the intervention of any ounscious will ut all; and the tramitiun
Irom tme case to nuoiher is so i;rad<jal. that it is impiuaible to (uppase that
there ha* li««n auv sbifiinjc uf tlte macliiuery employed fur cnrrying out tite
noTeuenl. 8u tnat a volitional origiu b not an Mtcniial feature uf tli«M
•o-called vuluoiary ninvemenu, and the raaohinery of the motor corl«x and
pyrsmtdal trad is available furiHher ihinj^ than pure voliltonal inpukaea.
I 684. The ))recedia;; dbicuation will enable us to be very brief cuncerning
a queatiaii which hai fruiu time U> linte bcvn much disciuMvil, iinil which hat
acquired |ierbii|H ftuiiiiou* im porta ix-e, vii., llw qucaiion n» la how voltiiunal
impuinn* leadm),' Iu volnntitry niovrmcnta iravvl along iho i^piiud cord. Tlie
oonclUMon at nhicb we linve arrived, nxmi^lr, ihai in ihe normal carrying
out of voluntary movements tliv chief put ia jilayvd by sdennt imputsus
883
TH« BBAIK.
(mning kIod); the pyramidal Irart, carries with it (lie ftoener tint Tolilionl
ioipulaes travel id Ihe spioal cord aluiif; tlw pirraini<lal tract.
In ibe dot;. ^° which tbe nholc pyramidal tract croeses al the <leoaMBiiia
of the pjrKiuida, we should ex{>evi lo 6nd that a br«ak id the p^TMiudil
tract of uDe aide of tbe cord al an^r poim aiun^' ita leotflli cauaM lo* tl
voluntary movement on the saiae «<le below Ibe level ul' the bremlc. Aod
ex|)eriu)eul8 as far as they go support Ihia view. Ho one it ia true bti
attempted to divide or otlierwtee cause a break in the pyramidal tract ■loar.
leaving; ibe real ul' the curd intact; and, indeed, even if an injury wtit
limited to the area marked out as tbe pyruaiidal tract, fibres other tkaa
pynimidal Hbrnt would be injured al the Mme lime, unce Uie tract b never
u " port- " one. But it hns been found thiii ■ secitun uf u lateral half of llw
cord, a luieral bcm'iseotion, or a teciion limited to the lateral eolumn of ew
aide baa for one of ita priDdjial ed'ecta Ium of roluntary raoTenieut on Hit
aame tide in tlie parU tupplied bv motor nerve* leaving the cord beluw t
level of tbe set-tiuu. Wc my " one of it* ]>Tindp(il eBcoU" because, \r^'
the coucomitaiit inicrrereticv with MiiitnlioDS coucerDtDs wbidi we shall i _
jirtvciitly, the Ium uf voluntary ■aovcnient i* not nbsututely confined lo ibt
same side ; tbera is some lo« oi |>ower on the croand side, at least in a large
number of casct. Wo mutt not lay alrea on thia croMe<) parnlyvia bcotim
it is probably one of the cibcta of ibo nitre operation, not a iwre "Ml-
ciency " phenomenon, and, indeed, appcan soon to poaa aw«y. But laking
into consideration what was nid ahovi; conoorning Ihe cflVvts of rentiirlu
conical arcuK, it is imjKtrtnnt to note thai in ihc experi*nce of many rxpov
lucuK-n th<: low of voluniiiry |H>w<ir on tbe opomled lido dimloialies alter a
while, nml Ihtit the niiimni if kept alive and In good htalth long enouli
appeara to regain nlRioel full voluiilnry power over tbo aflcctcd p«rt& In
such oaaee, ss m other operaliona on thu i;eDtral nervous •ystem, there it aa
regeueratiuu of nervoua tissue ; the two surfaces of ibe section unite by «»
■Mclive uoi nervous tissue, and tbe Iracts which as tbe result of tbe ie«lica
degenerate downward or upward arc permaueully lost, lleooe even if «c
admit that in the inlact animal a voluuiary tiiuvement is chieHy carried enl
by means of eHerenl impulses passio); along tbe pyramidal trail riKht don
to the motor niechaDisms of the conl inimediately conuccii^ with tlie moMr
uervea, we must also admit that the " will " under cbau(,tMl drnitnstauctt
can find other channels for gainin)c acoew to the same meehaniaais,
II lias been further obferve<l that if in the dog a bemiaeciion tw made u
one level, for instance in the loner ihorncic re);ioD of ibe cord, and thai,
alter wailing until tbe voluntary power over tbe hind-limb of that side hat
returned, a secuud bemiseviion. this lime on the other aide, lie made at a
higher lavcl, this second ogHTuLinn is followed by resolla similar lo ihowof
the 6nit ; there is loni of v»lijntnry jxiwcr on the *i<le o[ierate«l oo, with Mm
lota of jiowLT i>n lh(! crcKuml sidr, niid as in the lint case thia lata of power
not odily mi the saniu but also i.m Ihc cro«H.il ride ntay eveiilunlly dtmpptar.
This kluiws among other things thnl the recovery after tlic lint u|ieration wai
not duR to the n^nmining pyramidal tract doing tbe work of both, Purtlm,
the hcmi-ii-cttou may he repcaierl ii third tiuK-, the third Itcmisectioo bs^ng OD
Ihe same side as thv fint, with at least very uoiisitlvrablc retnru of ponr
over htilh limbs. That is to say, under auch abnornml circumstancce volun-
tary impuUrs may, so lo speak, thread their way in a ligiag manner fntBi
side to nilc along the mutilated cord until ihey reach tbo appropriate spioa]
motor mechanisms, ^^uch an abnormal nale of things docs nut. hovever,
really militate against tlie view that under normal circumstaocM rolitioaal
impuJsGt normally travel along the pyramidal tract ; but it does show, what,
indeed, has already been shown by tiie phenomena of strychnine poisoning.
flH VOLL'.NTAUY MUVIIUKNTD.
S8S
»
I
S 5S7, Uuu in llifi central norvoiis ■yKtfRi tlic [taMii|;i; of nervoiH inipulnM
luttOfT Ume nords in the p?nGml fwnac if chaiigv* p^rlpng1ll■^l nlong Dervuu*
inal«fialj ia not rigidiv and unsll«rably lixpH l>y thp ntiatomiml dislrilmtion
uf traoU of fibrw: in ill such drK'iiMiniti a* ihow in whicb we are etijfnMd
we mutt bear in luind tliat pbyttiuli'tfitnl (i>iidili'>nH w well as aiuatiiinical
eootinuilf ar« puient in deli>rmitiiii>; tbe [inatiige i>r itieie impuUiw.
$ 665. Wlieu we reflect on the fcreai prominunwuf the pyrumidal irm-i in
tht' !i|>innl ronl nl man aa compared with that uf ibe du^. "'^ nuir jii>lly inter
not only ihni tbe t'Vruniidal iraet is nnder normal eiri-'u instance* more exclu-
tivoly ilif. I'hanni^l id' volitioniil iiupiil&ca in man ihan in audi lower aniinitk.
but iiliMi, iM-iirin^ in mind (he dinL-iiMJoo in n previiuii eliuptcr, $ 51*2, con-
oeming tlic uctivitiM of Uic tipiaal oml of niao, thni the )K>uuitinl iiltcrna-
tivM pmMiml by iIm xpinnl cord of ih^ do); are (crvally reduced in that of
DiKO. And »uch clinical lunlorM of iliaeaite nr avetdeiitat iojnry in man as
W« iMMMMMipporl Ihitt cfinclintinu. Lenions i-oulincil to i>ne-hajrr>l' tbe cort),
or erini Inton* confined to tbc Inicrvl column of one -hair, a[tmutr to lead to
IcMi of viiluntnrr piiwt'r on the «nme ilile, and rlut irainc tide only, in the parts
below thi' Icvrl of the lr»iim ; ami the naint- Hvinpfxn* bare been observed to
accompany diaeaie limited apparently Ui the pyrnmidnl Iriicl of one side.
Mureorer, thougb com* of recovery- <if power have beat recorde<), ne iiare
aot Mob Mtisfactory cvi<teiK« as in aniinalx of the volitional impulses ulti-
mately making ihcir way along an alt>Tnaiiv<' route; hut here tbe sBnie
doubu may be euterlaint^d as were esprcswd in disL-nMiiiK the reflex acts of
the ronj in man.
When we nay that the Inm of Tolnntary power is seen on the aide of tbe
le«i><n oiiily, we ahotdd add that ibis statoineni a]>poara to apply chiefly to tbc
ihotacic and lower {mtis of tbe cord. We hare seen thai in man, in tbe
iiiijicr r«f[iofls of tbe card, the pyramidal tract is oolr jMUtly cniwexl ; a vari-
oolc but not inconndernhlL- uuml>ur of the ]tyrumiuul (ibn« do not cros> at
ihe decnesation of pyrami(K but running strwifrht down n* the direct pyra-
nidAl trad uAvct tbcir croiwin^ Inwt^r down In tne Cervical and up|)er tlmra-
dongMMW. Hence, weabould infer that a heraineetioo of, or a lesion conlioed
CO one side of the cervical cord, wniihl allect the voluntary DiovementA of the
gfoawd »ido as well us of the eanic mite, though not to tbe some extent. But
we have no cxncl information w to tbi» point. And, indeed, the pur|>oeeof
the iliriTt trad i# not clcnr; there is do mlei)iiato evidence for the view which
has lieen held that thc«e direct tihre» are dc«tine<) for the upper linihs nnd
upper fwri of the body ; since they are the last to cmas we should, a priori,
be inclined to suppose that they «ere distributed lu lower ratlier than tiigtwr
pans.
i 666. We KUiy now briefly Huinmariie what we know cooiieniiDz volun>
tarr inoremente. Awl it will be convenient to trace ibe events in ortler
backwanl.
Certain muscles arc thrown into a ooniraelitju which even in the krioAst
movemeuta is probably of tl>e nature of a tetanut. In aliuu't every inove-
meni wore thao one muitcle iu> defined by tl»e analotubu i> cngnged, and in
many mnvem«Di« a ftart of several muscle is empli>ye>l. and not tbe whole
of each. It i* perhafw partly owing t» the latter fad that a mtuclv which
hM iMOOtne tired in one kind of movcnieni, may show little or no Intigue
whw unptoyctl for another niovcmeol, though we must boar in miiul that in
m volnotary nxiventeni fatigue it much Biore of nervous than of niitscular
ar%in.
Boidai iba active niusalea, if we uuiy so call them, which directly carry
out the moveineiil, tho metabolism of which lupplios the energy- given -hiI as
work dune, uthcr musclM, •ome of which arc antagonistic to tlic active mu*-
U
TBB BltAt<7.
cles anil mmi.' nf uhicb tniiy W #)M>kcii of lu sdjuvaat. caler inUi tbr wlial*
act. Jo fl«sii)ti for iiutancu of thp fomrm ou lh« ami il is itot thf Boor
muBcles only but lh« cxienmra also nbich arc eDpiK«<l. Acconliuic tu tlir
inini«<l lately prccwlitiK poeilion and use of the arm, aixi according u tbt
kind and auiouni of flexion which is bi be tarried out, tJw exteoacm viU bt
either relaxed, ihat is to say, inhibited, or thruwD into a certain amciant al
vtmtrnvtiou. And in some of the more ooraplicnied voluut«ry niovvin*rm*
the part played hy adjuvant muaclea is oousiderabk. Henoe, in a voIudMi;
nunrem«ut tlje tvUl btis to uuin acee«» not uuly tu lli« a4:tivi; niuitclc*, but alia
tu the unttiifonUtic aud aiguvaal muscles; and uvcrv vnhintnry movcraiiit.
tvea one of the iiiiupleot kind, is a more or Icm ciinipli'x ain.
Tlie imi)ul«i9 nhicli lead l» tlie cuiitrautloa of the active niUMcUti r<.-nel) lb*
miiscl** along the fibreu of tlie auloriur ruoti (vie may for ihv snki- of um-
]ili<-ity uke ftpioal nervea alone, neglecting ihv pci^tilinr rranial n«rv«s). asii
lucb evidence as we )iaasen goes to aliuw llial the ira]>ul«ea governing the
antagonintic and adjuvant nniitdeii tmrel hy the aoterior roots mlao; the
question wb«>lh«r the iidiibiliun of the antagonistic ninsclca when il lak«
place, is carried out hy inhiliilnry iupulsca paning as #ucli alon^ the fibre*,
or eimjily by cuotml mhibiiion ^f jiravioiisly existing niot^ir inipuls«s need
not be conei'lcrcd now. Thntc luilerinr niots are connected, an we have smb.
with tlie gray matter of Ihu cord, and in each hypothetical segnieut of the
cord ire mav recopisc the existence of nn area of eray matter whicbt thoa^
ue coQuoc define lU limits, tve mny, led by the aouo)^ of the omnia) uervca.
call the niicWti of the nerve belonging to the aegineut ; and we luay further
recognixe in such a nucleus what wo may call its efferent and 'Ha uflervnl Mde.
Kvery voluntary niovoment, even the sinipleet. U, as »'i- have reneatnUy
ioriitted, a L-o'inlinaied movement, and in it^ courdinatiou atfcri.'nit itiipuhu
play an tmpuriant pan. The study of retlex nctioiu, § 590, haf led us l»
Buppoec thut i-Hcb 9)>inal oegtnent presriiln a uervout mechaiiisni in whlcJi >
certJiin nniounl nf coiirdinMiou in aln-nily pn-veni, in which «flhrent impubs
are ndjiittU'd in aSiirent iinpiihnai. But the mulls obtained by atirauullac
iteparale stitfrior nervf rootn itbow that, in the case of moat rotneleB al aii
events, the cfnin'inlly active niuncloi of tbo limb* for iitttAnic. each nmsckh
supplied by tiling coming' from more than (MW nerve root, that is to aay, iht
»ptnal nu<:leus. or nt I'uiHt llic npinal motor mechanigni for anv one tnnsds,
extends over two or three segments. Honce t /oriiori in a vofuntary mon-
ment, involving as this does in most case* more than one miude, the opial
Inechanism engnevd in the act spreads over nt least two or three sef^nHMs,
thus allowing of incratsed coordination. In that coOrdinaiiuu the tmiMho
serving as tne foundation of muscular sense piny an imjairtant (Ntrt, biH
other affbrcni impulses. such as tbuse from the adjoining skin, ulsu have thtir
■liarein the matter; and it is worthy of nolice that not only ia the skin ant-
lying a muscle served, broadly Bpcaking, by nerve roots of the same legmait
as tite rouBcle itself, aflferenl in one cafe, eflerent iu the other, but in tin- part*
of the body where coordination is especially complex, in the liagi-r« dir
(nslanoe, not only is each muscle supplied from more than i>w siytufnt, luit
also each piece of skin is supplied in tlie »iue way by the jMnicrinr nmu u(
more tban one nerve.
In the case of ttie fVog it b clear that iu reflex movemenin a larp.- amnuai
of coordination is carried out by thaw \'arious spinal meeluintsnu: aud as »v
have ur^t^, ne may safely infer that iu tlie voluntary movements of the ftnf,
ibe nill ni»k(« use of thU olrewly exietinK ciHiniinatioo, whatever be lb*
exact |iath hy which in thi.4 animal tlie will gaiii* aoc<a« to the spinal iDwhan-
iama. In llic 'Iok we may conclude ihnt in voluntary movraienl^ the sjiiaal
jBKhanwn*, with ojurdinating faocliunt, are also set in action, iu this < ~
ON robUKTAftV UOrBMKNTS.
835
I
I
I
/ImpiiUes ptunio;; !imi;;fai Trora Ui« cartel to ihe nieclniRuiiM lir ilio |iyrn-
luiilnl tract, tlioti]{h a|ipiir<-utly. in tli« absence oT the pvrumiil.il tract, I he
will am work uvum the oii-clianuiiM hv changes travelling thniu|ch other
pttru of the cereoru-4i|Huat tixiit. Ad<I in the ntitiiltej- aiiil laati, yubjwt to
ih« doubu alreuily vzpnMed aa to the iioii^iiLiiilititi* i>( tho human spinal
cord, mt may {imlKihly abo infer thni in mnh vuluiiinry marcmenl KOmo.
parhii|H <rv luiir Mty much, of th« cminliiinliim u currivd out by tho ef/iiinl
■DMhauitim wl iiilu nctioii through impultw nlnng the pyramidal tract. W«
may probublv further inror that a careful niljugluionl obtains botweeo the
baiginDinK» »i (h« prnimidal tract in the cortex and it« i-ndiniri in ih« conj,
Hj that tha topography of "nrca«"uf" foci "in thr cortex above ic an imagv
or prajvotion of the ttpinal mcchaniama bolovr.
Tba aompJaz eharact«r. on which vrc iniiated jiMt now, of alnioel eveiy
Toluntary mov(Mn(>nt ncccssilaica that in every such movememt a Urge area
of iipinal mechanism is involved. Bnt this L» not all. The muTenieuta of
•Djr pom of th« leigs for initanoc. niv not delermioed, nor is the oo&rdiuation
of the movements aflected, simply by what la i;oiiig on in the l«gs and the
part of the spinal cord belongin); tu theiu. The diacuiMioa iu a previous
Mctioa hoa sbown that much of the courdtnacioa uf the body is carried out
b/ the middle p'lrtions of th« brAiD, and oa theee the mo^r area mii«t have
tu hold an well as on the spinal mechantsnis.
The detail* of the nature of that hold are at preaeat uoknoiru In lu: but
it muKt be remembered tliat not all tht- libr^ uasnog down from th« motor
reciod. not all thoae even proceediui; from tiiu (U-uit<Mt nud mo<l uleurly
dehoeii motur areas, are prramidal tihre*. Witli the pyramidal Qbrej arv
minuted Aitr^ having other dcali nations, aud aoue of those probably pium
to toe thalamus and so join the Ertat Mgmeatal ref^on. Moreover, tb«
faotof region muot liave close liee with otlwr rcgiona of the ooricx wh«nM, as
wo hava saen, ^ ^'i'i. dbnt nasi to tho pons to make connections with the
eervbvllum. On the other band, as we baro Men, $ 113, tho corcbellum ts
vspccially conrtccuvl with what wc may fairly consider tlic aflorent >!!d« of
this e|>inal cord nnd bulb. Tli«a« facte must mereljr be lakeu as indicating;
ihe powbititiee by which th<r motor region is kept in toucli with the great
onordiDadog mecoanism ; it would be venturesome at present to say much
mora.
In an ordinary voluntary movement an intelUjceoi coaaciousaeas is an
eaiential elemeoL But many skilled movemenbt initiated and repented by
help of an intelligent conscious volition mar, when the uervons machinery
for earrying them out has acquired a oert^n Gicility i,niid in all the higher
prooeisCB 01 the brain ire must reeogniie that, iu nervoud losterial at all
events, action detenntnes structure, miauiing by structure moleoular arrao^-
ineut and dts|HMition) be carried out under appn>priat« circumWancw witli
ao little iut«rventtoii of distinct oonscioutnaai tliat the moveinesits are then
often >pokcn of an involuntary. All the argtiRientx which go to show that
the dtttinctly euuscious voluntary skilled movement U carrieil out by help
of the appropriate motor area, fpt tii *liow tliat the motor area oiust play its
part in thwe involuntary ^killetl movements also. So that diftind consaous-
nea* is nut a neoesaary adjunct to the activity of a motor area. And it Is
worthy of nolioe that aoaie of these, in their origin, purely voluntary' skilled
muTsmMits, which by lone-cimiinued training have Decome alauMt as purely
involuDtary, am hamfKrcit rullu-r ihAU uvintcd bv being "thooght aliout."
The wonl "training" »tiggw>ta the retlcclion that the phyaiologieal intor-
pretatiou of bocomiiu easy by practice is that nc<r jutbs are nuule. or tho
iRateria] of old [Mtb* tnads mors mobile by effort and uec. We liave already
1, 1 582, that the gray autter of Ihe spinal cord is a network, in whioli
83S
TUR BBAIS.
l(i« iwsaigo of impiilwn w il«ti»tniD«l b^ phynokwlfitl OKiililioat ntJwf than
BRBtoiDiciil coiiiiiiui(y. mtd lh« mnio coomaenUKim ribv oiih «ttll grauw
Turce be np[>)tc<l In thV brain. W« inual Hipp>iM> thai training proinatc* iIm*
groKth and mQlecular mnbitity of thi' motor areii and of nil ita vonDvilioU)
1 herv nre doubtlas limits to tbe cliangcs nhicb am he elTcctetl, but wilbn
tlwae )iinil« tbe will, blundering nl Rrst in the niaice of llie nervous network,
gradually eetabli?h«: ea^y {nibs; ihixt^li even to tbe end it blunders, in trt-
■nr to csrry out one movetneni it olieu aro'tnpliHlies anoitier.
ijutly, witbiiui nlieiiipiioK to enter iuu- iKViholnffknl (juciilioo>,we Diaval
K-Mt uiy that the birthplace of what ire call tbe " will," ia not enotemiaa**
with ibo motor area; the will arises ftom a owtnplox «eriea of «Teau,»iDenf
which ittke ptave iu mlier rewioria of the oonex, and pmhably in oiber parti
of the brain as well. With Ihese |Min« ibc motor arcji hax l\«* ■■nncrrneil un|
in tbe carrying ■>■![ of volition, but in the gwemtion uf ibi^ will. So ibii,
looking round on all sidts, it >« (tltviouc, m uc have aid. that ihr imtor am
is a nn-tv link in a complex ihnin. It », moreover, a link uf nich a klid,
that while iliv changn which the breaking of it makn in the daily life of %
lowly animal, <uch a« the dog. in whom tlw expi'rieon-of the individual otMl
relatively little In ihe tM.'n,'ouv and p>ychica1 »i"rehouEe iraitcmilled from hii
ancertors, can hardlv b« a|>prt.-ciatcd by a bvMaitiier, tbosp which tbe breai-
iot; of it make* in the daily life of a man, whu«e brain at any ninmenl ii not
only a machine fitted for present and Aitnre work bni a <'li«ely pucke<) reeorl
of bis pBBt life, are obvious not only to tbe individual himself, but lo hit
felloWB.
Os THK UEVBt-OPMKST WITHIN THK CkXTRAI. NeRTOIS SrSTKlt Of
Vli>UAI. A!iII»U»'!^>M>: iiTllltK 8ut«ATIO!CH
ritual Stntntim*.
^ 087. In the chain of events through which some influence broi)f>ht to bar
»u the periphery of a senfory nerve gives ri»e 1« a senealion, we are abb,
with uore or len success, to disliuguifib betweeJi those eveuts which are ilettr-
niined by ibe changee at ilie periphery and ibooe which are die cxpfoaica
of cItanKes induced in ihe central nen~»us system. Tbns when rertain njt
of light proci'e<liug from au object and iBlliiiji; upon the eye give rt>c in viraal
perceptiouuf the object, two aeUnfeveiiis hii{i|icu: ibe rays of light, by Mf
of tbe mecluuiisma of the eye. partly dinpirK-. partlr ncrvMis, give riw tu
wnnia chaiiM^ in the tihrcH uf the optic nerve, which we may mil vtraal
iro]>iit«ni: and lht«c vtMial ini{HilM« reaching tlic brain along tlie oMic iierft
give rim- to vtMial n-imtion.'' and m> to visual pcrcc|iti<>ii of the ohiecl. Vft
■hnll tnl<-r on, under tlie beuilingof "the *en«n," deal chiefly wilh the [leripb-
enil evenln. and havv now to ci'iiiiii1<<r »ime point* conwcled nilh the central
eicnw. lo learn what »i- know coiici-ming how the variou* K-n»'<ry iinjiuW
li.i veiling along the several kimlsof feiisory n<in'<* behave wilbin ihe central
nervou.< system. In doing so we shall have from lima to time to refer Id
peripheral events, but onh' occasionally, and never in any great detail. It
will be i-onrenieni to begin with the spocial sense of Rght.' and we must 6nl
briefly call allcniion to a few points which w« shall have to study in fuller
detail herealler.
Tbe eye is so constructed that image» of exicnial objeds are brougfat u> a
focns on the retina, the stimulation of which by light starts liie visual ira-
ilsea along the fibres of the i}|>tic nerve; and tbe distinctneai with abieb.
ty iDcaoB of the visual seii^iations arising out of these visual impalm, irv
vrSFAL Ayi> OTRKR StySATIOMS.
I
I
prrwirc extenwl objocU Ja ilepeodcut ou ihe ithnrpnMs of ihe retinal imogee.
Tlie ere is further so c(in>^ruc[etl that, in auy (Wftition of thir eye, the rsya of
lilfht (irooeediiiK ttom a portion only of the pxionml wiirlil fall upon tli«
rMtna; or in otlier wordi. in any one pasition of the eye only a {Mrtioii of
[b« external world ia visible at the same time. The |Mrtion to teen is tpokea
uf an Ihe runuif field for tluil pueilioo.
The iniafce tJ)n>wu od Ihe retina la au inverted utie, so that the top of an
HcKnil obJH't i.-< r«prf!<eDtM) by the lower, and the bouom by the upfier. |)art
of ibe tetiiuil iniflge ; similarly the a«ttud lefl^haud lide of thv reliiial ims)^
r(>rrTH|Mi(KU to the rijtbl -baud *i<le of the artoal obiM.-i. and the rigbl-liaud
niilo tu th« len-hanit side. Hence llie rigbt-liand half of the rinual li<-Id
cornwiMnfU li> tltv Ifd-haiid »i(le of (he retina, nnd ibe left-hand half to the
rigfat-liand *id<-.
The ■■vc can be movcl in vnrioun itiructioDv, and diner in ibc vimiuI field
thr (Mitlioii of cxlvmal nature n hich aio be wen ai the Aine lime dilfem
with i^ch dillirri'iit |H»iii<>n. ii large range of virion i* tliu* iRMured ; nii'l ihiH
CTUi Ik further iiit'reii»^'d by nioveroonUt of the l»-iid. Moreover, wc iic.r-
mally tankti u*o of twi> vy««: our normal riwon i* binocular, and lliv visual
(i«KI of tW ri|;ht pyi< diflvrs from tJial of the left ere. There n one »trikiiiff
diHcrcDcv u'hicb must always be br>nie in mind. A H-clion earned ihrougn
the eye in a vertical iin<l front-tu-back [ilaDe. through what ire tihall learn
to call Ihe optic axis (Fi|^. 'J1 1. 1>. r.) (the exact details of the plane nuiy be
left for tbe present^ will divide the rcltna into two lateral hsKes, an<l in
each rvlina one-half will be on the uaaal sitle ucxi to the noiie. and the other
half will lie on the malar or tenipornl side, next to the check or temple. It
roust be retuenibervd that (he na«il halves and lemporal halves of the tiro
retinas i|i> not occupy currM|)oodiiiK poeitiona in apace. The teniiuiral half
of tbe leA retina is »n the left tide of ila own eye, wherenja tl>e teni|M>ml half
uo tbo rieht retina \» not on the left, hut »ii the righl. side of iiM nye: and
M witb ine nami halv'v. Now, in lh« ri)tbt rye, (Ik- ri);hl-hniid niile of the
THual hi-ld oimsponiU lo the- nit--><l half of (lie. r4-ti»:i, and the Ict^-hnml vide
of tlw visual Held to tht tem|H>ral half of thr retina, wh<-rea» in (be left eye
ibe right-liand nide of the visual lichl «irr«!«|M'ndit (n the temporal half of
the mina, and tli<> left-hand niiie lo the niual half. This ia snovn in Pig.
'it I, where ihv left-haiul vtMial Hold ami the retina] area conocrnMl are shown
sbaiied in mch eye.
Wben we look at an object with iho two eyes, thoufjh two retinal images
are |>roducc<l, otM> in i>no eye and one in the oilier, we perceive one object
only, Uut two. This is the eswntial fact of binocular vision: wben certain
parte of eaeb retina are slimntaied at tW same lime ue are cviusejouS of one
MOMliou onlv, nut two ; and tbe parte of the two r«(i»as which, MiniulaiH
■t the same lime, f{ive the lo one senaation are apoken of as " rorrts|M>ndirig
parts." Froto ihc Airucture and relations of the two eveb It follows that ihv
temporal ud« of tin? rijfbt, awl Ihe nasal side of the left, ejre are such cor-
reapondini: tairts, while ihe narMtl aide uf tli« ri)cht eye corrw]M>ndM lo ibe
iem|Minil nidi? of the left eye. Itut the whole of each retina is not employed
in biniM'ular vtMon. Owin^' to ib<:! ihnition of the two eyes In relation to (he
noec, it come* ah<-)ut that un objccl field very much on one «ido, to the (o(t-
haoil aide for iniHaocc, while it is aipable of pruduc4ng an iiua^ on tJ>e
•xtmne aami nde of tlw left eye, and can be M«n, ttwrnforv, by Ihal eye.
cannot prodtice an Image on thv t«ni|ioral aide of the right eyo': the nose
blocks tlie way. It i>. tborefure, not eeeii br tbo right eye, and tbe vision
of it is monocular br tbe left eyo onlv. In Fig. 'Jll it may Im- seen that the
left visual tielil iif ine left ere < L. /'. L. ) extends more Ut the left, and is
larger than the left visual licjd of the right eye {L, K Jt.), and that the right
riaUAI, AND OTHKR SENSATIONS.
retinal nrvm. cnrnwpondiii^ lo iImk left risual field, extends furlhrr nloi)^ Uio
nasitl aide of tin' lirlt tide (o'l than It do« bIdii;; the tcmpciral sido of tbo
right ryti (a'>, thp diftvretiee beinR due lo the presence of the non {F.\.
I And MiniUr conditiooa obtaiti vitb regard to the extreme rij;bt-hand aide i>r
' the viMUftl Held.
h i 868. Xher ihcae preliminary stateiuenta. ire may now turn to consider
■MRHv anaioRiicfll facts eoiii'eniing thi.' ending of ihe optic nerve in t)ie
fbniiD.
; The optic aerve of each eye oouaiUa of nerve fibres comiDg from all paHa
; of tha retiaa of iliat eye: hut tlio two optic nervt!! iiioeL, ventral to tlie tloor
I of the itiird v-eulriclv, <-ruM each other lit thi; opiie rhiiumut i Fig. 'Jll. Op.
/Jr.\ sod are tlieocv ountiDued on umk-r the name uol of onlii; nerve*, hut
I of itfjiF <r»fU ( <tp. T.I. The decuwtutimi of fibre* which taKu.-< place iu the
cliiaaaM luu pet-uliar rliiinu'ten. Al their deciimlion (wc are spcukiug
DOW of man l the flbriat in tlio optic iicrvv Imloiiging tn tlie tenipoml half ol
tiie eye iu which the ncrvu ends paM into one optic tract, itiimrly, the uptic
trmet of tlie Mine viilc, while the fibrts beloDgioe to the nanl half ww ii)l»
another optic trmct, namely, the optic tract of trie opposite sidir. Thiiii the
Rhm of ihe tenipoml half of the right eye and of tne oaaal half of ttic \t(t
Cft pWH int'i the right optic tract, and the 6bre» of the nni|nl half of the
right eye and of the temporal half of the led eye pass into the \«fl optic
tta«t. Compare Fig. 'J11, in which the tibm forming the right optic triict
MC ihaded. white thiMo forming the letl optic tract are lelt nnshadcd. Non
tbv OMbI half of one retina and the temporal half of the other retina are
" cocTwpond i iig " parts. Hence, while each optic trad contains fibres be-
loo^ng to half of eai.'b eye, the two halves thuii re j) resented in each tract
are corresponding halves.
The amount and character of the decitiwation tukin)i; plai-e iu the opllc
chiuaia difiitRt in dificnuit aninuil lvp<«. ihc di<r(:rrnci.-ii having rvlation to
the amount of biixictilar viiiion, which in (urn de[>i-ndgi on tho pnoitioo of
the eye* in the head, that i«, on the prominence of the fiioe between the
evM. In the liah, for in«tanc<:, with hilcTnlly plni^od rym. n<> binocular
virion at all in pari ble. and the dccuwntion in complete-, the wholu optic
nerve id' each eye cronei over to the other optic tract. Betwwii this and
tha amngcnwnt in ninn, juat described, vanoiia stage; obtnin in various
aniomU.
The cJiiasma nlao eonlains nt it# hinder part fibres wbicli have no <-od-
nection with (be optic nerves or (ho eyes, but are limply comintwurel tracta
IWMing from one side of the brain, nantely, from the luealian corpiii geniuu-
atum ($ 631 ) along one optic (ract, through the ehiaema to the other optic
tract, and b» to the iiietlian cor|Mia geniciilatiim of the otlier side of (be
tM*ia«l>lil«nf UiartUravr (ber1sblCTD>adilien«Mltfilaaf llioi«Uniiof tae kA oyv: U K L
■ndr r. ff. Ik* trfl <l»a) IMdiirf lln Ml ■»« M|M pjw t«pntl**lV; ibo Ivo AtUiakl tk«|«ft>
e4 lb* ■••» nUniwwIinc «M*UUIiia ffaAara ttoloii mwt llM BtMi an thwlcO. Uwo*jMtala tlw
aiUdf Itilrtstil 4M« CI«1li( iVa lukniiiMC* Mn'. ■uit noitUMlatt iMdui laaKsal ■■'. Tlie ni|h(
mfUie limet h i»pr— aloJ ■• (lattiiK lii 'i I,, ilic l*l«i»l iinrtiii* VMkl«iitol*pi -. Ill Jv. ibo taliimv .
«hI lo A tf.tbc akleitnt rar|iM •iiiHlrleialnBiii, all tlim illiiiilfit : op. nul. ttK ofCIn ndlaUcn
(rum Itww bolm !•• It. Or. tba rffkl otvli^Ml lobr. mkiv •tln4wl nslDK Indlrals Ik* "vlnia
f it. ib( ~<tiniri" inrt to Ibtenmu: ••■. mcpu* rallixiiM, tWacmaai ibr>i<lmiuin -. i. :d.
\Ut IKHU or lb« IMonl (WMnale : llw kfl lUr Iu* bn» adUUei ■• ln<tln(c lU F. (liwM
Utn*. ibt (vnWI Mouw arw bir Uw ']ra ;/•-'. In-Uoiw Ik* |iMh fhHn ii t» III. IV. VI. tha
nwUI una* Iblnl. tuonh. ftn<l ill 111 Mim: |>. t. Ikt tnXirtnr l<nitltii>1inal biiiull*, •teffii a* a
b>uk«« llnr; .V C Uc nwttiu «u>lalin; t-V.ia« niiclaui iMMculatti: an.l T U.opHe UialatDiu
atumi InaMKno: f*'. Ibo l><in( M»b, ritf. Ik* kaaa, ami r^i. lb* blurt llnli OT lb* iliWroU rapaul*
7W«(llilll«ief Ik* limrili iviilrlvlik lib l^aMluTlhe inMctlar fotpon iiBdncMBlaa Siv •!»*■■
tir *toU*d ltn«^ ihai ol Uw bulb M •hown I? a iIim IIiw ; ji, Ibe tMnaal «IumL
■riian
m ciMUiecfe«l vid
whidi pK> h) (««j
M Uiai («n of I
iW rMiDJU uf tif I
in rkkAn.
icwllr in !
grmr Dutur.
«od to DtAtn I
iiy (if thr (W
of tibrrv pncM
•(■read iiai tau
■deimble lunbi.
■EtMT eornus quadrisMaaBS
^ _ _ MytiOlly In Uie nerwolk
If Mdr ionB of tbe ufiiic tnct i^
\we hieral enrpm EwicHhttB.
i|— liigirinmiiii PuHtwr, «« bn
pwtal ali eveuu ••'■■'■■■ /ray
I wilfc >ad. ID ■ <«ftAin >< -fidMB*
■«>5 fttiM ill mplik CTiilre it d^gcnenlw; k«
fU fflrlilMi aad Mm loa of tke ptripfaenl dtgvDcrating portion bM «^
(itivifMi* aAd on lb* tnftoe eeotre: wbn r >iniMl ncrv», for imUMv.tiH
i|i' I ' m tit* •fiiiul naftliM, tboufjli tbe a«r>'r Mow ili« «!c(n«4te*- ^
nnr iiikUhd anil tu pJMC of nerve in oWDPclion with it nmaiBfK*
itiiirli n« UfoTv, «« Itftve it, tiowerer. in oar pover to brinj; Abuut c1wh«*(
1>41
nni) wilier charnetiT. a c-nwotioti of Kn>vn\i nmountiii); to atro|>hT,
iltvi- iiiicrfi-Ti'iici- with ihtvuu* iilriii4iin« lictiiro ihcv nre fnll_T tle-
Vttlopcil. Tbiix, in nn mlult tiiiiiiiitl. ii >ri-ti(m nt* iin Dptic nerre or removal
of tlH- vyv Irjiils to ■k'^^iicrutinii in ihi- npiic m-rvr uml optic tnict : tbc aptto
Ibn* )iii\'<- lh<*ir lr>i)ihic mntri- in (NTlaiii ocllx iiI' lli<' rptina, nl' which «e
11 >|iruk in In-iiliti^ of viMoii, guiil cut bwhv from tli«l ccntni thcv (legeo-
lir lliu tiK-unn Ibr nitluiv »f tho oplic Hccumation in miimtilii, mm)
in mnn, ban lievn n«ccnniD«l. But it' tho oyra !« rcmovrd i; rroiiovnl
bittli erco bcinc dcNDtble on accoiini of tlip ctmraclera of iho optic rlwtw-
tiun) io n n«wboro wiimal. oot only do both th« opti<- ncrv» nnil llw
itirr |iKrt of hiith optic tracts ccatie to be further develo^wd and iK-gan*
titil the hodtm meiilioned nbuve, the two lateral corpora g""!*'" '**'*•
- ulvinu" on eneh side, and the Iwo unleriur corpora ifuadrigeflitaA, do
TuIIt develop: cctlain parM of tliviu undergo atrophy. The develop-
bpl of these oervous »lrucUire« seems therefore to be largely dopeudetil on
■r funrtiofial connection with the even by means of the optic tracts bikI
Brrea.
The Huriie meiliuil conSroia the view exprc»ed above, that the niodian
irfiiH i;eiiirulaLuiii ha« uo coiineriiim with viaiou. Wheu the eves of oew-
rn nniiualH arv extirpated, neither the mediou corpora gcnicuiuta nor the
iinrior corfMim ipiailrijceiuioa ihow any au|;ii of nlmphy, and tlw part of
optic irut-t which doea aot ilegenerate u the infcrinr commiMara eaa-
i«itJii)i: the two median corpora Kcni"«l*t«. Obviously th«e [Wtrta are aaao-
'ilcd with luiictiDnn of ihif bnun oth<^r than thoM of sight. The lateral
itjioni ipMiii-ulatii, th« pulvinar atid the aDlorior corpont<pisdr!grimina,HT«.
may mfx.-nt. ulone to be rcnrdod a* the cliicf oeBtral parts iit whieh tint
10 nvrviv ctid. Wc may alto rvpent that owing to the [wculiarity of tbv
lie deciisatidu each optic nerve thus finds lu eiraiDK* in buth sidiis of llie
in.
While th« optic cbiaania is. as we have aeeo. h«lpiiif[ to form the floor of
ihird ventricle, it ipvea off librea to the ouslerior |>erfi>rated api*. fNiine
tliew hav« been iiuppmwd to pa;M direetly in the wall of the ventricle to
nucleiM of the tiiiril (ooulo-motor) nerve, aiid to M^rve a« a channel fur
t Impulses, eausiiix oonstriclioii of the [itipil : but to this we shall
urn in dealing; heri-sAer with the mov«nivnti> of the pupild.
^ 671. Th<iii|;h iht' aIkivr three UkUm arv uDdoubt«dly the chief eudiiifcs
ika optic Dcrve. three primary visual centrvi, if «r« may ao call tliein, it
atw belicv(Hl that w>me fibrea of tlrt optic tract, makini; counectioiu with
'ihrr of iliw*- three hmlii*. jmih by iW cru» cerebri straight tv certain
of the cervhnti licmiNtihen- t Fig. *J1 1. <f ) ; but this fourth ending is by
mrans m> clcarlv lyliihliahoil as arc tW other thrvn.
And undoubtedly the nuiin oounection of th« cerebral hcinisphere with
optic tract is not a direct one, bui nn indirect one, through the thnM
ies in iiue^tion. Wc said (S&'M) thai (ibrea proceeding from the occipital
irtex bihI reaching the thalamus through the hind-limb of the internal
ipHile formed what was called the "optic radiation." Theu fibres be^n-
ng (or ending) in the eortei of the occipital region, end (or begin) fPig.
, i>;>. mil.) tu a Inrev extent in the pulvinar aiid in the lateral corpus
iculalum, but alw> in the anterior i>ir|>tu ifuadrixeniiuum. reaching it br
anit'rior braehiuin '^ t>^). When evuu in a grown nuimal the occipiial
cortex i- •lv<troye«l, not only ihe«e fibriy. but also [nrin of the pulvinar and
ril'Tnul corpus tirniriilaltini. iimb'rgi) dv)ceii('ralti>ti. and tliere !• some change
in the anterior corp«ii* iiuitdriK<"iiinuni. When the mmo cortex is deatntyed
in a newborn animal ibc .-anie paru ntmphy : and in nuch cbms the ontic
land ovr.':. which an- but link ndixitoil by the g|)er«tioa in the auult
MS
TUE UBAIX.
ftoiual.are aUo iiivolrvd in Ihtr atrophy. W« tiuiyailil llint rcinovml iirix^li
eyta iu lite ncwboni Knimul u nid tii l«iul, baulw tlic aimpliy of ib» tbw
ImdicB in qii«Mion, to dJininishcd ocvipiul lobo duv in Uick of while hmUct.
We may tberofvra conclude dial iu Uw oompk-x net of viviui) two oriwntt
central unptimtiu lira involved ; we insy epcak of iwo kindn of oeuUH for
vinioii — lae primarr or lover vitual centres supptiwi bv tlw thr«e bodiM of
which wo ara ({icakiDK. and « iMODdary or higher visiiitl oeatrr iupplitd hf
tli«onrt«x in the occi|>itaI region of tb« cerebruni. And pxperiineuial renlti
ncourd with this view.
Beli>r« we pn>oe«i to dimuB ihoee reoulu. one or two prvliminnry ubwm
Umm may piwra of use,
111 tW firet place. a« m; have prcvionply urfM. the iiiterp«vtrtlii>n of Uif
rcMiIta of an oxperiineiil. in vthicli ne Iww li> judp! of eeneory rfliictik *n
far more (in<«rtHia thiin whvn wr hnv« to judnco of motor etfeat», that u, rf
vouree, when ibe oxiierimcmt is conducted on iin animal. We can eatimali
the motor vtttict 'iiituitilAtively, w« can iDCmiiv and record the contnctioi'
of the tiiuitd<^ hut in tvlimstme n Msnsorr rflcct wo liave to depend on wgn
our interpnrlation of which w dii««1 on annlogi<w which may or may not tt
niialeadinfc. \Vc are on sater eround when wc can nptH'nl to man biinMtf ii
llie exjwrimentF instituted br dieeaBe : but the many aiivaiitagiTi ihu« Mcnid
are ofl«n more ihan con nteri)a lanced by the diflu«o cboractcni or the cumpkl
Gon»)niitnnl« of (he lesion. In dealiii;* with sensory ef&d* wc must eipM
and be content for tfa« niwent with (ytncluMons Icm definite and mon UM»
tain even than thoee gained by the study of motor efi«cta.
In the arcond place, in dealing with vision, it will be de^irahlv to kuo* tbt
n)c«iiiiig which we are attachuig (<i tlie words which we employ- By fating
mw, Ihni ifi, " complete " or " total " blindnew. we mean that the movHiMMl
and other aclioni of (he body are in no way al all iotltieiiced by the aauoM
of light falliuK on the retina. Of partial or incoiniilete or imperfect riaw,
tiaiug liie w<ira vifion in its widest sense, there arc many varieties: oailvs
may iltuitrnte some of the defects of the visual machinery, re|t*rdn' ■*•
whole, with its centra) as well as its peripheral parts, by rvlerring to cenaii
defect* of vtxion due to cban;^ in the eye itself The eyo may fall into talk
a condition that the mind can only apprecinle, and that to a varying ilefiM
the dilSuvncc betneon light and dorknew ; the mind is aware ihiu the rrtiM
(or it may be part of (he retinm is Iteing niitmliiied to a Ivm or KnaUr
dcereCi but cannot perceive that one pan of the retina is being ntiinuUlH in
n dilTerent way fl^m another part ; a sensation of lif;ht is excilix). hut not ■
ni of visnal MOuaUona eorrMpomling to the seta of {wnclln of Ininiiuxis nn.
which, reflet'tei) or emanating fnwu external objects in a definite ordw, sit
falling u|M>n the cyi-. The eye again niiiy fall into anotlter c>iniliticii, ia
which ^ich aeta of visual scnmlions are rxcttcil, but on account of diopUir
imiterfci-tioo!! or for other reasons the several weiiMttiiinti are not «>t«auiell
distinct : the mind ii> awnre through the rye of the esisteiic« of " thngK
but cannot uilcjUHtely recugniiie the cbarncters of thnac tbiii|{a ; tiw liimt
imof^ are hlurmi and indistinct. And a large nnmbcr of grailntvinn tn
poMibIc bdwecn the extreme condition in which only these otijecis srhWA
prCMDt tlte strongest contrast with their eiirronndings are visible to a ofliK-
lion which only just foils nhon of normal vision. Imperfi-ctioun of ihi* UbH.
of varying degree, may result from failure, not in the peripheral iipparatu*.
not in the retina, or onUe nerve or other parts of the eye. but in the cvntrsl
appMratus; the retinal image may be sharp, the retina and the njitic fibrw
may be dulv ri-Hin'onive, hut ft^ni something wrong in some jmrt or lAlwroT
the brain t)i4r vifunl scnsntions csoiled by ibe visual impuwi may tail in
dittinctncas, and that in varying dcgrr* : imperftctions of vismDt wlMtbBritf
I
I
irigiit. in which visiin
I'livrnllr «poken of undfr lli« iiol w
•tmAti/vjiia.
If line optic nerve be divided, tool btindneBs of odl' ryo will tvnilt ; but
if onir ii\*'u- tract be divided, it follows froiu whnt hns been wid ubovu that
hnlfbiiiidnvM in the oAmspondiiig: hiitveii ut' both eyes will rrsult. If. Ibr
iiutnnco. lii« ri^hl optic imet (Ki^. ^OU, Op. T.\ bi- divided, llie U}t vigiiitl
tivldii of both eyee will be hloiled uiil. The miui« condilioD will be brought
ttltoiit by failure in the optic Irai^t at iia rculral eliding, provided of <.'ourM-
ih« mischief be coofiDed lu tli« ending of tbe one tract. .Such a hiilf-blindiitiM
or half-vision is apolcen of on hnnianafuia oT heminu'ijiia or hemiopia : tlio
words lel\ and ri|{ht are tci-i": rally \ne*i in refnruicu to tl)« visual lieM ; thiiti,
left heiuiunu|Hia i» th« blottiii^ out of both K-lt vinual fidda ihrough failure
i4 the riyhl ofttic tract.
If, iii»i«iid i>f ihf whole optic nerve boing dividwl, oertaiu bundles only
were cut ac-rw*. iinrtial blindnnu woiihl be the rtwilt. a portion of the vieual
field would Im? blotted out. and miK^hk-f limine) to a few biindluof ooe oplic
tract would load cornvjoindiog blots in the corresponding hnve* of tlie visual
€«lda of both cy».
Furlb«, nn aflirction <'f h«lf the retina or of a limited arva in the retina
might iicvur of fuch a chantHcr as to lead, not to complete, but to |Mrual
bliodncM^ to a bemt aniblyopin or to a partial amblyopia. Thv pari of ibe
retina to Itfliwtnl might be central, or peripheral, or a quadrant, or any puteb
of any ■!/<■, form, iind relative portion. And we may further imagine it at
Inut iHDwible that mischief in the brain might be so limited as to protluce
any of the above partial eflects. though lh« reliDS, optic nerve, and Optic
tracts all remained inlacl.
The above visual imperfections we have illuHtrvted by changes to the peri-
pheral a|>paratu^ but there i» a kind of ini|)rrleolion which we may still call
a visual iMi[>erfcction, though it in of purely central origin. Id a norma)
>iate of thing* a vi-mnl scnMtion excited in the brain is or may be linked on
to a cbain of jMychirul evcnl*; we often, then, spnk of it as a visual i<lea.
When we see a dug the vinonl »cnMtion. or rather the jjroun of Mrnsations
miikiiij; up the vinual pcriTpiioij of the dog. does not exist l>y itwlf, apart
from all the »lher eveni)i of the brain ; it joins and aflects them, and among
the events which it hi atl'cct« may be and often are psychical events; the
vivual perception "cmeni into our thoughts" and modilics them. Between
the vixnal impulse, as it travels alon); the optic nerve or tract, nod ilji ulli-
mate pvychical eflect, a whole nriea of eveois intervene ; and we may take
it fcT granted that the chain may be broken or s|Kiill at any of its link*, at
the lairr as well as at the earlier one*. Wi? mav therefore consider it ptwn-
ble that the break or damage may occur at the links by which the fully
developed visual wnaatiixi JoioH on to {wychieal onerationu. We may sup-
jMH that an objed m »eeu and yet docs U*>t afliMt (be miiid ul all, or allk'la
It in an abnormal way.
Theae foregi^iug coiniiderattomi emphouM the diffimilty and unceriainly of
inlerprelioi; the visual coDditicin of un aiiimal which has U-cu rX|>i-rimeutRl
upon. Wbet), for inslan<-e, after an operation, an animal iwiuo lo Ite influ-
enced in ilH iireviiiuM normal manner by tbeTisual cH'rcts of external objects,
a moM careful twycliical annlyi<!» is often necessary to enabto u> lu judm
wheUlcr the newly introduced disrrgard of this or that objeel it du« to the
■HI* vumal denMlions being blurred or blunted, or to some failure lu the
nmrebical appreciation of the sensations : and in most eases S4ich an analysis
H Myond our reach. The i>realest caution is needtbl in draaiu); conclusions
from txperiineiiiB of this kiixl, especially iVoni such a^ ni>|iear to have been
H14
TnH BRAIN*.
liiMtil}- cxrriod out or lioMily oWrvcd ; and we must be coDtoiit hwe tndvvll
00 «om«' of thfl bruatler fouiiires only of the subject.
S 672. .Since we have iu this natter to trust eu mucb to 8nalogi«s with our
own experience, we roar turn at once to the monkey aa bein); more taltRl^
tire than any of the lower anunals. We have alrvady Mid that cl«tKll
escitaticin of the om|iital curlex behind the motor reKiou may prmlaca
luovemeiiu. but ih«i ihc-<e tuoremeuta arc in charaoivr difR.'retit rmm th«a
caused by ^limulaiion iif the motor r«)(io<i iMtlf. In lUt maiA < lUtka
nf partA of the iu-c;ij)ttiil rcj^ioo, the occipital lobe, ani) llu- au i lu, ftl
iiiManw. may glyc rim- to niovrmciiU nf ihr oyc«, of the fiyvlid*, w»l uf lbs
hi-nd. that L< ot the n<<ck. all tht- movcntt^ltt ^i pmiliic).''! Wing «uch ■* an
onliiiarilv (.■oiint^i'tnd with vtrioii. It will not W prutilable I" i*nli'r herr rain
tin; detail* poiiciiminE thi? t-xact topography of the rxcilabU' part* or nf (hs
niifcial chnractfirK of the inoveincale s« caUcd I'orlh. But it w important Iu
U'lle that thcsr niovetnculs are unlike the moveinontu cKcitttd by KlintnlaiMa
vf (he appropriate motor area, insHinuch as their occurrence is Oir b-m cri
tain, they need a Btron;>er stimulus to brinj; them out; when evokeal thiT ar*
fwble. Iiein]* easily autag^onixed by appropriate stimulation ul' the molir
urea, and they have a much loiif^r latent jieriod- They are not due In mtj
indirect stimulation of the motor area, ihroujth " nsMtciation " fibres oob-
neotin;: the aput siimulate<l with the motor area, or »therH-i_«e, since (btf
per^t atler removal of the motor area. Movemeuta of tbii kind may aM
be niiu&Med tn the Hug. They arc ohvioualy the nault of iiupulaa tn»
miltt-d in »oiiii> ilin^ct manufr fmm the ortex Inaome parb below, and nt
bo triki'ti AH III! indiriition that the pari* of the itiries in ijuiation arr id mv
way r>inii<.-eti:d wilh virion. The exact manner, howpvi-r. in whioh titer air
)iri>ij);lit abiHit i* at priwiit obscure The exphiuation nf ihcir ^n<Mii nblth
h fre(piently oflereil. namely, that the stimulation No hITivIa the i-nriit-al p»j
matter as to ^ive rise to visual fC'iiBalions, and thnl the ni'ivvRicole expiw
these seusalions, does not aeem Ml iii factory, For. if i( !>>.■ p'<uilile that the
KKIB9 chanjtei whidi the electric current seta Koins in the ortii-al [•'»y nalUf
can repMiluiv the |idych!c«l erenta which take place in that gray matter ia
iht^ normal uctinn of the brain, we ahoiild expect Biimulation of anyud
every part of ibe cortex to call forth wtme movement or other, ainoe it CUMt
Ik- iloublcil that every part of the cortex i* in mme way or other eDffiffd to
mychiojil operati<>n:i, nnd that cvcrv piyehical phaae tend* to express ibrif
in niovemciit. Whert^ny, ontiiido the motor region, with tbe exoeptiMiC Wl
Br<* now dixTUwing. tlic cortex i*. as we have •wtn, "" inexritable." and ewn
within the motor region itself the escitnble sub«lnuce is ncatteretl, wilh m-
cnuiog segreealinn as we advance aloni^ the animal lu'ale, amooj; iuexcitahit
aubitanoe. When we apeak of the region ur nubitaniv lu inexci table, ve 4o
n»i mean thai the electric current pn)duces no vlTeet ; we nnlv mvan that
the e6^i-t is not manifested by movement : the real diiren^ncv ItctwecD thr
ext-itable motor re|;iou and the inexcitable rest of the cuitcx in pnibablj
thai in tbe several mofir area* the current, playing upon the beginning* of
the pyramidal fibres, ia able to inauKumte simple motor impulw' or wiar
thing like them, whemui elaewhcre the molecular cbangiv indiioeil by iht
curreut arc loo oonfuKcd to reach tJieir normal expression. Thor« oao lac n<i
doubt, of ODUtw, that molecular change* in this or that part nf tlie bnin,
set going by proocmes other than acltial visual impulsee aloog il>e uptie
nerves, may <^ivo rise to visual sensation* ; and, aa we shall see in dealiRf
with the seikMs, the subject of lucli " vubjci^ive" aenuitions is unable to di*-
tinguiab them from sensations uf "objective" origin; but it is at leari
unlikely thai ihe coaree disturbances started by a letaniiing current *h"Ul>l
lake Hich a definite form. Morenver lh« vi«w in *iue9lion b disproved liv
VISUAL AND UTntCR SES'SATIOXS.
M5
I
lb« evperinteuUil mtill ilmt ilie aaiiie luoveiiieDta art' hiviut-ht aboitt hWu
thi- eortex n paTed aixir ami ilie clectruclcs nre applied to thi.- hu i>jnt-i.'iit
whiir nuiiier. Tbis reaiilt >iitj];;e«i(i tlie exiatencv of ertirrriii (racUi urbiiuill^i
tif M i|ieci«l kind, ilifleriii;; fn>ii) tbose of tlie pvramidul kind, itiough like
them making cunnecltonv witli the ooulnr utui oiltvr m\]»ekti : we hnvt, liow-
evt-r^u yel no ullicr cvideiico of iiucli Inict* vx'ti^iag.
% 873. Tbc rctullii <if iTiiioval vi' the cortex uUo iiiipjHtrl ttio mnfl g^DCnil
caitclukinn, iboiigli (kcre it much discordance untoog tbi.- various (ibMrwrs,
bdlh ai III tlw pnrrictilar result* nnd cspccuiilf mt to their iuifrpretAtion.
broftd tavt comts utit in all thr obcerration«, namely, thm tbe rrmovnl
or injury to tho bind rrgion of iho cnrtvx always prxHJuvce Butno dislurb-
of vHton, and pmduora dislurbnnct! of Ttcioa more surely and lo n
prvaUr exttoi tbun dom injnr}* to or removal of any other region of tliv
cortex ; l>ut beyond this broad laot there i» mnch dieput«, and we niuet bi-
K' litem here with n very brief sinlemenL
In tbc monkey some observere have found that n-moval of Ihe occipital
lobe on one side (the resion marked " viuon " in l-'igs- -0-1 and *J05) caiiwil
betniopia, the elf^ on the visual Heldfl lieinfc a cnMsed one ; when ilie right
lube wBM removed there was blindne^M in the leA visual fields, that ia, iu the
right halve* of the reiinaa of both eyes; in other wordn. the viaual impulst-n
|iMiiiin|j uloug ihe right otitic tract failed to produce l)it.'ir iiMial elfect, so thai
tbe animal aungaraed objects on Jlii MX baud sid«. We may remark tbnl
the decuseation of tbe optic nervo iu the muukev ih wry Minilar to ihnl iu
man. When both occipilnl lobe* wen removed, tutnl bliiidncM r«iulted.
But, and thiii is moat ini|Hir(aHt, not only wan thi? liemiopia canwd by the
nrnioval of one lobe ImiiMcnt, but alwi, arcordiug to aonic oliwrveni, the lent
vi»ion returned after the total removal of both IoIkw, though xomo impair-
tnrnt might be noticed lung afteraard. so lung in fact ao the aninnd *ae Kept
alive.
In the bands of other observera destruction of tli« angular gyrus of one
side (Fig. ^20t^) has led to hemionia, failure in the left (or right) visual fields,
indicaling failure iu tlie central endings of the right ior left) optic tnct.
Wing cfliKeil by removal of the rigbl (or IeA)gyrui(. and <le£tructiou of lioth
angular ^yri has led to total bliodnew, not only the heiutopiu. but tbe total
htiixliieM being, however, apparently trauaitory. And cases have hern ob-
H-rvi-d in which the transient blinduen due lo removal of the occipital lolie*
hwi l<r«D succeeded hr )<erniai>ent hemiotiln up<tii the sub»ei|ucni removal of
tliv angular gynu. Indeed, the general, but not uniform, tendency of tlie
nany experiments which have been made in to connect, in the monkey, botli
tbe occipital lobe and the angular gyrus nith vi*i»u.
In the dog, removal of portiotM of llie ocripitsl mrtex have also led to
p«irtial and tninnient blimturw, or, aceurdlDg lo Mime, to |>ennaurnt blind-
nest; but thi^ ilifficulties of judging the vixual condition of a dog an- vciy
considerable, and his vinion ts so dillbratt frt>m that of man, so much lt»s
binocular, for in»tunoo. than hi». that it would not Ik profilablc to relate at
length the resulls oblaincd in ibe dog or to diwuss the cooclufiona which
have been derived from ihcm. We will only say that some obterven have
lieeu led to think that the lateral part of the retina i« coiincclnl with Ibe
lateral part of the visual occipital area, the front part with tbe front part,
and mi on, the rdiua lieing, as it were, projecletl on tu tbe occipital cortex :
but ibv faciD are not clear enough lo make it worth while lo dwell upon tbem
Wre.
In man clinical histories hi far conform to Ibe Te»ults of experiments oil
the monkey oa to aaecoiate tite occipital cortex, and more particularly the
ctineus (irc ITiga. 207 and 209), with viNon. They liave, hoiiever, raised a
846
TTtK BRAIX.
point on u'hiub w« have nol yet touched. lu llie i.*X|wHmeata nn thr innnkM
((uotci) abuve, t1ie reoult (putting stide iruuieut elTecta <)<■» prob*bly i'>
" «hnck ") «r interferenoe wrilh one aide of tin! brain wk> huiuiojiiK ; und ihi*
in urliat we iniKkt expect from tlie auatomiciil nUtionN ; iht? uplic tract gm
Mtrnight to lh« K^iuiiiiil mHM« ot iu own dde, wtil ihr oplir nwliMaiB
puKS from thoM iDMwa to the occipital corux of tbr name aitle ; tlitn \t
no dwntaition. MV« of lh« fibres at tbv otitic norfc, or thry piui inia ibr
optic tract at tiMP chtuma. (Ninical )iisl<on« tench thr Mme Iflnooa «■ them
experiments on animnltt: lo«ion.'> limilod to th« occipital lobe ht\« fori
eyniptom heniiopia: anil tbi« i> Mud to be capcdully the rivult of tnbchirf
liiniied to the apex of thtr iiccipital lob<^ tJint is, to the cuoeus. Hut «x)itri-
nwotd on iD'inliers hnvi? been niiido in which destructimi of oDe uwular
grnia ba« produced, not hemiopia, hut cnimcd blinduen or crowed aniHTih
pia, that U to sar, ha* ntTttelcd tno whiile t)f the retina of oD«efe,U)d tbattW
cnmed eye. the eye of thi^ eame aide not being, or heinj* Ruppoaed not In li«,il
>ll afl'«cte'l ; similar results have al§a been stated to fullow u|hju removal <tf
on« ocd]»tal lobe. Aud a few clinical cases have Ikeen reoorilerl in whieb
diseiMe, etpocially of the aneular gyrug, seemed to affect thr vi«iuo of tk
whole of th« croaied'eye. (It must he reinemhered thai the aiiKular grrw
of man corrwponds ti> ii part only of the whole augjular gyriin of tlie munVn.
Cf. Pit;. '20^ with Fig 2<>T.) Si>m! authoH have, in eocardanoe with tliM.
put forward the tbeiiry that the occipital lobe serroi as a oortieal centre br
the optic trnct of iixiwu side only, aiul so fur iine-hatfufwch retina, while ii
fri>nt iif thi« on the angular gyrua it a oentn- in which hotli ojitic tracts ■»
rt-prr^nled. But the clininal hiitorie« hearinjc on thi* point oiDlM b*
rnguried as wholly ttti«factory ; and with rvfvrcnce If the experimeiHal
t«*ii1u we may once more iiuisl, and the warning appljv« perhaps with nv-
tjoular fbm- to these experinienta on vision, on the dan),'vr «( ooofoaiiiliDf
lltose immediate efTects of operative inlerfereiiw, which an of the oatniv*
'shock " in till- wide sense of that word, with tliose pure "< left e ion CT " iil»
Qomena which are alone the outcome of the loss of the part rcnuivut). It ii
diflleuU to Tivi*i the cunclusiou that much of the tnuisit«ry blindnnM whlck
u observed in thwc experiinenta belongs to the former L-aU-gory, lliat iW
effect b transient because It b of the nature of shock, and nut Iwcaitw tli
luM of faculty U supplied by some other cortical area heing HuhMi|i»Bally
gobrtituled for the on" removed. In the dog, injury to (ho fmnU) region ef
the cortex unaccompanied by any secondary mixcltivf in the occipital ngino
ha* IctI to inipaireil vision: and thin was probably an iiMtaDoe of'stiitck."
lor we hav« no other reiuon to connect the fiODta] r«vioD of Ihv uirtex with
vinoD. We must be very cnn-fiil in drawing the coocluriin) that, becaase u
operation produces iraasicnt hlitidiiiw, tlie )«rt operated oo haa ■ dlna
•hure in vision; and we may well hMitnte to accept the view that the wbole
retina t< repreKoted in the crossed hemtsohera.
Id codcIuhiou we may say llint. when all the many results which have lien
arrived at by fxjieriinent or by clinical obscM'alion are duly weighed, it will
h<? felt that while the evidence for tlic occipital li^, eBi)ccially the cunem,
being coiK-nrnnI in tite matter is ooiivincing. we cannot in the preacnt itai*
of otir knowlcflgc. dogmatically exclude the angular gyrvs. aud that heac*
the only clear and consbttent statement which can be mode with any eqa-
SdeDoe b the hrnad and Kinipte one that the bind region of tiie cortex m h
some way intimately cuncvrned in vision.
§ 874.'Siich an attitude becomes all the more ueoeuary when we ask gnr-
selve» the igiiestion. What b il wbich actuallr takee place ia the corlei
duriiij^ vision? Are we to conceive of il tm if a vi*uiil impulse set guttt
ailing the fibres of the optic tract underwent n» iwciitial cban^ until it
VISUAL AM> OTHER HBNSATIONS.
8i:
tliA <-ortex. u if it lli*i« su<I<lco1y developed Iiiln u " vixuitl (m*u-
' We can hanlly JU)i|)uie thti. Belween the (.-orlcx itoil ihr vgilio
the lower viiual ceutrea, the teciueiilal iqium*, inivn-euc ; aud we cnn
llr ■ii|>paBe tliiil imerfereDc« uitlt lliG«e tiDdiea (tTioducvs tli« «antc tSect
▼uiui) &!> Rimpte ieditiii of the uplic trmt. We have ic-n in ■ previou*
•rrlion ihat the ttog nnd the hird cerininly, luid nrvordiag |i:i •ome »b»ervvn
■Uii ihi- rabbit, Arv in the abtwnot! of the ixrehrnl homuphorce not totally
', Ihfir iiwvt'uieDln bciiiji ^iiiilnl )>y retinul iinpnwione; nnd cum are
d«d of tb« dog imtig ohviiiiiiJv *|UI i^iiidwl in eotm- nciHurc by rotiniil
after the Md^>itul lu^w hml b«eD wholly or almoai wholly
ramovcd. And, though thi> i» a nintlcr at pnKcnt ouI«ido exnct biiowkdge,
aiul I^MUffh it ii j>crhn)i« iHwhle Tor «imple aflbrcnt impulsoi to dvlertnine
area Oi>mpV-x nmvt^tiK-ot* without Lhu interveniioti of " roDB<.-iuusiies4," irv
are pmbiihlv juMilir'l in iiwuniing thul (he simple visuni impulwM, trBvellinfr
along ih*' fibrw of the optiir iriK't. undergo impiirtnnl tniD«foriiiatioiu iu th«
tapBCDial masMa. and that th« changes which are propagated aloDg the
flwflfl oT Ibe optic radiation, constitute something quite dilferent from the
itDpuIsM along the optic Irnct or nerve.
Judging from the noulogy of the oiolor region we may probably aasuai«
that in visiun llie cortical events are ptiychical in nature, and that ihefunctioQ
uf (he optiL- indiaiion is to fumiali whni we may call crude vuual eeouitionii
for further |)tiyi.-hical elaboration.
Ntir awfl this view oomfiel ua to supixxte lhat injury to or rciDoral of ihv
cortrx niuat |lrudul^e only psychical blindneaa or {MTobifisI impairinvnt ol'
Yiat-in, tbiiufth thin |ioinc has probably not beeii Mumclontly lieUI in view
during the variou* ex|>eriiiientj», nutTioii-nl care not having bcvn taken In
iDiDc biiw liir the litindniaa wan purely ptychical. Bearing in mind tbc
iMHtinn following upon IminnK of ihc occii>i(al iu>rt4:x, and the br-
ing cflcctK of any o[tcnition on (be brain, we may euppoae tlwt injury to
cortex nReclx the lower centres as well ; and B'>me of tho tnnaJeut ini-
IMinDciil of vision, on which we liave ju«t dwell, may perhapa he explaiued
ua (he eirect of ihe cortical injury on lite lower centm.
Although the mailer is thus in many of ita details at present ou(«ide our
Vftcf koow ledge, we may probably conclude ihal in the complex act ofcorn-
j>Iete vi*io(i. while pari. efi|>ecially the more psychical pari, is carried out in
(he ci>riex, moie particularly of the occipital region, part is afioomplldied in
ihe lower centres, Ibe lei;m«nial maasta. As to the several fUnctitmi) of the
ibree inawea, we know almoA alMolutely nothing. Kleclrio stiraulattnn, and
it ia *ud, luecbaiiiiral stimulalii>n alM. of the anterior corimra •{iimlrigcmiiia
■ iu uuunmaia, or lh« upttc hibt-i in lower auimaU calln fdrtli luuvcim^nla of (be
^^Hl and of varioun purL> <>f the biKly : and ruiuoval of th«ni oauaca blinit-
^^Hi and in vome cami Una of <M>unlinat)iiu of movements. Our knoirlclgc
■ III th«vc points iii not very exact; but fnmi lh« alwive facU aa wall as fr<>m
the ■■i>nno:lti>ii9 of thu nnlcrinr corpora ouadrigvwina with (be porta of the
brain behiixl wr may p<w*iblv NupixMHr tbai th<s« iKHlica are nora miecially
cniKCnivd wilb the part viiiual im|>ul>ca play in detvriuiaing the coordination
of iDorcmcntc^ Wr must reniombi^r, liowercr, that all three maam ara con-
aeetad with lh« cortex, aiul probably all tlireo play a part in vcrion even of
111* higbcat paychical kind,
StRMfMNu (^ Sm«U.
i 87S. The olfactory nerve, wbkh ia undoubtedly the nerve of smell, itanda
like Ihe optic nerve apart from the real of the eraoial nema; and a few
wordi H to ita auuGture and relaUons will be ntemmij.
THE BRAiy.
Lj'ing 'Jii ibv vnilml «iirluco of Um niitc-ridr rogioi) (if i-it<:h hriuiapbcA.
• m citch side of lhi> anterior liBMiro. k mod tlic (illiirtnrv Itiilli, vrliirh b pro-
longed diret'tly linckwiird tie the olfiu^Uinr' Imcl.comiiiR aiiiHin-iitly In kd ml
where ibe liim] mnr^Hii of the froiiliil lotw iibuto on Uit- nntt'rior ixTforkb*!
ttMCA in ttie Hoor of thi> Irani pHrt uf ihe thifl rvtiiricle. Tbp l>un<lla at
fibrm forming the nlfnctory norve nroper »j)ring from lliv bulb, whi<-h is cbtk
iniRiMliate c«rebml origin, both uitlb und tract being really pnrts of the
cerebrum. Jual as the fure*braia budi) otT on each side the oplic vesicle U
form theopliL- nerve, so eneh cerebral vesicle buds of! an olfaclory reride,
rhe front part of uhioh becoraw ihe ivmnded bulb aud ih* remaiuder Ihe
rounded Irigonsl Iracl or |)eduoole coDuectinK the hulb with the hetuispberr.
Ill man the original cavity of the vwcle is ooUtcTsted. beiii;; filled up with
neuro];lial gelatinous nuhi^tuuci.', but iu llie lower animals reiimiit^ an a liiitai
apnoe, ^e ventrii-li; uf the olfactory tract.
The bulb is a spvciallKed mataof gr«y nuttier, formini; a aorl uf cap lollw
eD(I of the tract, and prwcnia somo analojtie* with tlie corivx nf tlie ImbI'
sphere. Along the middle line Ikii the core uf neuroglial g«latioou> mth-
•tance; but the side uf tlic bulb ilorxil U> ihiit oire, in i-onlait with tbr
hemitf^ere, u much le« devirlnpi'il ihiiu ilitt tti<le lying ventral l» th« oort-.
next tu ihb cnbriflirm plate; nud wc may confine aunwlvM to lb* vmirkl
portion. Next Ui ihc nciirogliiil core lie> u lavin- of longitudinal mtidalUlrd
tibn.01, with which arv niin^lpfl mmc ncr^'c will*. Thi» layer which, Ibnnnh*
beginning of the tmct inside the bulb, ii ihinniatt al the ruund«d frvnl «i-
tremity nf the bulb and gmdunllv thicken* bnckwnnl. Next tu il liM a
" niicJi.iir " layer, compowd of small nuclear cells, arranged to ii Urg>- rxMI
iu loD^tudinnllydi^pcwed rovs. Fibrre from the proceding lay^'r |««
betwcwD the groups, which are moreovvr ecpnrntcd by interlacing bundlvsf
libref^ >'ext to tnis layer comis b eomewbiii thick one, which perhaps nn
be compared to the molecular layer of the cerebellum or to the pyramidal
layers of the cerebrum. It in coiufHiiied of a niole«ular gruund «ubaiaD«c
partly neuroglial in naiun.*, traversed by numerous fibrils and fibres, nan*
i>r thti latter being of the 6ne medullaied kind ; it also cnntaiiu, in no Inrp
number iu man, nerve cells, some of which from their triangular form SM
tapering branched procc«He« are not unlike the pyramidal oelh uf the wrtei.
TIk- larger of these cells are generally found near the nuclear larer. Nut
Ut ihi« mulecidar layer, or " gelatiuuus layer " a* it i« someiimea «alled, eono^
still working outward Inward the aurfuce, a cbanu-ierintic layer iu which are
found the " olfactory glomeruli ; " oml oni>idi; tbi* is tbe luyi;r of nllactiirf
Itbrcs pni[)er, that ix to say, tibns non-nitMlnllalod (S 70) hul bearing in
obvious neurilemma. Tht»e olfactory 1ibm< are arrangeil in a closettl
plexus, anil bundle of ftbriv gatliered np from the plcxuN ■( inlvrvaU pi^rot
Ihc pin nialer, which invent:^ trie bulb anil furnishm it with lui ample lupplf
of blooilveMels, to form the olfactory nvrt-c proper. Tlw strmttinr of IM
olfactory glonH-ruli, which arc nliout UM mm. in diaioMCr, has wrt T«t
boon fully made out; ihry nr«' describrd Bf Ix-ing formed by cuils of bit
olfactory fibres with small cells and bloodvcnwlsiatonipennd among the rail*;
ill the lower anininls a finely eninulnr grouiul subauuioft is proeetit. Flbm
from tbe layers benvtilh have been traoe^l to them. We may perhaps ooiiim
that they serve as the immediate origin of the olfactory- tibres; but lIioT
exact notations to the other layere of the bulb arc by no means oleftT.
The tract is compoeed partly of longitudinal Hbres, wilb which are miBgU
nerve cells, and partly of neuroglial gelatinous subaiADce. Tbo fibres b^
in the bulb, which apiieaiv to serve aa a relay between tbein and the fltini
of tbe olfectory nerve proper : and while some appear to end in <«IU in the
tnct itself, others are continued oil to tbe end of tlie tract, being joined by
VIStlAL AND OTHSB SENSATIONS.
S40
I
I
I
I
9 taking origin nlong lh<^ tnirt. Wc niny compiirc the Iiiil1> »□<! tho tract
l<> a fwrt or the reiiuu (bh n« iJiiill «u«, n purl cil' thi;! rvtiiiii <uriVM}H)n(la to
the iitfactiiry miicnUH n>i-mt)ran«) nu<l tlu^ ojilic ni^rvv.
Th« donal nurfucv <>( the tract k> itilhcixiit tn iinil continiious nith lb«
Mibatuwe of the cerebnU bcmwplicrc. in ■ eroor« of which it lies, hut tho
tract may be enciMderecI a« iDdvnriidciit of tho Itonii^phfre until it rmchu its
dkd, at which il hrcnk* up iiitn Immls of fihivs, sjHikcn ofao iti "roota," Tho
mnu c<>iu]>icu<iUH of ihciw is it liitcml ono, which ewccpiug Internllvncnwi tho
ftnt«riur pvrfDratril i>p(u», at the mouth of tlie liMuro of Sylvtii^ mny be
InK^ to the nuctcti) amygdnlti; (Fig. 194, .Vb), nn<I the junction of this with
tbe hip|Niciini)iBl or unciniitc gynu ',Fig. '20N) in tho Icmjionil lobo of th«
heiui9|>befv of the anmo liilc. A much smnllor laoilian one, which, hon-ever,
in aome of tbc lower unitnnU ia largo and coiupicuoufl, tnkce a median
iltrvctiou, pOMca into the anterior commiMtiro ($ ff'ifi) aiid ta rtacbes th6
olfnclory tract of the opjioxito side, Uther smnll ruota have abo beea
■leicribrfl.
S 676. In many animals ia whom th« senw of nnoU is ncuto, a portion of
the nirtfx, knoirn as ibe "pyriform lobe" or "hippoetimpal lobule," and
which is anatomically continuous with the front end of tbc hippocampal
gyrua (tho part to which the name UDcioale i;yrus ia ofleo restrieteu i,
acquires rvlniivcly large dimensioDs. This and tbe niintomical relatious juU
HKniioncd would lead us to suppose that a part of the cortex which b cou-
limious with the front end of the hippooanipal gyrus is in >om« way oon-
ncvtml with «uell. The argument from comparative anatomy, however, ii
oDo which must be used with otutiuD ; einc«, Widea the great difficulty of
determining the bomologiea of |>arts of the brain in dilferent ajiimata, rela-
tive iiicresM in the part in ijueBtiaa mifcbt be correlated to other thiniits than
the power of smell, and might be determined by cirtumstancce having nu
reJaiion toamell.
The esperinHntal cvidenoD, though on the whole it girai support to the
Tiew, it conflicting: and when tha dlfflcultv of deierniiniiig whether a
"duuih animal" cau or caiiDOl amell it bomc lu mind, thii will not Iw won-
derc-il ai. Tlie obavrTitliim that electrical stiiuulatinn of thn region ifl flues'
lion givt* riw! to movt-nK'nts uf tlic no*triU, n'lii<'b have liecn iuterpreieit aa
tnithitg in rexponn to subjective olfactory scnMilioiu, caiioot linvc much
aeight; ami while aotuc obwrvcni haw found that tlie removal of this
part of tli« brniii deatroya the senM of smell, others hare obtained nrgatire
rtvulla.
Thi- fi'w clinic-nl hiittorii« which hear iipiui the matter arc perhapa mor«
tnutmirlhy. Tlieiw seem to xhow ihnt a Wioii involving the cortex of this
ragioti, but leaving iha olfactory bulb and tract, as wvU iia other parts of
the brain, intact, may d«etray or greatly impair Miiell. And we may, per-
haps, give particular weight to the caws in nhieb epileptiform atlack», pre-
OHKd Dy an "aura" in the form of a peculiar smell, linro boen aa«ociat«d
with disenso limited to this region; for the phononKoa of "aura" seem to be
cnnneelcd with cortical proceaace.
Though the evidence on the whole goes to show that the cortex at the
front end of ibe hippocampal gyrus is especially connected with smell, and
wa have so marked it in Vig. itIU, yet the whole matter stands on a aoios-
what difl^roDC footing from tlte sense of sight. In man the relations of aniell
to the olber operatiuns of the brain (thotigb, as we shall we in dealing with
the seitsea, somewhat peeulinr) are far more limited than are ibuw of
TBion, aiMl (be psychical development of simple oUaetury seusaiium ia ex-
tremely scanty.
M
650 THE BRAIN,
Seiuali»iu cf Taste.
g 677. This special sense, ibough «o closely lUBOcialed with noell, bUim)),
togelbcr wiih the special eeuse of heariug, od x ditIer«o( footini; from the
two jirvciiiling special aens»,tUDce Itie nerves ooa<:enie<l Iwlong to ibe ale-
gor^ of ordiiiiir^ cranial nerves, and we lack, in reference to then, tlie
nnuioniiuil leading which is ulfered to us in the case of the optio and olbo-
turv iiurvfi^
\Vc shall tee in dealing with the senses that tlie fifth nerx-e and tlte glcoM-
ii)i«rjng«al nerve buve been ci>n*iden!d as nerves of taste, but that the inaiter
isonevubjcct to <-i>n trove ray ; thu giiAtatorr fuoctJou of the 6tih is attributed
to the pociilinr choriU tyni|iiini uuctc, iuiA otlMf (luestioDS hare been raissil.
Whatever view we luke, hnwtver, the nervet of lo.ite nre onlinarjr cntnial
Dcrvra, nad w have n<i uiiatninical guidance as to the fihrM uf i;ilher of ihs
n1>ovc two iiurvi'ii makiiiK «pei-'i)it ci>niH'cti<>ii» with nnv part uf ihc enrtcs.
Though iu;nfali(iuii uf tHHtt? i-tiler largi-ty into the life iif Biiinmlii, and ioilnd
of nifin hiinnvlr, nv have no Mitixfartory indications which will ennhle as to
Goiinvct lliii) Kpc'ial h-hm; with any part of the cortex ; the view, ind^'tid. Ihs
bven put forward that xvinc part of llir rortes in the lower portion of lb
tempiiral lobe, Dot lur from the ct-nlro fur imcll, nrvm as a centrr lie
liuu-; but thu arguments in favor of this view are not, as yd at l«sM|
convincing.
SenMiiont of Hearing.
^ 678. The cochlear division of the «glilh or atitlitnry oern nij k
nftiiinieii to be a nerve of the special sense of hearing, snil of that alcw;
the vf^tibuUr ilivUiou serves, as Ke have seen, for other functions than \iam
of hearin;^ (§ ti-lH), but as we Bball urge in dealing witJi the senses U nut tt
be regarded as whully useless for the purptisca of uiat »mm. The cnohlar
divisiMi we have Iruced ^| lilif) iutti the bulb, and the vertibular dirbita
into the lateral aiidili>ry nucleus (which, {>^rhup«, majr be rvganlcd ss s
ooiiliniintion ur segmental n^itetitiuu furward of the cun«at« ducIms ei rf
Iiarl of thai nucleuH), and iiil» the cerrlielliim, the cfrebcllar coniinusila
(cing probably lh« part uf the lu-rve whii^h m-ivfti for coordinating {\Uf^
tions. The conneclious of the auditory ovrw with the ccrvbral be(uis|>ken
hcl'iiig to tliv sanit' category as llione uf iitbcr air<:rrnl cranial, and w» WKf
add vpiiial, nerve*; we have ug very clear anatomical gutd* toward taj
particular part uf the curieK.
When wu turn lu the empirical result* furni»hud by cxperimeol Oil
clinical glNwrvationo, we liod that these, though even loai dcifinitcandka
accordant than in the caw uf the senses of sight and einoll, point tn poitgf
th« 6nt or superior temporal (tcmporo sphenoidal) convolution i Figs. Sf>l,
207, and 209) lying in the teinjiuml lobo lust rcntntl to the Sylvian lisiurr.
as being specially concerned in lionring in some such way as the ooci{>ital
lobe is concerned in vision.
Electrical Niimulnlion of thi» region of tJie cortex give:; rise to " prickiag
of tlic ran," and other movements such ae are frequently coooeeted wiu
auditory sensations : but such phenomenn are in ibis iusUnce, perhaps. In bt
depended upon even len than in other similar inslonoes. While sqm
ohwrA-era maintain that this conviilution. the ojwration includlns other |Mr-
tiuns of the temporal lube as well, may be remured from a monkey wttliott
prorlijcing any certain signs of deafness, other ob«ervers have Aiund Ihsl
reniot'al of it on one side alftoted the hearing uf the car on the oppuaite nde,
CUTAKBOCS AND SOUB OTnER SENSATIONS.
SSI
ttnfl rmmvnl on boili hiiIiv brouglit the uniiiikl into n coii'lition in which,
without being, [icrha|Kt. itlxu^luMlj^ cIohC, it ru(icte>l luiruril sriunil iu a vtty
im{>crri-n Riniin^r iiuWed, verjr iliiri-rotit fmm if n'lriiial ItchaTior. Tho
tcttuty clink'ul hi*t<irici bMring oii Cht« mnU«r iirt' not vt-rr <l<.viiuvc; tut
(bough dofttcH hw beeii oW-rvtil in cnnncciioii wilh ilixr-o*!: ulliscliiig llio
M|i«riDr I«in|Mnl cunvoliiliim, Xbv li^ion Ua^ usually invniltHi other [iiirt* A*
««ll, ami tho iliAfiiew hns bovn imiH'lBtf) vrith other «ymptoiiK, iioUtbly
opbuin. All auditory "niirii" hiu. hovrovcr, ai titiirs h^eii obicrviil in oon-
Bcccion with dinuwe nf this nvioii, u nlto n |HM-ulinr jwychicul fnilure,
known ta " vrurd-dcAfutsw," in vrnich, though «tuni)8 arc bvard, thai ia Co
Mf Mui'Uory wimtioas »n f«lt. il amy bo ovon as UMintl, thd porcoptivii or
paychkal npprtmtion ot' the tounils U tacking, and a ftpokea woia '» not
rvn<g»imi.
Iitutlr. we may add that though, ni we said, tho anatomical leftdiog is
Dot debtiitc, ob»ervet¥ have fouuil that in nenb'iro animiil*. on tho ono
band, destruction of iho fwrt of the ci>ri«x, |>rjl>ably corr«:«poniliiig to llie
ragioji mmtioncd above, lemla to atrophy of the median corpus gniiculatuiu,
and to mnte extent of the poBlerior corpus quadrigeminum ; aiid, on tho
other hand, destruction ot' the iulcrnnt ear lends to an atrtiiihy of part of
tho lateral fillet of t lie opiKieile criMsed side, wbJeb (BAjr b« traced to the
posterior corpus qua(lfi;(i;iuiauin, and llieiice to the median oorpiK genicu-
lalum ; and seetioa of the lateral fillet nu one side leads, among oltier re-
sults, to siruphy of tbettrin acustica; aud tuherculum acusiicutn (Smifjof
tho mnwuil sidu. Thia auicgeita that the path of auditory impulses ia along
Uw cochlear nerve to the uUeral fillet of the crossed side, and so by the poa-
lerior cor|His (jiiiulrigeuiiiiiim and median corpus geniculatuin to the cortex
of the teinponl lobe uf that croMcd side, the two later boiliei beariutf toward
bearing a relation aomewliai tike that borne towar«l eight by the anterior
eorpua quBdri)^:cininuiu and lateral corpus geniculainm. But tlm matter
needs funlier iurestigalion.
Tlierv remain* the special aenae of touch, but this wo bad better consider
in oonueciiuii niih mentations in gonerml.
Ox THR D(n-|CI,OPllBKT OP COTAXtOVB AJIO OOMB OTItER SkMSATIOS'S.
$ STB. Tho eenaaliom with which we hare juH dealt ariw through im-
pulsee [M)«ing along special nerves or partA of special nerve', ihe optic nerve,
tbo olfaetury nerve, etc.; ne have now to deal with aeitsaiious arising
through intiiuises alou); the nerves of the hudy generally. Those are of
several kinds. In tlie tlmt place there an aeusaiious which we may speak
uf MS "cutaneous seoautiotiii," the impulses giving rise to which are stnruil
iu the skin covering the budy, or iu toe so called mucous membrane lining
certain passages. Vbtae sentatioua, wbicb, as ne shall see in dealing vita
the MUM, are dependent on the existence of special terminal oi|[aiM in or
near tlw skin, are senaations of "touch," iu the narrow ueaaing of that
word, by which we a[>preciuie contact with and prenure ou the skin, and
the w-nsatiuns of " tcni|>enilure," which again we may, a* we shall aoo,
divide into <<-nBstiiin)i of "hrJit" and >cnuiti<>ii» of "cold." ThcM senaatioiM
may Iw cxciteil in varyinj: degree by impulwes pawing alimi; any nervo
branches of uhicli are siipiilird to the akin. Tlien lliere are the wnsaltoiH
ninitiltiting the " muKcular srniu-," to which we have alrcaily refi-rrol. and
tlH-*e again may bo excitcil in nny nerve having ounneotion* with ihv skrt«tal
uuKclo.
Aa nc shall see iu dealing nith the se^s(ll^ when a u«rve is liud bare and
652
TDB BRAIK.
it* dhmt arc fltimtilnlcd dircclly vith^r by pnmure, Kucb u plnchij);. or br
hmt, or by cold, or in other irny*, Iho E«»uiotii which are atused ilo nnt
fnablo UK to npprwisto wlii-thcr the elimulatioa is one ol' contact or pmniR,
or of tcnii>erntiire. "r of mime ollirr Itiml : w© only Piperience a ** fccliag."
which nt all events vthrn it rcncho.i certain intensity we speak of sa "pain."
And no have rctuon lo think that nt leti^t from lime (o time inipulaes alung
various n«T^'cs give Hkc to sciifations which liave b«en apoken of aia iboae o^
" <^ncrnl Bcneibiliiy," by which in addition to otfafir senaMiona, mcli U than
of loucli and of the niusciilnr ecnne, ne become aware of chan^ei in tbe con-
dition and circuni^tnnces of our body. When tlie stimulaUon of lli« akia
excee<)s a certain limit of intensity, the sen»e of touch or teuperature it Icat
in, that b lo «ay, ia not appreciated aa separate from, tbe seuae of [tain ; and
uiidcT abnormal circumstances acute aeusatiuns of pain nr« started by
ohangeit in pitrtsi, for example tendons, thi; condition of irhioh under nurmal
cJreiinisiancvs we are not conscious of appreciatioi; through any diatiact
HcniutlioN, though il may be that these parts <Io uorniallv givo rise to ftablt
impulses coiilrihuting to "^neral acnaibiliiy." It may Ihcrcfore ba debaud
whether " pnin " i# a phatw of all ML-iisations, or of ocncnd nntibtlity aioiM,
or a aeDsalioii itri ^mrrir. We »ball hnvu utiuetbmg further In aay od Um
iDBltcr vrheii we tn-at of titi- aenscn ; nieiiiiwhile it will be convciiifot fijr
prtacnt purpoew if we coniiitlcr ibiii thu M^imtinna wo have to deal with jort
now are the aenantion* nf toin-h imd of tcm|icrAturc tho«c of tb« muacular
Ecn^c, and ihoM of genera] wMisibilitv including thmw of rain.
S 680. The lairly convincing cvi({<rucv that the occipital cortex haa «|)«ciil
relatione nilh vi»i»ii, and tbo leas clvar evidciMo that other ret;miis hate
special relations with #mcll and hearing, suggest that opecial parts of ibr
cortex have spedal relations with tbo M-u»aiioii4 now uuder conaidermtiaD.
Rut in tbecoenof the senses of sight and smell we had a distinct anatomical
leading; and we have seen how iincertaia is the eviilence where auehoa
anai'inilcal leading fails, as in hearing and taste. In the cose of aenntiaaf
of llic body at large, the anatomical leading siniilarly fails. Moreover anr
attempt to piith the auahig)' of sight raises the Ibllowing (jueetion : If tlute
were Ino optic nervts on uarh side of the head, would tliere be iwo cortKal
nrtn.i, one lor each nerve, in each hemisphere, or one visual area only ? And
again, if the optic ucrve were the iostntmeot for some sense in ailditioa to
thai uf fight, ivotild thi^re be two oorlical areas, one for each actu>ation, or
one area only srrvin)( on the cortioal BtatJon, bo to af)eak, of ibe wholu nerve*
If wc push the nnulogy of sight it is open for ii*. sin<-e we cannot give a
definite answer to the nlMve (juoilion, to suppoee either that there \*, ouc afm
for touch, another area for iem|ieniture. ana so on, cueh fur the whole bu<lv,
or that there is an arvn liir sen^aiious of all kinda for i-Dch ufli-rcjiL nerve, xr,
that then? i» an intricate arrnngemerii which su])pli«» all the iinnbiiuUioaB of
the two uhich are re<inirud lor tbe lifi- of the individiuil. Of ibo thits
hvpothcwe the iHllcr i» the nicin- pmbable ; but ifto, it \» by it* very oatan
nfninst in*iieceptjble of en i>cri mental pro()f, e»|jeuially when we bear in mind
what wc have already (aid touching tliLMliKiiolty of jmlging tlw smsattoiM
of nninints. If the judgment nf visual senMilions is diHJcini.hnw much mon
difficult must bo the jiidgmmt of nenMiliotia of touch and lemperatureT
Indeed, sensations of poiii are the only tfciiutions of which we can tonn a
qnantitalive judgment in animals ; and our method of judging eveo these,
namely, by studying the movenivDta or other effects indirectly produoed, is ■
moat imi^rfoct one.
We can learn, therefore, almoat absolutely nothing in this malter frtitu
experimental stimulation oT the cortex in animals. As we have jirevioorlj
J&ViJt) urged, the absence of tnovemente when parlaof the consx oiberlbui
CtTTAA'BOas IXD 80MB OTHER SB::TSATI0N'S.
&fi3
the motor ngiom are Blimulaler] is no ovid«nce tlist tboslimulntion doa not i
^ive riM to mrcliical oveiiU iutu whicli scnsntii>ii« eiit«r; and movcmcoU ]
follov Btimulaitoii of tlie motor area, not beesim that area is wholly givMi
np to motor ereiiu, but because fi'Oin the bialolaxical arraDgemoot th«
stimulus gfitt rmdf tceem to relatively simple motor meohaniani*. TliJit ths
motor re|[ian has close oonn«otiona with sensory footon is not only almost
certain uo ibeorelical Krounil«. but h ibown in many ways, fur example by
the experiment, deMrribed in S Gtii, of exalting the leuaitireaess of a oiotor
area by generatioji peripbenil eeDwry impulseo.
Nor can the eflecta on Miiaation of reinovat of parts of Ike ooftox be
inierprvt^d with olearneu and oertainty. In thi-> monkey removal or
Heeiruciivii of die gyrus f(>rni<-utus{Fi{pr. 'iO-S nui] 205) on the meaial surfaos
of the braiu. veniral to the c'ull<i:to -ra^r^innl suI(-uh whiuh forriH on the meaial
aarfnce the ri>iitral limit of t]i« motor rf^i»ii fait operatiim of very great dif-
ficulty), ha-< brought die wfaulu iif the opjHuite »i<j<: nf the body toaoonditioil
which baa bceu <li>3irribty| nn aii anicsthvsia, that U a low of all oulaueoua
tactile Mneadon*, ainl an aunl^fvvin. that itt a lo« nf M-iuatiuiu of [lain, the
couilitioii ImIii^ acc!itin|ianivi] by littlr or no impiiirmi>iit uf vulinilary move-
mcnlii iiikI, ihnujjh npjtaruntly ilimininhinj; n» linir went on, lantiiij; iiulil the
(liMth of tlii> animal some wft<lc» nArrwunl. Ajfain. removal of the eon-
tiniiation nf tin* gyrus foniioHluii into the iiyrux hi]i|>i)campi hoa in other
ioManoea led to a nu>rv tran<ipnt annj^lhtvia also of the whole or greater jiart
of one iUki of the body. Ami it i« imi-rlcil ihut removal of no other rLVton
of thf cortex iDt<:rfcri.ii wilh irulnn^oiis and )«inful s«ii«atigns in «u striking
and laMtiiic a manner as docs the removal of parte, or of the wbulc of Uiu
mMtal r^oD.
ThcM ruulta, however, do not accord with clinical experience, which,
thougli aouity, seeniD as fur ns it gix* to shoiv that in man, when mischief
apparently limited to the cortex produces loss of aeiisatioDs. it is Ihu parietal
lobe eorret|H>iHliu>; to the mutor region which \a afTiwted ; but ihore appears
to be no recun) of any caM of a cortical lesion aSeedng aeiisaliaa without
aSet'iiu;; mi>i'eiiiuiil. We have praviiiusly called attention to the fact that
Uie temporary Iiku or iiapnirmeut of movement whivh follows removal of an
area iafr«()ueutly, if not alwuvii, mi-omjanled by an impairmeDt of cutaneous
sensations In the limb or part " (taralytod ; " an<l side by side with thU we
may put tlie expuriciicu that in the human cpilepliform attacks of cortical
origin, the *L-iiiirt: U at times ushered in by peculiar •enHations. calleil the
** aura," in the part moventents of which iiiauKuralv the iiian-h of oonvuliiv«
novemcnts. Rut these thin;^ do imiKhow thai the cortical area is tbe"aeafc
of sensaduiu,"thev rstber illiii>lratc what wt nutd ooncerninK tbo complexity
of the chain of wliicb the events lu the orlical area an- linlU, and tite cltMe
lie between sensory factor* and the diaracl^rislic elcuunls of the motor
nrioo.
la ibodog, while romoral of almost any oonsidcrable portion of Iba cortex
nflbotsanmdon, removal of |>arts in the frontal region producing per^iapa
l<M «6«ct than removal of parts in other regions, the loss or iiapairmeot of
•cnaatioa apfwars to be trannent, though having a duration brosldly pnpor-
liouu* to tM extent of cortex removod ; and when a very large portion of
tha cortex ta nmoved. some imporfeclion apiMutrs to rem lia to the end. ^Ve
have already nfcmd to the cim« of a dog from which the greater part of
both ccrtbru bemitphem hiui Imwm removed, but irhich remaiiwd capable of
currying out maet of the ordinary bodily moremcutB, ami that apparently in
a voluntary naninri in this ease the "blunting " of cutaneous scnatioiM
was perhaps more striking than the imperfcciiim of movement. It will be
854
THE QRAIN.
wadh vrhilc to consider the condition of this dog n liltlo cicely, on Drcnunt
of Ihp li|!ht vrhii-li it thron-e mi the {l^)b1em which irv nre now diKruwiiit'.
Clinit-ul vxpcrieticv shows thnt i» mnn (lie tnlpgriir of the cH-nrhnl hviui-
epherefl, and til' the conneclion of ihr hi7ini#phprcs nilh the rot of Ihi; i-eiitrnl
nerroua PTsleni, is e»^ntiiil to the full dcvc'opmi'nl of Bensnttana; Hiid ittnl
in this i'e»pei?t eavh heiniitphere is rclolcd to ihe crowed eid« of tbo bodr. A
very nminiitn form of pHrnlyfis or " itroke " is llim due to n Icwiou of dmiic
pnrl of one hpint<>pbere i the e\act poflilion of the lexion need not concera n
now f. freqiieiilly cuiiaed l>v rupture of a hloodvmsel, in which ibe pttMlit
lo^ea all [lower of v«lciutary movement and all senMlioM on the cmoaed fide
of hie body (.incluilio^' the face) ; lie is said lo he sulTeTinK from hcaii|defta,
"oiie^ded stroke." Not oulv )to vnluntarv iiuimlaea fail lo nach the mu»-
c)ea of the oflecleil side, but Bensory imjiuiaes. aiii-)i as those wbicb, starlcd
for iotilance in the !<kiu, would under nornml mnditioiti leiw) to aenaslioas of
touch, of heat ur cold, or of pain, fail to ede«t oouaciouaneM. when tiwr
originate on the alfected itide : the iiatient ceaiioi on tlinl aide feel a reugli
surrace, or a hot body, ur the prick of a niu. For the sake of c)e«iroeM it*
•uppoeie Ibe loes of ninvenient and »en««ti<in to he eoinplele, but it niif bl nf
CQune be pnriiaL Such u case Bhuvs we retieat llmt l\n< integrity of thr
oerobnil benii*phere, and of the eonneciiaoH ot thnt )i<:ml->pbcre, we mny mj
of (he ct>rtex ot that henil^phure, with the other parU of the nervous qntm.
i» cmcnlisl lo the develnnment nf the «enM(tion!t : l>ut it dnea not prove tbat
the oorlcx of ihu heminphere i* the " r<}al " nf thu wMiKjiltoru, it d(M« not ptvn
that ibi; a(IVr«'nt, and Miiimry impiilm^ olnrted in the skin, uodorgo M
Dintrrinl chiinge nnlil thfv mich the Diricx mid nn' ihi'n ^nddciilv convertrJ
into »i'n>alii>iiii : il njily priiv^'* ihal in the rnniplcx chain of <.'vents bv wbitk
ecntory inipiil^ick give riw In I'lill tmnsciDiin K-nwilioiw, thcevciila-in the conn
niroish an in<Iie|>enHible link. And the phi'nonicnn of the dog in qUMwa
on the one hand illiistrale how complex ihe chain is, and on the other hand
mi^gest that the contpletcDeas of the lorn of senHitian in tbe heniiple^e bmu
h not a pure '"deficiency" phenomenon, but is due to the Iwion atlecliDglha
diaiu of eventa in some way or other besides merely removing tbe link
Airnialied l>y means of the cortex. For, as we previotiiily urjce^l, the dog in
i)iit«tion, however curtailed ila psychical life may have been, wcmnl to a
nenal observer to feel and move much as usunl. Neglecting vimukI and
auditory sensations with which we are nol now d(!alii)g, it nertlol cnrrful
ob»ervaiiou lo ascerluin thai m>me of (he aniuiurx movi-meuts fpll iihort. llir
failure lieing apiuirently due to the lack of udeiuatelv energetic ciH>r<linnliiij;
tenMiry impiil>rH; n stnniger stimulus than iittunl find to be nppli*^] to tlm
•kin in unlcr tucjdl furih the usual moveinenis and utiter lokena tliai tlw
•linMihiii niu "frit." A* we have before urged, it is inipoMiiblc tosnupoM
that tbe mere (tump of cerebrum k-fl in thu ciuc i-otiM have tnkcn or nil iIm
funi'tinoK uf the loft heniLiphcri-M ; an<I making ax we have prvviouely don*
full allon'ance for thediflVrentiatinn butwcrn niiin and t\<r<j[, wc muri ooncludi-
Ihat in tbe more geneni seiimtinnit with which we arc now dt-olini^. ne wilh
the ninri' ^pircial vnrmil scUMlions, tbe full di^vrlopmcnt of a complete mdm-
tion is a complex act of more singes than one between thi- adereiit impulfc
along the nffi-renl nerve and the aflirction of cnnacioiMncm which we sub-
jectively rKognin! a» "the scnealion," llio cortical Bvcnt* are only mnw
among several. It fullnws that any nnal^igy bctwcon lh« cortical evenu
which play thi-ir part in a sensation and the corlical events which irnme-
diatcly precede ihe imuo of impniscs frooi tbo motor region along tbe flbra
of the pyramidal load U misleaiiini^ ; the hi);hly diRereiitinted molor hicallm-
tinn does not juslily ns in concluding thai ihere exists a aimilnr lojiograplilcal
' iisiribuiiou of Mneattons.
, . mav now Hlbick tin: juvililrin in * iliflcrvot nn)'. nml iimli'Kii
Wgiiining with Iho onrlcx ln-gin witli ntfcrcnl imptilww stnrtcH nluiiK nlfcrrol
iwTvcs from (heir pcrii>h<rnl piiiiinjpt, iitiil allrmpt to Inn^ ihctn twntnilwird.
Anil lint wo aiKv («ll to minil nhnt nnnlotnknt giiiiliiDcv wc |i[>{«Fn. (S 570.)
W« hRVt HCD tbkt tti« tilirrs of nnMcriur ^»<>li^ the chBnnMd of nSorcnl
iropalM*. cod in the Hpinni cord in «t lea»l two mun »»]*». Ono tiet nrr r^ii-
liiiucil "O. not lirokcn by nnv rvUys, lu (ho metlinn pn^lvrinr triict, and Uy
this Irnct ropn?80Dtativf» ot nil the s[>iiinl imtvc« arc coiinccl«l with lh«
gnu-ik nuclviifl in nhicli, ^fill, tli« nitxlinn {losCerior coiiinio ends. The
olhor fibres of a po&tcrior rum ap[>«sr to ciiil in the gray matter not fnx from
tbnr entrance ; but from the gray instter there Blarls the cerebcllnr tract,
which, though not conctusively iiroved to be, mar be aMumed lo be an sllercnt
tfkct. We may therefure pnibubly suppoee that afikrant irapulaca aloDi;
certain of the fibres of the pMlerior root make their way upward aloOf; the
cerebellar trad, and there are some reasoiM for re^rdiiig the TeeicuUr
cylinder nod tlw colls which rcpre*eTil thia where it in nut conspicuous in the
ref^ons of the cord, as a relay betuecn the Iwd ^yittems of Hbres. There are
alM the more scattered tibres of the ascending a ntenv lateral tract (ji &t>H),
which probttUy ia also an allerent tract, and therefore probably also cnn-
oecled with the poscerior roots ; but ns uo have seen mir koowled^ie iif thia
tract in imperfect, thou^'h, if an some ui>;e it coda in the resliforro body, we
may [tcrhupe oouoider it as similar at least to the cerebelJar tract, and treat
the two as luiev
Thus there iieem to be at least two main reci^gniied [mIIib, in the form of
tracts of 6bre^, for at&rent iiDimlnea aluu^; the ei>nl ; one along tJie median
MMArior column, the other aUnig the lateral column in the CLTcbellar truct.
The latter paasea straight up to the cerebellum by the nslifonn body, lmv>
elling along tl;« luimc ntdu of the cord, and any cro*>iiig ><f impulse* (Musiug
along lhi» tract munt toki.'. place brfore they mitrr the tract ; wc have, how-
•rer. no anatimiicnl giiidnoco lor viich u crowng. Tbo other pnth. along the
BMdian poMtcriiir [racl, comes to end in the grncilc iiucIcur; it has, imleed,
been nrgiil tlut the grncilc nuclei)* ic thus ciinncniect chirHy with the lower
limbs and lower pan of the body, and that the nniilngous posterior root
Gbres from the upper limb* and iic>-k past similarly inUi the cuncalc nucleus,
or at least into the median diviaion of that nurlvus, but this cannot be con-
ddsred n* proved. Moreover both the posterior columna, median and ex-
teroal, bring to thme nude! DbrcK which h:tvc started from mme relay in the
gray matter lower down, nnd which are not Kbrca coming atnii;;ht wiihuut
maj r»lay from the posterior roots; these, however, wc cannot diatiaguish
mm each other in their course beyond the nuclei. From the f^radla and
cuoeate nuclei the path onward is a double one, one broad, one narrow. The
broad patli, the one having must fibres and presumubly ctirrvin); m<«( im-
pillsii, leads to the cerebellum by the resliform body ; and lierv the path.
praviously continued exclusively along the same aide of the oord, heDimcs
partly craned tboitgll ninaining ;iarily uncroaseil, the aeiiaurr decasaMloo in
the bulb beioj; the oroased and the other fibres paisiu;;^ i^rom the nuebi
attmigbt to the resliform body bcjng the uncruaseil one ($ 613) ; the uncrossed
one we may, perhafw, look upon as really an upper part of the cerebellar
IracL The uariow path U die fillet (§ 63-5), by which some of the libres
from the nuclei am- co[itinue<l uo toward the cerebrum. This jnith is a
crossed one. the (-ntteing taking place in the sensory decuMalion, and it car-
liea relatively fuw impulw4, the chief increaM in the titjn <>f the fillet as it
paans onwiinl lieing iliie to tibmt coming from siniotun^ other than the
graf tll^ anil cum-ati; nuclei.
Ili'iicc of the MiuMry iinpultcs travelling along continuoiu tracts io ibe
I
THE BSAIS.
epinnl conl, these tracts nppftrenilv' kci^iiig alwAjr* to tho borib ^d«, tlie
creat mniDrity nnw to the L't^robclliicn; nnil i>r ihcw again the fmUer nu»
b^r, atl thoee alnng the corchclliu' tract, and some of ihoeo paMiHg llimuxk
the frracile nrtd cuneaie nuclei rcmnin uncrowcd to the end. Th« niily p«lb
by irbich nil these impnlsee lliiis paasinR to the cerebellum c«n gitin nrcm
to the cortex of the cerehrum, is by some or other of the lies betwcrii lb(
cei%belluD) and the cerebral cortex. The relatively feir impuleee which (■■
aloD); the fillet are for the most pnrt> landed id the rniddle paru of the hrmip,
for only a Hiiiall portion of the fillet paareH l» the cortex (( &3o), wmI it it
not clear that this part pf the fillet coiues from ibe gracile and cuiittlt
nuclei, ^a that ntoet of these impulses can cain sccos to tlie cortex only bj
the relays of these mid<)le parts of the brain.
Vvry striking:, indeed, are these conslanl relays oloii^ the path of anuoty
impulses ; in this reufiec-t the sensory impiilaea offer a atroag oonlrv.-'t \n tbt
TDutor iiupulsea. Itut a still tnore complex system of relays ha» t'> l>e tni»-
ttoned ; fi>r yet a third path ia open for sensory, aflbreot imgmbf* aloD)( tbt
o»rd. We miiM admit the pMsibility of alli-Tent Impalsee trnmllitie ainar
the ui'twufk 'if the Knir matter, their path lieiiig either ahaoluioly i-oDan«j
to the gray matter, or leaviDg ibo gray niattur at iiiten'ala, nuil }i>!iiin; it
again fav meiitin of thoae, toncer or thorter, commiMural or irifruunoil
fibres w^ich unite the longitudiiinl ti^mcntii of gray matter, and fors na
ioooosMeruhlc |>urtion of the vrhrile while nuitler of ihv corvl. Vft ha*f
seen (§ AS7) that under nhnoroial circiiai9inni:ni impnixv pais frwIyfaniJI
directionH aloD)|r the ^ray matter, nm) we may coucludv ihut under DurnuJ
eirciini^iniicu they can piuw ahmg it under ri-Ktricli«iiit and along linca d»
lortninv<) by phv.iiologii-'nl ccinditiiniii. Th<- Hbrm in the white matler wUlt
do not show either divci^iidiu^ or utccDding degcaontlion am pr»bub1y. a* n
have laid ($ M'2). intern itiiciul fibro*. oooneeting ■^meols of ;;ray noilM
in a longitudinal dinwtton: and, though we have no exact knowledije IMt^
ing this matter, wo may suppose that some of these ooovvy impubea Hptwdi
and others downward.
If, as some maintniD. the fibres of the ascending ant^ro-lateml lrac< «nd
not iu the cerebellum, but in the gray matter of the bulb, or hiifher up, m
have a lourlh path for sensory impulses, which, afler the (>rimury relay io
the acKniental gray nintter, pass slniighl up to the bulb.
% 6S!. ilow do experimental n»ulis and cliaioal bistoriei acoord irilbtueh
an anatomieal programme?
We may 6nl call attention to an experiment, wliicb, though sonMwbu
old, carried out on rabbits, and con&ned to one region only of th« oord, ibe
lower thoracic, has nevertheless a certaiu value on aeeounl of iu ■flhrdiiqt
mon or ledt dttilnctlj' i(us.utilative and meuavirnble reMltn. Wo ban seal
(S I7R) that alTereiit im|iul«« started in aflcrenl fil>n,«, in iltoac, for instaiiet,
of thi; ikiiatie nerve, so afleot the va«>muf>r (.-entrv in the bulb as b> caxuaa
rise of blodd-jiruxure, at least In an animal umler urari. Tboac aflVnot
impulses mugit \inia hr some nntli or other from iitv root* which «npi)lT tfai
sciatic nerves wil^ Htii-rent fihreH along the thoracic and orrvical conl ta ibr
bnlb. If the path lie Muclted. tliv Htimuhition of tht^ »cit>tic nerve will Glil
to prxiduce the usual riw! of blood -pnvMU re. Now in a rabbit the aiixmul •<!
rise of blood prewurc folhining u|Km thu (timuhition of ono sciatic oerw
with a certain strength of ciirnmt having beou asooflaiood, it is found that
a much lose rise of blood-pn^iirc or none at all folloira the same stimulalioa
after division of cerlain i>nrli< of the cord in the mid or upper ihoradc
region ; that is to «ay, the scclinn of the cnnl has partially or completely
blocked the path of the aAeront inipulwn. Further, the block is oonapirixiu*
when the lateral column is divided, and is not increased by other jiarts of
llio mni beiiiK dividiil nl llit? same tini«: when I>i>tli Inlcnil NKiiniia are
divkM, tli« block id almiuit complete. Ami furiher, AU]>pi»iiig udo ticialio,
my llie rinht, i» the oiw vrlik-b h Hltroulnli;!), m block oocun both nbt-u the
lateral cobimu nf llic Mimi;, nf^l, »ici« nn<l irhcii that of tho cnjMvil, left,
ri*le b <livi<le<), but \» tjrvntrr wbtni th« itivioiun in on the croHwd than when
it b i>u tli« Mine Hilr. Wv innr ill^<^^ tbtit tbc impiilMs, which rencb tbo
luRiliar inni bv tlic roota of tno Huatic iivrrc, travol up the oonl, ur f;iv«
riov wiihiu ibi- binibnr cnrA U> cevnU which wc maj cnmtmro to rKTViJU*
impulm-ii, ami which travel up the corH lo »uph a maniior thnt in Un- hiwer
Ibiiraric n-cirni lhi\r p*>« almnct excluiirolj «loti2 the 6hrr? of [Ik* InU'ral
ctilunin, Hoino hnviiig kept t'> the mow aide of tne conl. hul nii>r«- haviug
|. ornMiit over to tbo oppoailc Bilk' before rpat'hing the thi>riicic region.
Tbi* rr»iill ww> i>ht«inc<i in rabbilf, anii tbo eKperimciit whs mrrietl out
in ihv liiirrr thoracic region only ; the nmcbiMons to be drawn fmm it hold
|IO(hI fur that animal only, and lor that i>nrt onlj of its cord. Moreover,
the experiiiH-Dt only t<«t8 the path of euch im]>ul«c« as reach and atlrvt the
TMOinol'>r centre in the Ixilb. It ia. however, DXceedingl}* prolMible that the
inpulaea which, Reoeraled in seuBory nerves. alTi-'ct the vasomotor centre aro
impulare which, in the con«ciouB animal, f-ive me to eensationa of |>ain ; in
an intacl anitoal chan;^ in the vasomotor centre occasioned by the sliainln-
iMHi of eeniiory nervee are accompanied by eigns of nmre or leas [laia. And
indeed lliia is oonlirmcd by the fact that eimilar reeulls were obtained when,
Ike eiperiment beint; conducted iu a eimilar way, sigua at irain instead of
rariatioiia in blood-preaiure were taken as the t«kenii of ilie blockioj; of
inipul-M. Hence, aMuniin;; this, wo niny regard the experiment as indi-
cati(i){ that the impulses nhich form the basis of painful sensations paM by
the lateral columns iu the lower thomeic region of the L-ord of the rabbit,
and therefore, though thi» in a funhi^r atunniption, by the nine eolumna
aloiiif the tt'holc length of the cord. Wi> further may infer that whilv aome
of the impulaea keep to the 8anu> Mk of the conl, olhcrf, and indeed the
greater uumbcr, croat to the ojipiieiie Mo.
TheM eoDcluviona entail nssumpiiona, hut the main intcrprctnliou of the
whole eiperin>cul ontnil* a still greator aaninptioD. Th« tcitiog of the
influcnn' of llie tKiatie atimulation waa oanied out soon nfter the aoction of
the conl, and vet vin have asumod that the block of the imjiulfc* wti* due
to a pure drjieiency phenomenon, the aliM-noe of a UHial path. But wc
have no right to do thiit. It m ^lomiblo that the section produced, in aome
say or other, a dvpmxing or inhihitorv effect loner down in the cord,
otfectinf' etructnres other than the laternf columns; all our experic^icc, in-
deed, of the eflecia of o|NTntions on the cord would lead ua to expect thia.
It is further t>os»ihle that ii section of the lateral colitma might prodnee thia
depraaiing eliect, while oectiona of other parla did Dot, or might produce
mon e6ect than they could. It '» possible, for inataiice. that the section of
tlw thoracic lateral column inhibited, for the period during which the ex-
Gnmctit wa« carried out, the gray matter of the lumbar cord, and that the
ock nallv took place in this gray natter. Until the uneertaintiee thus
sttMtdinfE the interpretation are removed the ex|>erimeDt ia not valid as a
Eri»f that the lateral columns are the paths of alleretil impulsee; it would,
utrever. Htill aervo lo indicflte that the air<.-reiU ini(>ulse« reaching the onrd
aloii)! the sciatic nerve croawd over to a large extent before they iratne under
the iiitltience of the inhibition, aince we have uo evidence to »how that .-uch
ail iiihil^iury aelioii of the K«tioii would be exerted chiefly on tlie crooaed
tide.
Again, we liave aeen that the afferent impulse* nffiH^ting the vaaoinolor
oeutre gain ocota* to that conn without the liclp uf tlie |)aru of the brain
i
TUK BBAIV.
nlwvc (he bulb: th« ^xblenoe of iho vasomotor entire wns made oat (J) 176),
by combiDlog sttmiiliiiion nf n sciatic awvt nitti a twries of operaliow cnn*
mating in ninkin^ Micc««»)ve trnnsvene sections of the bulb from abuve
doiTiiwnrd ; an<l it whb not until the Beclions ivat^'lie^l liie rasomolor pvaire
that the blood-preseiire eflW-ts of tbo K'iatic etiriiulsii'>a were modtfted.
Hrat-p, if ibe ex]ierim«Dt be taken aa sboiriDg llui not only alTeiviit ia>-
nulsps alffrcliDK ihe vaeomotor centre, but otber aflbrant impolMs alau imrd
oy the tHicral cohiinDs. it would also seem to show that ihew otber impuba
raw ID like manner to the bulb, ami fpiin aeceM to lli« oonex lliruu|ib ibt
bulh. TbiiH iiiL-reases a <)i0icultj wbieh preaenia ilwlf vn^a when ilie afleml
ini|iiilses alfectiujc the vasomotor centre are alone oousiiltre^l. If tine es|icfv-
meat means anything, it means that tbe impulsa having in aaiaewajv
otla-r reached the lateral eoluinn, travel up that column by aome coniinuiNi
path, anil imlee^l b ifvnenilly lalton ax bavin:; thtii mmitiiig. Il<i( if a« pul
uoiiltT the very ihiiibtt'iil vievr that the aaceniliii^ nnliTri-hileral tract end* ia
the bulh. there In no <'iiiiliniii>ii> iilTiTcnt inict in ihr latent) column viuling
in the [>iilh ; the only ilelinitc oontintutiu ntFurMit tract in iho lateral cilania
of whirh nc have any clear knnwhilgc, nanitilv. ihi^ lyreUrllar tract. «*Ji
not it) Ihi' hiilh hill in thu cm-bclliim. Anil if itc attempt to gt>ttiat«l
Ihe ilitficuUy hy wippnninir that thiwc impnlnw at least which anoit iW
vasomotor centre, nfi«r truvrlling fur »omt iJii^tance in tbe cvrelwllar IrtiJ,
leave tbnl tract for xomr path Imiliii^ t» the bulb 'ami th« eervbellar tnd
AotB probnbly give "H' »* ivcll as receive Sbm along ils enun«^), we |irKti-
cally admit ihat the experiment docs not prove tbe existcnoe of a cuntiDaoM
piith.
A further difficultv i« raised by the fact that, ncenrding to tbe interiinta-
tinn which we are disctiinng, the section of the lateral column breaks ibi
pntlia of what we mav consider two kinds of impulses; those, tbe larfir
mimln-r, which have ulready cioased iVom one side <if the cxtnl to the nthsr,
and tbow; which have remained on tbe same side. For, as are bare alrvadr
wiid, we have evidence, in man at leiiat and aooie other nnimals, that alKreiU
impiiUc* crrm coin|ileiely over somewhere or other on ibcir path b«lcin
they arc develops) into full senMilionH : anil ae have alio cvidenoe, iboo^
bw strong, that they enui* not lon(f after their entrance into the ronl. But,
if wcmppiwo thin to bi- tbe cuse in tbe ntblnt aliio, it follow* that in ibi
experiment in tiuivlicm the inipiiUat whioli were blocked on their pOMUip
nloni; the lateral column of tbe aame sbk, whatever the way by which tblf
reiichcd that Intcnil column, were pursuing a tiath which would evonlutJljr
have Icil tlivm lo tbe other aide of the oonl. Hence the tei'tion of tM
lateral coluinn. in hrt-akinu their path. Iimke not a rotiiinnoun path kve|iini
to the IntomI column up the length nf the conl. but a niilb which *oon Irft
the lateral column lo pnwi cWwIicrc. Tbe experiment, tnercfbre. as flir a* iW
impulses passing up the xanie xide an- conci-nied, docs not prove that llw]r
purfuo a continuous |>ath along the lat^-ral column ; and if so what bewnMi
of ils validity of the cxiieriment as regards the impulses' eroming over trm
tbe other aii^e, for tbe experiment in itwif nmki'S no dirtinction beima
tbe two?
We may add, however, Ihat tluingh tbe pi>int has not been spet^'inlly Iv
vc«tiK»led, it is possible thnt in tlia rabbit, in wliuse hind-limbs bllatent
movements are so pnytominant, lher« n asanciated with the mnvemeala ■
bilnleral arrangement for senHaiioiu, and that thnae impulses whicb rsnisia
aloMif the Mime side of the coid as the nerve in which tbey origlnata, an
carrieil up to the brnin willioiii any crosfiug at all.
$SB3. The results of this vasomotor experiment then, though tbev are
fnnpiently cjuoted, do not nhen closely considered aiTord adetjuatc proo^tlial
•flVn^nt iinpu1«r« pumi« k cvnitimimi* |inlh ilnng the liitcrii) columns of (he
otti, ami mun.-»vfr lliti fiu'tK brought In light hv ihn cxpcrimeDt «li'iw hut
lilllv iu-c<inl uitli tlw anaromicfll imigrnmitK'. \Vi- hi>v« dwdl i>n it to long
»brmUM.* it i* more or ]««■ illiialrnlivc of tW mnnv iliffii^iilticx alt^Diliiiit the
iiilrr|in-(aiioii i>( i.>x|H-rimrnl» of ihis \tintl ; ami il i* in thru n'*|M'ot nil tlie
mort' valumltlfl tfM-nuM! the notiifil (■xprrimontni iyvuIIk mro sbnrp nml fvlciir.
Wr iiiHv i>M> oier mi>rr rnpiiliv' t)ic iinmi'rniis e<K|KrrinimL* on thi^ lower
naminiiM, >ti('h iu> rnUhit* itml dogs, in nhirh olWr inilicAtiotiB nF icDiutton
have hwi) miuli; ut of.rhii-My (hiwc which nrplhc*ipn»or|minriiI nMnnliooji ;
(h<»"' have h«i'w rarrini out in vnHnuB repnnn of (he cnnl, hut chirflr in tht
ihnracic n-^i<in. iin<l in ihem a like unccrTainlr of inl^rprelntton t* further
ini-nnw-d hv div want nf cxiicln«*s nnd aKivoinciil in (li« r«»nllfi.
If ire cnnlirnt nuKrlvps nilh innkinj^ no ilixlinrtiori lielneen the iliirrr<>nt
kinds of alTcmu impiilsrs, nnd in (h« cute of ihew animals it wnuttl h.inlly
be profiliihk to nttcnipt to moki? n ilixlinction. we niny say thflt thi* •■-vcrn)
■ •itxrinicnt)' m> fur nf^rvc ihnt they point to the laleml columns as hoing thi.^
uhiff paths of alTtrenl, fcnmry in)|m])>ra. or to spenk mure eiacllr. to tlio
■INMMge of these impulses bcine <»|>ei-ia]ly blockinl hy aeclion of the Inlpml
■eoldtnitB. tfome obefrvera find that in the dog and other loner nmmmale m
nction of the lateral colimin on one aide, or at least a liemisection of the
cord, produces " loss of sensation " on the oppoaite ride, (■reater than on
tbe ain« side, or confined to the oppojrite aide, and even accompanied
hv an exaltation of sensation, a hy|>erMthcsla, on the aiirae aide. Other
oWrvers a^in, and these certainly competent ohserver*. find that, in
the ilo}t, section of one aide affects sensation on both aidea, and. indeed,
rhi^'tly on the same aide. We may perhafs onoe more repeat th« vaining how
difficult a the ciuantitative and (|ualilativ-e dt.-t<Tnji nation ofM-nnitionH in such
an animal an the lUig : and may r«'ni(irk thnt in nil thc9« cases of unilati-ml
LsFclion the incrciiM-d blond Kupply dur to failure- of the normal vnoooon-
'■iriclar tune niuyl intliicn<v thn (ifriphoml development of aensury impnl***.
In ihete experimint.-', «» in ihtiw on voluntnri- movements, it is laoH im-
portant to diatiiiguinh hi'lw«N-n tmniiHliatt^ or tenipomnr and more lasting
tfTrclM: and obaerTcra hnvo found (hat the loss of sensation folloiring a
liemiM-ititin of th* rord. like the lo<« of rnluntary movement, is temporary
only, and eventually disappeiirs, though the recovery is slower and leai
eoninlcte than is the ckm with movements. As with voliiniary movement
(f G04) ao with Si-nsntion, recovery. tJiough leM eomplete than that of move*
meni. b pQiajhIe when n hemiscetion on one side has been at a later date
fultowed by a bemiscclion on the other side. We tnay, therefore, repeat
in rvfercDoc to sont>ations the remitrks which we then made in refer-
ence to inovenient; there is, however, an inijiortnnt diflerence between the
two cases; in respect to movement we have evidenoc that under normal
c<>ndition« the pyramidal tracts pinys an important part and thai any other
|NUh for volitional impulses is more or le*B an alternative one. whereas in
napect U> seoaation we have no anatomical or other distinct piuof of any
Boen Dormk) jMth.
Th* expennetita on monkeys are in tike manner oeilher aecordnot nor
deviaive; and even in thrae animals with their more varied Mgna of acow-
tiona, the interpretation of theae sij^ns is beset with fallaciet. fiome nb-
aervcn bare fuiund that a hemifcction (in tlie thoracic region) priMluce^) Ion
ofaenaaiioa on the croajeil side, accompanied by little or no loes on th«
Mme aide ; other ohser^'era acain have liailed lo obtain after a hemisectioo
Htbftetory proof of imy auch marked loss on the croEMed aiile. Further,
large portwna of the lateral column, the more internal pons adjacent i» the
gray matter being leJl, have been removed without any very obvious and
TUB BRAIX,
ceiiAiiily wiibout any la&tiag defects of aenaaiion on the one tide or tiD th*
oUwr,
£ 684- Tli« f^ltiiica) hUtories of diMases of the cploa] cinl in ntaa briDg
to light ill a fairly dear iiiauner a f«et of fODW ini(>'>rlHlic«, tumely, Uul
the Kvcntl im]iul96i which Airia t1i« hues of Uk Mvi-ral kiiiila afattauia
of touch, h«al. c»Ul, and |Miiu, and of iho muiiouinr ttaae, mn tranvmil
along the O^rd to dilTureiit wavo and pn^umnlilr bj diiRMVnt ■ininu
For dtMuuc may iiupuir one of ihmt muationt and leave the othwv iolacl.
ThiM ca»o> of :i|iiitiil ducuue are recorded, in which on one aide of the budf
or in one limh onHnary tnctilo nnwiti'Xiii M.fincl ti> 1>o liltU^ impurMl, nM
yet MnsatioQS of pain vrvn iil)«ciit ; ytbua n OMillc vras thrust intu lh« akia
no pain \tttf fdt, th'>iiii;h the pikUcnt waa airare that the ncc^lte hai bm
presN^d upon ibo i>kin at a particulnr spot; and converMlr in otiiercnMt
pain htie been frit upnn the in»crli»R of a Dcedle, thi>u);h mere ooataet with
or prMBiiro on the tkin ould not bo appreclAlcl. Attain, caaaa are reoonW
in which the ekin was teiisitivc to touch or pain, but not to rariatioM vt
lent perature : il i« furthi>r stated that cases have been met with in «hi^
cold eould be appreciated but nul heal, and vice reraa : and there are Mat
&cta which point to senaatiotu of pain heini; mire chicly a.«sociated wili
tlinae of heat, and utclile aensalion" with those of cold, than tliuae of pi'a
with thoee of touch or ihoae of bent with ifaoae of cnld. ('aoes of ffonl
diaeose are al»u remrded in which the nmseulnr seme ap)>cared to bea£bciel
apart from other senMtiooB. Weaball return to iheae loattr^n later oe a
dealing nith thoaensea: we refer to them now Htnply lu iihciwin:; thai iti»-
eaae, limited at far as ran )»e u^certaiiied to the 8[>iual oinl, may ulKsd lb*
aevenl at'naationa aepnratvly. and therefore a^ ta^ntttitig that the ttntH
kinds of inipiil;^, fiirmin)^ the lin»o» of the Nevcral kimU of aenxatiun. u*
traMniittod in diderent waya and f»lluw dilforont " paths " aleag the apioal
cord.
Clintca) histories mnreover agree, at lenal to large eitcni. in ahowin;; llM
nhen the lesion is conlined to one-half of the cord, ike sensaiiiiMt affected n
the parts below l)ie level of the leaion are chieBy or even exclusively ihm
of the crossed side. But there is not entire a<.'cordance, eajtecially at totla
cro8sin(> beini; complete. And with re^rd to the muscular sense there iis
distinct conflict of opinion ; the nmjorily of cnsee seem w show that in ait-
lateral disease or injury to the conl. the museular sense in company milk
the volnutary inovementa fails on llie *aroe side; but cases hare fam
recorded in which the niuaeular aanae in comiiany wiih other aennliio*
seemed to be affected on the cra«sed tide; it nitwL be rvmembered, howevfr,
that il is very didicull t» appreiTliite a dfticiency i»f muscular sente miacM
with deliciencie* in other utiiiHaiiuns, and wa should d priori expect the mia-
culnrftense tii run pnrulli^l with motnr impiiliu-M.
When, howcviT, wr appeal U> clioi<'al hi^toriu* or indicatiooa at to lb>
aeveral |<athii within the Kpiiml cord liiken by iheie several inipuUe*, tbf
answer it a miul uncrrlnin one, lu indeed might be expvctcfl fnnn tbe KM
often difTuKe chnnicKT of the li^i'ins uf dtseaw; and it \* [N)rliB|M not IM
much Id car that i>n Nttisfnctory d<v!uctioas at all can bo modo.
§685. Whcrihcr tlicii we turn to experiments on animalt or totboitedr
of difcaae. ihc loachings with regard to sensation, in contrast to thnse wito
regard to voluntary movement, are in the highest degree uncertain ao4
oDMure. A few general reBeotions will perhaps help us to appreoiali tbt
value of such facts aa we poataaa.
We have seen reason to think that in every movcmewl. whetlier voto
and of cortical origin, or involuntary and siarled either as n simple •.
redex or through Uie working of some part or otJwr of tbe brain, the i
I
I
CUTAXEOUB AVD SOUS OTDEB SENSATIONS. 8SI
iiniMiltww, which twcep dunn the motor fibres to tba rousclca. imuo mar-
sliBllnJ nitii cixViliiMtled tram l)i« gny maUer of Um eofd (lor lh« auke of
di-nmnt «r« nifljr otnit the cranial nerve*) from wlitit we hare cm]l«tl tbi:
■i)"i»r moduinisRia of the cord. Analoey would Iuhi) up to suppoM th»t tlic
nlli-rviii iiiipulM*, formiag th« baaea of Uie aaveral ktoiu of KDttatiuna, aiml-
Inrljp li;ll tW udercnt librts to join tba gmy mutter uf tlie cord in what we
naf call ibo Mosory mechanism. Am) ^uch anaiumica) lexliiiji iin we
pTMKM tttmt to Biipport thia view ; miih tbc escepiion of the mciliou po»-
K'Mor Intut, to which we irill return imuKNliatvly, nil the Shren of u piaicrlur
TDul M«m to end iu the gny matter not very fur from the eiilronce of
the root. We have seen tliul a ooOrdiuatv rcllex ntuvenitiut may be carried
out by at Ifast a few »e];mei)t« of the conl ; that a reSex movemi^nt may
be started by ctimuli uf vnriuuti kinda and therefoiv prcmmably by atTcront
impuUa of various kind* ; and that impuUvn forniinj; the baau of tli« iDns-
cufar matt »n Mwiiiial to thv ciHtnhmitUm »f tht; n]o<reiiMDt. All our
kii»«Iri]Ke Ko^ t>i show that in a r^tlex moveBienl carru'il <>ul by a low seg-
■ueiits lit ilic onl, the whole vhaiu of nv«ut» between thi; arrival of the
adervni ImpulaM along iba poaicrior mot and ibn inuc of eHl'rrnt impubet
along tli« anterior root may be carriv<l out by gray niatlor. and gray raattor
tlone. We may further infer tliat, while on ibo one band tlio hum; pro-
cedure miKht obtain not through a IVw w^gmi-tiit) only bnt alimg tlw whole
Itaglh ol the cord, there would bu an ndvaiila^^o. ni'iiiallr in rvupect to
the rapidity of lran«ni»ton, in employiiig inicriiuncial ImctB of tibraa b*-
tncen tlw teireral Kgroent«, the advantage being greater the more dittaiit
the Ngmenla which have to work togvlher.
W'e might furtlicr tuppose thai il would ho of ndvaotag« to poaMiai aomo
direct path bclwwn the oercbral cortex and the spinal Hanory mechanism
iume<lialely i-onncclcd with the posterior root, such as is atl4>r'led by tbe
p^vramidal tract between tbe cortex and the *puial motor mechantum iuinve-
ilMiely connected with the anterior root, but ito auatoniii-al evid«uce of
•ucb a trad ii fonhvoming ; and, od we have before remarked, along all the
traoU whieh seem to be aeDaory in nature, in controat to what takes place
in tite motor tracts, relays of gray matter are i-oiuiiiuuliy lieiiig iMter|ioLBted.
The HH-diau |KBie(tor tract, uace it gat hem up rtpienetilalivcii uf Miceeative
DATTM, prweuis ilM-lf as llt« nearest apptuHch to auch a »eiuury humologuc
of tba pyramidal tract, though it emJ* iu the bulh and it not continued
directly un to tbe cortex. And pursihly it di>e« play a xmieithut analogous
|Mrt. ill fo far uh it »erve> a» a Hjiecial CKnucction hclnwn lh«i brain aiul tlw
whole tericfl of Hpiiial iiervra. Hut we are wholly ignorant as to wlial il
mlly does; and whatever !« the exact nalurti ol' I'he part which it playii, it
proliably has relation* not to one kind of sensation only, hut to all tbe dittos
rnt kituls of sensation. It liiis indeed been suppiwoi by »nM to be csfic-
cially a tract liir ih« impulM-s of the muccular «en>e; but neither oxpctitneuL
nor clinical study aflbrds udiijtmle jiroof of this view. The condition knom
w locomotor ataxia, the sidient Iralure of which is 1«m or impairment of
muNular sense, i* avociated with diwaM of the posterior root and of its
entrance into the cord, not with discBK oonfinvd exelurively to tba nmliiiu
pottetior ouluma. Moreover, the tract cannot carry all the iropulMrs of
uiUBcular tense, since some of them must pas* at once into the gray msllcr to
take part in the c»r>nli nation of reHex movamento, aitd BitHi tbanftire travel
by librvs which do not form this tract. tSmilarly is tb«« BO idaqtHte proof
ol the Iract tK-iii^' an eiclueivo channel for tactiM or fbr pdabl MflMtiona.
We may alito. i^rliups. urge »iniilur conBiderattOlM with Ttffoi to the cere-
bellar tract, which. thi>ii;{b Htariiu); from a relay of gray nutter, is theoce
vtlWBXd to the Cerebellum a vuiititiuous tract. Tlus tract also has beeu au|»-
8A2
THE BRAIN.
pOMd to can? impulaee of a particular kind, and more {Mrticulnrly thowef
tuuacular aeiiee. rhere U Iras A priori olijectioii to this view, uiic* thr tnci
Mnrta iVoui the gray niatier, wher« the iiiipiilMS of muscular eenan uuiy hnn
ulrvuily (lime Uiuir, so to sgieak, loral work, and emis in lh(><wri'lH-tlum, wbirk,
na wv Imve Ktti, seems etJiiecially coiinei^lod nilli llie (VM>rdiiiu(iun of ixii>v*<
uieut*. Ilui M'illi ri>&pt^ct lii tliia tract also neillier expcriiiK-iit nor cliiiio)
study al!i)rds any clear and ilecialvc pruut' that it is «olely or even niwciallj
OMivi^rued witit the muscular eeii«.
With re;;ard lu the aDl*ri>-liiieral asccudinf; tract our knowledj^ is tea
iRi|>erfect ii> Juetil'y us in su{>fici»iiuj{ that it is the special or eKclutivc cbaniwl
for any one kind of seuiiatiuu, or imleed id drawing any cooclusiuti^ at all
concern ing il.
But wlieD we subtract from the while matter of tho cord ihme coDtinDOM
tracts (if asvcn<li»g de)(eueraliun of presumably sensory or atlerent functi-is.
and ibe continuous tracts of devceuJinj^ dc^nerniiun, irhich we may oooli-
dcntly s|>eiik ol' as tuotor or at least elfenait, tlH-re are leA only ibe fibrts
which we httve l§5S:J) Bup|iiiscd to l>e longitudinal oomrataMiral or lot^r-
iiunciul it\>n* between nurciwivc mij^metits. We are thus driven buck looar
titrnsr conclusioti, that seiuury impiiIiM<* ftam either by the ({my nialtn
alooc, or by a seriu uf tnvjm us it neii>, by relays of gray matter cuniMvtd
by internuucial tracts ut' lilins. whuni It-iigth «e eaiinul ssoertiun, but «liitli
niay hv nhort. That Hurh inli-niuncinl tracts intervene is rendervil [irubUiW
on ihi! one hiuid by the fnct tbitt sectiou of the while inalter, leaving lJb<
gray nmiicr iiutou<-hp<i, doni ntli-rt M-ii.-<ali<inK, nnil on the other h«ud by thr
lact that the acverul kiudn of n'li^titii'u apjH-ar tu iravtO along the card b«
sopurutc paths, or al ttuu^l may in: nrpiirnti'ly blockvil. It t* uf coursr, as as
have already urf^eil, p<irsilde thai the eUcct uf a section of n tniot of Hbm
may not be the mcn^ bluck due to Iinss uf continuity, but somv ac-tton tm lb
gray luallor nilh which r.ho tihre? are cttiinectcd, whereby that ^ray nulicf
lails of ite usual functiuns and ceases to carry unmnl the seosirry impiibn
reaching it from beion' ; it is alsii iHtwible that tliis or that leaioti of iliMwit
nrny, directly or indirectly, allect {larlicular parls of the gniy laittler st
allect llie gray mailer in a particular way, so that a ocrlain kind of maairj
impulse and nouo other is blocked. Ou the oilier hand, we have reasuu lii
ibink thai the rate at which impulses iravel along the gray uiaiter is rm
slow com|>are<l uiili ihat along nerve fibres; and in the striiftgic for lin
rapidity of Iransniiiviuu uf nervous imjiulscs is of great imi>ortaiice. ilesot
the view that ihe iulcinunciul lihrc= intervene has more to cuiumeitil it ; it
IK moreover to a certain extent dupportei) by clinical hinloriee. Itiii, it' «t
ucoeut litis view, ne iuii«t at the mitiie time admit that, in auiiiialj ai leau,
tlio Itne* provided by the iuiernuuciid tracts arc nut rigid, thai within limiu
and under circuiiHianccF alieniiiUve luules are poasible.
§ 686. We may here, [lerluiiM, laise ouce mora, and this time more pniai-
odly than hell>re, the duuhl whether we are jualilled in auiiming, a* «(
gcncndly do iin^uiiie, lliut the events which take place in ibe tiltrss ii»-
nceling relays ol gniy uiaiti-r wilhin tbe centr:U nervous *yaii-iii, arr exaetiT
the same nx tliow which take ]ilnce in tbe tibrtii of nerves outaiile thr nntrsl
system during the piuBiige nf what we call u iiervouH impiiW. Mini of v«r
knowledge ol a ncrvoua im|>uliM! ha* been gained by ibe «lndy of tbe tuudir
nen-e of a mnsclcnerve preparation. Our knowMgc uf llw pnicw^c" in at
lercut nerves is mucb mure im perl eel ; but llirreare nuuiy facts wbieb al Irhs
•nggoal tbat the molecular event* conslilnling an allen-ni impulse along sa
allvrcQt nerve arc dilfim.-til from, uud probably more cvmplicate^l tbun. ili^as
constituting an elfcreitl iinpulM; ntong an eHenMil nerve. And, with regard
to th« |>romiuu taking plac* in fibres witliin the central nervous system, a*
OPTANEOl'a AXD SOUS OTIIKR SENSATIONS.
863
I
ITC liArtllj imj exact oxpcriiDentol konnlcHgv nl nil, It luM l>oou matii-
tftiiwd by iDftoy nbwrrcn tbat not Olilj' ibc gray mntt^r, l>iil nUn tli<i Iriii-O
of wliilc loatUr in llio xjiinal cord, while tlii-y arc m|iiil>l<- »( conveying ini>
pulM-s in one (lireclioti ur the other, nrv int^iniihlo (if b<:iog so cxrio-it by
urtilicial sliniuli M to g«ncrflt« next inipulsca. Then obwrvera mntnliiin ihitl
nlifti niuvcmenis or sign* of »eii«iitioD fvllow tbo direct slimulKtion »f varimi*
parl« of (he cord, tho cfl(>c[ii arc due to ixcuiug motor I3br» or cnloriiig win*
Mtry libra bmving bei^u riiniiiliiled, and niit to a aiiinulBtion of thv intriniic
suntance of ib« puri^ tbomsckn; thoy pmpoM nccorditigly h> call ihoo
part« " kinesodic ' ami " iCsthModio " ra<(>ective]y, that is to say, serving m
patba lor inotiir oraemory impulses without being themsolnM either motor
or aensory, Tlie i<vid«ac« on Ihe nhole goes to show that this view in n mis-
tokeii ooe; that the vurious tracts of the spinnl cord, like the pynimitlal
tract, and iitdeed other [mMs of the bruin, are excitable toward ariitii-inl
stimuli. The question caiiuol, huuever, bo considered as definitely dosed ;
aad t\*e very fuut ttuil il lias been rnised illustnitea the point on which we
are now direlling. We may further quote, in similar illuairulion of the
«rae point, the fdlowing rt'tuarkable fact wiiich waa obmrved in the series
of ex|wriinenie referred to in ^ G6i un the eifecta of repeated hemieection of
the spinal eiird in duifs; The animal had partially recovered voluntary
muvemeuiii in hia hind-liiubit utter a ibinl heiuitiectiofl of the thoracic coru,
and yet uheii, at his ileatli, a strong tataniuos current iras directed tbrouch
lliu Bulb und cvrvical cord, no moveineats of tbe liiod-limbfl followed; tna
impuhea started by artificial sttinuUtion could not paaa the bridge which
auffioed fur rolillunal impuUea of natural origin. It is not too niucli to say
thot our exnerimeutal ItuowMve ta to the eveuls wlitcli aocoutpany tlie
activity of the strucIurM within (lie c«nirul nervous system it almost entirely
limitni to llx- rccn^iitiDii of the " currvJiIa of uclioii " referred ut in $ 651}.
We are already going beyond our lelhur wbvu we umunie on the straigth of
this that the proMwi* started in the fihrva of the pyramidal tract by artifi-
cial stimuliition nro in all rcspM^ts idculii-Jil with tli<>»c stattcd in the libres
of a motor nerve, \^'e arc going still more bcywd ■>ur tether when no
assuntv Uint the proccwcs started in the MiinC pyramidal librcs n» the outoimo
of nnturvl events in the motor cortex nrti of the same kind. Ilui thcw
anuniptions are trifles cunipnrcd with ilic nsfumption that the evvnl» taking
place in the fibres of tbe optic riulijtlion |Misung from (he pulvinar to tbe
ocxripiial cortex are identical with the events taking placo in thi.- tibn« of
the optic tract on the way to the pulvinar, or that the events Intvclliiig along
the Bpiiial cord to the bruin as tbe result of a prick of the little finger are
idfuticiil with those which the prick has HttLried in tho fibres of the ulnar
Derve. Of the latter events we know a little ; of tbe former events we knoiv
nest lo nothing. Aod we may here ask tbe question, WluU is the meaning
of these continual relays of gray matter alouj; tbe ecnsory tract, unlees it tie
that at each rtlay *>imtf tranlfuriualion, some further elaboration of tbe iiu-
pubea takes pUce. until what were the relatively, but only relatively, simple
impulsw along tbe libres of the peripheral nerve are by successive ste[iB
diangeil iu the complex events which we call a cAnscioua sensation? This
is what wu had iu mind when we gave ($ G63) a note of warning ooncerniug
ibcdungtr of considering- alt the events iu tlte central nervous system at cither
DMior or aensory in nature. It in jierhupa not an exugKeration tu rv{>n3iunt
the views of some olisen-ers a* if tti«y Huppa»d that ult'ercni impuli-es, suy
tactile impulMS, U>at i», impulsca eventually giving rue to tactile sensations,
travelled unchanged from the skin to tbe cortex, and there suddenly btus>
•omril into sensations. If such a view were trae, undoiibieally tlio chief liuk
of jihysiology. alnMst tbe uuly one, would be tu asc-rtnin Uiu trout uk
TSC BBAIX.
or patk* mak» tat*
praMcM* w to vktt an
which Uicae impukM {MUMd. Bat iC os ito Mkcr 1
urged biTe uijr real fi)uDiIaiii>ii. the humom of ti
inogtiiAcuiee eumpuvd KJih ih« altaoK gaiMAtJ
the KTeral ciuuigca bj which simple UBpalm arei
tiona, uid when and bow the diaag^ are eAend.
{U7. SeeiBg bow UBaiisfaMary ii oar pnaeM haaahiJm. wiik rvgaH M
tbetncUor pathaofaeocatiitiif id the rdaurelT uipla tfcMl eoid, u vooU
be maloi la aUenifit any dtanuoioa u to thcu- patha t> iW mch ■«■
eomplez brain. If it be ptDbaMe that tbe prntrnge it aSeeud bf wbiyi «f
Ef iBau«r in ihf. forruer, the aarae ra«th'vl ■• aocb Mea* frnfeaUa M tbt
«r; and if iKJiUer exiKriment itorclioical Modr lbr»«a wieb HBta«a
ibe pnih up to Uie bulb, theaa csimot be cxpcdM le ci«« flBScb Mpin
Ibe maxc oigny matter and Rbrei by wbldi tbe balb it jmad le tbe obRM.
Th« iwveral (lu'Sncd areas or collcOioD* of rnr matur. and tba antnl
■Inuuk und tract* of fibre* which w« briefly dcacribcd in a pcrrioda HOiiiB,
mtitl hare of courte a meaning ; but it may be deabted wbtcbv m 1mm
even to much a> ■ cnrrcet glinama of that nkcasiaf ta aoy ooe ease, ff m
txoept tboM which urn in immediala conaecUoa with tbe cranial iMfT«i lal
tbeir nuclei. Seeing that iho thalamus appean oo tbt ooe band to be am-
occtcd with all or nearly nil parte of the con«x, and en tlkt etbar ktai It
■trve a* the front of tbe tegmental ayatem. it it icmpltag lo aappoea Miai il
playa an imporunt pnrt in eeoaatloiia penaining to the body gaaerally, «
port of it, the pulvinar, cortninly 'loee with refcratee to tJie special Moaa of
sight ; bat we uve no decvivo indications as to what pan it play*. And
tiw part which it plnjrs, whatever that nay be, is not aa exdocirely eaMHy
one, noce both expenmental and uorlrid Iceioiis of the tbahtaae are apt to
produce disvrdera of movement as well as oth^r elTerent eflecta, We oo^
perha|)e to >ay tbe pArU which it plar«, fur it is a eonplex body. baTJpg
uanv tie* and prolialilr irerr«riiiiiig many dutiea.
Tne CUR* [lieu '•UK lillei u)cuiu, weming oa it does to be a special inlenmacii)
Imct conne<:lini[ what up|ienr tu be n*ure paHlcuUrly aUvreot or ecnsaer
ports of the bulb, such ■* tlie grucile nod euneate nuclei, with rartOM paiH
of tbe middle brnin and pruluiblv with the cnrtvs, preacnta iiarlf a* a
prolmblc piilli of xcnEatiifiu ol^ vuu kiud or anotlwr frooi tbe budy at latp;
tbC'iuirniw iMlh" of the uiutomicid programme (§ 681 r; but b reftrente
to this liKi. Govond its prubable oonuection with tbe auditory lefiaaliwi
(i 078i, we Ucit eridenee.
A i'»iv>{>ieuuus {nrt of ibc brain, namely the oercbellun), natarally anm
our attention on account of its large connecttoor with what appear to W
a^r«nt siructures ; in the luiit lorn leal progrmmmc, we called it " tbv bnod
pnlh." By the cerebellar Intel it has an uncraacil grip upon what ia pncH-
cally tbe whole )eii;;th of thu spinal oord ; by th« olher constituents u( Ik<
inferior peduncle it iins a like uncrossed grip upoti what appear to be aObtat
etrncturca in the bulb, the gracilc (uiu cuncalc niideitaa wdl as on iht
eighth (vcMibuIar) nerve and pnjbiibly rcprcscnlntives of other aStraal
cranial nerve* : it haj furllier a crosx^l grip thmiigh the grncilc and cuncat*
nuclei on the alTci^ut poaterior coluriius of the whole curd. Il is, of eounc,
possible thut tbe (>erel>ellur tract, though id itself oncruased. Inrs its hand,
DV nienni rtf the veaioutiir cytinder fur instunce, on impulses which ban
nlrendy rro*scil fh>n tht pveteriur roots of the other side ; fur as we hart
seen the evidence BS A whole ahows ilmt acnsory impulses do cross over : bet
neither has the craesing of the immilaee tieen definiiely pruvc<d, nor has the
path of tlie cruuiDg been clearly oeuoostrated ; while, on the contrary, tbt
bbree uf the audltury nerve which [>at«i to the eereliellum, and whidi us we
have >uj{geeted ($ Ul-'J' '»"}' I'' coni{i<iied to an outlying port of ifie oer^
>TaKB ABPICTS OF THS TOXCTIONS OP TUB BRAiy. 8fl3
bcllsr trad, ccrliiinir continue uncroceed into the peduncle of tl)« buiu« tlchi,
Wv mity coocludfl, tlierefore, tliat tlie ties of the cersbelluiu with tbc |io*-
irrior nM(» SI* botb cnaBM) and uncroeaei). And we may revard iIiIh dmiblu
ugrin of ibe cefebellum on ibe curd, (hid Rrip on botli tide* of it, an lui
Eduitionnl evidence that llie ties of tbe cereoelluni with the *|>iDal ooixl nn
"not iDcrelj- for tb« purpose of servinj; as the diaou«l fi>r tlie impultca uf
DiUEculiu- Hn«e, but are tbe means by ubicb the cerebellum tmnnfunus or
elnburales nnawy impidBcs, uot of luuaculur »en»e alone or cii'wi\y, but
pmhably of all Rjods, in order lliat ibtv mny lako part in cerebral upvrn-
lioDs. of nhicli the oodrdioacion of buddy juovemenia may be one, but
probably is only one of Mveral or even of many.
SOUE OTBER AsrECTB OF TBE PcNmOXS OF TIIK BlUIIC.
It ia diffioult toaaT anythiii]^ duiiniu; concvrniti); tbo Iransiaission
Bury impulaea and the dt^vrlojmimt of »en»Htiuni; it is siill more
<Iifficiilt io»ay auytbingderiDiio, bcvond what ha« l)e«ii already iucidenlully
aaid, cunrcrniaj; tbe parut [ilayvfl in the work of lb« brain l>y the various
aggTvgntkMU ol' sray matter and tracts of fibraa forming the middle part uf
llie broiH, Neiilier ex])tirim«nt nor clinical study has as yet nSurded any
ckar or i>ure lading.
To what bas alivndy been nid abuut the oerebolluin, we may ad<l the
fidlonini^:
Elrctni.'al stimulaiion of the surliice of the cervbelluoi, in the monkey
ud in olhvr luiiniulo, has le^ to movenuints of the est*, and of oth«r parts
of lh« h«ad ; but we cannot from aucb results draw any satiafactuiy iu>
fyttaeta.
TUe removal of various part* of tlte cerebellum, t«pectally of the medium
|»r1a, has led to a want of coitrdioatioD in liudily moveiii«ula ; and an uu-
ateady ijait, due to a like waui of Rde(|Uat« cu&rdiuution. i> a frequent
syoijnoiu of cerebellar di»«'ase. But tbe inco&rdinaiion which makta its
appearanco inioiediately after reinuval uf, or injury to, tlie cercbolluni iitay
ervniually disaptiear, even l)i<iu>:h lar^e [>ortiun» have been removnl ; and
maay caves of exlen^ive cerebellar disease have been recordnt iu which the
noat perfect ciHtrdJ nation of ninvetuents was retained. Hcncn the ttsults of
expeniuenial auil clinical ntuily. while un the whole aupporting tbe godcIu-
•ion that tbe cerebellum has in w>iue way lo do nitb coSrdination, throw
little or iw lif^ht on the exact nature of the part which ibeiH^ui plays in
the Complex proveas, hut f>erha|H rather vbow thai we ore at pnaoat Hholly
Ignorant of buw cuordiuatiun is bmuKht altout.
Many liyjiutbeMS have be«u put ti>rnard as to the work carried out by
the c^ivlwllum, but none of iheMi can be ttaid to have an adct]uate buais.
Am), iixleed. if there be any value in the rvdections wc have repeatedly
made iu previuua ]>s^, the ^diysiulogist ought not to use the words "func*
liana of Iliv eerebellunt." Iruni a phyMolngical jiuint of view it is, so to
rk, • mattvr of accident, that vaiiuus alructurx^. the mla of rariotis
li'ilugical pruct-aiK'a, have, front mur{iliulouical causes, been gathered
togvilii-r into llic hiidy which ■ualonii*li> call the cerebellum. The task of
iIm; [•byatiiliitcixt is lo unravel the tim binding these various cerebellar striic-
turea with oilier parts uf the ceJilral nervous systwrn, and so nilh various
parts of the hudy at lar;ge.
We must cmtieiit oureclvea Iwro with calling altenlion to two or tlinio
a4l anil suggealive facts ooncarning its »truct«ire and oonoectious.
iu the first place, one striking fact about Uk> cerebellum is tbe very Inrgt
M
ma
THE BRAIN.
development of comniu«ural fibres eonnecting tocher tbe fuputfidAl giwj
matter uf tbe !«-» beinuphem for tlie grudter port of Ihcir pstent, aai!
pnwrig, not onlv lliri>ut:h the ponir (^ t>^) v pan of the niiddlA pnliineir,
bill nl«o tlinxign llie ruedian vcmiui. Thi* gnml vmamimaro it noond onlt
to lliC c'^^t nilliiral eoiiimiMurc (if tlu- cenbriim: iiml fna th» fiut tku
ine<li)in inioii* uf ilie rtrrebt^lluro, th<js« nliicb iln niiicrt dnnun to thi* can-
miwiire, &rc lb« niii»t I'tTei'livc in enuning incourltniilion hdcT iomd nwr*-
menM. m- may infer (biti U in wimc wmr pUya nn hoportsnt part In w
ordinntion.
A »rcnnil striking fiict i.- nne on which w« have tilrend<r juel dmlCllw
connoclion, ebioHv nii inicronuMl one, thmugh tho inferior peduDcte, with tho
atfcrent slrucliirm tif llie bulb and »piniiT e'lrd. Wc mny no« mdd, Ihu
th« fibres of tbi» pediinelr iitiming into lh« centn^ of the tvhiie malU-rnr
Iho coreliflbtr heniinphcre i>t llic «ini« side encluM' the gray matter or ife
niiclci» (leiitatus nail appeiir birgcly to end in that boily, though aomtjm
on to Ihe vcniiin.
A third Mriking fact '» tho connection, Ibis being, m far as we Itan*,
wholly a cnvscd one, thiniigb the pons nnd pee, wiih tbe eerehral (i<nu,
both of the cxlreiDe frunial region, and of the U-mporo ocei|iiiitl regi'xi. nd
putU'ibly or even prnbnbly with lunrc scatlervd curtieal elenit^nta nf iW
parietal (motor) region. This connecliDn is one between eortex and cixtn.
or ul least between cerebral cortex imd cerel>elliir 6ii])eflicial ^rny natur,
for the fibrce of ibe middle peduncle pacing fraia the gray matter of th(
piius which Hervcs na a lelay end in the surface of ihe lateral bemioplitrc nf
the eerebellum. The frcinliil citrlical tibrea pacing to the pea have ■ d»
Bcendiug degeneration, that is from ihe cortex l(> the poait, and wn mat
pnbubly a»iim« that the similar tetDjwro-occipital fibres similarly derviMnM
downward to the puos (^fi.lS). Fmm thia it huti bei-n iulVrred iliHl lim
oenrbro- cerebellar roniiertion carries iinpulves from the ecnittr.ll mrux W
Ihc ccrvbelluin; and it had biH'U t\)rlber iDlVrreil that ihvw ini|)ul*e« are rf
the nature of motor iinpiil:ii;4^ A* we bare more than oiico tirgt<d, lb*
character uf drgvoenition, Ihnt ia whether "axcending" nr " dcKy-oding," il
not n vutiKfactory priinf of the dirrctinn taki-n by impnl»c« ; but il it pw-
bnpa of more imprinaiicc tn rvmember thnt, ax webarealiu^urgcvl. we liarww
right to aiuiime thai the tiiijnilmTS pa.-«ing along nich a tract m the omIb
tineilioa muKt be eilluT st'iisury nr lunlor, or indiwd that auL'h a tract gem
as an instrument fxr prinbu'iiiK etfecl* in one direction only.
That during life tUc tibrc-- ot which wc an- tipeaking »crve w an impr-rfoM
chain by whicn cerebral enriot ami i-iTelwllnm aflpct ihono Ihe oibcr, lli»fi
can be but HUlc ibmbl; but me Bri.i whully in the dark as to what rvallf
takes place along [he MIikk. Wr have Mvn i § -''.'A ) reason to think that lk>
brain may and nwf exert an inbibiiory initiienec over Iho npinal roni ; wl
Ihe niechaoical certainty with which nii iinimal deprived uf ita ceretnl
hemisphere* re»ponils to slimiili. in contratl lo the untTrlainty attending \kt
result of Miniuli applied lo an intact animal, as welt as all lh« ex|ierienoe«f
our own daily life. «howa that the Cvrebral coriex can work iu an inhilWLMT
manner on other nans of l he brain ; the remarkable ** forc«d raitveroenlf
on which wc dwelt in a previous section teem in some iustaneea lu be (In
r«snlt of the abrupt trnap of some inhibitory bond. Cunvcraelr all then-
[wrienco of our daily life, many of ihe pheoomena of lh« contlitiun ki>o«B
as hypnotism and of allied conditions, as well as rari^us esperitDeotal n-
suliii Kuch SB thot ijuoled in !i t'll'i, where a sensory impulse aeenia to tnhibli
Ihe aelivily of a motor area, show that the cortex may itself in turn be in-
hibiied by other iiarls of ilie central nervouaavslem. Uul we have at |itMMil
no eatiafaclnry indications us Iu the paths of inhibitory iinpulMs or M to bn
ASPECTS OP THE PUKCTION'S OF THE BRAtK.
inhibition it brought about ; nor linvc trc any proof that the ccrebo-ccrebellnr
livct U ■» iubibitorj one in either flirection.
Wo may ailtl (hat tomn of the Hbm of th« niich]le iiediincic appear to he
Dritlwr colli niiim ml imr cnnnortod «ith the conical tibrc* in iImi pn, but tu
rnd in other «n_vii : nnil Irncln buve Ih-i-ii (IcoLTiht^ n* continuing onward
•DliM of the ccn-tirllnr tiliri* nf the iniil<il<> jMriliincIo on the oim- hniiil upuiinl
bimird ihe cen'bmni, nint <in the »lhvr liftin) il'>wnw«r<l lowaril ihn «]>i[i»l
cord. It ha« Itc^n rurilur nrgnl ihul lb*»c tracts nm cflerr'nt in functioti.
Lastly, wc msy ciill iittt-nliiin t« ihc supcHor peduncles. Ties?, which as
ir« hai,** tfvti apiM-nr to com* largely from the gmv matter of the ducIcu»
dentatd* ami lo cnti in ihe teemnitum. largely in rho red mictciis, mny be
regardwl a* cmi'tiluiing ihroiijiD f he reluy of die fninl pnrl of the (cKraenliiTii
another tie, presumiihly of adilTcreiit naliirr from the foregoing, between the
nTcbellurii and the cortex ; ind^-ed it iiFcd to bo •'n)le<) the [ir^iceaaiifi u cer«-
bcllo ad cerebrum. It is an obvioui'ty cr<»guHl tnicl (Fig. If I, •SV) ; i I con-
nects one nueleim dcnlatun, and to presumably by that relay the fibres of the
inferior peduncle ending in that boily, and perhaps other fibres uroceeiling
from the suiierficinl gray mutter of one side of the eerebellum, with the red
nucleus and other parts of the tegmentum of the crowed side, and thui with-
the corleit of the crome'l side. It has been siippi^seU that the direction of
inipuba pvsing atong it is from ihe cerebrum lo the cerebellum, but we
hare no clear proof of thia; indeed, as to what it doei, ti'e have no aatis-
foctory evidence either exfwrimenlal or clinicnl.
We may here incideiiliilly remarlc that, in cio.tequence of alTcreiit tmcts
being trace<l to or toward the tegmentum and of the shiirp contrii»t pro*
•HilM betwc«u the tejfmemum and the ooD.tpiciiouMy nl'l^t^ pvramidal tract
in the pea. the vien' has ^ined ground that the tegnientiim u OMDtiallr n
•maory stru^tun^ lint there does not ap]irur to be iidei]uaie evidence eitner
cliiiiail or esiH-rimi-ntid for iiiich a cjorloMoii, The (balamus, which n'o
bnvv rvgnnled nt ibc front xo lo epenk of the tegmentum, euniiDt. as we have
already urge<J (SH«i), In- roniirU-m] cxcliiaively or especially sensory. And
many of the lie« of the tegmentum, such ns the nbrva from the corpora striata
ending in tW suUtantiii nigrn, fir (bin may be considered as piMjieriy belong-
ing to the tegmentum, are of the kind which we may suppose to be eSereni
or motor. Indeed we may pnibnbly reganl the wbow tegmentum as beioz
broadly the aiialogue in the forward acgmenta of the cerebro spinal axb m
butb tlie anterior and posterior gray matter of the spinal segments behind.
■ TboUEh we are thus in the dark coneerning what goes on in the cerebellum,
UtBUty M Worth while to call attetition onoe uion> to the remarkable char-
Ipetenof the superlieial gray matter ($049). The many [i»inu of rvM-m-
pUanoe between it and the eerebrAl cortex cannot but aujjgcit that th(< i>r\i-
Dtttat taking place ia it have aume analogies with cortical event*. And it is
at least a fact of annie significance that congenital deficiency, or etrojihy of
ttie cerebral heuii^pliere of one side, U fr^]uently ac<rompa»icd by a C«irrc-
■powiiDK delii-iituoy of the ef<i«sed ci'rebellur lieinMphere.
(688. Both the anterior and nnslerior <.'iir{>oraijuadri;{i^niina arc complos
tn atfucture; not nnly do ihi^y djSer from each other, but bIki in each the
grvv matter dilfen in dirfprmt pnrta, Ixith an lo its nature and nppeuraiiea
■na as to it* connci'lioaM witli tracts of Hbnat. If we have little right to
•peak of Ihe " functions of the cerebtrllum," wo have even Iwa right to speak
Of the •' fuactioiis of ihe corpora ((uadrigeminB" or of either pair of them.
The anterior pair, as wo have seen, ba* tu do in s»me way with vUion ; but
we have rranm to think that a part only of the whole b»<ly is thus concerned ;
and there is some foiindilion for the view that of thi4 pirt, ooe portion
balungs, ao to sfieak, to the optic tra^t an I another porltou to tlie ooftioal
I
668
TMK BtCAlN.
fibre* of tli« optic railUtion. Pomblv «lill aiMtber (Mit b coacwoed in
liriiigii));, ns we have ($ G7-1) eug^CBlcu, Ttniftl impulan to bear on the iw-
vnliiiHlion of itioveiii«iiu.
Sliiniilatioii of tlio Hurfnce of the poslcrior pair, boido giviDg ria* U
inovi-nii-Dt» of viiriouit |iuriB of Ihe bwly, baa in nrnnkeyB KM warn olhtr
piiinml.4, the siiijfiilur dlcct of prirxtudnK a vocal utteraDc« in tht (bran of ■
cry or b&rk. HuL ne oaniiot timke tnuch u&e of tbeee rcsulu tor tb« par
pots of ilniwitj); condusiiiiw bs lo nhat aliare ibme bodia Utko in tbr wliok
work of iht- hrnrii. In (be fn>x, ibe oiilic lobts correspond to tbe two pair
of oorponi iiiitiilri^uiiim l<)i;k.'thi-r ; una iho cry jufit mentioned majr perfatfi
be put side l>y fide with thv fuel tliat in tbe fro^ tbe optic iob» Memi*
turaioh a mecbanisni for cniukin^ ; nhen tbe optic lobes sr« deelroj«iI, tbt
n-flex viouking mrutioncd in ^ 6^il in done away witb. Tbe probable cowmc-
tioD of tbe piwlcrior citrjNTu (luudrigeniinn nith bearing b alao intereitiDg b
tbi* cnuui-ctirin; btil we have no MtiiKluciury evidence of iiajr special \M
bctnccn tbu liodicK in ijurrtion and cilhrr tlie coniral urea fur pbonMuaur
the viieal mfdniniARi in general ; tbe ovcurreiiuc of tbe cry remattu so turn
i»(ilntLil fact.
In frogm in which the cerebellum is very (mall, iho nptie lnbn> seota to Is
jiarlicLiinrly cnnccnicd in the coonlinniion of movcnwrnn. When tbebiib
ts rrmoviil by nimns of a section behind the optic lobes thv nnimnl loss tltf
|>uwer of balancing ilsclf ($U^U), which it ixwmwcm when tbe scctinn psiw
m front of the oplic lobes ; and injury to the optic lobes produce* innWiHitn-
tion of movement und iil\on " forced movcnH-nl*." It biis Ix-en luaiiiuiaal
ibnt ibe Iocs of coordinalion is in these cafes doc to remorai of or injarjli
the central gray molter in llie walls of ihc third veDiricle, and oat ts
niertr removal of or injury to the oftic lobes ; but the irhole evideocs p**
to abuw that in the fro;- and in tbe bird the opiio lobea d» play a pnri ia ibi
co{ir«)i nation of movement, though leniuus of ilie centml ^ray matter aroni
the third ventricle, or indeed of the thalamus or other parta of the i«p
lueiitum, may give rine to loss of coOnlinution or to " forced moveiocnti.''
In ilie mammal removal of or injury to tbe posterior corfwra i]u>dii-
j^miDa doeB not cau>c biindnew, but may, like a lesion of tbv nuicrior pai>.
give rise to loiut of ooOnlinulionor to foroeil moveaients; the edwot, bowcnr.
14 in niont inKiuucai very tcmponiry. The voiiueotioD of tfa* anteriot mil
with virion suggests a clue n> to how this pair tnkfi* part In cofirdtnUM*:
but us to huw the posterior pair could inlvrvcnein the mutter wo havobanfif
»o much ns a hint; fur, even if viif nilmii u eonneciion between ibcot ni
the senn: of hearing, and, rem<-mbiTitig ihiit a loud souud will on«n mm%
])e»»ii to rtvl, further mlmit that pun-ly auditory iinptilKV, lu diattnci tmrn
what wc have called am|>ullar impulM^, may lake ]mn in the ^Dcrale»
ordination of bodily movements and in the maintcnauoc of •qutlibriunmi
tbcy certainly do tn the ei>ecinl coordination of laryngeal move nients, (till
neare not miicb nearer un undcntttnding uf tbe matter. Wo mayaiMtyi
wcliiiD of the lateral lilkt, which appears as a consfncuous tiv beiW«M Us
|Hnteri<>r corpora qiiudrigeniina and the parts of llie nervous systom btbM
them, does not appear lu have any markeil cHect in producing inoo>>n]iliall|L
Id fine, beyond tlie broad facts on which wo dicell in a previonsseed^H
nanicly, that we muiuiain our equilibrium and carry out complex ridvcmmV
involving otlen several pnrlH of our bu<)y, lhroij);h what He call eodrdinatioa.
that allercnt impubes supply important faolors of this coord i ua tiuo, aoJ
thai the ccrcbcllniu. tbruiif^h the Tcsiiliiilar nervee in part at all ovtni*.
together with other portions of tbe middle brain, ar« in some itny it* cbNf
instrumcois, we as yel know very little. We have certainly uu adniualt
kiiowled^ OS to bow eitlter pair of corpora <iuadrigeminu axooUy ini
I
I
ASPECTS OP THE FUKCTIOlfS OF THE BRAIX.
in ihc mailer, or, indeed, as lo what other pirU thef play in the eeneral
work of the bntin.
With regard to other Imcu of Gbres or areu of f-mr mntior we have
nothiliK to my, except at rv^Hnli* tUnee which arv mora or lodi iinmediately
oMtDected uilh eertjuo of liie criuiiid tiervM. >ucli fur InslaDoe as the nurvea
for moTenionUof tbecye»,Hn<l tlieieit will be beat to coosider vrbea we have
to deal with the oervat tbemi<i>lv«*L
§690. Bt«i<lei the »oinatic ftinctionf which In prerioua dIacuMlnus nr«
have chi<-Ay had in view, thu liniiii lu a wboli- undoubtedly carriee out
aplandinic functioDH ; coiicernioi; lliMo, liowuver. vin miut be very brief.
Of the rttpiralory and vuoiDotor fnuctioiu of tbc bulb we have already
tmtrd (d (heir ap)ir»priulr place*, and we. have referKi) ($53l{) to the
experiinrntnl evtdriici^ iliiit n ln<ii>R of ihu ciirpiiH striatuni, or of the front
part uf tUi! oplic tlinlamtix luui n minnrknbli; infliivuce un tbii development
fif heat in tbe body, Wu hiivv furllmr »vv.n thiil thi- higher pnrU of the
brain, acllng through the bulb, cxprci*!- powerful iii(1iii--nci!< on nvpinilion, I
■la lli« vaAomotor syntem, nn I on iho bciil of tbc hcrtrl. Duly f):pcnL-uce
alT-trd! abundant in«t«ncc«iif nclinn* ench m those, iv well luof the inllurnce
of the brain on other organic fiinclions. We can brinj; our will to bi-ar on
tbe OKebaniun of micluritlon (§431) which i* almost wbnlly, nn<t on the
meobaDKin of defec«liun (S 271!) which is lar;gcly sjilnndinii- in nature.
Tticae movements, however, nn> nol skilled niovcmiintx : nml iw wu cxplntunt
ill dealing; witli them, the aciioo of the brain a.* rcgnrils them ancois liiiiit«<l
lo uii;;nteniiH;r or inhibilinj; the activitv of spiiiid centre*. Wcshould, thera>
fore, hardly expect them to be s|>ecially represented in the cortic-il motor
r^:M>n. But emutioua have a mii(.'h wider and more powerful tnHuonoe over |
the Rphuiclinic functions tlian has ttie will, and hare tbe poirer of affecting
thu Work of certain orjtanj, fur instance the heart and secreUng glands, which '
the will is umihic to toudi. And since ive have ever>' resson to believe that
the corlex is chuxrly aMor:iate<) with the emotions, we mav naturally infer
that elements of the curtex ■ii]i|)lr a link it) the chain thr»u;;h which an
emolioB intlucnm this or that splanchnic activity; we may, accorilinttty,
extieet to find that stimulation of some pnrt or ■■lUer of the cortex produces
»plaiichnic ctfecis. The riwiilts of cxpcrinu'nlal inveittgiitien. however, are |
Itoih scanty and difconlnnt ; but the greater weight should {>erbai)s be
attached to the positive rcnults. Thus, some obwrrvom find that atimulation
of tbe corlex, the locality being in the dut sonwr part of the siiimoiil jtyras, ,
priMlucen movements of the bladder ; and tncy trace the path of thin iullueuce
ibmufch tbe front rart of the thalamus and the tcgmcntam to the bulb and
aa In tbe curd, exuludin;; the oeretMllum, whiih other observent believoil to
be conceme«l in the mailer. Some observers a»iH find that stimulation of
the («irtex produces a tlow of " chorda saliva,' while others maintain that
ibe Mcreiiun, when it does occur, is an indirect and not a direct efloot of tlie
cortical stimulation ; and it may be remarked that the oortical area, which
iactainml to be a " salivalt>m area." IvIuk in the dnj; on Iho coavolntions
iloraal to and in front of the Sylvian Wure. is not either the area eonoecied
with the iiscial nerve, or that allotted lo lanie or smell.
i^milnrly, xtimulation of paru of tbe eortex has in the hands of various
obwtvcrs kxl to movemcntB or to arreat of movements of the inte«tiRa«, to
cfaaiuea in the h«*t of tin- Iwart. and to various vasomotor and other eflecla ;
hut It will not be profiiabh? to enter into an^ further detuiU. Wo ntajr.
however, add the remark that wliea the oorticul mutur area for a limb \i
reniove'l, or sutTcnt a Iwion. tbc temporary paralysis which is thereby
dosed is acompanicil by a ri*c '>f lctn|Wrature in the limb : Mf may be at
very great indeed ; in the moukvy tor {mtanoe, tbe band or fool on
B70
TBE BBAIN.
Ihr para1yz«<) oitlc may be a* inucli as 10" C. higher than (hut nf ibe i
»iiio. The cfTfi't
nnlv <lt
.Ily
riworoutor raralyviv, l>u(. osfMTial
fiilering that ibr niii»cli-( of the limb nrc rclatircly rjuitnivtil mikI *u pro-
ducing ivM lioiit ihiiii uKiml, ejiniiot Itf tiuti to thai ilonc The nrnmrhablt
rc-eiill niny lie luki-ii ii« "till fiitthcr iilu#iriuint! tint cwnplcxily of thv prw
ccwn Coiini.'d<-il v> itb Iho cgnicnl motor nn-n ; lh« nrrsi is in t>i)iii« way umt-
Gtntcd with ih<- vn^culnr nrriiii^omcnbi mid outHtiun of (he initM-lea «iik
who*o Diovcnitnl* il in CDncrructl.
S 691. Then: irmntn yet a few nords to tw Mid about the trortcx. Wi
rrgurd, and JLiHtly so, ihi; Hp)>niaucotin intriiigic iwlivitr of the brain an ihr
nio»t tlriliin); f^^iilurc of its life 'T\w nenrest npjiroac}) to it nhiiih are SnJ
ebewbero in the body, is perhaps the rhylhniii- Ireal of ibc heart. Tlir
analogy between the "regular aiilomationi" of the oiie, and the " irrwalar
autonialisRi '' of the othifr ia a Elrikin^ one; and iiideot our kno»lt(^«f
the relalively ttimple sponlaDeiLy of llie henri haa probably iDdueiired to ■
larse e.iteni our conceplions of ihe complex sjxHilaDeity of the bnuu. b
the heart the rhyihniic didclmrga of energy a chielly deternuiied by in-
trinuo cheniicnl clianges, hy the luciabiiliam of tbe cnnlioc- MilwiAnoe; ilw
influence of exIerDal circunutHiicea. ajwri fmni tboae which pnivida aa
adequate supply of pro|icr blond, \» wholly subsidiary aiu) aervm only M
roJse or to lower tbe intrinaic d>un;;e« from time to time, a* nccaiiroo ma
demand. And ihe analogy of the heart btu pvrhapa led us In cxwga«Mt
the part played in the bruin by itie like intrinKic cheinicAl tiwiamiMh
(We are liere, nf cnunw, viewing the at-iion of the brain from iba Mir
ataDdpiiiiit iidnii.i»i1ilc in llirae pag<'>, the iiun:ly phytii>lo«ical oih> ; bat wea
a mono of trt-iUuK'ul iim-» not prejudge olli<-r puinls of view.) Souk wrikn
use czpreMionii wliich i^ceRi to implv llin coiiccplioD that tbe iMrvoui
diaoges forming the btisis of tbe perchi(.-tt] and other procffiaes of llie brato
lire chicHy the direct outcome of the cbcraicnl mclaboliaiii of iho gray
mutter and e^pecinlly of the nerve ccIIh. They speak of " the diacharga m
energy" from these c«IU in the taiue wny that we can speak of ibe dil-
charge of energy from a cardiac fibre. But, to say nolhiitg of ibe luw rait
of oervotia metabolism as nienaurecl in terms of chemical eitergy. we bar*
no exnerimentni or other evidence uf nervous atibstauce in any part of th*
body neiiiji. like the cunliao substance, the seat of an important metabuli»
carried ou irrei|jei:tive of influences other than purely nulritive uuea. la
ibe case of nerve-cells interpolated along nerves cou)|>u4ed uf fibru* of iht
Mime kind, as in the sporadic ganglia, nil tbe instances where tlw nfrrc-tvUt
were supposed to initiate active jiriK-t&ies have, ou exaiuin.itioH, brokea
donu ; an wu have seen, the ganglia of tbe heart do not aupply tiia moving
cauite uf the heart Iteiit. It i!> only in the wiitral nerroiu mt«in whsi?
iiGrvei'^IU. a> |iart of |;ray niniter, are found at the meeting of utm-^r»
of ditrcreni kinds, that we have any evidi^nce uf " discharge of antrgy*
fruin the cl-Us-
Aa wtr |H>inteil nut (SAd^t) in speaking of tlie spinal cord, ibe ilischargt
of efferent impiiU«« fn>m the central iivrvoon AVEttvm. though it uodouM-
tdly must have a u^rtain chrmicAl bn»i*, nniiiely, tin; metabolism of th«
ner\'ou.i subntaiicc. in, in ilic fiwt lin", dei>pndwit <>a the itilvrnt of alfenal
impulses. Hut this, if true of the i>pinal cord, is i>till m<irv true of ibt
hrsin, wbiuh receivt« or may receive not only ell the impulses which reach
It through the cor^l. but cspecinliy [Kitcnt ond varied impulses iltnctlt
through the cranial nerves. All lifo long tbs Dcvcr-ncirang ohangea of tht
external world continually break us wavca on the peripheral eodings i>f Uis
afferent nerves, all life long nervous impulses, iion- more now fewer, are o«-
liuunily sweeping inward toward the centre ; and the nervous iuetab<'j!Uiii.
OTHSB ASrSCTS 07 TBI FUNCTIOXS OF TUB BRAIN, till
P
I
which ill the bndi of Dervou* ftctloD, tnuBt I>e at ]«aat u t&rjtoly depeDilent i^n
tb«w iiiDii«aca from irllhout, an on the lueru «beiiiioaI iiii|)]ilj' runmhiil by
the IiIohI.
We hiive «Icvel<>|ieil thii point becauM uf the iuflueoce it uusl have on
our coiMWpliMU of the (ihyritilneiml |)rucV!Wi.ii tttkiD^ iiIhcc Id the coflex.
If we tMKCpt Htv vivM jiut luid (liiwii, wv miint n-'jfiiril the *u|iereRiiiii.'Til
KClivitj of the corlox mid the churiii.'tcr.'' uf the proi^vMoa lukiiij^ place in it
«■ clui" uui lui much to itic iplriiiKic clK'niii-nl iiiiLiire nf iht' iiitvduh xub'litiK'O
n hi<'h i* built up into th<^ corlicnl gray inalter hh to the t'lit't that iniputx^
sre ctinlinuallj' itmmiug into it from «ll pint* of ihu hmly, ihnt alm><^E all
itiflucnco bnmgt to U^ar on ihe body mnkr ihi-miM-lvi* firlt hy it. To put
the tuntter in a ImiIiI vny w«i mity itek iho qiientioii. Whiit would Itappt-n in
lb« cortex if, iU urilinjirr nutritive dupiilv rcraniiiin); a» hi-fore, il wen? cut
wlrifl from nffercnt impulses of nil kimU? We run Inirdly doubt but lliat
vuliliiioml ami other [Mychical pn>e>«a'« would to»n ootoc to a atnndstill and
OonseiouMieM rnnHh. This is, iiidveil. roughly iiidienlcl by the remnrlcuble
CMC of a patient, whose iilmott only eomuiuiiictilinn with the estcmal nurld
wu by means of one eye, he beiog b'iii<l of the other eyo, deaf of both nan,
•ad Mflering from seiiernl niueethesin. Whenever the sound eye waa cloMd,
)w mat to sleep. It is further iiidireetly itlustimted hy che fonowiD;^ ex-
parimefltal reault. Wo have seen (!;(Sd.^) that a vertical inclsiun carried
tltroueh the depth of the },'ruy matter around nn area does not prevent
■tioiulation of toe surfaee of the area pnNliicing the usual movemeota.
Bat after aucb an iaeiaion the animal suHVrit s parnlysit of the Oioveiuoilla
ooonecied with the area, lik« that leaultiu^ fr>iiu the remural of the gray
mftlter of the area : aud the u|>«raii<>n i« said to be fulliiw(>d hy d>>>^uera-
live changn in the area, and dei;eueraiion of the pyrainiilAl libKtt «tar:iii^
from iL Home of thia eOcvl may lie due to nulrilive chan);i^ brought about
by iniury to th« pia mat«r and iliviriinn of bluodres«elN ; l)iit it cknniil be
wholly accountnl for in this way; it ap|>ear* aa if the life of the area il
euriailed whim ila nervous tin* arc broken.
We may conclude then that we nrv not juslilicd in s|X!nki»j{ of con-
•ciouancM or volition, or other iMychical prucuatui, even admitting mat tlieM
fail when the curtei \i removc<l, as being functioiu of the cortex in th« saute
■■y that we speak of the fuiictionii of other organ* : they arc mtUcr func-
UoM of the ooanecliniis of the cortex with the other parta of the coniral
nervous syatera.
We altould add that tliey are also functions of the oonucetions of the
several parts of the cortex with each other, All our knowledge goes to
allow ihat psychical pruceMea are dejteniient on, or are in eonie way aMo-
Otated with the cortex; but whatever c I nasi li cation of psychical fuuctious
w* adopt, w« are wholly unable to make out any localiution of functioni,
such aa we can make out for in •vemenla. visual len^ations and the like.
Even taking the broad aud elemeuiary division into " the emotions " and
" the intellect," ne canuut uitiafactorily allot eitlier diviuoa to any pariiuu-
lar |>art of the heoiiaphere. In d<i^. removal of [wrticular part* of tli«
bemiiipiiere* has iiKlced been i>b«erred to cbati^ce the character of the ani-
mal, cuuvcrtiiig for inilaiK'c a vicious, moriwe dog into a mild and iuiA'en-
aive one; aiul removal of the front part* of the beiuiipbcn; neenui to have
frei|ucutly a marked ctH-ct in reoderinx the animal mure improatioiutble artd
excitable; he Wcoaiut much more deiaoaitnitive and "gu»hinK" in his
behavior than Iwforv. But ihcM are mere hinbt, and the clinical hiatnriea
of dbacasc in man do not enable lU to snr much more. Such knowleilge aa
we (to poMoa latlwr tends to ahuw that tnc piycbical procwna in proportion
as they beoooN more complex involve a greolar Buntwr of nwvoua tactora,
THE BRAiy.
and, iherefbre, hare for tbdr miiterinl btuls a frreat«r wultli nf ucrvoin arat
nr in other wonb their lucnliuilidu bvoomM laM ileTiniu. Tbia while ws
may localiie tlic iK^'inniuj^ of h pfycliical procea*, a rinual Knntioa (ur
imiance, aod ont; of ibi U-rminal nuia auch a* the inue of Impubea sinntg
the |iyramitUI tract, we cnniioi put our finger »R the Mat of the iol«riiM-
<lint« tmii»actiou)i. Tht*e. Mvn in the HirnpWl procnM* inu«t b« o>i(n|i)ei.
aixl mii«l involve many rndiirs. Our timnltvl conceptions of the external
wnrli) are bumvl nii u cuinbinntion of vtfiinl trnmtiinx ami tn^lile aeiwatinQi.
It being granted that the vinual nunntion, in onti phase of iu developQWt,
i« conneotei] with <H-nnin ctuuiKcs in some spot of tti« ocdpjtal cort«s. thi<n
mtiat be mmc tif iNMnfcn this nod the corrMponding nvrmtM aeat of iW
tMtlle aemalioD wherever that may W, and further tU» between theae >«4
nther pans of the corU^x. Hence, m we said, the peychical pnniM b a
fimrllon of rnnncelions.
Many nf i\HfV. iiv* arc moft pmhalOy fiimishet) by the anoriation fibni
paMine from one part of the oorC«x U> a nei);>il)»ring purl. We muat alM
proluihly admit that impulses or to nsv a more general word, prooewn. may
travel laterally along the tangle of the corlieal gray mnlter, for ihi*. lin
the gray mnttcr of the spinal oord, seems to form a phyaiologicAl conlinoily,
DO more broken by the l)<eum than is the cord by ita segmental arniitc*>
meot ; and we know nothing a> to the limits whieh luUBt be plae«d on il»
distaDoe to whieh puch prooeases may travel from their focus of origiH,
Further, eeeiug how completely In the dark we are as to the reason why M
possess two hemisphere, and eopeeiaUy seeing that, as shown by speech, tte
whole of ench hemisphere h out identical in action with the whole nf lb
other, we may perhaps suppose that the fibn» of the corpus catloetini, wiAk
form BO large a part of the central white matter ol' the bemisphere^ ban
other duties than that of merely keeping the poinlii of one benil*phei«b
touch with the et>rrMii»Tiiltng poinln of tho other hemiupberi'. Ritt, wbca
we hnvir niiule rvery allownncir for all tho*e direct int<irc»rtiiiil coonecliuafc
wc arc drivt'n Id llx! cimc1u.->iiin that the in<lirc('t tie* betwvcn one part (tf
the cortex and ani>lhi-r thmugh the lower parts of the brain an) of no Isi^
pcrhapa of grentrr importJiiici^ Thi*, indn-it, ii> shown by the n-ln:i<iat t4
the motor region. We hnve already ur^d, that even as re^rds the nivie
carrying out of a skilloil movemt^nt (and we may add whether that b^^ rnl-
untary or iiiroliintnrr in the orJinnr}' eommon line of the words) tW n>ni*<
region must have olhcr lies with the (lart moved than merely ibo rfli'fMa
tie of the pyramidnl tibres; it muxt have scntory alTercnt ties, and the
course of these, including even perhaps those which belong to the mitseular
aense, we mav regard as an indirect Mie along the spinal cord and middle
parts of the brain, though the details at* as yet onknowa to us. It aiuil,
moreov-er, as we have also seen, have ties, at least in many cams, with parts
other than the part moved, for instance with the geuersl coonlioaliM
maehiiiery. And the ea>e with which some, not very ohviniie. chani;e. wiu
|>ermit the siimidation of a limited iriolor area to ^tart epilcptifiriti (.■oiirul-
mous. shows how ninoy and close are the ties in another direction- Further,
when we go beyond the final phases of the process in the motor cortex, to
tbiMe which precede the issue of the eSttreuL iniputsea, we find the ties mal-
tiplyiiig. For instance, since our mnvemenis an* so largely guideil by visual
BeoaatioDft, there mu»t he tiea between the mul^tr cortex and (he central
visual ap[>anitii«, it may he of the occipital cortex, but it may also be of
the lower rinual centn«. An we ituialeil. the motor are« is only a link in a
complex chain; and what we can see. dimly though It be, iu ref«rvui.'« lo
tlic oortical motor prooeves, probably holds good for those other n>Tlk«l
piMCMM as well, of vhose nervous g«nQ»is we know at jtrescut nothlag.
THK TtllE TAKEN UP RV CEREBRAL 0PEBAT10N8. 87S
I
I
I
I
ertn ibe higher iwyebtcal ereoto rannot truly be tpoken of "» fimo-
of the oori«x, ntuiiiig that they are 'irnpiy tlio outoome of nolcculRr
diugea in the OMtio*! gray matter ; thoy are r&lhor to be it^rded a» the
onteoae of eomplex pncMMa in which the parts of the brnia belnw lh«
o«niM pl«y a part no leaa iraportaat thaa that of ih* cnriex it«e1f. If 90,
the fibraa pMsio^ down from the eurbex to tlte mi'ldt? brain hare fbactJoiiB
by wliioh Ibey take part even in our paydiical life. fuDctiona for which
neither the words motor or wnaory are filtiog.
0» THK TtMK TAKKX I'P KY CkRKRRAI. OpEBATlOirs.
§ 698. We have nlrradv rttn ($ ASS) ihnt • cmi*iH«rnb1p liaie ia Inkcn up
ID n purely rv6«x act, nach an thiit nf wirikiiiif, though thi« is perluips the
moat rapid fnrmof n-fl«x nvivcnictit. WhcJi thr movtMnODl which ifl executed
in mimnae to n ftimiilu* invilvn cerebral npcrationit a «liH lonfcer time ts
needed ; and th« inlprvnl hclween the npplii'niiiin of the xlimulua and the
rnmnH-iicemciit of ih*- tnnoctilar coiilmction varin according to the natui? of
the mental labor involved.
The Mniple*! case in that in which n person makea a msnal immediately
that bi> p^rreivi:^ n ^tiniutiis— rj^. ^r., cloeefl or opens a f^tVaniC circuit tite
moRMtnt that he feels an iDttnctinn-sbock applied to ihe^kin, or (r«9 a l!a*h
of light, or bears a Bound. Ily arrangements similar to those cmplnriHl in
Rieiuutini; the velocity of nervous impulnee. the moment of the application
of Ifar Ktimuhia and the moment of the making of the Hi^Dal are both rveorded
on the Mine travelling surliiee. and the interval between them is carefidly
nmearrd. This interval, which has been called " the reaction period " or
"ntaeiion time," may be divided into three otagea: 1. The lime during
which afltrent impuiiM are generateit in the iieriphcral Muse organs and
transmitted along the iilfi^riMil nervca to the central nervoun nvvtem ; thb
may he ralle<l the "afTereiit stagv." 2. The lime during ithioh, through
the operations of the central nervous systctn. the affiireni imiMilio) are imn*-
formed into efferent inipu|jw»: this mnv he called the "central atage.**
3. Tbe tiiiiv taken tip hy ihv [isaMgc nt the elTi-rent tnipulacw along iha
aB^ot nervni uud tbe t ran* format inn of the nervoiu impulMv into muiicular
eoatractioiis ; t)ii« may lie called the " cflV-rnit Htage." Tn the rflert-Dl slaiire
the eveata are omiwnttivclr iiimpli'. and though not ahaotutelv e<in«tant, do
Dot vary largely ; nc an.- aide to furni a fairly tntitfnctorv citininte of its
duraiiitn, and m of the i>hnrc in llu- nhnlc nmetion period which may be
allottrd to iu The ovi'oIk n[ Lhe uflfri'nl ftage an? much more comples. and
tbe calimatea of ila dumtion, Ix-ing arrived nt in an indirect manner nod
chiefly baaed upon cah^latJont of tbe whole reaction time, are very unoer-
Uin. Henoe all atieiuptii to estimate the length of the " central " stage, the
"reduced mnction period," as it i* sometioiM called, by subtracting the
eScreot and aflrrenl staifca, must be rabject to mnch error. But a good deal
maybe Iranird by otudying the variationa under diflereat circoinsiaaoes of
tbe reaction iH-riml lut a whole.
Taking lirat of all tbn cases in which the eventa of tbe oeairal stage are
aim pic, such as tbnse where the xuhject baa merely to make a rignal uyon
fheling a MDsatian, we fin<l ihiit the length of tbe reaction period ■ depenatnt
OD iba {nlMHity of the stimulus, being shorter with tbe stronger stimultia.
But variations in the strvngth of the stiuiiilus. npectally in tbe nuei of
minimal etinnuli, have a much more striking eKcct iu determining the ivr-
tainlv nf the reaction than in nlfcctiug the laagth of llie period. Thus, when
the signal is made in r»|Mn>e to some visual Beoutiou, ujiou seeing an elec-
874
TQg BBAiy.
trie eimrk. for iiuUnoe, If tlic apark be a very weak one the subject of tke
experiment often fails lo make Ibe lipial iil all, tbougb be luay nireljr GUI it
(he ajwrk be a Ntron^ inie.
Suniv uf tbe in<ict mnrkcil viiriatioTw in tfae lenirlh of rhe re«c-tii>n (leriad
arc (lettTiiiinnl by ibe individualttv or thc< nubjonL Tliu«, vritb them*
6ltmiilii« ii{)(ili<'!'l iimlt-r thi' luinir circiiinnliinn*, ihe naoiiun perimloTMi*
|icr*rin will W fonml vrrv ilifTcrr'nt fma) tbiil iif UDOllier.
The IciigOi of the rcnclinn pcrim) vnriiv u\mi acv>r>liiiji t<> the luiturc lad
(lisfKMiiti'in or thr pcrifihrriil ort^iim »timtilittril. In i;i-ni>nil, il inny be mi4
ihal nitnm-ritip srriKatiotis priHliKiil by tbc :>iimiibi« of aii i-jivirii- *hiui
tLT>pli<''i l'> the »ltin i the »i^al, for inolnnoe, bt-ing mnilo by tbi- rijiht liuij
when tbp »Unc'k i» iHt by ihu k-ft hnml) nre followcii by a aborttT reariinn
prri<Hl tbnii nre auditory ernsnti'ins, whik- tbc period of tbne w ia tura
»b'>rirT than thnt of visual fLiitni inng pmrliict^it br luminous ol^ect*: on ih*
otbiT band, the shortest period of all i« aid to W Ibat of vImikI a^niialiaM
produced by direct electrical stimultilion of the retina. Itoughly Mimikinir,
w« may say thnt the Tcactiuii |>eriod is for cutaneous srnntwns oii»s*VMtii,
Cir hearing one sixth, and for ei^hl one-firth of n SMond.
Practice materially fihortene the reaction pcrind : indeed, after Xnog jvto-
lieo, makinff (lie aisiial. nt lir«t a distinct effirl of the vill, takes on (he char-
acters of a reBei act. with a corresponding;! r shortened interval. Ltially, tt
mar add that in the name individual and with the same stimulus, the toaylli
uf tlie period will varr actrurdin); to circu nidloncea, Mich as the time of year,
the weatber, and the like, aii well as accordiiif; lo the condition of the iacli-
vtdual, whether fresh or fatigued, fasting or replete, having taken more«t
leas alcohol, and the like.
The reaction [leriiwl of vision has long been known to astronomen. ll
iniB early found that nheii two oWrvem were watching the ajipranowt^
the ><am« star a con^idrrnbU' di-«-n^]>ancy i'xinli-<l l>etween lh«ir reapeOlTs
Tviiction pcHoilii. and th;ii. (he iiilf<'rcnc<-, forming the basis of tht KhoUM
" [leriHiiuil CQuntion." vnncd from tim« lo lime. According to Ifaa paftoml
conditions of (he observers.
§693. The events takini: place in tbeecntmlatagcarcnf courM,comp(tz,
and this stage mar be suhdividol into scvemi sti^ca. Without attrnpiiif
to enter into psychological ijocsiiong, we Diay at least rMogtitw oeriaia
elementarr distinctions. The adcrent impuImM started by the stimtiliis, wh•^
ever be their nature, when tbey r«ich the central no rvmts system undtrgs
changea. and, as n-e havcseen. probably compl«K changes. bci>>rotb*)y beoomt
aeiisatiouH; and further changes, now of a moredistintSy pavehicat rnaraelar.
are neceatar}- liefore the mind can duly approciale the cfMravU-rs of Uwa
senaaiioiiB and act acconlingty. Then come the psychical procuasca thnugb
which thetie ajipreciulcd sensations, or perceptions, or anpercepliona, as ihay
an eometimoa called, dclcrmino an act of volition. LMtly, there are thr
executive prooenee ■>f volitiou. the procemes which, payfillimi to bef in vitb.
end in the imie of cocirdinate motor impulsea, or, ia other words, start ili*
dUtiiictlv phvalolni-ical prucesBCfl of the efferent Mage. We (tiay ibiis sptok
of tlu! tictiL- re<|tiiri'd fur the [lerceplion of thestimulatJOD. of the time rei|uircd
for lb(? action of the will, aud of the time required for iIk oomplex iiaydiical
pmccKtM which link tlieee two together. Accepting this elementary atialyii*.
It is obvious that the totjil length of the central stage may be variod by itif>
ftircucM in the length of each of tbeae parts : and a more oompleto anolystt
would, of course. o[>eu the way for further ditttiuctions. Heoee, byattHlTbg
ihe variatiuiu uf the whole reaction time under varying forms or psycfuou
arlivity, we may form lui estimate of linic taken up by various ]isychital
processes.
I
I
THE TIMS TACBK UP BT CEBBBRAL OrBRATIO.VB. 875
Wr mn;' tnkc as nn intlsnce Ihe nt<« in vhich the wibjpct of tho i>xficri>
menl hni to Pi<rcu4> iliKriniinntion. ']'h« mode nf making tbo Bignitl Iwiitf;
tlie Hini«, and lh6 stimulus bcinf; of ihe Mine order id each trial — that \« to
Mijr, viMial, or culaneoira, or auditory. i-K;.— nmi getieral rirciimstiinc«
RnMBing Ute hidp, two diff«rent tlimuli are employed, and the subject v
required to mak^ a ^i^finl in respoiwe to the one stimulus, but not tn the
other; (he lubJeiM has li> diteriininate between the payebical elfecti of the
two stimuli. Suppose, r»r example, the flltmulua i> the sound of a spokeo or
sunt; vowel, and the subjecl is required lu make a siKual wlieo a is epokefl or
ranii. but imI uheii o is b|)oken or sung. If lli« subfeot's whole reaction
perioa) l»e deleriniiiet) (1) in tlve usual ttav, with riihLT a or«s|>oken (and
tbn result will be ftiund uul lo differ miit^rially nhrthcr n ur o be u»ed>, the
■vfajed knowing that imlr <i «r o»It o will be g^Mik^n, and then be determined
ajliiin (2) when he has ta dlscriniinalc in order tlitil he mnr make the sixnal
whvn II is spoken, hut not nhrn o \» KiHikrn. bi^ not knowing which is about
to bo spoken. th« uhnle miclinn period wilt 1h- found l» )h- diKlimaly louiter
la the scc«ud cn-ie. The exitt-rimrnt mav br variiil by makingimMif iill the
vowel nunds Inkcn irreiciilarlv m lh« otimulu*. the iiihjcct mpondintr by a
•icaal to one only, an urrangtsl lirfnn'hniid, And. of cuunM. other orders of
Mimulutt mar be iiicd, either vtfual. ihc [litEnal hf ing miwte wlii^n a nnl light
is sbi'wn but not when other t'olorx ur« ahown, ur tactile, tin' nignnl Wing
made when one part of thu bo<ly is touched but nut when other pitrts ars
tourhol. nnd the like.
In such cx|N.-riinent4> where the eubji-cl has (o disttnjruiBh, to discriminate
ItrtwfrD two or mora cvcdis. th« prtflnnmlion of the reaction period la
obviously due to Iht lonMr lime required fi>r the psychical procMWH takiag
plain (luring what w« have called the orotral slngv. In the two cows, one
vtthout ana lh« other wiib discrimination, not only are the aflercnt nnd
eflerent oiaKes the nme in br>lh. but we have no reason lo luppiue (hat in the
rentrnl stage is there any diflerence b^inecn the two c»»e» as lo (he time
taken up by the transformation of simplo feimary impulsca into perceptions,
or as to that taken up by the will in giiining accns lo the motor spparatiM
nnd HO staniiig the processes of the tlUn-iit stage; the delay takea place in
the [ciyiliinil procesBea interrening betneen th«o two piiri*. and the amount
nf delay it the meaHure of the time necileil for the pric™«« invntved in the
discrimination. Thiit " diicHiniuatiim jicrioi] " lim l>><cn fouitd to ditler in
tbe same individiml aci^ordirig' to the wnsn lion emptoyetl, viiual, nuditury,
etc, and accnnling to thi- kimi of dillervucc in tbeamntiot) which hoi to be
dii>criminal«l. for iniijincr in viiunl scn»ntioiis between oilors or butireca
ob;re(# in ililh'ri'nl narls of the tield of virion. In n tniric* of oWrvntiona
made in this way. tnc discrimimition periixl. i. p.. the prvilongntion of the
•implc reaction jicriod due to having to discriminate, was found to range
from 0 01 1 vcomd U> O.iWi svcoiid.
Another irries of obtrrvat ions tnny l)o made in the fiillnwing way: Tlw
signal being one mode with Iho hand, the simplo reaction period tnr n
stimulus is uclermiitcl with the signal given by the right hand. Two kind<
of stimuli are then employed, both nf the same order. Iiro vowel sounds tot
iostanoe, and tlie subject is directed to respond to one vowel with (ho right
hand and lo the other with the left hand. It is found, the subject being
right knoded, that the reaction perio-I i# greater when theagoal b made with
IM left haild. In this case the oetay take^ place not iu the KOOgnition of th«
etfbcts of the atimubts. nor in the Drooeves ihrougb which tlie will is lVm«<l
upon that recognition : these ore tiie same in the two ea^es ; it takes place in
■ he procMsn by which the wilt is brought to bear on the nervous motor ap-
I«r«Uia for making the tigujd, on the vortical origin, for exam|ii«, of the
878 TH« BBAIS.
pyntnidal tnct ; Iheae prooeaea take n Inntc^r time in ihe ctim of tite unic-
rtistomod left hnnil limn in the cue of the u«iinl nclil bend. In this way
we obuin k mea>ure of, W to Ipeak, the volitional sidi* of [isydiical |<ir>-
cewes.
In n »nm(!wl)iit ximilnr vny wti miir nhtntn a meanire of the time recuireJ
for ]M;rrrpttnn. A Mtmng mnMition fcillnvrini; ton cliwcly upon a weak one
will jiTTTont the p«vphi<^l rocofrnilinn of lhi> wmker onf. If, for inttanoe,
tven or ihrm Ictten in white on a lilnck hnck^ronnd W pnw-ntnl to iIip •■yr,
and n Inr)^ white siirfncp he pron-nlcd nfterwaH at an int^rvnl whitli h
mndi?i'ticcce!ivelT "hortcrnni) shnrler, it i» found ihnl when llip inti>mlli
nmdc very Iwief indeed the letters rannnt bo perceived al nil. In propnrliia
«» the tiitprnit is proinnired, tho recojmilion of the kttew incn^w*. nnlil at
an inlervnl of about 0.05 gpcond they are fully nod clearly retvi^initnH.
That is to my. the time rei)iiired for peroeplion U in such a ca«> nf ahotl
that lenirth.
The dnmtion of all these psveliical proeceaea. as of the simple rrarltniB
period itself, variea of course under different cireumstances. and the disi'Hmi-
nation period may beoonvcnienilj used for measurements of the varyiaj;
effects of HrcumilanoM. Pntclioe shortens ll»e diiwriiDinstioo periixl a* ii
dnes the simple r**!-!!!*!) jieriod. One of the most powerful inlliieneea is llial
of ellralioD. Aiid it is stated that the shnrteDinir of the period is creatft
when Ihe atirniion is cnni-enirated on the mnkinic of the aignnl than when il
to more MrwHally dirrdcil to r(-c<>|fttilir>n or the xiimuliis; in other ««nll,
the volitioijal prncessrv are more ampnabli? ibiin are the perceptive pmecMM
to the piivfhiml nctmn which we call atlUJiltun. On ihe other haiid.lht
perimi is dirtinftly prolnngwl if the i)liiicrvi>r h*- dUlnwieil by C(inc»niiiani
aensnitoDs. For ♦■namnlv, thi- perioil far diMriminatine lieiween two riml
aensaliona is prolonj^ if powerful auditory stinsat ions he eKnt4.il at the MSI
timf.
The *ame method of measurement may be used in other waya and anda
nther eircunislanoes with reference to psychical processes. It must bo mama-
bered. h'>wever. that all such nbaervations arc open to many fallaewaaaj
Deed particular eaution. It not unfrequently happens that falm reealt* u*
obtained: for ioRtance, the subje<H, eipeetinj; the stimulu* to be hrmitrhl t«
bear upon him and straioin^ his attention, makes the signal bcfir* iht
stimulus actually eonm off" And the interpretation of the roaulia obtaiaid
are in many anu-s. verv difBeult ; but it would be out of place to d<*«U vp»
tli«M matter* nay further hero.
TrB LtmI-IIATIC AltnAXUKUBHTS OP Tnii BHAtX AVD Spikai. Ooi&
! 684. The mrmbranet of (A« brain and ipinal eord. The oerebr»«pllMl
canal in lined br a tou(>fi lamellatcd membraiw, composed of cooiwrtit*
tiaauc with a small amount of elastic nctworlia, colleil the •iura mtUr, whMi.
Bomowhat closely adherent to the walls of the cranial cavity, is sepanud
fWtm those nF the vertebral canal by a oonslderable apace, contuinin); blood-
vewicls, especially l^ri.'e venous BinuMa, and some fat. Il mity be cousiderej
*» a development of the periosteam lining the wrehrunpinal cavity. It
•eiMb tubular sheaths for some distance along (he i>rver.il cranial and spiDal
nervn ; and forms between the cerebral hemisphere, in the longitudinal
IWure. a conspicuous sickte-iihaped vertical fold, tlui fiUf etrthrl, a> w«ll m
a smaller hnriK'mtal or ublir{Uo fold between the osrobellum and otrvbniai
known as the lenlarium.
The vascular pia iNa(«r b oloaely attached to the sur&ce of ibv brnia mi
•piiul cord, dipping down n> ite liare aecn into i)ie vontrni or nnuHor tiMiirc
ul' tbs cord M well lu Into iW tiwure* of tlic liruin. 8heiilli-like invmtmvDU
of pia mUcr nit: roDliniui] nlnng [be •everiil nerrcs u llicv Icjiw the ccKbro-
•|rilMi CBviljr ; nml iu ihig v<Tt«bnil cbiiiJ ui impvrfoct jmrtiUun Itnll'wmy
biniKa) llio donal and wutml nurlnci'* of tho cord is funiiiilieil br « moni-
bniw of oonnecliTe tiwuc which, ctniliiutoiiH iiloii); iiit whole length wiih iIk
pis mater, in atluched to Bn<l IiihkI with thi; dura mater at iiilervab only,
uninely, l»etwcrn the nucccMtve nerve r<joI)i. .Siinic iU outer etige lio* lhii» it
toolbod ajipeaniiice, ths mciubrtttii; i« ml led the li'ianieftnm dmlieulatiim.
»BalwMD ih« [Mft muter Dcxt lo tlie brain nnd curd niui tliv dora maler next
to ibo bony -HtiW* i* a ovily, which is divided into twu bj n thin membriiiic,
ttir nmi^fivid, cnnipo^ '•!' interwoven bundles of coiinet-Uve ti»ue. Tlie
tjisee between the nrnchniiid nnd LhcdurK mittf it culled tho mthdunU tpOM,
and the ipoee betiroen tho urschnoid and thd pin mtiler i» called the <«i6>
atwiinoiit (fpaet. When tho brain ii expt«eil by n-nioving the roof of thv
•kiill and Hlitting open lh« dura mnter, the siibdurid fjinco it Inid bitre. iind
iJie arachnoid i« seen »trelcl>ing over the pia muter ; to the vertebml ennnl
tbe arachnoid lies clom to tfa« dura mnter. so that UHiinlly, when the dura
nat«r is slit open and tum«d hack, the arachnoid i» carrieil with il and the
cavity exptwed is that of th« mbanichuoid epnee. The ani(;hiioid, like the
dura mater and the piit uiatcr, iaoontinucd for some distance over tbe nerves
•8 ibtty leure the een-bro- spinal eavity ; bo that each nerve at il« exit is sur'
KKloded by a tubular prolun|{atioD of the Bubdural Bpuce, nod wilbin tbiB a
aiiuilar tubular proluuKBiiou of the subaroohnoid apai%
The Bulxlural apace itt broken up to a alijtbt «Ktent only by bridles carry-
ini; uervea and bloodvedaeK especially venous sinuses, between the pia maler
and dura mater, and, over the surface of the brain, by villus-like prujectioua
ud'tbenranbnuid, ctilleil Pacchionian glands, some of wlUch pierce the venous
Mouses of the dura mater. It u lined throughout, both on itt dural and on
iu arachnoid irall, by an epithelium of Hat epithelioid oclls, nnd may Ik
oonipared lo B wroii* cavity such a* that of the peritoneum. I>ike tbeteroutt
cavitit* it <.-ontnint nonuiilly n small ijuantity iiiily of lluid, and ita nu la
potential nithrr than actual.
Tliv aubamchnoi'l tpnco on the other hand w, wpccinlly in certain rvgioua,
such as thi' donui) portion)! of the vertebral canal and the base of tlie l>rain,
iDucli broken up by bridles of coniMCtive tinxue pnniujf from it to tlie pia
malvr, ns well os by a network or s|ionge-like arrangement of bundle of
counectivu lii«ne lying immediately benntlh itself, and giving it when viewed
Irjoi below a honeycomb or fenestrated appearance. Ihe under surface of
the membrane itself iia well as all the trabecule of the tponge-irofic and the
bridl<« are cuvereil with an epithelium of flat epithelioid cells, wliicb is con-
tinued aim over the pia mater and the ligamentum denticulatum, anti lines
the tubular sheatb-like prolongations of the space along the issuin); nerve
rooit. Tbe subarachnoid space therefore, like the subdural space, may be
mpirded as a serous or large lymphatic spac«, but it is an actual not a mere
puleotiat apace 1 it always contains an appreciable quantity of Duid, whidi
however is not ordinary lymph, but is luniished in a particular way, and
devervei «[)eoial utudy. To understand tiie nature and origin of tliisemfcnt-
epinalfiai'l, u it ia i-all«d. we mutt turn to some special arraagninenu of tbe
pia mater.
I 685. The [na mater pro]>«r, conaiMing of interwoven buitdlee of con-
Deotive tiaiue, with wine dartio ISbrer and a considerable number of
coBiMCtivc-iiatuv ooqmaeba, senrM aa wo have taid a« the bearer of bluod-
to tbe nervous structures which it invests. The imnll arteries oa they
into the nervoua substance by tbe way of tlio septn nra surrounded by
878
TOE BRAlIT.
fieri viidcular lyniphuic rannlji with trhicli spocea in the tmroglU rtmibI*
Murk both of ibi; bniiii mid ii[iiual i-urd, cvpeciallf stisces eumiiiBtna): tta
larger nerre-cctlii, ore coniiiiiKiu*. A» i* ibc vote iriih other ib»i», m villi
lh« crntrnl nfrvrtii* *y>triti, tbu Mvcnil elrnient* of tlw iHWie »n batM
«itb Irniph ikrivcil ftom ihe bli>od; ami lliU, oo/ing througb the apMM lata
lh« ptrivn^ubir (■iiniiU ntiil tbi: nihrr lyinphutu* vnm;lii of ihv pla HMUr,
litakai it* way iolo ihc nubHiiichDoid vjihcv ; but tbi; fliiiti in the wibamdlBiM
tpacvi ha« oilier •ounx's bntiilcn.
Tbe roof of ibr fourth vnilriclc iii, aa wn hurt- aaii) ($ 602), rpducvd to a
nioglr InytT of 111)11 iivrvnii* I'oltininar cpilbcliiini. nbtch Bpp4-an aa a moc
liniii^lii lh« |>iii iimicr ovi'tlyiiig it. Jii ilu- hiiiiirr twrt oflbi^ vrtitri(-l« ihii
rniif i? ptrfiiratnl bv a iliMinct narr^iw oval otififx', the/oramrn «/ .UaJ-iJir.
hy lhi« orilicc, Hlik'h pti«c4« right tbroiigli bulh ibo pin rniilvr aiiri tlM
uiKtrrlving Inrvr ofrpillidiiim.llivmvitT of the Iburlh vctitridc. aiiiJ so iht
11 hole (criteof ciiviiicHiU'rivciJ fr> in the original mcdullariF' cnnal, the laiml
■od ihinl vcniridoK, Ibc ii'iiicdiicl. and tli« crntial cniial of tbi< apiual oord,
are niiide cunlinuoUN with tne »ubura<-bnoid Rp4ic«. Tli<rfv ant also other \tm
coRfeplciioUB cuniRiunii'nUone bclUM'D the eubainclinoid space an<l lh«-f'>unk
v«Diri('Ie. Il<'iii.'e (lie (erfliro-spiniil lluid in made coinn»on to all thcM obv{-
li«», and ia furnteh^d nd only by llie pia maier invealing the oulude of tlit
brain uod tpinat cord, but aim, and indeed probablj to a larger extent, bf
ibe epithelium lining 1 lie several i.'aviiie«of the cerebrO'Spinal itxit, tapveuXtj
)>erhii{ia by lh<«e porlinrift of that epiibeliuni wbifh coat the proonaea of fai
iiinttr priijectini; iitlo iljuHe cavities ut certain ptueea.
We eaw jirevjouBly (* IJU3> thai a lar^e l»ld of the pia mater, cnrryincto
with it the ibiu non-nervou»epilb4^liuni which alone frprniL-iila nl ibe fiaM
the oTi|-iDal wall of the medullary canal, is thruat iiinanl at thi- iran*Tprv
fiaaure of tbo brain, b<-iii-nih the toniin:, tn form thu velum iDturpuciium, iku
tnpplyiog a nuf in lb« ihinl ventricle, and that it ihanca pnfjvcta iota tttk
bili;ral reatricle an llie dioniid ploxuv of Mcb ntdu. reaching from tbo fuiv
RM-n of Mouru iu fmut along ibe ■■<Igr- i>f ttiv I'urnis to the li)i of tlw detoMd*
ing horn. The velum bi-irig a fold of the piii mat<T con«ifl< thcoreticall* af
two Inyen", ami bclvteiti the upper ilorMil Inyi-r ami the lower Tcntral laytr.
I'm a ihin bc<i of cinitieclivc tiasue carrying nrtoriea forward from the hind
Cilgc of ihv corpUR ciillu«um, and similarly carrying vvitu backvarU: them
vrawla »upply the choroid plexuK with an abundaot nipply of blood. In the
choroid plvxiia. the fold<:'il pin mater i« develofwl into n number of tiIIm-
like pniceMc*. the primary prorcnscx bi^iiring secondary onea. Kadi prucc*
coiisisia, like a villii*. of a bii«i« of curinvctive li<«ue, iu which the blomlveaadi
end in cloec-«et cii|>illary I»iips, covercil wtihan cpilheliuin. The epiiheliuin,
though conliouuu» with iIk' rest of the epithelium lining the lnlera.1 vi-nirictt,
and tbiiis oa wc have said, shutting off the lateral liNim the third veniridt
(except at tlic litrHaicn of Monrn^ and thoofjb like it deriveil front the wall
of the ori;;inaI medullary canal, in dillereoi in structure. Over the vnitiiele
generally the epithelium consiniii of ordinary short columnar, npparvatl*
viliatetl cells "ilh more or leso troiirpiireiit («U substance; the oolln uvrr tH
choroid ptexuH arc- cubical. uHcn irrej^iilur in form, and their cell aiihataDcc
ia loaded with granuleti. M>me of whic-b are pi|fineniary. Thejr have nrj
uach the apjiearanee of " active" necKtiug cella; aitd iodoed a branchM
proceoa of the plexun may bo coinpared to an everUvl alvc<>lu» of n ■veretiog
eland, wtib the epiihelium uutHiih! and iho bloo(tv>»L-lii within. It c^niiut
be doubted ibnl these i-elln play an impuriant iMrC in itecreting into the ority
of ibe ventricle tluid which, pawing tlK-iice dv the foramoD of Monm inUi
Die third and au into the fourth ventricle, lindi) ita niiy by tb* roraiuen itf
Uajeixlic into llw tiiibnracbnoid space.
TUB LTMPHATIC ARRANOEUKN'TD OF THE DRAIN. 679
At lh« r«liitn owrhRiiga llie (hiH ventricle it w&cils down Tnticnlljr two
iigitudiiial linctir rHii^ce. wlii<rti, n.-*cnil>liiig in alructure tli« chcroid plex-
uvnof the Imtenl vciilncic, nre cnllcd lite chonMd plciusm of the third v«n-
triele. From th« rooruf tho f<3imh vctilricle (here naogR down on mch side
S eimilnr lincnr tViiige, tlie choroid plexus of the fourth ventricle, which is
aii«cialljf develope^l al iio frunt end beD«a(h the overhsaging eeretiellum.
Tdcbc oulnidiary cboroiil proceoECfl doubtles ueisl in furnishing cerebro-
•funnl duid. but their share is snuiU coai{Mred with that of the main choroid
plexiita?8 of tlie lateral veotricte.
i 686. 'J'lie errrbro-rpiiuil JIuiJ, The apeeituens of cerebro-apituU fluid
which have been cxntnined aa to their cooipuitiituii are not quite ooiDparabte
wilb e«cli other, sioce while some (aueh aa i.liuoe obtained Irom cases where
• ftaotore of the haw of the skull has plaoe<) the subarachnoid epace at the
baae of the hraiii, where it u l>ri{ety <icrvelo|>ed, in onoimunicatiuD with the
exiemal mentuK aud the fluid e*c')i|>e(t by the ear) niar be reicarded as
Donual, otiient (such as ihone obtained from caseit of bvdruvephalua where
the vviitriclea contain an imusunl •[Uaoiity of tluid. or Iruni chhkh >if ajiinul
nmlfiirniitliiiiia I muHi be (-oiuidi^rKd ud abnormal. In iuihI of the more oini>
|iU-tr iiualyM^. ibi^ Hind i-xiiuiiii«<l hiia belonged to the lattvr ctiba ; nud the
lollooiiif; HtalciueuiH a|i|)ly. strii'tly s[>cakiQf[, to thvm uloite.
With tbii niiiiioii. we may say that cetehru-iiiitDal fluid ia a tramparenl,
wlotleMor very th^htlv yctlnuinh fluid, of fiiinl alkaline rMclion. frvv from
bbtQlogioal I'leiuciild. I'hp >|irt-ilio gravity t* about lOlU or Ium, thi- untouut
iT wilida Ix-in^' on an avrrni;)? 1 jirr c-cdI. Of the3« by far the ervalor part,
1.0 or U.S per oent., i* *u|i|))ii.'d by nalla, the total ijuanliiy of which a* well
ibe relative amount ol the iu;v<'ral vooittiluenlji being about the mine ni
in in bloud nnd lyn>|>h. The coinpamtivv dfficivncy of voUda i* due to
tin Kifilineas of the pnuviilA, which rarely exceed U.l per cent. Thoe are
cbieHy globulin kimI a form of nlbumoee. or crcn peptone ; albumin is taid
to be i^nerally ahwnt. The fluid, save ap{HireniIy in exceptional cases, does
Dot cloi, and conlniii<> iieilhcr tihrogenoua factors, nor flbrin ferment. It
very frequently coniatiis a substance which like dextrose reduces Fehling'a
•oiuiiou hut which is not a sugar ; it appears to be pyrocatechiu or a cloaely
allied iHMty.
Heeing that a fluid of such a coiupOsitioD is of a diflerent nature from
ordinal^ lymph, furnished entirely in the ordinary way, we luifcht be inilincd
lo inter that probably a very lar^e part of the whole maas of the fluid it
fuiniBbvd by the aecretiuK cpiihejiuiu of the choroid plexua. Bui it must
be Uirne in miitd. that tlie (uri-i;oii)>; unalyaes rvler chiefly lo duid appearing
under abnormal circuiiiBtaiKvt. aud il wuuld be hazardous to draw any wide
tafeiriioe from them. We have little or no exact esperiineDtal evidence aa
to hon much fluid is actually secreted by the choroid plexuiee; aud if tlie
flui4K which have been nimiyusl do repre*eut a mixture of ordiiurr lymph
aupphcd through the pia inuter with the jieculinr ttecretiun of the cboroiil
plexun and cerehro-ipinal omihI, aoiuc further change beyond tlie mere niin*
glingof the two fluids ia needed to explain the remarkable alMCnce of albumin
wliii'li liii-H liren to ittrungly insuted ujKin by various authort.
j) 687. We may fairly auppoto Uial during tite the duid ia oontiuually
being Mi)i(>lied, from the ono KXiroe or the other ; but we have no vrry exact
knowbngv ai (o the rate al which it ia funiLdietl. In the dog, the fluid haa
n ot>M;rve'l to i^ncJipe at ii rale varying very lar^gely under diderent cir-
cuinstitnccf. anil ranging fomi 1 c.c. in Tbrty minute* lo a« much a* 1 co-
in six ininutrs, the i4mil i|iinniity discharged in twenty-four hours varying
Irom '-Hi cc. to 'J4U c.c. In the casea of fracture of the base of the skull
lucuiiuued above, a very coosidcrabic flow haa been frequently obecrvcd ;
«■ ,1,
Bbeii
i
TRIE BBAIK.
but it may be d'>ubt«cl wliethi?r th« abnormal circumsdiDoes of such an
have not raited the Kcretioii above the Donoal. The rate of Sow was futunl
ID lh<! dug to be much increased hv the injection of aubsunoeB (nonuBl Mtliot
euluiiun) iuto the btood, but to be relaiively litlle infltienced bv anifical
heightening of arterial preaaure. This has been put fifrnard *a indinun^
that the lluid ia chielly fnniished as a eecrelioii sod oot ua au unUoai;
iraneudntion of lymjil) ; but it cannot be regarded u attoidiug » valid arga-
roeni. The preeaure under irhkh the fluid exists b also very variable ; it ii
olueely dependent on ibe viiscular arruo^emeuts of whkh we almll linTvto
speak presently. In ilie dof{ tliu averajj^ pressure baa b«eu cMitnaiMl u
aoout I'J mm. of mercury.
II' the fluid i« thus cunlinually formed it muHt alvmy* liiid a aummtl
eeeai>e. This is probably supplied by tfa« tubular pnjl<>ngnii<inN of thvaulK
arachnoid space along the iiervo i-oota , these are oontiimous with (hv lympb-
aiie vesulsof ibe nervts, and so uiih lli« lymphatics of the body gtoijwji
and in the skull, the |>asaagesof this kind aiou),' tliv cniiiial uvrvm, etpttiulj
uUiiig the two uulk- nerves into tli« urbiln, nllbrd a rculy tneuis of iKMa.
it i» all") ur);e(t that some of the lluid etcapes through the PacchioBiaa
^luDiU directly into the hlfwl of the venous nnuMw. In a d«ul budy flaiil
ininiduved iiito the sulrararhuoid ii|tncu through an opening owr lli« hulK
dbapiienrs at even a very low iin-murc with ^rcat raptdily. Thi- cimi*-
nlaiK'iii then are, however, not the mmc a* in lifv ; nnii the ftv pxperin>aita
mhich bttvo been made seem to «liow thai, during life, n sonienhat hi^
pri-Mtirtr in re(|uireil to swure the escape of (laid iiilnnlueeil in additioD It
that naturally inwn![«d. Thus it is Ktiilcd thai when in a dog normal mlm
lolulidQ i* inlniduocd into the subarachnoid caviiy at the loirer end of tk
epiuid cord very litlh^ ri.'«iir]>lion lak<'« place so long aa the pieswn
rontains b« low ns nbout 10 c. c. of mercury ; as the preraure b JncrtawJ
beyond this rcior]ition (juickly iucreoses. Ilut it may be do^ihtnl nbethfr
the resorption of added fluid is n fair leel of the a»cai>e of lluid nalurallt
preteut ; and (he exiwrinient Is of value mther as showing simply that thHC
are lueans of escape than as aClbrdtng a lueasure of the rate of caopfe B»
sides, the immediate effecis of applying pressure at lli« caudal end of i^
spinal curd are not the same as ihuse ol' applying the preauirt witbio lb
skull.
The rale of possible escape is not without importance urn regards lb
mechanical ioiportauce of the <«r(--hrO'SpiDal lluid. Thus it has b«aa lUgld
tJiat when an extra ({uantily of blood is driven into the skull, any injuiiow
intcrcmuial compreesion is prevented, not only by iImt tran^lerviic'e of a «t^
irspiinding quantity of cercbro-spinal fluid tfaTOUgk titc ft>rameji of Mintodit
fmni the cranium into ibe Mpiiial canal, the walls of which nru lees nfM^
complete, but also by the direct escape of tlie fluid from (he cavity of Ibe
tkull along ihct cranial nerves in the manner denrrilxMl. It has also hnn
urgtil Ihnl the fluid at Uw base of the »knll, in the largv miharnchnnid spaon
of nthich it gutheni in larger quanlily than elsewhere, ad* an n s»n of pr»
tcctive water cunhiun to the delicate cerebral substance, and thai, in gvnenl,
the prxiN'nn) of the fluid is mechanicully usoful to ihe wel&ro of the bnua,
rcHKivnl of the fluiil by aspiration being said lo lead to bemorrliage from the
pia maCrr and to various nervous divordcrs. But our knowledgo u lo the
part which the fluid playit is at present very imperfect ; and its very peculiar
chemical characteni suggest that ii has sorac chemical as well at least si
iMcbauical functions.
?B^TAflODLA^ERHANOBMBJ?T^^TH^BBAIN. 881
Tbe VAsc-vtuK Arranoemcntb op the Braik and BriKAI. COHD.
$ 098. Tlic bloodvcwels reach the nervous Blnictures by roeauii of the |na
nitiirr. In llie aiiiiial curd nrtcricii cuming fr»iii the vertebral, iDterfuaial,
anil utber arteriee, anil travelling iiloiij^ the nerve ruotd join the pis lualer,
anil Uk-r lhn>ui;h the liwiurcii and wptn n^ch all purls ut the oottl ; but, na
we have prrvi'iuisly rvmarked, tho capillary tietwurk t» much deiiaer, au<l
Ibcivriro the bloml mpply much f^rcater in llie gray than in tiie whil«
malUT. The x-faia, also giilhemi up iilong tliu iieptA aixl tiwurea into ibe
|tia niatiT, ih<i«>G cuniiug IWiin the gray matter forming, l>ffi)iv thrjr rtqu'h
Um extrrtifll pia niatcr, ii (■■>■■■ pi cuoii.i longtluilinal vein on eut'h Kidv nf tlio
|MMarior gray cuiuniiwure, puvK Ironi tho piu miilvr to the large renotia
MliuMa of lh« diiru niiiter nml »i> to ii(lji.)tniug vdna.
In the hrnin two iinporliiiil iVqitiirtv of th« dutribulioD uf the arteriw
<l«<TVe cpcriiil uiK-nlinn. In the tiril pinrr, tin- iiuiidruiile viipply by tbo
right anil Icti «'rli>hr«l and inlcrniil ciirolid nrttric" i» nia<lc nnv hy renuirk-
able aua»|i>ni<«<-» f'irniing the nrrtc of H'lllU. The right nttd li'iV vertebral
arleriea entering thi.- rerictiml cMtiiil nt the U-vcI of tli'^ sixth cer^'icnl vcr>
lehra ami running l'i>rwBrtl toward tJte bniiii join hcticath the ventral Mirbc«
of the hitlb to Torm the *ingle median liiwilur arirry. This, niti-r giving afl'
brmnehes to Ihv kult), cercUelhim. and {>ons, divides into the right and left
poeWriof ocrehnil iirteriiw, Each inicriinl carotid entering the skull readies
the blue of the brain in tho region of tho Door of the third ventricle, and,
pataiog ventral to and athvori too optic tract, gii-es otTlhc large and impur-
tutt middle cerebrnl artery along the ti^suro of rfylviut, and then, luroiug
forward and toward the DiMiBO line, puMes duraal to the optic nerve to eoa
in tbe anterior cerebral art«ry. Just, however, sa it giv«a oA' the middle
■rlery, it sends backward, inclining to the middle line, a relatively large
bnacb, the puaterior cxiiuniunicutiug uriery. which joinx the poetctior cere-
bral near the origin of thin fn>m the basilar uriery. Moreover, the tw4
anterior cerebral urteriu, amm after they have criMed th« optic nervei, jual
M tbey ure about to run straight forward ulong the ftontal lobes, are joined
lonelier by a dboft. ivide brunch, the anteriur comrauiiicaling artery. la
thw uny tlM vertebral arterieH through tbe Wilur artery join with ihcu
carotid artrrie* to furin uruiind the optic chiaxnia Ixiucath ihe floor of tbsr
tbinj Tcntricli- an arterial circle, Ihe circle uf Willis.
BIoihI can piiM along thin circle in vnrioii* way^i — from the bu^ilar arlerr
■long the right podteriur comniuntniting artery to the right internal cnruliu,
aii<l «o by tlie right anterior cerebral artery and anterior conimutiiaUiiig
uriery to the Icll Hide of tlie circle, and siniilurly from the basilar artery
along tbo led stile to tbe right, or from the right or frtrni llie lell canitid
through the circle, to the right band or to the lv(l hand in vuch ease. Sine* j
the channel of the circle is a (airlv wide one, the jMUwagc in various dimciioni ^
is an easy one ; all th« vrtvels nidmiing from the circle, including the biwilar i
artery nnd ils branchtw. can lie siipplirid by the carutida alone, or by tite ver-
lebrnls alKue, or oven by one rurutid or one vertebral alone. In this way an
ample Rupply of blood lo the brain is M-cnrc<l in the face of any hindrance
to the Hoir of blixnl along any one of tho four channels.
In what muy perhaps be consi<lered tbe usual arrangement, the («libr« <
ihe pislerior coiiiniiiiiicating arteries is rather smaller than tbe other pu
of iIh' circle, to that, oiher things being e^jnal. moat of the vertebral bla
will |iafe by the ptieterior cerebral arlenca, while the carotid blood ptMa tO ,
ihe middle und anterior cerebral urleries ; but many variations are met with.
We uny also here i>erha]>a cull to uiin<l the fuel that the kii carotid coming
M
882
THE BltAI^.
off from lb« top of the aorla ofTere a more »tnii)jht paib fur [Iw blood than
don llw right carolid which com» ofl' from lh« lODominiile arlerjr.
AniillKir Bpecial fenlure of the artenal itip])ly to the braia is thai 4)
three Innje cerebral arieriea — puBierior, middle, aod anterior — are diiuibf-
tod alnuMt exdmivelv lo tlie coriex sad to the sul^aeeat vhiie inaiter, «kilt
th« dr^jier twrtt of the heiuisphere, the nucleus raudatiis, llialainiiE, aM tht
like, with the cap»iile aiid oiher adjoiniiifc tibite nuilt«r, are mippliol hi
iniallur arteriffl i-»iiiiDf: direct fn>ni the circle uf WiHin, or fmrn the tmj
breinniiign of the llirte cerebriil art^rie*^ Ii in Mated that then: two Mrliflt
mnicu DO auafiiriiiiiveit with ench oiher; but thi» nnpeani to varr mtirh in diT-
fcrciit iiidividiiulB We iiiii)' add that lli« antvnur cerebral arti^rv riuiiplin
the c'lrtex uf the ilnraal HHpect iil' the Ircintal I»bc a» o-ell a* the fn>ni aid
niiddle pirtioDH of the whole ineajal Hurface of the bcmiiiphcre ; nhil*' the
middle cerebral, alwuvs lar^f, U diiitribut«ij f> the tide of th« brain, that a,
till; jiarietal lobe, wltb the ventral [lart of ibc frontal lobe and thr dunal
part uf tli« temporal lobe ; the uoiilcrinr cvrebral ■npplyiiig thv rot of ihr
cortex, that is to aay the occipital lobi;, inchidioK the bind part of the TonU
turfacv of hemidphere, toffcther with the ventrnf part of th« tOin|><>nil liiht.
The dixtribution of ihi.'M; ant^not ilivn-f'Te doea not correapood Ut fubctioul
diviirionii, for nbilc the middle ren-brul Kuppliiv a large part of tbe iDotw
region, it doen not mipply the whi.ile ol' it, and dot-* »(i)>|ily ]>ari« uuulde tl.
Though ihcRii.all arterial a« they run in the pia mater un the Hirfnc« iif th
cortex a]iiu>l<>ni(ii,f freely, there i« very little nDa»toiDOM> betvpeen lh« mall
artsrios wbieh, Waving the pia mater, dip dowD into th« mbataDca of tit
braio; bence, wlien tlK^e hiller nrleries are blocked, tbe nutrition of Ik
part of the cortex supplied by tbeni i« apl lo \k impaired.
S 699. The venous arrange men la uf the hrnin have v«ry epeciHl ehM-
acters.
Along the i)pf>er cimvex border of the siekle-shaped fold of dura BlUr.
tW fatx cerebii, in devel»ped a lar^e venous oiiiuti, the mperurr tongitu^ml
Wiititi. Tbia. triangular In Hx-liou, increueiujt in calibre from bviorv back-
wanl. \d a einiift. nm a vein ; it« ukIIh are |i>ro>ed of ii'rtbing but cwooeditt
tiMue lined with epithelium, musL-uIar elemeiita being cniirtly nbwm. Thuu);b
iti dianuel ia broken by bridlea i>f conn«ciive tintue pnMing acnaa it, it pw
•eanauo valves, and iudeed theee are a Wot from all the niniiiH'J' and reiurf
llie braiu. Miidt of the blooil r«tii ruing from the oortex and ttuhjaomt while
matter i» carried itito this ainua by veina, tbe mouths of wliicb arc fur tbt
moul part directed furnanl, that is tn oav, agaimil ll>e direction of the falooJ
Klmm. Aliiiig the lower concave bonier of tlw lalx in a ximilar liaiM, tL(
inftri'tr fon^iluiliiial rinut, which, however, b tmali, and into which rtla-
tircly few veins o|M-n.
From the deeper purl* of the bruin, and Ofperially from th<' ebttnid
plexuii. blood id coiiveycil bv the vritm uj Oalrn along the velum inlcrpoiitaiB
to the iransvcr:^ fismre. wbcn; the veins of Ctalen join ihc inferior longi-
tudiiial siiuis to foi III llie ••Irati/hl <ri'iiii*. This, running along tbu line formal
by the inter>ecti(in of the vertical lalx wiih the (mora or leasi bortiuwal
tentorium. Joins tb<- end uf the superior longitudinal nnus to form tW
twcrvoir or cellar, called the tortiilnr Hm/jtMi, j'nim which tlie l-tlrml timut,
pHEsing on each xide along the convex border of the tentorium and gallMTUg
veina from the cereliellum and hiiid regioDS, as well ns from the base of lb*
brain, delivcre the blood into the internal jugular vein.
It obould be added that veins from the noee and, through th« dpbtlialair
reioH, from the face join the veins and ainnaea of the braiD, and that tbe to-
called emiaaary veins pass through the crsuiiim from the scalp to tlie tuperiur
lou){iludtual and lateral sintites.
THE TASCOtAB ARBA:rOC]tEyTS OP TITB BRAIN*.
I
t
t
I
Tin flibanneb for (he venous blood of the braia are therefore nol veiiB, but
ftiDUMs; not to much tubas for muiDCainini; a uiiifarn curr«tit, w longi-
tudinul r«aervoirB, which, while affurding an ea^v onward path, cno also De
euily lillcd and eǤilir emplied, and io which the blood caa move to and tVo
withoul the reelrictions of valves. ThU arranj^enieol is conelated to thu
peculiar surrou»diuj^>s *if the brain, which is not like other or;pin9, protected
itiervlv bv dkiii or other exieiisible or eluaiic tuwue, bm id enciued by a fairly
r<iB)p(ele in«xlcnaibli! eiivelo{>e. the dkull. Aa a cutiMK|ueoce of this, wh«a
at any time an extra quantity of bluoil ia aeul fr»Ri Ihe heart to the brain
room muit be made for it by the iucreiiMMl exit of the fluids already present.
For any prcwurv on the brntn-HulMliiui-e beyond a ceriniu limit in injurioua
to its welfure and a^-tivitr. as is seen in certfliti maladies, where bluod pawing
bv rupture of th« bliHMlvenael.i uuC of ii« n'>rmiil chnnuels remains efTiiMd on
tbe surface of the bi«En or et«ewhere, and thu« takiu^c up the room of the
1tni]i«r hmin Hutxttanee lends, by "com]>r«iHioii," as it u callud, to pamlyvia,
• m» of coiwciouMiwv, or dnilh. Some room m»y. os we have Men f§ 6S7),
b«t providnl by the i^^ape of cerebrn-spinal ttuid from the tknll. But,
within the liuiila of the normal cerabnil einmlnticm. the characlerislio
venous iiiimttni i-Dpec^ially Morve to regiilalo the internal prMsurr ; they fcirtn
temporary nwrvoin from which a uomparalively lur^- quantity of blood
can Im rapidly divcharKi'd frnm the cranium, the flow from the sinnHo being
greatly aiviitMl by the low or nvgntive pnmure obtainin); in the veins of the
neck at each inspiratory aiovrnwnt i>f the chest.
S 700, The cupplf of hlootl to the hrsin sM-ms at first sight nol to corrs-
spund to the ira|>iirliince of this the chief organ of tbcbixlv. In the nibhil it
would amx-ar that hardly more than one per cent, of tde total ijimnTicy of
the bloo<l of lh*r tuxly is present at any ona time In the brain, a lunniily but
little more Ihau half that which is found in the kidneys; and white the
weight of bloml iu the brain at nity one time amounts to about live per cent,
of the total weight of the organ, beioj; about the same as in the miiaelee, in
the kidney it amounts to nearly twelve |*er cent., and in the liver lo as much
IIS nc«rly thirty per eetil. Mnking every allowance for the relative small
•iie and functional impurtUDce of the rabbit's brain, the bhHxIsupply of
even tl»e human brain must still be small ; and making every athiwanoe for
rapidity of current, the interchange between the blood and the nerrmti
alenienu luuat also be small. In otner wnrdA. tiM metahulism of the bnln-
•uhstAtie* b of impurliinee not so much on account of ita quantity m of it<
spMiRl qnmlitica.
The drvubtion in the brain may he ntudicd bv help of rarioiu in«th'wt*.
A ncnoineter may be connecied with the jieApkrral end of the ilivid.-d
inMnul carotid artery, n second maiiomrler being alluchiKl in the it^ual way
U> th* central |H>rti'JU. Since the pcriphL-nil manometer reconi* the blooil-
pnwure in the* circlii of Willis transrniiied alonjc the peripheral portion of
the carotid artrry, varinlion* of iiremin^ in llie ctrcle of Willis may thu* be
attidivi] ; ami a comparii>on of the pc'riplientl with the central manometer
will indinle what general changes are taking place in the circulation
ihrmigb the brain, Thus a fall of prtwure in the peripheral manoraoter
unanN>mpimicd by any corm|M>iiding fall in the centra) minomcter would
show that ib« " |)eriphcnil resistance " in the brain was being lowered, in
other words, that thw vessels were being dilated.
In another melhoil, in the dog, th« outflow of venous blood from the
lateral ndus through the posterior fncinl vein has been msMured. The
freetlom with wbii-h blood passes along the sinuses juslilios the ueamption
tliat the outflow through the opeu reiu gis-es an appruximaie ineuiire of the
THR BBAIS.
■SCI; I
mu of fl«v ttDdcr MUiiml cMtdiikM : «EB fk*
aad bnUm. Um coMivMd kw «r UmJ
A lUnl Mtbod b > pktbyHM^^kfe «M.
lb* rij^ banaf btMi nmvftd' br ibt
belt, Md tbt wmsnto of iW Mnb
utd Unror AneUy bj'aJew. layvaag
Ika of IW cnMO) aol jrH «mScd. b^ U
Aod it! noraanta noorwd in ■ KBilsr am
fluJ to ibc bde in B «ster-ti^bt Bwacr. U
btcnMi pr^nin : uxl oe bs^ uk« it fcr
mcd, that pester or lea pnnw* m 4m I» bor b
U> Um bnw- B«t ibe uaocnl «f ffBuia kcB^ls *• h
iiMtraowftt ■ill bImi (kiwsd on IM fLifiaf nib win
fluid ncapca frMD tbe cxTiif of tbc ilivll : if iha* W
ocape, or on tbe otber fautd sn i»crtBgJ fiaBly af
crtwe of wpplf of blood will piwiace ia oaa cm* a
inhaler norentat of ibe Icrcr. If tbe Btabcaae be
ilie hule BO as to allow free ocape of tbe
(ftatrtically taiinf; >m llie lurfsre of ibe a
rrconla Tariaiiom in the dorao-v«-ntnI diaaWer af
Ibne laaj be taken as neaHtriDK VBriaiima ia ibe
■0 in tbe blwrfl Huifpl^. Id n*-iih«r furtn. howercr, d«a iba
Eive (M all llie itifurmaiiuu «btL-li we want. Aa taiTCBa
rain, aod tberdore au «x[^quuiuii uf the Imia, and §« a
n^u^iag inXrtiiiiriit, rmv i««ilt ntbcr intm a fnUtw
fron hMrance Iv the vtooiw oalfluw ; tbe ibi
iDcat run. favorable to. ibe laiur alwaja aad liaiiaLily
aclii'iljr nT thi- iM'tvi>uit *(ruc-lur<i
lie rurrtiinl liv a liniiiliBiM^OB
tolfct
A:
benca tbe teacU*^ at tbe keer aa«
__ ubcvrvaticio of the naral arteria} ptwt
sod »r ibc blux) (irtwure in lbs vetna of tbt- ovvk. Muewntr. tbw aryaaw
wUicb wv uMtl 'i 4)^> in refrrenrc to the kidnrj bbj beafi|djed bmiad
I'tubably wilb cfual lurcr, namcljr, lint iho vaJoe of tba bmd aUaa to
tilt uutritioo uf ibe lirnuc is ikjiendrnt not atoae on tbe SBenM af Ua^
|>r«Mire, bat alto and n|iecially on tbe rapidiiy of tba Horn • inihid.lfca
Mcioond factor i* of parliculBr im[H>rlance in view at tba aeed of wipffliat
tbe nrrTOUB «kn»cnl> wilb an Bik((UBte inlercbaDp of gaega. Sam 4)i At
rapMlrty of flow tbe pin byBmograpbic method ata i^ve us iadifaei ii^ttmt-
lioB unljr.
t 701. Djr one or otber or nil of iliew method*, certaia inportaM ku
have lieen laode ouU Tbe volume of ibe brain, as deleriBined tif Ikt
BuiouDt or blood present in it, is coniiiiuouiy aadeifoing cbanga bnagk
nbiiut by varioaa cauHiL Kacli lieart-b««t makes hadf viable oa Ibe em-
liral BB ou the renal plelhy«iui>;;ra|>bic tracing, and as we ban ma in
•(■cakini; of rvtpirslion, tbe diminuiiuD of prraure in tbe great vciaa of tbr
ti*^-k during iiiapirmii»n lesils to a abrinkin);, and the reverie ebaage durinf
rxiniaiion lo a swelliii}( uf the bniin. Thv plelbyan(^rB|A alan shows vans-
ii<iiii>, Inrgvr and slower tbau tbe retpiratorv imduUiiou.-. and brouehl ab«al
by various causes, sucb as ibe position uf ilie beed in ri-laiion to wo Inink,
inovenienia of tbe liinbe, roodilicaiinua of ibe renpiraiurv moveintnta, »ai
upparruily pbnses of activity of ibe bniu ilvetf, a* in wading and tlcepiog;
iitidiilatione corrvspondinft to ibe Tmube-IIcnng variaiioua ($ 3$t(} «f
liluwI-prtBure ntaj not unfrequeully be glmer^ttl.
All ibe vartuua melliods Hbuw tbiit lliv flow through tbe brain is birg^f
dHcrmincd by u vatooMlor acliun of nonic kind ur another. And ibb vs
[AiN. 883
I
I
I
might initwd infer from onlinnrr eiperieiice. When tbe head is i^irdttcnly
ahiOed fmin the eKcl to a han^-iii;; ]io»iilioD, there muM be n t^ndencv tor
the blood to aceuii)ulaC« iu the cranial cavity, and Cfjnveniely when the head
M Baddenlr shiAed from a han^Jii^ to an erect piMitioQ, there mual be a
tendency fn the supply of blood within the cranium to be for a while Icm
than normal. K>tber ohaDg« of fxisitton. and especially perhajM the latter,
would Iea«l to («rebral dijilurbinces, which in turn would in oiirwlve) be
revealed by affectiouB of our conauiouaneia. Thai a jierleotly healthy and
especially youn^ or^iiuitu whi»e Tasoniotnr niechanlsinj are at nnoe eReotive
BD(I delicately reipi>iu,ive, can i>aas nwiflly frnni one jiu«iti<ia nf the head to
the other wiihuut ioctrnventein-e, whcreiut thoM iu whom the vii*imi>tor
mecfaauisma have by aic« or olherwiac boome imperfect are giddy when
Uiey attempt tuoh rupii) chani^, U in itaclf adequal* evidence of the ini-
|>»nnnc« of the vaaumntor arrangemenla sifbctinE the circulation throiij(h
iIh> bruin. Tho iMveral methods aeree in ahnwing that incrcaaed general
arterial pr^^Mure. Mich as that fir instance indiicol by NtimulHtion of a
•enitory ntrrc, leada Iu a gnmier How of blocul to the bniin ; the vnlume of
tbc bmin in increawd and the vcaoiin outflow by the lateral linus U
(lutckencd. Oinvervly, a lowcriiig of artt'rinl prctturc leads to a le«cne(l
flow of blood to thi' brain.
Seeing that thi- ccrfbrni arteries have wolMevelo|X!<l iniiicular coals, the
bcwilar artery- in fact l>eing ci>niiptcuouii in this rrspect, one would be led to
suppofc that th« hrnin pnewnseil snccinl vaj>om«tor norrrs of its own ; and
racogaixing the importance of Mima-supply to rapid functional activitv niw
wouM perhaps antidpnic lh.it by special vasomotor action, the siii>p1y of
bl<M<l to this or that particular part of the hrain luighl bo rvpnlnml apart
from changes in the general supply. The various observation", however,
which have hitlierto been made have faile<l to demonatrate vilh certainty
any such special vnaomotor nerves or fibres directly governing cerebral
vewels. Ii would be hazardous to insist too much on tliu negative result,
especially xiuce the obaervationa have been chieHy directed to the nerves of
the Deck, the exiierimental difficulties of investigating the presenoe of vaso-
motor fibres in the cranial niTvei being verv grml. Slill tt may be urged
and indeed has lieen urged that the flow of bluiKl through the hrain in »o
delicately respotuivc to the working of the general vasomotor mechaniitm
juKt W-ause it has no vasomotor ntTves of its own. In such an organ as
the kidiwy, an increase of geiwral blond-praaure, a* we have more than
once insisted, may or mar not lejiil to a jrKstcr flow through the kidney
nccordinjc a* tlic vtsMis or the kiilnry itsi-lf, through the action of the renal
vitmmolor nerves, are dihtol or c>n;«tnctcil; and. a* we have Men, a ooil*
•tridion of the n'.nal vrmiIs may be one of the conlributon to the incTMsod
genenit priw^urc. !u the hniin, on the other hand, an increase of general
arterial prwsure seems alwav* to lend to increase of flow. Thus in the
Traul>e>Henng undiitattons juxt mentioned, the expansions of the brain
arc coincident with the rises of the general prcMurc, whereas in the normal
kidney and in other organs the local Traubo- ITcring nn<hilat)on reverses tiui
Kviieral one, the slirinkings are synchronunn with the rises of nreasurc, the
U>ca) ooustriction being ona of the factors of the general rise. It is arguod,
that in the nlMcnoe of vasomotor nerves of their own, the cerebral vessels
are wholly, ro to sneak, in llie hands of the general vasomotor system, so
thsi when the blooa-presaure is high owing to a large vaso-eoiutrMtion in
the abdominal visoenti more blood must necessarily poM to the brain, and
when aesin the bload-j>reasure falls ihroug^h the opening of the splanchnic
rtiMxt-galee i,^ IT3) lets blooil ncceasnrity tlowa along tho cerebral vessels.
And indeed one may recognise here a sort of self regulating action ; for
THS BRAiy.
dimiiiif hiiig llw Biippi}' "f blcmi) to cite viuriniiftor ■.■rntro in ttie bulb mdf, m
we kooK, UBA powerful Hiinniliii in [producing va*<>-c<>ii»lnchiiii,wi<l m 1m4i
to a rieo nf blnotl-pn-murv ; but Uim very rwv of blixxl'pmwura drirot ni'iii
blood to th^ binin, including tbc bulb, und thit* the injtiriout «fliBct«tatU
brain thrcdicnpd bj: un ana'niic conditioD are wnrdcil uB' by the vtry br
Kinnine of the niin-mm itself. All (hive ndviint«gc# are. bnwevtr, <)ail«
compatible wilh the cik-x isle lire of >>p(rcial vnsixiKitnr mrrtmniNfUH.
S 708. Moreover the Hrjv,- of lilood to, iii)d eoDMSiuent ehangc iD tfae Iwlk
nf, the braiu, ninl iiid<-eil the tlow of IiIo'mI l)iri>U);)i the brain, br iDcanrfd
by the ventwe outfluw. inny Iw modifird iinlejK-ndenlty of chaugr* in i1m
general blood-prcwiire. For iiii^tiince, i-limiiltHiiMi of ihe motor n^-a of tli»
oOJtex (juickcn? the vetioiiB uulfliivr, without jirocliieing nnr ninrkcil chaitp
in thegenemi IiIinHl-prenure: this feature becomes vcr)- (inking at thr efutl
nf epilepliform conviilsione when these tiinko their appearance. It n difi
cult not Id connect iuch a result of functional activity with some special
vasomotor nrrvmi» arr]iu)jeii>ei)t comparable to that so obvious ia the rtM
of a secreling k'""''- A^aiii, it has oeen observed that eertnin drug« lian
an elfect on the volume of ihe brain, 'luile incomincnsiiratc with ib^r ei&d
on ihe vasomotor eyrtem ; thus in juirlienliir the iujectioii into the ^DetiJ
liIiHiil stream (if a weak acid |iroduce8 a large and immediate espaosioa 4
the brain, while the introduction of a weak alkali siniilarty givea rin t*
similar considerable ehnuking. It isHUggeeted that tlieseeffeetfi are pradwod
by the acid or alkali ncling directly on the muscular ooals uf the mioiiu
arteries and ao leading to relaxaliou or contraction respectively. In treaiiiu
of the chemistry of iiervuus substance (ST2) wo slated thai " ibe ptf
matter of tlie central nervous sysiein is snid to be ali|;hlly acid durioflift
niid to become im)re acid alter death." lieceni observations gi> to »\h*9 tbt
tiM! gray matter of ihe cnrtex in faintly alkaline diirin/ life and unHrr
normal (iinditioiiii. hut beconieti acid alter dcatli or wbni its l>loi'<|.*u|ifIy ii
interfered with ; and it hiu Wn urgi-d that nerrous gray mailer like mm-
culnr siihslance tlevulii[H acidity during ncttvity, a* well a* upon death, tk
acidity bciug pnibably dui' in uic-li cam; l« Mime form of lactic Ocid, AimI
just as it bm been xuggMtiil that the dilation of the miniil« artefitiaf a
skeletal muscle, accomimnying or f illowini; the contraction of ibe ffiiHtlfb
is btQUgbt about by the' acid gcneralcil during the coDtractioii eannf i
rel«xation of the uuis<:ular ojals of (hi; minute arteries, so it has been tap
gated that a (imilnr aiidity, the product of nervous aciivitv, similarly ]nm
in nervous tissue to n dilation of the veiKels of the part, 'flie exbteace «f
special vasomotor mechanisms would, bowevcr, afl^rd a more BBlbbdOTy
explanation uf these and other phenomena ; in sjtite of the negative radu
m far obtained, the matter is obviously one needing further invesiigalitA
Meanwhilewc bare abundant evidence that, however bn^Uf-ht about, tW
tlow of blood through the broin, and probably through particular pnntnf
[be brain, is varied in acconlanco with the needs of the brain itaelf himI tW
cvcnl6 taking place elsewhere in the body.
CHAPTER III.
eiOHT.
{Phytiolayieai AnnUimy of the Eye.
Tdr «jelMll is of ft •plierol<)iil itini>e. 1( ci)ii>>!h(« of two ■Mmentl
(lidtt«Dl-m»(l apbtTML The Inr^r M-Knit'tit is »ituiile<l poateriQrIjr, mmI
coitititiitn nbrMit fivcitisihit of the wntls <if ihv <>vciliull. Prom its frM
margin prujccb' thv samllcr »cgmci)t, whicli i* llint ni ti Kmuller xfihort. Thu
puatarior wgment b oompond of a urbitish, i}|iui(ue, firm wall, cuMi»liog uf
FM.aa.
Ita.»a
nn.311— INxuBUi or 1 llaiiiMWTti. StcnuforTitt Rviaiii. n. iiuwi or ■ctimUc tutt : d.Vht
mnta : a. mfdntlo M chomlltl c«i i ■. dlUtr UvuMiit . i. 'iiUry iiiwiai -. t. «ilt*r<r niu*rl«, and
/. ifit; (. iBDoiMai«(i*aiia,o(iMl9iiiMiawiiht)Mo;>UcnDTv«|jablnd, wlUtkAtrklvv'^Mda U:
p.lM>j LBMptiMnr llMiiwDl a< tba Im*: K vlucoui boilf ; h, bralitd mcaibfwM! t, ftmmiM
abBBlMi «.<miulirfPM><: r.ilntvclwalMUUIdli: iD|«la Banc Iki douri lUw IhfiNicb Om
•MiM to Um kofllBitla^ ■>» nr Ih* tall.
Pio. lU.— TniMUb SacnoM or TUK Dmku. a P*op*( Ubus of iba conMa: Jl,*nia*i«r (tuUa
taml^of ooreflk, wllh IMha ooDlnncllTat apHlMUiun on It; C <>Wlquaabna(i»m II lollw I»)«h
o((b««(nMB: X. p»H><M «lMtla Unim. wllh r, tfilUMUiiaaa ItoTtlM membaaa of Dononn :
9. Hutee Tl*w of Uw •pliboUvDi of lit* mtmbruio 4f Dvidmu.
three watt or (tiHtei — llie itlrrniie, the ehoroul, aiul retina. The nntcrior
•egmmt u coDtinuouR with tlip lolerolio coat. ( Fig. 212.) It it a trmiispnivnt,
•iHlic, *»QV«x ufK^n, called tlve «>r«M. The cornc* OOfUiM of ihrw Inrera
— an ftnurior ami pueterior eliutk lauJaa. hnritie h«(w««n lh«m ft tarer
which i« the pro]>er lusue of lh« or(^. Tliia niitFiDc hiycr is oompoeed of
8I0HT.
Fl.j JH.
about sixty siipenrnpo^cl tnniinii* of funfdrni fibroim ocIIr. In flic intCMlod
between the Inminn; nrc tViiini] tubular »jia<Mfi, wliicli contain a inuMmmi
fluid. Tbeanteriur nnd noitcrior citulic lnmiiimnn?siriiclun-IrM anil nigU;
elutic. When Kptiratell fmni the gimiMT eornc«l liiMie thev haw k gnM
lendcney to curl up. Tlii]> fact siiggota that ihme two laminie an> aotin
n^nts in th« retention of a [imper cnrrnlurc of the cornn. The oonMa »
covered on ile anterior surfaee by the onjiinclivnl mnrwii mem brnne. which
consists of three or four layers of )>nvonicnt epithelinm c«>ll«: llt<> deoper
layers of cells are obtont;. and placed per pen ilicii la Hy. (Fij;. 'Jl.'l.) TSt
conjunctiva at this point has no perraptinle bnsemenClMinbraiM. The p>«-
letior Btirface of llie cornea is covered hy a traiMparaQt HfOilS ntetnbraBC,
wbicb conaisU of a simple layer of iKily^imal pavement epitbeliunt oclb iw-
iiig on an etat>tic memhrane. This is tallH
the membrane of Demoura. Tlie mrori
has DO blootlveesels, and tlierefure dentroi
itflniilrinietit by diiriist<Ki.
The fi-lernfie roril is M> naniH n« accom
of the Hnimnis of iu texture ami h»r<liiCK
II forms the outer tunic uf the p(»iifMr
Mgnieiil. It i* whiiwh, opn(|ue, #mii4l,
excepting nt the poinia of aitacbmrat J
the nitiiulc* of the eyeball. It U comfwnj
of white (ibraua tinue, arranged tarwr v
luM in bundle*, which interlace each mW
in various din-ctioDS. Anienorly the iai»
lacomenta are in n cenemi tr:in»vcr8odin^
tiou; pwteriorly t^e dinrvtion is lonciM-
dinnl. This coat also coniaira tcIIov
elastic fibres and fuufum] nuclealed (elk
It is ranttniiouB anteriorly with the eoiiHi.
and posteriorly nith the perinvnriiimoflke
optic ner\'e. Ac the iutemat border of tke
junction with the oonieu \* a vrnuui (inn
called the lima di*uiariA iruU* uttwuUM
Sehlenuii. The ojtitc iterve ptcrcn it aUiui
2.6 mm. internal to the an tero- poster! or axis of the eyeball. At tbit poiat
the ooat is perforated by minute o^nings for the mmig« of lb* ntn^
ftlaments. One nf tliese openintts, which iit relatively large, gira MMi^la
tbe arfTui eentralu refina. Surrounding thin point of eninocc of tba ufrik
serve are many small opiiiugs fiir the poMnge of tbe ciliary ncrvnuJ
TCMels. The intenial surface of the nt-Ieru cniiuioa mhid pigment granalaL
It i» setiamieil fV»m th« tJwnnd coat by a dolicalc flocculcnt cellular tiamr,
called the Inmini Jfuea.
The tharuiii coat i« a vaitciilar menihrunc containing some pigmenl-vcraniilet-
The cxt<Tn»l portion ix omipHW^I printipnlly of bloodvesMb and nervH,
Between the vt-wfcli^ure found niiniurous xlcllale pigmcat-cells. which furBia
fibrous network. Th" interniil ourlacc. where it is adjoined to the pigmeM
layer of the retina, atif) conlaiux pigniont-cells. I'oateriorly it is pierwd b^
the optic ucrve ; anteriorlv it is conlioiious nith the ciliary proceaMe, and M
separated from the sclerotic coat by the cilinrii mtuele.
The cii'tanj f/roceurf are iirrnnged in the form of a ring. Tbey oooakt ef
about sixty to eigblv somewhat conical 'Shaped bo<lie», situate with their
bases internally, (^'g- •U) They are plaoetl posterior to the tru.nnd an
ailaehed by their tliicKened or internal extremities to t]tenM/lefuor]r^'JrltMea'
of tbe IcDB.
Imm Viiw «r the Kuuhtt or thi
Cmanni Otur wmi m ciumv Piio-
CBMS. IMD m Kll-K OF TIIK IHM
It. >D(«nor plMD of lb« chnrali) ecu:
A. cIlliiTy |irae—L'i: c, irU; d. >|ililMdrf
nf tlie pupil : (. bantllaof niimior Uic
aOklurof (boimpll-
PHT8IOL00ICAL AN'ATOUY OF THE KTB.
8S9
THc Oi/iary mvtcU ari«r« from the pniul i>f juiictinii of lh« •clemlic eont
and th« mrntw. Il c>iii>i»(s of two givniiinf — r nilinliiig or immdioiial, and
s cin-oliir larrr. The rnilintii)); fnsdouli Mrc ntiwtwl vzt«mnlly, umI linve n
riiliuual (lirvrliuii. I Fig. '21-5.) Froai thia Imytr nunwrouff laxciculi iiiter-
Ptn ;iv
\.V
I
Samoa of thi Quoy lltatoii or ink In ix Htt>.
a, mriMloiMl niarliMr bclcull or I1» miuruloi MHani: b.dcfptT'Nalnt mtlolnt flurliull :
t, t. r. aauulu |it*xiu : d aanulitr miurte of Mulla ; / mtHriilnr lamina on It)* [•■lerlur rarlVru
«rilm Ittn; a, miuriiliiTiiU-iaiiiihodllkrr honUror lti*tni^ r. ■nnulnr (WHlou r>r IIiu riiiiOuIui
MliarM. K ll(aiiien(uni peciIuaiiiDi.
Ince betwcvn the fnsciciili of the circulnr layer, which occupies nn iiitoninl
poeiliofi to the mdintin;; layer. The ciliary niiiacle ii inserted into the
CXlrrDal Hirfuce »f the aDterior portion of the 'cli<inii<l cimt. the tibn-« cxtfiid-
ioj Minewlint p"«leiior to I he aitierior ii>ai){>t) "f thi- retina. This inuiflc i*
• rery imporlnnt factor id itie mectianiaiu o( Bccotntnudiition.
no. ■a*.
i^
MocvLtm nRiH-rrsK or mi I«M or a vriim Riawr.
d^VHanuirf ilMpu)iU: b. a, laduuiif ftadruU ol dllatw iniwole; c, c, enaimltaa umbc u-IUi
The rri* is • fibromiifculur curljiii) whtcb is auipended bct«-c«D tbc
comcn iinil the criKtalliixt l«n«. It ia attnclH^ by lla dminiH-TVnuc to (he
ititenial vaU af lh« > imi» c. iridMi Id It* eenire u n ri>u»il |icrf<irali<>n nilln)
(Im- puj>il, which i" ■uwcptiHIe of n>(i>titeml>l<- vuriali'>ti« |a sIm, Thi* nii-m-
braiiv IB cx>i»|iow(l of > fibrv cuuncvlivi- tmuv having • gtntnil rnilifttJa
8I0BT.
no. SIT.
direction ttnm tbe pnpilliiry b(ml«r. Wiltiin ihia tissue are fuund pigRif^t-
relh ati<] unstnnlecl niuecular liaBue^ Tlie niui«<ilar-lMiti« eiraitiil r>ria*iiU
of rtitUnting hdiI piradar fasciculi. 'Fif!. 216.) Tbe firenhir fiucieuli fnna
a Bpbitioter nl the [iii|iillary margin; the miliaiini; faMrit-uH radiatp frnn
the B)ibiiict«r m tbe circumference. Ai llie drrumfereuce of tbe irialbt
■□einbmi)e lining tbe anlvrlor chamber form« 6brous pfxic«wee. which ai*
teniictl thv li'jiimtnlum iridi* ftectuuilum. Tb« puMerior wirfaoe ii <»ren4
vith a pij^iiL'iilary layer, wbicli is a oonlinuatton of the pigment layer of
Uk' rvlinu.
Tbc! retina or third coat cniuiata of two [tnrtioiu; the plfcmenlary man-
bniiM- anil lerminal elunn-nU nf tlia optic nervr. The pigmeittnrif Ufnknmt
or Kiterniif layer, which ba* Imwii ralleil tbe system nf tbe unki. coven tha
whole of tbe internal durfuiv cif the dliary prnciixes. ibciriii, aoH tlH>chura(4i
It consbtF of a eitiiftc layrr of hcxnji^inat nuclfnieil pi^nwnt-ix-ll* (Fie. -I')
of a dark-browii cohir. From the internal siirfuciMtf thinmrnihranctWltaiM
fibrw arc oonlinuc) bctwi'pii the cellular dcmi-nU of the itcrvnuii layor. It
is fmqueRtJY diMeclod with the choroid coat, and »{>nk«n uf a» kd* iif ill
Umiiw. 1^ color of thn iriit in rlifTcrent imiividuals i* ih-pcndirnt ii|nD iW
derwity of the fibro-ttmnwtivo tiwuc Bntcrii)r lo ihc myvi, and to thr anionni
of pi}.'ment gmniilci* in it. In pcr«oni> with dark eye« lb« pigment ia lUi
tiB>(ie ia relatively murv abuDdaiit.
Tbe iiitrntitl or nmwu layer of the retina it cnmptved ea«entially of Ik
terminal nerve eloiueots of the optic nerve. Bxiernally it ie corereJ with
the pigmentary layer; internally, it ia lined hy
A hoinoceDeoUB tranaparenl Blructurv callnl IM
htfolotd membrane. The Mruclure uf llie reliu
ia one of great ci-uiplexity. It conaists of nbc
diKliact layers M'.wn of whi<;h are layen of
nerve elenenlti. .\11 of the* layera are bouai)
toK^lher and »iippiiried by a eonnoctlvo tiwui
which contaiuN blood vi-itvl*. Thi» layer exttwb
from the entranoc of the oittic »i>rvc In a poiai
where the aiinidRr famHciili of tbe ciliary iniif
clenrefoiiDil ; at this point tbe nervoiiaclenwiib
ocaac to exist, nud the layer bna an irrr^lw
dcntalcil margin railed the vra t.rmUt. iteyqnd
thi^, the nervous layer i« coniinued as a mtn
librriu)' exlenuBlion.
The optic nerve piorcM the sclenHi'c and lit
choroid coats, and ine pii^rneiilary meinbranoif
the reliDa. when it rapidly divides into iMl
numbera of fibree, which conaisl alane uf llx
axisMjylinders or their ultiiDBte fibrillie. Tbit
layer of fibres is continuous over nearly tk*
wliiile of the internal surface, and ia called tbt
ffcand or oplie iierve-Jibre hi/er. On ttd inlerwl
aurface, between it and the byaloiil membrane, a a delicate Mructure callid
tbe firtt layer, or mrmhrana limilaii* inlerna. Tbe Oiird or ffonglion lofm It
oompoied of multipolar ganglion cctl*. itimilnr to thoae found in tbe cerehnl
•uhManoe. Id the [loiitvnor portion of the retina these ganj^lion celln art in
flereral layen ; at tlic mnriila tulea there are at many aa eight, and at tbt
anterior portion of the nlinn I here is but a Kingte layer. From each iif ibCK
cells tifarea are cuntiniicl li> llic fifl^ or uiternit/ ffntnulr. liii)tr, which eonntt
of granular cellii with nucl'-i. nHwcen the thiril and lifth layers ia a lam
of veaienUr nmttvr containing iicrve-lihrils of extreme minulenes. tVa
r«aHit>rT-<au« or nii Ktiaiiu.,
iKOLUKKII.)
A, immincd pl(iiicnl.<iBlk of Itic
<4ia<ilil runt; K. fr>iiit tivii of tliv
hgHWIPFIfc* QollJiof tho lAgmKnuiry
iiMOibniio.
rOTSIOLOOICAL ANATOHT OF THE KYE.
en
b the fourth or inl«nial ^ranulaUii or mo/<-rn/<ir hi/rr. Tti<- liilh or
rxtrrii'il yranutiiltil or i>w/wii/ar /oyer uouista itl' |>nriillcl inU'cliived lilinv,
ootiliiiiiini: nuclei miH stuonth cell*. TW MirritfA »r extrrnai iiniinilt Imtr il
r*rr aimiliir lo llioyi/lA. The n'jrAfA luyer o>inRi#t» of n •.lelicnlo ntctnbimDe
>»f vi>nDecliT9 ti»u«, called lh<: mtmbrnHa UmiOxn* ejtfrna. The iii'h'A or
biidllury lajwr, or ftij^r <>f rid* aoi) coNm. or Jae«b't mrmbrauf, is mniputcl
of Iwo etomenla, ibe roj* nod conr*. The roU« are cvlindricul hudies, cHch
Flo. :!I9.
niLKD.
i
Tta. tia,— OnoatHHjtiii' KKnnccTji'moi nr tiib OiiniBi-niiia or im JCiaTx-nnitn i* TWt
l;rmi. l. iDomliraDB llmiiKU inlema : 2. ApUc a«rr*-nbt« laivr; S. ■>)'» utiiUNlllolmll*: I. Ib-
IkhmI BTUiiilXtd or tnolfvular Uyu : \ liilonul gritniilt l*rtr ; '• astna&l ■mnitlaWd or uiolvcsltr
■■fcr: T, cxtaraal rminlcUrtri «i mvaitiniiu Uinlumcilertor; >^ tHclIltir Urcr. or Itfer uf rod*
PMl. tl».— Rob nan C^iwii riuui txk Itcnx* cv M *^. [inwnrnl In o tm rrr niu. mliiUrai ol
pcnanteaeld lodtow tbt diKlllmiioritionrlkn. biiJ ihc iIKTitcih koiihturibi [nlpmkl •(■«•»(.
1%« oalor nsmentol ilio tmu la tm>li«4i up liilo <tl>kji. xblnb. Iinwimr, ar* Ulll wUwiWit M ono
■■Mk*r; slUit taaauriht ooiui are iMn alMr KnirhiLln. tx IIMDitluiielen.)
FN. aa.—DuaMim>UTK RrpunoiTtRtHi or ms CnsvBcnvi Tmtk er nu Rmi>* i^ «uk
sKiBmitMuilKHKirt. TbwnuiabaneatiMinnille tboMorUwnvtnlUyenodbvRaiMahoitn
Inric us
rnding cxicmnlly in a truncated, flattened eitrcniity, and internully as an
■llvnualiM] GttiT, which probnbly communieatw vritb the de«|ter lujer of ]:;nn-
eiiitn ovIU. The rones, as ibcir names indicate, ure o)iii<.'al-Bfaa|>ed budies.
VMch ciiiisisla nf twu jKirtions, a cmical btxly havin;> prujeclinK rroin iU ajwx
A rud like wgment, ubich sppeare in all n«pecU like Ihe ruda. Tbis kic-
■Mot is callcathe con« rod. Tlie terniioal eitrcinities nf llie cone rods no
not extend us far externally hb the extremities of tb« rods. The rods and
cones have been demonstrsled to vrimiat of iwo Kegmentg ur liniba, which are
CMupoaed of filawents, granular matter, and uugloi.
-1
893
SIOHT.
Tli« optic nerve, where it pierces the co*ls of the eve, projocl« •)m««)M
keyoDcl the Burfitce of the reliiia, as a ;>'i/)i//'i ; here ihe «MBeDti&l nern'<l(>
ments of the n-lina nre absent, and luminoua raya are iinperocived : hnt*,
it is i^alleiJ the blind tpot. About 'i.6 riiq. external to the point of eotnum
of the optic nerve, and in the exact wntre of the retinal surfiice correepoDil-
in^ to the tmtero puecerior axis of the e^ve, ia the " yellow spot of SonitCT-
riog" OT mtKula lutti. (I'tj;. 2'Jl.) It n an elliptieal-ahapetl apot, haran
its long diameter traiHverio. In the nenire of the macula luie« m a ilepfw-
aiou called ihefmvi e^ntralU. At thi.'> iminl the nervous layer of ll>e niiu
U very much nKHlificd in ciimpiMiiitDn or the dilferenl Inin-Tn. At the niBmb
Itik-a the nervou* layer ia much thii^lccr than at any other juart of the in«in'
braii«. The f^nglioU ({/iini) and the extcninl Kraiiulatcd (*itih) arc tlir
Kio. ai.
Fid. SZ2.
rin 2S
llRibndiiloiit. >iiti«rp|imriil(il byatisitlnit. uid In Ita mMJlclLiviUrlitinl tto itw InMM »<* H Oi
iicnt. Kith lu acciiiiiianiiiiic lulerr. (Aftor Soxhriiiii xn, )
Pio. m.— WtiiNKicD VnmciLEiETiosoriMcRinaiiMlvned (Nm KoiuKKn). *, mlonMW
■ppMMnw of the ouicr (urfai'tf of tba rctlnii am Ilia jrcltuw iput, hIicto iIhi* uc oniT pj—.l
tppnuanni of liirr n>(liu imr nm fvllox ipnl— a ulncla drcla tX rxli BununiiiMBC (UB Sdw[«
>pC**rinM or Ihu ml'l-llpaf Ibercllna, n iust numticrar rola ■nrruiiuilloKAicti ■•••. In aD IM*
Ditiins IDC lunpu rliiii> rapnsuiit [li» ouuia. mul Idu •iiiuUvr uiini tbi Mil Moa vnlvu*. lUa
Kl.ii^.l
Fin ::a.-~AKzritnncT(TKiHorniiL.tHiJ[XM ji Oahm^rd Lna. a.ikci»uc)«iB;b,«
teniiiiiv
RiMt thiclc<'ricd. The ;;iin^]imi layer Ruiiitist< of six or e!;;hl InmiiDO of eeUk
Tin- rod* of the ninth layer nrr absent, and nro rcpUoed bv ocxie*. In th*
f'lvnn ftitti^ili» the inlcrnni gnmulnlcd (Joarlh), the inteniaf grnuuli? (Mi\
mid the optic iicrvt-fibrc (trcond), nre wantine. The gBiig1ii>n ocll (rtrn/^
thi! external gr«nitliited (lizth), and the external granule (mm->i(A) lajronan
incrc3iM-d in thickiic^a. The giLn^Uun layer of oelU in the fovea eootbttif
thrcv laininn. In all nortions ot'tne nervous layer the nxlc grMtly pmUimfr
nalc in number aver tne codu, exoepling in the macula lutes, where ihtj
arc entirely absent. The retina is much thicker posteriorly, becomiDp thinner
a* it ext«n<le forward ; the nervous layer gradually disapitenring in the dsic*
rior pnriiun of the membrane.
The inloriur of the eyeball is divided into two portions by the OfyaUlliM
kiu and ite aiupensory ligament. The anterior portion ountaine the oftrnw
humor, the [wwierior contains the vitreoiu body.
The fr'j'latline lnu meuiurea about 7 mm. in transverse diameter, as4
nlMHit 4 mm. anleM-poiterior diameter. It Is a transpArenl biconvex bi)dr,
eomoirhnt Aatletied anleriorlr. It oonslsis of a uunilier of ngmenb whick
radiate from llie centre, similar lu the segments uf an i>ran^. TheM Mf>
ments are cnmpostHl of superimpoMd laminae of varyinjir density. The la-et
BuperHcial are soil and gelationut; the deeper ara relatively hard, so that
ravsioLooiCAL anatohv or tiik f.vk.
8D3
I
I
I
thev furni ■ kcnicl or ntKleua. The laminEe arc made uji of parallel fibres,
niili an iiitilulnttng ctnirw, ihe convesiti«« and roDcavilieaul ihc nttjoiaitig
fibita filling a<:curnlctv miu citch iillier. The kiis U covered with n cai^fiule
eouiMiDg ol* H Iniimjiiiretit, eluatic, frH),-iIe lueinbrnue, which lins a icnOeucy
Iti cur) U)>, wilh ill I'sU-nuil jiiirface tuiifriuiiBt.
thu MUfpevM/rir liytmrnl of the U-ii-* U I'onDCil br a conliituatioo of tliu
hjralotd iiii'tiibrnnc irhk'h lines the vilrcoti* biidy. The hvataid meinlirtine is
n <k-lic«te lniii*|'iirt'iit Mriictiire lituated hclw«ea the vitKKtus body aiid tueiu*
brana liiiiitan* inlcrnn of ibc rvliuu. Il i» continued lu fri>iil to the orn
•rmi/'i. wlierc il dtvidvt into two In^'on. The jHinti'ri'ir is lUinched Ui ihc
pc«I»rior portion of th« eti])tul« of the I<;iik; the nnterior [loriion jcradunllv
boeoRica Ihickcr a» il cxtendx forwnnl behind the cilimn,- firocettiai, and u
mtavhcd to the atiU-riur Burimcc of the cufMule. Thiv thickciii'd porti^u of
ibe mnnbrane, which is orrugatcd where il haa ailHcheil the ciliary pro-
coan, h ctillci) Uic I'mr of Zinn. ThcM: twu hiycra L-oiivtitiite the nuvjit'nwry
ligamenL Brtvrreti Ihem is n irinntnitar capal, wilh il* bune ci)rtv«{<oiiitiug
lu the cmlwlline Iciis. This is cnlksl tho cniitil ol' Pctil.
The ri'trro»» bodi/ w coDiaiiicd wilhia llic cavity Airmvd by lh« hyaloid
HMinbrane and the posterior siirfnce of th<.- Iuhk. It coiwinU ot'u clear, odor-
len, altKiniinotis Hiiiil. hitving an extremely drlicnlc- inlvrlMOnieiit of fibres
extending in all directik'iis through il. Thi'^ tilire« are not disCVTHible in
the adult, but can readily be seen in iho fixtus.
The aqitam* liumor is contained within the spac« formed by ihe poslorior
turface of the cornea, and the anterior surface of the lens. The space, which
b divided into luo chniubera by the iris, is filled wilh n clear, colorless,
limpid fluid conioing uiline and pruteid substances in solution, This Uuid
CvDMilUlea the a^unuA hujnor.
The anterior external portion of tlie eyeball, compritint; the surface of tlw
cornea and about 6 or 8 mm. of the sclerotic coat, is covered by the oonjunc-
livnl mucous membrane.]
A my of light falling on Ihe retina (tires rise to what we cull n sensation
of light; bul in order that diMinct vi>:on of any object may b« gained, nn
iniugc ol the object niiiiit be funned on the retina, and the Iwltrr define*! ibu
itiiu]je the niorti dinlinct will be the vision. Hence, in studying the pbyaiology
of vision, our firel duly is to examine into llie arrungt-iiients by whicb Inn
formation of a wlislintnry ininge on ibe retinn i» cHcictcd ; ihc^ we may
call brk-lly the ilioplric merlumitims. \Vu shall then have to iu'juire into
the laws according to which r«ys of light impinging' on the retina give rise
to sensory impulfM, and those according lo which the impulses thus gvncr-
•ted give rise in lum to sensalionitL Here wc rhall n>mo upon ibtdithculty
uf distinguisbing between the unconscious or |>hysical an<l the consciuiis or
[»ychicai liiclors. And we shall find our dilrieultin increaae<l by iho fact.
iliai in ap)>c«ling to our own consciousness ne are apt to full into error by
confuuniling primary and direct sensations with states uf consciousness which
are producvd by the wcAving of ihese primary scnMlions with other opera-
lioDs of the central nervous system, or. in fumiinr language, by confounding
what we see with what wc think we ace. Thwe two things we will hrictly
ditilioguish as visual sensations and visual juilgmenls ; and we shall find tlial
both in TuioD with one eye, but more («|>eciAlly in binocular vision, viaun)
judgnwnis form a very lai^fe part of what we frc<iuenlly epeak of as our
sight.
894
SIGHT.
Dioptric Milchaxisus.
Tht Formaiien o/ fAc Image.
T1i« eve is & «inK>rn. cotifieting of k wrw« of Icnwe mid idpiIir >m&gal
ii) a (lark chiimlicr, the iris serving ns a diophriigm : miil llie objret nT i1k
OjitHLrnlos It tu furm on the reliiiu a rlialiiict ininj^i^of extcrual •ihjei'ta. Thai
a (liiiliiiL'l iiimgp le I'trintd on the rcliiia may )>e a«cerlainci.l by iviiioi'ia|E l^
Bclerutic from llic bntk of on pye, nnd |i>i>|{)ii)( at the hindor etirtaee w ibe
IraiiBparenl reliua white raye of light jiroi^eediiig fri>ni nuy external objeci
are allowed to tall on ilie coniea.
A dioptric apparatus in its simplest form cnoaista uf two media wparutj
bjr a [spherical i aurfuee ; and the o|)iti-»l propcrtia of such an a|>p«niut
<lepend iipoo (I) the curvature of the surface, (2) the relative refradin
{Hiwer of the media. The eve cnnwsia of aevera] media, bounded bj mi
lacee which are approximately apbcriail, l>ut of diSereuL curvature. Tte
turfacee are all centred on a line called the optic ttxit, which meeU the rctiu
at a poiot Boiueu'hat above and to ilie inner (naaal) ride of the fuven rm-
tralis. In pawinx fruru the outer surfnct- of the ctrDea to tbe rvtuu ibr
rnys of lii;ht traver^ in sucecMion the cimien, the aqueniu humor, the Ua\
add the vitreous humor. Kefrnetion tulte* place al all the •urfaoe« hmoA-
ilig ihrae several media, but pArliailarly at the anterktr Aurfwre or Ik
<i>ri)ea, and at bmh the nuterior ao'l juBlvrior siirfnt^eti nf the l<-aiL i^iw*
the anieriur ami poslenur !<iirfiioc« i>l ih<- ("irnea are [innilli-l, nr very noiIy
(to, tlie rays i>f li^ht would snlTer littli- or mi ohaiige of dinn^iinn in piMii^
ihnMigb the i-driiua, if it wt^re bounded <-u linth rUUr* by ihr Knmr mrdian.
Tlw direction of llie riiy» of light in the n<|U(.i>ufi humor iroiiM. iherHbn,
remain the Mimv if the cornea wvrc iimdc exceedingly tbiii ; if. in fiict. in
two surliu'cn wvn: nindi- into one. fonning a single anterior surface to ibt
a^ucuiii> humor ; or, nhich comes t^i the same thing in the end, since (be
refractive power of the substance of llie cornea la almost exactly the nnM
as that of the a<iucoui> humor, lite rcfmction at the poelerior surface of the
conKt) may l>e ncclcclcd altogether. Tims the luo surlaoes of the curaea
are practically rrtliiccd to on«. I'he lens varies in density in dilfercut partt,
tlie refractive power of the centrnl [lorliona being (greater than ibal uf tke
external layers; hut tlie refractive ]>oiver of the vhole may, with'iut aay
serious error, be lueiimcd to be uniform. The refractive power uf the rilmai
humor is almost exactly the same aj) that of the aijueous humor.
Thus the apparently complicated natural eye may be simi>1il1i>d into a
"diagrammatic eye," in which the refraciiiiK surfaces are rwluct-cl to llint
via. : {I ) the nnlerior surface of the cornea, (2) the anterior aurfaoe uf lii»
le«9 separating the lens from the aqueima humor, and (3 f the poatvrinr n^
face of the lens separating the lenii fnim the vitreouB humor. The medii
will similarly be reilucetl to two: the subslancfi of the lens and thi' a'luvMi
or vitreous humor. Tliia "diagmmmatic eye" ia of {;reai ute in the VSTM
calculations which become neceat^iry in studying physiolugicn) nptlca: fm
the macnitudea which are derived by ailculation from it n--|^>m>cn[ tlip cut-
reaponoing magnitudes in an avemge natural eye with Kuffirnni accuracy in
•crve for all practical purpoM;*. Tlie vulum adofHcd hy Ltsttntf forth*
onvtanis of tltis " diaRram malic eye," and to him we are iudubtcd for lb
introduction of it, are oa folluw :
DIOPTBIO MECnAMSXS.
895
I
I
I
K*4ltu* ot curvature of coram 8 tttm.
of wurior furrsoe or tea* . 10 "
of poalerior ■' " . . . tt "
It«rnit1iv«index oraquMiMor vltrmua boinor . , ■ W
MfMii rrrmciivc index of len* Jf
I'lsuni^o rnmi aotrriar aurfncc nf comen (o aiili-fiur ^nrfMH of
lens -I mm.
Thitknmsor lena 4 "
Tbe culculute<l poxilioii nf [he principal p/mterior Jiteut, i.e., the {>oiiit at
which all nt}^ fulling "■> the ooriiea jwrnKel lo the opiic axU an brought to
sfiwutiU ia thetiiauruiiiiiiaiic eve 14.ri4(i) mm. livbioU tbe |>t)iiierlor Hurfiuw
of tlie lens, or 2'J.b4tO miu. behind the univrior nurliue of tbe cornea.
That i* to say, tbe fovea i-etiiralb iduu occupy thin pusiiiun m onler that a
•JiatiiK-t iiDave of » ilislant uttjeet may be fiiruir<) ufioii it. It mult be
URilemt)»tl uinl tbcwc vulues rvfer to the eye nhcii at reil, t. c, wh«n U b
not uudergoiiiK any utrain of ucrommutlatiuii.
AeeanmoiiaUon.
WhcD an object, a lens, and a screen lo receive the iinege are bo arranjretl
ill rel'erence In each uther that tbe iraajte falln upou dm vcreeit in esiiel
r<icii9. the rays ul' \'\^\\\ proceeding IVom each luminous point of ihe object
an.- hr»u]-ht iiii» f'n'us nn tbe screen in a |>uint of tbe iina^ corre»]ninitiog
til tbe [Hiint of (he ubjeci. If the object he then remuvcd further uirrnv
frutn the letis. the rayit proceetliog in a |>encil from each luminou;! |H>iHi will
b« bn>ii)(hl to a Ak-ua ai a jHiint in front of the Mcrveii, and, •uhni-igueDtly
(Itverving, will &I1 u|>un tlie screen at u circular imtRh ouoiiHveil of n Mri^
of ciivlcb. the to-called difftttion eirelai, arraiiireil cunccniricKllj miind the
priiicijMil rav of the pencil. If the object be removed, not further, but
neanr the ivtu, the pvocil of raya will meet tlie Krven befur« they have
bc«D brvtight to fiicnn in a {luiiit, nmt coiiwiiucntlv trill in (hi? n>«e nIfO
giv» riM to tliR'ueion rirclca. When an object i< placed before the eye. so
that the image falls into exact fucUH on the rclitia, and the pencil* of ravs
proccrding fnim each luminous point of the ohjcci are brought into focus
iu iNtinl* on the rclina, the seiiiatioii called furlh is that of a distinct iniagv.
When, oil the contrary, the object is too far avay, so thnt the focus lies in
front of tbe retina, or Ion near, to that the I'ocit^ liea behind the retina, and
tbe pencils fall on the retina nut at |K)iDt«. hut as syeteraBof ditfiision circleo,
the wiintioii produced is ihac of an indistinct and blurr«d in^. In order
that objecU both near and distant may be ^«eu with eqaal ilutiueiueas by
the tame dioptric apparalus, the focal arrang«menla of ihe anpoTUus iDUat
he artommodaltti to the dislaocc of Uie object, either by chaagiDi; the refVac-
live jMwer of Ihe lens, or by allcriDg the distance between the lenv and tltc
•creeu.
I'liai the eye does pomeea such a power of acconiuiodnti<Hi i:* sliown by
e%'ery-day experience. If two ne«<lles be lixe<l upright aunie two feet or •!>
aMii into a long piece of wood, and the wood be held before Ihe eye so that
the Detdlei) are nearly in a line, it will be found ihat if aiteiiiioii Iw direvtcd
lo tlie lar needle, ihe uear one apf>eant blurred and ijidiatinct, and tluU cun-
vciMtly, 11 hen the near one b dblinct, the far one apiiMLni blurred. By an
effort of Ihe will we can at pleaaure make either the lar one or the near one
diftincl ; but uoi both at the «ime lime. When the eye b anangcd m that
Ihe far neolle ajijieani diatinct, the iuuige of that needle falls exactly on tlw
relimi, and ■■ach pencil frrmi i-nch luminuun [lotnt of LIm lieeille unilei> in n
point u|>on ihv rctiiut; but wlten this is tlte chm ibe fociU of the near needle
SIOBT.
lies beJiintt lIu) retina, mxl vncli pencil from mch tuminuut )i»ini <if ihit
needle iitlla upon the rerina in n acrttx or itifTitflon circles, i^imiinrlr. ohen
tb« eje is nrrangvd so th»t tiiB nuir nrc<IIo w diMincC, the imaj^ uf Uial
Deedle fulls upon the reliiin in encli a way, tbat eacli |iciicil of rnys rrmn
eAvh luiuiaoUB point of the nccilk- unites in a point on the retina, wltili' Mck
pencil I'roiu encii luii)ino<iH |>oint of the far nec^lie unites at a pi'int in /nml
of llie retina, and ibcu divLTgin^f attain fdlls on ilie retina in a iteries uT dif
fusion circles. It' the near needle be (jradLisily brought nearer and nearer
tu the eye, it will lie fuunil that greater and greater etiort is re<)uin!d to tee
it diatinctlr. nnd at hiel a jioiut is reached at vrhicli nu eObrt c«U nialie tlie
iaiai^e of ibe needle api>e.ii' anvlliin<j but blurred. Tli« diDiance of ihit
point from the vye murks Ifie timil of aeconiiuodaliiM
riant. for near ubjccie. .Similurij-. if the pentou Im short-
eigliled, the far needle nmj be moved aw&y trwa iIm
eye, unlit a. point is reaebed nl which it cemaet lo be
seen difltinctly. and apjiears blurred. In tb« ooe cut
the eye, with all it^ jHiner, in unable to briuj; the ItiMgc
uf the needle aulBcienily forward to fall on the reiin*;
the fucuH lien perinaucntly l>vhiii<l the rt-tina. Id Ik*
other the aye cannot bring tliv iuta^ ntiHicivnlly haci-
ward to fall on the retina ; lh« focu» lin {rrtnaneatlf
in front of the Ktina. In both ca»ei< the [KuriU uf ra;(
from the needlea strilcc the retina in dilfu^ion drelo.
The «ttnic [ihenomena may Iw ibowti with grcaicr
nitety by what iit called Hcbuiner'» FIx|>orimmt. If
two itnioDlb hotf« 1)0 pricknl in a card, at a diatowr
from cJich otlKT Icsx than the diouKtcr of the pu]iil,
and the card be held up before one eye. with the hula
horizontal, and a needle jilaccd vertically be looked M
tlirough the holes, the fullovrin)* facts may be ufai«mi):
When attention is directed lo the needle itself, the iau|e
of the needle appenre sii));le. Whenever the gue »
directed to a inure diKtanl object, so that tlw eyu it m
luDjter accommodated for the needle, the iningv uppMii
double and at the same time blurred. It nl»u spjmn
double and blurred when the eye is accumiiiodatvit for
a distance nearer than that of Ute iieedli'. When unly
one needle is seen, and the eye lliervfore in pnjpcHy b»
comimiduled fur the diniauce of the nceiilv, no wlMit it
produced by blockinic U]> one hole of the card, rxocpt
that the whole Held of vision ieetna diiumvr. Wbcii,
however, the image is double oii nccount of tb« rre be-
ing acciKiiaKidated for a divtanee grtMer than that i4
the needle, blocking the left-hand hole vauen a d'mf-
pearoiice of the righl-bund or opjHifilf ima);e, vul
blocking tbo right hand bohr niiiiea the lell-lniml imiigt' to diMipfx-ar. \Vb<u
the eye it aoconimodated lor n divtanco nenrrj than thiit of ihv needle, block-
ing cither hole causes the iningo nn the nmo udv «i \'nnisli. 'flte aitoK
dingrsiii "ill explain how time results ara brought nbouL
Let u( Fig. '2'Hf be a Inniinnii* point in tlte nettllv. and » r, n/tJw extreoM
right-liand and led-hand ray« of ihc pencil of rays proceeding froni ll, and
passing resjiectively tbrough the right-hand e, and letl-liand /, boloa In llie
oard. (The figure ia stimwtMd to be a burlsunlal section of the eye.) Wbtn
the eye b accummodaieil for n, the rays « and / meet together in the |>o(di ',
UiD rettDB occujiying tJi« inwitiou of the plane x n ; llie lumiuoiu (wiut a^
DuonAM or Hrnmiis'B
ZxrauHot
DIOPTRIC MEOUAKISMS.
807
I
pmn w nnr iHiint, Hill] iliv iwc(lt« will apiHtir u one occdlo. When llio ejro
■> MoMninoliilcxl for ■ ilinliinrc be.vniiil •!, lln- rottna mny bo cuit>i<U'rfd t» Ire'
no Im^fCr *l n n, Ixit iM-urvr the \iiu*, nt in >n for cxitniplc: the mp a e will
Ml th» plftne nt p. ami Ihe raj:* a/Mif; heDC« tlio luminous point will no
loDgtT ft^i|H-iir •iiiglu, bill will In? xnn lu tvco poinbt, or rnihor a* twn «vs[ein»
cif SitTiiHuii circl««. iiod the sin^to iivetilv will iip[icnr m two blurrvi] dmhIIw.
The ray» niu»ing thr»ui;h the right-hniHl hole «, will cut the rvliun at » — iV..
oQ ttto rignt hanil »Mv of the ojitii.- axis ; but, u ne shnll •«) in #|>oflKiiig of
(he Judgtnontii pFri«intiig to vuion, the imagtt on the righl-hniiij stdi; of the
ntina '» rrjerral kv tht mind to an object on the Icft-lintiil lidc of the dlt-
•on; bunco the attectiou of the rt-tina ut p. j>rtKluccd by the nip a r lulling
Ml it tlirrv, give* riie tii nn iinngc of the sput a at /', and »ii»ilarly the led*
luind »]Mit 7 corresponds tu the right-hand '/. UWking the Ictt-haiid hole,
lhcrrfi>ri-, caUMX a disnppcnrnnce of the right-hand tningv, and eier verM,
Sintiiarir. when the eve is nccoinmoduled for a distance nearer than the
OMdlt, la* Klina may (k 8uppi«ed to be removed to i I, and Uw right hand a c
■nd teft-haiul a f rays. a(Wr uniting at <r, will iliveryo again, ana strike thq
retina at p' and '/. ' Tiie blucking of tbe bole e, will now cause the disap-
pearance of the image ij' on the Ifltl-band aide of the rvliua, and this will oe
rererre<l by the mind to the righl-hund aide, so thut <^ will seem to vanish.
If tbe needle l»e bmughi grmlually Dearer and nearer lo tbe eye. a point
trill be reached within which (he image b alway* double. This i>uinC marks
Willi oooiiderable exactitude the near limit «f ac(»>miuo)}aliun. With aburt-
eighted [ler^ms. if the needle be removed further and further away. n point is
rMebed lieyund which llie image ia always double; Uils marks toe litr limit
oT Hooramodalion.
The experiment may also be |>erfurmed with the needle placed boriiou tally.
Id vbich case the hulcs in the cnnl «liuiild bo vorticnl.
The adjiutnictii iif the oytj for near or fiir diitancM may be a«i»tml by
usinj; two necditm, iintt ueiir and one liir. In this case one needle should \h!
vertical, and the other horiminlat, and the cjinl tnmoil nund so tbnl the
ho)ca lie horixontally or vcrticnlly according to wliethor the vcrii^ud or
hnHx»n<al ncrdle in bring mailo l<> appear double,
III wlini may hv regarded iis tbe normal eye. the so-called emmHrapk tvn
[Fig. fJO], the near limit of accommodation is about lU or Vi cm. and the
XHairiiunc Sts. FASAUit, mi.n ruiv«u or t«i anal*.)
far I'lmil may be put for practical purposes at an infinito di#lancc. [Fig.
S'id.] Tbe " range of distinct vision." therefore. (<ir the emmetropic eye is
very great. In the tnyopie, or short-sighted eye f Fig. '£"£7} the near limit Is
bruuglit much closer (5 or 0 cm.) to the oomea ; and the far limit is at a
Tcriable. but not very great distance, so that the rays of light proceeding
frofn an object not many leet away are brought to a focus, not on tbe rfltiM.
■ or«oufH.n th»aoiiwi tro. «■ «r« *t»tl m*. aamrnaalMMNt
■■4 DMtTMialHnUniiaiWfaUtkmarilieMUtiai Ht IDr«
|HM tlM IWIBB to M monU.
67
. tatiboud lijackuiM UthaWai.
OMnataaM «fcSb *b nsf turn lap-
SIGHT.
but in ibc TitreouH humor. Tho ran^ of ilintiixit virion b, tbc^^f^rp, in ih^
myopic eye wry limiird. lo the kgptnuriropie IFig. -IMIJ, or lonK-Nk'tilal
eyv, thv rft)'H or tight comiDg from cvtm nti infinite ilisuincv *re, in the |m»-
Rvo BtAto of ifae eye, brought to a IWcw bpyood the mfoa. Th« nrar hmit
[Fm-sa
DMmono Etk. Tbc iloit4>d lluo* iliowliic bow HconuuvlKtJan lut iba iUtmvI'C bj* i' mv
«t4«0titi«SkCiM]
of >ccmimo()utioo is «t tnme flistADcc olf, nm) a hr liniil of nrcomimalAliM
doff not exict. The pr^jiAijnyjiV eye. or tho long aight ofohl f<»plu, nMild>hl
tbc hvp<Tiiii-trx>pic eye in the liistuiw of the near {xfiul nl' nn'ommndMioa
but dittvn from it innnmuch a* the former i« tai w«cutially defective ooadi-
ine. m.
m
UVDiic Rte.|
lion of the aocoRimodation ine«hnniain, vhereu in the Inlt^r the povtrtf
accumuodatioD may he good, and yet, flrom (he tnternal arningemenis itf ll»
eye, be unable to bring the iiuH^ of a new object on to tl»e retina- Wb«
m Dorcnal eye hecoiues presbyopic-, thu fi>r limit may remnin the Mtnc, but
llTrKHHimioric Evi,]
since the power of accommodating for near nbiecta is traakened or loit, tbt
chaogtr is di*lincl)y b reduction of tho r«0}^ of disliuct viiiun. In th» Mr
ma) emmetropic eye, vihen no effort of accommodstion b nude, the priDei]Mt
focus of the eye lite on the retina, in the myopic eye in front of it. and ii
the hypermetropic eye behind il.
Mtthtnitm of atvommod'ttiein. Id directing unr aUeolion from a fiir Ua
very near objeci, we arc codecIous of a ilii»[inct eSbrt, nod fv«l that «0M
change has taken place in llie eye; when we turn from a very near to a &'
object, if we are cuoBcious of any chanj^ id iii« eye^ it m one of a diflfenni
kind. The former is the ien»c of aii active acoamawdatjan for near otyetfi:
the LaUer, when it b felt, is the seuse of relaxation after esortiou.
DIOPTRIC JtECHANISSra.
899
Siuct the fkr liratl <if :
I <lislnnr«, no ituch
r-ii)tiui(iYi|iic v.yt u ii
notivr nrmmn) (Mint mil llirfitnlii'tiiiu'cs miil viiNt. The. niily chiingc
ihat will liik^pliK'i-'iii th«: ovc in turning rmni nvjirtu liiniliJMtswttl lien atvm
|wiwvvui»daiDgortbcHCGODiiiiod«ti<in)>r«viou»tyiua*lvfortbcne«robjeicU And
Uikt no »ueh Kotive Mcomoiodation Tor fur dbuuiM tnkt* jilaci- in tliown t>v the
IkrU-tUftt the eve, nhen opened nlUTlx-ingelosod for some time, is fount) not
in mealium Rinli.', bitt adjusted for illMnnc* ; tbnt when ibo aeoom modal iun
n>n-liiini>m "f ihc eve i« purnlrtnl liy lUropinr or ncrvou* Hi»cnw, the acconi-
ni<xliilii>ii fur (lii>iiiiit objn-l!i i# iiniiHcr'Irfl ; iind llinl v,c «re coiueious of do
cflbrt in Itirning from m<Hl<-ralcly ilisliint (« for ili«liint objcrW, The senw
of effiin ol^cn «p<)ken of by mvr>nic pcnionB a» being frit when tlwy al-
leni|)4 to BM- tilings nt or bevnnJ the Inr limit of their range teeniH to arise
from a movcmeni of the eyelid*, and not from aiij' iiileniar changM taking
place in the eye.
NVIint, (lien, are the changes vrhieh take place in the ere. when we accom-
nodnle for iK-nr objeels? It niighl lie thought, nnil.inilced.onoowaa thought
thni the curvatures of the eorne« was chnnged, lH-i.iiming more convex, with
■ shorter ivdius of curvaliire, for near ohjeets. Ynuiig. however, showed
that aotrom modal ion took place as usual when I he eve (and head) is immeraed
ID water. Since the refractive powerv of BC|UeoUB liuntur and water are very
nearly alike, the cornea with its parallel surfaoee, placed between these two
fluids. caD have little or no effect on the direction of the rays passing ihruugli
it when the eye is immersed in water. And accurate ineasuremeiitJi of the
dimensions of an image on the oomea have shown that these undergo no
change during aceoniinodation. and that therefore the curvature of the
coniea is »i>l altered. Nor is there any change iu the form of the bulb;
for any variation iu this would necessarily uroduce an alieratiuu in the
curvature of the cornva, and prtmurc on the Dulb would act iujunounl}- by
rendering the retina ansemic and >0 leM aeniitlve. In fact, tbvm an- only
iwo cbangrw of ini|iorliince which can he nnccrtaiDcd to take place in the
eye durin;; nt-coriinioilniion fur near objects.
Our i* that ih<- iiuiiil ['onlriii-ts. Wlii-n we look at near ol>jC4:l«, the pupil
becunuw sniBll ; irlu-n wc turn tn djuiant iilijvclf, it diliittv. fhin, hnwovvr,
cannot have more than an indirect intluenoir on the lormatiun of the image ;
the chief lue of the contracltoii of the pupil in nccomniodauon for near
ubjecta is to cut olf (he more divergent circuralvrrntial rnys of lighL
Tbe other and renlly efficient ehnnjn- i« that the anterior surface of the
lens beramca more convex. If a light Iw held before the eye, three reflected
iniagi-s may, with care and under pri>])er [ireeautions, be seen by a bystander;
one a VL-ry bright one cwused hv the anterior surface of the cornea, a sevund
Im> bright, by tlie anterior surface of the lens, and n third very dim, by the
posterior eurlace of the lens ; when the imagM arv thusc of an object, such
Bs a candle, in which a top and bottom can be recogniKod, the two former
unages tre aeea to be ereirt, but the third inverlerl. Wbeulheeye is aecoiu-
mwuMd for near objects, no change is observed in either tfae first or the
third of tliese imagiM ; but the second, that from the anterior sur&ce of the
lens, ie seen to Wcunie dislinetly smnller, showing that the surface has beoooie
more oiuvex, When, on the contrary, vision is directed from near to far
iibjects, the image from the anterior surface of the lens gron larger, indi-
cating that the convexity of the surface ban diminiebetT while no change
takea place in the nirvature either of the cornea or of the posterior aurfaea
of the lens. And aceumte measurements of the size of the image from the
anterior surface of tite leas have shown that the varisUoos iu curvature
which do take place are an flicient toacoount furthe power of acoommodalion
which ike eye |)
SIOIIT.
Thsobavrvntionof ttiMprvRccW IihicmU raci1ilal«d bylbcalaiple inftnimtnl
Intrnducril by llclinhollx iitid cnlleU « phakotcopc. It comiata of a pirhII, ilifk
i.'liiimti«r, wiih n|ifTiurM fur the bbMrv(>.l ami obwrving tyw, a nerdlf i* ttud
at a ilKiri iliHiuiice in Tront of lli« fomuT, to Ri>rv« h a acsr ol^fct. Tor «liicli
HGcoinmnHalioi) him to bi> inatl<- : and a lamp iir cnndlc i« *o dUpoaed ■• to tlunw
an imnf^ »ii rni-h of the Ihrrc Kurfncn of ti)« utnerrej tje. Since lb« dlaUst*
between tau iinHgeii is more rmdily a|ipreclat«il tliaa !■ a ai(n|>t« change of ru
or a iinglt lintige, two prisms an employed m> tut lo lliroir a double imagv of iht
lamp on each ot the ilireo fiitrrac«>. Wlien the anterior iiurfB«c of Dm tm
become* more convex the iir» imagtv rctlec.ied from that surface appnadi twk
oilier, when it becunin lew efiives they retire from t»vh other.
Tli«te obeervaiions leare do doubt tlml llt« etaential clmnee by wliici
ncc<Jium(j<)ftiiou is eHi>cled, U lui alteration of the convesiiy of the ititervir
iturrB(<e of the lena. And that llie lens id the ageDt of ac^iminodatioii b
Turtber sliuwu br (he fact that aAer removal of the lens, as iii the o[)eniiic
for (fitiirurt, the power of accouiiundalioii h lo4t. lu the ca^ee which faan
lieeti recorded, where ey(4 from wliii-h the lenH lind b«fii removed eecnttil
>lill to puniteM vome iii'i-(>iim)o<lBtioii, «« niuni tupixii^ that titj reul iccutBii»
dalioii tuok placo, Init that the jiupll crmiruciea when u neur object km
liiokeil at, mill xo arwiptcii in ninkiiif; viiiiiin more diatinci.
Thi» incrraw; of the convexity ot the li-ioi \ih» b«oii »upp<jsed la be doele
n coniprtsvioii of the circuniferc'ncyr of the Iciu by a conlntctiuti uf ikv farii;
but this is disproved by thp fact tbnt accoiniiKulation may tukc place in IJV
from wliich the iris u miigeiulallv abwiit It ha« aim bwii attribulei) U
VRRumotor eJianges, lo incrcjiM'tl liiloen of Lh<- vimvU of (he irii or ciliar;
procMMs, niRoundine ihc Icno : but th>« alv) in disproved bjr ihc fuvt llnl
nccomtnodntion may Tic effictcd. nftcr death in an cyi> whicli i* nrai-ticallj
bloodless, by GiiniuIaiitiK the ciliary (runglioii or ciliari' nerves with an inla-
runted currcuc or by other oiennB. The real nature of the mecliBtiiatu mvim
la De ae I'uUuws.
The lens when eiamiued afler removal from the eye i» found to be a hcrft
of con aide ra hie ehisiiciiy. When tlie curvature of the anterior aurfamnt
the lens ia determined, as may he done by appropriate means, in iia naturtl
pufeition iu the eye at real, and then again detcTTUincil. nl\er ilie k-iiH bait hcM
removed IVom the eye, the aiuerior surCuce is found to tie more ci>nvex in tlv
latter than in the furiuer case. There seems to be. in the eye in ii» natvtil
4<oiiditiun, winie ajtencv ul work, keeiiiug ihe anterior surface of ibe tfM
•omewhat fiattened. The siixpeuaory ligament, attached to the choroiil Mi
viliar)' procctwe* behind, and paMing over the fntnt of the lena, b jutt «utb
a fttnictiire aa would iiroduce this efltcl. In the iiatitrnl pu«it)ou of ibt
choroid thi« ligament u tcnxc, and leml.-i to lliilten the front nf (he leM
When iho choroid u pulled forward, ibe lignmeut becomes alack and lb*
knit bulge out forward. Further the cilian' niuacle nitachod im tb«a*
band to a fairly fixed region, the junction ol the fcleroiic anil cornea, aai
on the other U> the loiner and more movable chnroid, would naturnllv, abn
thronn int<> cinilniclion, pull foruard the choroid and «n nlacken the aaann-
wiry liganivMi. and hence i>ermit the elastic lena to bulge out furwnn). AaJ
we have cxiurinH-nlii! evidfOfc. carried out on lower imimata, that nimla-
lion of the ciliary ganglion or of ilf »o^called radix brevia, tlott lead oB lU
one hniid lu a conimciioii of the ciliary nin«elc and pulling forward of tbi
choroid, and on ihv other hand to an iucren«ed ewrvaturc <>f the aniemr
Burfaco of tbo lens. Hence ire may conclude that aeooioiDodaiiiin fur oca)
objects consists e«Kiitinlly in n contr.iction of the ciliary tnutole. which, by
pulling forward the choroid coat and the ciliary process, sliK'keiia the nuipco-
sorv ligament, and allows the lens to bulge forward by viKue of il^ elutidly,
HDa eo to iDcrcato the convexity of its anterior eurfitea
DIOrTRIC yECHASlSM8.
£101
I
I
AccOfumoiliiii'Hi ia in moat cnwa a voltininry hcI ; »inc«. homsTar, thv
cbuigfi ill thi- kiM is ulirnrs accomiHiDicd l>y movvtikOtite in tho iris, it will
Im coovcuieBt to cooaitler the latter, oefore wq disuiMs tho nerrous mechaDism
of th« «liol» u-t-
JiwaiunU of thf pirpit. Thriu;>h by makin); ihe pflbrts required Tor ac-
CDtDiDodatioo ire can at pl«D»ur« contract or diiaiv tlit- {>iii)il, it is not in our
txia'pr to brinj; the will to act dirwtly on the iris by it»elf. Thia I'sct alone
udkatca tbat tiie nervoue meebaniam of the pupil 14 of a peculiar ubaracter,
and such iodmd we 6nd it to be. Tbe pupil ia coiilnteteti (1) when Ui«
miaa (or optic nerve) h atiinulutod, aa wlieo li^ht falls on tbe retina, the
brighter tbe li^bt tbe greater being tbe coniraotion, (2) when we Rccommo-
dale fur ne«r ubjects. The pupil is also contracted when the eyeball is turned
inward, when the aijueous humor is defirient, in the vat\y mma of imianuiiifc
hf cblorofonn, alcuhol, «tc. : in nearly all stufcea of poisuning hj morphine.
pnTaoMiginiue, and aume otli«r <lrugii, and iu deep slumber. The pupil ia
diatUd (1) when Hliniulntiiiu of the retina (or uplEc nerve) ia diniinulwd or
atmied ai in piiaaiuf; frum a bright into a dim light or intn dnrkueai, (2)
when the eve u adjusted f»r far r>l)jvi'tA. Dilation alan (Hvum wht-n tht-re i*
an exccMofaqtieoua humur, duriiif;dy!>pnwa, during vinluiil mti^cular tflurta.
aa lh<^ ruiult of a Htiinulaiiun of Miiisurv nrrv<«, ua an cffcci of emutiona, in
tlir liUi-r vtiigcs (if jiniaiittini; by clilurutiiriii, etc., and in all stages of ptiiaun-
iiig by atrnpini! nnd aiimc other clnt|ni.
(.'ufi tract ion of iho pupil ia cnuain] by <vintraclioii of the circular fi\>x*t or
sphincter of the iria. Dilatiun is caused by cuntnirli'>n of the radial fibrot
of the iris; for though the cxialence of radial librcs boa been denieil by
many ohsarrcn, the preponderance of ovidoncc i* clearly in bvor of their
bdog really preacnt.
Coinidt-ring liow vascular the iris is, it does not seem unreasonnbU! v* in-
terpret some of the varialious in the cuditioD of tlie pupil as the results of
aimple vascular lur^feoceace or of depleiion brou^bt about by vasomotor
action or olherwiac, the small or coiitracled pupil corresponding to the ili-
)atcd and filled, and tbe liirge or dilated pupil to the constricted and emptied
condition of the bloodvosela. Tliua slight iiscillationaof the pupil may l>e
observed syuchmnous with the heurt-lieut and others eynchrououf with the
rapiratory movements, llut the variatioUB in the pupil seem too marked to
be merely tliv rlfi-cta of vaacular chnngi^, and indeed that consttctiun nt tlw
pupil <«iiBot be wholly (he nwult of lurgescenctf. nor dilation wholly the
rvault of depletion uf ilie vca*«l« uf tbe irii, ia shown by the fncU thai both
ibeae event* tour be wilm-Mwd in a pvrfvi-tiv blnodleaa eye, and tliat the
movement* of tli« pupil ulxen brought about by agents whidi aliu nlTtvt the
liloodvcawlt, begin a»ni« time bvl<)re the chaiig<ai in the cidibre nf the hloud-
v«taeU,sDd ind««il may 1w over brtorcllicac have arrived iit tlietr maximum.
Moreover the fibr« of the aymimlhciic, which. aa we shall ace, arc ooniwnKtl
in cauiing dilation of tho pupil, run a (otncwhat diflcrent murac from those
which govern the bloodvcaacla of the eye. We may, ibcrtfirrc, adhere to tlie
view that the main cliangea of tho pupil in the dirt«ti»n of nnrrowing and
widening tirv brought about by contmctJona of the plain inuecular fibres in
the iris.
.Muscninr contractions lending to changn of the niipil may be obferved in
the eye fvmovcd from the body, and indeed in Inc extirpated iris. The
plain mUKuIar tibreeof the iris like other plain muscular fibres are remark-
ably sensitive to variations in leni|>erHlur?. Betides this there seems to be,
in ccrtaio anioiaU nt leant, a connection within the eye between the iris and
retina of such a kiuil, that 1i;;lil falling into an extirpated eye will lead to
uarrowiug of tbe [lupil. Putting aside, however, sucb exceptional events
aiOHT,
ri<i.9sa.
wc mar lar down tli<! bnmd priDciple iliat coulraction of ibe pupil, Itrmiilit
•boiit hy Ufcht ralliiii: i>d tbe I'etiua, U a reflex aei, of nliich tbe 'ipiic b ibe
•flb'eiii n«rve, tlw lliinl or Dculii-mctor the eRl-reiit atrve, and ihe owtn
•otnc jiiirtiiiii ■>!' ilw liriiiu lyiiix beluw ibe i-orpont riuudrigcniina in the frtni
pHit or ihe ficicir of ihe aciucduct of Brlriun. Tbi* iit pnived by the follow-
tug fnvi»: When the optic tierve is dividt-il. iliv fulling of ligbl un th» rettna
iH) li'Dgcr vutiri-H n vuutructiuii of (lie pupil. When the third at-rte it
'livideil, xlimulntiiin uf the reiUia of of
ihv (ipiic nerw iio lnDjcer cuiuc* nift-
Irni-tiou ; but direct uliniuliitioi) uf tb»
[loripbc-ml iHirtimi nf tbi- ilividMi ibifd
iKTVt CHUMS' fxtrvRW coatrudi'io tf
the pupil. If tbc region of tbe brwa
ofHikcn uf iilxtvr a« a centre be «■!«■
fully itiinulmi-d contimctigii at tb
pupil will tnkr pIiKi>: vwu in tbe ab-
wnnr of light iiiid n)U-r divi^iim of tb
optic norvc. A(1>t rcinoral uf tk
Mine oenlrc slimulniinn of tlM rdiaa
is inctft-ctunl in Rnrnning th« pupil
Bill if tbe centre aixl iu cnnoetiiw
with ihe optic nerve and ihiid Mm
be left iDtad and in tburougbly kmbJ
condition, contraction of the pupil viO
-7c.
U
'-f—s-c.
occur aa a reeult of liifht falling on ^mJ
r~— -w
^wT"
PinnnuiHinc BirnDiENTiTrrrN or mx
XUVM (ImiKNUIU THE PCIir.,
reiina, tbou);b all other nenous
be removed.
The Dervoti* centre ia n<it n dnuhls
centre with tim> completely indepcndeal
htilve«. one for each eVK ; thcro ■> a n^
tiiiii amount of funcliimiil c<nniiuDaiM
l)etM'cvn |h«< two xidcs, »•> ihut aLtn
one rctinii in Hlimolaled iKiib pniii^
contrucl. It might be imagined ihst
ihif cerebral oenire acted to- a laaie
ccnlre, wbofe actitMi wM dimply !>■
crcaMd not orijjiiiated bv the •Itniula-
lion of the retina ; but this is disprivn)
hjr the fiict ihiil, if the optic nerve l»
divi()cil.subc«i(uent rection of the thin)
Dcr\-e produces no further dilation-
In omiMdcring tbe tnovenieuta nf tbc
pupil, however, we bavo to deal Ml
only with a narrowinf; of tbe pupil
thus brou;>bt about in a rcHex wav \^
cotilrnctioD of tbe circular aphiiicttf
fibres, and with llie abeeoce ol' eurbi
uarroniuK- but also with active diW
linn due to a conlructinn of tlie railial dilator fibres, and Ibia tenders the
whole nuittcr niiicli m'>rc cuinpleK ihan might be auppuiied to be ibveat
from the simpb- itiitcinciit just made.
The iris ia «upplicd, iu CDniDion with Uve ciliary' mtiacle and cboroiit, bj
Ibc abort ciliary ncrvm (Fis. 229, «.e.) comiu;: fruin tbe o|ihihu]mic ur
tcnticular (ciliarv) enuglion (lo.) which In Bounvcled by iu mou wilb tbt
third nerve (r.fr.\ we cervical lympnihetio nerve (jiym.), and with tbe
IX, opiic Dorrv : 1.9, IcoiIcoMi gkiigliun :
t.h. II* tliun max ftom til: oc.ki, iblH at
oculooiolar nirre : fjnn. \i» >)rinp«ltiiiUr nail :
r.L IB Iddr nwl nnm V. opWAn. Uir twutl
Irnnch at Ibv u^hUialnlc dlvlilun of ilit DRb
nrfTS : t. c, Uiff *hait olllurr nnrvn Dotn llis
IcoUoabr (uiBtlon ; I.e. tbe loot tillirr
n«rn Cmm ib* raal bmach of Uw opbitial-
Dilc ittTMon at ibo nftb Mm,
DIOPTRIC HBCHANISUS.
903
inch ftftlic nitlithnlmic di
I (r. (.). The t\*an dli«ry
I
I
I
ncrvr» «nt, mnrvuviT. iicc<ini{iHnir<I by llir Inti^ cilinrjr ii«rrc« (I.e.) coming
from iIh' Miiiui DsmI branch of iho Diihthnlmic iliviRion of (ho fifth n«rvo,
Wbat uri; Um asM of theae Mv«ral nvrvnt In rrliuion lo the ixijiil T
If the ctrvical ■yDipKth«tic in thv neck Iw ilividrd, nil otner portioDS of
the nwTOUs inocbitDtaiTi bein^ iiiinct. n conirnction of the pupil (not nlvayt
wry well mnrkcil) tiikw place, and if llip pcriphcrnl portion (i. r.. tlic iipfwr
portion utill <!<innecleil witli the hend > be Htimiilftted, n well -<lo vol o)icd diln-
lion w tki' mull. The ayoip&thetic hu, it will be oh'K'rveil. an rfTt'Ct on
ibe iria, tbir oppcaite of that whicli it eiercisea on the blundvcssiiU : when it
i» Mimiilnied llie pupils are dilated while the blooilvoMcU nrv onelncted.
Tbta iJiluini; iiilluenee of the Bynipaihetic may, as in the cue of the vaao-
motor action of the aame oeri-e, be traced back down the nock to the
npper thoracic ganglion and thence aloof; the rami commuoicantes and
raote of the h>wer cerviciil and lirtit dimal or (wo Hrst dorm) spinal oervee,
to a rej^n in llie lower cervical and upper doreal cord (called by some
ftulbors the eenlrum ciUiytpini/e inferiiu), and fnitn iheuce un ihriiiu^h the
medulla obloo);ata to a centre, wliich ap|>eare to be [ilaced in t(i« floor of the
front part of the B(|iiediict of Sylviiid not far from and appsreatJy on nlber
aide of the centre f»r cimtmction of ihe pupil.
The dilaiiiin of the pupil which in wiineMte<l in dyipncea. and that whidi
nsulu from •tiuiululinn of n'nitory ncrvvit and from emoliuna, appeara to he
bn>U){hl alxiiit by ihr action of the aympiuhetic. thu venous blood, or the
aOMory impuiiwii or the ciiioiiomil impiilws »>> iifll-cliui; llie dilating; centre
U to an^nni-iit th<' lUliilin^ iinpiilvN procevirui^ from it hIoii]|C the xyni|M-
ihetic. The i-xiiitcii(.i' of the snbnnlimite centro in the ci-rvical or doml
eoni, *{wkpii of just now, i* Kuppwcil to be indicalrd by the fiict that after
divinon of the niaduUn oblonj.'ata, and conWi)uent Deverancc of the eHurenl
paths from the centre in the aqueiliict of Svlvitie, dilation of the pupil may
still b« brought about, in some animals at feast, by dvBpnwa or by adequate
stimnlatioo of sensory nervea. A question ia raised herv in fact somewhat
similar to that raised in connection with (he me<lullary respiratory centre
{)>. 47iJ); and here as there we may pmhably conclude that the independent
action of such a spinal c«utre is of Hubordiiiaie im[<ortanoe.
The pnpil then «reuia to be under (he doruiniun of two antaifanlslic
neobanuiDa : one n ciJiitracling m«chanl*ni, rellex in nature, (be third nerro
•ervinft as the efr<'ri-iil, iiud th« optic as (be afferent tract : the I'tUvr a iltlul-
ing methaniim, appun-ntly tonic in naltire, but mbjucl to au;:wpntation
ftnm variouH causca, and of tin* tlve cervical aym]iia(betic it the pITcrcnt
channel. Hi-nci.-, when the (binl or optic nerve u divitlw), not only doe*
contraction of (bo puiiil coue to bo manifest, but active dilation occurs, on
account of (be tonic dilniiiig tnflurnoc of (bo sympathetic beloK left froo to
work. When, on tite other hiuid, the sympathetic is divideil, this Ionic
dilating tnlluenrc falls away, and contraction re»iilt«- When the optic or
thini nrrvt is 'limulnleH, ibe dilating dfect of the symjinlbelie is overcame,
and cuDinK-tion results: and when the syinpnthotic is atiwulated, any oon-
traciinic inHLieiiec of the third nerve which may bo prweat ia oreroomo, and
dilation en sum.
But (berc are oonsidcradons which show that the matter ia Mill DH>re
complex than this. A small quantity of ntrtipine introduced into the eve or
into tlie system causes a dilation of the pupil, This might be atlrihuted to
a paralysin of the third neri'o, and, indeed, it is found that after alropiae ha«
ptijdueed its effects the fulling of li^ht on the retina no loo^i.'er cauMa outi-
tnetHHi of the pupil. A ditfiouliy, however, is introduced by the fact lltat
wbto the third nerve ia divided, and whea, thorefore, the conlnicUug oflkvli
SH3BT.
of Ml! mil In lion of tli« relina are )j)ac«il enliral)^ ou one itde, anil titerc
noUiiui{ lu prnttut tbe ity in pat be lie pmOucing its <iii«iin^ vffccta to ibc
utinoKl, dilntion it mill further iucreawHl liy atruniob When )>bvs(»li|pDiM
in inlroiliiix'il into the tyn iv ftyitcin, ouTilniclion of ll>c puful i* omad.
wh«tli(T tho ihinl iiorve he dividMl or nut ; nnd irbni tbe doM h nSkJCiUjr
Htrong t)i<; <'<>n tract ion in so urmt tbat it <ntiiiul \}c urercomc by stimulalita
of tli€ M-nipnthriie. The dilHlion which la cKUM-d by a »ufli(-i«ii <lcwo(
fitrupiiip nmy he grciitiT than ihnt which am onlinarily 1>K priducvil by
Kimiiliilinn uf the sympniholic. nud the ('<inlmetion cnuoed l>r ■ *ullid«M
dose of }ilmi»tigni)ii<- miiv )w grvitlrr thnn ihnt vibioh is orilinaril; nnr
ilticed in n rrtlfx iimimtT i>^ j^iRiulatinn of tW optic ntrve, or C%'«D tun
that produced bv dirrcl eliniulniion of tliv third nerve. Ei-idt^ntly llio*
druee net either directly on the pinin niUK-iilnr tibrts of tbo irin or on ««■«
looal mechanism, the one in Mich n wiiv tin to cause diimtiun, tli« olber in
■udl ■ wsy as to canse contmclinn. 8i)cn ■ local niechanisiD cnnnot, how-
ever, tie in the ophlhalroic ;^nglion, for both dru(^ continue to prndoof
tha<e tllecls in a most marked de;;rcv aller ihe gBD>;lion has been I'XerciMd.
We mutt suppose, ihercfort', that the niechantsm if it exists is sitiiatnl a
the iris itself or in the choroid, where, indeed, jtaDKltonic uer\'e cells an
abuiidaiil. Tlie niovemenU of the iria in the extirpaletl eye. afwlieu of jmI
tiow, may perhaiw be altribuled lo the same local mechanism. Further il
Ih siati-il that with i!timuUtion of the ermpaihetic, the latent |>eriod, ■'.<-., Ue
)>vriiHl iuierveninK between dte betfinninj; of aiimulatiuit and the liegiDiiia|
of the movement of the iris, is much k'V'>I^ tl"i" **'<'h Hiintulaliou of ibt
thirtl ucrve, iiidicattiij; that tbe former acta tlirouiih a local iue4,<lmuiain b«
tbe tmicr more directly tin the muscular llbrea. Tbe nbole rpieatioii, hint-
•wr. of thi* local mcchaniam, and of the exact nmleoractioa nf tbe varinui
drup and of lh« cbangei in ihi^ body which luad lo (wntrac-tion or 'lilntM
respectively of the pupil, netnli' fuller di«cuHiuu than we cui affofl to gin
to It here. Wc may ad<l thiil the local ncliou of nUopinv in oonlm^i muj
action on the rcnbrnl centro is well illiiKlrnleil by applying atn- ^
eye locally- Tbe pupil of that eye dilnti-a widt'lr: i<> ci>ni>i.->(i< ;«
light linlls un tbe retina, nnd this ro uflecis the cerebral centre, ivbich aa mt
have M-en is not slrietly unilalemi hut in communion with its iVIlufr, UiU
increaitud constricting impnises pan from both centres, and tbwc. iho«igh in-
ellectiial in the airvpinitcd eye, lead in the untouched eye to an iacreaued
narrowing of ihe pupil.
Th(! share of the Mh nerve in the work of the iris aeeme to be in parta
•entory one; the iris is sensitive, and the sensory impuUce which are fta-
eratcd in it pass from it along tbe fibres of the tilth nerve- Moreover On
lidb in peculiarly rvluled lo the dilating eflecis of tbe syinpathetir. Far
though t^e opbtbulmic ganglion does receive libres directly frooi Ihe carerti-
ouf pilous <if tbe dympaiheiic. the dilating aOion of tbe aympatbctic
woulil m.'cm to lie carried out oat by thate UhivA but by Bbrea jiiiriiaii; the
GIt.h nvrve. and {)na»ing to the iris not by the ganjjlioa but by ibti ophihil-
mic hritiich nnd the lung ciliary nerves Tbe vil-mi motor fibres of ihv ■>*»'
pathetic, and tbngw which dilate the iris, al^vr riiiining together in the ribib
Oorvioal vyoipatlietic chain, purt compiiny higher up, the latter piuuuox I*
UiaOflMerian j^nglioo. and thus nuicliing the niiMil braudi of |]ia nphtSll-
mic division nf thi- fiAh nvrvc. Sumir ntti^TveRi maintain that in addition
to tiKVo dilating Hbrc^ of [hi; nv in pathetic joined to it, tbe fifih coiitun*
fibre iif iis own which bIho am able to dilate tb*; pupil.
We ninv mm up the ncrvims nievbanivm of llw pupil then Miaowbal •*
f<jIlow«. The «alivnt and miHt frcigiiently rviiDalcil event, tl>c oontrvctioo nf
Ihe pupil, upon cxpa«urc to liglit. is a t«fl«x act. (be centre of which i*
placed 10 the brain ; and the correlative widening of the pupil upon dinii-
DIOPTRIC XECUA.NISMS.
906
I
I
I
nnttnn of liglit in iliie to ili« innicHciion of th»ityin|MUbelic mnkiiiii iiwlf
Mt upnu the nmiiiiii of \u nuinpiuiiil. Tho «;iiiilrftctwn of Uio piipii in the
nrliiT utatrra of lUc Hctiiiii uf bIooIioI ■m) chlorofonn kik) in Hiunilwr U
ptulxililv <lit<) to Ml iiicnWwNl acliiiii of llu- cootmcling t.-mm. ImiI lh«
uBrniK ]>ii|iil cnu»oil by viicli ilrugx M morphiiK* unit phvuneli^iiiio i» iluo.
chivAj' III l*-n»t. to K Idcnl nclion. The (jilaliiig i>trvrt)> of fueli dniev ne
Urouine nn- ■!»» luri^cly due to a locnl nctinii, but in t)i« «'i4leiK'<l pupil of
iba lutvr ttiu^ of kloohol puisoniag anil of dyspnoMi we c&ii prubnbly trace
cIm dTccta of au exhiiuslion of ibo cpivbral coiilrading cei>ir«, MHsted poe-
liblv by sn tncraiecd actiritv of lh« ililatiBs centre.
Tlwn* iviuniit^ h notd to be said conceruiiiK the conlraclion cf the pupil
which takes place when [he eye is acci>iniii(Hlat«(J f<)r near obj^cta, aiid when
the |Mipil it lurneil inward (the two beiii); clusely allied, »ince ihe eyes {am-
verite li> aee uear uhjeclH). and ihe return to the more itilaled condition
nhtn tlio eye returns to rwi uud re^tii"* 'he accommodation fitr far ohjecta.
Tkcae are instances of what are called "asaociated muvemi.'nt«." Two
motenieuts are Ibua apoken of aa "Mandated" wheii the a]>«cial central
MTTOoa nieclianbni empluyctj in carrying out the one act ia to connected bf
nerruui tiw of some kind or othvr with that eiiifil»yed in carrying out tlw
oiber, that nben we «et the one inei^hani«iii in aetiou «re unintentionally act
the olhiT in ai'lion ulwi. Tlie ciliary niiiT«-leii wliieh brlus abimC acoiniDO-
dali'in arc governed in ihii action by fibres which may lie traoed. through
the luliniy Dervir:< and Iriitieular Ka»u:li<>ii. alonj; the t1iir>) or <h~i]1>i- motor
Dvrvv, to a ci-iilrv which Iic9 (in ihigii) in the hinit part of ihe door of the
thir<l ventricle, and which in enpei'ially oonni^clcd with the mo»t anl<-rii>r
btiadle* iif the r«iiit:< of the third nerve. Thin centre i« nnder the command
of our will; witeii wc wish tn nccnmmodntti for near objecia wo throw it
into action, aud it, when in action, chHd also into action by " aMocintion "
the Nnire fur the ooutraction of tlie puf^l ; wlien the action of the accom-
modation centre OMM* and the eye falls back to the condition of rest, in
which it is accoinmodated for far objects, the action of the piipil-contnictio([
cetitre ccasM also, and the pupil therefore widens.
The iiieclianisni of accommndAlion may al«o l>e effected in a local manner.
And tite ilruKS which have a sjiecial action on the piifHl. fiuch us atropine
aiid Calabar lieau, also alfect Ihe mechaniani of acconiraudalion. Atropine
liatwlyxca it, to that the eye remains adjusted for far objetis; uml phyio-
•tlKnunc tbnm'a the eye int<i a <-ondition of furceil accommodation for near
objeeU. Tliia double acl»<in luia been explained by the HUppovitiou that ubilo
■Iropine paralyti^n. pliyanalig;n)ine tlirows into tonic or tetjtnic oontrtietiim.nn
the one hand the circular mnacla of th« iris and on tlie otltcr the cili-try
louK-h*; bttt the pheiiomeDH, on inquiry, appear loo complicated to ba
expUiaed in aoaimplv a manner.
We can BceommiMlatc at will ; hut few peraoDS can eflect the ni.-cr«i«ar]r
cbangeio the eye unlr» they direct their attention loaome neJtr or far oliject,
aa ibe case may be, and thus avisl their will by visual M-JiMtionK. Ry jinM>
lice, however, the aid of external object* may tie diipcnaed with : aix) it i<t
wlieo this is achieved that the pupil may »ccm to bo nuuk to dilate or con-
tract at pleaaure, accommodation being cSoclod without the eye being turnod
to any pantcuLar object.
Imperfeetioiu I'n Ihe Diaplrie App(tfiilH$,
The emnwtrojHC eye may l>e taken as the normal eye, The myopic and
hjrpermelnjpii- c\e* mav be cunaidered as imperfect eyes, lhoui;h the former
oerliuu ail van itt^ over the n'>rmal eye. An eye mighl be inyupic
810 DT.
fw>m too i^TeMt a conrexiiy of tlie cornea, or of the anterior iurfiiw iif '
lens. OT fnun permnn^nt »r>ni)ni of tUe accomiMnd«ii"ii-mech«nwni, or fn
ton Kreut a len|[lh "f f hi- ]<mic axl* of the eyeball, TUe last ajipear* to b»
the UBital (-iiuw. Siiniiurly, mo»t hy|K'rmi'trrt|>ic eyea powctb loo cliort a hiilh.
Moreover in Ihetttronglymarkwl inyopi*- pve there w fWquentlv hypertnifihT
of t]>e lonKitiidinnl (meridiniial) lihrts nf ihi- ctltiiry mu»cle, often upokwi (/
«xeluure]y lu the cilinry rawwle, nod atrophv or alm-iice of ihf rimiUr
fibrai; ill the hri>Tr(H'trcipii" pvc, on the othi-r Ituiii). tho rirrtilar flhr« art
well devdopct! luid the mrriilionrtl Rhm si^nty. Tlii' |tml>y<i|)ie eye ia. h
we havp it-en. iin cvd normnllv conallliitivl in mhit-h th« inwer of m-wwniwi-
dntion hiiN iMt'n lo«t, nr is fiiilini; thmnch increioing wenknen nf the inliary
mi>«dc or a loM of elaalicity in the lens, or through tbe pArta beconinf
ripiH.
Sphtrwal abemfiott. In a Fpherical l«i» the rayn which iinplnice on th#
rircumrerence are brong^ht to n fociia soonirr tlian thoM which p«M nnutr
tlie wntre, nti<l the rsys praceedin^ from n himiDoii« pntnl nrr- no Inasfr
brmigbt to a mnt^le focus at one point hut form a number of fiict ml fUffinMil
distanflos. Hence when rays nre allowed to fall on the vhole of ihi- k-iin. thi
imaiie formed on n ecreeti plat.'ed in the focus of the mora c<-niml ranfi
blurred by the diil\tB ion -circles cait«ed by the circumferentiiit raw whiA
have been br»u(rht to a premature f»cii!i. In an ordinary nr>t>i.-(il in'tninMet
Bph«ricsl aberration is obviuled bv a diai>hragni which ■hTiiii nfT the iwn
circiiniferential ra«. In the eye llw iri* ia an adiiirtabtc diaphmi^a ; laJ
when llie pupil contracts in near vision tlie more diversent ravs priM-efdinc
from a near ohj eel, which ten<i to fall on the circumferential parts of tHa lot
are cut off. As. however, the rvfriiclive jwiwer of the lens doee not IncrctM
renularly and progrewircly fn>m the wnire to the circumfertnee. bul rari*
mo«t irrec'ilarlr, the purpooe of the nnrmvioi; nf the pupil cannot be timplt
to obviate Fplierical aberration : and iniWd ilic other <)[ttieal imperfectKM
of the cvo nre so srrtt. that wich spherics) alHrralioia na are caused by lh(
lens produce no obvious effeol on vision.
Antiirmnliun. We have hithenu treated Ih* ere as if its dioptric mtAm
were all pnnsof iwrfeot spherical auriiicen. In reality thi* i< rarely the cmm,
either with the lens or wiib the cornea. Slight doviatioii!' do not prndun
anv marked effect, but there is one deviation, knowD aa regular a*ti|croaii>"<<
wbieti, preeenl to a certain extent in most cv«», very lar^ty developed fat
Bome. fre(|uenlly leads to very imperfect vision. Thia delect la d>ie to the
dioptric surface beinj; not spherical but mora convex alont: one meridiaa
than another, more convex, for instance, along the Tcrlicnl than alone th*
horizontal meridian. When this v the case the rays pmceuding from ■
luminous |MHnt nro not brounht to a ein);le focus at a point, but puawwi t*«
linear foci, otie nearer than the uormal and correspond injr to ihi- laurr iiinvat
surface, the other further than the uormal tociis and cornvponttiii),' In thn ha
convex Burface. If the vertical meridians of the mrfaca be n>on> oonva
than (he lioriruntal, then the nearer Huear focus will be horiconlnl and tla
farther linear focua will lie vertical, and ptcr verfn. <This can tw shorn
mud) more effectually on a modet than in a diagram in which we are limiltJ
(A IwodimeusionH.) Now, iu order to see a vertical line distinctly, it ia muoh
more iiuportanl that the rays which diver^^elrom the line in a acrica nf har>-
innlal planes tihould be brought to a focus properly than thow which divet^
in the vertical plane of the line itaelf ; and nmilarly, in order to see a hcn-
lontal line diMinctly, it is much more im^iortani thai the rays which (liret)^
from the line in a seriea of vertical planee should be brought to a fociu
properly, tlian those which ilivertie in the borixontal plane of the Hue tiAtf
Hcuoe a hoiitonial line held before an aitigniatio dioptric aurface, idmC
DIOrtRIC XBCBAinSUS.
807
i
ronvox in rlir vcrfical rnpHdiniif, uil) ^ve riiw to ihp iniocv of a h'lrin'iital
line nt iW iioin-r t'uciie, ihe veMical rny* iliverKinj,' ftoni Inc liur being lirrw
btDOgbl to a liiwar hi.iri«iiilal I'ocitB. Hiiuilarly, a verlicnl line h«li) bofurt;
the aBni« enrface will give rise lo an intake of a vertical line at the furihcr
fucuf, llie borizAiitnl mrn iJi\er),'ii](; t'toiu llie vertical line bein^ bete bntiight
to B linear verlieal fueua. In other words, with a dioptric surlnce nmet
convex in the renicAl meridtuDa, horizonlal liuee are broughi to a focua
aoooer than are vertical Viote.
Slnst eye» are thus nior« or lew aflticroaltc. and (tenerall}' with a frrealor
cnnvexity aUng the vertical lueriOians. If a ua of horito^iial or vertical
lines b« IooIcmI bC or if the nt^ar piiiai of Hrc'iuiuioilatiiiD be deierDiiDed by
Scbeiuer's experimeul (p. 8U0K for the iietslle plactd firac horiiuin tally and
then vertically, the hnriziontal li»« or nM'dle will be duttoctly visible at a
tliorter distance frum the eye tlinn the vertical tinea or needle. Hiniilartr,
the vertical line niunt be furihcr from the eye thun a horiituDtal one, if both
tit to be seen dtMinelly at the wine time. Tin- oaiiite of asiiK>nati»ui is, in
tbflgmit niuj'irity of ciiseii, the uni'i)iiMl curvature iif the mniea ; but some-
timea the fault lit* in the lens, its was the caM with Yoiin){.
WbeD the curvature of the c-nira or lens dilfcn not in two meridiana
uiily but in wveral, irregular B.-<tik!mBti.ini in th« result. A oertain amount
of irrefcular asti|[matisin cxisU in tii'Hit leniv^ thus causing the image of a
bright (U'iut, such as n star, t» be nut a circle hut ii radinic figure.
Chrrtmnlie ahrrrnUon. The diflercnt my» of tho «|ieclriim nri- of dilfemil
rrfningihiliiy, tlitsM: townril the violet end of (hv sDectrum Ix-ing hroiigbl to
n f(K-tU wHiocr thun those near the red end, This in optical instruiueiil* it
oltviateil hy using conit>ou»d lenses made up of various kinds of gliiM. In
iIm oyc wo have no evirlenoe that the lens is so conslitutcd as to correct tbb
faolt ; Mill the total dispetvivc power of the instruinctit i? so smnll, that such
itSOUBt of chmmalic aberration as does exist attracts litlle notic*. Never-
tiMtcHsoRiaslijihl aberration may be ilelec'ied by c«rel\)l observation. Wlion
the sfm'triiiii is observed at some disliitice the violet end will not be ecwd In
focuB at the Mime time as the re<l. If a liiniinuua |)oint Iw hioked ai through
a narrow orifice coveie^l hy a piece of violet glnsa, which while shutting out
the yellow and green allows the rei] and blue rays to nuw thd'ugh. there will
be seen allernnlcly an image hiiviiig n blue centre with a re<l fringe, or a r»l
centre with a blue fringe, ac(^or(ling us the iuuige of the point Itmked at is
iJirOfra on iin«aide orotber of the true focus. Thus suppoting /l Fig. %\fi.)
to be ihe plane of tlm mean focu* of A, the violet rays will be brought lo a
Via. ro.
DiioaAM lU.ivnuTDKi CiiRminr AsissinMi,
(t i> il>i itlapifl« (Diftcs : trnt*wsta ibc blun. >nit Srihe r<1 ajn: flstbglbesl |dinaar tlia
blur. Jt uf Ills red nn ,
toKW in llie plane V. and the nA rays in the plane ft. If the nija be aujp-
pus(^l to fall >ti\ the retina between I'Bnd/, the diverpng or blue ray* will
form A crntre xurroundeil by ilie still converging rrd rays; whereas if the
ray* fall •■u the reitnu bi-ixeeu/and H, the (Mtnverging red raya will form n
ccDtfv with (lie still divcrginif blue rajra furmiug a fringe around tiMBk
SieHT.
tbe mrs Tnll an lh« reiinn ni/.tbo two kiniU ufmn will he misM togtlbir:
U will be Mt« fniii) th« ttgun, ihc circiimlvmilmr Mill ooavttrging nd n;
A r M it outc ilic plniiu of th« rvtina ii, in ordinary vistua, ■coumpnniiic) tif
the divrrfpng vidIh rny A (i. Mud thus by n sort of compmniion, mt m
logctlirr even (lit- myit which <liffbr moet in rvfniction.
Eiilojidr phi"iom«nii. Tli« various inmliu of the eye are not itnifotmlT
IniiiitfiHmit; thi? rays of light in pRuing thnm^h thein iiiidrrgo local
iihKi>r|>li<>n und rtfrnclioD, imd ihus various (hndows nre ihniwn on lh«
n.-linii, of nbtcb we bocoine cioitci'ius lu iiiiporfection? in ihc ft«ld of vinoa.
Mpecially wbcn the ey« is dir(!<.-te<(l to a iinifonnly illuiniunlcd aurbce
Tbew tin spoken of as eotoplic pheoonieua, aoil are very varied, iniuiy forw
having been deecribed.
Tlifl ruoBl cominon are tbcee caused by th« pf«sence of fiualiog bodis in
tlte vitreous humor, the so'called tini»tat I'oiilanla. Tbeao are reatlily wn
when the eye ia lunied toward a uiiifurni atirfuce, and nre freiiuently vttf
troubleeonie iu Wkinji through a niiL'ro«c(i|>e. Tbey are opectnllr obvioui
when diverKCDi rays full ui>oa the eye. They aaiume the firm of ru«i aad
S [roups of l>eadit, of single ttesda, of sireakit, {mtchts, and )(nuiule», and aaj
le reco^'nixHl by their alnioal conliuunl tnoveiDuiL, ca|>ei-tally when the htat
nr eye ia moved up and duwu. When nti atti^inpt \t made ui fis tbe vwica
upon thi-m. tbey immeiliately float awar. Tearw on tbe conit-n. t»iu|¥mry
uueveiiiiciw na the aiilcrior surface of the eonie* al\or tbe eyelid tuu btas
pnneil on il, and iiujwrfci-tinii* in the ten* i>r its capsule, kIimi give rise tn
visual tinngiii. Not uul'rei|iicntlv a rndiitt« liKure corr<-fl{K>nding f> Uw
■rrangciDunt of the fihns nf llie Icnii makcgi t|.-< appi-nnincv.
Iin|ierfoctioii.'> in the luarj^in of the pupil a|ipi'fir in the rhadour of the irit
which bound' the field nf vinun ; and the mdvonieiilji >>r the irii' in nne «y4
may b« reiidereil visible by looking at n bright point or luminous surfact
through a pin-bole iu a cnrd pluml close in the front of the eye, in the
anterior foous in fact, and then alternat«ly elosinj; and o[)ening ibe other
eye ; the tiehl of the Hrtt may be ohserved to contract when li^ht enler^ ami
to eipnnd when the light is shut otf from tbe seoond. The iiie<lia of iheeVD
are fluoreseent : a condition which favura the pereeplioii of the ullra-TiuW
rajs. If a white sheet or white cloud be looked at in dAyligbt tbruuifb a
Nieol's prUm, a somewhat bright double cone or double tuft, with the apion
loucliiog, of a faint blue color, is seen in ihe centre of the field uf vitian,
crosaed oy a Mmilar double cone of a somewhat yellow darker color. Tb<M
are spoken of as Haidinger's brushes ; they rotate as tiie prism ia rotated,
and are nuTiifised to he due to tht? nne<|uul absorption nf the iNilarJuHl li^hl
in the ycthin ^[lot. The primti tuu»<t l>e freijuently rotaicil. n» wlten tW
prism reiuniuft at rest t)ie |ilien»ineiiu fade. LuXly, the otxioil nrnDge-
nietils have a further imperfecliou in that the dioptric turfnctt nre nut Ire)*
ceutjed uu the uptic axin.
Visual SESSATioxii
Light falling on the retina excitca seiuory itnpuUet, and the«e paaung op
the optic niTve to certain parts of the hmin. produce ohaonB in ccnail
cerehrul i>(riir;tur«'«, and thuH tpve riwe to wbtit we call a teiuttluut. In a sta-
aatiou we ought to be ahlu to diHtinguiiih brlwvca the events ihrougb whid
tbe impact of the ray* of light <yn the retina is enable<l to geiicnite etOMfy
impulser, and the evenU, nr rather Mries of events, through which ibat
■cnwfy impulsra ifor. jud^'ing hy tbe analogy of motor nerves, wc have on
rauDtl to think that they undergo any fiindanieulal changes id psetiing nhiaj
tbe optic nerve), by the ageney of the cerebral arrungemeoia, (Hvelu|i iuix >
TTSCAf. SENSATION'S.
909
rinuntioii. Huch an nniilrtiii*, however, h, at present nt lea>t. in mort pnrtlcii-
Un, iiuili- l>evi>niJ oiir )iawer ; iind ne muiil therefore treat ut' the seDBaiioils
tia n wliolr. (Ii*lin);iii>hiiif; iK'tucr-ii ih« fverijibenil ami ceiitrul pheuootena, on
th<; mn- octuai'im when nv arc iible to dg m.
The OrijfiH of Vutml ImpuUet,
Of primnr;- importfluni to lh« uiKientamling nf Ihf wnj in vhich lumin-
am andiilationa giro riM to tboec ncrv»ii» change* which pa» alon^ the opiic
nerve ua vignnl itnpulwfl, n the fnct thai tho rays of light pmdiicv iWir em<ct
by acting tiot on (lie optic nerve iiielf hut on ir» terminiil organs. Thrj pnu
ihrotigh tbc anterior lareiv of the retina iippdrently without inducing any
edcct: it is not till they hQve reached the region of the n)d» and cones that
they set up the cbangrs ciincemed in the gfneration of visual impnlpcs; and
Ibe inpnUM here geiiemtr<l tmvel hack to the layer >if fibres in the anterior
■iTftoe of tbe rellDft and ibonce pass along the optic ner\-e. Thitl the optic
fibres are themselves tnscnuble to light and that vixoal impulses begin in
the region of rods and cones is shown by the phenomena of the blind ep>it
and of I'urkJDJ^'s figures respectively.
Jilinii »pol. There is one pan of the retina on which rays of light fall-
ing give rise to uo sensations ; this is the entrance of the optio nerve, and
Ibe cnrrespniiding area in the field of visioD is called the Mind spot. If the
vinal axis of one eye, tbe right for instance, the other being chised, be fixed
on ft black s|>oi in a white slbeet of paper, and a small black object, such as
tbe point of m (|uill jwu dinped in ink, be moved fEmdually sideways over
the paper away to tbe iniiaide of the liehl of vitiou, at a certain distunee tbe
black point nf the <]iiill irill tiimppear from view. On ci>nliDuing the move-
nH'nt (till further outward tbc point will agaiu cnme into view and cnntinnv
in sight until it is Inst in the |H'riphery of the field of vision. If the pen be
used In make it mark on th<> paper at the moment when it is lost to view,
and lU the moment when it comes into sight again ; and if simitar marks bt
niKiia along the other meridians as well as the horizontal, an irrcgiitar out-
line will be drawn circnmscribing an arm of tlic field of vision within which
nya of light produce no visual sensation. This is tbc blind spot. The
dimeDsions of the figure drawn vary, of courw, with the distance of the paper
fironi the eye. If this distance be known, (he sire as well w the position of
ibearea of tlie retina corresponding to the blind xpot maybe calculated
from the dingrammatic eye (p. Mt4v. The prisiiion exactly coincides with
tbe entrance of the optic nerve, and the dimensions {shout l.>^ mm. diameter)
•Im correspond. While drawing the outline as above directed tbe indiea-
tioM of the large branches of the retinal vessels as thev diverge Iram the
cntnuce of the nerve can frequently be recogniied. Tne existence of the
blind sjwl is also shown by llie fad that an image of light, sufficiently small,
thrown upon the optic nerve by means of the ophthalmoKope, gives rise to
no sensattuns.
Tbe existence of the blind a{)ot proves that the optfc fibres themselves are
InseMible to light: it is unlv through the agency of the retinal ex|>ausiuit
that these can be atimuhited (>y luminous vihrations.
Pitrtitijf'4 fipin*. If one eutcn a dark room with a candle, and while
looking at a plaiu (not iiarti- colored) wall, muvea the candle up and down,
holding it on a level witn tbe eyt-s by the aide of the head, there will appear
in the Arid nf vision of the eye of ihv same iide, [ifojecie>d on the wall, an
image of tlw n<linal vmmIs, Qtlilc similar t» that seen im luokini; into an vye
with the ojditludiBOMOfie. Tli« field of vidian is illuuiinutvtl with u glare.
dio
SIOUT.
and on tliia th« branched retinnl reMela apiwnr lu KbiuUou,-!. In tlii* mocU
of eipcrimeiitiiif; the light eot«n lh« eve ilimiigh the (^inii>n. nixl nn traue
of thi! nuulle ia furinei) on the duhI mle of (he retina ; guid it i* thn ligSl
eniniintinfi from this image which throwt ihadoirA of the rttiomi vmhIi on lo
ibo read of the reiitia. A fur better mttluxl b for « Moood penon to omicm-
tratc the rays of light, with n lent of hiw |iowcr, on to the uutatdo of tb«
sckrotio juHt behind the cornMi; tlic tight in this cb»o rtDBiiiitw frooa tk
illiiniii)ated S[Hit on tbc Hdi^ntic nml [loiHing Mritiebt thmiigh the rilpeow
humor throWH a direct vhiubiw of the vtmel* on to the retina. Thus the nji
ptw<ing tbrtiugh the tclerotic at A, Fig. 231, in thediredion br, will throw i
ebsilow i>f the Te««et ' on to the rctinnat .'; thi# will np))«ar iwadark lincu
11 in the gluri' nf On: li<,-]d ijf vi;<irti). Thin provv tbal the Elmcture* a
which vL'<ua] impulMw iirigiiinle mtiiil tin behind th« retinal vewcts, otfaemin
the »lindi>WN of tbc^c could not be porceivod.
If the light he mciveil from A to a, the «hndow on the retina will mon
fhtm a to ", and the dark Iin« in the field of viiion will more from B to X.
If the distance B A be nMiosured when the whole image is projected at a
known distance, t B from the eye; k being the o]>tical centre,' then, knowing
rm. III. FiQ. at!
JfeoRBllI nXnTRATDN Tlti FOUATIOH At
^gaai)i»> nDiiRMwan rut lLi,niis(Tio:<
nuoaut iLuenuTDM m Tnnuneal
i-i iiKisuf • navnta wimi nta lummm
Ik IiiBscrxD nnociMi not COwtB*.
the distance it i$ in the dingrammutJc ere. the distance ,' « can be calcabUel
But if the distance ,^« be thuv estimated, and the dbtanoo &a be directlf
mnuurtd, the distances ^I'.nv.i-i. <iv can be calculated, and if the appearaoo*
in the field of vision is rmllv caused by the shadow of '' falling on ,i, thew
diiitanccs ought to correepoiid to the diHtancesoflhe retinal veeeeUi fronlh*
sclerotic b on the one band, aii<l from that |>»rt of the retina ,i where Ttwid
impr.-mons begin, on the other. H. Miiller found thai the dialanix- ,) i- thai
calciiliilcd corresponded to the distance of the retinal reanU fr<>iu the lam
of rods and cones. Thus PurktnJ^-'H figures prove in the first plui.-« that tbt
•ensory impulees which form the commencement of visual lensatinus oHgi-
Im» ncBf tu iicHuiloc (nri^eo. at about It mm. 4Uuac< iniai (bi rcdaa.
-I
TIBVAL BBXSATtOKS.
911
naie
pttK of tlie rc4inn Itdiiod tlic rrliiiiil vcmels.
tome pttK ot tlie rc4inn tiohiDij the rrUiiiil vcmels, i. p.. loniewhora
lielweeo (Ih>iu aad the c-Iiorui*! coal : and 11. Mullt^r'it cak'uUliunf go Ikr to
^ow liiat tliey originate hi ibe niuitt pusl«rior or exirtnal pun uf tliu Trlina,
vie, itie layer of rods and cones. It muel be lultniiml, tiowcvvr. thttl 11.
Muller B TMuliB were Dot sutBcitrDtly exact to allow any great ecnws to be
plac-etl on tbis arKumcni.
Id tlie secuod inctiiod of exftcrimentiD); tlie imoKe always inovea in the
OBine direclioo bs tbe livht, aa it obviuutily uiuat do. Id llie flrai nivlbod,
«b«re the lifilit eoiera tlin>Uftb tbe cornea, the iioafice luovee iji the unie
direction u tbe lijjlit wIivd tbe li|{bt in moved froni tifi,bi to lell, pruvideil
tbe moveiDeiit does not exiotid l>ey<tud tbe middle of tbe vorDea, but iii ibe
(ippowte direclion lo the light when ib« latter it aioved up aud dowo. In
I-ig. 232, whieh rei>r<*ciitii u liurix»utnl aeetiun of no eye, il a be tuoved tu ii,
b Dill utDVe to ,i, tbe dlinduw on tbe rvliiiii c to r, otxl tbe iniuge d tu i. If.
un ibo other hand, k be Hiijipiwol tu iiiuve ubove the plane u I tbe jiajwr, fr
will move below, lu funi>et(uttii'c c will tuove above. Mid </ will a{ii)iear U>
niovc Ix-'low, t. «., d will aitik nn a ri.'>cii.
It it dtairwblc in thii»: cii--<u> to move tbe lt){bt to and fro, i-M[>erially in tho
fitat tDCthwt. all the relitiH nnun In'i-omi-n lircd, ami the inrnj^u liidiv awiijr.
Sonic ubKrverv can rvcognirj- in ihc axiit of virion a tnitit hImiIow corrv-
•pODdiug tu iht ed^ of ibe ile|iri«iiun of Lin- fovm cmiliulis.
Tb« retinal vewcU niny iilao lie remlervd Tisiblo b; looking through a
■mull ontiov. nieh a* n |>iii-bole in a i-nrd plaeed vluM to tbe «ye, at a bngbl
Ituld uncb a* tlie Ay, ami moving the uritiw very nipi<illy from ^e to Mdo
or up oud down. If the Riovcnicnt be from nitlc to «ide, the v«mgI« which
run vsnical will be iK-en ; if up and down, tbe horizontal vcmcU. The liDft
capillary vtvmI* are seen mom niHly in tbis way than by Piirkinji-'» method.
'Vliu same uppcnraocee may also bo produced by luoktog through a micro-
Kope from which the objective has bL>en removed and the eyepiece only left
lor in irbtch at least tber^ is no object dialiDOtly in focus lo the field;, aud
moving the bead rapidly from side to aide or backwanl and forward. Or
the inii-rosco|ke ii«ell uiny be iiivvv<l ; a circular movement of tbe Held will
iben brinj; both tbe vertically aud hurit<uulaUy directed ^'eoMU into view at
tbe Mme time.
TiK pkolt-riiemiAtiy nf llie rdhia. In AeekiDK to understand how it ia thai
ra)a ul liuht lalliUK upm tbe ru^iou of the hkU aud ivimw can ^nve ri»e to
wniurjr, viiuial inpuUM Id tbe optic ncrvo, we may adopt one or other <if
two viewa. Od tlie one hand, we may »uppiw« that the vibratioiiH of the
Ctber arv able, tlirough the uiimna of tbe rctmal ap|iaratutt of the rod* and
ounea fur exiiiuple, lo give rum in mme way or »thcr to molecular vibration*
whieh are the Oegiaiiiiig of the nervuua impulwMt in tbe optic nerve. Ho
Mtuifactary explauuiioti of how such a cluinge can be hnrnghl about faai
been oHufvd, and indeed tbe dilficullitv of such a cuuceptioD are very great.
On the other haml, we may more naturally luni to a dicmical explaiuUion.
We are familiur with tbe lact that rays of light are nbic to bring about tbo
doouniptMUou <il' very many clwmiical Bubflttnoc*, and »'e aocoruingly sprak
ot' llune mbslancei as bciog sctwitive to light. All Iho bcU dwolt on in this
book illu«trale the groat complexity and cormtponding ioRtabiliiy of the
coni}io»ilioa of protoplasm. And we might rca>^'<iiiibly »iip|H«« that proto-
plwm itnelf woubl bo e«nsiiive to light: that is lo say, mat rays of light
Miling ou cTcii uudiffvrcnlinted protoplasm might set up a d(>compueiliou of
thai protoplasm, and so inaugurate a molecular diaiurbancc; iu other words,
that light nii^bl act as a direct stimulus to proloplBem. As a matter of lact,
however, such evidence as we at present poaPCua goes to show that native
uuditfereutiated pruiuplaem id u» a rule not teoMtive to li|{bt (iliat_
913
SIOHT.
tbon particular waves which whuti ibcy full oa our reliui giro ri«c in u» Ui
tb« Mneation ut' liiflii), tliuiigh in tbe cnw of some lowly or^imtDS, »Iiqi«
I(r«to[>tuni Bjchibits very liltip (liH'enmtiaiion mh] in narlimlar cnotailu nv
pigment, a jvnsilivoiieai to light hits been obef>rved. >or caa we be mrprind
St this i n (I i (Terence of prut"|il»sni when we rcDect tlisl wliat we may call
pure prutuplasm is remarkable liir its traii»]>arenoy, ifaat is to say. the raji
of light paM tbratigb it with the Blijfliieei passible alnorptioa. But in order ^
thai liKhl may pruduoe obemtoal eDei-u, ic iuii»t be amorbed ; it mun Jj^H
apetit in i}oii>j{ the obemioal work. Acorilincly, the first aitrp tow-ard ll^H
liitmaii'in uI' an Dr^an of Tteiun is the diirereoiiaitou of a |>oniuu of fiiotir
iilaam into a ptgiiitiii at once capable iif ahsi>rbing littht aiid eeaiiure la
ii;ht. r. e,, undt^rguinj; deoouipositiuii upoii ex|Wflur« to light. An arj^anina,
a iwrtinu of whinte pruioplasoi bml thus beoxne diflVrvutiatcl intotucfas
pigment, would be able to reaut toward tight. Tbe light fiillinj; im tU
ui^Dism wuuld be to part absorbed by tbe {Mgrnenl, and th*^ nya thai
aW)rl>tHl would produce a chemical acIJuu anil iM frw chutnicul aiibaUUitn
whirh lii^foiv wt^re iiol pnnciil. Wv have only to suppoiM tliat the cbiMnial
siihiiliinciit are ul' siicli a naiure aa to act lu a *timiilu9 to the prtiioplnaio «l
ullicr jiurti< ot'lbi^ orf^ni»m (and we have manilulii evidvnoc of th<' ex<|iiiHla
»en»iiivi-ii(vi» of prtilnplnaiii in general to cbeniical slimMli), in ontrrtiifo
how my." or li^ht falling on the organiani might excite movouirnta in il,iir
modify moveiuenin which were being enrricil on, or might othorwiao alftrt
the organism in whuli-orin part.
iSuch c>ii»idcralii>n» a« thi; t'uregoing may be applied to even the cumpJei
organ of virion of tbe higher nnimnln. If we suppose that the actual ler-
niinaiiiinB of tlic optic uervt^ arv onrnxindod by siibatances sensitive to li^i,
thto it becomes my to imagine how light falling on iheae seii^itire Mb-
BlMicee should set free chemical bodies pueMased of tbe pru|>erty of aciiiif
aa stimuli to the actual nerve eadinga, and thus give rise to vbunl ln)(mlfn
111 the optic llbrea. ^V^e say " easy lu ima^ii'^," hut we are at prtnt &r
From being al>le In give definite proofs that such an explanation of the
orij^n of visuul impukes is the true one, prolwble and eniiciu|i M it may
apiwnr.
One iif the mi»t striking features in the stnictute of the ratios I* tl^J
abuniiancv of black pigment iu th« retinal, or aa it la aometinMt C^^^H
choroidal, epithelium. It in liilboult In suppose that tho sole tuneiloB fl^^
this pigment i* to absorb iJii? auportliioiiH raya of light, and thai the layi
tbiisahvorbcd are put to no mw but oimnly wantcil. Aim! indeed it baabwn
ihown ibiit the pigmroit is acnsiltve to light ; but the ohangca in it iadtiaid
by light an- excenirely alow. Mort'ovcr, ita pnwncc cannot be of funita-
tnciitiil impDrtanee, since virion is not only pomihle but lairly distinct witk
nibinof. in which this pigment is aliscnl.
Then agflin. in the viwl majority of vcrtehrato animals, tbe outer limbs if
the rxids are sutfuoed with a pnrplish-n'd pigment, the co-called visual purpk,
whidi is so eminently sensitive »> light that ininges of external objects Day
by appropriate means be pholographcti iti it on the retina. When ibe «j*
of a frog or of a rabbit is examined in an ordinary way, with full ex|>0Mn
to tight, the retina appetirs colorlem. lluc if the eye be kept in the dork
for some lime before it is examined, the retinii, if removed rapidly, will W
fuutid to be of a beauiit'ul purplish-red color. Upon ex|vjsurc to light tW
color changes lo yellow and then fades away, leaving, however, the rtfioa
Dot only while, hut mure opaque than it was before. ITjion cxaininadoa
with the micriwcojie it is louud that the purple oolor is oonlined exdoainlf
to the ntds and to the outer limb* of the rotU, the inner limbs beiog whoUj
deroi<l of it.
he color of the rndx u <lu<- tn thi- presoDc* of a (lisliitct tiigmciit. lh«
' tmub) purpk," (liSiiai'd ihnxigli llir «ih«Unc« of the uuirr limhf ; siimI this
may be rxiniclcti rmm (Ih; rud» by dinolvitig thcM in Bii ti')Uoius wkitiun of
bil« hIib. a dnr nurpUt soliition u thus obUiinexl, which m cu|wb!« of
beitiK blenched by ttis nctiuti of light, iin<l tn ils genernl features aixl l>e-
■ liaviur is ttimilnr lo the pigmiitl ae it iintuniily exiele id th« retina.
I Visual purple is fountl, ai wo have eai<l. eseluNvely in the nuier limbs of
Hlie ni(U: it Imih never yet been fouiKl in the eniies, and it is acconliD]{ly
^WMCDt from the retinaa (such aa thott of anakce}, which are comjiooed of
4ooea only, and from the macula lutea and fovea centralis of the retinas of
nan and iIm ape. The intensity of the coloration variee in ititferent animals,
and iW retinaa even of »t>me animaU pooeeasing rocla (bal, dore, hen j aeom
ta be whollv devoid of the visual purple; it is i^nerally well nmrketl in
Rtioaa in wiiiuh the ouler limbs of the rods are viell developed, (la abaeaoe
or pmeiice is not demndetii im uuclurnal habits, since Ihe iutense oulor of
the retina cif the owl is id atrung contmst lo the abaenoe of cqIut id the bat.
It has lii'i-ii found tn lli« reiJoa of the embryo.
The vioiial purple b bleaclied unl ooly by wliite. but alaci by luonocbr't-
mntic. light. Of ibe various prismaUo ran Ine moat iieiive are ihe jfreeniili-
vvllow m)'v, thuae to the blue side t>{ tlitat' onming next, the leuot active
bring thi- red. Now it is preci;elr the green isti-ycllow rays which arc mo«t
readily alwurhed by ibo eol'ir ilaoff. A naiurul colored retina or n nolulion
of TwiiRl iiuqilc eivta n diffune tpeelruiu nilhuui any defined alsorptioii
bandi, aiui acvorUinx tn the amount of culunug inalerial through which the
light pBWM, ahwrpttoo i* aeen either to bo limited to Ihe grveoiah ycllotr
part of the cpectrum or to iprcad thenco toward the blue, and, to a mueh
Ira fitenl, toward the retl. Thtm the variouji priomalic riiy* produce n
ttWo-chemical eDiL'el oo the viaual purjilv ia pro[>orlii)ii as they nre uhajrbed
ly iL L'lider the action of light tbc vihuhI purple, whether in solution or
in iia ualural coiiditinn in the ruds, pH«s» tnrixigh a purplinborauge to a
yellow, and tiiially becomes coloricaa; and we appear to be JiiMilicd in speak-
ing of a "visual yellow" and "visual white" ae products of the plwlo*
ehemit-al chatl(^ UoderBOne by the visual purple.
Fur (be nUutmtion ef the visual purple, atler it haa been destroyed by
light, llie maintensDce of ttie nrculation of the blood through the tueuea ^
tbe eye is not eaieiilial. The choroidal epithelium has by itiielf, provided
that it atill retains its tiaeue life, the power of regenerating the purple. If
a |mrlioii of ihe retina of an excised eye be raised from its epithelial bc«l,
bleached, and then carefully reatored to its natural poeititui, the purple will
.return if tbu eye be kept in the dark. The churuidal epithelium way, iji
Ukii, be Kpoken of aa a " pnrpurogenoua" membrane.
W If the image of iorae bright object, such aaa lamp or a window, be thrown
[mi to the retina, eith«r of an eye lu its natural piMitioa or of one recently
•xcNcd. care having been takeii to keep the retina for some time proviouM
■way i'roiu any raya of light, tbe portion of the retina od which the raya
have (alien will lie found to be bleaohe<l, the reat of tbe retina ttnaiuing
purple. In fiK-t, an " opuigrum " uf external ohjecta may thus be obtained ;
aixl if lh<t n-tina be removed and treatc<l wtlh a four pereenL aolutioa of
tintit^h alum lictore ihe choroidal epithelium haa bad time to obllieraU tba
ilcachiii^' rH'X'iR. ihe retina may n-niain [iern)ui>cntly in tltal cotiditiuii; Ibe
jihiito^clieniicai eilcrt may, a« the phiito^:rMpIivr* aay, be " fixed."
It aetmed very tcni[niug. vapiwially upon the first diseovery o( it, to sup-
poae that tbil viaual purple is directly confirmed in vi»ion. If ne aupptiw
that visual purple iL»clf i» inert towurd thv ending* of the ofttiv iMrvc, but
,liiat eilber vuual yellow or vintul whik% i. c, »oiiur product of ibe action o(
M
'J
914
SIGHT.
light on visual purpItN may net as • ■timulua to tboee endingg, tfae wkj Mtmi
Of)flwd Ift uwleniiaiidin}; liou- rays of li(>lit can i;ire riae lo eeoaorr ini|Mt«
in Ui« npiie nerve. UnJbrtunatel.v visual jMirii'o is nlioeiit trota tlie <»■«
nixl rmm (lie Tuvea ceniraliit. whU-b, as we ahall see. l> the r«Ki"i <*f diiUaa
it i# lurilier eutirely wautitif; in aonw aDiiuale wliioii umloubtedlt
VBion :
see vcr)- well ; nod, lufttly , auinmU, well u frags, iiaiurallj poanasuig Ik
pigment, coiiLiniie to nee rerv well, and even ajtparenlljr to see cnlors, trb»
tbcir vixiiiil purjile liaii htxii ulMi)Iiitel}' blcaehttil, »» it mav be by prohaied
rspiHurL- of tlie vyti to Hlmug liu;hl. W« wniiot thereiure, ut preeenl il
kn*t, i-xfilniii the origia of vlsiinl Tcapulsea bjr the help uf rkual piur|de. ii
tlio winie time tiH history BU{[f|:e«ta that some sabttancea, sensitive like it to
light, htil tiulikf it, coWlnai utid tltervdciTe esea^Hiig oWrvalion, may nil,
and liy phultt-chomical chatij^ be ibc mean* of exciting tttv optic ornw.
And. u« we vJinll tee later on, one iJicory of color vinkui is bawd oa lis
nwiiniptimi that viiton is carried on in aonie way or otiter by changes ia
wbut Riay be called the visual wiibttancn preaent in the n-tiiia, tbtae h^
sUin<:t-9 bi-iiig uM-il lip mill rvguneratnl tw vision ia going oii.
But even nitmiitiiig an prooable tlic exi»lenec of (hew sensitive visual mI»-
stances, the ohnrigtv iii whicli lead tu stimulation of th<' teni endings of tlM
TCtimd Dcrvoii* Rieehaiiium, ne CAniiot nt pNaent state anytbiag d«(iBtt« eea-
cerniiig thoH- ncrvc-rndings or the maimer of ihoir stimuluttuti. It may U
that even the outer limlw of the rods and cones, iu spite of the appaisM
break of conliriuiiy lictitecn the outer ami inner limlw. are really ntmm
in nnlurc. It mny he, on the other band, thai the outer limbs are dllMT
purely dioplHc in tiinction, or are associated with the seusitive Ttsual M^
stances in such n vrsy thiit the purely nervous structures mtnl be cnnsidatal
as extending no further ut least than the inner limbs. We oiunutasyM
make any deKuite statement in the one direction or tlie other.
In i»nnection with the origin of visual impulses, we may |ierhafa call
attention to the remarkable changes which the eella of the retinal pignmH
cmthelium undergo under the intiuence uf light. Wlien an eye ha* bsa
siiut (fir from nil jiglit for tume little time tlie piginetii is oou'cvotnitsil is
the budiei) of the ceils, and the remarkable lilamcntou^ proonsm of tits artk
with llie pigment granules or ervdtaU which tbey carry, eihaiid a slight da-
tBn<« onfy between tbe limbs of the roils and c»nes (about ooe-lhird ihni
the length of Uie outer limbn of tbe nnla). Under the influence uf 1^
theae proccMca, loaileil with |)i;cmeHt, thrust them.ielvn a much lunger as;
down tiiwani tbe external limiting mvnihnine-, in ci>i>s«(|uenoe n i iniijilitalti
(luantity of pigment is found niiiweil between the outer mad even ilie iaMr
limbs of tbe rods and coiiea ; iiul(«d, tho initer limbs of the ruds fweliing si
the same lime become jammed, an it were, In-twocn the niiuiea of figm&i,
cuiuing the epitbelial layer to adhere very cloaely to tho layer of rmli sad
cone*.
Thi- retina and optic nerve, like other iiervoits struetunv. develop elccliie
currenlH, whieh may he sjioken of us currents of re«t and currents of acOM.
Tlwy may be shown by j>lneiiig one electrode on the retina of a bisecteij syi;
or on the cornea of a wholo one, and the other on tbe optic nerve, or b af
part of the eyeball, or even nn some dixtant part of the ImkIj. Tbcrsit
also maiiifestnl by the iM>lat«<l retina ilsolf. The pbenomena appear soar
what complicated by the appenrnnce now of positive, now ol ncgainr,
variations ; but this fact eomi-s out clearly that the inrideocc of ligh. m
the irritable retina develops an electric change, the ma)^nittide of which titi
a certain extent prn]iorti<>nntc to tho intensity of the light acting aa a 4i»-
ulus- The changes acconiingly diniint«Ji and cease to appear as th« rrluu
iufllly loMS Iti irritability after >leiiih. We may iidil tliat th<M tJeoirie
iKifDuna apjienr tu Im ijuil*: loilopciuleni of the uuitilitiuu arihsVimal
purpk.
k .SimpU SentaUoM.
RtlttloH* of thr Mfiuation. to the tiimiihu. If vt put ii»i<l* for ihfl nrcmiit
kit <|u«Uun)> o( rulor, we may my that Hght., vi«wi:)l «' a *(iniu)u) nltcctiiig
Um retina, varin in iolcn^ilv, that if, iti iliu rni-niv vf the Itiniiaoiix vibrn-
vlinns u maoifmn) by Ihdr ampliliiHv, ami in iliiration, that it, in titi! ivnelh
BbT tiiiiv a diirccfeion of wares conliiiim In full u]mhi tlir retina. Tho ofleot
nf ibv light will also (toneiid oti tho t-xlcnt of retinal mrfiicc cxpomd to the
taminmM vibrations lU tne mme lime, Taking a tiioiiiiiiiis point, in order to
«UtBinat« iht buier circiiniRl.inco. we mny makr; the TDllowing •tatemonls :
The aRMUton has a duration miicli irrrnli^r tlinn that of ihu stimulus, and
in this rMpwt is comparable to n nmsciilur contraction c»iise<l by such a
stimulus ns a sio^de induction slim-k. Tlic eontailion iif a flash of nght. for
iaatanc«, laaU for a much longer time Ihatt that during whiL-h luminous vibra-
tioits are &lling on the retina. Ht-'ncc. when Iwo rtimiili. such as two flashes
of light, f^dlitw each other at a sufficiently short interval, the two sensations
are lii«ed into iine : and a luminous point moving rapidly round in a circle
l^ivea rise to the sensation of a continuous circle of light. This again is
quite comparable to muscular leuiniis. The interval at which fusion takes
place, that is, the interval l)etween successive stimuli which must be exceeded
ID order that succcasive distint-l sensations msy be produced, variee acconlinj^
to the intensity of the light, being shorter with the stronger light; with a
faiui light it ia about ^ second, with a strong light ^g or ifg second. This
nuiy be shown tty rotating rapidly before the eye a <li» urruDged villi alter'
nate blatik and white aector* of ei|inil width. With a faint ilfu mi nation the
flickering, indintive of the succeioive M^usatioim frum thi? white Mctors not
bring eompI«t«ly fuM.'d, etmist when the rotation beonnw >o rapid lluil each
pair of black and white iieclon> takes only ^ aeemid in pawing before the
bcye When a brighter illumination i« useil the rapidity munt be increiuwd
Pbrlort the flickering rlisa[i[H^m. That jwrt of the H.-iii(utioo which is rw:gg-
niM^I an lasiinu: aAer the ccwaticm uf thenlimulus is frei^ucntly s[)ok«n of na
llin " atU-r-imiigi'."
Though th(! duration of the afterimage is loiigt-r with the sirongtf light
(that caused by looking even niomentarily at the sun lasting for some lima).
lite cxnimenceiDCiit of the di-dine of ibn s-^nsntion hcginn relativdy Mrlitr,
hence th« greater dilficullr in the c^Niplole fusinn of siicecasive atnaations
with the strongrr light. The interval at which fuoiim ink*" plaew ditfers
with dilTentit colors, being shortett with yollow, iulenudiate with red, and
loomt with bliK.
'Die duration of a stimulus necessary to call Jortb a eenntlon U exc«od-
ingly short: thus iheshortcat ponible flash, such ns that of unolcelncs|»srk.
gives RM to a sensation of light.
Objacta to motion when illuminated by a single electric spark appear
motionless, the stimulus of the light n>fl«cied from ihem ceasing before they
oan make an appreciable change m their jHwitioo, When a moving body is
illtimiDaied by several rapid tfnshes in sucoessioD, sercral distinct ituagea
oorrespondisg to the positions of the body during the several flashes are
geocrated : the images of the body correapoodiog to the several flashes fall
on different pario ot the retina.
TUe inteiuity of tlie aauation varies with the luminous intensity of the
a nux candle appeara bri]{hler ibau a ruAhltgfai, Tli« ratio, h»w-
»l<t
BIGHT.
ov«r. of tlie fTDMiion tn the (limtilua u Dot a timjAt one. IT tlie luiiiiu<«itr
of uu ubjrot bo graduHlly increaMd Truia n retf fteble unge to a T«ry brishl
one. it will 1>c fniiml ihnt thoiigti ihecurn^ponding Komliona IlkeniMgrnu-
nlly inori'dK-, thr iiicn-meiits of Itic Mtuation^ due to increnMrnU of lli»
liiiiiitiiLi«.it}' grndunllv ilimiiiieli : and ut Initl nn ioereiiM.- of the luraiBontjr
prndiicrs till npiiirniilile incivxEc nf M-iwntioD ; n li^ht. wbni it rcMlia ■
ccrtiiin lirighlnnK. ii|i[)('Krs m liright tlint nu mnmil trll vrh^-u H bteoaii
■»r l>rightri. Hviice it is niucli c-imivr to iltHlingiiixh n xli^ht dilfvmMlof
brfghlnw« bct*wn two feeble lighlji tliiui ihu Hinic liitrcrmoc tH-1«Ma t«u
bright lighls; wc mil ciwily tell tbc diHi-wiicc iM-twet-ii a mvblJghl and ■
WHXrniiilb': l>tit two f^unn, or <-vcn Imt bnglit liiiii|», mir of nbicfi dilrml
from the orhcr nirn-lv l>y juKl tbe iiiinilirr i>f luniinoii* nty* whkb n nt
(-BiidlFi-niili in iiditiiion to thorn- uriii furib by a ni*bligbt. would aiipMrb)
Uf to luivc oxnclly llio rame brighinoK. In a diirktnnl r<H>in nn olijcd
plnwd Iwfon^ H cHiiillu will throw what wr considiT a dpv|i slindaw ao ■
sheet of pnpt^r or any wbilc Miriam. If, bi>w«ver. tunlight bu aJknnd lii
fall (in tlic pnperat tlic vnnio timo fmm the oppmilv fidv the nhoilnw » du
longer visible. The ditfercnoe between the total light n-fU-cled fmni tktl
part of tlie |>ii]>er where the «hndi>wwni>, and whieli i» illiinitiiAli.tl byibrtui
iiloiie, nml ibiil rcHocleii frnm the nvt of the [injier which if illuminntal lit
the cfliidle nt well ne by the sun, renitiins tbc aame; yet we rail Do UiDgn
apprecinio (hnl difTcrGiice.
Un the other hnnd. if tistd^ two ruehliglits we throw two sbodowa on a
while surface and iDiive oue nmbligbt nway until the shadow uitiatd b^ k
ceRses to bo vigible, and, hftTinc noteil the disUiuee to nbieh it bad in bt
moved, repeal the same experiiiieut with two wax caudte». we shall fiai
that the wtix candle has to be moved juat aa far as the rusbliglil. Iti fol
it id (bund by careful ubi««rvaiioii that, nilhin tolerably wtilc limits, lb*
■mallcat difl^rence of ltjj;lit which we can atiiirbciate by visual a^UMtion* isi
ronstant fraction (about ito'^) of the total lunuDutiiyeniplnyed. TI>e«aM
law linldagoud with regard to the other Koaca aawcll. Thennialli'st diflirmo
iu length w« can deivct between two linea, one an ineh lon>; and the otW •
little Inw thauan inch, iit the >ame fi action of au Inch, thai thr »lIlalIrftdiir^
L-nce in length we lan dcieet Iwtwecu a liui; a fool long nn<l one a lUlJa \m
than a foot, ia of a fnut. Put in a more gcn«rnl form, then, i)m law. wbieb
b ofti^i called ^Veller'^ law,' in aa follows: When a aiiniulua n coaUauaUf
incrcafeil, the increase of tttiiuubin neeiioary to eflll forth the aia>llMt m»
rivhic increase of aeiisatiun nlwuyH hvan th« mux proportiuu la tb« aUs
Miniulu».
Itiffinrtion and Jyimtni of Kntaiimt*. When tight falls on a large portte
oF (he retina the total teimtiou prcMluced » grisirr iu iim-tuut ihaa wbw*
nnnll portion only of ih« retina is aflccted ; a large pi«<-i- nl whita |a}ai
pnxliat-o » gn-aler (ulal etfect on our conntioiisness than a siimll one, Uwup,
If the mrlaces be uniforndy and equally illiiminaieil, the inUnrity o( the
sensation is in eneh case the same; the small piei-c id juaper B]>i>ean ai
bright or nii " while " a« the laiga ono. If the images uf two luminous ub-
jeets Itill on the rclina at sufficient diuances apart, (bu const<)uent aenwlim
are distinct, and ihe iuiiiuity of each sensntion will dcjwnd solely u|iaa tbc
luniiniiily of the conx-sjiondin); object. If, however, the two nbjects an
made lo approach each other, a point will be reached at which the two ica-
sntioDB are fused into one. When this occui? ibe iuluiiuiy uf the total
fensation produced will be greatber tbsn that of either of'lhc M-nMthiiu
> riMD wblidi Fcrliaa, bjanuaummloB. ol>l«liuaaiM'linwltHit*>i'>nwloa«(-lbtaiul>.«U(fe
Is MmtUnua iHMmellr tio^" "' ■* PMhiivri law.
bj the einj^le olijecU. A number of luminous p linU seatu-rcij ovi-r
I iurfiiw irould appear each to hnve n oeriain Imj^hlQcas ; euch would
give me to a e^iualioo of a certain inlemilv. IT tbey irere all gathered
into one spot, that ijiot would appear far briifhter than anv of the previntis
pointa ; the intcnaitr of the sensatiun would bo Rreater. We may theroforo
wppoaa the retiM to bo divided into nr^aa cirreapoudiog to aensatioiiil
unita. If the iiiMj(«a tkwn two luRtinnus ohjecU fall on separate tiiuaI areas,
if we may *» mil them, two distinct aen-talinnA will be produoed; if. on the
cmtrary. thev both fall nit the aarae visual area, mie aensatioa only will be
produced. Where the senuttions are sepAtate, the intensity of the one
(with est-eptions hemtler to he raentioiiei)) ia nut aHiK^ted by the preaeoM of
t)>e other ; hut whcr« they becnroe fuied the ititeuitty r>r the united senaa-
tiooa U KreatiT than either of, thoui;h not e()iial to the «uin of, the aJQ^Ie
■entaiion*. The exiatence of theae >en«aiii>aal uiiii« t* the basis of distinct
Tbiuu. When wc ^enk of the sinallett iiiee visible or ilL>tin;.'Liidliable. we
are referring to thi* <linvusio[i< of tin- retinal arviu iMrrai|>oiiiliiu[ to theae
aenaatiiuial unita. The retinal arva mu«t be carrfiilly dialloguiafaed ftofa the
■anaatkMUil unit, for the NenxatiDu ic. tv we have .-ven. a praceaa whoM arena
atrMdica fri>m the retina to ciTtain purls nf the bmin, and the eireunojorip-
tion of the *eiimtiimNl unit, though it muM b«^in aa a retinal area, must aUi>
be cntitinuei) m a <vrehnil aiva in the brain, tlic latter oorratiMndiux to,
and U'ing an it were the proj«i(:ti»n of, the former. With matt people two
atani apix^r as a Kingin »tar wbeii tlw distanoo between tbein aubteadi an
auRlv of len than 6i> Kcondn ; and the best ey«a g«n«rally lail to diaiinguish
iwo (tarallel white sUeakc whttn the distance mjtwoen tbe two, measured
from the middle of each, 8uhicn<li> an angle of less than 7S aeoonds. Some,
howeveT, i-nn >li#tinffuish obioits •'iO sccitn'Is distant from ea^h other. An
■Di;le of 73 focoiids in aq object i-urreapond^ iu the diagram mat ic eye (see
|>t H94 1 to the lenjctb of 5J^ /• in the retiual image', and one of AO seooiKls
to 3.ti.i ...
In the human eye 50 ooum may be counted alonj; a line nt'iQOii in length
drawn thruu(;h the centre i)f the yellow spot ; this would give 4^ fur tlii^
disianoe between the cenirea of two adjoining cones in tbe yellow spot, the
•venjce diameter ufa cuoo at its widest part beinir •If and there bein;; ilight
interva'a between ueiithboring coaea. Hence if we take the centre of a
eoae a» the centre of an anatomical retinal area, these aoatoTiiieal areas oir-
respond very fairlv to the phr^ioli'i^ical visual areas aa determined ahove.
That is to tav. if two pi>int« of the retinal iuia^e are leas than 4^ apart,
ibey iiiav bi>ih lie within the area of a single cone ; and it b Jnst when they
uv leM than about 4 >< apart that they oeiise lo cive rtae to two distinct aen-
•Uiam. It must he remembered, however, that the fusion or distinction of
I the wnaationa ts ultimately determined hy the brain and not by the retina.
[Twk p»inis Iff the retinal image Idu than 4» apart might lie botli within
'^the area t>( a sln;;le eune; but tbe reaaon why, under such circuiustauccH,
they give rise m one senMllon »idy is nut because one cone fibre only ■■
•timulaied. Twu poinla of a rrtiual iniavi' mi^ht He, one on the amn of ouo
cone and another «n the area of an a.lJDiniiiu time, and still be lot? that
4>i npart; iu sui-h a ra«! Ino conn-fihrct wouhl be stimulated, and yet only
OMatHMlion Willi Id \i- ]iniduc«d. 8o also in the lea wcntilire |>eripheral
psrta of the r<<iinii («•> p.iints of the rMtiiial imtfO mi^lit stimulate Lwn
CMiea a ooii«i<leral>]u diMancv apnrt. and yet give rini to one senotion only.
In the CMS where IIm two pointJi lie entirely within tbe area of a single
f, it b Axeeodingly probable that, oven if the adjacent cones or cone-
> Br ^ !■ nwsDt oi.»<tioia«a4tk or ■ uiUliUMtn.
918
SIGHT.
flbrce in the relinn nri* not il thv Mine lioic MttinnlnKvl. iniTiuliM radiilf
frnm llie ceretirni riKlini; of the exnt«l cono inlo l)i« iivigliboring cpnrlinl
enctinKB of tlic neighlwriiig conts; in othrr words, the Hnsniion-nrpa iu iKc
bntio doee out exactly corrarpond to and is ni>t fJinrjily dctintti \ikt lit
reiinal area, but gradiintly mdee away into n«i};hlit>ritif; eeDMitton-ami
We niay imagine Ivo poiiil« of the retinal image to tkr apart ihnt rrtti tir
extreme niargioH of Ineir respective cerfbnil eeniinlion-an'a6 do not imicfc
each other in il>e least ; in Hucti a cane ihere cnn b« no duubl about ibe t««
{>oin[a giving riiw to two sensations. We niiplit. Iiowever. inia;;ine a Mcu>i
CBie wh«re two [loinU were jual so fnr a|iart lliat tbeir respective waulii*-
arena ehouM coalesce at ibeir maTginb, and yM thai, in ptBaiaK fnioitW
centre of oue senBniioii-area to the centre of the other, ire should find im
cxainiitalioii a cxjnHiderable full of een^lioii al the jnoction of (be too
iireaH : ami in a tliird ciute we might imagine the two centrei to be » thw
to euch other that iu paaiiii^ from one to the other no appreciable ritabii-
liou of ieuaatioD could be discovered. In the la^t osae lliere wouM be \m
one Msnsation. in the aecond there might uill be two Moaatioju if ibe inar-
final fall were great euou^h, even though tbe areas inrtiallj ooalwaA
liiiK, though the nnwiic of rods and oooea ia the baais of dbtinct tiaion, the
di*tincti(iii or Albion <if two viaion Impultea b ultimately determiued bv tbt
diMpu»itiiMi and (.-oudition of the eerenral ociitmi. Hence tliv panabiliiy of
incrcaaing by exerci«e the faculty of dixtin'p^whing iw>i M-naaiiwiu, itDee bf
uw the cerebral teiiaation-areua teeoine more and okiK' diflen-ntiaird. Tbih
liowcvcr, ia evcii more atrikiiigly HhowR in touch thnn in aight.
Color SntMiionn.
When we atlow aanHght reflected from a cloud or sheet of l>nper to bll
Into the ere, ite have a sentaiion which we call a wnsation of white li^rlii
When we look at the »iuie light through a prism, and allow difftrmt [mru
of the spectrum to fall in Duccemion into the ere, we have ^DsatioDS iiliidi
we cmU noipectively sensations of red. orange, yellow, ^reeo, blue, violet, ftc
ligtit. In other words, rayu of light falling on the retina fcive risa to diStt-
ent MCDsntiono, according to ihe wave'lengihs of llie rays. Though we tpnt
of the »i>ectriini as cuiiniatiug of a tew colors, kuch as re<l, urangv, tie.
there are an nlinoxt iiiliiiile iiumher of internieiliBte tiiilt tn the ^WCtrui
iLt«ir; and wv pcrcHrc in external nature a largo number of eolot«. mrhai
purplr. brown, gniy, etc.. whi<'h do iHit corrrifioiid to any of tbo color tn-
tations giiiiXKl by rcganling the succi^wive pari* of the N[icciruni, We lU,
bowerer. «n fxaininatiou, that certain diatincl coW seoMtioiis, not con»
Spooding to any of thi^ ci)l"r» of ihe spectrum, may be obt«iiM>d by the
fbtrion of the nrnoalion.* cnuM-d hy two or more of lb* prisnmtio colon.
Thne pnrplc. which i" nut pm>ent in the 8|iectnim, may be at once prodaoal
by furirig the «ciituiliiini< of blue and red in proper pnip>'riioiM. Monvnr.
many of the vnriouf tint* and >hadcii of nature may Ih- imitntiil by Aaiitg a
prtii-ular color scntuition with the scniuiiioii of while, or by allowing a etr-
tain ((uanlity of light of a particular color to fall Rparaely over (he arra at
ihe retina, which is nl ihe tame liinc protected from the aceecw of anr nthrr
light, i.e., aa we ray, by mixing Ihe color with black. TIiuh tbe hmaM
of nature reeult from various admixtures of yellow, red. white and blaH>;
and a small quantity of white light, K^iieied over a large area of il>e retiaa,
t. e., white largely mixed with black. fDriiia a gray. In fact, the ijualitin nf
a color depend (1) on the nature of the prbraatic color or colon. «. t-,cn the
wavfrlengtlks of the onsiitueni rays, falling on a ^ren area of tbe rvtiua :
VISUAL 8SNSATI0NS.
»I0
I
I
(2) oil i)ie nmount of LliU eoloreil liglil witidi fmllt mi Ihe nrvn uf ibi^ rfliiiK
in a |[iwn linx! ; iukI (3) on the tniouot nf whilv li;;lit fkllin){ mi thr iiitno
arc* nl the woe UnM). Whru ray* corr>Mi|)iiMiliii!{ to a prUiniitic i^oltir fall
upon llip rclina uDiLCO(»m|>aiii«l hv imy wliili- light, the color !< «iii(l tii bv
" Hiliinilnl : " oikI n color in k|hik<'ii (if tu uivttv. or \vtf niiliirnUil nccordini;
aa it i» itiixiil with lnw or mori' white li^'liL WbL-ii wv air liil Id ihwcribc a
culur (IN iH-ine "I* s"ch a lint or hue, w an ga\i\vd hv ihu lirvl of (ho abova
cx>ui)itioi)». Bill Vic hiivr no common jihrmHs br wiiich «ir ilistineuuh the
aecootl of thv above ronditionii fmtu (he thin). Thv wort) *' pnlu," il ii tnie,
i* nixt frraucatly utcd to expmM n color \-orjr slightly ntiirnti.-J : biK iho
ironb"ricli" or "(t«c|>" are iisod fomclinii-s aa nwoniae hichlj: Milnrated,
■ooiCtimM u ni*«ninp simply that a large (lunniily of light ot the particular
hua il paMing into the eye. .So also witn (lie phm«« " bright' ; liiis we
oft«o iiH nbeo a largp amount of colored nntl whilo light fall nl (he samo
tima on the fame retinal nren, but we eoioeliinc)' alw uw it to vx]>r(«e the
narv iiiteoMty of the senaation.
The ))«■( mrlboil of ru.ilii); color aeaMtions b that adopted br MaKiell, of
alloniniE two iliirereiit purL* of the Bpuutruui to full on the amui' pui't ur ihc rrtiiia
at ill* Ntmi! timt. The \u* of th« piir<r |iri4mai)c colon ellmfniilM riror* nliiob
arue wImh pijnncnbt, the colorn of which nrt not piife. but mixnl, nfi- cmploywi.
And wlwre t>iKm«(iI> nic ut«l, it in the ■cnMliont to which the pi|;racn(ji kiv ri*a
whii'h must be miinl and ii'jt th« [>i|;uieiili theinielvtii. Tbiu wliite llit- senna-
tioiM ^in«d by looking at KAinboee vellow and indigo respFctively when fiiMd
gfT« riM lo a Hnsation of white, gamboge and indigo thcmwlvn when mixed
appear gmn. The color at tlio mixed pigment in clue to tlie fuel that tlie rays
nhii-h reach the eye from the mixture are thoM which are leail atiwrlied liy the
tiro pigment*. The gamboge absorhs ibe blue rnyit very largely, but ilie green to
■ much lemt extent; while the indieo ah»orh« the trd and yellow rii<» very
largely, bill alt" aVuoriw very liiile of the green. Menco green is the predmninant
huaofthe mixture. When pure piKmenla. ■'. e., pigment* correajtonding iw cli«ely
HiNMdble to the priHmHtic colon, are ujted, utUfaelory teaulta may be HEaiiieti,
dtber by lulng th« reOecled Image of one pigment, aud ariiueiiig bo that it falh
on tho retina al the same spot u the direct imaici' of the otlxT pigment, or by
allowing ih' image of one pigment to fall on thr retina before tbe M'aiation pro*
dace<l tiy the other hai paiM away. The fir*i mult ii ejwily reached by Helm-
bollx'a MOiple metliod of pincliig Iwo niece« of colored paper a little dlntanoe
apaitoa a table, one on ench «ideof aglaxipliiie inclined at an angle. By looking
mtb line eye doWn on the glwu plate the rcBocted inijue of the one pH|>er may
be oiaile to coincide witli ihc direct Imago of tbe other, tJie angle which Ibr Klnm
pinie maketi with tbe Utile being adJnatM lo the diatance' between the piecee of
paper. In Ihe hccoimI method, tlia "color (op" is used: Mclor* of ihe colon lo
be inteotiialed are placed on a disc made tt rotate rery rapidly, and tbe image
of one coTur ia (bus broagbt to bear on (be retina ao toon alter the imsge of
another, that the two itenuiiona are f^ued into one.
When the Bonsstijms corrcajKmdtng to Ihe several prismatic cnlora are
fuwd tog«her iu various oocab mat ions, tlie follovriDg remarkable results are
brought about :
1. When red and yellow in certain pro|>orliom are mixed together the
result is a iteniwlioti of oran}^. luitc iDdiBtiiigutshnble from (he onin;:;e of the
apcclrum ilaelf. Now the latter is produced by rays of certain wnTe-lcnjrtha,
whereas llie rays of red and of yelluw are reajiectively of rpiitc different wnro-
l«ngtbs. Tbe orange of Ike tpeetrtim cannot be made up by any mixture of
(lie rerl and tht i/dlote of the ^>telrvin in (he seuse that the red ami yellow mjs
can unile together lo form rays of the same wavo-lenj;llia as (he orange rays ;
tii« (hrte things are abwlutely diRerciit. It ii iimply the mixed tentalitut of
the red and yellow whioh bao like tite Mrtttniion of oraoge; (be inixUiiti b
SIOHT.
entirelr unl obMlutely n plijViologioal ona. Id ibe ma» wbj we lUf hf
Appn>))riut« mixMma produce the Kmstto(w(!ormpnD<tIng to other parUof
th«#|V'Ctruni. Now we must suppose that ra^ of ililTcrviit wnvi;- knjctli
Sfive Tim III iliirimiil fenwiry injpul*ee; ihat. fur uManw, the u-amry iai]H(lM
KCuemtis) hy oningn ravH nrn ilifliireut rrum thoM gen«ratc<J by reil ml
hv yvlluw ny*. Hence we nre luH by the fiict or mixoil Kiiinli'MU bdif
uM-ntii'iil nil.li other iipi>areiilly itimjik sritKntionii to infer ihut the Momj
impuiw* which iiny my nrigiimle* tire cither theniMlvc* ul* b crMiiplcs dm-
Mt«r, or )M hitciiniiiig mnvvrlLfJ into tciMntioD* give riM (a complex or miitl
tci)nti<>n»; ih^t, Cor iiiKtoiicc, ihc inipiilM or acnution wbiofa ■ ray in ik«
niitMlv of the ornnge gives ri»o tn. is not » nmpio impulse or Miitttjin
iiii«wcniigttxclasi\'el^ to thecolorof thitt rnr, but that Ibe my ptx» rw
«itli<T to 11 complex impulM which bceomet ciinvortoH into a cniapkx mm-
tioii. or lo n iimplp iaipiilsc which evenMinilr <)ev«lo]is into n mixnl .4
complex tH-iiention, into ine cumpwitiun of which In each cue other itnop
liniF nml shs'kF of ntl niiij yellow enter.
'Z. Wlicn certain color? are mixed locether in pain in oortAin definite |tn>-
portionm, the ratult is while. Then oolore are:
KH (near a),' nml Bluc(5rcrn {wja F) ;
Ornnge (nenr Ck end Biue (ln-iitccn F and O);
V«11ow <iiriir D), HQi) Iiidi^'O'Blue (near 0);
Oreen'Yellow (near E), and Violet (between G and 11),
and are ntd to he " wmplcmentarv" to each olhrr. To tboee ml^l b»
add«d the peculior uun-pniiriiiitic coVir purple, which with gn!«n abo give*
while.
3. If we select arbitmrily anv ihrve colom corresponding to nay tliTM
parts of lira fpcclrum euflicicntly far apart— ear, for iustanco, red. gm%,
naU blue— we enn, by n proper adjustment of the pjoportioDs of each, |ir^
diice while. Further, iheee three coinn e&n be taken in 6uch proportioM m
with a proper addition, if necessary, of white to produce the eenaaiinns uf all
other ci>lur«.' That is to say, given three itanditrd sonsatwoB, »ll the other
MiiMlioOH may he ^-aiDeil by the proper mixture of these.
It i." obvious JWiih the fore|toing that our real color eeosUioiu are miKk
fewer in number than ihuee which we appear to huve wlien we look on ihf
cotoni of tho ^jicctrum or of nature ; that rays of li|;iit awake in aa certaia
simple w-nnation*. which n)ixe<) in variotis proportions repnxluoe all our •«•
salions. Ami the oueaiion ari»e!>. What in the nature or what are ibe char
acler« <>f thn«' tiimpk- ttcoHiitiriii!! ?
When w(< i-xumine our own neuititlioiis of light \re find that certain of tliMe
seem to be (piile diolinnl in nature from i-jit'h other, so that each I* aonotbiagi
eui genrrit, wlicrens wt^ i-iuiily fiTngniie all other mdmiJoiui as various nix-
tures of thciie. Thu.i rcl and yoHnw are to U!t iguite dUlinci; wa ila !>•*
recn{;nize nnvlhing coninion to the two; hut orange u ohvlouvlv n niixun
of red and yellow. The scnHttions tniutieit by dilTurcnl kimU of light whiA
(huB api>enr to 11s distinct, ami which we may oiieak nf an " fuuilaint'ntal mti'
salions. ' are white, bWk, nx), yellow, green, bine. Kucii of thew mwmi 10
us to have nothing in common with n»y of the otlten, wherens in all olhsr
colon we cnn recogniu! a mixture of two or more of these.
Thin re«ult of common cxpericiicofiJggc«is the idea tliut thiiwe fuDdamratal
Hcnoalion* are the primary or simple ecnaalions, epokon of above a« tho«e ovi
of which all other tencaiions miiy be supposed to be compounded. And a
I "not Idtcn refer to Fimimnlx.fi-c* lliin.,
■ A Oiw hlfkir HiuniifO Fi.lon nnixii ri> v> rvfrnliircil. Mil ■ nililoiriir niir no* of IfesaaSk
whlNoui, Ws fDH) . ivrli«». ihrnrlbiv 'fiett M tlioc hiumM-I rrii-n « latlyc "nwntiwial If*
iniiptiuiilllilnsllfn nrtlm TTiTT*«rMlrirlly mlntint nilnrr wUh Ibc loMnirMnB ur«ait*
921
»
tli«>urT hits h(>en propoM"! (o nwMirilo lbi> vBrioii» ftcK of color tMod, with
i)iv »iippne)(ti>ii ihnt we pnmnw lhrv» fix fiiii'liim^iiMl KantloM. This
llicorv. ktioMii lu that of lleriiiif. in fumicwhnt m follnvri : Thr *ix M-fi«ili»n»
readily fall into three nairv. th« members of esch pnir havini; ana)<ii^ni* reU-
tioBs to each otiier. White mid btaek nalnrnltr pi tojrolhcr, the oni; lining
the nnlaf^iiifllic or correlative of the other. There iit n niniiinr c>nni!ic'tioii
hetireeii reil and )treen, the one beini; the coni|>lemenliiry of the oth'T. nnd
bptweeu yellow and hlue, which are aiinitarlv com piemen tarv. Wo mw
rcaaon. a tlioii time bark (p. SN), for believing; that vision orieinulM in
the cluinjiei lakiiii; place in certain Tuiial sulMtanoea for a naunl ■ohfllnnoo)
in the r«lina. Aud the (hcorv of which we are speakini; mippoicfl tbiit lh«rs
exist in the retiua, or at leiwt Momewhere in the visual apparatus, three dtB-
tinct TiMial (iibiilnneet which iiri! coiitinually uudericoin? a (luiible luiJlab-
olbm, one cf>iwlfu<niri>. of muiinilation or buildiuK up, and the olh«r
dMnictive. of diiwinitlnlii'ii or hrcnkiuv <lown. One nt thew auhalADcei ii
funhi-r of Hiich a nature ihnt whrii di.viinilution in in exocM of BSMailBUao
we have ■ t«-iiMif ion of whit<>. and whrn awniiluljon la In exoSM B Mnsfttion
of black. With ■ itcciiml milNttunn^ excMi of diMtmilnlJoii pmvnkea red, of
aminilaiioit grr^n ; and uiih tho ihiril oiibstance. yclt'iw and blue rMpect-
ively. Wbnt in th«! lallvr two ntjliatancaw dtMiniilatton and a:«imiliLtiikn are
exactly equal, no rlfcct i# |ir>Mluccl : hut ivith thr. fir^t Milwlamre thin oin-
dilinii pnxluna in ua tli* aflbct of gmr. Furtht;r. thc«c HulistAiicc* are of
Mich a kind thai while the first or whitr-blm-k *ul»taniT in influenced by
ny ahing the whok rnngc of the siwctniro. the two other wilinlnijcen are
difliTttntly influintnt by mv* of different wave-length. Thw* in the jwrt of
tlw aptdroni which wo call red. the rays promote n rapid di<t>!mihii'>n of
tin Rd-green «iib«t«i|oo with com pa rati rely vlieht eflecl in either dirMlion
on the Tcllon-hhie aulistance; hence our aeDsalion of reel. In that part of
the apc^niiu which we call yell.iw the rays effeot a marked dis*inii]ation of
lh« yellow-blue titbaisnre, but their action on the re(l-Kre*n substance if
equal in tite directi»n of both awimilati<>n and dMsimflation : hence our sen-
wtiun of yelUiw. The itrcen mn. axaiu, promote aasimilalion of the red-
green Mibsiance. leaving the ajatrntlation of the yellnw-blue nuWanee eiiual
to the diMimiliiiiou. and aiinilarly blue ray* cwnae aniinilation of the yclhyw-
blue sulMlancc. and leave llic rvd-irn-cn nnbitlnncc neiitnl. Finallr at tlie
extreme blue end of the uporlrum. the rayn once more provoke dtK<imilatii>n
of ilie ml-graen suhKlanire. When omngi- rarx fall on the ritinii. there
ia an exccM of disiiinihition of l>nth the red-grren and the yellow-bltie
•ufasUnre; when greenish blue ray» are ncrreiwd tborv is an exec** of
Maimilaliun of Ir^h thftic tub<tanees: nnd other ini«rmediale tintn com-'
apond to variable amounta of disiimilation or animilntion of two or moi« of
tltCM Hubttniicc*.
When all the rav* together fjll on the rctinn. the red-green and yellow*
Idut vulMtance remain in equilibrium, but the white-black auhitium Ii
v(oI«ntlr di«>imilnie{l ; and we say the liiiht i« white-
Anotltcr ibcorT ( known a« the Young-Ilelmholti theory, beeaiiw it WU
intnxluiTil by Young and more fully elaborated by Helmholtx) strirci In
rodui^p the matter V* still further »impliciiy. Starling fVum the fact men-
lk>nci| a Fhort tinie since, that all co'or sensations, including the sentutlioti of
whil<\ may be obtained by the apprt>printe mixture of thr«« standuni iicnin-
tioiiH, this theory leacha that our visual apparatus ii m> eonsliluteil a-*, wheii
excited, to give rise 1" three primary senaationa. and that these primary aen-
aaiions are called forth in diffi^rent degre«a by diflereni ray* of light, •» that
each ray givea rise to ■ dilfereot mixture of the three, ^veral aetn of thiee
■neb (irtmarj seuantiom might be cltoaen. which would wliafy the oondiiiiH
BSS SIORT.
of ^viiig; Hm, by npproprialv ntixturc. Ut all soBMiiona n( vnlor, includiBi
wbiU:; bul for rmwtiu into whii-h wc catiix)! enter fully hero, tbe SKimigM
which mik}- thii» be tukcD » priinnry seiistuioiiit aitpear to dfrnwpoad bi tm
•enwittonH of re<l, gmn, ouil blue ur viulel. Such n mw uf three (trimiT
cukir wnvntioiu it repraaonted in the dingraoi (Fig. 233). Tbiie llie m
rra. at
^
I
X 0 f i-i-
VfOMikm or Tunc* f'KlHlR^ v
ll«lhP(D-r«lIi?4"i«d.~ :"K'W"/' ■ml a " dclut " n. 'Uilan S. O. r. ct&. nfil*
•inC IhetTd. unnic. ftllow. r[c., wlor oT Ibe ii|«rtmm. uid Uic dlavum •hmrt bytW btlrklrflli
citnain cavh i-nv, ■■» wlul cileiil Um wvurkl |«liiiArr tulot MOMltsa* on Tt>|<aiatvlT «MIM tf
TlbmUou ofilllhivul ■■Tc-lvugllu.
prioiiuy wnMUion, cxcitcil to a cbH^ extent by the nr* *t ihf vxtrrnic ml
VD<I, it mtM powtirfully oBbcted by lltv nivs lU n liltlv di«tiincr from the vat,
tJio rays from this point onward tuwiird tin? blue end producing lev aail hs
eflect. The curv« of the green primary eensiition begins later, sad reaeha
ile ntaxiiDum in the green of lli« epet-irum, while the blue or viulet primarr
MDBnlion IB Blill later, am) only reaches iu inaxinmin totrari! the blue end it
l)ie e{H.'clrum. Each ray euth furtb eucb sensaliuii, bul in a ditlereut dccRC,
and ibe lulnl reault uf each lay. or •.(' each group of rays, is deti^rniiiMU bj
tJte proportionate auinunl of the three eeuaalioiis. Thus the aeoMtiaii A
orange (O in the ligiira) is brou;;ht about by a mixture of a great deal tf
the primary nn] with inucli lew of the primary (trwn, and haixlly any oftk*
iriinarv blut!; the »rung« oc-iiMlitin is converted into a yellow •«nMiliati lif
iminisliing tl)e primary red and largely increasing; il>e priomry ;;rr<rn. thl
pritoar}' blue unitergoin^ iiUo sunit; dli((ht increase. And uniilur ly with all Ht
other (cnnt ions. When ench uf the primary Memotiona ia excitet) lo a tiiasi-
rouin, a» when ordinary light falls uu the retiua, the leault ii a artyaliua of
nbiie. Atvonling to this theory, block is Bim])iy (he abtencc of i)inTiiiii
frriRi the vituni nppiiriiluK.
In the view, as urigtnallv put forward by Young, the three primary no-
aations ncrc suppowu to )» reiirrH-iilcd by three m.1* of Bbr«a, encfa m4 of
Hhree being dirtcrrntly allcctcd Ly ditllrent reyii of lii{hl. and the impulHl
SitBsing to the brain along each »ct a»akoniiig a diiitiuct teoMtion. No such
Mlinclioti of fibres can be found in the retina ; bul an auatomicnl boiit of
thi» kind is not neecamry for the Lhcsiry ; we can ewily eoDoeive of iheMiw
librr- transmitting three ili«tinct kitt'U of impubea; or we uty supfMee thai
thv visual lubstancw are three iu number instead dfxix, the cnongee in Mwh
Hulntanco proroking a primary wnsatiun.
titich are the two nmin theories of color vinon ; and miieb may be mU ia
tavoruf both of tbom; at the same time both of tliem prevent raanv diffleal*
liea. To diacuas litem fully is a task beyond the limita of this booV. and lo
discu.« them in any but a full manuer would be unsatiafacUiry. We muiI
S
I
I
iti*8cd, tkcrrrfore, witb l!i« furring simple stnl^'iiirni nf tin* mo twms-
TtHlefMlDdcDll.T of any thcon-, however, we mny remciiit*r 1 1 t ihfll nil tliu
MMiMili»iii> wliicli Kc nxjwricDov uihIpt the sclioii i>r light of whnii-ver kind
amy hv ri-<liicc() to sii, while, binck, rrd, vdlow, gr^o. nid blue : nitd (2)
ibit ibcar m«y Iw all rrprrMlnced by various niiiKurM of three •liinilard mu*
MlioilR, if bluck be alhtwitl to indicate tbo absence of all wiHstion. Thmo
»tt nintlrreof fact, whal ta nl present ile baled is whether the six rundnmrDial
■Biintions are the outennie of l.hree primary censalioiis or whether they rcpre*
wot lix diitiiiet conditions of the visual apparatus.
Chtor hiiuilneit. Persons vary niueb iii ibdr power of BpprecKitin^ sn<l
discritninaliDg fiolor, »'■ «,, in the inleusiiy nn<t accuracv of ineir color »en«a<
lions: and »onie peonle reKanI a^ similar, t'olun which (o ninst people arc
i;larin;;ly dirtiiid ; tliese taller are i>ai<l to he " color-bliud." The most com-
mon form of cilur-htioduess is that of (wrsooH unable to diiUiuKuish j;rcen
aD<l red from each other As iii the case of Daltim. ther tell a red gown
lyinn <«i a (jre*" Ifraw plot, or a red chernr aumu): the ([r*eii leaves, by its
foru), and not by its color. Tli«y confound nol only red, green, and wrlain
rorma of brown, hut also rme, jiurplt^, utiil blue. Such |>erso»s ure oflcn
vpoken of oa " re<l blind." On the Herin); theory ihey lack the red-icrcen
visual substance : bene*, nil t)ie color lu-nwitionii tbey piiaMW must he lliose of
yellow Hud blue fre^l from all mixture of nil ur ftrceu; and such ai-counts
as have li«en itivi'ii of their »cnsuli»ns by those pi-nons who are "red blind'*
in one eve, but piMtn* normal vi»iun with the other, acconi with tliw ronclu-
■jon. On the Young- Hetinholii theorv, «uch persons Invlc the priinnry red
MDMlion; and bdico tbo femntious wtiicb thev have miiri be niixturta of
grwn and blue alone, our yellow appearing to ihvni n bright ^r«cn, and our
gnrn-bluc a kind of gray.
All such red-blind peupla ought, on either tlieorr. to be len aifeded thao
are persons with nonnal eves, by the red end of the spectrum ; this ought
with (l>ein to be •bortenea and obscure. In a certain number of persons
who confound red and green, this b the case ; but in aoroa instances no such
lack of appreciaiitinuf the red end of (he spectrum can be aacertained. Kuch
esses have lieen suppcaed to be t-r^n blind, ibat b. lacking the primary sen-
Mliuti of Ereen. According tit the Ueriiit; rhei>ry >;re«o blindness apart from
retl blindiifw bimpooiible, the only two piMeible color defects being rc>i*);reen
and bltie-yellow bliuducw. And ihe exbienee of dblioci green blindness
faaa been held to cotiinidkt that theory. On the oilier hand, the Ilering
theory admit* the pomibiliiy of total color hi indm-M. ■'. e., ihe inability lu see
anylbinir but white and black ; and thim. on the Yuting HelmholtE theory, U
impomibUt, *iuoe for viii-m t» exist at all, oue of the throe pnuiary iwtisations
muit l)c prMH'nt ; n man to «ee at all must aee things in various shades o(
(itbitr rvd, or of ^rren, or of viidet, though he mav confouml this sinKlc-
eolONd vioion with the normal vision of white of didtrrnt inienstliea. Bui,
indeed, n full examination of color- blindnras ratlier increusva tbati diminisbea
the difficullies of deciding Wlwetui the two rival theories. *
Injiuimce of the piffmeiil of the ^Uow »fiol. In thi> macula lulea, wbidi
(MM of tbe retina we um> chiHly for vision, images fulling on other jMrtx of
ihp rrlina bting wiid to give riw to " indirect virion," the yellow pigment
■ImoHi" some of the grccnishbtue rayii. Hcnoo, the Mnsation which we re-
ceive from objects which wo art- in the hnbil of calling white i« that whidi,
if this pigiuetit were absent, wc should receive from ohipcl* more or l(w yel-
low. We may use thb feature of tliti yellow xpot for the purpose of mnking
ibe S|iot, so to speak, rinble to onnelvn*. by an experiment euggcHt<i| by
MuKwell. A solution of chrome nlum, which only triinsmits rod and grem-
i(b-blue rayv, it bdd up between the eye aud n white cloud. The greenbh-
tm
BIODT.
blue myt ar« alMurbeil by tk« yellow sji<it, ami here the liifbt ifittt riae la a
MliMtiaii of red ; wheroiu in C\)t ml i>{ tbft field of viaioii. ibe Mosuim «
tliKt onlioiirily iiroducuil by ihv piirpltih solution. The yellow spot MCOBK-
<|U«iiLl3r Dtnrkuil oui u» » riMy jintch. ThM very Ktna. buwerer, ai«« ftWay.
In sptflkin^ urMiiiuition lu a I'oiictinn i>f tliu riiitiiiiu*, p. !)15, we nfetn<\
t(i wbit£ liglit i>iily: biil the dilTcruut oi>]ur* urv tincuuiil la the reUli4U
biinui by thit tnlviixily »!' ihti utiniiiliin, to the amauiil of AOiutitiuD proilaeel
Thii* iJie lann: rcfrnn^biv bliut nyt procljoe K winwtioa more r«ia<li1y ihin
thii jolloiv nr rdl riijA. Henco, in dim lighu,iu th>wB of evening nu>l oiiMa-
ligbt, the bluet Drupnnili^-iiv, itml the ti^U iinH yellov* arc le»* olivi'iiis. Sr
ftlnQ u'hm a UekW-mjili i« viitwcil tbrougb a yvlloiv glim, the yellow hue Wf
gi-siA to the mind bright sunlight ninJ auniiner •toiktber, ailbimgh tlie acwal
illiiniii)nli«n which reaches the eve i* dimini^hoil by the gloM. C>iiV«rMlir.
when the e-jino kad^i-itpe i« vioiceil through u bliio glius the iilvn of miioDlisil
or winter ie tuggesk-d.
The iheiiry ol* three printnry color wnsattons mny bo usol to explain wliy
nDV coliired light, if iniidu BuHieiently iut«o»e. appears white. Thii* n riultl
light of iDodenite intensity npiieiirs violet bocAiise it e]ccit4» the primary Mn-
Balion of violet much more thiin ihiisu of green and rod. If thu slintului ha
inerejised the mosiitium of violet stimulation trill be reaehed, while tbe«U*0'
latioH of green will coutiauc lo be In^-reaKd aod even that of reil In ■ slij^t
degree. The result will be that the li;rhl apjieitn violet niixe*! with gTMO.
that is blue. If tbe BtimuluB be Mill t<ii'th«r inereosed while the f;reeii and
violet are bi>tb excited [o tbe iiiaxiniiiiii, the red atimulatiuu mar b« increainl
uniil ilie rt-tiult U viol^i, greea. aiid red iu the pruportioua whteb malu) wUlt
light. And s> with li;(hl of other colora.
After- imiige*. We have alreadv se^u that in virion the sensation ImK
much longer than the «tiniu)ii:>. Under eeniiiii cireum.ilanccri, iiueh a» p■^
ticular C'luditiim* iif the eve; aa inli-ii-te *tiinulu*, etc., the acnv^tiim it m
pmlongi-d, that it is ypokcn of n« an nfu-r iniii/u. Thut, if the evi' b«t tllrecteH
to tlic NUn, ibe imn^ of ihnt hi>dy is prcM^it for u Itxig whilv itfter ; aixl \S,
<tn varly waking, ine eye be dirccteii to the window fur an in.itaiit guid then
vl'ifCfl, nil iiuai^i- of tbe window wiib ils brighl ikioi.'ii and ilarkfr «a<h4. thi
vari'>iis parts being of the same color as the object, will remain fur nu Bpfr»
dable time. These images, which are simply continuatiovs of tho sodMUioa.
ftre spokea of as potitive <i)Ur-iinaoa*. They are best aeon after a awmeatsry
wiiiosure of the eye to the slimulus,
VVhen, however, the eye has boon lor some time subject to a stimulus, the
aeusatioii which follows the withdrawal of the stimulus » of a ilidoroDt kivl:
what H called a neijatine aflfrimaiff, or n^galivr tinijre, is pruducod. If. »&»f
looking sleadfaslly at a while patch on u black gri>uu<l, to* *y» he turned Is
a white ground, n );ruy |Httch b seen fur some little lima. A olnck Datch <m
a white ground similarly gives rise on a yray ground to a uegatiw imagsffl
the form of a white patch. Thi« may be eX|dnitR'd as the result of exltaot-
tion. Whcn'the white patch htis been looked at steadily for SOnM time, \it$
part of the retina on which the ima<;e of the pntch fell beixmos tired : h«aM.
llie white light, coming from the white ground siibjeriueiitly Uiokei) at, which
falls on this part of the retina. d.>ej not produce »i much seuMtioQ as in
otl>er parts of the retina ; and ihu ima-;e. c lo^e jueutlj, appean gr>«y. AnJ
•0 in the other iu^tance. the whide of the retina is tirea, except at the patch;
hare the rettua n for a while man seoiiiire, aD<l lienee the white negativt
image.
When a red patch fs looked at, the negative image is a green*blue. that u.
the color of the negative image la complementary to that of tlie objeot. Thai
aUo, orange produces a blue, green a pink, yellow an indigv-blue, oagaliti
»
I
nnsf!«, iind so nn. Tliis, loo, can be exjilBinof) ns n mult of exhauUion on
either liriMlbettis of color vifion. When the colored patch it limked aC. one
of llie itir«« jiriniary color ecttsationa h niiicb cxlinutitd, ami ihc ii<her Iwn
leu au, it) vKryiiis proporiioos, according to the csaci natuic of the color of
Ibc |«tcb ; and Ine less exhauBied Moeaiiona become promineut iii tli« afrer-
imatt^. Thus, tli« red jialch eiliauels lli« rrd fcDfatiun, and ibe negative
imaxe ia made ui> chiefly of green and blue aeunilionF, tbal is. appears lo W
Cnh-blue, ur uluttb Kreen, according lo the lint of the red. Oa the other
ilbeiia, «e ins;^' luppoee that, owing to Uie continned eifect of lookiue at
ifie red patch. diFxiniilaliou of the red'){re<'D auhatance becumea len aud Tcm,
lemdiu^ to a pnimioeoce and, indeed, to an actual iDcreaae of tbe proccaa of
nfr*iuiiiaii<in ol tbe name lubaiauce; h<-n<«, ibeteiiMitiou uf green dominating
in ihr nrfTiilive iinaire.
SioilUrly, «beii the eve. after lookiii); at a colored |«[ich, is turned to a
colored gruoud, the efli-cia may tuiily hv explained by relereuoc to the cum-
parsiire exbauatioii ul' ilie cnliir peiieatiima rxciied br ihe jMitch and the
ground re)>{)eciively ; if u yelton- gnmnd be cbiit-n after lunkini; at a ^r«vu
object, the aegntire imnge will iip|ieur nl' n reilii(>h yelluw. and *ii uo.
The theory of three primarv KnrntiutiH dn«-n not to readily ex[>lain why
tifK*l>ve iiuagei nhould ninki; tm-ir apjicnranco without nay >ul tieifiienl Mimu-
latioii of the relitiB. When the eye* nre tliut and all nceew n) liuhl, evt-ti
ibroMgh ibc eyelid*, carefully avuidcd, tb« field uf vi»iou i* not abtulutcly
diu-k : there ii *till u iKiiHtioo of light, the •u-called " proper light " of iIk
retina. Ifa white patch on a black ground be looked nt tor «oni« time, and
lh« cyca then shut, a negative ^black) image of ihc ajwl will be Men on th«
grouitd of the " proper light " of Ihe retina, having in its immediate neigh-
EnriHod a ijiecially briebt corvtiii. iyo. alM>, ir a window be looked at and
lb* tjtt then clorcd, iTie pmitive afler-iniafic with bright jmiirs and dark
Miches gives rise lo a ne^ativo alter-imsge nilli hright sastiefl nod dark panca;
and similar eflecis appear wiih culora. I'hcc^ and amiUr facte have been
largely u»rd in support of tbe Uerinj; tfaeurr. When the eye has been
looking at red, and to has raUMd diiainitlaliou of llw red-green substance
mere re»l. as on thutiing the erts, favurs as»iniiIntion of the same substance
and thus leads to a aeniuition iif green. And tbe rbylhuiio oscillations from
one color to its correlative and batk again, fre<|uentty obeerved under theee
conditions aud irbich puint to asimilatiuii and Uiafeimilntion alietnately
ffmiag the upper band, are nut witbuutanalogiea in other commoo iuataocee
uf {iroiDplaiiiiie metabolism.
VnOAL Pekceptioxs.
I Hilherlo we have studied fenfaiions only. and have conndercd an external
' object, fueh ea a tree, a* rimply a aourct of «o many dtttinct •ensntioiii*,
diflering from each other in inicn»ily and kind (color). In Ihe mind theia
eviisatioiu are ooordinalcti into n pcixeption. We are not unly voiHCtoua l>f
a number »r «cnsaiions uf bright and dim lightv.ofgrem.brunn, black, etc.,
but tJicM M-HMitiuDs are so rclatetl to each other and by virtue of eeRbrai
Moctwie m fashioned into a whole, that we " ew A tree." Wo aooaeUnee, in
lUwtntion of such «n effect, speak of an image or picture in tb* oiIim) cor-
KHKniding to the phvbicnl image on the retina.
E When we look upon the external world, a variety of imagee are Ibroied at
' tbe aame time on tbe retina, and give rise lo a number of contempMWWoua
visual sensations. Tho sum of thc«c sensations eonetilntea "the field of
viaioo," vhieh varies, uf voume, with every luovemeni of tbe eye. Tliie field
926
SIQBT.
(if %'ifUMi, bein^ id ranlilr an iiegrvgatc of tramionfi, is of omirM « mijttliwi
nmUor ; biit we arc in tlici hnbit of uHDg ih* nme phrase to denote ih« Mm
of cstemul objvcts which give Hno to the wgregat« of vi<ual semttiuiu; in
common langutti^ the field of viatoD is " alllhiil we can Eoe " in iinv (raaib<«
of ihc eye, and wc have n field of VMon for csch eye separately and for th«
im) «ym combioed.
Using for the present tbc ironia in their Bubjevtive Benae, w« may renark,
(hat we are nble to neaigo lo caeh conatititenl ^Titration tta place aiDoag Uw
nggrei^Rlc of senaatiooa const i tilling the field of viaion ; we can, aa we Hf,
toealiie the senaalion. We chusiiv n'liether itbelooga to (what we regard ai)
the right hand or leA hand, the upper or the lower port, of the 6eUI «(
viiion. We are able to disliogiiish the relative poeitiona of any two diaiocl
senaaltona; and the relniive pusitions, together wiih the relative inteuilio
sud qualitM (oolor) of the Benaaiiona arising frifiii any i>hje<Tt determiiw our
perception of the object. Il need huriily be remurke^l that this lucalitatMa
IB purely aubjective. We simply determine the p<»idoii of the M/uarion ia
ihc lieli] of rikion (which is itself a wholly tuhjective matter) ; we do not
deleroiiDe the pi»ilion of the oijKf. The cuniieettoo botwevu the jioikim
of the object iu the external world and the iKwiilon of tiic Mnmtion in tha
field of vision, cannot be determined liy visual uboervatimi alonr. All tb»
inlbnoalion which can be joined bv the eye is timitci) to tbe field nf virion,
iind provided that the relative pnsilion of ihe wnxnitiuis iii the fivM of vixnii
reiaained the kame, the actual p<»itioii of exiunial nbjccu miglit,iufiuH
tUod is ooncenicd. be chikuited without our bciiijc airiire of it.
Asa matter of fact the lield of vimon in one im|M>rtant gittrticuIaT ika
not correapond tij the field of external oinects. The inittgv on ihr rrtina if
inv«rtMl; the mvH of light proceeilinj; <r<ini an object which by touc^ «i
know to be on wlial we call our right hniid, full on the lefthund »i<Ie uf tht
retina. If. therefore, the firld nf viiiion currcMtHindcxI to the retinal inaM,
the object would be aeen oti the lef^' hnnd. U c, hoirever, see il on the ri^
hand, bceanse we invariiibly aiuiociate right-hand tactile localiutiuo aitk
left-hand visual looilixiition ; thai ii to wr, niir lield of piston, nhen inur-
pretcd by touch, is n re-inver»i»n of the reiinnl image.
The diraeosions of the fichl of vision of a single eye ar« abntit 146 Afgna
for the horizonlnl and lUO degrees for the vertical meriilian, the former
being distinctly grenler than tho latter. The lioriiontal dinieiision of lb*
field of vision for the two eyw is aboni 11*0 degrees. By moTemenin ii(" the
ejfcf, however, eveu apart frum those uf the bead, tiw utent may be coD'
siderably increased.
The satinfactory perception of external objects re<)uirea distinct tUm;
and uf this, aa wo have already said, the formation ol a distinvl Etnagt mi
the retina is an caeeotial condition. We can receive visual sensation' uf all
kinds with the moBt imperfect dioptric appamtus, but our pnrceptioo of aa
object is precise in proportion lo the cleanieaa of the image oa th« ntina.
Rtgion of distinct niti'iii. If we take two pointa. nieh at Iwu bJack iatk
only just so far apart that lliey can be seen disiinelly as two wbu ptacii)
near the axis of virion, and then, keeping the axis Bxed, move i)i« two mfiui
out into the circumferenlinl parts of the field of tIbod, it will Ix- fonnii llial
the two soon apjfear as one. The two sensations beeome fiin.tl.aii ihvy wwM
do if bnmght nearer ti> each other iu the center of the field. Thu funhn
away fmni the centre of the field, the further apart must two pi>iDt« be la
order that ihey may be seen ns two. In other ironb, visioa Is much mart
distinct in the centre of the lield than lowanl the dreuniferetice. Pradi-
cally tli« region of distinct vision may be said to ho timilnl to the macuU
lutea, or even lo the fovea centralis ; by continual aiOTemeiila of the eye ire
»
lire comlnntlr bringiiig nny object nhidi vc niih (o mc ia mrh b po«iiiuii
thst lis itns^ fftlla no ihis rcjiiou of tlic rotinn.
The diiiiinutiun of diHtmcln«» does ui)t lukc plucr c<)iinlly frum tli« ct'iitro
to the eirciimlereiK-'e nlong *ll ni«ridian>. Tlie outline di'schbed \ty a line
uniliiic ihv points where tvo spots oeaM to be ttca ns tn'u nhcii ii)ov«tt alunj;
di6ei«Ri rami from the centre, ia a very irrq;"'"' figure.
The seoMtioDB of color are much more distinct in the centre of the retina,
tban towsrd the circumference. If the visual axis be Bxed and a pi«ce of
colored paper be moved tunard (he outside of the field of riaton, the culur
tinderp'ea cbanfrn and ia eventually lost, red disapitearinj; llru, and blue
1a»t, the objwt renmininH visible, though with very indistiurt ouilinei. when
tt» »>lor can be no Um^r recoitnitied. A purple color bec<>itie# blue, an<l a
riitie color n tihiul) nbit«. In fact, t lie re seems to be a certain aiDount uf
r«d-bUndi>e«* iu tbe |>eri(iheral parts of all retiuHS.
Modifieit Pentptiont.
Since oar perception of external objects b bued on the dinlinclntM of the
misations which so to form the pcrci-cition, it might be cxiwlcd that when
ao image of an object it formed on tntr retina ine Mmitory iinjmlM^ would
coiTMpond to the retinal imii^, ihv^ •cuMtlionN cirntponil to the sensory im-
pubnanil the i>crccpt)on cnrrtvpond in the M'nNilivnt, and ihnt, therefore,
(he mrnitil ot<naiti(iii mullinji IViim our Incikiii^ at any aii}t*:t or view would
corm|wnd csnctly (o the n.-tinal imsEC. Wc hnd, however, thai this is not
the cnre. The Mnentions and probably even the simple setimry iinpubea
produced by an image n;nct upon Mcb other, and thefe reactions modily our
mrcvplions, independently of tb» pbysicnl conditions of tbe retinal imase.
There ariae oeriaiu diacrcpanciea between the retinal image and ili« jiercep-
t*(>n, some having tlieir siiurce in the retina, some in the bruin, uiid <ith«n
beiuji (if Bucb a nature, that it is difficult to say where tbe irrelevancy is
introduced.
Jiraiiialion. A white patch on a dark ground appears larger, and a dark
patch 00 a white ground Soulier, than it really is. Tbio t* e»pecially»o
when the object is somewhat out of focus, and may. in this case, be partly
explainet] by tbe difluuon circles nhich, in each case, encroach from the
white u]Mm the dark. But over and beyuml ihi«. any Mu:»ati<in, coming
from a given retinal area, oocufdea a larg<^r nbare of tlie livid of vijtioti, wban
the rt:>t of tlic rclina atid central visual apjiarstus are at reel, than wli«d
they are simultancoHsly excited. It is as it tbe neijfbbonn^, either retinal
or cuebrvl, structures were sym|>aiheuca]ly thrown into aouon at tbe saow
titnsL
Omtrtul. If ft white strip be placed between two black ilri|M, ibe edges
of the while strip, nnr to ine black, will np[>ear whiter than iu median por-
tion; and if a while cross lie placed on a Mack iMckgniuod, the centre of
tbs cron will appear somctimoi su dim, compared witb the }»rt» cluee U* the
black, as to seem shaded. This occurs even when tbo object is well in foous ;
tbe increased tcDsation of light which cause* the apparent greater whttoiea
of the borders of the cross is the result of tbe " contrast " wiih tbe Uaok
place<l imrordiately clwe to iu Still more curious results are seen with
colored objecls. If a small piece of gray paper be placed on a sheet of
green )>Di>cr. and both covered with a ihret of thin ttteue paper, the wny
paper will appear of a tunk color, the complementary of the green. Thn
eOeet of contrast is lar legs ttriking, ur even wholly absent, when the snuill
piece of paper >« white iutMd of gray, iiud geDCralljr disappears Nben the
siauT.
tbin covering of tiwuc jMpcr i^ removed. It ibo Taniibc* if a hold, brnul
binck liDci Ik <lrHwn round the small piece of {wpcr, m m to iaulAle ii frum
Ihe ground color. If s hook, or pcni-il, be placed vertically on s (bNt u(
while paper, nncl illuminated on ooe Fidv Ity the mu. and the otlwr tij a
candle, two shaHonK will be produced, one Trum the auo which will bv Ql»
niinnieil by the yelluwifh ligtil of the candle, imd tlte other frvni Ihe caaiU*
nbich will in turn IwillDminiUcd bv the nhilelii;hl of the sun. The fnmtr
Datiirully appear* yelloir; the Inttcr. however. ap|>eara not while hut blue:
it anunics, by contrast, a color eoin piemen 1 a ry lo that of the eBuilli.'-li^
nbtch eurrounds it. Iftbo candle be removed, or ii» lif;hl shut Kfr by a
ccrceo. the blue lint diiappeare, but retiirDs when liie caudle i» a^^ain allimed
lo produce ile shadow. If, before the candle a brought baek, a vim»u W
directed through a Durrow blackened luhe at some part lallin^ entirely wiiJuB
the Biea of nhat will be the candle'f shadow. tiK arem, nhich in the ahaoiTC
of the candle appears while, will mniinue to appear while when the caaUe
is made to cast iis shadow, and it is not until the directiun of tb« tube b
changed so as to cover pan of the ground outside the shaduw, ua well at put
of the shadow, iliat the latter aaHuines its blue tint.
FUling vfi Hie blind *poE. Though, as ae have seeu, that |iartof the nitaa
which corresfionda to the entrance of the optic iierre io quite (iMooaibfal*
light, we are conscious of no h)ank in the tleld of ruion. When in lo"kitt
at B paue of print we llx the visual axis so thai aume of the prim luuM fiifl
till ll)« blind s|M>l. no gap in [M.-rceived. We ciiuld not cx|hh-i to ncv a black
patch, Itecau^e what we call bluuk is the ubteneeof the T^tioaiiun of ligU
ln>ni sirnetur^-a which are »en»itive to light; we must have visual urgaMt*
■i:c hinc'k. But (here are no vimihI nrgauM in Uiu blind sjwt, aud ci'iiM-qucotl;
we are in no n-ay at all nltbcieii by the nip of li^ht which tkll mi it. Tlwn
Ik in our subj<-c-tivv field of vision no gap cormpundinfr !» the gap in I bt
retinal image. We rclcr ihc venial i>in:> coming from two piiiiiis nf ihe i
lying cm uppostlc margins uf the blind spot lo two points lying cloae to
(jnce we hnw no indication of the spitcc which Hcpirates thcui. Om
the efleclE which are pn>duccd when an object in Ihe field of view paa
the region of the blind spot there boa been much diccuniua. In (irdiaarr
f bion, of course, the existence of the blind sp>t is of little fnonient sltiWB
is outside I he region u»ed for distinct vision, and besides ihe image uf ■■
ubjecl does not lall on the blind spola of b(rlh eyes at the same tinier
Oeu/iir ^peef ivi. Ho far from our perceptions exactly currespuiiding toi^
•rransemenla of the luminous ra^s which fidl on the retina, «c may han
visaaTMn 1^1 ions and |ierceplions in the entire obsence of light. Any fiiiaif
lullon ol ihe retina ur of the optic nerve »utHcientlv inteuae will gini riw ID
M visual sinsatiun, (irndual prei^ure on the eyeball otusei a •enaaliun ol
rings ol i-oloicd light, the socalk-d jiboephcuee; a tiudden bluw ou the (yt
caust^ a M-n^aiiun of tlasbes of lij^-hi, and the «eeming idL-uiity nf the vwisl
»en.'«tions m brought about with vinual tvnxatiotiB prcMJuced hy light b aril
illuMtnili'd by ihe i>laii'iui-nt once gravirly made in a German cmrt uf law, by
a niluwo nho atraertfil that on a pilchdark night he recognirjil an awailaal
by help of Ihc flaah of light cauM-d hy the iiaMMlant'v hand ciiuiiig in viiiitBl
coniaot with his eye. Klectrical MiniulalioD of the eye or wpiic nerve will
•Iw give rise to vitual seoMitloDs.
Tlio acnntioiu nhicli muy ariiw wilhout any ligbl falling on the Rtias
need not necessarily he undctintd ; on the contrary they may Iw moai cinHy
dcBnctl. C-omplex and coherent visual imagex or peicvptiuna may ariie is
llw brain without any corresponding objective luminoiu cause 'l''- — •»■
called ocular spectra or phantoms, which are the nuult of an inin u-
lation uf some (probably cerebral) part of the visual npparutin, litv a li*-
stogetbt^
Vxacer^^H
MssesMP
TIBIAL flBOEPTIOIfS.
llnc(nr«t wliivh givca them nn nppnrcnt olijrclivc mlity qiiiu- ns strikiiiK as
ihBt oroniiiinn' viRual pvrcepliniiB. Thoy mny occnsionallv bcseen with th«
ev<9 tifH-n (iind ihenefure wliilo ordinary viiiinl pfrccpliitiis arv being gcn-
emied»at veil hc when (lie eyes aio closed. Tney sometimes becuaie so
ft«<)uent Bnd ol>lni»iTe as lo be di«(rr««in|;, tind J'onu an important elanSHll
io sunie kinds of delirium, such as delirium tremens.
A}iiTftiulion of apttarenl tite. Uy the eye alone we can only estimate the
appaifitl tiie of an ottjecl, ire can only tell what space it lakes in the Held of
Tieion, «e can rinly percvive the dimensions of the retiDnl image, ami there-
fore have a tijiitl only lu speak of the angle which the diameter of the object
•ul>teii<ls. The rent sizu of an object niuat be determined by other means.
But our [)eiv«iilion of even the appareiil eixe of an object is so modified by
coiKurrttil ciintnislnnces that in many cases it cannot be relied on. The
RpptircDi >iieof the muon nmat be the aame to erery eye, and yet while some
peraouB will be found ready to compare the moon in mid-lieaven^ with a three-
penny iiie<-i>, Mtliere will liken i( to a cart-wheel; that is to say, the ooffle
auklriMcil by ihenioooseemaioiheoneto he about eigoal totbataubteDded oy
a lfaree-t<eiiuy piece held at the diainnce from the eye at ivhich it u most
commonly hmked at, ami i» the other about equal tu thnt Aubiended by a
curt-whevl similarly vivHfil at the dislance at which it is luMt commonly
Inokt'd at. If a line niich an Ai', Fix- 23'l' ^e divided into twoe4]ual narta,
A B, /> C. nixl A H lie divided bydiMinctiuarks iuiowvernl parts, aa U sDuwn
in the livnro, while li V he Icit eiilire, the distance A ti will always appear
{;ivii|pr ilinn C'B. Su nlno, if two eijunl squnrea be marke<l, one with biiri-
■uiital and iIm oilier wiib vertical alternate dark and light banils, lh« former
rto.su.
I
wHI appmr higher, and the latter hmader, than it really is, Ilcnce short
pprvnus uili-ci drtm-A liuriznmally tiriped ju order to increase their apparent
WiKlii, aod veiy sioiit persons avoid lon)[itudiual slripee. Two pencctly
IiHrallel l>tii-» or bands, each of uhivh is crowed by slanting pamltel short
inn, will api^ar nut parallel, but dirergiiig or converging according to ilie
direction of the vrtm-linea.
AKKin, when n short jierson is placed title by fide with a lall {)er»(»), the
fortner aiipeara sborier and the liilier taller than cacti rually h. The muuii
(Ml the hiiriuiD ap|>fara larger than when at the xciiilh, becaune in the lint
ptMiiiuD it ran be ntwt ea>ily cuitipiired wiib icrmtrial ohjecis. The absence
uf CQm|«r1iK>n may, boHever. ctjnirilHiic to nn opposite effect, a* when a
pet*oi|i Innks larger in a f<>x; hciugsveti inili*tinclly,hu is judgeil to be furtJier
oil' than he ivally i«, and >o ap^it-nni larger ihnu he natuinllv would do at the
(ivtani'e at which 1h- ii> suppiurd tu Itc. Ht>, onnvcmly, duumt niountaiiia,
when Kcn dtitiuclly in n L-lear alnKupbcrc appear small, because on aocouni
of their di>i!ncin««a they are judged to be nearer than they really arc. Indeed,
«<ir daily lite is full of inMancea in which unr direct perception is inodifitd
bv circuniM niters'. Among ibo*c circiinmtnncca previous experience i* one of
lUt most |H>irnt, and thus simple {icrceplions btoouM mingled with « hat arc
In rralily judcmeots, though freqiirutly made unconsciously. But this
intniKion ft lui-.! eicpericnco inin present perceptions and sensation* la must
' "iotia in binocular vision, lo which we now turn.
930
STOUT.
rw.3».
BiKociri^AS VmoM.
CbrrapondiHy or Jtimtieal PotnU.
Though we bav« two eyes, nod must therorore r«ceirp froRi every ohJM
IwQMU uf seniiatioDB, our perceplian of any object is under ordinHry cimiK-
rtunrex > bihk's one ; we aee one object, not two. By mitting vilher eve imo
nn nnueual jKisitiun, as by BquioliDg, we can render (tie perception aoabl«;
1TC eee two objects where one only cxisrs. From which it is eri))etii thit
tinfjleness nf perception depends on tbe imM
ol' the object Imlling on certain jmuIa of ttm
(B reliiis at the same time, these parts betOf K
relaied to eaeh irtber that the Aenxatidna tnm
eu'h are blended into one perceptioQ : and il
is also evident that the inovetneau of the ej^
balk are adapted to brinf; the inia^ of ibe
object to fall on these " eorrMpui)di»ff " or
" tdeutical " parts, as they are called, of each
1 IVV I retina.
When we look at an object wiili one cyv
the visual axis of that eye ts direcied tn Ui«
object, and when we use two eyes tbe ridul
axei of the two eyw couverjje at ih« ol^M,
tbe erebalU innving accordiujtiy. Tli«
spondiiif; points or tbe two retinBH arv tU^H
wvHutKo iwrm.
t, Uu> lea; K. tlie tWbl Of*: K,
tt»(>|M(«)a>Dn«:<ti,lk,.(^ftN (wlnu
in thi rtf he •TV eornaimidluit ii> iha
pot nil n. (I. cin lUalatt efc Tlietwo
flipua bcloiF arc ptQlacaaa* ttl L. Ui«
Ivfl, anil a, iliw r^t inlnii li will
t« imii llwl 0,1111 Uju muter udpoft.
curmpuuila lu Oi, ou (be nuMi aide
at B.
OU which the two images of ihr nbjwt
when ihe visual axes cuuverKe at tbe ubycct
ThuB in Fig. 2Sr>. if Ce, C'c, be (bo two viMul
axe*, c, e, being tbo ocnirm of the fuvw>«n-
Iralva of the two eyes, then, tbe object ACS
btttig seen single, the point a on ibe oo«
rciina will " corr4«|Kin<( " tn or be " ideotfaal"
with the prant a, »n the other, and tbe raial
& in theoiK^ toihcpointi,in theoth«r. tunct
a p')ini lying anywhere on tli« right side of dm
retina hu:' il* cQrrcsi>on<Iing point i>n tbe n^
side of the other rc-tiiia, ami the f>oiu{s oo ibt
left of one correspond with tboso oo tb« l«ft of tbe other. Thus, while tbf
U|>p!r half of tbe retina of the left eye corresponds to the upi^er ball' of tin
reima of the right eye, and the lower to the lower, tbo hkwo/ side of ihe llA
eye correiipiinili> wilh the malar side of the right, and tbe m-ttnr *it ths
left with flic natal vide of the right.
Tbe bliMiding of the two sensations into one only occurs when lbs tws
images of nil oliject fall on thrifo corresponding points of tbe two rptiaK
HeDoo it is obvious that in single vision with two eyes the ordinaiy
iDoveinenIs of the eyeballs mtiiit be such as to bring the Visual axes to aB-
verge at the ohjuci so ttnil the two images may fall on corresponding poiala
When the visual axes do not so converge, and when, tliereli>n<, tbe 111114H
do not tall ou corns>[KMidiDg points, the two senaattoui ar« not blended ioU
one porecptton and vision becomes double.
MovanenU 1^ the Ej/ebalU.
Tbe eye is virtually a ball placed in a socket, the bulb and tbt orU
forming a bull-and-aouket joint. la its socket-joint the optic ball is capsUr
BINOCULAB TI8I0X.
031
P
PW.SM.
ciU^,
\
M.i
of A viiriet]r of nioT«iii«ate. but it cnntiut bv anv vijliinlftry eflbii b« moved
i>iil of ilB soctivl. It is slainl tbut by n vcrj torrible (iponiog of tbo ey^'
l\dt lite cycbull may bo rligblly |milruiJctl ; but tins iriSine lowniniion timy
be n«Klcct4Ml. By (IImiuc, howirvcr, th« position of the eyvuull in thv auckel
DiNV tK niat«rinlly cbaomd.
Emcii eyebnll is opable of routing rount) wi immobile crntre of rotAlion,
which line been fuiiiKl to i>r< placed a little (1.77 nimj Wliind the centre of
the ere; but the tnoveiui'iils of the oy« round tbo centre are limited in a
C'culinr way. The shonlder- joint is uImi a Iwll-and'foqkct joint ; and we
DOW that we can not only move tlic arm u|i and down round a borixuntal
ftxw paaeiDK through the centre of rotation uf tbo bend of the humctiiit, and
from «id« to ndo round a vertical axis, but w« can abo rotate it round its
own longitudinal axi«. When, however, we come to examine clo^ly the
tuoTemenU oftbc eyeball we lind ibut tliougb wo can move it up an^l down
round a horiionlal axiti, aa when with tiied bea<) we direct our viaion to
the heavens or to the ui^untl. and fmiit side to side, as when we look to lefl
or ri);ht, stul tbuuub by combining iheee two movemeota we cau give the
eyeball a rariely iil inclinalluns, vtv cannot, by a voluntary effort, rotate tlie
eyeball rouiid its lougitudinal visual axia. The urraiigeuieui of tlie mutclet
itf the eyeball will permit of such a
tuovemeut, but we ciinuut by any direct
effort uf will bring it about by itMlil
la certain rauveucnl.-i of the eye, rota-
tion uf the eyeball doea take place ; an<l
by brineiog al)i>ut tl)«<e movement, we
can imiiroctly cauw- mtnlion ; but wc
cannot rotate tlw eynhnll except thuti
iiidirvclly Uf a part of these movement:^
If, wh<-R vision in directed to any ob-
je<'t, the head be mnvei) from eido to
siile, tile «yc« dn not mi.ive with it ; ibcv
app<-nr to reimiin stationary, very much
as the needle of asbip'scompnsa remains
Blationary when tbe head of the ship is
turned. Tbc change in the pOBilionof
the viMial axes to which the moveuenl
of the head would naturally ^-ive rise is
met by compeosatiD)' moveiuvnis of the
eyeballs; were It not mi, itteadineas of
TisioD would be iaipuatilhle.
There is one posiiiuu of tbe eyes whid)
ha* been called tbe firimarg jiafilioti. It
oorrcaponds to that which may be at-
tained by looking at the dlntaot borixoii
with the head vertical and tliv body up-
right ; but ltd exact deterniiiiution re-
i]utrea sjieciul prc^-autious. The visual
axes are then jmralkd to i,iu:b other and
to tla- meiliiui J'Iiuh: of the iH-ad. All other pcwitiviu of tho eyea are called
teoondatTi ftotUioiu.
U*mim «/ lAc tgtiaU. Hie eyeball b mover) by six muscles, the rttti
tM^tnOf,*mtrittr,iuiernu*,mDd exleruiu, and the oltliftii infr-rior and niperwr.
It b fixilM by calculation from the attachments and din-dioMH uf tli« mus*
clei'. ajid Ooufirmcd by actual obtervatioii, that the six muscJeri may bo con*
sidurcd u three pain, each pair rotating the eye round u particuUu- axis.
Mi
IMtnaiH cor ra* ATTucacm nr na
UlHLI* ur THE SYC tHS OT THDM Am
or lUir^iluK : thg t*iwr totBH npramiMd
b>- duitdl lla«L
Til* Kill llT nULiIII .if III* MOIU* PlMIDUt
and inHmuii. b«in( |cr|wndluulBr (o tht
pUnuff lhc|wp«. atnnotMihawn. (AMcr
Fiisi
m
SIOHT.
The rclaiive nitnchiueuts aci] ttie nxta of rolalMi are diflgmuimduiDT
vlwiwii ill Fig. 236. The reclus Miperinr and ihe reciiis inferior rotate tlit
eye nxini) ali»ri£nntu1 axid, uliich isdirected from the upper end uf ihf dok
tr> ilic icnipIiT; the olitiijuti^ Mi|iennr aod<il>1ii|uue inferior rL>u ml a liofitooul
nxixlin'i-U'd rriiin the renire i<f tbe eyvlmll luihe occiput : and tli« rectm lo-
t«niUK and recttiH cxUTiiiin rfiiind n vt-rlical axis (uhich, being at right aa^kl
lo iW plants i;fthc paiifr. ninnoL be »lionu in llie diaKram). [MHini; thrati^
lb« reiitrc ')f riitiiiinn nt'llic evrbiill |uinillcl (o ihe meilian plane of ttin hiM
when (lie liciid i.-> vrrlinil. Thus the Inller pair aclin;; uluoe woiilil lurn
lh« eye fn>m itide l» r-ide. the other *trnight jwir iivtinf; alone would tanft
Ihe oii'o up and down, whili^ the tibli'iuo niiudea nrliug alone wuuM pn
Ui« eye an obli'jue movement. The rcctiiv exteniitfi acting alooe wouM
turn (be eye lo ihe miilnr side, the internuK to the »aMl »de, the mOui
eupcnor upward, the rectng interior downwunl. the obliijue Miperiordnvv-
ward and mitnanl. and the inferior tipwnrd and outward. Tbe rrcti ftiperiit
ntid inferior in nioviDgtheeycnpnnddowiialtu^iiirn iiKiiuenhat tnvratdud
al the some tjme give it a *lieht amonnl of rotation ; Ixit thin i.-i corrcoH
if the obliijiie niiiteK-« net »t ine Mime tinK< ; and it in found ibiit lh« rectiu
enperior ncling with the obliquus interior move* tbe eye upward, and tkt
rectus inlerior with the obliiiuiis superior ilownwnrd in a verticnl directina
Iti oblique movements also, the obliqui are ulweyo ataumietl with tW
reeti. Hence tbe various muvctDenis of the cyolwll may bv nmnged ai
follows :
E3cTation,
iMprcaaton,
Adduction to
nasal side.
Adductiuii to
maUr side.
Elevation with
adduetion.
Depreidion
nitb adduction.
KleTAtion with
■brluetiun.
Deprextloii
wilb ahdnction.
Reetui •u|>erior and nbliquu* Inferior.
Keelua interior and obliquus lajreriur.
ileetu* iQteniu*.
Elect lu extern us.
Kcctiut superior and internns with obliiiuu*
inferior.
Rectus la ferhtf and internns wilb obli<|Uus
superior,
Iteelus RUperiof and cxivrous with oblii|uui
inl'eriur.
BectiiH inferior and exiernus with oblI(]Uu«
superior.
Oidrrfinotton of vimal movetumlt. Thus «t«i in the moT«nienb uf i
aingleeycn eonsidcmble amount of coordination takes place. When tits
eye is movml in iinr uilier than the verlu-ul and horiznninl meridiaai, iw
pnlse? must descend t<> at least tbrcv (uusc[<«, and in »u<'h relative energy to
eaeh of the three as to product) tbe rc|iiired invlinalirm of the visual aim.
Uut the cDiirdiniition unevrvfl in binocular vision is more striking still. If
the movements of i>ny {lereon's eyes be wacehe'l it will be aeva llint the Iwv
eyes move alike. If ihe right eye moves to the ri(;hl, «> ili»s alwi the UJl :
and, if tbe object loukctl at Iw a disUint one, exactly to tlie aume extent; U
the right eye looks up, the tefk eye hxiks up also. aiKl H in e^^-ry iitbur
direction, very lew pereuns nie able by a direct eHort of the will ui taitn
one eye independently of the otber ; though some, and iinioiiji; them one ills-
linguibbed both as a pbysiftlogisi and an oculist, have ncttuireil Ihu po«4f.
lu fact, the moventeiils of the two e^'ea are so arran^d that in the variMi
nioveineutB the images of any object »liould fall on the i.MrrfKpiii)dirtg [miaU
uf tbe two reiiiuu, mud that thus sin^^le viniou should reitili. We isnMit ht
anr direct efK>rt of our will nlaee our eyex in nii'b a {Muiiion that the ran lif
liglil pniceetling from any object hball he bniu^lit lo a focua on parts ut tb
BirroccLAR visros.
I
L
two rMmu which <Iu not cornupnii(), and thii^ i;ive rise to twa <lmini>t
visunl inuges. \\> cna brin^: the visual kxea of the Iwu eyea fraoi ft conili-
lion uT jHinllelieni to ooe of great convereeDoe, but we cttonol, without
special aMtslance, bnn)[ them froin a ooudition of pnrallelisiu to one of
divergent. Tlie iierei»oa|>e will enable us to create a diverKencft If in a
il«rcoei»|HO picture the diAiance Iwlween ihe plattirea be invroaaed verjr
gradaally so a* carefully lo luiiiatain the impreMioii of a aiDgle obieot, the
risual ax«a may be bmuithi to diverge- (^imitarly if a distant object be
loultcd at n itb a prism hi-fore one eye, and the tmajre of ibe object be kept
mrrfiilly HinKle, while the pridu ia turned ven' !)l»wlr up or down, tbeni on
suihlenly reiuovinj^ the pri^m n duuhl« iniajie id fur a luooieDt aeon ; ahow-
inK thai llieeye before whi<-h ihe prium vnt placed bad moved in dUacnord-
anoe with the <ilhcr. Tli« d>iul>le ima^', hoirever, lu a fvw oocond* af\tii- lh«
reitiiival of the pri»tu beoomat klnglu, un aoouuni of llie eyca omiiif; into
a4.-c»n lance.
It ia <>nlv wliri) luM of oourilination ooRur*, n* in varioiia diMaaea and in
alcnhulic or nihrr (H)iw>Riojit, that the miivctnail* of the two cv«t oeaM lo
agrew wiUi uu'h iilhrr. It in cvidtint, th«n, tlial wh«ii wv lonk at an itlijtx't
(o lli« right, WQM) wf thereby ahdtiot lh« right oyc and ailduct ibo l«ft. we
tlirnw into actiim the rvctus cxicriiun of tlie rieht eye and the r«cluii int«rniu
of the Icit, and vimilxrly whoi vc look to the left wo titc tli« roctna eslcrnua
of the left and the rectus iiitcniiis of the right ore. On the other baml,
when we look at a near (i<>ji:i't, and therefore converge the visual axes, wc
U*e the rrvti intenii of huth eye* ; and wht'ii wc look at a diiilant ohje<;t, and
bring Ihe »xct from (vuivrr^cnce tiiwanl pninllelisai, we »tv the recti esterni
of b-itb eyoB. In the varimis muvPiiiGiils of thv< erv there is therefora, »o to
•peak, the moit delicalp picking and choasing of t)l« muscular instrunienu.
Buriug this in mind, it cannot be wondered at that the variuus niovementa
of the eye are dejiendeiit fur their cnuMttion uu visual senaatiims. lu ordur
to BiAve our eyiv we muj^i either hMk at or fur an otjeot : when we wish to
oonTOge our tix-^ ne l'H)k nt some near object, real or iiiiaKiiinry. and the
cnnvugence uf ibi- axi.-« i» iisuatly accompanied by all the ivitidiliuns nf near
Tisioo,auvb n incrautci] nccumnwdation and wntraciiun of the pupil. And
Mk with other loovemrnts. The doae auocialion of the movcroenls of the
eye may tie illtictrnl<H) by the fidlt)win); au»: RuppuM the eyca, to Mart with,
i|ir<Ht(^) fi>r thi; far diDtancr. and thai it is deaired lo direct attention in a
nearer iHiint lying in the vi*i:nl linr of the right eye. In ihia case no move-
toeiit of the right eye is roi|uircd ; all that is neoeatary is for the IcH tyv lo
he turoeil U> the right, that is, for the rectus internua of ihu left eye to lio
thrown into acii^m. But in onlinary motrcnicnta the contradinn of this
miiacte in always associated with eitlier the recttis eitornu* of tlic right t>ye,
as whm both eyes arv Uimcd to the right, or the roclns internus of tluil tw,
aa in i-onvvr|;on<x ; tltc muscle is (piitc unacciittomed to act nluno. This
would lead oa to suppose ihnt in the civv in luoslion the cinlractifin of the
reclua inlamus of tne lel\ eye is uecKOtpunied by a otMitmction of both recti
exiernus and inlernua of the right eye, keeping that eve in lateral i-(|i)i>
librium. And the peculiar oscillating movements seen in the right eye, as
well at the seoM of efllirta in the right eye which ia felt by the person, show
ibia to he the case.
Hueh a coinples CHirdinaiiou re(|uirea for its carrying nut a distinct
nervous wHcliiuFry,and wc have rea^tons for thinking that such a machinery
esitU in certain purls of the corpora ipiadrigeniina or in the underlying
structurm. In the nate* there apttcani lo be a cumnion eentie for both evos,
stiuiulation of the right side prwiucing niovemenis of both eves to ibe ieii,
•if the led side luavemeiits to (be right ; while sliiuulaljoa iu tlie middle lino
SS4
sianT.
Iwhiod cauMa s dowmiruni iiiovement of bold eyea with conv«rKen<w of ilit
flxe«. sill) in Ibe fmnt nil unwiird nvovciKiil nitb return to parol Id irai, hotk
acouiiipauied by lh« iintuntlly awi^ctaletl inoreitients tit the paf^il. SiimaW
tioD of variouH paru uf the mats muH^ti rnrioua ntaveineols. <lepei>Hin|: an
tlie ]>08ilioD of ibe »prii ttiraulnted. AAt-r nu inciuon in the middle ItM,
BtimulatMHi of Ibe ncrvi^uN centra ou one ude producea iDorenMinl* io tW
ey« of the aame aide ou}y.
Ttif Horopter,
When we look nt nny object wo direct to it the vmwA kxv*. t« tliat wtini
the object is stnnll, tbt? " corr««ni>nding" pnm of the two rrriiiiwi. nn «hi«l
\hf two itniigos of the ubjcct fill), lie in their mpective fo7(w ccntrnln. But
«bile we nrv looking iit the pnTticiiUr object, the images of other ntjeeit
surrounding it fall nn ibe retina ma-
no- W. rounding the fov««, and tbu« go Xu tarn
e nhat i« cnllcd indirect vision. And il ii
ohviougly of advantage that thefe ima/n
also fihontd fall on " cor res pond tag" |«rti
in the two eyea. Now for any given pea
tion of ihe eyee there exisls in the field tf
vision n certain line or aurfnce of sucka
kind that ibe inmgex of the |xiint8 in it
all fall ou correfip-mding jwiint* of the re-
tina. A liiio or siirfuct' having ilib prop-
erly is ea1le<l a horopter. The hofvpitr
ia, in fact, Ihe aggregate »f all thoM' jxitDli
in Kpac- nhicb arc projevUil ita tv com-
aponding |>otntii of the nrtina; beoM ill
deieniiiitntion in nny pnrlicttlnr ean it
ainiply B matter of geomi-lrical calculi-
lion. In wMiiQ incUinces it becnn»C8 a rar;
oomplrenicd figure. The rnM- wh<«a ftai-
lire* arc m™t «»ily gmsped ia a cifdt
drawn in llic jiliincof the Itro vistial <i«
tbrough Ihe point uf the convergence *t
the nxen luid (he optic centres of ibe !■•
eye«. tt k obvinn* from geometrical rchtlions that in Fig. '237 the iaifa
of any point in the circle will fall on correeponding [H>jni« ot the two r^
liua^ When n*e Mnnd upright mid Imik at the dislanl horixon, the horopitr
is (appraxininiely, for nnrmni eminetropic evce) a plane drawn ihroufb Mr
leei, that ia to say, ia the ground on wbieh we stand ; ibe advantage of Oil
tt Qbrioii&
VtSUAL JviH)MEyT&
Binocular vision is of use to us, inasmuch ns the one eye is able to till iif
ibe gitp und imjierfeclionii of tbe other. Fur example, over and abuvo lb
monocular filling up of the blind spot, of which we spfike on pan 9S^,
since the two blind xpota of the two eyes, being each on ibe nanal nA§, irr
not "correnpondinu:' part*, the one eye suppliM thiU part of th« Caitrf
vision which i» lacking in tbe olbrr. And olber im|MTfecti»n« ara atmilarh
mode good. Rut ihn great uw of liiixiciilur viiinu t» to nlTiiF>l us iBcaa*
of forming visual jui^menia oonciTtiiiig the form, mec, and dtstance «f
bbjecis.
nUCllUI ILIIvruTDM A 8111 ri.i
tloMuma-
Wlran tli»TUu>l US nHnvisaiil C. Ihe
nimgn 0. a of anr |otnl A t'U Ibc i^lrvlc
dnoii IbriMiiib C mid ihu a|<Ucal cvatna
1. 1 «1I1 flill on a>iiT*|niulInK p'4n».
vtscAL 3t:D0ue:;Ts.
»83
I
I
Judgment of diManee ami tite. Th« pempiiont which «r« gain ■imply
Mtd tMt\y by our lield of virion concern two dimensions only. We ciin
berontc aware of the apparent site of any part of the tield cormpoO'ling tn
anv particular object, and of its topofiraphicnl reletiona to the re«t of the
field, but DO more. Had we nolhtn); more to depend on, our Btght woiild be
aJmost valueleaa aa far as aiiy exact infornmtton of lh« eztenial world wai
ouneemed. By aaaociation of the visual Bensations with Mnntions nf loiioh,
and with scusiiliouf derived fr<>m the movementa of ihe eyeballs rei'ilred to
make any such part uf the field as corrceponda to a [larlictilar object dis-
tinct, we are led to form ju(l;;menls. t. r. (o draw conchiatriDs concernini; the
esletoal w<>rli) by meona of an inlerprctalioo of our visual perce|itionB.
Lfwkin;; Itef-ire u». we say we see a cennin object nf a certain cnW neArty
in fn>ul of us. or much un our ri);ht hand or much od our left ; (hat ia to
aay, nc juilp.- such an object tii he in such a ptisition because fwm the con-
■Ulutiuu of our brnin, str«U]i:thcue<I by ull our ex|)erience, we associate such
a pan of our field of viBion with such an <)bje<-t. The subjective visual
coRi|ilcx a^nMUon or )<er«.-cj)lion is to us a syinbul of (he external object.
K\'ea with one eyo we caD, to a certain extent, form a judf;ment. n'>t only
aa tu the position of the object iu a plane at rifcht angles to our visual axis,
hut almi an to its dixtuncc fn>in us nlont; tlte visual axis. If the object is
near to UK, we have to acc'iuimoilate for near visioa ; if far from us, to rvlnx
Mir acoum mild nti»n inechnni«m so that the ere becoroei mliuHled f»r ilistance.
The mtitcular meivH: (iif which we shall »\)t:tM preaently) of this L-lTort enables
»s lu form a judj^mcnl whi.>thi:r the object is far or near. Se<-int; the narrow
twige uf our acoommcxlalioii. and the- ^li|{ht muscular etforl wliicb it vntaiU,
kll nonocnlar judi;meul* of diiitanoi! must b<^ lulijuct t» much crmr. ?^vvry*
one who has tried to threnil n neeilli; without unin;^ both oyc«, knows how
grmt these errors may be. When, on the other hnnd, wc use two eyea, we
have still llie vanalions in aci^ommoilntion. and, in addition, have all the
ai><>i»la[ice which arises from ihe muscular effort of so diroctinf; the two eyes
on tlie object that single vision «hall reuilt. When the object i^ near, we
cimverge our visual axes; when distant, we brinj; them hack tuwar>l jwral-
lelisni. Thin necessary contraction of the ocular muscles affords a muscular
sense, by the help of which we form a judgment us to ihe distance of the
object- Hence, when by any means the convergence which lb aeoeaaary to
bnog the object into single vision is lessened, the object seems to beoone
roore ilisiant, when increased, to move toward us: as may be seen in the
■lereoaouite.
The jiM^meut of siie ia closely connected with that of distance. Our |)«r>
ce|itioiu, ([aiiied exclusively fK>m the Held of vision, f^o no further than the
apparent late of the ima^, i.a, of the angle subtended by the uliji-ct. The
real sixe of the object can only be gathered from th« apitarfut Mf uf the
Image when the distance of the object from the eye is known. Thus per-
crivin^ directly the a|>iiun:nt niw of the lniB}(e, we jud^ the di*lanc>« of the
object ^'iving the image, and upon that oome to t conclusion as lo its size.
Ami, cxinvcriwlr, when we sec an object, of whose real Hi7.c wc nrr uttuii aim
aware, or arc led to think wc are aware, our judgnx-ut of its ilisl.ioce it
inSuCDC^ by its apimrcnl sixc. Thus when in our ticid of vision there ap'
pears the inuigc of a mim, kniiwint; otlicrwtse the ordinary siw of a man, we
inftr, if the imagv be \nrv small, tlmt the man is far off. The reason of the
innge being small may W Wcnuoc the man is far off, in which case nor
judgmcnl is correct ; it may bo, however, bocJiuan the image has been lee>
sonrd by nrti6eial dioptric means, as when the man is lookeil at through an
invcrlcti telescope, in which case our judgment becomes a deluaion. Ho nbo
an image on aecreeo when gradually enlarged swins lo cooie forward, nlien
9S9
SIOQT.
eraduallr (liiniuiitlicd, Monu to recede. In Umm omh tbe inflnwio* oa *Kr
jwJgin«D[ of tbc iiiuRciilnr hcmo at biuooular adjiMinoot. or Bwnooalir
accnmnkodstiun, n ihwariid hy the man ilinx'l iiillucnoe of ibe iwofiiittin
belirocD rite kdcI (ttXiiDcv.
Judfftiifnt oj ntliiilty. Wbon wo look at n siunll cjrc'o, ill porU of tW
cirole aro nt the tamv ilislaooc from »*, all pjirta are e<|[iul)y di»liticE al \i»
tamo time, vbctbor ne luok at it will) ono ojrcor with Ihd eyes. VVlico.aa
the oilier hnod, wu I<H>k n( a. sfilicro, tbo rariuuB pnrl« of wliich &r« al dUEn-
eiit <li»lanciit frum u*, n tvmie of the aocoiniiiodatiun, but much more a taut
of Lh« binocular a<^iiatfDeiit. of the coDvei^eiwe or the oppoaito of tbe t«fl
ei-o, required tomalte Uie various parta«uceewivelyd)»tii)ct, makes uaawm
that thi^ TftriouB parte of the s|>liere are uue<]uall_v <li»laoi ; aod froni that «r
forai upudgnicnt of ita solidity. As with diatatic-e of ohjeoU, so wilb toltd-
ily, which is nt bottom a matter of diaUuioe of the parts of an object. ««
CM) form a judgment with one eye atone; but our ideas beooioe niucti man
exact and trustworthy when two eyes are use*]. And we are much
by iho ell«4.'is produced by the redeotion of li^ht from the various mi
of a solid object ; eo much so, tlwt raised surfaces may be made to a[ __
depressed, or tiiK t<n-«>i, and tlat surfaces either raised ur depressed, by
proprijiie arraD^Dieuts of shadings aud shadow.
no. SM.
B
f^^c"^
\j
/5x
R
BioM^ular vUion, iimreoTer, aflbrd* us a meant «f judxios of thir aiiliditi
of obj<.-ct«, ina-MDuuh iin the image of any wjlid ubji-ct which falls oil lo tbi
right eye cannot he csiu'tly like Oint which iklls oo iho lelV, tJ>ough both in
coiiibiDcrl in (be sindo purccption of the two eyes. Thus, when wc took M
a tniucated pyramid placeil in tlie middle line before u», the iroAM wkidi
falls on the right eye is »f Iho kind repremited in Fig. '£Mi, R, white that
which falls on Iho lefV cyo has the form of Fig. 2^^, h; yet the peraeptita
{^ined fn>m the two inin^s tngclhor C(inwpciDdi< to the form of wbidi Fifr
',!38. It. is the projcclinn. Whenever wc thus oomhiiM* in one perceptiou tsa
disfiimitar iniiigirx. one nf the one, and tho otlier of tlie otlier eye, we jadp
that tlie object t:iviiig riMt to the images i* solid.
This is the simple principli- nf the sicreoscope. in which two sligbtJy dls
simihir pictures, such as would correspond to the vision of each eye stp-
«»taly, »tv, by meisns of rvAecling mirn>rs, as in \Vhe«t»U>[ie*s urMinal
tiwtrmDent, or by prisms, as in the ftrni introduced by Brewster, made M
coat images on. corresponding pnrLi of the two retinas, so ss t>i produDS ■
Stable perception. Though each picture is a surface of two diinea^iiiiiis uslr,
tbe resulting perception is ibe eaciie ns if a single object, or group of objccu.
of three dimeusions had been looked at.
It might be 6Up|>osed that the judgmeut of solidity which arises vrboa iv*
diasimilttr images are thus combined iu one perception, was due to tbs fact
thai all purls of tbe two ima^ cannot fall on correspond ioR parts of iW
two mtiiuis at the same time, and that therefore the combination uf iIm in
needi) mmie niovenieut of ilie eyes. Thus, if we superimpose K on L {Fit
2S8j, it it flviilent that wheu the bases ouiuvide tlie imucated ajiioea viill uod
MM rena; ht^iioc, whrn tba biwa* fiill rni oormpoDdiii}: [wirts, ihv ti|M«a
will Dot be C'lmbiiiMl in <>av imsf^, kn<l i''<v ivrrM ; id order thiU boUi inH)*
b« cotnliiiK'f}, tlicrv niuit bp n slight, rapid Bi'ivrmciit of the cvm from ibo
OM lo the uthiT. That, hnncvrr, ni) Kiich movumMit ia aetxmstry for -itdi
portieuiarwe is •Itowii by the fad that ^1 id obj<-(?C« itppMrna iuch irbai
illuminatnl bj >n eledric ipprk, tho diirulinii of which h ton short to iwrmil
of any n)ovenenl4 of the <>rM. If rhe fln«h occurred at the mnmcnt (hat
lb» •yen «r«re hinocuiarly Biljustcd for the bases of thp |>yrnmid». tho two
apion ni)t fnllinj; on exadlr comspn^Hlin;; piirts noulil f>ive rise to two
pom^pliona. and thp whole object ouuht to aTtpear caiifuseil. That it do«e
ant, but, on the cnntrary. appears a single mlU). must be the result of cere-
bral operations, res'dliiis in what we have called a ji>d)>meai.
Stmg'jU of th( two fiti'U of vitioH. If the iioajte^ of two surfaces, one
btMik and the other white, are made lo fall on eorrespoudins paitsxf the
•y«,m as to be united iulo a Bincle perceirtion, the result it not always a
mixtnre of the two inipreMinnB. that ix. a unir, but, io many cases, a seosa-
tion similar to that pr^xliiced when a piilished Hurface, auch m |fluiu)>as^, is
)ook«l at: the surface uppean brillianl. The reasun, probablr, is Iwcau'e
when ire l<><>k at a p<>lishe<l surlaci?. the aiiiniint of reflected li;;ht which falls
upon the relitm n Keuerally iliirefKnt iu Uie two eyes ; aiiil hence we anociilte
an une(|ual stimiilalion of the two retina:) with the idea of a jmlished sur-
face. Si), aU). when the iiiipruAiiiin.t of twi> c»lorfl are united in binricular
%-isioD, the rMult it, in most casea, not a mixture of the two eolont, as when
th* nmc tno impressions are brought to bear toi^etber nl the name time on
a single retiita. but a ilruKKl*^ between the two culon, now one, and now the
other, becomiii)t prominent, iiili-rmediate tintu, howerer, bein^ frei|uently
pasaed ihrouirh. Thiit niayariM- fn^m the difficulty of uccommoilatinK at thv
Mtme time fur the two ililTcrent »:>lunt fsoe {t. 907 1 : if two cyn, one uf wliinh
ii looking at red, and the other at blue. I>c both «c«omniod>lcd f<>r rvd rays,
tbt red KiiMition will overpower the blue, and Ww i-crra. It may be how-
pTcr, that \hv tonilciicy to rhythmic action, so munifcsl in olhvr nimpler
mauifnttalinns of protoplasmic aclivitv, niakoa its »{)|>caraacc als^ in thu
higher cvrebral l&bora of binocular vision.
The Pbotectew Mix-SAStsua of toe Ete.
Tlie eyelmll ia protecteid by the eyelids, which are capable of moTement*
called rrsjiectively opening and shuttin;; the eye. The eye is shut by the
contraction of the orbicularis muscle, carrinl out either as a reflex or volun-
tary act, by raeans of the faciid nerve. The eye is opened chiefly by tbe
raising of tbe upjwr eyelid, tliroufib the contracliun of the levator puljiebne
carried out by meana of llie thir«l nerve. The up|>er eyelid ia also raised
and tbe lower depressed, (he eye heiu^ thus o|M>uc-d, by means of plain mus-
cular flbreii •.•sisiinif in the two eyelids and ic>iverned by the cervical sympa-
thetic- Tlie shuttinn of the eye as in winking >* in general eflected more
rapidly than the ii|)pntUK.
The eye i> ki-|it continually mnkit partly by the necrotion of the gland* in
the conjunctiva, and of the Meibomian t;laod.i, but chiefly by tbe scorvtioii
of llw lachrymal gland, fndcr unlinary cirfuru-itanct* tlx* fluid thun lunued
is carrietl away by thi- Inohrviuul cniDiU inln ibr noMil ><ac and thus into tlie
(■avily of the n>iH!. Whi'u tht- M>i.'n.'lii<n liecDnHs to<> ahunilant to eacape in
this way it i<vt.-rlli>ws mi In thi' i*lm'k:> in the form of ii-am.
If a i(uun(ily uf ii^ir- 1 1 i i' d, thcv arc found lo liirni a clear, faintly
alkaline fluid, in mnit) tl^i-.^j;.? itko tuliva. conlaiDing alwtit 1 pvr eyau of
eiOHT.
aniids, oC wliiirh a small {lart m proteiil In nmturc. Among Ute mIu pnccat
eodium chloride is coDSpicuou^.
Tlie nervDua nicchanisni ol' the wcretion of tWB, in many respeoU. mnn*
Mea llint uC the aecKlion of BalivA. A flow is usually bronchi about <*thft
in a rvHex iiiaDner by sliiiiuli applied to the conjunctiva, ibe iiitasi mncOM
metntnune, toti^-ue, tiptic nerve, etc., or mora direallv bv eiii<>li->nB. VewMW
OODgeuion of itie heait ia alao said to came u flow. Tlie •^Iferenl uervc* beloac
ftilher to the oerebr&^ioal aysteo) (the laohrymal and orbital brsneW m
the fifth nerve) or arise ftuni the cervical ftympatlielic, the aflureni aenrn
varviuj; aofi^rdirii; to the enciliiii; caii§e,
The avt of blinking undi^ubteilly fuvtirs the pttuotfe of team thmiiith tb«
laohrymal cunaU into the nnsiil hac. and hence when the orhirulari* ii
jMUalyiwd tvani do not pa^ ao r«a(ti<y ad uaual into the oo«e ; but iliv veacI
inecbaniHui by whbb this is ettboted has been inucJi duputod. Acmnliaf
ta tome auihont, the conlraction of the urbicularis pre«e« th<i fluid onward
out of the canals which u|H>n the relaxation of the orbiouliirin, iltlataaaj
r«!ceiv« a fmh i)UBnl!ty. Othera maintain that a apecial arranifrini^nt i>f miw-
cular fibrUH keupi the eauaU open even durinii the chuingnf the ItiI^.iui lliil
the premiin; of the omtractl'm of the orbluularia U aUe to haw lull eAea ia
drlviDg the teur* thniugh (he oauaU.
CHAPTER IV.
IIEARINO, SUELL, AND TASTE.
Uearino.
[Phytio(ag'u)al Anatomtj o/lhe Kar.
TiiReur.orofg&Dof liekriug, iflc>iiipooeilorthree j)arU,cnlIeii tho nrftfnur/,
Mtiidle, at internal ear.
TtM aUmiU ear ooiwbt* of an oulfir |>n)jec(iii{( porliim, culled thv pinnn,
mntl tlw HudltorT' cmiul, or nttatii* auditoriu* ejierntu. (Fig. 2.t5l.) Tlic
|>iiitiH !■ a »oii)cnh«t ubiong fHun«l-«biiiiMi orgau, tbe ttDinllcr portioo of lh«
Flo. at.
f.V4
;i->^-V
fAiii
:«,•■
Vumcjti aWTiOH or n* llR«Tt« AVBROmn jNb Trurii'iTa. (9urpi.|
o, cofSlavliKiai p«rt of tbe mtuim : i. owtow ti*n>DO ; <*. maulmBa iini«nl i d, nrtv of the
■rmpAunio ; t, Ewleelilaa tub*.
funii«l beini; attaohed to tbe alciill by li):aiiii>iit»iiH tivue, l)ie larger portion
Bervin^ lo wlleot Utd CODVey lb« wimmius iimlulmtiunfi to tbe niVHlus. It is
composed of cartilsfie covered by iuieftuaittit. It* iturface is irre^larly
ctirvnt aod d«|>reaaed. The outvr nr(>)«<:tiii^ ritii i» lh« hetix : anterior to
the helit ia a tecood elevaiiuii. railed ib« antihtUx, wbirh dmcribea a curve
pnnialljr around a <Iee)> tleiircMioD wliicb IcwU to ilie tnealus, called Ibe
coneAii. Between the Iielii luid aiilthelix m thf/ounorike helix. Tlie anti-
helix biAircateaat iissui>eri«r portion, and eitdoK* tbe/iMm ofthe aulibelix.
ProjectiiiK [xutienurly fnun the sntrrior portion of the ciuicha it n papillary
pmmincix^ called thi^ tragn*: ^luiterior to ibin, scparut<Hl by n fiaitirv, ia ibo
anlilmffut, which i» a lymtiiiuiiiioii of the helix. Op the inferior portion of
010
HEARIKG, 3HELI., A:fD TJlSTS.
tbe piniw in m «ofl prmliiluu* iinrtioii, tenoed the lobute. Th« niMlai
leada trom Iho coacba to the mtilillc utr, from which it is aepxrai^ Hr •>'
Ijmfno'ic membrane. lUi direction iit forwnrJ. iowtrd, and slighdv u|i' .
]u Inircr turfiiM being loogor thnn the upper, oa socount of tM oblii|uiu "i
thi; puMitiun [>r thfl tympsDic mcmlirune. The canal considta of »a ezUnul
mevthmiM earlUnginmi* porliou, whicli in contiuuoua wilb llic piuna, >n>l *o
liilvnial OMCWM pnrtion funned by the ma-iluid bone- In ibe etwmni p-ir-
tiiiii of the cnnnl arc found nuiiivn>ii» bain nod aebuceoua (ctaarb: in ibt
iiitcrnnl portiiin iire found the ftratuHotu glantU, vbioh Becrete ft jwcului
KiilMtniico commanly knono ns lfa« carwax.
no. 240.
*f-1
i^l
7Ut. 311— IKNXII ViKV or Till UCKDMllll
Ti-MTAXi a ran P-mx vmi nil MtLLEn
ATTjiCHtn. n. iiinnliniiivurilruin ■<( ihalfnt-
|«Duni; A. nullciia: e, t«uil orelmuUi llhrci
■(tb«olrcuiD(^nTnci!:il,liif«r1or.<uidr.fU|«riur
l]nii|asi« m.ntrj : /, trmcanlo boo*.
Fw, MU-PUX nr THt OMCiai nc remnem m TKiTvMi'ixt:ii.«int tntik MoiFtM.
of lAa ijiiipuiain: A, iiMiabmii& trwioni. '■ EutUiliUn l«bc: < BMllrta; r. limii/iiipM;]
Ujouw tjmiiiaiiliDiuol*; A. teiinnrtyuiiAiil: 1. •upedhu.
The niidiiU Mr or tymfritiian \a an irre^lur tlnit«ned cavity, aiitiotnl ta
the petrous portion of the teuporal bone, and lined irith a mucous iseRi-
bran«. It b «ej>arate<I from ibe meatiu by a ineiuhratiuui diaphrB|[m, which
H the ti/mfinnic menUifane ; uud from tbu iiil«(tiat ear by an oaseo-membranoB*
partition, nbich turnis a eommim wall fur Wh. Tbroufcb tJK Kugtachiai
lube it oominunicales with tbe pharynx. On iu poaterior iraii ars Man
orifices of the inaaluid ceUn. The tympanic membrane is a setnitnuuiKimU.
oval nicmbranr, cimcave uu it* exlrnial, nnd cmvex on ita internal surface,
where it hiLi ntliiohed tbe long iiroccw of (he iiiulluuit, one uf the oosielw. It
is plaotKl tti an ubliipii; prwilion, stuping dowunard, forward, and inwanl iH
•n angle »f nl>(>ut 4o degrot*. Its circiimforoooe U altadicd (n a gruiitre in
Ibe temporal bono. In the fietii* tlii« iiortinti of thr bone exiHia aa a oepant*
piece, callud ibef^mp'intc 6on« (Fig. 240>. but it alWivar<l becomes omfisd
^< tbe temporal. The tympanic mcmhrano oonibta of three layers — lh(
cxlcnuil, middle, and internal. The external » a continiialJoa uf ibe in-
tvgiimvnt covering the mentns : th*; internjil is a cimlinuuliun of the miwou*
membrane lining the lympaniiTu ; the miildl* layer, which is the bust
importani, is tense, slrong. and tibrous, made up of circular and nuliatiag
fibnM, nith a small amount of clastic tissue intcmtixeil.
In the int«riiat wall of the tympanum are tnrismiill o|>oiiinga— Uie /r A'rfni
»ni/M Mad Jenetira rottuw/u — which communicate wilh the labyrintb. Tb~
pnrsioLomcAL akatouy or the ear.
9-11
f«nr*lnif«tiindii8clo«d by a mcfobniue^ ExUndinj; belwc«n ih" (vnipnnic
fti«-mbrann boiI the feneMra ovitlis nr« the omMm. ammtwii, at ihrc« small
bona, which Conn b eyeteni of levcre, nicse reticles are lerntCfl, t'njm Ihrir
rtMnbUiioe lo particular objects, ihv malltut, iueiu, aod tUipe§. (Fie, :;41.J
Tbn lualli-iia cnneieu of a head, iipck. lotip and abort ]>rocCM, and Mudle.
Tln^ hi-a-l aniculateB wlUi tho nmf of the ijinpanum, aw) in a di-premion of
iha iiH-ua ; ll>e linndle i* direcltd downward and altachcil by iti> whole Icii^-lh
to the trmpaiiio niembranc ; tb« long procas ( pnce»nu rjTnnli»\ » dir^i'led
furwam and has atlai-h^I the insertion of the lazator tympani mutrlt : tlie
»h»rt proceas, which is at llie bwe of tbe lonjt !>««»»», has BtUK-hwi the In-
■wrtioD of ibe Unwrtumfi'ini muivlf. The immi* coDeiata of a body, a long
and ibort procesa. Thu budy of the incus has a depmeton in which anicu-
laivB the bead of the mallcut; the thorl )>rocew is ariachvd to the |ioeieriur
t'l'^ 'Jt2.
Fiiksn.
Kainr*! ■)».
('. *wUtiHlo^ or, kiiuilucl lit tha wMlbulei o. lam btnivllii'tir* : r, ftm« ntinlttibtrlra ; A
i^tfcUdilHMiM]*- t.niiwrtut: p. pMcMor: l,harjwn(sl: a.n.a. ■b*iiD|ifiItari>iuciuit]r(rfncb;
C mtlitM -. «*, aqtudiun of iho rooblw : •>. (in*oiu tone or the lamipa >|dnlu. 41iot» nhlch It Uit
•rBltTatlMUmnnrnmlcBiingaiibtbuTntlbuIvi A Ksla ifmisnl bclcnr iha*i4Ml lamlii*.
wall of tbv lympaiium; tbe long proctTa(knl[cular procoa) n placnl almost
vertically, and at its rnd if a rounded proceaa (the ot orhimhrr), which
aniciilalei with llic head of ihc Mi[nt. Ilie fiapn i-oDtislii of a bend, neck,
two crura, and a lin/e. The head nrlicuIait'H with ihe long pron-tn of ihe
incus : lht> neek «cn-fs lis a tioini of iDN-rtioD of the »iarH-iiin» muiclc; llio
cnira divcri'e from the aixK end iinilt' wilh tW ovnl bHM> at il» trreatcct
diameter. The bun- ia 6xtii ia Ihe feneatra ovaliit by attiu'hmenis formed
by the lining nitnibiiiPta of Wb Ibe tytnjuinii in and iulcnial ewr. Tliew
vasiolea arc eonneclcil with eat h other and lo the wnlla of the tymi nnum bv
ligaiDMils, and at iheir artionlattons they are rurnixhi-d with eartiliiges nvA
synovial meinbrnDes. They are cnTelo)>4il by pri>)<'ngatii>ii« of ibe nincous
Btembrane lining tbe tynipnnum.
Tbe iutmiol tar or folryrinlh ia ibe mc«t enenlial portion of the aadilory
api>araiu». Il cunsiaia of three ptirtiona — the irHihtitt, (■rniiciVcufcir rano/f,
and eofhlea — aud U situated within the peln>us portion of the teninornl boue.
Within ihe omcouh labyriuib ia a nienibranona Inbyrinih to wbicn the audi-
tory ncTTc is distributed. The vestibule is un irregular chamber which
M
nSARITTO, SHELL, ANT> TASTB.
ecrvM H a odidihoo mnuis of oaiumunication betvit«ti ihe ivinpuiuiB uJ
tKc twniicirciilu- caiuU nod oodiliM. Oii iti* cxuruul wall u tbe fencttn
ovuli». cIowm) by the bMe of the atJk|>««. On iu intvrDBl wall ia a dqn«MH
ciilird the fitvf'i iKmiff^ierica, which 'm MfornlvKl by i»inut49 upentan for the
Mupagc of uuditur^ iivrve-filitmeiiiM. Abov« nud piMerlor to tliN mpntmrn
u MiotJivr, tlic/o'vn krm'itltipiiea. Poslorior t<> th« fovoa benii»phericB b the
oriGco of tlM> nqiitsluctu* vusUbuIi. In the {Koiterior wall are fire opcnioa
loading to the tcinicirculnr octnalR. Anteriorly, it wmniuuicateB with IM
cochlea by tho npvriurfe tenia veAlihitli taeJiinr^ TUt tenieireular entmU an
three io immbor — l\iK*iiprr'wr,po*U-Tiaror inferior. mjtAhorixQiUaL Thn fam
tbo grtnier purtion of a circle, nnd cotnmiinicHtc with tbe TMlibula oj In
PM.1M.
<ti
f
AKPitDccTiTioii D> thk SmictRiTtt It CiKiu i.MjRiicu. iFiuiu k mndri la Vhtfanlir fijllni
Uiiwum.l
«, (Dpoilor Tonlnl . t^ poaierlor or Inbuiat tvrual : ma t. buriaiiiiu oanal j 4, mmbmb ai*»
lD|uf iliu two tvnliAl r*i'*U: K \*n ul iliu vciUtiuUr favlty;/, opMins of Um uiandBct at Oa
vcitlbuli.
openingi, on« of which is common to tlie eiiperior and horiioiital caoak
Ihe superior caiitil ia situated vcrticallir and at right onglee with tlie pos-
terior surface of th« petrous bone; the posterior eanal is also vertical aid
parallel with the posterior surface of the petrous bone ; (he inferior canal il
placed hurizontnlfy aud at right angles to the others. At the oonnDOiee-
ment of each of inese eanabi is a dilated portioD, called tiie ampulla.
The ooehiea occupies the anterior portion of the labyrinth. lu boae, whicb
wrreaponds to the iriternal auditory mftxiua, \s perforated by laauy mimite
orifices for the |)anagc of tilaments of the cochlear branch of ibc auditory
nerve. The cochlea consists of a central axis, or moJInltu, which liai a
SpinJ canal wound around it. Tbie canal makes two and a half compleu
luriH, and lerminateB in the a^>ei of the cochlea iu an expansion Leraied the
iu/itndlliu/um. (Fig. 'H').) i'he modioluH is »>inewliat cone^baped, tai
furiuB the internal wall of tbe canal, being [lerforaled in iU centre and M>l«t
by aitertura for the ponage of the filaiaeuta of the auditory uerve. Tbe
canal ia divided into tKO paasBgee or eeata by a »e|rtuiu called the lamimt
tpinlii, which is partly oweuus and partly tueiubraiiuus. Tbe ouaecm por
Uon projecta fruui tbe ciiodiulua, uiidnay across the canal ; il coneista of M
laiuiuie, between which ilie nerve filaineuUt run. The meuihranoua portiM
exienib fruui the external margin of the ua*e<ju» laiutua t > tbe external wall
rilTSlOLOGlCAL ANATOMY OF TQB KAB.
913
A.
V-,
e.e.
at tW raoa). Tt contiuitH of liro lay^n; ibe superior i» (he ii)i-iiilintii« of
Corli. or mirnifcrnu'i trrinria ; ihn iiifurior the tnan&ran'i baeU'iri*, wliicli k
•ttacbcd i^xlrnintlj to thr pl'inum trmilimnrr, Tli<;M mctnbmiKii lire iilacvH
psrmltel «ilh cncli i>th«r un<l cunlniii U-tw<ri:ii tliciu the or^nn 0/ Corii, wliicli
KstB on Uie bAsilnr;^' rovmbniiut. (Tie- ^Iti.)
Th*«raAi mrfifrif/i'coDimiiniciitra Mow wiili thcvwtibule by tho apertura
$eafa vrsti^uU rorhlta; the lower prnwi^, iir #r>i/a lymptini, cominuntcfttoi
with lh« tyinjniniim bv tht> fctiostni nitiimln. Thtc« ecuin communicate al
th« a{>ex of the otchmi by an o|K-iiiii(; liTiDnl the hialtu or hr/ifotrema,
which exists iti coDKijunnco of 11 tlctimncy or the Ital half turn of the
■ lamina Bpirnlis.
r SuTlod III>r>l-Gll IMI C^XrllLU. (IlMnniEr.)
«, axil wtUi lU canali: i, liiriiiullliuluiii nt •nlni^l uppw sad of Iho ub: c, H|-t"'a ot tl»
eothlM 1 iL aiMnlmna ttf Ouni : t, mambmns of Bcibdh ; /, tilMHi or ballaomiui 1 •>, antla
The ooKous portion of the lamina spiralis baa on its superior exiomat
Firlion a dcniioiilaled cartila^initus eubstanci* called the (antiit'i lifniiruOttii.
rom the superior surface of the spinil luiuiua, and internal 10 the lamina
denliculaia. la a delicate tuembrvDe esi»i<)iu}: upward and outirard at an
Ugle ul* about 45 dfyrees to tho cxli-nml wall of ihe seal*. Thia w called
the membrane of Reiamer. It divides the acala into Iwo paeoag^, the lower
of wliicb ia tbe (fiurFiu ewhUar'u. ThiA duel eodd in the apex of tho cocble*
in actvca, and coaimunicat«« at the haae with the aaccule by the ditettu
revnitnt; h contains the easenlial |)urtiun of the auditory apparatus of llie
coeblea, anil is a part of tbe menibranouit labyrinth.
The oeyin 0/ t'orti reata ujwd the traiiilury inembrane. It contnsta of Lhc
inner and outer hair-cell.i, and two rows of elongaied cells, placed paralld
with each cfther. bavin); an incliuin);; piitiliun tti that their free extrentitica
rcat againm each other and tbua form tbe ardi of Corti, which oovcrs Uie
ecftlrtii rjMiee, (Ki)(. '2\fi.) ThcM; rMwii are caller] tJie inurr and otttrr rods,
orpillaniol'Curti. Fnmi the itu]ierinr extremity of both the inner and outer
rods, Gnf^er-like proeewn project externally. At ibetr \mm» c<irrv9|Hjtiding
to tl»e oentral »|)ince are Mnj^le mwii of nitcleatcd cells. On tbe iuiprnal oide
of Ihe inn«'r riHl i* a »ing}e row. and on the external aide of the outer roda
■re tlirve rows nf elongated cilialol cella. Extending neniM the top of tho
oi^gan of Corti, from the iuner hair-oelU to the external wall ul' tlu- onal, t»
■ very delicate alructurc called the rtlievhr memhrane. The auditory nenro-
< Ttw tiMtr Matx b iJlililMl inEo two iwn to ■ menilinniiut ntUlhMi, tka uqa
aallaa >te •»>■ lotituu . Uia oibw, duciw cueiiksti* iri§. Mt).
of wklcb ti
RBABINO, BMBLL, A!fD TA8TB.
H
^
fibuiMtato probcblj lerminalc in Dw ciliated oella, bring (tiliroueljr onnsceud
itilh the riliii.
Tbo OHM'oii" ItilijriDth i» Matii br ■ fibro-M-rniu ntvitibrimr whieb WcMn
u wutcjy fluiil called Utc ptnlgmpk. Tbc perilymph fills ihe tcaJa of iJU
PM.Mt.
ADuo«AXor4«iCTK«orniiTiuiirnii:C^.iiitiKiuM.Hi. (UuHAoI tmm lUmt
«r, «■!• Tvtlbull : ST. wala l;in|iinl ; ft; «ml nf Ibc rorlik* ; I, neuitn** uf tt^mm
MVhlwr ■•ni'i-buf ibgaa<llr->r;DMT«: t. Iiini1iu«|<lnillii«n>! 1. i-tanon •Mn'iiinni': a. W
flnUruiMa; b.ii>1<.-'»i|iInLll<: M7iD|«nl>'ll|.atihi'(iilp<ii trJnlM: ■!. iBiKrtodc^ uidl : ».«
mdi«f Cunt;/. IkiulB* rvlirutul*. i. In»vr halrcvlti; «b, B«mlinH taitlulii mT.\
(MnUp.oululMlT-nllti M.cvutml mbp* LtUKmih* m*.
F>o.;n.
FnWt.
Fm mt.— Pxraotii iMoii Piinr Bntcrm n saov nil iiai>um>n Livruiini ra n^a
IBatMaiir.) n.niiilt mc: ft, lixitalub-. r.duotui camimi; ■!. U>a« wo or atiMv: i. Iliiilttil
/. ■iDIolIaiy «DUii<niiMil>oiiaMBlolm>Ju lube: g. •MDlaln''ilit ioIk.
now.— p«muMmooi«r!tiBna toTiii umitiwiisLdiiivnjyTit. iiiBmnn.i «,u«n«i»tt>
Minute: t, Dcm rMMtDf ib> umiiuUiirT tmlBrtniviii rm ■ «imlcltriilar lute: «,bn«(b4f »
ncm •ulartni tUa bitci n< o> nirttlt.
PHYSIOLOGICAL ANATOllV OP TtIK KAK.
»4fi
eoebles, nnd nirrouods the liueluf foMrnriA nnd th« tDembranous portions of
llie Ubyrinih, which an iiUiatcd in the vo*iibule and sttmicirciilu- c&aala.
The memlmiHoug fubyrinth ia a closed sac conuattog of the ecmictreular
oDals, D vestibular porlioD. and the ductus oochlMlia of the cochlea, llie
f«iiiicireu]nr caiuila arc of the bbido fom U tht OOMOUfi canals, and arc coii*
taiueil u'ithin ihera. The vestibular portion OOnKSto of an expanded body,
the ufriWtr, and a smaller body, the meevh. The utricle is eitualed at ibe
fuvi^ h«a)iellipiica; the eeniicircular canals open on ils internal surface.
The saccule lies at the fovea heuiispherica ; it ia connected with the ductui
oocbtearts by the diiettu rcunifiu. In tlie walla of the saccule and utricle
are two calcareous bodies called the oUilUht. The walla of tlie auipullu,
according to Bowman, alao contain aome graiua of a HiuiUr aubUaiice. The
walls of the inembranoua labyrinth couaiat of a fibrous ti»ue. lined by pave-
-^^^
:* -t
PUOUHOFTUSllOIISOrTSHIinitTtOHOrTHI ACDITOKT KKM t IX TVK A>rri.Uia UD JUlTVU.
1. Uwnllorthcaoipulla: l.nraotarelaslMMiiwnt-iiMtiibtiiae: >.(taulil]r«iiil(MFNl nenv-Bbta
I a I In I Wllilw UBTenliia llie baxnieiit-meiubmnc . ^ pIcilDirui iiDloii ot itac n(rT»6lMcs <rllb
Inimfaiwa nwW : a. ftullbrni Mil*, wliti micJuiw uul ituk fibni In (bait Iniolai T.nppgrtlai
«lUi«.andUMTt«lm.
mcnl nucleated epilWium ot-lls, having a elmcLurcIuK baMnuint' membrane.
These cpilhelial cells arc much niodilicii at the placo of CDtranc« of the tibrca
of thii BuUilor}- nerve. The vestibular branches of the auditory nerve are
distributed to the ampuUie. utricle, and saccule. ( Pig. 24^.) In tV utricle
and the saccule the fibres terminate in oral plates, called \\w m'lfida ttetutica,
which are more or lees colored by the de]>oeitioii of yellow pigment. In the
anpalba the libree terminate in elevations called the rrube onuti'oce. After
tbe ncrve-fiUtnent pierces the membranous wall at these poiole, the aiig*
cylinder alone pooetratee the baaement-membrane ; it then forms a pleiusof
delicate oerve-flbrw with nuclei, and finally terminates in fusiform epithelial
celb whicb have tenuinal cilia called the auditory hairs. (Fig. 249.) I1ien
dliaicd cells are suptiorted by ooluniuar epithelium.,
00
HG
HBARIKO, SMBI-t, AND TASTE.
The tnembninous Inbyrinlh \* lined by pulrgonal oucl«ftle<l e{HUieiiua,
vhidi mcfCtM Ihe em/o/ym/i/i wliioli &\l» ib'e aac]
At in the ejc, ao in tbc eiir, vre have to deal fini with » mrrv of tpttkl
Mnr, the MliinulatiDii »! which givM rite t« ■ RjHTiul •rnmlion : woundiT,
vith lerniiiinl orgaon ihrougli which the ph^cal channs (impt-r In iW
spodal «-ntK> arr rnalilcd to act mi (hv i>vrve; nod ihirdU. with luUidiuy
nplioniluR, by whifh tbc MMrulncM of the tviwr i» incnrnwil. TIm- n'lilnl
ciinni'diniif "f the nitilttory ii«r\-c are nuch thnt whenever llie auililitry libra
tin iitiniiilnlol, whclher by inc«n!> of the tcrmioal (>rj{iin» in lite usual an
or by ibi; din-rt Eifi|ilicali(in i>f utinnili, t-lcclrical, ntcrhiiniml. vie., thv tnult
a always a Ki-iioiitiQD of •ouiid. .Iimt n» etimulalion of ibc <mlic fibtn ft>
ducw nn other Mnxattun than llmt of li^hl, foRtimulatton of ihe auilituq
fibre;' prixliiiT* nn nlhcr M-nntiou tlinii that iif fumul,' Tbc terminal orgui
of ihc niidilory ncr\'c arp of two kinds : the com plica lod orgno of Corti ia
the oocblca, and ihc epithelial urningcmcntff of the macula.- and cristv ui*-
ticir in otber pnrti'of the Inbyrinlh. Wavc»of »oiinil falling on the auililoi;
nerve itself produce no cHect nhalever; it is only when br tbe mediiin m
the endolymph Ibey nn- brought to bear on tbc dcticalc aiMl peculiar i!|hU»
lium eel's whicli coiislilule the peripheral tcrininaiions of the nerve, titf
Mnsallone of sound arise. Such delicate sliuclures are fur the sake of pof
tcction natiiraily withdrawn from tbc surface of the body where lliey wmiM
be sulnecC lo injury. Hence, the necessity of an acoustic apparatus, fbrnuDf
ihfl middle and external ear, by which itie waves of sound Afe most Bdns-
lageously conveyed to the terminal organs.
Thf. Aeovjilic Appnraiui.
Waves of sound can and <lu reauh the cndolym)ib of tbo labyrioih b)
direct oou'luciion through thi- xkull. Since, however, aononwB vibnitiua»an
IraDsmitled with great ilitlicuUy from the air lo solids and Uquidit. and iMal
sounds come to us (hroiigb Ihe air,soRie special ap|iaratus is required to Ina*-
fer the aerial vibnUions to the li<iuid« of Ihe internal esir. This apparatiu
is enpplie<] by the tympanum miu il* appendages.
The condia. The use of tbis, as far as bearing is cuncerned, b lo coIIm
the wtivm of sound coming in various direclions, and lo direct then un to the
nicinhrann lympHni. Inourwlvta of niodemteserTkeualy, in niAnyaiuMali
il is of great imjiortanco.
Tlir mnnbrana tpiipani. It IS a chnracieristic protierty of slreuJied mem-
hninrs that they are reiulily thrown into vibraltuu hy ai-rial wavca ufwaad
The membrann tympsni, from iis peculiar conformation, being funnel-tfaa^
will) a deprcsevd centre surrounded bv sides gently convex outwan). ia pecs-
liarly susceptible to sonorous vibrationii, and is ni'if't readily IhroKn tDb>
corresponding mnveoieula when waves of sound reach il by the meatus. H
has. moreover, this useful feature, ibat unlike olher mrelclied metnltnina,It
has no marked note of its own. It h uul thmn'o inio vibraliona by want
of a pnrlicular length more readily tliau by othcn. It answer* eqaallj aiM
within a coiisiderabit! range, to vibraliuus of very difierent ware-teogik.
Had it a fundamental tone of iu own, we should bJde>iract4>d by ibc pruM-
i>encc of thb note in mi»t of the sounds we here. When sounds iuijiiu]^ o
the solids of tbe bead, as when a watch b held between tbc teeth, tbe nina-
I It will b* M«n Ut»t on itikt llinv an- rraKini for llilnkliw ihsi impnlaa [i*M>ia slMaM
sndllOTT Mnr may fiva na« U Mktr Mncu tlun auiiiiary MnMltont,
rOTSIOLOOICAL AITATOMT OF THE BAR.
I
I
I
liniw lymponi u Mill fiini:ti<iiiiil. VibnttiDiM ara cunvoyM rrnm the trm-
|>onil IxiCie to it anil bciici- pnm in th« tiituiit wny, in lulililiuu lo thotu
tmoeioitlvd dirvclly iVoni ihv ixinv to (W [>orilrm|i)i.
T/ie audilori/ tuvii-Je*. Tho niallou*, tlio IihoHIc of which ilMConding for-
wanl Mid inwBnl, it itllBchcd lu the meinbrnnn tympntii, unti the incus, whose
long proccM i* connected l>y meaoH of its ob orblculure or leiiliculnr procee)
aoo lh« ilat>es to tli« leneMru avalis, form toj^lhcr n body which rotates
rouod lui MM, poaiiDg through lh« ahurt proccn of the incus, tb« bodiiw of
the inciM nod nislleus, and the proccMua ^racilia of the mullou?. When the
nallciiii IB curried inuard, iho incus mtive^ inward too, nnd when ihe midleu*
returns to its p«ilion, the iiieiia returns iiiih it. lh« peculiar andille-aliaiie'l
joint uith iia catch ie«ih ])erniitlin^ thin maremenl readily, hut preveniinjj
tb« stajiM Itcin^; jmlled buiked when (he membranu lympaiii "ith the iukI-
leua is, fur any reason, purthvd oulwanl more than usual; the juini thep
gttpw, so as to permit the malleus to be ii)i>ve<l alone. Various lignmeuta.
the superior or suspeoii'iry, tinlerior, mid vxterual, also aerve to keen the
iDalleuB in place. The whok- Kcrit-* of ONiicled luaj be resorded bb a aliigle-
anii«<l lever, moving on the li^'amenlal lUtuchmenl of the wort proceM of Ihe
iticii!^ lo the poDieriur wall of thu tynuiununi, the weight lieing bruu)[ht lo
lieor at tli« end of the hmg pruuiM of tlie Incua, nad tM puwer at the «ad of
ibe batidte uf the inallcu*. Tbu long, uwllual arm of thu l«ver is utiout i>^
tnni., the abort, «tapvd)a1, 61 Diai. in length; htiice, the movcmcnla of the
Ma|)e* ore lew than thcwc of the tyiiifmniim ; but the Uw* in nmplitudv b
toade up by a gain €>f force, which is in itcvlf no obviotia odvuiitAge.
Thus e^ery movrDieat of the tvnipniiic membrane i» traiismitti»l tlirough
ikik chain of omiclva to the mcnibmnc of the fenestra ovaliii, and so to tb«
iwrilymjih of ihe luliyrinth: the vibraHoiis of the tympanic membrane nno
conveyed with Increu^U iut^usity, though with dimiiiiBhed amplitude, to the
latter. That the boms thua miive tn nuute bas beeu proved by reconliuj;
their moTcmcnIs in the usual L'raphie melhod. A very light style attacheal
to the iaeua or atspM is mudi- to write on a travelling surfaoe; wheu the
niembrsna tymiwiii u thrown iuf f vibratiouit hy a sound, the curves dewribed
by the style indicate that the chain of bouai murea with erery vibration of
llie lynipanuni. Uu the other hand, the comparatively looee attach men bt of
the several buiiM is an i)lj»iuele to the mulecuUr traiisuiisaiou of wmoroua
vibrations ibruu;{h them. Moniivcr, auuoruui vibrattona can only l>e truna-
initied lo or puiu ahmg such bndie* as either are verv long compared fu the
length of tile souiid-»avn, ht, us in the owe of membranes and striun, have
one dimeoaiou very luuvh smaller thim thi; ullivra. Now the bun«A in ijuoi-
tioD are not e«[wcially thin in any one dimvnsiim, but are in all their diinen-
sioos exceedingly small c»m|uircd with ihu length of the vibratious of even
the shrillest sounds we arc capable of hcjiring; hvucc, they inusl be usdesa
fur the molecular propngnlimi of vibrations.
Tltt lauor lymjtani mu*i-lf even in a ipii<9cent state is uf um In prevmiing
the ntcinbnina tympani being pushed out fur. When it cunlravt* it render'
the ni<-mbran« lympaui nioro ten*c and Ix-ncc haa tiaen *up|Hwci1 to act m
a ilam|K-r lenseniDg ihc amount of vihrnliou of the memlirone iu the esse of
too |K>werful soumls ; it is aaid to be rotidily thrown into contraction at the
commencement of a sound or noise, but to return to ri-sl during tiK conlinu-
BUee of a mufical note. Klferenc impulses roach it lhr\iiigh tibr«s uf the
liflb nerve, and ila activity is regulated by a reticle action. In some pemoa
Ihc muscle eeeoM to l>e partly under the dominion of the will, since % peou*
liar crackling noise which these persons can produce u pleasure appears 1»
bo caused by a contraction of the tensor tympani.
94S
IIEARINO, SUEItL, A>-D TASTC.
The so-called lasator ivinpani ia coDBidervd to be not a miucle at kII. hui
a |iKtt of the ligainentoUfi Buppvirtu of th« lunlleuA.
The ttaptdioM muneU h su^i^i-iAed to regulate the movcineats of the alapo,
iiud t^jienally to preveut its baae bein^ driven too far into the feuestn ovilii
during liirgu or Auddea uovenneDtB of tne EDerabniiia tyiu|iani. h u ((nnnal
by flbfee fmrn the fm-M uen-e.
The JCiutac/iian tiibr. ThU wn'M to niointain an equilibriuio of pmnn
between the extenml ntr and ih»t wiihhi the tyiuiMinuin, aod to serve ■* u
exit for the secrelionH of that cavity. Were the tyoipaiiuto peraaiMntlT
clcmed the viliratiitnB uf (he ineiulirana tyiiipaiii would l>o irijiiriouKlyaAdel
by variatioDs uf preraure occiirrlu^ either itnide i)r outnidc. Tbe KaHtacklu
tube i» undoubtedly open during twatloviug, but it if Mill iliii|iuieit wfaeiber
it remaina permaDeiitly oiieii, or it* opctied ouly ut inierTiil* ; probably i| a,
at imift liitieri, iieilhcr widely open nor clowly *buu
Auditory Snuationt.
tUit-U vthimtion ommuiiictiled by the itapes to iho poritympb trareltaii
ware ovpr the vrftibulc, tbt- Ronii circular canals, luid other p«rte of the labr-
riuih ; and from (he iwrilymph is iranBioittcd through llic menibrunous wbOi
to the endolymph. From the vestibule it passes on iuto the t^i\a vcetibuli of
the cochlea, and drsc!cndiiig the scnln lympani, ends as an impulse agaiail
the iiieiiibrano of the fruentrn rotunda. lu the regions of the luaculie aol
erialie the vibrations of tbe endolymph are supposed to throw into eon*-
spi'iidinj; vibrations the Bo-called auditory bain. In tbe ODcblen tbe vibn-
tiona of the perilymph are suppo§eil to throw into vibmliooB lite hB«lK
membrane with the ituperini{)0»&d orj^u of Card, consisunc of il
Corli with the inner and outer hair-oelU. Tbe vibrationii iIiu.h tr
to these ttructurts zive ritie to nervou* iuipulstd in the term I nii tiona
auditorv nerves, and thew impulMV reaching certain pariH of t)Mi braio fl*-
duov wfiat we call uudif)ry Kunatioii:^ We are aovuatooied to diviile oar
auditory aeinuitioni into ihrwe caiiited by noiso* and ihottt cauwcd by miwial
mund*. It L> the charactcriittio of the littler that the vibraiiona wludi eaa-
»litntv them ariT |n-rti>[lical ; they occur and recur at rt'gular inierVKli. Win
no mnrkt'd jii-Hndicilv in jircseiil in the ribration*^ when tbe repntltioa of tbr
wvcrnl vibnilicinM U irregular, or the |>erio<l »n complex Ha Dot U> ht nti/Sij
apprecintt'd, ihr upmrntiun produced i» that of a miisr. Tliere u, bow«T>r.tW
abrupt line belwom the two. Itelneeii a pun; nud rimple musical SMtai
jiroduocd by a iK'rii-a nf vibnittons each of which hnit exactly the same wb»»-
length, lutd a hnn>h noi»e in which no conHUCtilivr vibniiioiis may be atikft
there are numerous intcrmt^dlale #tagea.
In l)oih tioi«m and musical «oiind« in nongiiiite a character wbicb w« all
loudncM. This I* dctertuiocd by the amplitttdr of the vibrations ; i^ gnaut
the distiirbanoe of tbe air (or other ined!i)ni) the louder tbe sound, la ■
luusical sound we reeogniiu? also a character which wc call pitch. Thit II
determined by the wavedcnglh uf the vibrations ; tbe shorter tbe waf»
length, ibe larger the number of consecutive vibratioiM which fall unoutfe
ear iu a second, the higher the jiilch. We are able to sj)euk of a wbide ssis
of tones or musical snunchi of diSerenl pitch, from tbe lowml to the ht][bcC
audible tone. And even in many noises we can, to a certain extent, nof
uixe a fAuih, indicating that among the uultifarioiu vibratlonii tliure its
periodicity of certain groups of rtbratiaiis,
l,aDtIy, we diatiuguiah mtuical luunds by their quality; tfao wune Ml*
sounded on a piano and on a vioUu produce \'ery dtlfereut seoMtions, «na
PHTSIOIOOICAL ASATOJCT OF THE BAR.
ul)«ii H arrir* of vihratimu having in vnc\i ctvte Lhfi Kuav periorl of rrpctitioii
b wt lulling- Thi« nrJMfl JVnni the fuct tlmt the mimical soiiods gcaeratcti \ty
noM miuiciO ingtniinrnls lire not siniplo but coniDDUti'l vibnuiooa. Whcti
lb« notv (' ill the treble for instiiiico is rtriick on tne piano, xni) wo anmlym
ihv tolnl MHiDi), u'c liml tlinl it cnn be resolved pnrtly into w, terics uf vibra-
liuiw with a period cbumcteristic of th<3 pur« time of tbo treble C, and
MTtly into other aerie* of vibration* witb perimle charact«ristic of tbe C
in the nclave above, of the G nbov^ that, of the C in the next octave, and
nf the E niMve tiiaL And the Mii»tiuii which we anooiate with ihe wiund
of the treble (' on the piano is detemiiDcd by the characleri of the complex
vibration arising out of these eeveral eonetitueiit simple vibrations, A-lmoM
nil musical sound* are thus composed of what is called a " lilDdaneatal
tone " ncoomjMtnied bv a unniber oi " overlones." And the overto&M tary-
in;r in number and relalive promiiience in diflbrent inslrumeDta, f;ive rise to
a diflerenoe in the sensation caused by ibe whole t^me. So that while the
fundamental tone determines the pitch of the aound, the quality of the sound
is detennitied by the numlier and relative prominence iif the overtones. In
a sontewhnt similar way we di^tiui;niiib ibe <|ualiLy of noises, sucb as a han;;-
iag, crackling, or rustliDji; iKiise, by an ajipreciaiinn of siiilden or irregular
cfaanges in tlie amplitude and |>oriijd of the cimstituent vibrations.
Since we have a very considerable nppredalittn, eaiiable by exercise of
aotonisiiiflK enlar^niem, of the loudu^wi. piieh, au<l ijualitr of a wide ron^
at noises aiid umsieul »)undd, it is dear iliut, within the limits of hearing,
each Tibration or serieit uf vibrations must prixJucc its tSix% on the auditory
Dcrree, according to the measure of its intensity and period. Out of those
eiffierU, out of the sensiiry im^iiilsM to nhicb the several vibratiuas thus give
rise, are geuenttud our sensiitums of the n'liso or of the sound.
The vibrations of a musical sound (und tinoo noisea an oo tmperf<Ktljr
undentiXN), we may, with benefit. cbieHy coulin« outwivca to rousicu
sounds), as they pa» through the air (or other medium i are not discrete ;
the vibration* rorrwpoDding to the fundnmenlnl lone and ovsrtooes du not
travel as so many separate wave* : ihey all toj^ethor f^irni one complex dis-
turbance of the medium ; and it ia M one miapo*it>- ware that the sound blla
on the membraaa tympani, and paasing through the auditory npptiratus,
bnaka on the terminations of the auditory nerve. vVnd when two or more
munotl sounds are heard at the Mme time, the same fusion of tlie waves
occurs. iJince we can di»tinguidh several tooea reaching our ear at the same
time, it ia clear that we must poeseaa in our minds or in our ean some
ninna of aaalriiiigtheee composite wares of sound whieb full on our acoustic
omins, and of sorting out their constituent vibrations.
riiere is at liand a simple and easy physical method of analysing eom-
jMiaite Buumls. If a person standing before an open piano sings out any
note, it will tie observed that a number uf the strings of the piano will be
t]in)«n into vibration, and uti esamin^tiou it will be found thai tb'^-' strings
which are thus set gi'ing e<>rn-ii)ioi)il in pitch to the fundamental tone and to
the several overtones of the note ^uug. The note sung reaches the strinj^
oa a complex wave, but tha»e string* are able to anolyxe the wave into iia
OMHtituent vtbrutiouD. uich siring taking up those vibrations aud those vibra-
tlofwonly which Iwlung to the IuUl- giveu forth by itself whun stniek. If
va s«[)|mmn; that tuich tenniunl fibril of the nuditiirr ner\-o is connected with
an organ so far like a piauo-slnng that it will readily vibrate iu raspODse to
■ Strics of vibralin;^ iiiinulnugi uf u given |ieriod and to none other, and that
we poascw a numbrr ot nuch terminnl urgiuu :iutBcicnl for tl»e Bnal)rsis of
all the M>nDds which wv can analyM-, and that each terminal urnn au
affected by particular vibrations give* rite to aaensory impulse aud thus to
9fiO
HKAHINO, S»EI.L, AND TASTE.
a ectiBnIton of a rfUtincl ch«i«rter — if we Mipp-Jse thew ftrfrnna toexitt, nur
a j>| I red at ion of soumlii in in a larpv moasHre cxitlaiiiHl. In Um Offu ol
( orti w* find Mnictures lUe nrraniieiueiil of wliioli irmutiblr sajuilll to m
lliiit tli«e lire tlie organs »'e are sei^ltiog. We have only to BiiiiiHoe thai nf
the long series ff mda iif Ciini. vnryiog rejrulariv as these ih> mmi ib« }**■
t'lm to the top of the spral, in length and in the a|ian of their arch, radi
|)air vrill vibrate in rm>i>ii»e m a iiarticular lone, and the wlioI« aiaiitt
•cenu explaiiiei). But the more the imhject i* tiiqiiired into, the more do*-
]>lex and diliicrult it n|i[ii-ani ; unc} we are obliged lo conclude that the ma
pUved by the r»dH of Uorti is only a (ubordintite part of th« Ainclion uttb
whole organ of C'orti.
!n dit firM place, it is rlifficnit ta see hnw the rmfe of Corti. even If lh»f
are thmnn into vi1miti«ii, ran rirSginaie wntory irapulKS. for th<< fihriU of
the aiidiUiry ucrve (erniii)Bte in the inner and niiter hnir-ecll*. 'nd illth
thCM cell». nnil not along the oonrw of Uio fibril* w lliey |MMt umler ui
b«tiroCD the rodn of Corti, thai ihc »en»ory impitlRi* must iMtrin, In tl4
Mvoml place, the variation in length of the fibrm along the »ork» is itmfl-
rient for the work neHgiioi) to tnem. Monrover. they appear not lo bt
elastic. La«lly. they are wholly alMrnt in binK who vcn' cWrty can appn^
date muRical Rounds. This Wt fact proves indubitably ihni the rocli ia
ijucilion arc not nbsolutelv cswniial for the rconiftiition of tones. In tk
Aice of those ditlicult)i.>s it "has been sii^estcd that tlie h«eilar membruu.
which ifi present in hirdt as well as in mammals, and whieh, beioj; tow
radially but loose longitudinally, i. e., along th« spiral of the oocbW, mat
be considered as congisting of a number of parallel ndial strings, eacn
capable of inde|>endent vibr«linns, is the soughl-for organ of nnnlyDtt: fcr
it may be shown niathematically that a menibranA to stretched in oae dinc-
liou only is copablu of vibrating in such a manner. And tlH> mitial disKO-
duns of tho basilar membniue irive a much greater range of ditTcrence tlun
do the riuU of Corti, diminishiu); in nmn doirnwanl fh>m 0.495 mm. at thi
Uip to (l.(M12S mm. near the bottoiu of the spiral, whnn^ut tbc diirerrnu ia
kngth i>r the latter 'a iiimnly thut lielweeii O.lMFt and 0 085 mm. for the inatr,
and betwt«n0.01!> and 0.08,1 mm. fur the out^r libra. Aocordtngti) this rirw.a
parlioular Kimpli- viliration rciidiiu^ the s>;nla tympaui of the cochlea iknxi
into sympullu-tic vibmtii<ti!> a Ktniill purlion of the b^iiilar membcaop. ih*
f-ibrat40DB of which in turn so affvet [be stnictun^ overlying it. that acnaan
impulses arc gvncnitrd. The K-nfory impulses reaching the brain gite Hm
to a corresponding scnmlion of n particular lone.
The remarltabfo reticular ni<^nibrnnc which has such jieculiar relatloai
with the hair-cells, and thri>ugh them with the bA*ilar membrane, must, am
might imagine, have s>~>mG specini funi'tion ; but it is impis*ib]e at preatui
to assign to it niiy satisfactory duty. The structural arrangemeuls te*m,H
anything, to indicate, that when a segment of (he basilar niembraiie h
ihniWD into vibrations, the overlying hair-cells, reticular meuibmuc. aad
rods of Corti vibrate en miate with it. Itul this renderst Ite whole inaiur
still more diflicult. Indeed the whole subject is in the highest ileartw nb-
Kctire, and the moat we can say is that the organ of Corti as a whole wnw
lo be in Home way cannected with the appredalion of tones, but thai tf
present it in very hanirdoua to attempt lo explain bow it acta, or to naign
particular ftiDetinna to puriinitar pans. The disliuciiun between the iiMr
and outer hair-cells seeiua to be very parallel to that between the rode and
the concjt uf the retina : but even tbia analogy may be n fallacious one.
It hit* l>een observed thai among the niiditor^- hair* of tho cnis
•Mnc will vibrate to particular notes; hut the auditory luiini of tbe ma
I
I
■ra fur loo muL-ti of tlie Mine length to permit the BUppoeition tliat tlioy can
set ■■ or|;ni» of ftnalnis.
If the organ of C<>rli ia the mmnB bj nhicli m B|>prcciiitB tones, it ia
pviilent that by it also «e must bv able to e«tiniate loutlocas. fur the quality
of a muHiral sound is dependent on the relative intenBity. ae well oa ou the
tiaiure. of the overtones. Anil since noise ii nl best bill omtu^ niueic-, the
cochlea iuuh be n nieaiii of apfireciatin); noises as well lu si>iiada. Uut this
wonid leave iiochiuj; whatever fur the rest of the labyiinlh to do in respect
In the appreciation of sound Aave to far as the diKerenoe in etruc'Liire betwoeo
tbe blir-eella of CVrti. with ilieir short, thiek nnia, and the hair-Iteariiifc
Mruciurei in the inartilw and criHlie with their ihiu, deliral« hair^, may
pceeiblv iiidieale a diflerence of fuuciion. the latter bein); more eui<i.'e|itible
to ihe irreitiilar vibratinua of iioiMs. That the veHlibule and sciuiclrcular
Cuiab ari-, however, conterned iu hearing U Bhona by ils heiai; tbe only
Auditory orKan in the icrhthvupHiila. iiuledH wo su[)|ioee that in the Ui;rhi-r
verHferatea ita Amotion has l>e«n wholly iransferreil tu the cuchlm. That
tbe wrakircular canala may have dutiuo a|wn fr>i(u luinriiig we ahull abow
later nn.
Concerning the flinction of the other |Huia of the luleroa) ear we linaw
very Utile. The oiolitlm have t^wn »up[)uiieil to iiiteinufy the vihrntioiis of
the ondnlyniph ; but ninco apimrunlty they are l<Hl|cvd in a (luanltty iif rouaix
it u probable that ibcy really act bh damjierK. A aimilur daiupiug action
haa iieeii Migfniaivd for the ni«n)brane of (^irli (inanfimiut leciart'i) over-
han^nit the fibre* aixl hair-cells ; ao<l some wrilcni have auppoMd that mu*-
eular fibres prewnt in the plaiinin frmiluiinre nmy by tightening the basilar
Bwitibrani; serve iw a M>rt of ni^iminiKlation me^'hanixni.
(It mart, however, he borne in mind that creii niaktnt; the fullest aliow-
ance for the assistance alfor^led ut by the orfran of Cjrli, the appreciation
of any sotind is ultimately a mental act. The analysis of ibe ribraiions by
tbe fibres of Corti or tbe basilar membmne is simply |)reHtiiiimry to a
syntbeeis of the sensory inpulMS ao generated into a complex seiuatioo.
W« do not receive a distinct aeriei of specilic auditory impulses rMulliun in
a apei-iflc Kneation for every possible variation in the wave-lenj;Lli of sonor-
ous vibrations any wore than we receive a distinct series of speciKc visual
impul.-es fur everv noasible wave-leuicth of luminous vibralioos. In enoh
^cate «e have nrobauly a number of primary sensations, from the various ^^
mingtinK of width, in ditlerent proporljona. our varied complex seoMtioaN ^H
aride: the dilli^reiivc- between the eye and the ear bein^ that whereas iu tlie ^^
former tbe number of primary senaatiuna a[rpeurs f> be limited to thrve or
tat least to six. in the latter, inanks to tbe or^n of Oirli. the number is
very lar^^: wliat the exact number i« we t«iinot at prewnt tell. Oiirap-
preeiaiiuu lor a sound b ul bottom an appreciation of tbe CDmbinol etlvct
pniduced by Uie relative iuten»itic» ti> which the iirimnry auditory wiuatiuos
tare, with thu be'ii nf ihi.- orifitii of 4*i)rti, excited bv the sound.
Whatever Im.' inc exjdaiiatitHi of tbe manner in which our iltntinct auditory
•en««tiun:> ariM.-, the ranjcv and pn-i-iiiion of our aiiprei-intion of mutical
•iruniU i« vi-ry threat. Vibration* wilb a rocurreiici* Wlow 'Mi a itocond' are
aoable fi pr'nIuM) a scuutioii of snuiiit ; if thi- waves an: [Hincrfiil enough
wa may firl tbem, but we do not hear ibcm if the vibrations aa- Hmplc, and
•uoli M would givo rise to a pure lone : if the fundamv^ntal tone U aocom-
putiod by overtones wo may hear ihi-w. an<l are thus apt to say we hear thfl
fonavr itdsu in reality we only hear the lntl4T. The nota of the HMbet
organ pipa, 99 Tibrations a secund. gives us the seasalion of a droning
I B} KmM kutlKin tha llmli M plutil M tuit u U w sm It ■ •reoiri.
u
Qr>2
nSARIK'O, SUEM., AND TASTE.
nound. A tone of 40 vibraliana is. lMwev«r, qui{« distinct. la Um oUsr
direction it is pa»ib)e to bear a iioK> caused by 38,000 vibnttoM • moaai,
though tlie limit for roost persoti^ ia far lower— abmit 16,000. Some foum
he*r grave souihIb more easilr than liigb ones and vwe vena. Thb idat be
•o pn>nouD«ed u to juatifV tJie subjecu being ifwken of as de«f to gnn ur
hrjtn loneo reflgwctively. 'TIic rangt in difTerent aniinala is vewj difliervaL
Tbe {Miwcr of iliitiiiiifuiabiti;; <>n« tvAf (torn another varies, aa is well knon,
in diffeKDi iudividiiitl*. aeconliiig as ihejr liave or hare not a " mu>i«l tar."
A well-lmiiiix) enr can iltmiiiguiiiti ttie dillenenoeof a nngle orerenofi
half ribntimi u !i>h:oiiiI, aixl tlint ilir»ii);h n long range of DOtci- Tbemafi
of an ofdiiiary apprecialiou uf um«vi lie* between 40 and 4000 Tibratioiu ■
t)econ<), t. e.. ik-twmi tiii> luKist baas C' (C, 33 vibrations) and the faigltM
treble C (C* ■l^'i4 vibnitions^ nf llie piano; toi>es above and below Umse,
eveii whei) aiKlibk-, tieiu^ iliatiiigiiiali<-d from each other wiili ^reat diffieolljr.
When twu conwicutivi; noiiiitU fulluw each other at a suffimeotly short is-
terval the teiiiwtiuuii ore fiiwd into one. lu this respect auditory sensatiua
are of shorter duratioo than ocular si'iiMtiuus. Wben oeular aeoMtlons an
reiKntMl ten tinm in a wtcood tliej l)ecoine tuiexl (p 915), whereas the licb
of a [wnduliini Ix^ting 100 in a fcmnd are readily andibloas distinct autiiMk
When two tiiiiing-rnrks not i^iiitt-* in luni-urtTtitnick toeMberlbo interfirreiui
of tliu vibrntioiiii i^vt* riw tmtn idli-rnalin^ riso and fall of the sound, knotn
as "beats." Whvo the tHitU fullnw i-iicli 'Hhor ii^i rapidlj as 1^*2 in a seooad
they nnse to be recovgnisetl, that u to say, the sensotioas wliicb ibey csim
become fu^cd. BeJoro ihov diKapp<-.ar Ihcy give a peculiar diigresahk
roughnm to the sound. Th« pleasure given by musical sounds depcadi
larvely on the sbaenco of this incomplete rtiuon of lenMtions.
(. orretpoiiding to cntojitie phennmenti there are various c»Mie phraom-
ena, sen Rations or niodiflcaliaDB ot HeitsatioDS orifcinolin;; in the tvmpaauni
or iu iho lai)Trinth ; moreover scnsatioDs of sound may riee in the auditosj
nerve or in the brain itself, without any vibration whatever falltug on llic
lahyriDlli.
AnJiiori/ JudrjnenU.
in leekiiig for the cause of our visual scosatioM «v invariably rvfer to Hon
external world. The sensiili^in caused by a direct •timnlalioii of tbo opir
iterve or retina by a blow or ■ galvanic current, we ideiilifr with that caustH
by a flash of light. A sensation arising from any stimulation of the leA
Hide of nur reliria we regard as caused by oonie object on tlic right-hand sid«
of our exteninl visible world. In a »imiliir way. but to a luis exietil, «•
project otir niidilary sensations into the worl'l outside us, and when the aiwU-
lory nerve is alfeclcd we seek the cniifc in vibrations starting nt a gre«ter cr
l«n distance from us. Wc do not think of the sound us originating in Um
car ilM'ir.
Tiu* mental projection of the sound is much more compleie wlien the ctl
is stimulated by vibrations renching it tlirough the n>embTanH tv|iai)i thn
wlten thv vibratioiis arc conducted by the BoUds of the head directly to ihl
perilymph of the labyrinth When the meatus extemua is tilled with duiJ
and the vibmtionsof the membraoa tvmpani are in consequence inivrCntJ
with, the apparent outwardncas of sounds is to a very large extent kal:
soun<ls. however caused, seem under these circti Distances to arise in tbe eu.
Hence it would eeem that the vibrations of the membraua lyiiipani. sr
poesibly the action of the muscles attached to the ouieula. givK rise l»
obscure sensations of nbioh, by themselvM, wc arc not distiuctly conscKHU.
953
l>ul nliich DcverthelNS lend u< to juilgc thnl tliG K)uni]ii lieanl bj- tncans of
th« lyiDpaiiiim come from i>iil«ide theenr.
Our judgment of th« tlitttinee of 8oiind» is very liinit<^l. A s'lnnd whose
rh*ncl«n ira know ap|)eare to ua nenr whon it is loud, mid Tar olF wtien it is
Aint. A blindfold pereon will be usable to dinlingimh bctni-ca llie difier-
ence of inteneiiy pnidiiccd on the one bund by n tuning-furlc l)«in}[ held
befitre liim, finit with the broad ed)(e of (he fork toward liim and lliCD with
tbe narrow edfte, and the diHerenoe on the other liand cauaod 1>y the removal
of Ihe trmini;-fork lo a diatanee. We can, on tbe whole, bctt«r apj)rti<.'iat«
tbe distance of DoiseetbaD of iniiueal sounds.
Our juii^nieiit of tbe dlrretion of i)ouo<U is also rerv limited. Our chief
■id in this u the pontiou in whk-h we have to place tne bend in order that
we may bearthe Bound to the best advantage. If a tuning-fork be h<"ld in
the n>edian vertical plane over the heiid, llioiiah it is easy to roconpiiH.- it an
beinic in the median pluDc. it hecomea very difEcull when the cyn' are »lnit
to say what is ita piwitiMii in that iilanci, i. e., whether it is more t«nard th«
ftoDl or luck uf tbe heail. In tlii:> re»i>ect, loo, our appreciation it more
rate in the case of awstv thnn nf niUHical tonnds, with the exn-plion of
pveu out by the humnii voice, the ilirectiou of which can be judged
better Ibaii even that of a ooise.
[Phytioiogietii Anatomy «f the yatat Ftma.
Tho nnnl foMse arc two irregular caritiea nhich oommiinipate anteriorly
with tho air, through the anterior nares and jMjateriorly with the pharynx
through the posterior nares. The fo'aie are partially divided into upper,
mitMIe, and lower air-pawauea or chamlwm by the Buperii>r, midiilc, and
inferior torbioated buncs. They are lined by the Rcbneiiterian or pituitary
niueoos ineinbrane, which ia oontinuntis uulcrinrly with Uie integument and
poateriorl^ with the mucoua membniDe of the pharynx ; and witli the mem-
Drsne lininjc tbe ductannd siuuKt* connected with the foam. At the podtion
I of the <Iiatril>ution of the nlfnclory nerve fi la men l« it u much ibickiT, more
vascular, jiigmetitecl. and Vtuvd by (x>liininnr nucleated cpitlu'liuni crib : tbe
nmaining |>urtiun of llir nieniliranu covering tbe fimue, excepting near the
■Bierior narai, in lined by columnar ciliated epithelium. Thin monihrane
coataina racemow mucous glandM, which mctHc mucus for the jiurpoae of
keening the membrane oonataotly moiat, which i» a condition cswential lo
perfect ulfnction.
Tlie olfactory tract i« a prolongation of the cerebrum, which terminates
anteriorly in a hulb>iu* expansion, the olfactory ganglion. It cunrista prin-
ciinlly of gnty malU-r. Thi^ ganglion rats upon the cribriR>rm plate of the
cinmoid In'uc, and in thiif position tends about twenty filaments, which con-
sist of gray mutter nione. thtvugh the cribriform plate to be diMributed to
the piluitnrv membrutic of the upper third of tbe septum naai, tite upper
portion of the roof of the no«e, Iho superior, an<l a portion of the niiddle
turbinated bones, i Fig. 2d0.) The wliole surface correapouding to the dis-
tribution of the olliictory nerre is colored brownish by the pigment in the
epithelial cells of the mucous glands and merohnmei This nig men ted revion
is ealkd the rryio ot/uetoria, and is tbe eneottal portion ut Ine niual foMH
ooncemed in olfaction.
According to .Schullie, the epithelium of the regio etfattoriit u of two
kimla: The tirat (Fig. 261, a J couaisia of yellow nucleated pntoptatniic eel'
9t
RKARIN'G, 8USLL, AXD TA8TE.
which linvc a cylimlricnl boilr ipmiinaiing Al iu Tree extremitr m k •quuvl
Iruncatctl Mirfiic-c ; tliu 'lUier cxtrt'tnilv of ihv kxly is vlrvHcboc) owl m » fiW
in«Dtou» prolongiUioii, which cxpamls inin n irian^ilnr phitc nii it tpmmtim
the eiihmiicoiis liwiio. Fmin ihc luue of thin plaiv ii numbor of nUntili
nre ^vcii ofl', which nro pniKm^^t into the mbiuiimii* ItMiio. Th* wtcBti
They nre »imilur (o thoso just d4v<:ribc>]. exco|icing thiit ibeir frm titrhmk
covered with cilia. Between the epithelium rolls the oIlHvtofy oerrcM unt
Fio. aU. run. m.
V
I
Pio. asi).— VitRTit*!. ftK-nnx Of IliuHT N*ui. rnnu.
Mioimlu OITKK 8ii>i or ronu. 1, ulDirtoiy irnct: 'j.
oinwiory ncrva: 3, mlddls lurblBktcil taoo: i. lo«*r
(iirliliisM br>ne: A. Iinnrlim Itain Ida dflh n«rTt>.
Hnncba or tbo nfUi an kin (liowti In Uio ADWrtoc
inrllon. (ARor arhoui.)
Kn. Ifil.— CiLu or nu ULrifTOHir Uccoi'i HtuinajiM. (o. fr. <; kAor acHCLas; t,Af, *■<■
LocKium CtAkKt.)
oat«. These tennioal (ilanieota (Jb.f) are lonj;. delicate aiructare*. mUA
have a niimher nf fusifurm espanainnii along their courw: in lh« lar^
eipanaiou is fuuiid an uval nucteu*. The tcrniiiiul GUnieaUi an calM lb*
el/aetory relU. As yet im ouiineiUiun hftwMn the Mibeiillbdiml and iat«npt
tbelial iicrve-filanieiiid hait lioeiidtiriionitrAMr!. Thuepithutiul C4:ll«(if toAt)
in tli^ above figure are «h(>wii crxinucTU'i] with the itubcpilhelial liitfue. YW
fillh iiiTvc siippHui the truau: with ni-ii^ury filnmciil*.]
Odorous pHTtieW pitaent in Uiu laipired air pMting throuch th« Inmr
lineal cliaiiihi^n (liH\i:>e into th« uppur nasal chnmbws, and lafling i>b ibt
olfiiutiir)- fpiihi-lioto pnidiice nnmrr iuipuIsM whieb, ascending to Ihc brain,
give ri'c In nctiwilicnis of son-It. Wt- rnuv pn>4iime thai the wnsorv iFiipulm
arv ori^iiiulvd hy the cuntiu't of tlie nitDroiis piirlicW with tb? peiniliar n^-
nhaficd iilfiielory ot'lla di^crihi-d by Max SchiilUe ; btit wo arc ad maeh ill
the durk about this mutter as ah iiit tho development of risual scoaory im-
]>ul*ea in the roit» and cones or of auditorv sensory irapuIsM in the organ of
Corti.
The »tih*idTnry appuratns of smell is exceedingly meagre. Br the (onwd
nasal inspi ration , called siiiHing, we draw air so fitrtibly ihroiigh the nostrih
that ciim'tits pans up into the upper rh well as the lower naaal chamber*
and tbuM a more complete contact of the odorous partiotes with ibe olfaoiorj
membrane than that nupplicd by mere diffoBion \t provided for.
Wc bave ever}- reason lo think that any Btimulus applied to the olfactor;
9VRI.t.
955
nerve «!!! ppMliice t\>e seitRHliim of aniell : but the proof of this !■ not to
ir M ill the caae of tW ojMit- lad nadilurr iictvm. Wv are. hoirever, sub-
ject to ■enaatioM of wnell nnl cautM^t Uy objei^tiTC rxlon. I1ie olfaclory
inembniiM it ibo onir part of tlin bi»(y in which mlon an iiuch can give rise
to any *«nntio»s: am) thit iM>n!4iliiin« to which t)i<-r pve nae nr« nlwajs
ikoaf! of irtndl. The miicouti incmhrnnc n( the. niiie i*. howerer, also do
tlDHnim«nt for the (level opnif^nt of nflirrmt impulxM other than the specific
otfllclofy ()fm^ Chrnicnl utimiilation orthfi nlfni*ton' merabrnae by pungeut
subaiancca such it* Bfiitniiniu cives riar to aienMition (itnlinct fn>in that of
flnall, iMiiHilion which nfrorr)* us no informnii'-n c<>nc<Tiiing th« ehcmical
nalun&T lh« stimiilus, and which is ini)istinK>ii*hnblF fmm tli<- MmutioBS
producc«l by chemicnl ^limtitniion i>f mhcr pari* "f tli»! muwl inembmne as
well a»of other Biirfacp* enuHllyseiisiiiveloehemk-nl sclioti. It is jimhable
Ihat thcM Ivn kinds of sonmi'iiiB thus arisinc <n the olfactory- mcnibnioe
are conveyed by iliUi-n-iit ncrvca, IIm former by the ollnclory, the latier by
Ibe fifth ncr^T.
For the tlevelopment of bouAI it appeflrs necessary lhat the odorou* par-
ticles should be conveyed lo the uasal merobrane in a (gaseous medium, nr at
teaat that the surface of rhe membrane should not be exposed nt the same
lime to the action of floids. Tlina. when the nostril ia filled with rosewaler,
the odor of roses is not perceived ; and simply Gllinjt the noetrils with dis-
tilled water suspends fur a time all atnell, the sense reluminf; jfraduatly after
the water has been removed ; the water apparently aoU iDJuriously on the
delicate olfactory cells.
Kach substance that we smell causes a speeiBc Mnsatlon, and we are not
nul^ able to reoo;;uize a miiltittide of distinet odurii, but a]m> to distinguish
individual odora in a mixed «mel1.
A« in the previmi» sciiitcit, we pn^jccl our Rcn«ation inia th« external wr>rM ;
the Stncll ap|ican to Ih.- not in ikur ikhh;, but Snmnwherv outside ux. We can
Judm of itif ponilion of the mlur, however, even lesw definitely that we ean
of liiat of a sound.
The Miwatlon laka some time to develop after the contact of the slimtilui
with tlie olfactory membrane, and may Inst very long. When the stimulus
is rrpeaicil the scnwilon very S(«>n dies out ; the sensory termtnnl organ*
spM^fily Iwcorae exhausted. Menta) anraeiations cluster more strongly niunil
sensations of smell than ruuiiil any other impressioas we receive from without.
And rrHex cllbct* arc very frcfjiicut. manv people fainting in coiKojucuec of
the contact of a few odoniiis paKiclcs with their olfactory cells.
Apparently the Inrgrr the surface the more intense the aenaation : animals
with neule •cent hiiving a proportionately large area of olfactory monibrano.
The quantity of maleriul re<iiiired to produce an ollnctory senaation may be.
M in the case of mudk. nlmivt immeasurably small.
When two different odors ore presented to the two nostrils, an oecillatiftO
of sensation similar to that spoken of in binocular vision (p. 9^7) takw
place.
The assertion that the oUiictory nerve is the nerve of smell baa besn db-
puled. Cases have been recorded of penous who appeared to have poaseMttd
the sense uf smell, and yet in whom the olfactory lobee were found after
death to be abtent, Direct experiments on animals, however, show that )oa>
uf the olfactory lobea entails loss of smell. On the other hand, it is stateil
lhat section or injury of the fifth nerve causes a hiu of smell though the
ol&etory nerve remains intact ; but in ihcee Maes it has uot t>cen shown that
the olfactory membrune remains intact, and it is <iuit« poaaible that, as in
the case of the eye, changes may uike place in the nasal meaibrane as tho
retull of the injury Ui iliv Uflh nerve, Mifiicieut to prevent its performing its
usual funcliuHs.
reabin'g, shell, attd taste.
Taste.
[nt/swlo<ficat AnaUnni/ of th« -Oiulalorif iluoaiu Jlemtnitt,
The peripheral organs conceniod in tlic »cti»o of Mate tn locklbnl in Um
mucous Bwrnbraoe cohering tlKnIursum oftlic tongue, tlw ftuc«>.wft pdau.
kimI uvnifl, aod poaeiblj'ti portion of the iippor putt of the phurynx. Tlii<
tncmbrane is analoeous in Mructurc to other membmnea of it* typv, «x«cfl|
OD Ute doreuiu uf tbe tongue, where )t« rtritcUiro i* limilar to that of tit
integument. At this position it cod»*i« of ii eorium, with a pupilhrj and a
BUperiiciai epitheUid lat/er,
PIC.2U.
VnncAi. soman or not Qacvmrtu^im fjinu-i. iProm Kolukck )
A, the iMltllli 1 fi. tbc tumunilhis vrati: (i.lbac|ilIheUalcuv«1nRi A, llieiiorrca u((W |«|4Uiuil
wiUl «|inwilliiu lowuil tiM luiftica ; r. tha ■NODdaiy inplllir. y>.
The Klructurc of tlie ooritim is nmilar to iJtnl of the Mn, bgl i» tJiinan
mid IcM compact. It mrvea as a point of insertiuo of tbe inubcular fibre* of
tbe tongue.
The pojpUla are thicklj* distributed over the whole donnl surface, bet
more part tell I nrly marki^d in the anterior two-thinls. Thcr project M mtnMc
pRMUinencce, wliich jjive tbe tongue a rougheDed. charactL-ristic appearaaoa
Plil.311.
iTs^
Manei jii(i> SccnoK oe Tn> PrxoiriMii """i », (Vmui Kumikkk, ■ml nfUr Tom tart
tklWMJIV.I
J. th« (irrfltn or > ruDdflmn (aplllji. imnlally dcnudnl M )(• ndUHdluo, V ; a, i(<l<liaUnii. J
■!r<lanDrBfuiiNtfijrmiitiiniB«ri(bllHtUooit<n«Mtaln]aaUdio.uMT; r. Ma i r, mfUImt )«•■ •«
iluido |a|4lljc In liig nsiKbbartaa ■trncturaar Uto tDagua.
The |>apillni are of two Icindf, tbo timpU and eompound. The nimpU paptllc
are aiinilar to tiinse found in the akin; tliey are found GCOttereil over the
whole dorsal surface, between the compound pa{Hll». Tbej an bum
numerous in the poatortor portion of the oi^an. fne eompound papjllm an
957
of thiTc vnrieli«i>; the pnpilln- maximii: or circumvalliitje, (be pnpilln modin
or ftiDgifonDra, ntii) (he [inpillif minintn.' or filifurtim.
The papula rimimrfillatrr ( Fig. WJ t, which arc the lnrpr»l. »re «btuil
«i^hl or ten in number ttnd form n V-ehapcd row nt llic junction of the
middle and posterior two-thirds of the tongue. Thev consist of a central,
bnxid papilla, surrounded bjr an annular rins or wall of about the Mmv
deration, and se;>nraled from tbe centre impiMn by a circular liwiire. The
cCQiral papilla, aa well as tbe eurroundine wall, is covered b_v siniplo papilliw.
Each of tneni re>ceives one or more cupilTarv loops and nerve-filnmeDta.
The papillix funyihrmtr (Fig. '253) are found principally on the tip and
sides of the tongue, allhouf-h srallered fparwlyover the whole of the anterior
two thirds. These are so named from ihetr fungiform shape, being espandcil
at llieir free eslreniity and projecting on a sliori, thick pedicle, Thev artt
vuTered by simple papille, and contain plexuses of TeaeeU and nerves.
a h A r a
4I, TWHOftl MMlua n««r the iiiltUio uf Uir dnnBl tutkciof Owtotifao: n.a, ftmgtfcfm ia)4llB:
h, tOXttna fBMIbt. with ikrii t>*ir lltr pnt-om, <,riiiinvoBWdeprl*«d of umi •pUbenuD : me-
MBld 3 OlMftftn*; S. aiininn (»id[iuudi1 |a|41lE^ a, mittrf . r. vtiD; e, aplllu; loofBOf Iba
•MOnduTiav"'*- fr. Un»orbueiuciil-iDCiiilirBDt-.(l,M004i>l(rTP''l>">»^<le|>ii*«dsff.(. Iblf^ltll*-
■an i /. I)■l^llka pmw«i or •(dltidtum a*fi<tiis Ui» dinpls |M[riiln, uuwDlllal 3» dluuctm : f.
M|anMdaa(iteudp«nl«k>ot«tlibotluiu,pu«fitlM3CO<lluii«Mn. LXbatnAmndoatMiurflire
of UM loiuM : 1, m, andii of bali-llk* «|4I&«1M procfM. tkamlmt *wMIn la lb* iMtrtcMad
ainDE*inent 0/ tb* laiUeha. bill la 4II • aMt««MUB erf llM pMOehi towitd ibapttnltXMHMSa
KiH hair, roainlbed IWi1luii«tcn~ (AflsrlVDoaDdBciwMtV.I
The papilla filiforme* (Fig, 254) are by far tbe mctt nnmeroua, and are
found thickly distributed ove^ the entire surface of tbe anterior two-thirds of
th« tongue. They are minute, conical in aha[)e, autl generally arranged in
nEAntKO, ByBLL, Airs TADTB.
bii)eniiirorin rows, which are more or less panillel wilh the two ronf of
rapilln.' circuDivalIa(a>. Tlieir free eurface is covered "ilk »ini[ili? impillc
The cjiklieUum coverinj; them ia greatly modilied, aad afijinire in ihc for«
of hnir-like procefees. (Fi^. 254.) These procetwa Hr« bathed in aiani^
arc iiioval>le. and have a geiii;ral ioclinalion (miDtiOK backward. The eiaiU
enceof ihfce hair-like pr»ret»cB oil the fllilorm papilla> luggeetA that ikii
variety of impillie is iniiumtely cotineoted with the taotile aeasibility of ibt
tongue, a»tl not with gu^lnii'^ii. In caroivora and herbivon theDc proonm
are of a horny structure, and iterforiu an active fuuctioa in ilio attriiKia nml
preluntiioii ol fund. In man iheir iiipecial functiun appears lhrmi((fa ihtir
inliniairr (sinocciiou with the tactile aca»e, to Kuid^ the tuiij^ue in ita vuiaMi
and coiiii'Iicate') inovetnMits.
Thi: uitininte (enniiintiouii of the gu»iatury nerves are yet envelvpeil il
olneurity. According ti> Kngulmann, the kI<wo- pharyngeal nervea IcnniHCt
in flaek-shapcd orgnns. which are tcrim-d tUc gwtntnry ttuthn or Intte bndt
(Fig. 235.) ThcH- hulbs are found pHncipiilly in the pMpillnry stirftcvaf
the vrall of ihu circumvallHlr pnpillic. They arc aUa found in the fuDglAni
Flo. 3iS.
^
On»iONT Bi^lM rsoH iiii l-inH*i. UtnAToRv Oants or mi lunHr.
pnpilla), but are Icse niimerDiiB. They coimiat of a fla«k-ahaiie<l fundus, which
r«eU upon the »ubepithe]itd li^Bue, and a mouth w hich opens upon the sutftre
of the mucous tnemhranc. The mouth ia known at the guHaiory ftore. Tbe
fundus of the flask is computed nf two vnrietHS of cells; the outer ur
iiivetiiti^ eetU are fusiforDi, nucleated, and eranular, plnec*! nnmlM ud
arranged conrcntricnlly in a direction tram Uic haw to the neck : iticy Um
form a wall which cncloees elongated niicleuied ccIIh with filainenti<u» jm-
MflKx, which extend through the gustatory pore and prnjert an very Kntty*
pointed or truncated extremities. Theae inner cell* are calleil the gndaiaij
tetls, and are supposed to he the essential terminal elements concvned b
gustation. Their relation to thegii-italory nerven has not as yet )K<en ckarif
demonstrated, but they are evidently connected with (he ganglionic pIcxoM
of nerve-tibres at the papillary Im.-M's. The gusiaiory nerves are nlsu lup-
poeed to tefiiiinale in the epithelium of the papilln.]
The word taste is fre^nenlly used whcii the word smell ought U> be cs-
ployed. We sjieak of " taiiiing" i^ihirifvroiis riibitiancni, siic}i as an onwi.
wines, etc.. when in reality we only smell them as wo hold them in iwr
mouth; this is proved by the fact 'thai the so cnllcd tuetr nf thctv thiap
is lost when the nose is held, or the nasal membrane rendered inert liys
catairh.
The terminal organs of tlu wiue of taate thus more etriotly d«finei) *n
' «n(lin((S At lh« glMao-pburyn^fll iiiid liiigiiiil mrvM tn Itie tniico4ia luem-
bniMtof tkc tongue and paluli;, iIickh; nn\n srrviiig ni tliu ttiie^'tal ncrva
of iMtC \Vh«t)i«r llie Ki-callftl gutlatory budii cut) be r«giiMi-<l tUBjiixrlllo
orglMof liut« npiican (Inublfiil. Thv tiibuMiiry njipiiniluk u cnntiiicd to
ll>i' toiigiic mill li)>ii, uliicli by thi'ir movonicnin tisrut in bringing tbe Mt|itil
, ■iibiiliinnw into nniitnct with l)i« inucou) nivmbntDe "f lh« mouth.
f Tbougli uc can hnrdly be mid 10 project our »en»atiun of tanic into the
Cllernil worKl, vrc uMign to It do »ulij«ciivc lornlixalioD. When nc piMe
<)uiuiu)' in cMir month, llie rrculting •euiNitJon of tiiNte givM ns no intbrniiitiail
■a to wlK-ro the (|uinin« U, though wc mtiy Icnrn thnt by oonconiitant gcneiml
Momtion* aming in the buccal mucous membrane.
IWeVccugniic a niultitudr of diiitiuct Instc, which may be broiullj' claiw-
fied into acid, Mnline, hitler, nn<i sweet tniilM. .Sii|>id subttaiiceH have the
■tower of producing thwe M-nsntions by virtue "t tlieir chemicnl nuture.
but olber HiiHuli will iilso giw rise to aenifnliona of inste. When the tongue
» tapped, a tnoie ii> lelt; anil when a eonatjint current is piimed through the
luoutb, an alkaline or, in «onie perwns, a hiltcr metnllic tusle is di:velo{>e(l
wbcn ih*- an<xle, and ao add taste when the kathode, u plnce<) un the tongue.
It it prohuhto (hiit in theee cases the terniinnl organs ure iiidir*.-i-tlr utlcc-ted
by the ciirrcnl. Whi-n hot or pungent sub:<tuncea lire intmdue<nl into the
muulh, scoHitioni' of general feeling nre exciiiit, wh)<'h oWurc miy strictly
gtuUlory •ensatioDs which may bo prescot at the siinic lime.
'iltough analogy would leitd ns lo supposv thnt a siimulus applied to any
part of the course of the reni gustatory h'brcs of cither the glomt pharyngeal
or lingiial nerves iruuld ^ive rite to a sensiitiun of tiute and nothing else,
the proof is not forthcoming; since both these nerves ore mixed nerves con-
taining other alferent libm as well as thi»e of taste.
' When the constant current is used u a meuiB ot exciting taste, gustatory
aensaliuns are found to be developed in the back, edges, and tip of the
tongue, the wifi palule, tlie anterior pillur of the fauces, und n email tract
of the jxNilerior part of the hard palute. They are nb»eul IWim the anterior
and middle durval, uud under aurlaoe of ih« tongue, the front portion of the
hani puliile, the [)C6terior oiilan of the fuueea, the gums, anil the lipa.
t^ajiid BuliBlAUcca are unsuitable as a test lor thii purpoite, ou aei^ount of their
rapid itilluiiiio. Bitter sulMtonces pruditce imi^t eflcet when pliiced on the
back of. and sweet !<ul>dt«nce]i when placed on the tip of, the tonjfLic ; but the
(■Mintf ]x>wer of the tip of the tongue variiw very much in dilft'ruiil iudi*
viduai.i, and in iimny Mreou aliiiuit entirely HhM'nt. It in mid lluit iicid* nro
best npprtdni'il by the edge of llie tenguc.
It w iMxniiMl for the ilerelopment ortaste thnt the Mihtlnnce to be tasted
should lie ilixnilved, nnd the effect U Incrcnacd hy rHclion. The larger (he
surl'nee (he more in|t-ii»c the Mamtlon. The scnsniion lakes some lime t^j
develop, and endures for a loog time, though this may be in part due to tiie
stimulus remaining in ooniaet with the terminal orgniw. A temperaluns of
about W is tlic one nio>t favorable fur the prvduclion of ihe sensntion. At
temperatum much above or Wlow this, tatte is much impaired. The nerves
of taste are, as we have t«i<l, the glosao- pharyngeal and the lingual or gus-
tatory. The former supplies the back of the tongue, and section of it
destroys taste in that regiin). The latter is dt»tribtiled to the front of the
tuogue, and section of it similarly <leprivce the tip of the tongue of taale.
There is no reason for doubting thot the gustatory fibres in Ihe gloaso-
pharyngeal arc prowr fibres of that nerve : but it has been urged by nuuir
that the Kunlaiory nbres of the lingual arc derived from the chorda lympani,
and that those fihrea of the lingual which come from the tifth are eniployeil
exclusively in the sensotions of touch and leoliug ; the evidence in lavor of
this view M, however, inconcluiive.
CHAPTER V
FEELISa AND TOUCH.
GeXSRAL 6&N8IBir.rTY AM> TaCTILK pRItCKPTtOXS.
Wk have ukcn the foTcgalne sense* Bret in t]i« ordsr of dtKiUMA «
avcounl of ihcir being ominentlj 8[iccilic Tho tyn givM u> onlr vimi
muatioDs, ihc cnr odI y auditory scnsutione. The Mitntioas lire pruduocd io
each CUM by H|wciJic xtimuli : the eye is only ntTcctod by light and tlwaj
only by sound. Moreover, ibc informtitioii they iitTonl us is c«>ii(ia«*l totk
extomal world ; they tell us nothing iibout oiireelvt«. Tbe tkHoim tjimI
wnntions tvhich ariee in our retiua are referred by us not to tho retina iuelC
but to some real or imnginnry objefl in the world without (including m pan
of the cxlernnl world euch jKiritons of our own bodi« as are vi.-qble to ow-
tdvn). .Sueh, nlao, with diminishing pnecisioii is the iofonnalioD gainol iij
boring, taEl«, and smell.
All the other ailereut nerves of the body, ronlripetal impiiUci along w\dtk
an) able to aOcct our conscioueuesi, are the means of conveyiog to us is-
lormatioo concerning ourselves. The aenHilione, arising in them from tbn
action of varioue stimuli, are referred by ub to apprupriate parts of oar
own body. When auv l>ody conies in contact with our linger, we know tlut
it is our liug«r which Iiaji been loucheii ; from tbe resultant seoaotinn we ad
only learu the existence of oertain ({ualiliea in the object touched, but n
also are led Ui vounect the oognitaaou of theae <iualiUca with a particular
part of our own body.
Like tbe inure sjwi'itjc *eii»cii previou»1y studic<l, (he Aeiuationa of wbid
we are aav tpeiikiu^, and which may be referred to under tlte namt «f
touch, using tlmt wunl for llie prevent in a wide mtsuiing, requlr* for thdr
produdlou terniinal urpmx ; ami the chief but not excluttve ornui of toad
IS to b« foun<l in the epuli^rmin of the nkiu nud cvrlain underlying Danrooi
Structure!). F«r ihe ilrvelDpmcut of sjn'cific tactile sciMttionjt the*o tcfsiinsl
OlfianB arc a* cwenlial as arc the t4-rniinal orvnns of tbe eye for sight orrf
the car for hearing. Cimtuct of the «kiu with a hard or with n hot boJf
gives rwv to a distinct sensation, wh<;rrl>y we rcco^isc tbitt <r<^ have taucbfd
n bard or a hot body. But (be a])plication of cither body or uf any o(bn
stimulus to A ncn-c-lrunk gives rise to a ««isatiun uf getural frelin} oalf,
cornviiutiding lo the Mniplc sensation uf light whtch is pro4luc6tl by dinci
stiinulati<m of tho optic nerve. Wo have no more tartilr- prnxption iifi
body which is in contai't with a ncrvc-lrunk Ihan we could have i'muo/ ptf-
CfptioH of any luminous obj<ct, ibe rays proceeding from which were «Uvng
enough to excite sensory impulses when directed on to the opiic nerve is-
stend of on to tlie retina, suppusinj^ such a thing to be pomblo. It is furUwt
ebarncteristic of these ordintiry nerves of general feeling, that the aeiuatiaM
csusrd by any stimulation of them beyond a certain degreo devdop thai
slate of conscioueneM which we are in tbe habit of u[)eaking of as " paio.'
Putting a^ide the general feeling which many paru of the eye pnHiA,a
very strong luminous stimulation of the retina is rei|uire<l li> prt>duce >
eenaatioQ of pain, if indeed it can be at all brought about ; wbere'ju a vtr;
TACTlI.lt SKNSATIONS.
9rtl
I
I
moderate stiinalMMn or lh« Bkio, and itlnxwt every BliniuUtioD of an ordi-
aarr nerre-troiik, U said bv us to be {xiinful.
1* IioiikI) iho skin is the clivef on(Sn of touch, the mucous membrane lining
ibi; vnriuus paBBAgeaof lb« b<nli|- also serves ss an iDsLinneDl for (he same
teti»e, but onlv ^rashutl distance frnm the re6|)eclive orilices. We can
re<.-<>f>nii« harO or hot bodies with our lijia «r moulh, but a hot liijuid, nben
it ba« reaobt^I the o»ophaaus or atoinacn, umply sivea rise to a wn«atiou of
pain : wc i-aimot distinguisjt the seou^on caund by it from the seasHtion
cnutii-d by a draught of a luu acid fluid.
Ffini |>ar(M aud tiiwtiet of the iKidy other than the akin aud tlie portions
of RiucoiiK ititimhraiie juat uicnliotied wc liave ubacure avuMlitina of Keuerol
tircliiig. by which we are mtuit: vaguely aware of the geiiiTal coiiditJim of
our ImmIv, though our jud^menta in thin matter are chiefly iuAui-nced by vhat
me aliall have to vpoalt of directly as a muiu'ular !>ense. In all |iarl« of the
body, however, on uct'aaioiii* all ton fntfueiit, thin gem-ral fcelintf may bccoma
promiDCat aj> pain.
Tb« stimuli which, irheu nppli<.-d to ihenkin, give riiw to tactile percO]>lJott6
are of two kinds only: (1) inechaDical. that in. the contact of bodies exert-
ing varying degrees of prraiurv: and (21 thermal, i. c, the misiiig or lowar-
ing of the temperature of the skin by the anprDarh or conlnct of hot or coltl
belies. Wc can judge of (he weight and of Ihc (emporaturo of a body.
because i>c can. through touch, pefxeive how much it precHS when alJoirod
to reft on our skin or hoK hot it is. But wc can through touch derive no
other nerceptiong ond form no other judgments. An electric shock sent
througli ihc «kiii will give rise to a scosatiDU, but the sensation is an iiidefi-
Dite one, because the elcclHc current ads not on the terminal organs of
touch, but on the fine nerve- branches of the skin. We cannot distinguish
the sensation so caused from a mechanical prick of siniilar ioteusitr, wo
OUlDut perceive that the tennatiou Ls caused by an electric carreot. ^imi-
krly ocrtain chemical stil>nlanc«>, tuch ai> a strung acid, will give ri»e to a
•HMUioD, but we cannot [tcnvive the aci<l, nc can form ui> judgniciit of its
BUUre Aich as we could If «e ta.ilc<t it; and if the add dutv not twrraealo
the skin tn an to act directly and chemically on the fttni iivrvo -ubrat. wo
cannot distingiiivh the acid Croni any other liquid giving riMi to the samo
aiinplc contact impnwou*. The terminal organs of (he skin are sudi ••
are nnly nUbcled by j>rr0ure or by temiMTnturc. Conversely, preamire or s
varinliou in t'nipeniiuri- brought to l>e«r ou a nerve-trunk, instead of on (bo
terminal crgauo. produce* no >jireific tactile sensations of pressure or tcm|)cr-
ature, but merely general tensations of levling rapidly rising into pain.
TaCTII.K t^KKSATIONlt,
Setitativtu of Pnttvre.
■ As with visual, so with tactile, and, indeed, with all other aensattoDS, the
I intensity of the Mtmilion luaimaina that general reladou to the intensity of
■ the etimuluB which ue mike of at i>. 916 as being fonnulaled under Webtr'a
P law. We ran di.'tinguisb the diflerence of preieure IxHneeu one and t«o
grmtamt* as rvadily a* we can that between ten and twenty or one hundred
and two hundnxl.
Whrn two wnvalionx fnlhiw each other in the Mme apot at a tufficically
short interval, (bey arc fumil into one : ihiiK, if the ItiiKer be brought to bear
lightly on a rotating cnnl having a serirs of holes in it. ibe botes Mas* to be
wit a« «uch whi-ii liicy foiloa each other at a ra|>idiiy of about 1600 in a
81
962
riBLiiTG AXD Torcn.
sooood. Tltc vibrnlioiu uf ■ Cird ociuw In bi; a|>(in.-oiHbt« bv l4iucli min
Uief reach tli« snini.- ranHitr. Wbcu wiuniiMw ure generalpd ■! txiiuu nf
Ike pkiii bio clone t<igctni><r Uiey booomo fuMd into one; l>ut Ui tbU (wiiu *•
shall rctuni pnwmtly.
Tbe mnsniion cnusod bv prauun Ja at its innxiniunt sooii iift«r iu btfig*
Ding, fttid tbencclorward tfiminiibes. 11i« mnrv middi-iilf tliv pnaMira b W
crcaeed, tbc grenler llio senuttion ; and il' I)k' incrnwe hn sulEcieotlir gndual,
even very great preMittv timy be applied n-ith'viit givinc riaa Co anjTMiHi'
tiou. A genealioii in any e|H>t is increased by contrast WMR Um iiarraUB£a|
arena are nut eubjcicl lo prcMurc Thus, if the linger be dipp*-)] inti nut-
cury. the pmsuro will be felt mo»t at the turfaoe of the Huid : and if iIm
finger be drawn up and down, the Benaation caused will be that of a riii|
ingviug along the linger.
All parla of the akin are not equally positive lo preamre ; email diflrr-
encea uf simple preaiure are more readily appreciated wlien brouf;bi (a bear
on the jialniar i>urfac.-v <>{ the finger, or on tiie tVtreheiai). than oo ilip anoot
on tbe Hide uf the fool. In makittg ibeM deterniinalioiu, all luuwular man-
nouti sliould be avoided in order tu eliminaii; tbe muscular aeaae, of wkieb
wc Mhnll (peak pr««iilly: and tbr area Nlimubilud «hi>nld be aa arnall and
tbe anrlatx* in coniacl il- uniform an pi'vible. Iu a nimilnr manner, vnall,
conwcutivc varinli'His of jirawuri:, il> in wnnting a pulw, are imin: nadily
apprrcinUHl W c<:rtuin part* of the ^kia ihan by ocbeni : and Uie minimnm of
prcmnrr whiuli can he Ml difliini in ilittercut pari*. In all casu, vnristtMM
of prrwiirr nrv niorr ciwily ilixtingiiiithvx) when ihry arp nacocmTD tluui whni
they arc MRiitltandouB.
Sriifatlfint of Tirmparatun.
When the tcmpenture of the skin i» raised or lnirere<i in anjr nioi. w
riM-cive aeiiHatioiu of beat and cobl respectively ; ami by tbeac MimiioM «f
the tiMnperalure of our own skin, we fiirm judgments of tlio lcin|>onUurv of
b'fdicH in wutaet with it. llotijes of exactly tne tuiine teni|>eTmture ai tlit
region of tbe akiu l» nhicb they are applied produce no duoh thrrmal seiua-
tionit, thougb ae can, from tbe very aDsenee of aenMttioDs, l<>rm a judgOMt
US to their temperature ; and good oonductora of hi^t appear ra^ie«tinly
butter and culder than bad condiictoni rattted to tlio sanu) teraperstur«.
Wc may consider tbe tikiu ai having at any given time and in any ([irea
Spot a normal temperature at which tlu- wunaliou of temporature is at tarn;
fot under ordinarv c-ircunistanceo wc ar*- not dirodly contcious of thr !*••
peralurc ol' our skin ; it t« only wht^n the normal temperature at the met il
raiaed or lowerod that wo have n winjuitioii of beat or cold napedinly.
This normal tcra[)eraturo may bo at (he Muao time dilTerenl iu diflVrent tmM
of the body ; ihus, at a time when DCJlhor the forehead n'>r the band art
ainog riso to any sensation of t«mperaturo. we may, by pauiug tbe hand to
Aefwebead, fro^uctHly Iwl the fjrmcr hoc or cold bdcau^e the nirmul (am-
pentures of the two parts diflor. The iionnal teniporalure iu unr vpat oitf
alao vary from time ti) lime. Thus, when iha hand b placed m a wars
medium fiir soms time, the aensation of wnttnih cea4«i; a turn nunnal urn-
peratane it esublisbed with thfl sero of sen<aiiuQ at a higber lavaL a da|xw-
tfOD or elevaiiou of thia iieir temperature giving rite, howerur, •» It^f..:
aensatious of heal and cold respectively. Thai it is tlie changei) cm It;, m
and not tbe obanjjre itself, of which we are c»uKii)o«, u ahown by tho (tei
that when a portion of tbe skiu i« cooled, by brief coninct with a c^ild -t: ■ "
ibr liiManoe we are still conscious of the xpot being cobl aftitr iW "
remove
ptn^ li*-uiei] fay ili« Bummmling wanner tlwuet,
Tbe (.-liaii^re iu leiu|>eriuurv of (lie Kkin neecaiBrT lo produce k teiMtUoD
miM have a certain rapiililr; uml ih« iiiDre gm^ual the chnn^ Ui« k
{■itciiKC Uie «en»uti'>ii. T)i« refieiitvil ili|i|>iiii; of tho liand iiilo hut n'ii(>.-r
prwlueci n jircater neitMti'ni tliun vrlii>ii ihc liimd ir iillowwl k> irinain nil
ihc tini« in llie wnirr, tltougli in tin* liittvr curt! ihe teiii|HTitltirv nf (lie 'kiu
i» Riiwt atTrdcd. The ciTccu uf oouLriut urc also «ucd in titiMi (Cimtiutu m
in tbiDipof (tremiirA.
Wf can nith Mitiii- nceumcy diitinguisli varinlions of lcni|WratuT«. npo-
dally ihiwc lyi»eiii;urthe ii»rmal leinpsratiireof tlir ekin. Thno wnrntiuDi,
in fact, folliivt Wt'hiir'* luir, ilioueh apmranlly Mtieatioiu of slight culd )ir«
mutv virid than tlio«<' of >>liglit nmt, the range i>( taixt m-cunitc KiMUtion
<t>tuitig to lie botwom 'JT ' luid 3-1''.
The regions of the »kin nuwl ttcimitive to variations in t«mperutun< nro not
iticHl with {ho*v m<>st '"nsilive tovariationt in pnmiire. TIiub the chrrks,
vlids. tcniptcs nnd lips iirv mora muitivc than ino hands- The Icael sensi-
tive parts are th« \v^. iind front iind bni-k of the trunk.
Tm simplest view which can be taken with re;^ril to the distinction be-
tween prcMuni wnnlions nml lemnenilure aeueatioos, and wliicli is sug-
peted Dir ihe fiicta just nidiliikncHt. u to suppose that two di^lincl kia<U of
t«r'iiinal orj^am exist in the skin, one »( which is atlect^d only by prepare,
and the other only by viirialious in teniiK-ruliire: and that the two kiiub of
»|wripheni) organs are ronnected with dilTereiil piirla of the central sensory
or};aii> by scimrate nervelibres, CertiiiD pnthu1»givHl coses liave been
JnottA fti showing not only that tlua is the case, but that the two seta of
brta pursue diflereiit courses iu the spinal cord, 'ilius in oertiiin diseases
or iDturies lo the brain or spinal cciH, hrpenettbcsia as rt^nb tein|)ernt(ire
has ncen nb*erred unaec'>in|wtiied \tv an nugmi-ntation of avniitiri-tieM to
propnire ; and, conwwelv, inHtnnora have Ikth wen where the patient could
tell when he w«.t t/iucbcnl, bill could not diatingiiinh between hot aiul cot^
On tho other iiaiid. then- arc faclJf which «how a clow dcjM'ndcJici- heln-caa
I 111' -I ii-ni .iiiii of prr-.'Oiiire and tcin|)criiturri. When each <ftiniiili(f h brought
1 1 I' 1' 1 .1 vnry liiiiiuil urcn.tht two scn«ntiun« are frecpicnlly confouiiji'd,
a|>cclitlly 111 ihnce regions of the body whom sensalions are not n<rutv. Ho,
alo, II [icnny eoole<) down nearly to zero, and placed on the Ihrchcad, will
lie jiidgnl t>y most people to be ns hoavv or even heavier thnn two penniee
of the tt'mponttiirc of the forehead itseff; and. convcr«i^ly, ti body warmer
than Uic skin will often nppear heavier than a budy of the same weight, but
of the snmv lempcrvturo as the skin. Storvivcr. cases have been recorded
where a hoi body, such m a healcil spoon, was felt, though the applicctjon of
the same spoon at the temponiture of the body produced no aeasattona, and
yet the lieatod spiion was not recogniied M a tut tiody, but anpeared to be
simply something touching the skin. It may be argued ibal tneee initancos
■how nothing more than tbc changes in the akiu. whatever tbey be, which
riv« ria* (o sensations of pmsuiv, are modified by the temperature of the
Ain for the linie l)eiiig. whereby the judgment as lu the prewure which is
lieiog eiertc<I is rendered faulty ; hul ihey may also be taken to indicate
thai variations in pressure and tcmfteratiire nfiect the same leriniiisl orgaw,
and the same nurvc-Ubrcs, though slTi-clin;; them iu a ilifTfreni wav. and
generatiui; ucrvoits impulMM so far different ihat they give rise to di^erant
•eiMUiuus. And we may hert- note ihnt <re certainly cannot speak of nerves
of warmth in the same seiL:r« iu which we sneak of nerves of sight or of
iag. A «*.imulus (of whatever kind) applied to an optic or auditory*
t. If a<lu>|iinta, give:i rise, lu we have seen, to a scuiuition of li^ht or of
FEELIN'O ASD TOOCH.
wund : ft Blimulus, oii tlio oiher hniiil, npgilinl ut th« trunk of * cuUmoui
iwrvp, ipvee riw oiilv to gcnrnil livliiig i>r pain ; tltmigh ihr nerro etruiiilf
cunuins tibres by uliich ^cnnitions of pnwtirv imil of lc»)|>rraitini nwli iw
brsio, the general feeling which i-timulniion of tlxi trunk causw u akin
n«itli«r to aensntioDs of pnrssiiro nor to tboae uf warmth.
The nipidily with which hot or col<l bodiis brought into c'ootact villi tW
skin Kive riite lo Mosnlions of tt^nipcmture, Higgesia (Iml the terniinnl ■{>pi'
raiUB fur ^neraliiii,' thetie ^onNition^, irhnb-Ter h« ita nauirv, is pliioed iii the
epi([«rmiH. Mid indeed as iiettr ns powibic to the siirfnoo. I'nvure, on iht
other hand, (.-ud be rendily trna^iiiilled through even a thick layer of ikk.
And thoite who nmiiilniu the esistenoe of different lenuiual oiKUU tat pirn-
Bureaod le[n{>er&tu re, regard ihe uerreendiof^ in the epidermic a^ the Utirr,
and the eorpu»cula tacltid, en<l't>u IIm, and allied ortpuutte the foru>er. Bwt
tiw evidence we poaaew concerning thia loatler is at protcnt iuooncluiive.
TAcrn.B Perckptioks asd JuixiiiRiiTS.
When a body pruMM on any part of our skin, or when th« t«oiporatiu<i oT
tbo skill at that spot is raiMxi, wu uro not nuly codmIous of prtvuiv « rf
beat, bill perceive that a purtii^ular part of mir body hn« b«eo touched nr
htatcd. W« refer the lU'nuaiiniii* to their nhuK of uri^iii, uod we thia tw
touch perceive ihe rtlalionn to mireelve* of tlic hodv whifh ^Ivn rise tn tlw
tactile #eii«alions, iti the ^nmc way m» io our vi«ual |>er4<eplion of extMnal
objei'ts wc refer Ki exleni.il niidire ihcienwlioiu origiiuitiue in certain parti
of the retina. Wheu wc nre touched on the finger and on llie l>itck wc nlrf
the sensations to the tiuger and to the back respectively, and when wt ai*
touched ut two platen on the tuime liuger at th« same time we r«fer thea^^
aationa to two iiiiin<» of Ibe finijer. In thii; way we caii localise our >'^^fl
tions, and are tnus assisted in perceiviu;; iJie epaco relations of objects inlf
which we come in euntacl. "
This l>f>wer of localiuuK prendre sensations varies in diSkreat paiu uf th*
body. The following table, Jkini Weber, gives the distaiicv at which in
points of H pairof conipa.ises nmsl be held apart, so that when the twopotnti
arc in contjict with the »kin ihe two consequent seiisationa can tie local ivd
with sufficient accuracy lo be referrtKl to two points of the liody, and not eos-
foiuxhil tog«lher as one:
Tit) uf tongue I.I aiin.
Piliu of last phalanx of finger S.2 **
Palm nf iocond " " 4A "
Tip of Hon- fl.6 '•
White psrt of Hiw 8.S "*
Bsck or seeoiid pliHlniix uf finger 11.1 "
Skin over malar hone 16.4 "
Rw-fc of hand 29.8 "
l''orcnnu S9.8 "
^teniuin 44.0 "
lUck «6.0 -
And an analogous di«tribuiion has b«en observed in reference to the toosli-
lation of MDsatiutis of leniiieTatu rr. As a general nilr. it may bo said thai
the more mobile purts are ihiwe by which wccan ihu* discriuiinato semUioa
roost readily. Tbe lijghter the prcasure used to give rise to the eefuatiow
tlie niore easily are two sensations distinguished : thus two points whiob, wbn
touobing the »kin lightly, appear as two, may, wlwn tinnly proetod, giveriis
to one ^eniation only. Tho distinction between the soiiM^utis is obscured by
TACTI1.K PBBCKPTIOX8 AN'D JTTDOMCJfTS. 085
llii'riiii! Mnwttion; tiri«in^ Kt the i>anii> time. Ttiii.i. two |iiiiiitA iir(i(i);hl
tr wi(hiti a ring of hifAvj mnlitl prcMMnv on Utc »kiii, Hn: niailUy cm-
ftined into one. And il noc^l Iiimlly be wij tlint ihe»c litctile pcrocptinn*.
I like nil olhcr pcrcoplmiia. bto imoicniwlv incrcw«() bj- «svreMc.
I Our " lipid of loucb," if n-c may be nUowciI lh« nxprvnainn, i» coinpowd ftf
tactile KrcDis or iinilA. in the wlqic vmy thnl viir fio.M of vUion is )Xinipi.i«CK] of
viviinl arras or unil?. The tactile tcnxntinti ic, like the vtsiitil MnmltoR, •
^^mbol to m of eoiue esteninl «vcitl. nnd w<- r^'frr tin? •■.■nMtiun to i(e nppro*
priate place to tbe lield of touch. All thtii hw Ix^-n mid (n. tilT) conwrn-
L iDf; the subjective nature of tbe limits of visiinl Hreas, uiipliM cijtially n-oll,
I nuMu miil-iH'liti, to laclilo nreiu. When Iwo points ot Ih« cotnpnawA are
C firll as two di-lJoot iteni'nlioiis, it \s not noocveary tbnt rwi, and only two,
I Berre-iibreialiould beatiiuulHlod ; all thnl is necessary is I hit tliH two cerebral
Ltensnli oil-areas ftliould Dot be ton coniplot4.dy fii*«d ^l'.;l?llle^ The iniprorc-
r Bieni by exercise of the seote of touch umet he explniiied not by tin increaaeil
development of tbe t«riutiuil organs. Dot by a h'f>'Wth of iicw nerve-flbrea iD
the skiD, but hj a Diare exact limilalioD of the ««Dsatii>nal areas in the braiii,
by the develnpuient of a reustauce which limita thi: mdiation lakio;; place
from tbe wntreti •>( the several areas.
Uj a multitudt? of siniulliincuua uod cuDscciiIive tactile seoanUoiis tbua
coDverieil into perccptiiinn ne an-- able to make ouraelvee aoi|uaiDt«d with
the form of external objocu. We <mn I«l] by varialiun« of pretmire whether
I a lurfavc m rou^h or )>ni<iiith, plane >ir curved, what variations of surface a
i body preaeiits, ami how far it is heavy or li^lit; and fmrn tbe iDformation
thus gained we build up judt^menls a» to the form and nature of obiccta,
judgments, however, which arc idohC iiitimatt-Iy bound up with viaual judg-
ments, the knowledge deriveit by one aensc oonecling nnd completing tbil
obtuin<;>l by tlie other. As to other seDB«», so in this, oiirnnntions may
mi!*l«ul us and cause us to form erroneous jiidf^i-nt»~ This is well illns-
tnitc<l br the so-called experiment of Aristotle. It !s impciwible in an ordi-
Dary poMtion of the Kn^rs to bring the radial side of the middle tin^-er and
the alnar side of the ring finger to bear at the same time on ■ small object,
eucli oa a marble. Heooe, when with the eyes shut wo cross one fiager over
the otiier, and place a marble between tliem so thai it touehea the radial side
of the ooe and the uloar side of the other, we recogniie that the object is
euch as could not under ordinary conditions be louohe<l at the Nime lime by
ihete two portions of our skin, and therefore jud^e thai we are Uiuebing not
one but two marblesi T.'|ioii repetitiou, however, we are able to ourreoi <Hir
judgment, ainl the illusion dlaap|>eaia.
Distinct tactile sensations are, as we have seen, prodnocd only when a
stiiuuliis is applied to a terminal orj^n. When seusatioiu or affections of
general seitsibility other tliau tbe dntinct tactile sensntions are developed in
the terminatHHi of a nerve, we are sdll ablei, though with leu exairtiiu>b<, to
refer tbe sensation to a |>articular |)«n of the Ixxly. Tlius, when we are
jirieket] or burneil, we can feel whrre tbi; prtek or burn i*. When a sensory
iicrve-triink 'a alimulateil, tb<- weDKiilion L> alwuyx rvfi-rreil tit iIk' [Kriphenil
teriuiuati'in-A of ihii nerve. Thii.-i a blon- on tin- ulnar nerve ill thi' cltiuw is
felt ns a tingling in thv little and riit^ finjfun eorrc^jrinding l<> ihr •liatrilHi-
tion of thci ncrvi-, nnil lu'ii^ntioni' *Uirliil in Ihe slump nf an ■T»piiliiti.--1 limb
aro rofbrmi to Ibv alx<«Dt ntombcr. When o<ild i* apprivol t» toe ulliovr it m
felt as cold in the skin of the elbow; but a cmling of ihv ulnar nerve at this
spot, •incc stimulation of a ucrve-tnink girca rise to general Moaations only,
(imply gives rise to juiin which is referred to the ulnar tide of tlw liaod
ana arm.
■
see
PKKLIXG JiS'U TOCCII.
Wln-ii we ooine into citiitaci with eslernal bodiee we are eoDsiioui M
only 111* il>e ]>reMui« exerteii bv tlie objett oa our skin, l>iil alsr. of tbf iii»-
sure vh'ich we exert on ilie object. 11 we \t\nce ihc liiiiul ami iirui flal <« %
table, we eau itiliniiile the prtwure cxerteil by liudii? n^tiii): mi llie |<almW
die ban^, Btid i>u vamt Uia evnoliuiuD a* tu their neiKblJ*; in ibia men
are cmiiciouH only of tho pn>Mtm! «xerte«l by the body on our ikin- I(
however, wo hold itie bmly iti llie hiiail. wo not only ^0 the pnwaiire «f th
bu()y, bill we iire alw) aware of the mii«mlar (.•simiun n^juiml to nfpm
and lift it. We jHuaew a miiRciilar m-dm-; mkI wu linil by cxpmeoc* tbM
when wc tni«t to ihlt inuK'iilur tiaifV- lu w«ll a* to ttnwiiiinn of [inwu)^
ire CUD furni much niori' tuxiunite juilgniciila conocrainv the wviglit ofbtxlis
limn when wc r«ily on iu.'iiMilioni> of prmtiirc ulone. \Vben we want to idl
how hcKvy II biiily >*, wc nrr not in the hnbit «■' allowing it nimi'lv to gim
on the hunil ljii<l tliil nn n Inlilc : wc holt] it in our Itanil mn<l lilt tt i][i ami
down. Wc Bjiiiciil to our nuiM^iilnr m-ii#c to inform tn of tin- nmoiini ot
exertion ii(-i->-t«iiry lo move it, and )>y help of that, jutigs of ita wvi^
Aod in all the inovcincntti of our body we are enidcti, even lo an ulonatt-
ing dtffW of nci-iiriicy. mn i« well fecn in iIk- diectiKiioni! conirrniing visitjo.
by an spprcciiitiiin, more or len dietinclly <ron»ejoiu, of the amount ol' ilit
contmclion tu which wc arc pntliiig our Hiuscles. In autnc wny or other «t
are made nwiirc of nhnt [larliciilar muscle* or groups of miifclcs are hein;
thrown into nclioo, and to what cxIcDt that action i» being curried. We air
aba conseioup of the rurying cunditinn of our inuscln, even when ihef an
al reel; the tired and especially the paralyzed limb is said to " feel " heavy,
lu iliia way the stale of our uiuHcle.i largely determines our gcDeml fMliu|
of heultli and vi^r, nf wcariniM*, ill healtlt atnl feeblcoeaa.
It ha« beiii tugKio^d that «ince iniuele iiuMeMU little or no genrrvl m-
tihility, compamtrvely little [lain ttuing fell fur imilai>ce when inunoloait
col, oiir miiiK-ular >eU)io i» eJiietly denvtsl fWiui the traetioQ of the e<U>traei-
ing mii»('le "U it» attachmeDln; und nDiliiiihlinlly ui many iiMianc^ of rrwi^
the ]wiin ii> ehii-flv fell at thu joiiik*; and, an we know. Pacinian bodita in
aIniiidiiiiC aroiiiid the. joiiilx. Atrerrnt nerve«, honcver, having a diderent i!i>-
pontion fri'iii llie cmlinnry motor ncrvei' which li-nniimle in eiid-plalea, htn
been deocribiil n» jircneiiT iu nitiwli-; and annlogy wixild lend un lo euppMf
that tliew nifcrctil (il>rt». though [HJMCMing a low niDOral sODMbility. bi^
l>e eiiPilr exeit<rd in a vpceilic miiniier by a inuaciiTar contracliMi ; bat ^ll^
Uier invtatigmtionH are nec^wary Itoforu tJinc can (m Hccofttod as tbe tnt
IKrvea of tbo mu«cular >ensc.
In faror of the view thai the niii>ciilar khmc it {icriphcral and &■>( ivoinl
ia origio, may be urged the fact that the m-um is felt when the muscle* an
tbrown into contraction by direct galvanic stimulalMHi instead uf by tbi
air«noy of the will. Slany autliors, even vhile admittioc tite exiateDoe uf •
muiicular wD^e of iicripbcral origin, contend that we also noaseas uid an
very lar|;ely guided in our luorenieDta by what aiif^bt be culled a " neunt'
aenite of <7eutnd origin. Thai U to say, the ohan|[e0 in the central nervMI
■yMeiu involved in iuiiiaiiiiK and carrying out a movement of tW lajdy, «
aflect our ei>iiKriousriewi, thai we have a scnite of the elliirl ilcelf.
Il ha« been olieervetl that when the pueierior mote are divideil. inoveowMI
lieeonie lees orderly, nb if Ihey lacked the guidajioe of a muscular wnm;
aud all)ic>iit;h tlie imjutirnii-nl uf tbe muveiitenls may he due in [utrt tu ibn
coiuciilcnl luni of [aclile ieiiMitionn, it U [ir<ibahlc that it ts incrvitMi) by lli*
Ion of tbo muscular seme. There it a malady or rntlier a condition atieii>r
THE Mf3CUl.AR SENSE. 907
JDg vsrious dUeased states of the central nervous system called locomotor
fttaxia, the chaiacterletic feature of nhith is Ibat, though there is no loss of
direct poorer over the muscles, the various bodily movemeDts are effected
imperfectly and with difficulty, from waat of proper coordination. In euch
diaeasee the pathological mischief is frequently found in the posterior col-
umns of the spinal cord and the posterior roots of the spinal nerves, that is
in distinctly afferent structures ; and the phenomena seem in certain cases at
least to be due to inefficient coordination caused by the loss both of the
muscular sense and of ordinary tactile sensations. The patients walk with
difficulty, because thcv have imperfect sensations both of the condition of
their muscles and of the contact of their feet with the ground. In many
of their movements they have to depend largely on visual sensations ; hence
vfaen their eyes are shut, they become singularly helpless. In other cases
again ataxia may be present without any impairment of touch; but a dis-
cussion of the varied phenomena of this class of maladies cannot be entered
into here.
CUjVPTEK VI.
SPECIAL MUHCULAR MECHANISMS.
TiiK Voice.
^Thr Phi/fiotogieal Aii'itoviy of the Laryiu.
TnR larynx ig a iiiumbniuo-cartilaKtnouB chamber, broader kbov« tku
below, mid nluaiiil iti tlio ■iiti.'nor ineriiau porlioit nf the nevk. It coMiai
of ft number of carciUgM. irhich are articufaied with each other, ooniMMed
bf ligameiita, morcd by k nutabcr Df i»iuclc«, aikI lined by a mueoui mtm
braoe.
The principal cartilagee arc the thyroid, cricoid, the twn irjrteoojd. u»i
tli« epiglottis.
- ■>
^.>a*.««
V=="
UlDUs ADTnoK i» Muimi. NwK. I'liinrHx. ami Immytx.
«, Mpmn at nait. titlCHt It. trcLlou nf hu4 |i«]«le : b. loDfue: t. ■gMlos ot v«(iin inOHM
t«l>U: 4.<Upi; H. utuU: r. ■lurrloi irrh »r )4Um ef fanvs . i. loocrlor anb, t btfill.r.
phatyax: A.b)raUbaiu; t, Ih^miil ivitillkt* : n. artoold mf1lln«i>i <.f|Mii1oiiki . r, clutUt; Kr*^
rior oianlni or Ihc nan*^ I, Mliiuru fnuuXiB l {. mfcttia nfeniBf 04 Ufyni ; V fMMff Ml
<Moptal(tIII (iMMlbofriKhl Eiutantiiaa tobn.
The thyroid cartilaij^ U the larf{«tt niid coosIMs of lirn ((UMlrilaUril
nlnlee of nl», which are <M>ntiiinoui with '-iich other in front, where thtf
jonn ilie i>rv<iniiiencc calk-il thi- iK>iiiiini Adaiiii. Tbo poeteriur hordenaT
tbe tliyruid cartilaKe aerve n.-> n pmiii of nl.tndiiaeat of tlia stylo ptuirynffMU
aod pclato-pliaryni^ui) niuKcIn. Tlic uppnr purl of nich of tbeM ^raM
leriniiiala m n superior oonii), which articulat«s ititb the Uymd haite ; U*
»
sr portion MrrDioAt** in thi- iiifeHor cornu, which artioutaua trilh the
criooJd cartil*g«. The uii]H'r Imnlcr bcLwecn tlio cornuK is oonnected with
tlM hvoMt buu« ]>y thu idyro-lijoi*! membninu. The Inwer border ia coD-
DMted with the L-ricuid cartilugv by the ibyro-CTionid inoinbrmne at the
nediui line, aixl at the hiIcm hv thn orio-thyniid niutoltfx.
The cricoid cartilagi- is Mtiint«d bcbit the thyroid i-nrtilug« with it« brood
jKiriiou iKMeriurly. Al the ii[i|)cr pnrt of its broud portion atv two Moooth
nirfaves on which the or/tciiDid cartilngw lutiouluic.
na-W.
riij. a«-
FUI. SO.
Pml aw.— Vint or nil Liiivsi jum rikt <» tii> T«*ciiki nmn BniiM' hitii «« Miua,n
ft, Ik* hatf of (he lijxilit b«m : r, •f(tk*tt> ; i, ihc laUttur bofden of tte ihn"ld a^i-
»: «, tlie BtdMB hdt« of Uw orKoM : a,app«r |aut nr ifa* arrtviKdilr ■. iilitml ui oiMorUn
•UlqM hvieUIl uT lbs vjtUuM ouuela; b, Utl poMac oitetMiyUDold idumIc: t<id> of IbO tn-
«g|u(M( canlUglBiAit rln^ of Ui« Inobm i I. Hbroiu mcntnitc craBlnc tlie lack oT Ibe ttuchta ;
a, nwoulu nn* cxpool ■■ ■ |iut. (FMu Quoi^ .Iwiuiiir.)
riD. JH^ViBw OT TiiK LiatT'i nooi uen. l.apnfura at rtrtUt: l afrtmotil nraingM: t,
VMkI oonli ; I. pnatHlnr rTtA>«r7<0nnH iDiBpln : S. laHnl fdeOMHixM moiclQ <f IlCtil iMc. thM
«( Islt aMe rniunicU . 4. •lyli-notd niucla : t, thjTixryMttoJil mliaala of lolt dila. Ui*l of Hehl (Ido
nnwTfdi *. Uiyn*dr*»llJtKD: *. vntuiiltanibitrviU. inMcnarcrlaHUTUiwilil llgMnML WllhUw
cin>|ium of ilM arxUcnb] dukI*, Uili dtiBnm U ■ (»|7 fRiin Mt. WlUU'* Ocni*-
Pw. nft,— View ur ttu mm* lunr or tbi I^uitsx a toM by hum nr rai LkBTKoaKora
mwiKi' TUK t'TnB4H(««r i Orati Xotk <.PidsiauU: <. Uio carlllicTC of 8Hib>rltil| a,ftfj1*-
The mrYlLiiiHcl nrlilngn xre pvnmiidnl in form and orticulnta on the
upper siiriiicp of the criwid. Eac^ cartilage ha« ao external, posterior, aad
intenml (nmliai)) tturliioe, an apox and a biMo. The apex Js pointe<l baclt-
WBr<l and inwnrd, and it siirmudDtod hy a fniall cartilaeiaoua tutiercle,
vailed the carlilnRe uf SanU>rini ( Fi)^. 'J59). The b«se, which arti<Tulatee
with the cricoid cartJIaKe, presents at iu external iiiternul aD^le a projectioD
calk-d the firoe*u*u iwu/u. Ai the posterior internal uiiu'le ia a aeoaod pro-
jection, lailleil tlie froetatua nntteitlarU,
The «U|N-rior o)H.-iiiuic of the larynx is formed anteriorly by ilie epijthMtii,
poNteriorly hr iho uptvea of Lite arytenuid i-artilavu*. and laterally br the
■rjtooo^^wltidwu) fohU atretcliing Wiween thtsM: puinia. Tbv inferior
J
970
SFECTAT. MCSCrLAH VKCHAlCtSUb.
ntmiing i-orn!9i[i>inilii tn ihe iiifi-rior border of thp crimid ctirtilnR^. Bt1««rg
lli(*e jMiinU )i> tbf ci'.'ilii i)t' Ihe UryDX. wliicli has Htr«lehiiii^ ornw ilt «i(In
tho Titcnl c<MiU. Thu vm-al cofdn ooiuut of two |Mint : the eii|>en»r or faW
t-iK-nl ciirilit Km Tni:iiitiriLiii)-ligameiih»i^ biiaiU, nhicU exKiiil from the nttA
iii^ iingk of iho thjrnjid lu the. extiTiial aurihoea of lb« iin-t«n"i<l <nni
Ugn; ihr infiTiiir ur lrii« vo<:nl oonJa (cAffrdi iv>(vi/*j> iiro membratwi-hn-
lueiKotiH biiDdx which Mrddi soron the cuvitv of tho Inryns fnm tht
rnvdin^ tinglr i<f ili<t thyroii] to the provwu* tucaI«* of l\» tirytttwiA cu-
lilagt*. Rrtwccn ihr bitnlrni nf th<! true aiid talte vomlI mrd* Ui no eMiptinl
oMntDjf, th<^ rinUiflr, whirb Imils to ■ Kiianp ruiiiiing upK'iiril nnil hiiiioi
litv fiilwj voail curclo, fidluil lliu *hiciJi« tnrvti'jif. Th« mumti* iDi-aihniE
lining IhiM sic coniiiiiis t> ^'n^il niimtH?r of folliciilnr ^ImiiU wbiob ^\\^f\^ucr
n iDueuits wcraiioii for the inir|Hwc of luliricjitiii)' thr trii« vwnl i-nnlv
Between tht- trui.- vocnl rnH« is nil optming nlircU is callnl the riniit yl«i-
(trfiA Th« f(inD of tht L'lotti)! Vftrics verr much tiolh in th« tii.'>|iiniloi7 «»J
exwnton' nvU, mid in tbc act of nhouuIioD.
Th« tnusde«f of the larynx are iliviilctl nnatnmimllf Jntn the iiitrioiic wA
i-xtrineic. The former are niiie in oumber. fixir of tWin Itcin^ in fain
They are ihe eKential luiiscica cooceriieil in the luovi'metitfl of tli« arytraniil
carii)a)!efi an') chorda vnciilca. Tlie exlrioBic nitiBcles counccl the lanat
with adjacent [wrtM, and ai-e for the moat |>arC couoerDcd in tho elevatimi
&n<l dei>ri?tiaiiiii iif the orpiti.
The larynx ih lined with » inueoiu meiubrant- whidi is Cdnliniioua Aam
wiUi (bat lining ibe jiharynx aiid mouth, and l>eh)w with that Winat Iht
trachea. Alxivo Uie chonia voailtiH it u lined with pavement epithelim,
excepting at the lower anterior portion, where it U nliated; iKiaw tk
chorda voealcei ihe epitbebuiii U of a nliatml (idiininnr varieitT. Tke
nitUMiis nifmbrani^ enntninti niiniy niticoiis glandf. which i\tv pn:Uy ani-
furitdy di»tribut«i): thvy ua-, hitw^rer, very ubuDdaul in the |Mrt of tk«
nM-inbmiic lining tho >uti-cu1ii« Inryngis.]
A blast of air. driven by ii mure ur Ims prolongetl expiratory movenata^
ihmwa into vibmiions two elaalic niembraues— the fAonkt- iwxUr*. TbeM
imparl their vihnilion§ to the column of air above tbem, and m give Hm la
the ioimd which we call the voice. Since tfae sound h generated iD tbf
vocal cords, we may speak of them and of those parts of tbe larynx wharh
decidnlly airect their condition as constituting tbettoential vocal apparalu;
while tlie chamber above the vocal cords. comprtBinn; the vcnlricles of tb*
larynx with the false vocal cords, the pharynx and the cavity of the muutk.
the latter var^-iiig much in fnrm,couHtituteu!iiiMdiaryap^iaratusaftheiMliirt
of a resonance liibt!. modifying itie uound originating in the vocal corii.
In tJie voice, us itt otlier sounds, wu di»tingiiith: 1, I^udiMOi. "nil
dcpiuds on Ihe Hlrength of the expiratory blaiiu 2. Pilch. Thi* depnlt
o« the kni-lh iind IwiMiou of the vocal oirdn. Their teni/lfi nmy Ih.' regatdd
as conslani, or varying only with age. It conscipirtilly itt-trnniuca tlw
range only of the voice, and not th<^ [lartioular nolv giv<-n out at any um
time. The thrill voire of iho child m ■Iflfmiincd by the nhorlniwi of ihr
cords in infancv, iind lliti vute^ of a wipriinri, Icnor. and iMritoiM' an all
dependent i>n the r<«pcrtiv« k'ligtlis of ihi.'ir voeal conU. Thfir ViuMail
Ml the conlrnri' variable : aikd lh« ebiof uroblein* conrnTlol with llu> n<i9
reler lo variiitioiia in the Icmion of the v«cal conU. 'i. Qiinlily. Tib
de(iend.-> on tin-- numWr and character of the uvertoiies accoiii)tanyiiis "»<
Aindamcntal notr aiundcd. and is detenuineil by a variety of circunislj
ch'tef among which it the physical c|uiility of lie cord*.
Tfae vocal curdjf, atUched in front to the thyruld cartilage, end bduiiill
971
the i-roTrtwi* vocalts <i( thi? arvl«n>iiil (yipliliiges. Hence n disttiirltoii \m*
btva iJruwii lietui-ni ibi- rium vucsIU, i.r., llie ojwoitiK boumkil liilcrally by
tbe viral (.■•inU, Mill iW Hniii reapiniEnria, or Kfoice oHvven the nnrCMi'iiil
nrtilnm lit-binil thu priK-aMi) v«<-jil«: thi!t«e uaoxs, bowevor, are not I'rvo
fro«» objootio». Ill qukt hmthiiig (Fig. 260, B) the two ronn togdher %
nu.sn.
u
n
<^-
'^
TUB LAan>x a sax kt UttAH* ot-im Lilimiiumra iji innnKwi Cuhmthim o* rnt (iijoma.
I Pnon Qt'ilVi .1 nlwy. ■<>« C'uciihai.i
J, wblla tfnflDV a Mib aatr: S. la quM toi«lUii«. Cilurla«a dMfi tniHntkn, n* (Ofn-
q«adiii( dlifWi— tie Ikun* .«'. V. c*. UluAmla llio rtauna ia fodtlon of llie utwmU (•rUlffO
■aillli«fcfBorilicnini>«<>n>Uii>ii<lrtinHn»|<lnlnrt*lnlk(abi)T«Uin<«(aMlkloDt- L lh( Intaof Uw
■Docac. c.ilieiij>i*r ft«[«rloI ILe «[•1|l^>(ll■; t', Uic iDteida OT raAlan «f tbe Cpltldtl'* : P^l*n
a(UH aorlortM skU oSthe (ilitrinx IhIlIiiiI Ihu kryux ^ u. imtllBKln Ifta ar]naii>«T4(MililiaB Rilil
tanwdbylkcainllKgeof WhttKtc: «. •*«lllniln>'v*<l br lb« oaitUica t' 8a<>I«**nl ; a. (hn niuiMU
•dbuanvaold aitltaKe: or. iha iraovnal oonli: tii. iW libi toMl o«f<l>: ir. lb* UMbBft wRk
IB rtot* I ti Ik* t«o tiraiiobi at Uitdr ooniionicwmviiL
V-abii|K4l »iMioi', wlik'b. rm m« havi* tt-cii (p. 449). in <lwj> hif|iinilHO is
vri(li>ni.-(l inlu n tliumlwiilal (ifK'iiing hy tliv divor^noG of tho pmceant*
vornliv < Pig. 360, C). Wlirn a nol« » uImmii to lio ultcrrd. the rocnl cKinls
•re l>T ihtf Kpnroximntinn of th» pfoivMiis vocal** broii);ht into n puniion
pamllH \a mud other, and the whol« Htnn b narrowed (Kig. 'ifVX A\ lif
their parallelism am) hy ih« nirruwDon of tlw intrrrnl beln-ren tli«m tlie
cords are renilpn-d moro Miwvritihle of be)D|; thrown into vihnilion by a
moderate blast of air. The prolilcinf we have to oomider are. tiret. liy nlial
neana are the conk liriioght near i<i each otherordnwu asunder a«ocaiHiim
demaixtf ; nod. wcoudly, by what lueaiis b tlie tension of ihe cords made to
vary. We may apeak of the«e two ncliong aa narrowing or widening of th«
gkuia. and tijfhleniiii* or relaxation of the vocal conl*.
JVorTOtnay oj Iht ^lolli*. The change of fvru of the glotti» b beat itutlof
072
SPECIAL Ml-SCrtAB ItECHAKISMS.
Alood ichcn it Is bornp in mind tlint eneli nn-teuuid curtila^ is, wlien
boriziontal eecliun <Fig. 2'iU), someuliftt of the form uf s triMn)tle>i
internnl or tudllan, an esl«riiat, and h ]>oeteri(ir side, tbe prooeeEus ratalk
beioj; piaoed in the anterior Annie al tlie junctioo of th« meoion wad ezteral
sidM. When the cartilages are ho placeil tlist the processus Tuoilca »•»
proximated to each other, and The internal diirfacts of the carttlsM MU^
parallel, the ;:-lottis in narrowed. Wlieu on the eontrarr the oartilagt* U
wheeled round «!i the pivDtd iil' their ariieulntiotts. so ihnt the pracnwu tuoIm
diverjiL-, niid the iniemal snrl'uccB of the oartiln^ fonn un angle (ritli nA
iHlier. the j-iuttJH ia iridened.
There are Heveral muaulea forming tuKethcra group, which li» bven cmlled
byHeule the itphincier uf the liirynx. Tliew are: 1, the(Ayrri-<trv-r/ii>7/o(finii.
pnxweding from the inner siirfuce of the llijroid cartilagi; iinil Iruni th« arr-
tenoid <-pi>(hiUideiLn liniuciit, tiud nwerning rotind thf outvr ridgt of Uw
nrytenoid (-jirula)j;e of fta own riide 111 Itft iDHcrie'l into the procCMUB inasai-
laris of the aryi<muid curtilage of the other »ide ; '2, the lA^ro-nrt/lmoi'li^
ftcfeniM*, piumng from the rci-utnmt angle nf ihx tli^roiil eartilni^ \o W
inserted into Un'. outer edge of the arrteiiifid enrlilikgc of the rame Hile ; X
ibe AgronTiflrnolilriu tntcrnuK, puming from the Miglc of tin- thyroiil at-
tjlsgo to the proi^wnus vorali* and niiler vide of the arytenoid eHTlilage ; 4, tht
arj|i«H0id«tM (po'tien*), pa««ing t.rnnarenH-ly from one iirf tenoid enrtilan tu
ODOthor. All thew mujscle*, when they Mci logetlicr, graep round tlio glotdi
and tend to cIimc it up ; ami eacJi of them, acting alone, has. wirU the ex<
caption of the last-nnniiM) (iirylcnoideiis\ the MJue efleet. In additioa la
Ibese. the cfifo-iinil^nouifiij' tnttrijlit, whidi nasses from the lateral iMirder
of the cricoid Mirlds^^e upward and haclcwanl to the outer aii^fle of Uie tn-
tenoid. by pulling this outer angle forward throws the pmoewus vecsb
inwanl, and ao aUo narrows the glottis.
l)V'jfcii(«y of tlu 'jloltis. The rrieo-ary>enoidfH* ptwlifu, patting ftooi tbi
posterior surface ol' the cricoid cartilage to tbc Otller angle of the artrteaml
cartilage behind the nltachiuent of tlie lat«m) crico-arvleiioideuii, pufli bad
thU outer angle, aiwl so cauBiDg the proomus vocalii to move oatvanl
wiik^ns the giottift. The arytmotdeu* pottinHs, acting atooo, ba* a tiaiUir
vitect.
Tightrning oflJie mm! eorttt. The crMvjAyroi(f«>M pulls ihv lliyruid do«fr
wanl and forward, and *<i iiicreasai the diHtnnee between that cartilage and
the aryti'iKiidH when the liittcr are fixed. Suppucing, then, the aryt<'D'>*Ii'w
and eri«)-arytcnr)idi;ii» jioilicus to lis the arvlt-noid-i, tlw effijct uf lh« cm-
lracti<in of the erini-thyroideo* wnuld 1m: to tighten tbc vocal cords.
Sinckraitiy of ihr xtmil mrd*. T\\\» i» cflbcted by the nholu sphincter groua
just niciilioiicu, but more i?<|M;i.-ially by the thyro^nlrnoiitfi r^lrmur SH
inferNiw; throe, acting nlonc. viippieong the arvtcnoid i.-artilagus to be fixel
would pull the thrr'tid ciirtilngc iipwan! and iiacknurd. and so sfaortssi lbs
dtKtaiicv between tlic procOMUs vocales and tlini body.
Thui^ almost every movement uf the larynx i" alTiftcd not by one musck
only, but by several, or ni least by more than one, acting in concert. TW
movcn>oiit« which ^vo rife to the voice are iirccmincntly combined aoilei^
ordiaate moveinenta. ^Vhen we reineml>er now a very slight variatitm h
the teoaon of the vocal cords niu»l give rise to a marked dillbrence in thi
pilcli of the note uttered, and yet wliat a mullilude of fine dtHereoces ef
pilch are at the command of a singer of eren motlcTati' ability, it »i-)nn 1
exceedingly probable that the various muscular combiaatioiis reiji.
jiroduce tlie posuible variations in pitch are of such u kind thai fn-'
part only, possibly a few libres only, of a particular muscle, may Ix
Contraction, while all tlie rest of the muscle renmiiu <|Utei. iiiKiug m.v
THE VOICE.
973
I
ir t!ie great rnnRe iif jiitch poesMBed by even oommoii voiceit. iia
' ihi- ]H«iail)lt! vumliDtis III' lenuoD nf u'lik'h lh« vocal fDriU iti
tbdr nnluml li-nglh uri' (-ii)iiitiU-. it Iibm l>e«u BU^igesled that Mine uf tbe librt*
of thi^ thyni nrvhriioiilrun inliTtiu>, wliieli, puung eilber froin tlie tbyn>i>l or
from tb>- antctimi), Hp|nutr U> end to lh« vocal coixU iheioselvea, wiij, by
fixing |Mrti<'uUr (initilM nf tbe cords, so to H|>eak, "Mop" tbem : nii'd by
Uiu* nrtilicially ohorleiiiD^' tin- length actually lUrown into vibrntinti, pro-
dtico higher inAct than the corda iu their natiiml lengtli itrc tiiiiablo of
prmlucing. It hiu hwti kIm auggnl^Hl that tlie prooMtUK vocalm way
overlnp ciwh otiior, and thereby shorten llie length of conl urailable for
vibration.
Thcw varioiu muRcIts are supplied by therneiiii ner^'e, or rnlber by spinal
■ooMSory tibr« running in the vagiM trunk. The «(i[>erior laryngeal is tlie
afl^rent nerve supplying the miicoiis meiubnine, but it aim containf ibe
motor fibres diBlribiited to the crico-thyroid miisele; heniv. wfavn ibis nerve
ia divided on ooe aide tbe oonwpondingvoad cord i» mluxed and high notca
become iiDpoaaJblek It n wortJiy uf nutice thai thi«, the ebief leoaor, and
therefiire the tnoM important, inuacle of the luryns, hiu ii Mpnrnlc and dia-
tinct twrvoUB ailpply. Acoordinj; [<i some uuthors the nrytenoiduus posltcua
ftlao receivca its nervous tupply from this nerve; but this b denied by
others.
The inferior Inryngeal or recurrent brandl supplies nil the other tnutcles.
Wtien ibiH nerve ia divided tJie voioe ia lost, since the approximation and
paralleliam of the vocal oorda can no longer be efleeted. When in ii living
BDimal both recurrent nerves are divided, the glutlia ia seen to becimie im-
mobile and iMtrtiuUy dilated, the vocal curds assnmin]; tbe ponitiDn in wfiich
tbey are fuuod in tbe body after death, and which may be coa«i<Ien;iI as the
ooDdilton of iHiuilibriuni betireen the dilating and coaMricting niusckv.
During forcible inapinUion ibo gloltia puafcsi from thin condition in the
difvcliun of more comnlete dilation; dnnng forcible cxpirution, tbe change
is one of constriction. When the peripheral portion of one re>.-nrr«nt nerve is
■timulotod, tho voeal oord of the tatma sidv !■> approximatrd to the middle
litio: when both nerves arc stimulated, tho vocal cords atv brought loj;etlier
and the glottis is uanowci]. Though the nerve is distributed tn bolhdilaiiug
and euoslHcling muscles, the latter overcome the former when the nerve u
artificially elimolated. In the complete closure of the glolli», iihicb ia so
important a piiri of the act of coughing (p. 'tiyO). tlie group of mtiaclca
which we have spoken of as constituting a sphincter is thrown into tonTible
GODirsctioDS by tne recurrent Isryngenl nerve.
Though fundamentally a voluntary act, the utterance of a given note is
not affected by tbe direct psasagA of aimple volitional impulses down to tbe
latynfces) muscles. Bo eomplex and co5rdiuat« a movement as that of sound-
ing even a simple and natural note re^nirw a ooordioatiug nervous raeolian-
isiu in whidi, oa in other complex muacuUr actiooa. allerent impulses play
an im{Kirtant part. Auditory sensationa. if nut m important for uu oocuralo
manattenieDt of the voioe as are ritual aettaationa fur the tiKiveinenia of tho
eye, are yet of prime importonee. Thin is recognized wlten we auy that audi
and auch a one wboM power over bis Isryugeul iDUSctea is im|>orfcct " has
no vuT."
A penon may apeak or ains in two kinda uf voioe. In the one the sounds
are full and strong, and tSe nwmanco chamber which is aupplifd by
the trachea. bn>uclii, nod indeed by the wbol« chMt, is throws (nio
powerful and piil|>»ble vibriilioni; banco this vdvo b spoken of as tbe ctiMt-
voi<v. The i>lhvr kind of voice, called tho blsvtto, in (bin and ptwir, dssls
cbiedy with high Dol4S,and is act aecompanied by tlui muuc cxHi«|iicuiKis
flT4
SPECIAI. Vt^SCCLAR M'SCHAXISUS.
viliraiioiia of llie chwi. Much coiilrover^y has Uikm place m to ttwana
miuiner Id which th««e tno voiL-es aro r«|)«Gt)vely iMudaced. Tba nnmflinf
opinion lencheH that in the chesl-voive ibe Tix>al mnls are MinM'n'linl iIiwIl
iMir HubaUuice beinu ihritel inwanl toward llw inealinn lino liy ibi' motnr-
tiqo of the lhvrii-aryleii<>ii]«i i-xierui luuaclw, ami Ihe oiwning brtnoeo tli«n,
Bnincliniea ftu Darrow im ii> he aliiiit^I liiiemr. exictxtii iil»ng ihvir nbiilc Irtutb.
Ill the fiibettn voice, on tbi^ i>th«r ham), tin* vocal iMnli) are Mtd hi br tliit
and nif.mhraiKiut, and the note to be given (arth by a vibnitian. tml <•( tb
whoK* Kidth up tli« eunh, a.-< in tbe obait-v<^c«, but of (titt exlmtii' nb«
only, till' lateral |>arts, tUoujih not HbHuIiiUdv at r<i>t, vibmling with a iHif
«iil rhythm. Tliuiigb the vi\wh kryrix in tlii- lalNi-tlM vuicu i> stratcbad In
Ihe nnlcro-iHMieriur direcliun, ami thr vncn] curds eumMpupdinjily otoocMtl,
the ritna ro(»ili» diMs uul cxtt'nd nlon^' tbi-ir wholi- leugtn ; nl their mMC
jtarl Ihi' i'iir<l* an ill oODtnt-t, uml indcxid, uc-cnrding in ttjtne naltion,)
iiijjh fal»olti> noltw are pntdiK-id by a i«irt of "' it»iif>iiii; " of the curdo.
acnse of eflort which ai-i'onip:init« ihr fal>«'ltavoi<-o iiiilicalee that the d
id the larynx which bring it about arc c-Aected by eonio opc^-iftl mu
mnDanTrea. m U n\m titggnUxl by the fact that iW Mae with whicb (alMla
aulefl can be utlorcd is readily inrrpasod by pmclice. Tlie cbau;» ff^im iW
I'ht^t to the &lsetlo voir? is an abrupt one, nm) the cuiubiued ranee mar W
very fSlenwve. a« in the i-ase of pcrMns who can oirry on ii duel. «iigtDf
ultt'rnatfly, for InelaRce. in a tenor (chest) and a aoprano (falM-llo) voift.
Thn reulriolcs of Mor^f;ni are apjuircDtly of use iu K'^'ins ^ ^'"^
cnrdM iiutBcifint room for Iheir vibraiioDs, and |terhai« supply a MtcrMiM by
which the vocal cords are kept adequately iiioiai. The purpose of ih< &W
vocal cnrd» u not exactly known. Sume authors think thM In tbc fabMIs
voice ibcy ai« brought down into conUuit with, and tfau* wrvc to stop, lia
I rue vocal cords.
At the ago of ginbcrly a rapid development of the larynx l«kea pUcft
leading to a chnngt- in the range of the voicv. Thi.' peculiar handuewW
the voice when il i» thii» " brcnkinir " wwinn 1o br due to a tcm|w>mry («•■
grated and swollen coaditinu nf the mucouR tnembranc of tbe vocal curdi
aoconipanyinR the activo growth of the whole larynx. Tbe cbaitE« in ikv
niuvoiM niennbrano niiy conic on quite suddenly, the voice "breaUng" f«
lasunoe in the counv of a inghL
VowiM.
Rvcry aound. or even- doI« (for all vocal Muiid* irb«Q cnocUlMvd h
tbcmwdvea are niusical sounds), caused by the vlbriUluaa uf the Toonl ourih
beaidca iU loudness due to the force <if the expiratory blajK, uitd it* pitci
duB to tlie tensioD of tbe cords, has a ipinlitv •>f lU own, du« to the ouaihn
mm) reUtirc prominence of tlie overtontw winch aoironiimny thv ('iiitdamMit^
ton«. Some of thcee fealnrea irbiob make op the c(ualiiy aru inipueed oa lis
DDtv by ttx^ naiun? nf the vocal oords, but ntill mon- urino fritm variowi amb-
ficationH whirh (he relaiivc inleuiilia* of thn overtonrn undergo ihruiigh th*
reaonanci' of tbc cJivity of the tnouth and throat. Whenever ire hear m nnu
Munded by the larynx we ore able to n'cnj'niu< in it featHRo which etinU*
us Lo Etntv (but oiic or other of tbc " vowxl* ' is iH'ing uttcrvd. Voirel Hiun4>
arc in Aict onlv cxtrrmc aun* uf <|uality, •>xln-tnc prominence of orttaia
overtonn brou^lil about by thvi*liapciu*umcd by ihc buc<(nl and pharynpaJ
jtaaM{«8 bihI onlicM, a>< Hit- vibrations pan through thcin. Kach vowVlliw
SPEEfir.
970
I
I
I
lU ii|>|troprIal« tnd oniiulirc diaposition o4' llieee parts. When i (ee in Toet )
■0 B<>u»iic<l, llie Bounding-lube of the upper air-paangn b made u sfi'irt n^
poft*ilile, ili« larvDX w raiued and the lioa are retracted, the wliole cavity t<f
the uioutli inkinx on t)i« funu uf it brood flask with a narrou neck. During
the k'^'i^K '"" "^^ ^ ^0 '" 'i*^) ^'''^ *hnve of ihe nioiitlt in aituilar, but «ume-
wliat l<>tit:«r. For Ifae priiduclion ol u ibm in fuibtir) Ibe inoiith ts widely
(>)MMi. H> that the buccal cavity ig uf ilie (ihai)e uf a fuiiii«I willi lite ujm-x at
lfa« phan-nx. With u, tlie buccal caviiy \» iiauiu tliink :>liu|N.-d. nith die
iiH>uth more rlimet) Uuid jii a. but llie liji». iuxtCiad «f K-iu^ retnu-ied aa in i
and e, are Mxiiewliat {irotrudett, *■> that the auuniliD^-tube tit i>niKiU)/ed. The
greaiviAt leii^rth of the tube i» rcacheil in u (i>»), in wbirli the Taryux U
oepreiM»l and the lija protruded ax much a* pixtnililo. Whlln the two latter
vowel* ara being utivlvd, the general liirm of th<! buccitl CHvlty Is that of a
flask with a ihgrt neck and a KiniiU u|)eiiing, tin- qrifiw.- bdug stnaller lor u
than liir o.
Kitcli of th«te variou* " vowel " furnid <»f the inoutli poaencs a note of ila
own, DIM) tnwitrd which it netH a* it rvoonance chnmlrar. Thil*, if MVeral
tuaiog-forks »f various nllch Ik- behl while iiiiunditis before a m»uth vthtch
fcaa MWiatd the iwrticulnr form iiec^vary for •ounmng U, it will be found
that tbc nmaanoo will Iw imKicnUrly grmt wilh the fork having the pitch
of tho bat* ^ 'flaL Similarfy, ••ilier nud higher nule« will \k iiilciinilird wh'.'ii
Uw nioiilb if iiinuldi'd to tutor the other romila. And it is the rspcrieiicc of
ringers that each vowel ia Kiiiig with pocnliar eau on a note having a pronii-
sent overtone correepondiiig to the tone proper to (ho mouth when moulded
to litter the vowel. The precise nature of the vowel sounds is, however, still
diflfxited.
Aa the ribiBtions are travelling thron^^h the phBrgngMl and buccal cari-
lieB. the posterior nar<« are closet! bv the soft palate ; and it may be shoKD,
by holding a Uaiiie livfrjiv the itoMril, that ui> current of air iMues ttom the
nuae wlwn n vowel in pn.i[)erly »aid ar aung. When the puHterior narus are
no* ollvctiialty cliwed the sound aotiutnat a nii^al (ihnructer. The i«ine hap*
|ienH wlivn lh« anterior nam are closnl, lu nhiai tb<^ niup i> held bttlweun
tbe lingcre, the nnwtl chnmlK-r tli4-u forming a cavity of resonance.
OuuonmU,
Vowels are, as their name imiiliM. the only real vncnl ^nuiids ; it is only
on a vowel that a note can he mkI or sung. Our speech, however, is nuuH
up not only of vowels hiil also of oonaonaDls, i. e,, of noiiiide which are |»rv
duce<l not by the vibralion» of the vocal cords but hy the expiratory blast
being in varioiiH ways interrupted orotharwisatDodilicd in il# courac through
the ureal and mouth.
The distinction between the two b, faovever, not an abeoluM one, iinc«, as
we have seen, the characters of the sereral vowels depend on the tbrni of the
itMHilb, and in the prolucttou of •one consonaDts ' B, U, M, X, etc.) vibra-
tioDB of tbe vocal cords form a uaoenary though adjuvant factor.
CooaoDauta have been dassiSed according to the place at whicli the obar-
aoteristic uiierru|>tioa or oKNliKcation takea place. Thus it ruay occur :
1. At the lipa, by tiie lunvement or pmitton of ilie li|» in reference to each
other or to the teeth, giving rite to lahial eonsouants.
2. At the te«th, hy the nuivenii-iilor poaitionof the fhint mrt of the tongue
in If fervuei: tn the leeih or the hiinl palate, giving rise to iUnlut ooiiaonanla.
3. In the throat, by the movement or i>aiilKMi of the root of Uw tnogue iji
reference to llw aoA [wJate or pharyux, giving rise to guUMrai cottSMiauU.
976
SPKCIAL UUSCl'I-AR MKC11ANISM9
Amooff the ileiilab BffMU nay lie d!xtin^uiHhe<l Lho rientnbi mminoaljr «
oiillw}, luch Hit T. lh« itibilnuu iiiiub lu S, itnil the lio^unl L. nil dtflVritig (a
the relative pnriii»n nf the tmiifiie, tvcth, nrnJ jiiitnte.
CotiMmniits limy hI»ii be rliMiiliei) aci'Driliri^ bi the cbftracter of tlwBon-
mcnis wiucli givf- riw In ibi-in. Tl]iii< lti(:y niiiy be citlirr <^lMm w
1. Exiit'i'ivt*. In tbcM.' the chiinKt«r( nre ^ven to thtr maml by thv miiUm
utfthlUhtiicTii i.'T romoval ol'thf Rpprnpriiit*- interriiplioti. Tbiif. in uUmiu
Uk tuhiiil P, iho lijiH iin firvl rkueil. ihnii iin exfiiriilorr <-iim'nt of kir ■
driven iigntiii't them, nod U)>on tbcir beiug «iidilcnly <>pcnp<l. the aouod i
ectKTiit<^]. Similnrly, the dontal T \t genornk^d by the eiiddcD roionv&l li
(iw tnlcrniptinn c»iiBrd by the npprosiiuatioD if ihc tip nf th^r tongiti; (o li*
front »!' iho hard palate, and the guttural K bv the «id<len removal of tbt
ititerniptioD cau»ra by the approximation of the roof of the tongue to Ihi
soil null) I p.
The labial It ilttferv from 1*, inasmuch as it is aceompHDied by vibralica
of the vocal cordg (that it, a von'el sound is uttered at the aaiue time), ui
th«e vibrations (.'ontinue after the removal of the intemiptioD. Hence, B
is often spoken of as bein;; uttered with voice and 1' irithout voice: and D
aiid U (hard) vfilh voice bear the same relaiioo to T and K witliMit ToittL
The emiiinwMs conaooants may further be divided loto—
2. Jv/iirfifc*. In these the souitil is Kciit^niled by a ni»h of air throa|{)i ■
constriction formeil by the [lartial closure of the li|Kt, or by the miiiinvof tHr
toiigue Kgiunst the hard or soft palate, etc. Thun. F ts sotindtil vih«u itc
lipe are nrDught into partial, ajid not as in P and II into compbtr nppruit
matioD, and a i-iirrt'nt of atr in driven iliri>UKb tbi- Darniw«)i <>|n-i)in^. Fit
Uttered without any accumpanyiug vibration of tlie vocal corila. u r., willmal
TOicCL With voice it become* V.
Thd nhilant ^> is formed by a rii»h of air post an olietnietioD nuseil bf
the partial closure of the tcrlh, the Iront of the taogue being <lepr«na) M
the same time ; and S aceoropunicd with vibrations of the vocal nrdi
becomes Z.
In Sh lh<- dor«al surface of the tongue is raMtA m as tu namw tk
Oo between that organ and the palate for a considerable porligu of iu
■
Tb is formed by placing tlie tongue between the two partially open vm
of teeth ; and the hard aiid soft Th bear to each other tne sanie rtdat>aD>
do I' and 11.
L is produced when the passage is closed in the middle bv preasiu]; the tif
of the tongue against the hard palate and Uie air is allowet) to escape U iw
sides of the tongue.
When the constriction in an arpirnte b fonned by the approximalino iif
the root of the tongue to ibe srdl iialate. ne bare the gutturwl CH iw is
loeh) without voice and OH (as iu lough) with voice.
■'{. Befjnanti or tiataltf. iu tl)e»e, nil nf nliirh must have ^HbratSoMrf
the vocal eonli- as a basin, the ueual paasage ilirouxh ttie mouth is cJtMd
either in a labial, dental, or guttural, fa^iiou am) tbe peculiar chandab
given lo the kiuwI by the nasal cbambert> acting na a reauoance (»vilf.
Thus in M, Ibe pustai^ce Li dotted by tlic apjiroximatlon of the li{«, in N, bf
the appraximoiiuD of tbe tongue to the hnnl palate, and in NG by the a^
pruximation of the noi of the tongue to the *(ift palato.
4. Tbe various forms of R nre oflru spoken of as tvfrro/ory, tbe characteristic
aouml* l>ring caufM by tbe vibration of eon»o or other of llie parts furaitnc
a c»ni>lricti<>n in ibe vocal paMagCi Thus the Ordinary U is pn-Itiivl by
vibrations of tite jioiiit of the t^wguc elevated against the hard pnlate, tin
I.00OMOTOB MROnANISHS.
977
rguUunil K by the vibrnlinnH of tli<- uviilii nr other purU of i1k> wiillci of lh«
pbliryns ; itod id touw Inoguagcs Ihvrc Mvcms to be kd R prvrlurvd by die
ribratioDs of the lipe.
11 ie caused by che rush of air tbrough the nidvly open glottin. Whm.
in Boanding n vowel, the eooud voincidoi with n sudden chftiig*^ in th« |iti«i-
don of the vocnt corda from one of divergence to ouc of nppniximation, tUc
vovtet id |m>nouiiced with the gpirilu* 'ujier. WbiMi the vucnl oonia kit
bruucbt tu|;cth«r belore the blast of air begins, the Towel u nronouiiced with
the «7>in'twi /rnu. The Arabic II Is prudiiced by closing lli« rima vocAlis,
the fjiijfKitlis aud fulae vocal cords being deprased, and eeodiog a blast of
air thniu^-h Uir rimti rusgiiruturia.
On nuiiiy uf tlui abuve ixiitils, bowevor, there are gnat diflerencee of
ofitniitii. ihc dbicuMiiiH of uhtcb Hs neli as of other more rare cousoiiaDlal
■ouiida would l««d us too far away from the purpose nf thid book. The fol-
lowing tabular •taionient must, iber«furt:, be regarded ad introduced for cou-
venienrv only.
Rxi-UHIVES. Latiait, witliout voice .... P.
JMiaU, wilh mice D.
Drittal; without vnicP . . . . T.
DtnlaU. with voice D.
f}vltnriiU. Tt'nUoMi soiix .... K(bardC).
QillHriiU. nilh voice Q (hant).
Abpibati». lAbialt, without voiw .... P.
Ijabialt, with voice V.
DatiaU, without voice .... L. 8, (toft C), Sh. Th (biinl),
Detttatt, wilh voice 2. ZU (Ediuhiv, ihePreoch jl,
Th (soft).
Oniffum/f, without voice .... Cll (as in /ee&).
Oaffwnib, with vcMce Oil (ss in ft>«ijFA),
OXAim. Jjobial. H.
itaiioA N,
Oyltumt, >■(),
ViBXATOXT. labial, not known in Riimpean «|>eeoh.
Vrulal, K (common).
fMtvral, K (guttural |.
Whitperinj; ia s|)cech irilWit any emjiloynHTnt i>f ihi; vocal cords, and is
rllM-ted chiefly by Ihe li|ti niid tongue. Hence, in whimpering the diitinelioii
bt'twcen omiwiiiuMl.* needing and tlxine not nceiling voice, soch as B and I',
bvcomea for the niovt |iaTt lost.
IjOCOhotor MK(;iiAN[eH6.
The skeletal muttclea are for the most pnrt amnged to act on Ihe bonea
and cartihigi'ti a* on levers, exnniplra of the first kind of lover being miv,
and ihosc of the third kind, where tho poncr is applied umer li> the fulcrum
than is the iTaght> being more common than tlio second. ThU arines from
the fact ihnt i1m> movemcDts of the body are chiefly directed ti> moving oom-
Saratively light weights through a great distance, or through a certain
istaoce Kith great precifion, rather tluui to moving heavy wctgfau through
a short distance. Ihe l\itcTiini is generally supplied br a (perwct or imper-
fect) joint, and one end of the acting muscle is made ust by t>eing attachcil
either to a fixed i>oinl, or to some |Hjint rendered fixed for the time bejiig by
the contra«tion of other miuclet^ There are few movement* of the body in
vhich one miucle only it concerned ; iu the majority of cases Kveml muscles
act together in concert ; nearly all our moveuenti an coordiiuUo iuov«iu«Dt».
62
978
SPECIAL MrSCl'LAH MRCIIAMnMS.
^Vltere gravity or the elastic reifltion of tlie pnrt» mrted on dim not ailiirf ■
Millideni tuiIaKunum tu the coulni'tioa of a (nuMl6or group nf iiia»cl«s, ibl
return lo tbc oocHlitiou uf ei|uilibniim i^ providMl for by lfa« Bctiiiii piUur
ctiutic ur ooulractile of a set of aniu^oislic lumcka ; thii ia Men in tht- ca«
of ibe fiu'e.
Tlie ertct pnflarr, in which the weight of tlii' body is borne by the jiUdlu
arcbck>, Li tlie rt>aiilt of a Miries of coDtracttoiiM »f the iim»i>l«« >tt tlie iruid
Hii(t legH, hiiriu^ for their object the keepiiij^ tht- bixly in Mudi a poiitlnn tbi
the liiiL' (if gravity falU within the iircu of the fwt. That ibis Aom ntuii*
mui>ciilur L'XcrtioD ii aliDwn by tlw facU, that a iienoii uIh-ii titAOiJing p«-
ffN'tly lit rtwt iu a oumpletely hiiluiiceil ]in«ilt<tn fall* nhen hv become* ub-
miwriciuN, ant! thatadMuI Kxly cannot Witetun lU feet. The liooof gnrilT
vf Ihf ititail latN in front of the iwcdpttal articulation, a« ■• abown bjr thii
nodiling iif the liMid in ntcvp. Tlx' trnlw i>f (gravity of the coiobiiMd nnd
and trunk lic« at uhnut the level »f the entiifi>rm cartilnicc, in front of tbf
tenth dorail vertebra, and tbc line of gravity draun fmui it |in««* behimlt
line (oining the ceiitri« of the two hip-j<>iiil«. xi that t)K> <irec-t Iw>)y would
fall Mckwairl were it not for tl)e action of the must'li'si pftsning from tk
thighs to the ]w\\u aMistod by the anterior tiganieDie vf the hii>-ji>iDl& Tht
line of gravity of the combined hcaid. trunk and thighs falls niorrovtri
little behind the knee-joints, m that «om«. thiKigh little, muscular osertioa it
ret|uired to prevent the knees from being bent. Lastly, the tine of gtanij
of tile wliolo body iitii^se^ in front of the line drawn betireeu the two anklv-
joints, the centiv ol (gravity of the wliole bo<)y being plaoe'l at the end u(
the sacrum ; hence some exertion of the musclm of the calvea is r»|uirad la
prevent the body falling forward.
Iu uiiUiuff, tliere is iu each stop a moment at which the body rwts vcfti-
vallv on the foot of oue, Miy the right leg, while tlie other, tlie Ivft kg, ii
iueliue'l ubli<]uelr behind niih the hvel raiMid and the (oc mtin|;oaiW
grouud. The len lej;. xlightly llexeil to a\-(>id eonuci with the grognd. »
then swung forward like a [wnduluni. (he length of the >wing ur atap inhf
d«t«rmi»«<) by thv length of the l«g; aiul th« left tov' is hninghl to ikt
ground. Ou thi« lull ton as a fulcrum, the body is moved forward, the oanir*
M enivity of tbc bo<ly divcribin^ a cnr^'e the convoxity of which is upwud
ana tbc left Ic^ necrssarity becoming straight and rigid. As the Ifody iniini
forward, a pom! n-ill b«i reached similar to that with which wv anupoMd ik
step to be started, the body resting vertically on the \ct\ foot, and tbe rij^
Ifig being directed behind i^n an oblir|ne position. The movement on lh« wl
Jbot, however, carries the body beyond this point, and in doing »u swinj;! ih*
right leg forward until it is the length of a step in wlvanc« of its nreriNi
poattoD. and its toe in turn forms a fulcrum on which the iKHty, and with it
tlie lefl leg, is again swung forward. Meuee in succeasive steps the eoolR
of gravity, and with it the top of the head, describes a series of ooosecutin
curves, witli their convexities upward, very simitar to the line of flight of
many birds.
Sinee in standing on both feet the line of gravity faUs betweeu the im
feet, a lateral diiplacement gf the centre of gravity ia necenary in nrder w
balance the body on one foot. Heuee iu walltiag the centre of gravity lU-
■oriliea not only a aerie* of rerlleal, but aiwi a series of hortxoiital eurvw,
inaamuch as at encb ntep the liii'- »f gravity t« made to fall altematelr iia
each siandlug foot. While the left le^ is swinging, the line uf gravity fslU
withiu the area of the right font, and the centre «f gravity la ou the right
side of the pelvij. As the toll foot bucomi» the standing foot, the ceoiiw tif
■ Tlililn<n<al«>iaTlM|B«kiitiliuuMI)e<knatanierlkuinMactaBlinBWce; niuH|i««>,>lsraItM
ti^l lo III* crauiul fliM. Uitt wnUiA wait Uw tat i-oalnt kiMt.
tOCOMOTOR MECRAyiSIIS.
I
I
I
itrnvily U stiifttNl lii tliu left *ii]e of the pelvis. Th« ftotud curve iKvcriboJ
hf \\w wnirc of f^ivity U, lliercf in>, ii Romcwliiit compHc&Ucl one, Wing
eompowd of vcrticnl ainl h»ri/.i>iitHl factor:). Th« iiAluml tlcji i« llic ono
whioh >■ ilct«rmini-jl hy lliu leii^i.U of tliu awinjiii^ \cg, shioe till* ncU il* R
prndiilum ; nn<l hvna: th«i Kt«p of n loiiK-lvegoil pcnun it nnturully Imigur
tlinn tliiitof ft purs-)!) wilh short tefr*- The length of the t'jep. hoiruvdr, may
b* diminished or iDcivaiBi) hy n direct intiiiciniir cflr>rt, u* when a linQ of
Boldicrs kocp Mcp in Mpite of ihcir hnving Icn of diftorcnt length*. Such n
mwle of mnrching must obviously tij fiit)sniu|, inumuch m tt involves au
unDeoesBary expc^nditurv of energy*.
In aloir »-nlkiiig thorv it nn approciablo time, during which, while ona foil
M alreadv in pnsilioii in serve iis u fiilcmm, thf other, stringing, f )o: h» not
nt left tne ground. In fiwt wnlking this period is so much reduced Chatooe
Mot leaves tn« gniund thir monieni the other tiitiche^ it ; hence (here is prao-
tMMlly no period during which both foot ar« on ihe ground together.
When llie body b swung forward an the one foot acting as a futoriiru
will) Huch energv that thU fout leaves the ground before the other, swinging,
foot has mched the ground, there being nn interval during which neither
foot a on the ground, the penon is said to be rumiina, not walltiog.
Id jumping, this proi>tilsion of the body lakee plooe od both foet at the
attoie tim«; in ho|^ing, it is effi^cied on one foot only.
The loooiDotJi>norfour-foot«d aniiuale m neeensrily morecompliuat«d tiian
thai ofiDnn. The ninple walk, such as that of the horse, is exeout«d in four
liniM, witli II dittgonal lucoeMion ; thus, right forc-IeK, left hind-leg, left for«>
l«g> right bind.lvg. In the auhle, such as that of the camel, the two fe«t of
the Mine *iili' are put dowu at one and the Mme time, this movi-mmt being
foilowfv) by a similar mnvument of the ollmr two legs ; it enrrcapomU, there-
fore, v«ry closely [i> bumim walking. In the Irot, which orrctpondt to
hiiiiian running, the two diagonally opposite fiict arc brought to the ground
at the same time, nnd the body is propellml fnrnard on them. Concemiux
this, however, as well oonceroilig the still mora complicated gallop and
canier, observers are not ftgreod, ftnd much disousion ha^ arisen.
The other problems connectod with the action of thit various skeletal
muaclee of the body are loo s[>ecial to he canei^lered here.
BOOK IV.
THC TimVS AND MhrilANISMS OK RKPRODUOTION.
CHAPTER I.
I
I
I
ORGANS OP BEPBODUCTIOS.
Manv of ibe indivIJukl constituent pnrU uf the b9dr nm ca|<able uf
Kprodiiction — iV., tlicy vnn givu rim tu parla like ihciDWIveR: or thtj are
ca|>abl« of ifgvncnUin— 1.«., their places ciin b< taken bj* nev ptrU rnire
or len clowlr irm-mbliiig tlinm»i>lirei^ Tbe ctemeiiury tiwuea undergo iJiiring
liTe II vciY large amuuDt of rogencnition. ThuD. (he old epitlielium ocntei
irbich fnllBwar from Ibp Biirfnca of the body nre Hucce«<:lod by now st-alM
from tbe uiHlerlyiiig liiri^ra of the opidermiit : old bk>iid>oorpuSGlea give place
to avif onra; norD-uiit miiHclos, or tlutse wbith have fuled fh>ni diswse, an
r«ii«ir«l by the lu.'omtioo uf frceb Hbra ; divided iiorvei graw agaio ; brukpD
boD« arc unilod ; connective tinue aeenu to diMppear and appear atni'»t
without limit ; new mcTvtiog cells take tbe plan of tlM old unea which are
cofet ofl*; in fact, with the exception of aome cne*. sueb a* cartilage, and
then doubtful esceptioiu, all iho-e fuufUmcDlAl tiMue* of the body, ubicb
do not f»nn pan of highly diflvnTiitiatod organs, arv, within limit-t (ixeil
morr by bulk lima by anything ol«o. capable of regeneration. That re-
generation by NibnitutioD of molecules, which is the basta of all lifu, U
aooomiwnicd by a r^neraUon by aubstttutioo of m«M.
In tne higher animals njffenermtion of whole ori;aiu and membsra, evan
of ihoM nhoM conliniicd njDCtional aetivitr is not cwential to the ««U>
being of the body, is never witneseed, lliuugK it may be wea in ibo lowir
animals: the dii;ii« uf a uewt may be restored by growth, but not lb«M of
H man. And ilio n-pair which folluwa even |>artial <leurueliofi of highly
differentia led organ*, such a* tlif retina, i* in the hifjher animals very
imperfect.
In the higher animals thv repnxluciion of the irhole individual can be
eflTected In no otlier way than by the pruci^** of Mxiial Mnerulion, through
which ihe female ropmtcnlativt^ element or uvum i*. under lh« influrncip uf
th<> male reproMntative or •permato»)on, developed into an adult indi-
vidual.
We do ni>i purpose lo enter herv into any of the moqiholof^cal probleins
CJMtDMted with the eorie« of changes thpjitgh wlitrh the ovum becomes (he
adult being : or into the obscure binl'>gicar iniiuiry as to how the simple,
nU-but-^tniclurelcas ovnm contains within ilMlf, in |»teDtialily, all its (btiire
developinenls, and as to what is the raseniial nature of the mate acUou.
These problems and quealiomi are fully discusted eUewhere; they do not
Eroperly enter into a work on physiolojjy, eioept under tbe view that all
iological problems nre, when imslied far enouirh, pliysiolo^iicnl probleuu.
We Bnall limit ourtclvet to a lirief survey at the more imparljuit phyaio*
logionl phenomena attendant ou the impregnation of tbe ovum, and oa the
tliitritton and birth of the emhrvo.
THK Tiasues AND MBCUAM8MS OP RSPRODCCTIOX.
Th« frmale orffatif of ijtnffalion lire nnntoniiciillv rlivideid Into ihf inv
and exteroa) organs. I'ho Inltcr conifiriM* Utu Inliin untjorn hhiI minora, lla
clitoria, the hriiu'n, tlio luentiu urinanus, tlic viilvii-viiciTiul glumi*. awl ita
tiiurous aD*J £«bnccoii8 elands wliifli arediHiriUiitnl in tlic rmioiuii lurmbriiM
covering ihe parts. The rxtcmal organs piny n vpry milwiiiiiiry |mri io lh»
function of reproductian. and tli«}- will bo jniwrd \>y witli lhi*( brief doUc*.
The inieroal argana comprise the vajpiia, uteni«, PaUupiui toboi,ud
oraries.
The vagiDa is a niuBculo-meiubraDous canal, about four to six incbn laa^
directed obli(|ue1y upward and backward, and i-xteodiD); from (he hymCBM
ibe cen,'ix iileri, where it is atiached at a poiat a alwrt distance aboTc iht
Km. Ml.
U
W
^-
U
V
intoBUounc V»a ormE I'lsniB isum AiTB9iiw«n.ul
(Frou Qutol.) HMir niKunlilM.
Ttt elcniiBU-lu|>i«iiMrlof tht ntpjin hoTV boi'n UM o|vn br nmnvliig lli*p(Mnlor«*U. IM
tUiupJin luV. rmiiKl llsuiniit, •OiloTurlM IIkuuuiI Iuk Ivcn cnlilwn.and Uio Imad UH'"*
mnorr-l in itic kn iidc; u. ilic ii|>(wr i«n il ilii ownit; t. Ibv tcnli epK*"" *-^ i» iiiiimin
Uw lilUJu^ilui >t>«)HiiJt llivulcTliii>i-«Tit)^|] thciim, atiil Wm dllstlon oC the conloil cntlr i>Bkai
ragii, urmaj arbor tHk- r. upr*r pan nf Ibcvaicluii; ml, F*IlD|4iu lube or otIiIufI : Uw iim-d*
oniiiMinlcalkiD of lu carJIjr vlth iBmoI UiacORiunr lliv iitpni t oo «a(k >lila Um«D: t. nniigtn^
iiwDI:l».IISUiieiilrtr lliBonry; 0,<ntTf . i wide ourlcrput or Uic Tl«l<t 1^nflf*ui lti*«,lL<a
nnitelitl*<l*alnmlt]'-. jin. twoTBrtuin ; Koavotxtia br«MI>b Ovq<in>ll) rxml tiDniuriad vnili
bmul UfKIDCDI.
OS uteri. Its iralli cotiMsit of an cslenial cost of limgiiudiital mutcultf
fibres, a niiddie eri'dilu c<^at, iind an internal mucwis coal. The miiOia
nK'Uilinaiie id cviulinumi.i hclow with ihat covering the extcruitl geuiult, s»l
alxivc with the munius nii'nibrsne lining ibc uicniB. The anterior nntl ym-
tvrii>r «urla(.-CR are marked bv longituiliual folds or raphe, from wliichs
numlwr of trdjisvcne fi'l«ls are giveji od'. Tliifi niembraue Is pruvi<ie(l nith
niuctMis gliiiidx, ami is lliickir covertd with senailive papillcc.
TIh! uterus is a flattened, pmforni. muscular or-^an (Fig. ^61). Aaa^
lumically it is divided into the fuoduf, neck, and cervix. TIh- nevk indicnM
the point of division between tbe lower coDslrided purlioii, which i» th
cen'ix, and the upptr expanded portion, tlie fundun. Tk« cervix, which
extends from (he neck (o the end of tbe orxan, projects into Uic va|;iiM,il
which point it is marked by a triiDsvene ftMiirc, uallvtl the o> iteri.
The cavity of the uterus is soinewlinl triangular in shn|>c, and rery modi
ANATOMY OP THE UUOAX8 OF OKNBRATlOy,
U83
Rlten^l •nlPDJ |«iBipriorly. Tlie iaft-rinr aDKl" of ibe cavity i» ontiuuous
nilh till- i-iwhI niniiiii); tbrciuglt llie M-rvix bi tbe vajfins. The superior
anwlMiireMlIci) ilicconiua: iit tlii-boiioia itfradi iian orificeof a Fallopian
tUM. Tbe utcriM i* coaiixMi'd nf three coats : a veroiiii (fomied by tlie ]ieri-
tmictiD)). n niuHrtilnr, ituA ii miioiitiii iviat. The miicoua rant a L>outiDUi>iiit
uilh tliitt lining ilm Fiillopisn Iij1>n> nn<l vnxiiiii. It u (.'4iv«r«il nith
■ cilumnnr cilinlt'il r|>ithuliiim, itod, if cxniuinvd with u I«dm, thv openiiig^ of
ft ibo mucotic fullidcs will W mou to be very timfuiwlv (ti«iribul4>a over the
I Riirfiiec. If n vertical wction be in«'l«, aa in Fig. 26'X thv t»biil«« will be
Piu. Ma
W>
;l
*s
"m
RniKW or ma Ukinii Mumnavi: «■ a liini> I iu>a «> mi. PuiKdi «r UuiacMciiM Thm-
■ulTT. lAMttiis ihc ■mniFiDini* wiil othrt iimilMrUnof ihc (lanih. ■(.<(. -il. with ItwhoiaM*.
«.«. n,OB IhclnEcmai larHu'ol' Ibconwn. Twiue ibu nalunlilTC.
I
Been to be amnijet] pcrpeDdicularly to the aurfaw, having a wary coiinr.
In ih* iinprexDaied iitenis lli«y become much swollen aod eolarfpfd. Th«
niucoui membrane lining the cerm, on account of il« peculiar a(>)ie»nii)ce,
is (rallcJ the 'irfti/r viltt uicrinaa.
The Fallupinn tubon (Fig. '261) are abont four iuehca in length, nml
vXlMid from the ciimua of the uterut tu the uvariet, irberc thcv en<) in
enlarge*! rxpnndnl extn'mitic, the niarginn nf which are noven^l bv long.
•trader pmi'nwi^, oni- of thcni l>ciug connected In the nvary. Tlii« jKirtitin
of the tube if rallc-il tin- finihrintecl extremity. The luhw nrw cnmpoerit of n
M!rou», muMuIiir. mid iniicoun liiyor. The miicnnp momhmne i» coverwl
with cilintrd oitutniinr t^pilhdium.
The uvarim I. Fig. 'Jiil ) nrc Unliencd, ovotclal I>odic«, which are situated
On« on each aide of the iilerii», and rncliwrd in thn f»li)# of the hnrnd li(^-
menla. They are each ponnpclcd with ihr nlerux by a lignineDt. and with
ihc Fallopinn tube by one of itc timbrin;. They each connirt of a lilin)U8 cual
((■(fii'fii alhuijiiii-a I which encloses the *truniB <if the organ, i, I'*ig. 'Hi'l. 't The
■troma is coniprHed of a soft, vaiteiilar fibroiis tiMiie. having imiiedded in it a
nutnber of miall bodies, called Grnalian vesicles, which arc in divenr dtagea
of devel4^Rient. Theee veifeiclee cinintence their development in the deeper
portions of the orary. and as they approMch maturity gradually make their
way to the surface, where Uiey project as pmniinences, and tlieir capmile
finally rujrfuring, diachar^je ibeir cunieuts into the Fallopian tube. Each
vesicle ct>u»iets of an external coat furnieil bv the ovary, an inicnial c<>at or
capsule, and niihin this a layer of cella, wliich constitutes iIk- metibraHa
r/rniiulomi. The interior of the veuicle O'usisis of an albnniiiimu lluld, in
which is sUHpendetl the uvule.
Tlif mnU /fenrntlic ortfint. Tlie name physlologicnl intere*! in not centred
in the ninle orgniu uf gruL-ration as in tboee of the femnle, the priuc)|>al
iotercM being conoculratvd uiiou the organs which lu'cretc the mali- duid by
which the ovuli- in impri'giinied. Our remarks will, then-fore, be alnvNt
■Biirely cunlim-il to the organ* oooccmed in the wcrvtion uf ihi« llui<I.
084 TUB TISaUBS and MECHANISIIS or ItKPItOntTCTIOX.
Th« ntalfl ornns comprise the penis, or orgnii f>f <.'o|)ulmion, iI)l- prcwUie
hhI C'onpiir'i gluidA, the teeticles, aad vaea defcfWiiia tuul vwicuuB ivin)-
■titles.
Tlic pnwtHlti Klaod kiirrrtuiKb the neck of llie bioiMor and commenoeHieM
i>f tlio iin-lhni iViic. '2GA ]. Il wi-rolvs ii milky l!ui<l, irhich u oonveyet) Itj
(hu proctutic iliietd M Iho Hour of ibi: urHbra. Cowper's f^laixla nmao
rtusll glaods wbiebaresiliintcd beliccvii ili« layers »f ihe deep perinea) Cue)*
at the aiiU'rior jiiirt nf llie mombranoiu urolhrn. Ttiev secf«t« a visdd Huij,
whidi is wiivrj-t-d bv ducts to tbo floor of tlie iiretbra.
Tlw tcatus or Uwttclea are tiro small. Ilatleurd. oToittul );litu<ta. nbleli an
VmopiMxnimur ^aiPMicnOVAiiTorTitRt'AT. lAnn^iiniiH.' i^
t. onler covtrlnc aud not hunUc of Iha ororr ; t'. alUdbnl bnnSn; z llf i-nrun Mn^«. imrf
lIV ■ Uliioiu bIkI nuiiiUr itnitflupr; A. KminKiir «»)Biiinra lylnit cili>ni*l Wlluf nimiiv •
bee: CoilowiiK ulil(i|| birr Iwyun ui crtluricr vij [o low inon iVviilir liilnllw uruy : 7. aictcm
ruuiii] nlilrli ilii- llnstMn (i.itllnir uiid tiiiiloi rnniiilm are iinin Itonaal. •■■i) uliVk Mn (Wiri
w>mvnrhiildot|iariniiriLvivrafy BL>d Arv»iirr<i:>aiiilDi]b^UJontbnm'ktrQTua; K iUiweartrAiioHtOtiMa*
MIUcIo wltli llii' litiitii tntxvMiiil ill III.' lnlTruf •-v1l> roiKIIIIiUnc Ibi- pntllBPraV' <U<v ^ t, ntm^
•Anuucd lOtllvli! ixiiiUIiilOK till' iiTiim. clo. ; 9'. m Rilll«lc rrnm wUcli the ovum hiB wnMNMIV
wcBlqJ; lU. oirpiu luteiiin.
«ilu&led ill a niiinciilo-menibniuoiix ngueb. cullwl tbe Krulum, and watfteaiel
by th« epormiitic cntih. 1-::ich lf>ttid« contltla of t«o paru : tb« glaud jDfnpw
and tbo opi<li<)ymw. The ginnd f Fig. 205) in oompMod of an otitor nbraw
coal, tho hini'i^fi •i/hu^tnm, thin being covered by a scroti* mombnuic tbt
(tiJitoa wyini/u. Tbosubetanoeof th«Klai)d pnnnii'ts nfa uuiub«r of pynm-
ida] lobular divioioiis, which are Mtuat«(t wilh their bnsce tOwArd tlleH^
fac«. Ench lubule i»ci>nii)OSod of servral convMlud'J Itibiili urmini/cri.aai
arc oeparated from adjninitig l»bul« by prolon^Htinii of fibrous lixuie fraa
tbc tunica albuginea. Tfie tubules arc ootuixk'hI of a b»n)ng«aet>u> biistiBttl
iuiM)]l>rane, which is lined by granular nucleated epillKlium. Id tbo >t|i<^
of tbc lobules tliey have a stnizht caur«e. and form tli« vom rrcln. Ther
then cnl«r the flbroua tiHUe of the mediastinum (Fig. '.^6-3), and fvrn s
)>l«xu* of tubed calleil the rfU UitU. nhich end in the upper pan of lb
medianlinuro na the iimu rjfrrenlia, and iheiw becoming verr rauoh convoluted
form ihc gtobia major or head of the epididymis. The tubule? of the (•lobw
iniijor iiniiu to form a Hn)[le tube, which i» very much convoluteil. and cio-
liiitut*^ the Irudy and ylobiu minor uf the epididymis, and is then cHitinusd
I'rum the glnbu* minor to lh« baae of lli« bladder as tbo excreturr dflcf or
III* deferea*.
The vt* dtjrreiu, eoURienciog at the j^tobus minor, ascends (a tlio potttriot
tWATOlCT OP THB OBOAH8 OP OByBBATION.
9SS
part oftbeapeniiitic c^>^l (I)ri>u2h the .Hpinniticcaimt iFil»thepelvii. wheni
It niDM lo tli« Imw of lli<- )tlnd<[i;r imj becumei culniyeJ, uciiulaUi), anJ
narruw«<t,Mii] j[>iu«<rilh tlikMltirl or (ho vuicula Ne mill alii to fcirm a commuii
ejsrataton- tlucl. Thv walU of the vii» ilefomuare OLimjx^wd of librauautd
mtueular (iMuft, which i« lined by a nmcou* mcmbninc with oolumnar cpi-
iheliuni.
n.1. TH. na. «a.
9
r>9 >M.~TklE OLMnr Till M >!.■ UUIMIKK, WITM THK VRKOBia HWUULM JtXD PBOtTtnGLUIC.
I'tAflfT MllUU. I. <IM orlnuT blaililer : 'l.lh« l(>n(>lu4lnftl lajvr Of muMiular HbtM i l.tha|inaUW
1: 4, BiMBltnuiaiu potUoa at ihs uniiim: X the urcMn: A. bloodnacU : T. lei) ; I, itgbt iw
mm: >. !■« HiBlnnl nwila in tM iMtunl inriuuni iu,4u(iiu4>oaUior1ui«f tlia IcA *lil«
tnTTMnt (he prattaM cUnil i II. ilgW wnlnil *«ol«la InjBttod and unrartllvd i 12, U. blind
» winba <if *Mleu1» -. II. rl(lki<luDiiu«fuuktunBili«T«nlncUie|if(M«o.
ne.SSL— «, lobuIiM: b. tom rvou: «. nwdlaMlnuui ^ <f, vua tfltanuiU: r. b^T f^ cpldldjDtU:
/ nu loMoi 1 1. ctobiH minor ; A, ni dchicM : I, lauli'K albu^iMH ■nd II* Inicrlobalar rtHeellon* :
E /. ■Ii>taimijar.
■ A rin. y« t<
1 -t • ' •
■ 4, 8FMK110M1 rnoic TIM tlnuM Vm UNnanin. lAiivr Xau4s«N.>
' I. nwyii»rt »" ■■■ — *" ■ ^MitDllk>lMndlaMMtn:a. mUwiUc;k(lomkbn<»
li.8rc>iuTKCKUjiA)<tiiMMUi«aa<iorTnaliuu.UiniuHMira Pktiuu-mivt.
(Alitr Kmuku^ M-1.
l,it«ia«llo otOk wHkOMM taw Dudol. una oftkiatfawr; 1.1, hot ■uakC witb (wciaMM
lU IbnMlni : I. tlw BlawHIM ttaa^Uii mH ih* bodf ttldtMil : 4. MamtWi Mitrlr <1amd(aa.
le iwnoN&SMmitwifMaratwuQtoiigauJ MceultCAd badiet,pUccd oxtonul
to the vaM defereotia. Tbo structure of the aomlDil T«gicl«a b simiUr to
tlial of the maa deferentia, cuiui«tiiig of a fibra-musctibir «ral1 linod with a
986 THE TISSUES AND UECnANlSVS OF BEPBODtJCTIrtt.
inucou* ni(iiibriin«, which u covi^red by (jranuliir, nucleated, poly§:ocial qj-
ihi'lium ocllf. Tbive orgnnn Acrve as rei-epliclea for tbe BeoiiRnl Quid
ri-eivti-d by lli<^ t<-Klci>, luxl iiL the »aitie time prnluce a secretion of thnr own
wbicb in mUlt'il to It. Thd vjii<:ulnlury duc-ts, wbicli are tbrmed by tb« unk«
of tbo duct* i>f tbo viiMi d4;ti:n-iititi und veeiculx teniinaltd, open inlo ibe
prwrtatic portmn of lh<- iin-lbrn. Tbrlr ettatti are ihiiiiier, but bare aawDluUjr
th« nmv xltucltin- »» tb<i vii>a dcfrn-tilin, uitli which tbey are conUnuoui.
Tli« grmina! jtui'l in n (imiplcx wcrrlioD, being cc>ni[Htf(^l of the anMunical
clrmeiit* nf rprTinaioioa, which are IbrnM'ri in the tntei. luid of the aecrelMiu
of ihc VM* ucforcniia, vwiciihi,i wniiwiiUv, the prwtiili' and Cowiwr's glaadi,
and iho niueoutt glnmU of thi? iin-lhm. The Mmiiiul Siiid ia of a thMc,
wblliih, Ftrinlcd nppcnriinrc, and, if cxninimd niicrtiw.-opic«llT, is teen to
rotilain innumcntblr bodicv nbicli iim in aclivp moliim. Thcac *n tW
t>po^^mtol;oid^ and nn ibc cwcniinl mali? clomrnt* cinrrrui'd in ihi; ftiounda-
tion of the ovule. Kiich of Ituve bodies (Pig. '2fi^> c<iu»iel« nf a flaUtoed.
OToidal hc&d, havin)^ at Us bo«e u tapering caiidntt- appcfldue in acttre
vibniiilc motion. Theee aiialoniicfll elem<-nt« were nl tirM caoriwrcl aniioot-
cuJA, but they are now looked upon us free maseee of protoplasm with ciliu;
apneiidsges, which endow them with the power of migration.
The fipcrmatoEOA are developed from the nuclei of veeiclea which an
formed in ibe tubules of the teatea, Tbe nuclei are metamorphosed iau
the heads of the spcrmnloxoa, the ciliary appcaidages being afterward derd-
oped as a eon of ouigriiwih. Different stages of tbe developroeni ui
other interesting features are »liown in the atiuve figure.]
CHAPTER II.
MENSTRUATION.
From pub«rty, which orcure nt from 13 lo 17 yrtin it H|re, li> ihu
climftcUric, whkli arnvi-» nt frum •l-'i to 'tO years of iige> ihi.- humau reninlv
is Mihieci to n nKiiithly <li«cliurgc of ova fniin the ovnrw». iiccoiiipanicd by
special ehmiftM, not ooly in lhu»e orf^nOB but fll«o io ibr FallupiaD IuIh*
and iitenjii, ni well ns hy general cIihii^fh in tbe body at Inrgc, ilie wlioir
eooMilutiiig " nirnslniatian." The e«Bential event in nitrnitruntioD is tlw
jK nt HI) ovum froDi iIb GraafisD Tollicle [Fig- 26T]. The whole ov&ry
at this tiau) bccomce coDg»ud, anil the ripe follicle bulgrs from ilt> eurfacv.
The mcwt projecring portion of the wall of the fnlliclc, whM'h has previuuely
|Fl., JIT,
Fw. aw.
I riii.'JW, 8»rnoetorG»<L*funlbuji.xaor * Uuuui. iARh VoxKiiia.) t. iinnuaurtlieuTafj
B atili tJM4*WMl*: %p(ritoaeuui S and 1. Urtx at Uie ciitniBl ouiof ilieiinuOiui ftiUk'lc: 3>.
I ntiMlWIii (nankai -. C niUd of Itiv (tnulbiii nilUcle- : T. imDUliir oiiir. ur iKirut irtoltttfvt, von-
■ lalnlnallMovnIvOi.
FM. IM^Ovcu or TtiK Soir. lAIUf Ai.B«v.) I. inffmliKl •pol ; I, fennliid nalcia ; R. folk , 1.
M(M fdlutlila : \ dlKLO* |irellgm* ; It, Kilbennl gmnuh* or nlla-]
becoine exeeatiirely Ihin, is now ruptuT4<l, Bn<l the ovum, nhic-h liaviuii l»ft
its enrlipr potiiltoii. in lying cImo untli-r thv projeclinK Mirfare of ibc roUiele,
eaoupet, together with the cclli uf lh« dinnm jiroii'jiivi [Fif[. 26d], into ibp
Fallopjaii lube. How ihc ciitrani-i^ <-f (he oviiiit info llir Fnllopian ttiW U
MCuml in not extti'tly kiiouii, 8<>ni4' mainlnin ihal ihr oriiry ia graniwil by
the lrum[M.-i-*hapc>(I fiinliriati'<l moiilh of thu I'altopian tube. ilrHf liir^icl
HD<I oitifctflvd ; tlw iiiovnnonlB nccw»arj- to bring ihi» about bi-ing elfccUd
by the |tlain ffiii«nilar tilirca pment in the nM>ui1i of lh<' inbff, (^herv, ro-
jetting this vii'ff, ami Rm:iling that the turecscencv of the lube dooa not
ucair nntil aAer the ovum has Iwcome safely lodged in the tube, euggesls
that tbcovuin is carri<tl in (he propiT direction by currents in the peritoneal
cavity »el up bv the action of tho eilialed epitheliinn lining ibe luW, cur-
rents whose drreclioii and strength *ttm, as shown by cipeiinient. lo b«
nde()iialo to carrv into the uterus particles present in the ptritnncal flnid.
Arrived in the lube, llie ovum travels downward, very slowly, by the actinn
probably of ibe cilia liuJii;; lite lulie. thtinph poMihly its prt'grets niay
oocoaioaally he aMialed by the peii^laltic eonlraclii'ne of the niuccular walls.
The alay of Ihe nvuni in the Fallupiau tube may extend to tevcral day*.
There ia an (truBiim of blotid into the ruptured follicle.whieh is §uhfci|urnily
tiillowed by histologitul changtA in ibe eoaU of ihe follicle letulllng in a
MRKSTRUATIOK'.
imrnin tuteum' [Pig. '2H!>J. The disebargo of tho ovum i« ncompMiw)
only by a coiigcetion or erection of the ovary and Fallopian tuUr. bul abti*
by marked chun^efi in the uterus, e!^[>ecially Id ifae ut«ria« tniiiv>ut cd«'Iii
brnne. While the whnle orffao bscomea oangeeted and enlarged, tha niacKii
membrane, ikml wpei^iallv the uterine glano*. are distinctly hypcrlropht»l.
The swollen iiileriial aiirfdce is thfDwn into fohU which almost ouliUir4tr iW
csvilv : anil a heiunrrhuifio dlsi-h^r^'e, oden cimiidenble in extent, couftilnl-
ing the meiuimal or catumeuial tinw, takes place from the greater port of
ita Burfaoe. The blond lu !t pna^ei thmngh the vazina beoomaa auai««hit
altered by the acid secretions nf that pauage, and when soaaty coa|[(ilaUi
(Be-
lu«^ » VKimiui.eaLinair>. Al n kahnwii ibc lUU »f (ho IblUrit IsmnlluHr aAar tlwctinl-
■l(in of Uw ariitft It* Mflty bolni nllvJ w<ili lii>»l aiil no ui*n«lile Inirawe of Hi artlMW
UiiUw litvlns yM ■■t*Dpliipe . ai (s IMakonliif ot ttitt hnlnx bu binotna ai-parfsl: ti f ittafW
(11 pn^t ruldi wbic'h nrf <|.>i i>iiii<il ut il, sitl ilio clulnf (ibtod b ibaortL-] ^t" >'»'>. anil MO*
MinallfMiJwulnriivd ; ■ Piiilla'i(i>i» nf Uif urns I'luivo. ai *iu>«ii at r, /, u, *, tjtiiu ih> lui^
luMom. nllb Ite dalloate d«lrlx.]
but eliehtly; when the How, however, is considerable, distinct dot* may
nuJce t^wr'appea.rani«. The iwolleu and hyperlrophicd mncoaB niopikiane
then uudergoea a rapid de^nenitiun, and b shed, passing away aomctirao ia
distinct ninaaa). forming the latter part of the raeDHtrual flow. The lift •>(
the mucous Dienibnute U nn complete, Uiat the buses only of the iitenne
Slandf aru \e(i, and fniin tho epithelial oella lining theec the regeneration uf
i« new iiieinbrano is #nid to take place. It ia not certain that meostnia-
tion, in the liumiin subject nt all evemtat, in always accompnnied byaillt-
charge of an ovum ; inJocd ca#cs have been reconled in which menBtraatiao
oontioued after what appi^arcil to bo complete removal of both ovariea Ati
(■ Tb«litl'>afD(labulatitiUEiB:tit tiy lUtibQ liipi
■nil lilMUa cf Ibc ooQ-pnf iiaui ai'l |i'''V>o"i n"iu:
<lw pclMl|itl dllEfl
bM
At Itr. nul </ MrM
■imfei.
OM lamA.
AnauMftL
COkm U-TBi'it or PuoMawT.
1
i-y«iT< Iji-cei'u or Mi:-iyrHL'Anoc(.
Thnw-iiiiinin at mi liu'li III •llaiii'
wall trttclit nt- ! I«Tirirr, toutoIuimI nlUi brtrU f*
iiltik. I to*, dot •1111 rwMltli.
lt>-i'".-fi v.;r^r,pniliUMioIan liitlj. , 9o*«»«ljlMlia of an ln«h iH *adK»«.
iilAuul uloaMi- WWcJillaa «all ■■'Vt^l r*^' . f*^
BiKl ^^' ' - ■■» li rni of MM*
Alwiit.
AbMM
I on,. I. , .., ,■! .-,. Maml
ci.. ,-ln<l>»
In iiilanli)}
any bctfM tbIIo* «Dl»r, i
MKNSTKl' ATIU:'.
'Jiii>
II weni» pr(ihnb?e bIvo ibnt under ccrtnin drcuDittKiuefi, e. g., coilui, s dig-
cbarge ol an uvuiu uiay lake place at otber tmn iban at the niemuual
jxTioi). Siiico, biiwe\<^r. ilio lini« during wbich both tb« oviini and rhe
([tcntisiouyiii may rfniiiiu In (be lenale patftgEtt aliv« aitil imtciibDally
atimblu i« coaaide ruble, probubly cxtendinc to soiiie days, loiiuit elTecicd
ritWr Mine time aHer or foin« time before the menhir mil ebtat>t- ul aii ovuiu
nii^bt Irud to iti]|)K)tiiiilioii aud tubKQucnl devi'liijimciil oI' an ttnbryo;
hrnci* tbc fiii-l tlint iiiifin-^nution may follow upun wiiub ut tuaw time alter
or hefiirr intnotruntiuD \» no very co^'ent Hr)(Ui»fDt in favvr of ibe view that
eucb u 0(iitii> bay caused an ludti^udiDl eicajio of an ovum. Tlie etcape nf
Ibe ovum i» mid to i>rende, ralbrr iban coineidc witb nr fotlow, the min-
mcnial flow. If no fpcrmntoxim vnmv in contact with ibL' ovum it din, ihv
uterine mrmbraixi rt'lurns to it* normal condition, and »■> trace of ilic di*-
i'liar^ of an ovum it lell, escept the corpUN luteum in tbc ovmty.
It ie obviou* thnl in lbf»^ jibrnummii of menxir nation we nnve to deal
with complicated rrllcx avlionx alfi-cling not only tbc vnKiilar supply but,
apparently in n dirrct mnnner, iKc nutritive cbanpt of the organu con-
cerned. Our studies on the nmotis action of M'en-tion render it co»y for us
to conceive in n general nay how ihe stveni event* are hroiiehl iiboui. It
is DO more difficult to Mipj^ne Ihnt the xiimulut' of ihe cnlnrpcnient of a
Grasflui folliele chufcb nutritive as well as vuscular chtmges in lh« uterine
mooona iDemhraue, iban it is to suppose ihnt the stimulua of food in the
alimentary canal i-auav tboce uuirilive changes in the Mlivary Klund* or
paucrea* nbieb couslilule fecrelii>n. In ihe laller ease we ciui (n r-uine
extent Inire out live chain of events; in the former caee we bariily know
more than that ihe miiinttnHnce of the lumbar coid is tudicieui, nn far na
the ceuiral nervous Hvalcni ia concerned, for tbe carrying on of ihe work.
In the cB*e of a d<i^ in which tbe ejiinal curd had been completely divided
in cbe dotml tegi4>n while t ho animal wmk a* yei a mere puppy, "beat'
or mcnatniation look place as iivual.
CHAPTER III.
IMPEEOSATIOX.
Ik onitus the iJiR-linrge of tho eonipn coiilnining ibe spermxtoioa U idmI
probabtr eRocloil by mi-iinH of iIiq perislultic coo (racl ions of ihp vtncala
Mini inn les nn<J ritsn ilcli-ivtitia. UMiat**'! by rlifthmicnl oHitraction! of tlw
bulbu-caverDitiUS tnusck'. the whole being a redes acl. the oenlre of which
apjieiir- to be in the lumbnr apiual cord. In the do; etDMStoo of Mmen can
l>e brought iibout by stiniulntion of the gisas peiiis after oomplet* diviaua
of the sptual cord in the dorMtl rtgioD. fhe emtaetoo uf seuea b pracedcd
by All erection of the peiiig. This we have already seen. p. 274, is in part at
ImuI due to an iacreased vascular supply broujihl about by nieans of lb»
nervi erigeotea : it is iirobAble, hourever, tltat the coiidtiion is further Mcurcil
by a comproasion of the efn-rent veins of the corpora earenioM bj neaw af
amootli muscular tibres prateni in tho^ boiliea. The aemcD bttog KoainJ
into the female or>;aiL.-. which are ai the time in a Male of tui^(«ao«aea n-
•ombling lUo erertioD uf the iwiii*, but \na marked, ibo «^<ermata8M flail
their way into the Fall<>|iian tiitK^uixl hf.rr (protmbly lu il« uppar part)
Come in contact with the ovum. In Lhr c-aiw of niiinc aiiimaU itnprwnaiiia
tuny lake place at the ovary itM^lf. The piu«tiigc of the vjHTRlRloiiua u miHl
probably effected nwinly by ihuir mvn nhralilc nctirity : but in •orae antnali
a retrograde pfiri«tnltic movement IrnvelliDg from the uterus alou^ tbe Fal-
lopian tiibm liiH been iibfcrved ; this might aa&iat in brin;;inj; the semea bi
the ovum, but innfmucb sa these movementa are prvibably pirls of the act
of cniiuB and iniprcguatinn may W deferred till some time after that event,
no Bnai atreer can be Intd upon them.
A3 ttie reeuU of the iictioD of the spermatoioa on the ovum, ibe Inlur,
liutcad of dying as wheu impregnatiuu fuib, awakes to ^reat nutriiin
activity accompanied by remarkublc Diorpholo^oal chaqgea; it enlarges and
develops into an embryo,
[Preccdini; the time of the occurrence of the entrance of tbe apenaatOKNB
into the ej^'g, certain anatomical cbaiigea have l>eeu observed to OOCiir, wJ
in order thoroughly to under^Iatid these, &s well as the chanim* whlti
follow iu tbe ovum, it will first he neccaiary to review the anatomy of tlie
Tlie ovule is a minute cell, tlie wall bciti^ form«d by a Mruolumlew, traiw-
]]ditpiit membrane, cnlted tbe tana nrUiielih, or vileltiiic menibnuti'. ^VItUa
thi* i.i the yolk, or rilvlhiit. wbieli coiui^l* of n graaular wmi-Huid mam,
having dUiipeiideil in it a iiuc/t^iu or ijfrminai w^ida, containiDS a niMMlu
or iirrminiil spat. The genuiuBl vesicle c»R«i«t« of a very di^cate, traas-
jiaivni. homogeneous membrane, which enelons a fliit<l nitb granule^
and MUtiieuded in it an eccentric nucleolus of a granular and librillated
•Iniciurc.
Previous to the occurrence of the impregnation of tbe ovule a very ia-
t'-renting seriua of changoi linvc been obaorriKl to take place. Acoordinif t"
Itfllivtir,' the fini iiiten»tiug polot to be noticed is the migratioD of thr
■wflt^iunarlyJoanalur 3illrnM«op)-,<irtiit>:>t. wc
IMPREGNATION.
IK)1
gwratinsl vesicle towani l)i<? cell nnll. Tbti vcsiciilur wnll (h«n beooiuea
mvj aiiJ K'^""'!)' UUujigtearA, while iil the uuav tiuio iho tiiiclMlua or
gM-ntlDBl ipuL bas UDdargoue iiietaiinirplKiti), <o that whiit ramnins or tbrne
Nruc4ur<* u a spindle aluiped maw. One cKtrvmit)- of this nuM fcrniljally
prnjcoU tlirougb the nil wall nDil ii thmwn nfTiu a potur ivxieU. Frum the
olh«r ranainiiix j>i>rLioii a lecond |>»lar rueiclu is forniocl. th« pnrt of the
mnis tbon remiiiiiiujj; in the uvulf Ix-inj: )ii-rmiiicnt. luirl m calleil the hmtir
pfwiMd«ttf. Tho iw>\i dinugv ol»orv<''l i» iliu ti|>|H?nraiici< ofa xoDe, of radial
Dtrifl) anxiDd itiu iirimiiolriin ami iu ii)i);rati<)n lo the centre nf the Cfi^. The
i>t)ernHitoz<iuD ihcm iteuutrato tho wall iif thii ovule, probably ai the p(uut of
trie formation of ihe polar vtriclnw. Thr tail of the Hperniatozoun beooma
alMortx^l. and the bcai) u motaniorphotec) intu the male piVHUie/etut. Fram
the mnle prunueleiiK a niiniber of nMialine Rtriie arv glvtat off in all dirco-
tion», itnd it tlivD migrates toward iho fflinala proiiuoleus. and aflcrwanl ftuM
with it, forming n single or c/mn-uyf nufiem.
Cloaragc or segmentation of the viielliia then begins ( Fig. 2701, by whidi
prooeos the niicleui thus forniDil divides iatu iwo pur(«, eaeh taking with it
Fla.n«l
I
l^i Bl
intoKUM or ni Vtuou* Biu.a or ciJuvAiit ur nit Y«U. i&Rer DtLna-j
liair uf lh« vit«Uin(! (iiiiMa. Tbeie two divide into fuar, and thene four into
eight, and »> on indi^liaitelv until au ag^Itinierate mass of uuelttaled cells
rewlto, oocb of which euotaiiu a yt^n of the oIeavag« uuoleuH. Tltia maw
of celb is called the mulberry iuom. and the rain
OMutituiint> it arrange theinntlvea aifkut the ioierior
of lh« (U'i'i fttlliiridii and furin the blntlodenaie ttsieie
or m^mhran'r. Thifi inrnihrauc then BpliU up into
two laverri. llie cxteni.il stiil internal ; a third ur middle
layer iieiiij; nllerward forinwi lu'twcvn iheiu.
Iniui«<liiitv1y after the furmiUtnii i)f the two lay«r»
of UMtiMli'rfii. Ml opoqu* roundiul e»llectt>iu uf tuall
oelkoocurn, (.-ailed the nnea ffefmat'itim ur embryonic
Spot. (Kig. '^71. 1 This epol then liouvnim <>ltin|{iilul,
vad in it* luU',>iuidiaaI axb the lint trace of llie
«fnl>ry» apiicant *a a fuint line, termed lh« primiltve
trtter, thu being in the inidsl of a clear oloDgat«d duu>
of oallSi (he orta peUtieidi, which is itMlf surrotinded
bv a more opaauo wae.
' In front of the priuiilivt- iraee two Pilil« aw formed
from which a groove U pr<doDged Iwckirnrd in a )in«
with Ihe primitive traoe. Tlteae folds gnulually rxtoad
along the entire length of (hi gro.ive, anil f jrm the Vimnit 4ar»aU*, which.
by growing, projeet more and mir« abjve the groove, and, gradiially ap-
proaching eocb other, ooaledce and enclcMO thi? neural canal, which will
AAerward contain the oerebro-BpinsI aiii. At abxit th<- same inriod eor-
tai-«ue^~ III )■ . Aim
Cuai)KK<i.ii> •! <■ r<inii«-
n»0«ini UM ■MBCBflM-
■tilra or aniliry«ale«pot,
Uw mn* tmitmOk, uul
IMPnBOItATIOK.
mpondini; to tbe deveIo|)iiient of ibo (lonBl laminw aiaiiliir lamiia
given ofl* IVont the under surface of th« blji«to(l«rni. Thne iiro tlui lam
tUHlraltt, wbk-li. by craduttlly enlarging nn<l finally L-imlcMing, eitclow At
abdominal cavitv. I)ene«lb the Hoor of the groove alMve di-scHbed a tMi-
oale, wliltisb collection of cells app«ara. Tli» is (be rhtrrda dormtit or not*
chord, around which are afterward developed lh« bodies and pruonH* vl
tli« vertebra.
During this perKxl other cbang«8 have also taken pttuv. The oepkalie
and ritudal etiremitiea bav« become flexed and form tlie cephalic and caniial
fifx*iTf4: and the einbrjro hLm being <!urve<) u|)OQ itself laierally. the vil«lliiw
tour appvnrv i>i;paratrd fn>in it by a connlrii'ti^iii. Thbi cijtiiitrirliou gradl^
atlr iDcmioing, finiilly MpanitM the rilelliiiu niat» as a vcnicn'Kr body.U
beme ooimrctcil with tbe l>ody of the cnihryo by the viiellitK^ (iucU (jTvii.
278.) The vninilur btidy thus fonntd w tailed tb^ wmMinU vtMr. Thi>
l)tAV*»M>tnC MCtVDI MIOWIMU TUK UKMYMM DI * MtMHAL tMI MI K Mut
rmimvii AUDiirTtav <*i)»i. mn Tin MuuiMXn i>«' tkr Unti.
TiMMBff* npraxnipd is iliki 4l*(nin vonaii-inl* M Uimi ••■ Uiv nnvttiili ur anvalf^U di| U
Uw hoaia nntiTy<^ |ff«vhiiw lu ilie«i]«irtMiof Uirallaiiuti: r. ilic HUaw ctHnloa : ■>, nu iBoM >
m'.Xim plM» of c«or<(gcncv nt ilie uonlon and ranotlan of ike fain annlra, a'tf.m natum
isaniaw»Uy«r: c,lhi> licailaiul irontof Uwmtrro, raoti^Mnflli* l«lMilUv- '. . iir
•Hul uan: i,t. Uir ili«>|il« •IIibmimt w^l <» »> "PHr *^ lomT paf\'- ■
MdUUmI Tolclo : rl. Ibv vlWillM duel ; ■. tlH Hllknlnla M«n«rtnl lo arnlii k' i-iiti im iLaiiir
Uon til Uw rUsuuiUut e*ii«l.
at first comaiunicaiea with the iiile*tina] cavity, but as <)evvlopiaeDi pioenA
tbe duct of coinmttnimiion Wcotuc* cluscl, and the voicU- is oierrly Utachnl
by a pcdiclt!, and finally diimp]x-jira alttigcther. At the lirue of thn iln«l«^
luent of the bkiudvewcls, vcMcIa appear on itir furl'nce of \' ' ''ia *
vcnicle, coRMtiluling tliu vattmlar areit, lb« cbirf vt^rctA being :
ini-Henieria iirtnrii-» and veiiw. The ve>ve!s of Ibe V8i>ctilar anu ubtuib
nutritive niulerial C( nluincd ailhin the vcmcIg and ctiuvcv it lu (be ttldMyi'
Sot ilN suateQanou.
rMPaBQNATIOX.
993
Sboftlv afW the occurretK;^ of the oommeneeiDeut of tiia formation of (be
umbilical %-«jricle, double f»l(l«, furtneH of the extertiul iHVor uf the. l)ln«tu-
dttrm, are ^iviin off {nun iho cH^plialic and caudal vxtrviuitiot aud Utcrally,
wbioti eurTi! artiuiKl iirer ihc doriinl aiirfiK^o of cbo embryo, where ibcy iDiWt
and coalcwf, and their point of junction buonming nb»or1>ocl, form the
amuiotic vavity. (Fig*. 272. 273. 274. «inl 'il^t.'i The outer Wrr at th«
full), or fabe namiun, graduxltr os)Mind< and curers tlio whole of tbu internal
I
fm. m.
riihiTt.
f(,
X
■■\
f
H «, otwrloii wKL villi. Tlio villi arr <li"Uii u- In Uol ileivlutml (il Il»> |An <<l itw rriiiriaii b> wlilck
W |k« alUnKil* It tileiiiltng : Ihl* purllmi olltmsUIr liMunui Ihe iilnoenta. 'i, utaioc tniiioiii iholm)
UT*n lit llw amnion ; e. uunloUc cavity ; •!, 11101111)11 of Ibc InmOuc, ■bmrloc lla Miiiii(ii:Uan wIlD
tbc nuiUlloil uNlpttN (. uuiUllPal (wd.'Iv ; /, tlluallnii of limn auJ nHwh. 9, albiiUb^ lAflct
Tairn ut4 Boinuii.i
I
•urftce of th« vitelliiu! mwiibrMW, which it uliimiitely replucos ; tbe iuitcr
Ufer, or true amnion, u conlinuou* wilh tin- skin of th« embryo nt the
UlubilicuB, and closely envelops it. The nmniotic cavity or sac ihua Fumicd
becomes filled with the liquor ttninii. which gradimlly iucreaws in utuintUy
M pregnancy advaoooe, up to about tbi- fifth or §ixth ni<mtli, wneu tbo
(jiinntiiy gmiluKUy decreases up to tbe time of Ubar.
FH-ZM.
ruKVM.
rM. I?V— ln*u«AM M fKimuTtt Bn. (AAh lui.nn). •.imlriMMl<rBKh:Kamnliclc«Birlt)ri
«.»lkuiol^
Fm. sn— riKL'Niiiin> Ens witn ALLurrow >Euii.v ivwruiK. a, Inaar larrr uf ■•aiil<iuc 6M :
k, «UM layer u( diuii . r. lutlW olur* Uio KtoiiliiUr (dU> t»iu« lu coaUnl. Tbe allanlDt* \t mtm
IwKUatlngtBimai tbe wMMaiHl Inner InjtBnnribcanuiinilc foldi. TliUII«un>. whloh reintMOU
(Ml) IM uwlMlB MkaBd Uir |aiM ■likiii rbem, iboalil be oani|an.>-l nlib Pint. matHlm. Ib
<iMi4i will tolMnid llu MnKRim viomal in i>»« MUk
At about tbe time orthnc-nmuienonnMtnt of tfaederelopinentof the amnion
N new orgnu, tbv itUaitoin. aiiiivnnt ah h pyriform maw of c«lU at a point iin-
niiNlintely )uM(i>ri«>r to tho vitdline duct and projeolin^ throusb the aaine
oj>«niu||c.' (FiK- -''■') '^■■■< mtwof oi'Ilti nndergow npid growth, 8;>reaidi]ig
■;;(
fiD-i
IMPBRONATION.
ilwir hclvrccu tli(> true iiuil rnls« Kmniiilic foldi, finallj c(im|ile<ely eoclumDjt
tbe ctnbrvo nntl amnion < Fij;. -T6j, bccDniiD)j; M the name tinw uiljoiatd to
the fniM nniDioii, wht-n it U (Icvvlopcd intu the Ime cfmriou. During tW
SroccM nf llic develop mriit <if thi; allniitoiN. il lias Iwromo vcn vwcuIm-; ll
ret there aro two arlcritv and two vcini), altcrirnn) aiif of iw) Tcbw diiik»-
pear*. Thcao vcneU coiirtitlilo the umbiliral vritwl!?. fomiitig part of ihr
umbilicn) cord nbiofa oniDecIs tbo iillnnoiU with ihp nnbr^o. lluring thr
devclopiDpnl of the Allniilniit it prc»rril» thrrtr distinct nnatamical uortMin*: a
portion n hich bccnnirii ron'trictrd nif, nit il wen?, I'roiii (he ntt anu fbrna iW
urioary blitddcr : tbp outer portion forms the cborion, tbe intvrinnliabc |Dir-
tiou fomiiiiR tile iimbilicnl cord.
During the development of tbe enihn,'o up to tbta titnc, tbe lir^ eluifioa
waa formed by villoeitiei formed on tbe ritellioe membrane : au<J fulliiwinf
that by villoeilies developed ii]K>n tbe false amnion. The allantni' then
becoming developed, completely eovere tbe internal eiirface of the &1m
aniDioD, nbieh tben gmdually disupwarB as a dialiaet itructure. Th«(r«r
ehorioti is ihen formed by the allantois, which becomes cuvered by a ^rmlii
of a multitude of vascular shaggy tufts or villi lFi(£. 277), Theac villi, »t
rif:T.l.—t>n\*» Hi HIM (Vii H Df li'.idit Wiia, ililMii Ittna til kUKlli iii»t wUailic IW
tuflr>i (bvmtlhM of Ibvrliuilndi |<«nlr ■ammli niid iHnli mtdawt •Iwoq b^ I lie cnmlhi^ nSt
FlQ. fin— nMma* or hHk op tiiK Kimi. N'iili. ■I>u»I io Innu |«r1 iir ikv |'Un>nta. IU(V) au
nllnA n.SL tUM'11ulkrnri<ilIiv-4&,''.«, ilTlcAfiilrefMlf. •■.•■. lli lH»li<ilnkniit>-ilir tlw<r
tint, are dtdlribulcd over tbe entire mrface of Ibe organ, but lltvy aoon t*a>-
nwnre dJMppeanne, except at a small area eorrebpi-Ddin;; to tbe nlUchiami
of the pedicio nbi^ eonnects (he allantois wilb the embryo. At this |i<»iii
(bey beenniv gn-ally Ini-reaMxl in nuiiitker, and also in size and viueularilr.
Thr»c villi arc cijnipi«cd of a fibrograDular mjilrix. in which are nucaenw
cniiillary loi)pii. niid arc covered with a laver uf vpiiht^lial t-ells ( Fig. 2iS'
Tliis I'ortifiu uf the cburioti forms tbe fcelnf portion of tbe placvain.
tVPREOVATlON.
No MMXh'r, howctCTt IWTa itivm- cliiiQ',;(w Itei^iin iti thf oriim thnn ourrela-
tiv< ob>Dg(». brought kboul iiroWblr by rrHcx action, but at prmeol most
I ofawnro En ihtir cauMtion, mkc plnm hi the iKcnH. Thci raucous inembnuie
of thii organ, whether lh« coittis mulling iii impre)itiati')ii \k coiaciileni
with a m«nslrtial (wrioH or Dot, becomes coug(St«<l, an<t n rapid };ruwth takes
fibtfr, churHctcriM-il bv a rnpii) proliferation of the epilbelial nnd subcpilbe-
inl iHHUcs. Unlike the ciiee of nwnHlriintiou, however, this new (;rowih iloett
not give irar to imnieiliiile dccny and b.viiiorrba^e. bill rcmaias, aiid luav be
di«tiogiii«li«a as n dcw temporary liciiitg lo the uirnie. the »o-called decidua.
Into this decidua iho oviini, oii its dccoent from lti« Fallopian tube, in which
it baa undergone d^velopoienlal thaDgta. extending most probably m far. ill
least, as tlie f'orntaliou of the blastoderm, if not further, id received ; luitl in
tlii» it bet-oaiet) imbedded, the new growth cloeiog in over it. (Fij{s. 27!', 2>*l>.)
H«aiiwbi1e the k61 at tlie uterine Htnictnr^, i^i|iecially the muacular tisMiie,
become alw^ niuib enlar^eil ; od prc-u;niiucy iidvaiicett a lar^ uuml>er of new
mnacular libreu are formed. Aa iLl' »vum i-ouiinuea to inoreaw in alw, it
bulgea iDto the cnrily of the uteruii, carrying nitli it llie (lortioa of Uw
wut. zn.
RI1.3M.
h
»
IM,m— Ptan' trioK or tkk ronaAiiiiN ui ina Uniur* Retilia laurxu nit oii'v.
Fto-Mt-Hiiaa AirtiMati i^m* nr t>nniii:« ItEnau,
<lrct<liia which luu closed over it. Honcrforwanl, accortlinzl]^. a dirtinctiun
is madt' iu the now welldevclopMl dccidnn bdvrcvD ttin rtMiiiw rrttrxn.or
thai part of the uiembraue whii-h covers the projecting ovum, and the
deddua nrrn, or the re*t of the membrane lining thv nivity of the uterus,
the two being cuntinuoua around the bam of the projecting ovum. That jmrt
of tlie deciilua which iiit«rvene« between the ovum and the nearest iitOTin«
wall is frei|u«ntly spiiken of aj< ihe litfidiia wrntina. .\» the nviitn developa
■Dio tlie fu.-iuit Willi its menibraiiea, tbe decidua reHexa Iwcumea pushed
agaiuitt tbe dei-idua vera; about the end of tbe Ibini uMuth. in the ailiuan
subject, tlie two come int«> complete contact all over, and ultimately tlie dis-
tinction bt-twern them U loet. In the rei;iou of tbe decidua serotioa, the
allaiitoie \i-e»<i--lii nf the fortua tievelop a placenta.
In the cariicM ntnjpv of tlie ()evelo]>iueDt of the placenta, the delicaie
tiIImu pruooMoi of tlii> oburion insinuate ihentielvM into tbe hypertropbieil
follkkw of the decidua Mnitina. The villi thtni un(l<^r'„'» a rapid incretue iu
•lie and va»rtilanty, Ix^miiug branched into Mtcnnilary and lertiarv rnmifica-
tiooa ; while, at th<- tame time, corroipuiiding ehnnfct*" ore taking place in the
follick*, by which tlicy Iwcomc greatly iucrcA-wi] iu »\ix ami vasctiluHly, and
al the HUiie time forming diverticula in which an' irnilH-ildei) the mniiRcaiion^
of the villi. The villi and fullicl*-* thui grow liiiiuliaueouily, and finally
I>e0(>ine blended with each uther and an- im hmgrr frparate sinictnrM. Tbe
fcliii-ular bloodvcewla lirM form cagiillary plexuM^ ; the!« veawla, howevar.
996
IHPRBONATIOK.
Iwcouie ea\»i^ed, fortuiDz frei^ucnt nniwhiiiiixteti. nod Knally conlwHni to
form ivnotu Miiitm ( Pig. iHl ), m nhicli nrc Imthml the ftvul villi. There U
Vl'.. »\
AscTMM lit A MMiDxar * FrLi.r Kohhli- ri.i< ir»T*. Hiik iiii- i«fi«( tlw atcnw (a ■h»«ab
lIienin)MalT«B»l>: .1. li. rutllnc iir1crl» i>r lb* uwrii*.
nocoutiiiuity e«liil)li»lied heiwe«a tb« mnliiriial and fcuu] blood; tbe laWf
rhangQ of nutritive inat«riul iteoeftutrv fur thi* er'iwtli and developmeiit l(
ibv &!twf tnkiT pliicc llie dcilinilu nnfu »r tlic villi.]
For further account of the vnrioii* clmtiee* bv which thme efenu tn
hraaght nl>oii[. if well ii9 of the hinlorv of Ihc eriibrvo ilwlf, we rutut refer
the reader to ntiaiomica] iKHtiMW.
DvKi.vo tbc ilcvttlopni^nl of the chick wilhin the hcnV egg lb^^ miiritiv«
tnaicriul needed fur tlii^ grovrlh fini of the bltutotkriD, nod njb«e'|u«-ntl]r of
the ecnbryo, is supplied bv the ridk. while the oxjnu of thi- iiir piimiog
ttw\y through the |ior<iiie shell, gnios aecrm to all toe liwiKv tKiiti of the
cmbryu uid yolk, either dircclly or by the iotervenlion of Ihv uUnDioic vo-
s^s. The nnuaiDaiiiut embryo, during the period which pivm)<.« the •xton*
sion of the atlnntoie vewcis into the cxviliea of the uterine wnlls tA fnnn tho
pltic«iiM, ni<|]>t be nourished by direct diffusion, linl from the cont«iiI> of thft
rRllopiaa tube and subsequently from the deuidun ; and its supply of oxygvD
iDint oomc from Ihe same sdurcM^. All aiinlo^ would leftd us to suppose
thai, from the very first, uxidtilinn is gatog on in the blasloderniic an<l cm-
bryooic structures ; but the urnount af oxygen aotUftlly wilhdmviii from
without is probably BXc«e«liDf{ly small in the early 8laKc:°, seeing that nearly
ibe wlHfle onerey of the metalwlism going on is directed to the buililing up
of structures, the cxjieuditure of euerg}' m the fiirni i)f either heat or eitet^
ual work btiuK exireiuely small. The markei) iiicn-'atte of bulk wliich lakets
pjftoe durini; the (.■ouvcmioii of the luulbcrry maw into thL' hliuttudermic
vwkle, ahovis that at this epuch a n-laiivtly speaking lai^ quaulity of vraier
at leoAt. and probably of uiilritive mativr. niu.it pan flrom without into the
ovum : and nutiM'iiu cutty, though the bluslnilvrm and embiyn may for ihime
time draw ihr ninterial for tlieir continued ponstnietion at lint hand fruni
the yolk-Mc or umhilicml vmtcic, both this and ihcy ctmtinur prohahty until
the idlantoiit is formcil to rr wive frwb malarial frwm the niothrr by dirvct
tlittusion.
As (he thin-vrallerf nllnjitoie vesm-ls come into doer and fuller connNlioo
with the maternal uterine siuiises, until at Inst in tho fully formed plarenta
the former arv freely bathed in the blood streaming through ihi; latltr, th«
nutrition of tho eiubryo becomes more and more cnnlinctf to this tpKial
channel. The blood of the fattus Hovring along the umbilical artMioa aUbol*
exebanges with tlie venous blood of the mother, and leaves tho placenta by
the umbilieal vein richer in oxygen and uuiritivc material lu^l jioorer in
oarboDic acid and excretory products than when it iasued fr»m the fo4us.
As far as the gain of oxy^o and the loss of carbonic acid are concenml
tbeae are the results of simple diffusion. Venous blooil, as we have slniidy
■etn, always contains a (luantily of oxy-hiemoglobin, and the i(uantity of this
sub«lance prcMni in the blood of the uterine veinit is sufficjeot to supply all
tlie oxygeu that the embryo ueetls ; the blood of the fetua, containuig teas
oxygen than even the veuoua blond of the niutfaer, will take up a cerinin
ttioui;h small quautity. The fietal blood iFavelltnf; in tho uubiliotl artery
must, in proporlibu u> the extent of the nutritive chanoea goiBg on in the
cuhryo, powea* a higher carbonic ten*iou than tlial in the nmbilical rain or
uterine nnuit; and br dilTunon gel* rid uf ihi« lurplua during its stay in th«
pliuvnta. Tlte blowi in the umbilical arteries and veins i«, therefore, r«lB-
tivelv tjwakingi venous and arterial rr!i|Hrctively. though the simill exrun of
uxj-AMlogloblD in Uie bUxKl of the umbilical voin is luiufildeiil to give it it
dhtlnctly aiurial oQlor, or to diftinguish it a* sharply from tlie more wnoua
:
MS
THX KITTRITIOX OP TBK KHUItTU.
tilood of the iimbilirnl artery, u u nrdimn' arterial from ordinair .
blood. TbuB, the ra>lu» breathes by tneans of the maternal hlikod. in
aaine war that a fieli breathe* t>r menR<> iif the water in which U dwells.
The blood of the fietus w very ])oor in biemo|{lobiii cormpondioe lo iti
low oxjffiu coaftumplioii. Wlieii the mother 'm a>iihyxiiiled. the Iietiu ii
■Hphyxiated too. the oxygen of the latter paffiiag Iraek u;,iiin into the blood
of the former; aiid the a«nhysia thus produoetl in ilie fotue ii taiich moi«
rapid thau that which results when the oxy^.'en U u^'<d up by tht^ tianciar
the fwtua alotie, ok when the iiuibilicns is Itjtulured and the fa-tii;^ ni>i allowad
in breathe.
If oxytfcti a»d carbonic acid thus pa^ by difliuiou to and from the raiAhir
nnd the ht-tuf, one uii^'ht fnirlv expect that ilitTuaihk- »ilu. jmiti^ida. and cv-
bobydratea wuuld be conveyeil tu the latter, aiid dilfuniblr rxcn^^ton* tarrinl
away to the formur, in the uime way : and if fatM ■■an pniv ilin>c<ly tiif* ibr
portal blood durinK ordinary dif^tion, tjicn; can Im: no mwoii fiir doiihtii^
that this chiM of fdOil-stiiflii alNo would fin<l it* way to the fii'tus ttimugb llir
placrnlal etmclurti*. We du Icuow fnmi cxperintcnl that diHiifililo tulMlaam
will jHMi both from the mother to the frUiK. niiil from the Iwtits tu thr
mother ; but we have no delinilc. kaowleilgv w to the exact fumi anil raaoDir
ill which, during nnrmal intru-nU'rine lif<;, nutritive miiteriaU an- convwyH
to or cxcrelioiw loiivrvfd fnini the grnwing young. The placenta is remark-
able for the great development of cclliitar slnicLiirw, apjiArenily of an c|n-
thelial DaUire,oii the border-Intid hoiween the maternal and fa-tal eleiunlf;
and it baa been aug(n:»t.cd that cbcac form n temporary' digestive and secrtfocy
(excretory) organ. But we have no exact knowledge of what actually doei
lake place in theee atructuree. From the cotyledons of niiuiiiaiiid iiur ht
obtained a while creamy-looking fluid, which from maov ftaiurva o^ lli
obeniicid compcaition might be almiMt npuken of at a " utcnne milk."
Bnmking broadly, the fcetus liree on the blood of it< fflother, very mucb
in the »awo way an nil the tJttoue of any animal live oo the blood of th(
Imdr of which ihey are ll)« |mrt«.
For a long lime all tlie euiDryouie tiMuet arp " protoplasnio " io cbanelir:
that u. the gradually difTtrbtiliating vlemeiiU of the Mveml tiaauea reiaaia
still embedded, ao to speak, in undif&rvniiated |irotoi>la«m ; and during tlut
period ibcre mu«t be a gvnerid nniilarity in the mctabolUm going oti id rari-
out part« of the body. As iIiir<Tcniiation bocom« more and more tnarkad,
it obviously would be an economical advantage tor partially elaborated mai^
rial to be stored up in various tintal li»u«», so as to he rendv for immecUHi
lue when a demand arose for it, rather than for a apacinl call to be made U
cadi occasion upon the mother for compamtively raw material iwediog nk-
sequent preparatory changes. .Avci^nlingly. we And the tinaee (^ the (mat
at a very eurly ]ieriod lundi^l with glycogrn. The miiKlee are eapecially tiA
iu tbiB :9tiii9tanoe, but it ocL'ur^ in other tissues as well. The abundance of 11
in the fornter may be explained partly by the fact thai they form a vm
iArge proportioa of tlic total nin^ of the festal body, and partly by the ba
that, white during the praaence of the glycogen ihoy contain mucli undirter-
eutiat«d pnxoplasui. tney are exactly the organs which will ultimalelj
tindergoa largeamount of dilferontiation. aDd.lberefire, need a largvamuott
of material for the motaholUm which the diirerenltation entails. It i« nnl
iiutil the later Mages of intra-iilennt' life, at about the lifth moiilh, whniil
i» larmly diwippearing from the musclot, that the glyi^^'i.'en itogiii* to tw
depnaitea in the liver. Ity thid time bUloloncal diflhrentiatioti hax advaiUMl
lar^ly, and tin- u*e of the glycogen to ine economy luis 1iu<h>iii[> that to
whicfa it ix put in the ordiuar;' life of the animal ; hvniv, ne liud it ilejuMilvl
in the iMial place. BeKtdej being pr«ai-nl in tlif fuMal. glycugen is found ^
TUK NUTRITION OK TUK KUMBYO.
»mi
I
I
in ihij |>Iiicvntnl stniuiiit'cs: but here pnihiiblr it k of qbc, not fur th« fetii>>,
b(il for tlio nutrUioi] und f^rou'th of ibc pliicoiilnl HtniL'turos lhftiii»elv«t. W«
do not knoH' Uow much t'arbohydratp uiiiterial (iod? its way inin th« umbilical
Teiii : Slid we cannot, thcrclbre, »tate wlial is the source of the lislul f;ly<.'ug«n ;
but it it ul Unat pi)Mibl«, not to sav pr>)l>nble, that it arises, in part ai all
eveois, from a splitting tip of pmeifl mitterial.
L'oDccrfiing th« riw and development of the fuoclioDitl activities of th«
eiubryu, our iiuuwledi^ in abuost a blank. We know ecarcelr aavthtug
about the variouB steps by which the (trimary fuudaniuuial <|ualttie« of the
[>nit(>|>|jisiii of the ovum ure dilfureutiuted into the complex phenomena which
w« have iittempte*) in lhi» book to expound. We can hardly slate more
than that uhile tuuscular contractility buconiea early d«velupecl, and die
heart probably, as in the chiuk. heaU even before the bhiod-corpuselea ar«
formed, loovementa of the fo.-tuA do not, iu the human subject, become pro-
nounoed until uflvr the tiflh muiith ; I'roiu that ttuie forward tber increase
anil NulMuqueiitly become very marked. They are <if^n s[>okeu of a« reflex
in i-Uaractur; but only a preoonoelrcd btiu would [irevunt ihein fntm being
rrgiinim) aa lar^ly niitomado. The digentive fiinctioDii are uatumlly, in
tbe nlxH-ncc uf all fotnl from the alimeutnry cudbI, in al)Gyau<'(\ Though
nvpHin may be fiitnid in the giiwtric membmnc nl about the fixirlh month, il
IS doubtful whether n truly |N;]itii' giuHric juice i-' nccrtttwl during intra-
nterine life ; Iryp'in nppeiir* in the |uiiicreiu somewhat later, but an umyl-
olylic fcrnwnt cnnnot fw obiaim-d fn>m that urgiui till nftor birth. The dato.
hi.weTcr, at which thi-w »i'vcnil fdrmcnts make their itpixmriuico in the em-
bryo appcan to iliUbr in dilfcrcnt animnU. The excretory functions of the
liver nrv developed early, and about the third month bile pi;;ment and bile
salts find their way into the int«8line. The ijuantity of bile wcreled durini;
inira-uterinc life, accumulates in the inlostine nnd especially in the rectum,
formiutc. tojrether with the itnmller secretion of the rest of the canal, aixl
•ome d«^uamated epitlielium. ilie so-called me<^«nitiai. Bile aalla, both un-
altered and varioiitily chaD;.-e<l, the usual bUe pigments, and cholestorin, are
all pnseni in the meeoiiium. The distinct formation of btle ti au indtcatioa
that (he products of fmial metaboliam are no luUij^ wholly carried off by
l)i<^ matt-mal circulation; a»d tu tlie excretory function uf the Uvur thcrv
are now luliled tbow: uf the akin and kidney. The substaaom etcopUtg by
lh49>* orgnns liu<l their way into the allantuhi or into tlie amnion, aconrding
tu the arninp-mi'jit of the fecial membrnnai in diflereul cloMe* of un!tnal# ;
in both ibi-Bi- tluidi urea or allied bmlie* have been found lui wdl as the
onlinary nlinc i.-oniilitne[iL->: the taller may ur may n'lt have been aettially
eecreten. From thx allantoic fluid of ruminants the body allantoin ha»
t>een obtainod, and human and other nniniolic Huiilx have bren found to oon-
tnin urea. It is maintained by some, honever, ihnt the tluid in the ■mnion
is wcrebn) by the mother nnd that benc« the eiibetanocs prwcnl in it am of
maternal origin.
About the middle of intra-ulcrine life, when the ftelal clreulatiun [Kig.
*i92) i* in AilI developRienit, the blood flowing along the umbilical vein is
carried chiefly by the ductus veuneus into the inferior vena cava and an into
the right auricle. Thenoe it i» directed by the valve of Eustachius through
the foramen ovale into the left auricle. pasMng from which into the led ren-
triele it is driven into the aorta, ['art of the nmbilinki Mood, honever,
instead of pa^itiK dinx-tly to the inferior cava, enteral by tli« portal vein into
tbe beiMtic eirculatinn. from which il returns lo the inferior cava by the
bepiUic veins. Tlie inferior eava aUu contains blood coming from the lower
limhe and lower trunk. Hence tbe blond which tnMing from the right
aurtclc into the Idl auricle through the fotviumi ovale i* utMribuieil by the
1000
THE SUTBITIOS OF THE EVDBTO.
]vi\ veiilTicle through the aortic nrch, tltciiieh chicllj blood comiDg ilimt
from tlie plncentn, is itl»o blood wliich on lit way Ironi tbc placenta ki*
paaeed ihrniigh Iho liver otid blood drmctl fnim the timin of tb* lomr
part ol* the body of tho ftotun. Tbo blood dncondin); «!• fivtnl vODun btood
tnm ibe head and limbs bv Uh- imponor vcnn mfn dovs iK>t n>iii);I>' «itb thai
of th« inferior vena cava. !}iit fnlU tntu the right ventricle, rruni which it ■
diseliarwd ihr><ii);h ihc ducliiF nrlcrimiis < ItoialU) into ll»e norm. iN-tit* ih*
arch, whence it tlows tiartly to ihc lower trunk and liinl>». but cbiclly hi Uir
umbilical arleri«« to the plarcntn, A smntl qiiaoiity only of ibr cijutrni* li
th« nKhl ventricle tinds ile nay ititu the lungt^ Now ihi- bloud wbi^i c«am
from the [daceoca by lfa« nnibilical vein direct into (lie ri(;hl auricle ia. a* ttr
U the fdelua ia concerned, arterial blood ; aod the poniouuf umbilical hloei
fHa.Vt
:v
\f \f
I, (be unbUlnii mod. mtKtuiiiu ■>! ili
win kwl iwu umlillml *ii' i 1l«v l^'
lli« iilMiniU ci^ 1. >■>« ii'ii' ' iIkMhi
Inta dim bnuclin, t«n il. 4< !'• (k •iiiintninl tt
IliE llTvr, inrl oil* il), ibe duriii* vT-umnk. urMik
ralm the latrrka *«a* mn if>); *. Um |m«I
vctu. nluinlitji tlu) blood A«M UM lnlnUDovMl
iii>lIliii(wlitiUi«rlHhIhc|*<>'' '"-"'' >. thirty
/j iinntlw . ibpwntix' nf Itr t>l -nlblfli
•■ ftrrov proccvfUDf fivm ^ lu " ..< rttH H
lb* loft Ti'nlrlrlc; Uid htnol Mlmmu lb* «nn
Id (III- arvli nl Ibc aona ill), to Xm dUUWM
inniHKli Ihc bnodie* (ittMoiT bT (tic ana ti Oa
IiMil and itpiiMRilmDitiM; Ibf arrow* Uuif II
a'pr*»-iii Djc- mnm ofibc feJvod ftnai ite >«<
anij iiiiiHi riiinulUt* Ibroucb \br ]ac«lu •*<
wibelstmii teln" In lb- ■'it^rt-^' •-"• •■« iUt,ti>
ilw rlul'i niirlcli! i<i. HI MlManiM
IlimtUlll Illli n||hl mill: ' I l< II" lAllUMMfl
■rttlT illil; ]•. Ibc -IiiFlii* ■rltrtii-u*. vWtb If
l«an liitsa |icn|4:t ttciUiiuiiIlon -^ ■)>• (•■InHatn
unrrj; (In- iifmn al ™cb 'i-l"' •'■ ••'
loH iiiIUHirnrj- •n«l'.'« rui oir 1< t
knnit Jii|[» It"' ilis«adlD« aonii .-. i- -wi^
•llililva l«t" <■"' ri'iiiHua ItiiuK. (»■] Ibas ihloik
liitcnial UUa. w hkb bnyoDr <t» i.i.iM^i.uUrua*
(lU), aii-1 Ti'lani Ihc liliio] ■> i •«)
lo Ibo I'lutnla. Hiid Ibvpii- i-*
■tv noitlnafl tobi ibv bvHK •'ilivioliuai tto
■rmnit n< ibi> trfaliialVin ••! Uiibc i«ii«Ii ■■■
llic niluniot iba mvoot UcBd by itwtWfc^
li'It'nnt i-rnacnri.j
which irareraea the liver probiihly hi^es "t tbit epoch very littbr i.x;
during il» iraiiail throuith that gland, the liver Iwing at tht» |H-n.id « *"«
excretory rather than iiii actively melnbolic orjjan. Hcnei- the bhwd of i«
inferior vioa cavil. ihwiKh mixed, la od the whole arterial blood; ai»l il a
thi« blood which i« wni by the left Tontricle through the arch of the nUia
into the canrtid and tuboluvian arteriea. Thuit the bead of the fastne 'u \mi-
vidofi with blocMl cominaratiTelT rich in oxygen. The blood drtt-eiiiiiui* fiMl
Ihc head and upper limli* by 'tbo superior vena cava ia dieiincUy v<Im«:
and thin |>nf>ing fmm tbi^ right v<niride by the duclixs aneriiiauii b ilrimi
along ili« dwccnding aorta, and together with BOme of the blood pnaKUj;
THE A'L'TRITIDK OP TRB KHBRYO.
1001
I U>e left vcniriek touhi) the sortie arch falls inlo tbe tinibilicnl arleriM
aad 80 reaelic« ihe pUceDtn. The fielal circulaiinn lh*D i." ta amiiij,i;<l. iliac
while Ihe most (ti^tinetl^ veDous blood is drivi-n hv ihe rifjht venlrkle back
to the jtlacenlu lo be uxyceoaleil, the luiuit (lixiint-lly UTUriiil (hut still niixeil)
blood i^ driven hy ibe left veotnde to llii? wrehrol wlructurft*. wliieh have
iDore Dt«d of osyKi-ii thnii have the other tiwiicM. Cotitran- to what lakw
ptaee afterward, tJie uurk of the right venirlt'le i« iii the fd'tu!) ^Tcatcr than
thai "f Ihe left; and, iiCd'Hin^lv, ihat greater thiolcnrin nf tW left ven-
tricular wall». to ch anicteric tic of the adult, dom not Ih-cviuci iimrlced until
^clo•e uiwu t>iith.
In the later Hufcm i>f pregnane}' the mixture i>f the various kiocU of blood
in the rieht aiiriole iiifrvaiu.it jm-ptiralory to the changea taking place at
birth. But during thi! wholi^ timrof intni-iitcrine life tjiaainouulofcxygeti
in the Moid [ituoln^ fmni tlu- aortic tirrh to the medulla oblongata ia outfi-
cicut to prvveni any inapiralnrr itiipiilM-a being orifiuatm) in the medullary
rtv^iiralory cx-Hlre. Thii^ during the whole pcricxl elapsing betwwn the date
uf iIk olrii'-liiral mtahlinhint-nl. or rtUhrr the consequent full dGvelopmoiit of
tilii irritability. un<l the v.poeh of birtb, remains donnnnt ; the oxygvn Rupply
tf> tb«' pnilopltL-ni of it« iierTe-cclli is never brought so low lu* to ml going
the mpiralory molecular explosions. As soon, however, a^ the iuiercourM
b«t«een tW maleninl and umbilical blood ia interrupted by separation of
the plueenia or by ligatiirf of the umbilical cord, or when, as by the death
of the mother, the umbilical blood censes to be replenished with oxygen by
the malemal blood, or when in any other way blood of sufBcientlv arterial
aualitj txnm* to find its way by the left veutricle to the medulla oblongata.
IM mpply of oxygen in the respiratory centre sinks, and when the fall has
reachra a certain point an impulse of inspinitiuH ia )tenerated and the ftetus
for the first time breathes. This action of llie Kitjiirati^iry ci-iilre may be
•MlMed bjr adjuvant impulws reaching thv centrx> alone varioim nflireut
ncrvea, auch ua thoae started by exposure of the body to ihi; air, or to ewld :
but th<4« are oubordinnte. not i^vn'nlint. A retarded fine brentb may be
hurried on by dasliing water on the face of the newborn infant ; but on the
ollwr hand, the fa-tus, upon tbs ccmition of the niacental circulation, will
make it» lint rtmiratory moremcnts nbilc it is still invesitil with the intact
membrane* ao<l thu« sheltered from the air and intleed from all external
stunuli.
Before this first bmilh is taken the pulmotinry alveoli contain no air, and
the luo^ when thrown into nnler sink at once; they arc then said to be
" atalecUUic" After the tint breath, the alveidi i-outain air and the lunga
float when thrown into water. A striking dillereiice. however, exists hetweau
the lungs uf a nenbom infant and ihoee of an older petrvon. When tlie
Sleural cavity of the former is opened. Ihe lungs do not collapse, no air is
riven out by the trachea; that partial dietendon of ibe luugs, and negative
tboraciepreaaure, apfiears not to be established immediately upon Urth. That
porlioD of the reuiiual air in the lungs of the adult, which remaining after
the mi»t forcible expiration, is Mill driven from Ihe lung* u[>uu the pleural
cavity bdiig laid open, and which might be called "colla|Be air." is wanting
in the newlturn infant. When the change from uue L'onditii>n to the other
is eili-eted in not at present known : it may puuibly ariw from the growth of
the cheat outMtripoing lhat of the lungv.
When the lirsl hn-alh U taken, aa iiuder normal circumidaucn it is, wilii
free arcew to the atmcxpbere, the \ttng* become lilk-d with air, the seantT
supplv of MoihI which at tlie moment was |iM»iiig from ll>e right ventricle
along the pulmonary artery returns to the left auricle brighter and richer in
oxygen than ever was thv ftrlal blood Iwfore. With the diminniion of re-
k
1002
THB HCTBITIOK OF TBK KMBRTO.
MMUice in (he pulmiiiiarT cirviilalinii Rniucil 1>y lh« cxpdiiiiiion of tbt ibofit,
u lai]ger ttipply "f h1ui)([ p«Mcw inin lliv pulraonBry Brtdrr in»loiMl nf iaia
tl>o micUiis arU-riwup, anil this dorivnliuii of tli« (.imtcntii uf Um> ri^u vrn-
tricl« iacrewiiug wilh lh« coiitiiincd mpiraLcir}- mnvomcnU, Uie ourrrni
Ihroueh the Inttcr rniin] Ht Iwi cckms ulto^-tlicr, nnd ii» cliannal ibortlj
mfmr birili bcconics nhlitarnloij- Currcaponding tn th? greUor flow iuUi tlir
pulmonnri- nriery. » lar^tir mikI Urgar <|uaiilily of blixid returns fmin ibr
piilnionnrir- vciii't iiilo l)ie le)1 iiunclc. At the Mine limp Uie current thnMifh
the diiclits vciiomis from the uinhilicnl win hsvio); cnued. the flow frqcu ih
inferior earn hue HiiiiiiiUbeiJ : and tlif bloml of the richl aurieic findini
litllc re^istitnoo in l)i(> ilirectioii of the ventricle, whieli now ivodily di^
fhargra tie cdiiIodU inli] ihe ptilniouary artery, but fiixlin^- in lh« lefl auride,
which b L'ouliniially beii)); tilled from the lungs, an nWacle to iu MjMgc
through the foramen ovale, wasea to take that course. Anv r«turo of blood
from the now vigorous and active left auricle ialo the riglii auricle ta pi»-
vented hy the valve which, during the latter stages of iDtru-ulerine life, haa
been growing up in the left auricle over the foranieD ovale. At binh thi
edge of this valve is to a oertain extent free so that, Ju csae of au emersieDCT,
as when the pulmonary circulation U olutructed, a direct wcape nf hhoi
into llie left auricle from the over-burdenei) ri)(ht auricle oan take pbm.
Kveutually, iu the (.-oiinv of the 6r>t year, adhraiou talc<« plaoo. and tbr
Mjiaration »f the two uuricl<» become* cumpluh-. \^'itli it* htrmr supplr «f
blood and jtreater work tlie left vontride acijuiret the greater liiiclcDon ebv-
a«leri«ic of it during life. Thin tli<- fn-tal circulatiou, in coDMqiwaotoril»
rM*{>initory miivvmuiitv ti> wliich il* interruption giv«i riw, cbuigtg its ~
into that diiiractemtic i>f the adult.
(
CHAPTER V.
I
PARTURITION.
■pile of ihe incrensifl^; dialentioD of Jtfl cavity, iHe ulenu ramniilB
quinoeni, tut far an nuy lunrkeil tnuHculiir tuiiiraL'iionii ure iMiXM-meit, uuiil
K <«rlttin timt- liits been run. lo tlie humati aai>ject the period of Ke^taltiiu
gencrullr ImW from 'Si^ w '280 days. i'. c, abmit 40 weeks, Uie h'^ueral
cuiOom (leiiig to expect parturition at about 'i-SO davH fr»m the latti meiisitru-
aiioii. Seeing ihmt id nuiii}- vium it in unci'rtaiii wlieiber lh<- ovum wliich
develop* into the embryo [^{t Ihu ovary at llie niL'tMlniation prin'tnliiig or
•uoeeeaiog ooltun, or, a* itcime have iiruiuJ, indopciidunt of mcu^tntatiou, by
nuon of tlic roiluti ilaetf, un exact dcteriniiiaiioii of the diinilinn of preg-
nancy » impomiblc.
In ifaeeoir tlio period of KMtatUin la ntiout to 280 days, in the mari' aboiti IVAt,
•beep about IM dny*. dog almut >\''> dnyn. rabbit about no dayi.
The extruBioit of the fielus U brotixht about, partly by rhychmicnl can-
intctiona of the ut«ru» iUelf and partly by a preiuurc exerted bv the coti-
irootion of the abdomiiul mitmlea, *imilar M that deirribtii] in dcfecatina.
The conlrartionH of tlie uteru« are the Ant to apjiear. and their fint oRcct \»
lo bring about a ilitalinn nftlieot uteri ; it ix not till iho latttrRtagv* oriabi>r,
while the fd-tus i* piu^ing into the vagina, that the nbdoninal mnsclM are
brought into play.
The whole (iroceee of parluritimi may be broadly considered a reflex ayt,
the nervous eeniri> beiu^ pliiccd in the lumbar cord. In a do;;, whoae dorul
cord bad be«n coiiiplelely severed, {Htrl urilioD took place as usual ; and (be
Iket that, in the human siibjei^t, labor will progreoa quite naturally while the
patient is unconscious from itie ad ruin Ihi ration of cbloraform, shavs that in
woman also the whole matter is an involuntary aoliun, however much it may
be aoaiued by direct volltioDal ellorts. That the uterus is capable of being
thrown into contractions thniuuh retlex avtion. excited by stimuli applied to
various aflereut iierviH. in well-known. The oonlractioo uf the uieriu, which
is so necf^tsary for the prevention of lieiuorriiage after deliverv, may fnt-
nni^tlr be bmught about by exerting prossttre or by dashing cofil water on
tm abdomen, bv the introduoliuiu of foreign bodies into the vagina, and es-
pecialiy by putting the child to Ihe nipple. And we learn from oxporimcnta
on animal« that rhythmio iviniractioos of the itterui, n^Mnibling at Ictut thnoe
of parturition, may I>a brought ahiKit in a n;ll<;x ntnnnrr by ftiniulating
various affereni ocrviw. Similar movemenla raay be induced by direct
stimulation of the »pinal coni along its whole length, as well as of varions
parts of the bniin; but thrrc arc rcowini for thinking that in these cawa the
impulses starl4;d in the brain and upper part of the spinal <.-ord praduoe their
flAola by working upon what may br called a " parluriliou " (^..-nlre in the
Upper bimbar Fegioas of the cord. And it would ajipaar that (he uterine
oontractionn which are indtic«l hy such drugs as ergot, as well as thoae caused
by aapbyxia, are, at all events in |Mirl, brought about by (he agency of Ihe
tame lumbar centre. From thi< centre (he path* for (he cStren( Impulses
appear (in the dog) to Ik twofold ; one along syinpithetic iracu, by nerve*
passing from tite inferior mpsenteric ganglion to tlte hypogastric plexus, and
1004
PABTUItlTlON.
tho oihcr iilou^ t<|iiTia1 tmctH liy Iirnnclieii of tUc tncral aervtB ti> tlic ninr
iil«xiJK. Il in Kiiiletl tliul tlic cliitrucien of tbo luavemeuls induwti by slimu-
luting iIicm: two traiTbi lire itniucwluit ilifTurcui.aiiii moreover itial tlie synipa-
Ui«LiL- truc-t in vnnii-fiinstriclor unit (lit; <[iiittil tract vowHlilator iu DMuk;
bill the iimtli^r huA ddI yi-t \n\-n fiillr worked oiit.
Wu nre, however, hiki-illy jii«liHr<l in CDDxideriii^ tbe rliylltmival codUbc-
tions of l.hv ut<;ru» during jiarturilion niiNn){ilc nrlli-x acU excited by tbepft^
cncc of Uk- fwluK. Wr lire iillcrly in llic durk a» lo why tho uterus, after n-
mBiiiine npiinrciitly perfectly ()iiit.-sc%ni (or witli cnntniclion* m< iltght m tn b»
with dinic-iiily n]>|>nTiiil*Hl) for months, i" »u<l<l*uly thrown into ui-ti'iii, akJ
nilhin it miiy In- for n few houm or even Iww g«U rid of tbi- biintni il bat
borne with kucIi ((ilirnnce for to li.mg » time ; iiou*^ of tlie vuriuu.i hy|x>tli^^M
wliicli hnve bcTti put forwiirtl caii he eun«idci'ed id> !aiti»lii<.-t(>ry. Anil uq^H
we know what 8tnrii> ihv iietive phiiee, ne shall reinuin ii> ignoraiKV of the '
exact iniian«riii wliicrh llie netiviiy is broiight aboot. The pevtiliHr rbyibmic
character of the c>'llt^leti<>n^ eaeh " piiiu " beginning fn'bly. rifling to m BUx-
imuni, then dcetiniug. iind lioiilly dying nwny altogether, to bo mioomuM
after n pauee by a similar pain just liko il«elf, pain following pain like tiit
tardy long-drawn beats of a alowl y beating heart, BUgg«tts that the chun *f
the rhythmie contraction ia leated, like that of the rhythmic beat of Ik
bearl. iii tlie organ it«clf. And this view is 8np|X)rted by the lact that e«-
tnctions of the uterus similar to thi»e of parturition hare been observed ia
aoimalfi even al\er cuinplete daitructton ot the »pinal cor«l : and tlie Btott-
nieut^ induced by aitpliyxia teem iu part, and those cao^ by sune dn^
auch aA ammonia, seem lo be wliully due to nn inlrinBic action of tbe iit«n»
il£elf. Neveiheleaa, general evidence aup|K>rta (be conduniun that, is >
normal slate of things at all erenta, the coniruetiuiia nf the nt^nia. liketluH
of tbu lymph- hear Is, are largely dc|>i;ndc<nt on the aplnnl eord.
Thir occurrenoe of controctionN in eoRM'tiiienee of an aAphyxiatnd oDodilMi
uf the btiiud explaisa why, when pregnant luiimaU are tuphvxiaUK]. an «!■
trusion of tbe fcetu* frequently tnki? pinec. There u no evjjenve, bnwim.
that the oiiact of labor it, aiuaod by n gradual diminution uf oxygeu in lb*
blood, n'nching at List to n climax. Nor nre there HiHieii^ii facta to oooiwi
jwrturition with any condition uf the ovary resembling that uf lueiutraatH*.
Tho action of the ahdominnl miiseleif in parturition i«, on the otbtr band,
obviously a rellcx net carried out by means of the spinal cord, tlie uvvamn
stimulns being supplied by the pressure of the fcetus in the vagina or by tW
contraction of the uterus. Uence the whol« act of parturition nuiy with
reason l)c eonxidered as a rcHex one.
Whether it Ira wholly a r«flex or partly an automatic one, (he net oa
readily be inhibited by tbe action of the eenlral uervmis systeni. Tbui
emotions ar« a ver^ freoue&t cause of tbe progress of pariuiitiou beinx ni-
denly stopped ; as is welt knairo, the entrance inln the bedroom of a MnafH
often cau!«8 for a time the sudden and alwolute ce^^atiuu of '* labor" paiaa,
which previouftlv may have lieea even violent. .Iud>;int; from tbe aittlosy
of mieiuriiiuu, Wwe«n which and purturition there are many |toiiK* d
reaemblsnee, we may 8upiiiK>e that this iuhibtlion of uterinv (-ouiractioa* I*
brouffht about by an inhibition of the centre in the lumbar cord.
Aiier tbe expuUion uf the lii^lni', the f<i-l»l plair^mta acparatea from ikt
uterine walln, and \», together with the rcmnunta of the luembruiu*. cs)«Ual
after it. Tbe uteru* then fall* into a lirm tonic oonliaclion ainiilar to tte
of tbe emptied bladtler, by which inirans hKinorrhase from the ^wwcU Inra
by the Mpataiion of tbe placvutii in avoided. The Fining incmbnuic of tha
uterua it gradually retitored, [liv mn»eular elements arv reduced by a n|ai
Iktty d^eoeratioa, and in a short time tbe whole organ has rclunied to iHt
aormai eonditton.
J
CHAPTEK VI.
THE PHASES OF LIFE.
I'he child bas at birth, ou nn nvoraffp, rmtber leM than one-thinl tbc dux-
iroum length, and about one-tweiiliclh (bo maximum weight, (o which in
future Tears it nill nttain.
Tb« cttin|Miiitioii of the body of the newboi^ babe, itf compared with that
of the adult, will be seen from the foUowiog table, in which the details are
more full tlian ihoie f;<veu on p. ii'ZH :
Wllglit of ouii In peiTvnlur
of iKlilj-I'Vltht
Era . . . .0.28 oi)2«
Bnin .... 14.34 2..^
KUmts .0.88 0.48
SUn . . . tlJ 8.3
Li*w .... 4.38 2.71
lleitrl .... 0.89 UjS2
Stomach and intculine . 2.S3 iM
Lunpt .... S.16 101
Skploton . . Iti.T tS.30
>Iu»:)c», etc. . 2.XA 43.1
Testicle . ami 0.8
WirtEbt of uniiin In
■it nil, u oompftred
wKb mat of neotani
tAtv uivn M I-
«n
It will be obeerved that the brain and eyes are. relatively to the whole
iMilr-wetKhi, very much lairer in the babe than in the adult, aa b also,
though t(> a leas extent, the fiver. This disproportion is a very niurlced em-
bryonic ft*lure, and, aa far aa the brain and eye are cniiix-rned at least, haa
a llK>T]>bolof[ical or phylojteoic, aa well aa a phyaioloj^cal or teleol<<i;ieal. aig-
nificADce. Inasmuch aa ibe amaller b'xly haa relatively the larji-er niitfoce,
ibe akin is luiturully prci|)ortioiiately greatwr in the Imbe. ' It is chiefly by the
■ociimtilatiiui of muscle or flwh. properly so-called, that the child acotiirca
the Ittilk and wi'ight uf man. the skeletal fraroewurk, in spite of ita ncing
■(wcilically liifhlvr in its earlier cartiliiginous »>ni)ition. mainbuninj; thrviugh-
oiit lift) about the Mine relative weight.
The incn-*M< in stature is very rapid tn curly infancy, proceeding, ho««vef,
by dnn-asing inerementM. During or shortly before nunerty, there is a^in
a KMntwhal Ruilden rise, with a suli*ei|ucni more aieMy bat diminiiibing in-
crwa* up to about the twenty -Rfth year. From tlwnce to about fifty ytan
of age the hcjght rcmaint ntHtionary. aft«r which th^rc may lx> a docTWM,
eepecinllv in extn-niv <ili) ng<i.
The inrwwK- in weight U also very rnpiil at first, and proceeding, like the
beif;ht. with itiiiiinl<bine iDcr«nicnl>, mar eontindo till altmit the fortieth
year, .\flcr the *ixiictn T«ir a decline of variable e-xlent i« generolly wit-
neaani. It i^ a remarkable JVicl, hunevtir. thai in the (irft few days of life,
an far from thrre lieing an increaK-, tWrv i^ an actual dwrmsc uf wei];ht,
so that, even on the seventh day the weight riill euntintiee to be lea than at
birth.
The saliva of the babe is active on tinrch.and tta gastric jmce, unlike that
1006
THK PHASES OF hlTK.
raai
1
uf ninny oewboni aDimah, hoa t(ood peplii? powers, fmm whiirli we mny iutrt
UiAl iu iligottive pr((C(w» in general are identical witli that nf the a<luli;
but lh« foots of lb<- infant contaiu, beaidcs owiniierable quauliir uf iioili-
mstod f(M>d (fat, cnwiii, etc.), unaltered bile-|>i(;iuenl, and utide(WiB|Kavd
Dil«-«ttltfl.
Tilt hunrt of tlic balw («eo Tabic. |i. 1005) !», relatively to it* body-«tjg)iu
larger tlinii tbc ndult, niul tlie friijiient?}' of tite bmrl-bcut Riiicli tffeaicr,
vii., nliixit nil or NO per miiiiilc, fiiUiriK In nbmit I 111 tn tlii; M^^ioa ri«r.
and idimil 90 tn ihc tcntli yimr. Vonr^poniVing tn thv smiiller bulk oi tlia
body, the wliiilv circuit of the blimd nvFlom i* truvcrwti in a i>hnrt«r tiat
than in the adult (12 »ccond#ne ii};iiini>t 'i'^); »nd.e"ii*r<jiieni]r. thr- natwal
of the blooii in the liwuc* iicxccr.lingly m|iid. The n-»pimti<>ii of ibt bab*
is quicker thiin ihnl of the ndiill. King at first nboiit $'> per minitle, fallii
to is in the second yenr, to it) in the tilth yenr. and to onuard. Tb«
apimtory Rork. while it incrcnsee aheotiilcly as the body grows, is. relati
to tfae body-11 eight, grcnicat in the enrlier yo^re. It is worthy of iMtice
tbe abeorplinn of oxygen is snid to be relatively more active than tbe
duolioB of carbonic acid : ihni is to say. there ib n continued acvuniuUJiaa
of capital ill tbe form of a store of oxyKen-holdinR explosive ooinpuuiKb
(•ee p. 472). This, indeed, is tbe Btrikioj; feature o( infant meiabolisn. It
i« a nietabolisiu directed largely In constructive ends. The food taken repR-
tents, uiidoubt«dly. so mucn potential energy; hut before that euerKy ou
aiHunie n vitid mode, the food iiiuat Ite convened into limue; and, in auflba
cymvemion, morpholueical and molecular, a larife ainouiil of ener)a:y maXfe
expended. The metiibolic activities of llie Infant are more pnniouocMl than
thon of the iidull. for the ^ake, not xi mueli of cnergtea which arc npi-oi m
the vorld without, as uf en«r|^c» which are for o wliife burieil in the rapidly
iocreafing muM of (Ivsh. Thus, the infant roqaire^ over and ubuvo the waiit*
of the man. not only nn income of energy corresponding to il>o eoet^ oftbr
tleeh actually laid on, but nUo nn income cormpondinK t« the energy nail
tip iu making thnl living iiculpturcd Hesh out of tnedcad nmDrpboit? ptnlnii,
fnls. c&rhoIiydnitc» and it»lts, which serve as fowl. I.)ver and above thL*,lkt
infant needs a more rapid inetabolisni to keep up tbe narnial hodilv temper-
ature. This, which is no lefs, indeed, slightly t'l.:t°) higher, than thai i>f tfct
adnlt. requires a greater exi>eiidilure, inasmuch as the infant with its rels-
lively far Larger surface, and its cclreincly vasctdar skin, htsea heat f ■ a pto-
portionately much greater degree than doee tbe grown-up man. It is a matw
of cnmuion experience thai cbUdren are mure aBfacted by culd Lban an
ftdults.
Thin rapid metaboliara is. however, not ninnifeBl inimediatelv upon birtk
During the firai few days, Mines pi wdiug to tbe l«a of weight QientiOMil
nbdvc. the respiratory aciivititw <>f tbe tiHaiicn ar« feeble; the cutbryocir
hebilK u*'" us ret uul t<< Imvc U-i-n cimiplotely ibrowii ofi^ and aa waaMaMl
on [1. 4!)'l, newbiim nniinitU hiuir with impunity a deprivation of oxyEte,
which wiitiid be fatal In diL-ni Inter ou in life.
The quantity of tirine paMMit, though scanty in the Knit Itvo ilaya, rio*
rapidly at Ihe end of the first wer.k, and in youth the quantity of nriiK
paired i*. rehitively to the body-weight, larger than in lululi life. Tltie nuy
M, at Icaol in quite early life. |Mirtly due to the more liijiiid nature of llif
food, but is bIko in jmrl the result of the more Hclivc metabolism. For u>4
only is the quantity of urine passed, but nlm the Hinounl of urea uul eoiw
other urinary cunslilncms excreted, rclntively to the body-weight, greater la
the child tbiui in the adiilt. The presence of uric, uf oxalic, and. nccordiBS
toeome, of hippuHc adds 'in unusual iiunutiltef is a frequent chamrterollo
of the urine of children. It is slated that calcic phoe)>liales. and, iodani
THK fHASKS OF I.IFIt.
IWT
I
I
\
tili<i«plmte§ i^nerully, are deficirat, beiDg reuiiie>l iu tlw l>Ml3r fur the
buildiuK ■'[' "' l-"^ oi^seoua skeleton.
AHodiili'd |;>n.)t>a1>lr witU these const nivtit-e labors of tlie jirciwiii^ ftvme
b the pruuiiueiioe at (he lymiihiilk srsteni. Not only lire the lyiu|>hiilt('
gUD<h largely developed uiil mnre nctJve (as is pivitHibly Hbown by ihcir
tendency (a diseoM in ynuthi, but the i|iiHUtily or lyiiifih ein-ulaiimi in
gniitcr than in Inler vHin. ('hHrHutertni<-<if youih ix the niie cit'tbe thyiiiiix
body, vthieh iiu'reaiifti up to the ^ecvmil yi-ar, itnd tiiuy ihcu remain Inr n
while ctatioiiary, but KeiterHlly belhre pubertv, btw vultered ii retrocrxn^ive
iDCtatnoriiho«iii, nnd frenueiilly hnnJiy u vcyligc ol' it rviniiiDii beliiDil. The
tbyruid [mdv i» alio relnlivoly ^reiilcr in the bnbc ihaii in Uh' lutiilt: the
flplMti. on ihc olhfr hand, which growi' rnptdly in turly infancy, i« not only
aWiliilely, but aloo reklively, ^n-ater in tne mhilt. It nwd luirdly be said
that the recuperative power of nifnm-y and early youth i* very itiarlced.
It woidd bo beyond the Mstpe n( tWn work (<■ enter iiilu the jwyehieal
condition of the bnbe or the child, ami our kuowled^' of the details of the
working of the nervous ■^tlcm in inlinney i» too meagre to permit of any
profitable disciiwion. It ie hardly of dm to i>iiy that in the young the whole
■errouB eystem \a more irritable or more excitable ihiiii in later years: by
which we probably to a great extent nieou that it is lesK rigid, lera marked
out iato nhat, in preceding portions of this work, we have fl{>okeii of aa
nerviiua luecbanisms. It may be meHtionetl that etimulaiion of ibe varioua
cvrebral areas, in newborn animal*, does not give ri»e to the usual tocalJKed
movemeiits. The lense of touch, both as regards prewure and temperature,
•ppean well derelofwd in the infant, aa does also the sense of taste, and,
pomibly, though this is dispute*!, that of smell. The pupil (larger in the
infant than in (he man) acts fully, nnd Donders observed normal binocular
movenienitf of tlje cyea in an in&nt lets thun hour old. The eye b (in nutn)
from the oiiLnt fully *eniiittre to light, though of courM riiiual nerccptions
are imperfect. A* regard* h«ariu);, on the other hand, very littV reaction
follows upon Mouudit — I'.i-., auditory •enxationii term to bo ilull duriiif; the
firil few day of life: ihts may bo partly, iit lea»l, due to a))M!iice of air from
tbe tympanum and w tumid condition of tho tympunir iniiennK ni«nibrnne.
As the cnild grows up his feii»e« rapidly i.iilminal(:, and in bis early yoar« he
poMMMB a general acuteiie«> of »>ghl. hearing and tuueb, which frequently
PffOllH* blunted its his pychical life become tnller. Children, however, are
•ud to be ten apt nt distinguishing colors than in sighting objcrls; but it
doM OOt MMiear whether this antes from a aunt of |>ereeptive discrimination
or from tbeir being actually less sensitive (o variations in hue. ,\ ehjiraeter-
islic of the nervous avst«m in childhood, the result, probably, of the more
active niclabulism of the body, is the necesnty for long or frvijuent and deep
slumWr.
Dentition marks the first epoch of the new life. At about seven months
the two central iiieiaon of the lower jaw make tbeir way tlimugh the tciiiu,
followed iromefliately by the corrMnonding teeth in the upper jaw. Tbe
lateral inciaoni. lirHt of the lower anu (heti of (lie upoer jaw, appear at abntit
tlic ninth month, the first molars ut about the twelflli month, the eaiiiuee nl
fliHtut a year and a half, anil tbe tem|M>rar>' dentition is completed by tito
npiH-ainnee of the M'ciind midan usually I>ef»re tbe end of (be •eooiid
year.
About the aixlh year the [lermaneiit dentition cmninenvea by (lie ap>
nearance of the first pi-nuanent molmr beyond the secowl Umponxj molar;
in the seventh year ihc central [HTmaneul inrlsors replace their temporary
representatives, followeii in tlie n«st year by tlH- lateral iociaor*. In lli'v
ninth year the tem|Mirury tirst molars arw mjiliu'ed by tbe first bictiapida.
TRE PRASSa or tIPE.
aud iu the (eitlli year tbe second toniporurj itioinn «re simiUrlr rr|ilu«d
by ilie secuud bkuspids. The caiiiiie!) are exchanged hIkiuC tbc eleventlt or
Lwdfth )'ear, and t)io ^coiid |)eriaaiienl tuolurs ate t'lii ahout thr iir«lfUi <it
iliirleeath year. There is iheu a lung pause, llie iliml or vrltdom iMrtb dm
nialciug ita s[i|ieunuice till the seventeeuth. or evpn twenty-fifth yoor, or ia
Mime c&aen a»t a|ij>eiiriut; at all.
r>horlly alter the cmoluwm of the )>erniaiieat dcnliliou (thv wisdottt teHk
vxMpteil) the oix-urrenoe of {luhcrly nmrlu the hejriniiiti^ of • n«w plunaf
life; aud the dilleratioe between the itese*, hiihi-tin merely |H>ic4iiia]. m«
becoDiea functional. In both Kxei th« maturation qI' ih<- gi-tii-ratiw ongaai
M nocotupHDied by the well-known ehiingea in the body at tar^: but tbe
ovrntM are iitii<;h aiore churHe[«mtic in the typiual feoinie thiin in th« abet-
mnt male. Tliuu^h in the buy, the breaking of tbo voiw sad the ni|iid
growth »{ ihe binrd which iiociini|iaiiy the itpnearonce "f avtive Sperntate-
20a, nrv xtrikinf; fnitum, yet tliry are. nfltT all, superficial. The cuma of
his inrreuing weight and hi-iglii, iind of iho other ev«nt« uf lib econooiy.
pitraur for a while longer nii unchanged raiim : the boy does not beeurn* a
miui till Rome vcnrs uAcr puberty ; and tho decline of hia functional inu
hood is so gra^unl that frequently it c«ii«es only when diaeuae put* ma wmi
to a ripe old agei Wilb ihe occurrence of iDeust rust ion. on the otbor haad,
at from thirteen to neventecn years of ajio, the girl atmuM at oooc becoim
a woman, and her functional womanhood ceaaee suddenly at the dimaclMir
in the fifth decennium. During the whnle of the child-l>earinK period Wr
organism is iii a coniparatirelr slatiouary condition. While tM-iurr tba tgt
of puberty, up lo nbout the eleventh or twelfth year, the girl u lightarai
shorter than the hoy of the same &Ke. in tlie next few yean tier raw itf
growth exceeds his ; but she hns then nearly reached her niaximnm. while
tie coiiliniu^ to fftovi. Her curve i)f weight fmni the ninvtwutb year ■•owaril
to the cliiuacliinc, reinainii stationary, being fol lowed imbsequenttv by a kl«
inereitse, so that while the inan reaches his ruaxiniuiu of weigh i ai ahmt
fnrty, the wumiin iii at her gn-iilent weight about tiflv.
Of the ■tiitieiil tlttferenci-s of iwx. loinfl, »iii'h as the formation nf On
pelvis, luiil the ci»tnl nicehani^in of n^ptration. arc directly conuecdv) with
Ibo act of chilli-bearing, while uthen have only an indirect relation to lh»t
duty: "nd i ml i cations, at knst, of nearly all the ehnracteriatic diffli>reiK»
are seea at birth. The baby boy is heavier and taller than the bnby girl,
aitd Ihe maiden of live breathes with her ribs in the same Huy n.i ikmi ihf
matron of forty. The woman is lighter and shorter tliai the nian, tb* lim-
its in the case of tbe former being from 1.444 to 1.740 luetrtii of height, aiul
from Sa.f^ lo DS.d kilos of weight, in tbe latter from t.4fiT t» 1.8!KI n? heiglil,
in the latter from 1.467 tu I.HDU of height, and from 49.1 to 94.& klhie of
weight. The muscular system and skeleton are both abdoluMlv and rela-
tively less in w<iniiin. and her brain ia lighter and smaller than tiiat of ntsa.
being about I27'J grammes to 1424. uer inetabuliiuu, n» moa«ur«d by lb*
respinilory and urinary excreta, is also not only ahaoluiuly but relalirely 1*
tbe body-weight less, and her blood is not only leas In <iuiuility. but alin of
liubler specific gravity, and contains a snuiUer proportion of n-d cornnscha
liierBtKOgtb is to that of man as about 5 to 9, and the relative lenirtn nf btf
sl^aslOUOto lIo7.
Prom birth onward (and indeetl from early inlra-ulerine litet the incr»
ineut of growth pi-OKmaively dimiuishM. At Inrt a point is rvacfaed at
which tbe curve cnte the abstiiiwa lini-. nixl tbo incnmtenl becomM a dteia-
ment. After the mltiiination of iiiaiihnix] at forty and uf wooiaohnod at Um
eliinacteric, Ibe ptiuie uf life deelinei into old age. The meidiidic aetivitt
of the body, wbteh ut first irn* suliii'ienl not only to cover ifae daily wada.
»
»
it to ii'M now iDHlorUI, Inrcr on is able nnlv lu tui>el the iliiilr waiiln, ami
nl t«>t in tii'> iiiiiN-rlVYt even Ui puHtain in ilt vniircty the exutliis I'rnm^.
Nrillirr ili n-gnnli vigor lunl riinctionul npni'ity, nur im rogurtl" nt-ijchl nnil
bulk, i|[< iIk' itirnin^'txiint* of Ou: sa-vrnil liftiniv iiD'l organ* <-<)iii('i<li- «ilWr
nith ciivb other or with that ol' thr bmiy iil targo. W'c hnvf ulraiuly (VU
lb«t the lilo i>( tticl) an crgnn ns lhi> thymii* i* tinr Hh^rtfr than that vf itK.
pOMCMor. The eye i» in [t« ilinplric prime iu phildhom!, when it* molia nru
cImpM nnd its niiioi'iilnr mrchiinitiRiv mi«t niutiiU-, ami then it fi>r the riuwl
pan Mn-n as n tiiy ; in Inlor vmre, when it couM bo i>t' the gn.«t4wi >N.Tvir«
to m i>till avlivv bntin. it hu« almcly fullen iiilo » clomled unci rigid old a^p).
The )ikrl«tou rciich«« itF limit very ncnrly lit the »anie time n» Uu) wbolft
fnuD* mehcs it< niaxininm of heijihl, the cuiilcscvncc ul' th« various eDiphysM
being pn-tiy w«ll eimipkled by about lh« tnenly-filth year. 8iniilariv tbo
mnwiilar «T«leni in it« incraisc tallies nith the weight of th« whole (khIv.
The brain, in ipile ul'thc inereiiHing eoniplexily nOtructuiv and function to
whieh it continue* to nttaiii even in mitldle liTe. enrly reaches it# linitt of
bulk and weight. At about seven years of ago it Attiiiiis whut may Iw eon-
sidere«l as it' tir*l timil. lor though it may inerejisc somewhat up to twenty,
thirty, or even Inter yeare, ils progresi is mueli more sli>w after than before
•etreo. Tbe va«cular and digestive oreans as a whole may conliniMi to
iDCrMM even to a very lale period, trom these facl« it is obvious that
though the phenoneDa of old auc are, at bottom, the result of tho individual
decline of lite MTeral tiMiiea, ibey owe many of their features to the tlis-
anrnogeraent of th« whole orgaoisin produced by the premature docav or dis-
appearance of one or olh«r of tlio coDStitueut bodily liiclors. Tliua, for
iDsiaoee, it is clear that were there no natural intrinsio limit to the life of
ih« nii)H.'ular and nervous f)-«tems, they would nevenbelees come to an end
in c-oitr«<(ueuce of tlie tiutrilivif diHturbanoee cuuM-d by the lowi of the teeth.
Aod what ii> into of th<- teeili ia probably true of niauy other organii, with
llie addition that llieM euniuit, liko the tovlli, be rvplaoed by mechanical
ronlrivanon. Thus thi,i tenn of life which is ollottetl to a mu.tcle by virtue
of its niolecular ounnlttulion, luid which it vouhl not exceeil werv it alway*
pluciil umh'r tbu mint favomblt* nnlrilivo eoudilicm*, is, in tlw- oi^nism,
delerniiiivi) by ihv similar lifi-ltrnw of nthrr tiiwuw; the future declino of
tbe briiin i* pmliably involved in the eiirly deray of the thymus.
Twodiangas ctiaractericticof old age are the fo-eullcd cniearcoitsnml fatty
dagCDCnitiuiM. Thcae are seen in a completely typical form in eiiriiUg^, as.
fur irwtancc. in the rlha; here the protoplasm of the cartilagororiMiscIo
becomes hardly more than an envelope of fat glob«il«s, and the supple matrix
ia n-nd(Ti-i| rigid with amorphous deposits of calcic phoM>hale« ami cHrhn-
UBicx. whit-h are at tlu- rame time ibe signs of past and the cause of ftitnre
nutritive decline. And what is obvious in the case of cartilage is more or
less evident in other tissues. Kverywhere we see ■ disposition on tlw part
of protoplasm to tall Imck upon the easier task of forming lal mlher than Ui
nny on tbo more aniuous duty nf niflnufaeluriuK new malerini like ilself:
everywlwre almost we see a tendency to ihu replacement of a structure"!
matrix by a dejicait of atnoq>boiu waterial. In no part of the svstein la this
uwre evi<)eni than in the arteries; one conimoa feature of ol<l af^ ts ilie
oonvertion by siirh a cliauxe of tlie supple elaalic tubes into rigid ehannela.
whereby the supply to the variouu tiieueaof Dulrilive material is rendeiv<l
iofireasingly more diHlcuU, and their intriuric decay prvtportionately hurrie<l,
Of tbe various tii4u«e of the body the muscular and nervous are, however,
ibose iu whidi tbe fuiK-tional decline, if not rfruciural devuy, beeotoes aoonest
apparent. Thedyoaiuic coeAicieDt of ilwtkeleial muoclea dimioiabw rapidly
alter thirty or fbrty yeata of life, and anmilurwant of power ooiuea over the
61
1010
THE rnASES or life.
ra tbt
1
pUiti iimtwulnr fittrr* al»n; iW heart, (bouKli it may not (liiniobk, or
tamy slill iiicrciuw in wi-ij;ii(, poMciwiw luM awl lex force, aiul the amvw
uf the InUstine, l>Iii<l<i<.T, aixl utiier orxauH, <llmi»ii*li iu vigor. In the ocrr-
Mia Hj>l«iii, tbo liiicw of mtiittaiicfi, winch, a» it« hav« ao«u, belp In map <M
ibe centnU orgiui« iutu inechaniftmK, und ho u> produce iU iiiulufiiri«n
. actioiM, bocomo at Imtt hiii<lnuuT9 to tlM puMigc of nerroua tiupubo* tn
dirDOtiuD, wbitfl at ihi? »uan: timv the molecular cneisj' of iIm impuUei
•elv«e becomca lott. Tli<- oyc l>vci>mM fnblo, tint »iily fnim nl»ii<Iina« of
medta oihI prMbyofHC mtiAcular inabilitjr. btit aUu I'mm ihc wry bluDUtOMof
thoretiiin; tlic vni^^ory aixl motor irapulmv pom with iorrvMug alownca ta
and from thv cpiilriil iutvouh vyvtcni, an<l ib<; bmin brcouun A mors and
noro rigid mttSf of pr<>l»plik»ia, tlic molcculiir linc« of which riitlior mark ibi
hlrtori,' of past iiclioii* rnlhcir ihiui errvc n« indi<:«tions of prixeut |iot«Bc;F-
The cjiitbc-iiul glunditlnr vlBiuaDts seoni to be tho« whum powers ara tbt
loiigCHt prcMtr\'vd ; iiiiil bunce tJio man whu iu the prinH' of hia manhood
a " nartyr to dj^spepfia " by reason of the wiisitivriic» of ^tric nerni
the reflex iiihiuitory and other rceulu of their irritation, in hia lator ~'
when his nerves are bhinted, nail when, therefore. hi» peptic cells iirc &!
pursue their chemical work uudisturbcd by extrinsic nervous worrios,
and drink.* wttb the oouro^ and sucooas of a bov.
Within the range of a litelime are eomprieetl many periods of ■ dxkv or
test frequent recurrence. In spite of the aids of a complei civilization, all
tending to render the eonditloiui of his life more and more equable, man still
sbowB in his economy the eflects of the seasans. 8ome of ibeae are the dirM
requite of varying temperature, hut some probably, eticb as the K*'" **f «eubt
in winter and ibe loss in summer, arc habits aoijuired by iloacenl. Witfia
tlie year, an approximately monthly period is oianifefited iu the fejaali
meoBtruatiuu, though there h no ejcact evidence of eren a latent
cycle in the male. The jibeuuinena of recurrent diMiam!^, and ibe
critical <Uys of many other malwliee, may bu regnr<lod a.* pointing to cy^l
of amaller duration than that of ilie mouu'a revolutiuu. unltm nc admit ibf
view urv«<l by ttome authum that in tlieK cascu tbe roeurrcuctt i" U> lie attrib-
uted rather to periodical pboM!) in the diwdue-pruduciuf; germ lUvlf, than W
variittjoiu in the medium of the disease.
Pnimiitenl Hmon;i: ull other eycHeul event:> i:t the fiict that mmi animah
poWMiing a ncll-develupeil nerviMi* iiyalem, mtwt, nigbt afWr night, nr day
after day, nr at leaM linn: alV-r time, lay lliem dona to sleep. The siUmM
feature of sleep i* Uia ottwition of the nuloaiatic activity of the bnin : it it
th« diastole of the cerebral bent. Kui the condition t» not eonEnwl h> tfar
cerebral hemisiihcres ; all part* of the hmly either directly or indirectlv takt
sliare iu it. The plicnumuna of sleep are |K:rha|w aocn in their simpleat form
ill tlt« winter-sleep of hyhnrnation, to which cwpecUUy cold-blouded a^'iimk
hut alsii li> some cxtrjit warm-bloiKle^l animals, are subject. In tbew cm
the cold of winter slackens the vibrations and Icasens tno axplosiooa nt tlw
pm(u|»Ia»m, not only of nervous but also of imiscubtraiM) glandular sme
turvs: indeed ilic activity of tbo whole body is lowered, in 8on»e res]wob
almost t» actual arrest. At the same time that the labor of tbe oerebtll
muIceuW liocomes inaiifficicnt to develop cofudousneas, the respiratory oeotrt
is either wholly quiescent or discharges fooble impulun at rare iotervaKanl
tbo heart boats with a slow, infrequent stroko, not by rea»on of any inlubt'
torr restraint, hut because its very substance in its slow nwleoular travail
esn gather hnul for ex plosions only after long |iausc« of rest. And sud
few ami distant lnonts as do ooctir are amply sufficient to meet the needs of
tbe tboble metahoiiain of tbe several ItMues. The sleep of every day difin
ttom the sleep of winter>cnld chiefly because the slackening of motecaUr
TUK P0A8K5 OP Lirs.
1011
I
I
I
ICtivitMM n <)ur in tho termer not to oxtrinKic' hiii to intriniiic oiutM. not to
chaogM iu tlic medium, hut to oshituglion i>r the »iibjuet, nnii bocautc lli«
plMQomeoi Bra Ui^geljr confiuod to the cerebral hcn]i«pUi.Tm. It it tnii; thnt
tbe whole body sham id the eondition. Tho piilsQ unA breathing niv stom>r,
the intc^tnc niict »lhor iiileninl miiwulsr mechnnianis arc more or l«n at
rcflt, the eccrcliDi; orj^iiRs acv ic« active, some appari?iittv being whollir qoi-
est'ent, and lh« sici'iior »n Wiiikiiig nib« his erM to brini^ hack to his ninjunc-
dva its needed Dioislure. Indeed the who]o mclabolism and tbe domndent
tamperatiire of tlie body are lowered ; but we i-annot say at prewnt how tkr
tbeee are the indirect rranlts of the condition of the nervous syalem. or bow
ftr iliey indicate a partial «[umbeiini; of tbe several tissues.
Thoracic respiration is said to become more prominent than diaphraf^inatic
reapiralion during sleep, and the Cheyne^toxes rliythoi of respiration (see
p. 491) b frw|ucntly observed. Uurin); sleep the pupil ie contracted, during
atep ileep exceedingly so : and dilation, uflen unaocompanied by any viable
morements of the hinhs or body, takes plaue when any sensitive Kir&oe it
Mlnulnted: on awokine also the pupiU dilate. The eyeballs have been
guieimUy described as being during sleep directed upward and converging
or, Bccording lo aome authunt, dirergjng; but others maintain (hat iu true
sleeri the visual axes ore parallel aud directed to tbe br distance. The eyes
of children have been deecribed as ctintiDually executing duriui; dieep move-
iuent», tilVeu irregular and uatymmi'trlcul and unocoompanied by cbKiigee
in tliv pupiU.
We arc not at |>rt«eni iu a pwitinn to tnux out the ovciil* which culmi>
luic in thU inactivity of the cerebral ttructurok It ha* Ix-en urged thai
during sleep the hruin ix augpmiu; but uven if thi* aim-niin is a conatODt
accompaniment of itlet^i, it niuitt, like the vascular condition of n gland or
any olner active iirgnn, be rc^nlvd as an effect, or nl least as a «uli«idiarv
event, ralhcr than iis a primarv causes Nor can the view which regartts
sleep OS the r<«ultof a shifting ul' the mechanical arrangements of the cranial
circulation bo considered as satisfitctorV' The oxplanalion of the condition
is rather to be sought in purely molecular changes; and the analogy betweeo
the systole and diasti^le uf the heart, and the waking and sleeping of the
brain, may be profitably pushed to a verv considerabM extent. The sleep-
ing brain in many rSBpeols closely reeenibles a quiawent but still living veo-
Iricle. Bolh are, a> far as outward niauifestalions are concerned, at rest, but
both way be awakened to ocUvilr by an adequately powerful stimulus.
Both, though (|uie»cent, are irritable, in both the quieooeDee will uliimalely
give place to aoiivity. and iu both lui appropriate tumulus applied at the
ritchi time will detertuiue the cliangv from ml to action. Just as a single
pnok will under certain ciraumttance* awake a ventricle, which for some
seeoads haa been luotionlew, into a rhythmic activity of many beats, so a
loud noise will start a man from uleeii loiu a long day's wakefulueas. And
just OS in tbe heart the canliac irriiaoililv b lowcat at tbe beginning of the
dfautole aud iucreiues onwanl till a beat Mirata out, so b xleep deepot at its
«oumenc«meDt after tbe day V labor; thenee onward slighter and slighter
itimuli are needed to wake the sleeper. Forjudging uf (he depth of ordi-
nary nocturnal iilecp by the intomty of thv notM; requirMi to wake the
sleeper, it may K- tv>iiclu<bil that, inereoaii^ very mpidiv at finit, it reacliee
iu maximum within thu tint Itour; from thence it diministiM, at first rapidly,
but afterwaid more »liin'ly.
We caouot, however, at present make any definite sint>-mont)> concerning
Iht oaurc of tbe molecular changes which ilctermine Uiix rhythmic rise aad
Ml of cerebral irritability. The &ct ihat tbo produci* of protopboiBio
■divily wbe« ihcy oocumulalc within tbo protoplasm appear to booome In
1012
TII& rrtASBS or tirs.
ealijU
f3
tbe end an obftruclion lo thnt nctiviiy, hnxsugcvetcd the i(Ii.-nlliai tW ]irr»rOf«
in ibe cerebral ligeiip {>f uu excc«e nf the prmlitcU nf ncrrmis in<-Uiti'ilirm n
Uie <'au»e of ilecp. In'ieed luetic ncid, thi> incnuue of vhich uriu sapfwMil
to be llie caUf« of the acid reaclion of muecular and nervous ttuum aiier
exercise, haa been eBpecially pointed to in this coilDectioti ; but. iia me hm
wen, the acid reactiou iu quefelioii appears Dot to be due to any incrraMd
Sroductiou of lactic acid. Bcfiidee. if the sccuniulntion of metabolic [inv
ucW of any kind were the caune of sleep, it is not cleor why we ehiMiU
ever have any hope of uakin}," More may he said in f*vor of the c*>ai
lion that during the waking hoitia the expenditure of oxyeeo exceedi
invoine and thai the (|iiic«cence. which we call tieep. comes trntn lhi< < '
lion of the body's ttore of iisygen, more esjwcially of that " intnt-mol
oxjften of which we spoke, in d^aliug Vfilh th« revptratioii of (he
Uut tn this view luii-il be added some hyjiotheBia, auch ih th« liyplny ofnov
iuhibiiory medianiitm, « hereby tbe rellpirutut^■ cetitru ijt not mtuied to ib-
created activity by thin lack »f oxygen, for ai we have itrfu the lirrnihtitj
sliarea in the dluiuber of the body, lliouj[h continuing in piny with u
amount of enervy, which |icrnii Id a gradual mioraiion nf th(.> lott itoreaf
oxygen aud do finally brinic!* on the awakening vrliidi end* thtr sleep. And
tbe uccrafiity fur such u cnniplication indioatCN thai tlu; cxplntiatioo i*.Bl
prewnt nl leuM, inadi-ipjali.'.
The phciiumc-Jiu nf xlcej) nhow very clearly to liow litrg« au exiMit lo
apparent anioinaliiim iit lIic idlimatv outcome iif the elTrclB of nntcceilnl
Mimulation. Whr-n wu wioh to fio to idcep we withdraw oiir automatic bniD
H much fl.-> [KiHsibKr from the inflncnoc of all CxtTin«ic •tiniuli ; and an ia-
1crv»tin^ CH'c i» rrctinlcd nf n lad wlioi>i- connection with ihr external norid
was, from a complicated anicsthevia, limited to that aOordcd by a singlecyi
and a single ear, and who could be sent to sleep at will by cioaing wcf^e
and slopping the ear.
The cycle of the day ia, however, nianife^ed in many other w«ya than bf
tho allerualion of eWping an<l waking, with all tlie indirect eH^cl* of ibo*
two conditions There is a diurnal curve of temperature (tw p. 6-19), ap-
parently independent of all immediate circuiuslances, the bereilitnry iniprva
of a long and andent «ei|uence of days and nights. Even thv puUc. «
•enaitive lo all bodily chan^cB. sluiwa, running ibmugh all the immeiliaif
eflTedn of the changee of ihe minute and the hour, the working of a diarvil
influence wbidi cannot lie ai-counled for by waking and alet'ping. by «urk-
ing and reMiiiK, by rneulu and aholiuence between meals. And the mmt
may be «aid concerning the rliythni of mpirallgn, ao<l the ■inxlucta of pal-
monar^-, cutaneous antf urinary excretion. There seema to Iw a dailr iwrw
ofbodily melabulitim, which in not the product of the day'a avBOts. 'XVitUa
the day we have the narroni-r rhythm of the mpiratory ceiitrv with tlw
accompanying Hue and full of nclivitv in the viiiuimoUir oenln-d. .\ail
lastly, ihi-n.' stiind" out the fundamental fa<'1 nf all lioilily |wriiMliirily. ikit
alicrnnlinn of the heart's fiyttole and iliiistok- which cascf onlv at lUnlli.
Though, as wc! have *crn, the intcnnitlcnl flow in ihi- arlcnt-ji 'u loueil ilm
in tht ciipillaricM to nn appnrrnlly (•oiiiinni.Hi" How. slill the coii-^tnMtK iw
I>enlrd cycle of tho cardiac (huttic must leave iln nmrk ihr.. Iif
whole web of the bwiy's hfe. One mwins of invt^igalion m .r.
riill too groM lo pcnnit us to truck out ile iultucn<-c. .Still lit« are hi «I
present in a nceition to ^ay how far (he fundiimenial rhythm of the* heart
ilsclf, that rhythm which is intlucnccd. but not cmilcd. by the ohaon ft
tW body of which it is the centre, is iho mult nf ccvniical cfaanf;>ei, lae rr
Hcction as it were in little of the cycles of the univerM>. or how liar it b ih*
outcome of the iahercut vibrations of the molecules which make up iu nh-
Htanec
J
CHAPTER VII.
DEATH
Wbek t)iv aiiitiiul kiiigiloni is surveyed from a bmwl iiiaii(l[M>iiit. U be-
cnfnf* ohTicM-i thikt lliit nviini. or its cuitelHtin' (be »pr.rm»Uy):.nna, \» the giwl
of nn iiiili villi) 111 i;xi»lri]cc; lliut life is u cycle beginning in nu ovum itnd
eoRiittg round to nn uvum tigiiiit. Tlin gnwter pnrt of the ■clton* which,
lookiDK from a amr point i>l view at the bii;bt>r niiitnal* nlno«, wo nn; apt
In rorutidcr m iiminriitly tbc piirposw for which niiimikls ounii! inli) exi»t«noG,
whi-n viowwl frmit ihr dixtiiul oiitlcdk whcniv the whole living worlil u tar-
Toywl. fiulc uwny into thd likaieM of the rauro by|)lnv of uTum-boitriDg
orguinnit. The Rnininl body i* in reality a v«]>icl« for ova ; and nfldr the
liw of Iho paivnt has become polAntially renewed in the offspring, the body
r«iDain« iii^ ■ cm i-ofT envelope who^ future U hiii to rlic.
Wcft- liu' nnimal friinie not lh« conifdi'wicit mnchinc wc hnvcwonit to
be, death might come lis a »implo and gmdiinl itimiliilion. the " wins overj*
thing" U'lii^' the laat etoge of the iiiicceHive Iuh of fumlamcntal ponren.
Ad it iw, however, death is always nioro or len violent ; the machine comes
to aa end bv reowMi of the disoraer caused by the breaking d'lwn of one of |
its {NUU. Life oenses not because the molecular powers of the whole body
tlackeo and are lost, but be(»u»e a weaknefe in one or other part of the
lunchinery dirows it* whul« workins nui of gear.
We hav« seen that tlie central factor of life ii the circulation of tiie
blood, but we have aita m«ii that blood b not only umIm*. but injurious,
UolaiH it i* <luly oxygenated ; and we have further m>«ii that in the higher
animal* the Dsygcnation of the blooil can only t>e duly olTecleil by meant
■ >f th«' rv9pinitt>ry muncnlar u)ccliiini*m, prtsHdetl over by the molulla ohloD-
gum. ThuK thf life of a c<impW nniinal li, when r<<ducc<l to a simple form,
eom[NiH-d of Ihm' factor* : the muinteniini-v of the circulation, the aocoai of
air to the luGoiogluhin of the blixiil, and the funclional activity of th« !«•
tiMratory centre; and denth mny eouie from ibc nrrcrt of cither of these. Al'^
Itirhal [nil it. death tiikw jilacc liy the hcjirl or by the lungs or by tlM
liniin. In rviilitv, liowcvvr, when nv pioh the Bnalysis further, the ooolntl
tact of death is tlie stoppage of the heart, and the conee<)Ucut srn^t of the
cir«ulatiun : the tisMet then all die. because thry lose tWr internal medium.
Tlw- failure of the heart nmy arise in itself, on account of some failure in its
iicrvouH or muscular elements, or by reason of some miecliief aSecling its
mechanical working. Or its stopiwgv may bo duo to sotne fault in Its inter-
nal milium, such lor instance as ■ waul of oxygenation of the blood, wbioh
in turn may be caused by either a change in the blood itself, ai) in carbonic
oxide imisoninf , or by a failure in the mechanical c-onditioua of reapiratioa,
or by ■ ceesatlOD of the action of the rapirutory centre. The failure of
this centre, and indceil that of the bean iuelf, may be eauaeil by nervous
uifluencci> itruceediiig from the brain, or brought into o|>eraiion by meana of
the central iicrvoua system ; it may. on the other hand, be due to an imper-
fei.-t tttate of btuod. and thia iu turn may arise from the imjiurfeict or |>or-
verse action of various ceoretory or other tiMues. The mmlo of death ar« in
reality as DumennHi aa are the poaible nMdlfications of the various Gurfom
L
1014
dkatu.
of life : l>ul they aII rod in k Mnppnge of tbo dmilation, uk) tbc «itli-
dnivnl frnm lh« listut^t of tlirir ini«riinl niMium. Hroce w« c»ni*> (o cdc
sidcr the ilenlh of iho hody n» niBrkc<l hy the cefcation of tbc bntrt'a bt*l.
ft CMMilion frotn which no recovery i* possible ; ntid bv ihie no are nulilcil
lo lix an exact lime at whieli W(i toy the body ts deatf. We cnn, bovrvvrr,
Ux DO BUch exact time to (h« denth of the individual litsiies. They are not
mechanisni!!, and their death it a gradual \iist of power. In the caw of tb*
coDtrnclile tissue;', ire have apparently ■□ rigor mortia a fixed tenn, b^
which we can mark the exact time of their death. If we admit that mfba
onset of rigor morlia recovery of irritability is imposuble, then a ripd
muKle is 0D6 permoDrotly dead, lu the case of the other tiiisueA w« btt*
no Bitch i>bjective eij-n, aince tbe rigor morlU of aimplo pruioplaani nutni-
festa itself chiefly by obscure cbemical ngns. And id all ca:«s it is ubriow
that the piiBttibility of recovery, depending »a it docs on ibe skill and knml-
edge of the experinieDtcr, is a wholly artificial siuo of dealli. Vet «« an
draw DO other sharp line belween ilie secniioglr dead tinuie wh<«v lifv h*
lliekered <lowti into a Hmoulderitig ember wbicli cwi still he fanned bark
a){ain iulo flame, nud the bandfiiT of dimt. the n^regalA of chirntiiail Nib-
Btaiicea iutu which ihe dccoiniKising tiMiii<> finally cruml>leiL
Moreover, tbc fiiiliire of ihe heart itMlf u at Imtton 1<M of irriubQitf.
mid the prunitiiljly of recovery hr.re aluo n-sto, a« far as is kDUWii at prcMi^
tiu till! nkill and kntiwledge of lUwv vho iiltempl to recover. .So that ttUr
all Ihc I'igiii' of tbo death of the whole body are as artificial M ib<M> of tW
dcutli uf the coDslitUGDt lis»tic>^
APPENDIX.
ON THE CHKMICAI, BASIS OF THE ANIMAL IIODI
iiKsalniBl bodj, rr»m n cbeuiical pointof ci«w. mity he reeardcd uaniixlaro
, €>f rRiioDK rcjirpMDtitilvtH <if ibre« large oImhm o( cliemkal HDbaUncM, tIk., pro-
leida. <:>rbohT<lnil«s tind fnt*. iii nmocidtJon with ■mtillrr qaanlitio of vmioun
••line and other uyttallinc b»tli««. Ily pratciil* are mmat bodin coDtuiniug
«ubon, oxygen, hyJroKni iiiid uitroKen tii a oeruin praponton, rarying within
DaiToir liiDtlt.aii(I havlii)! crriaiii general fmtorea; tbeyarc frf^iufnuf iiiolLpa of
aa albumiiKHd*. B,v carliohydrsi«« iin< mranl MaKhea and •iigiim and tbtir iiJIio.
We have also Krrn ihnt thr animnl body may be conaidcred aa an aMemblage r>r
protoplaani uiidtrT viirinut ni«clifii!iiiLon<i'nndi}rniimeroaa praductaof protnplaamic
adivity. We di) nut nt prtoeui know tiiiylhing deflntle about Iho Diotecular oom-
poritMMi oriictivc living protoplasm ; but nben we ■iibmit )irotoplaani to cheioka)
aoaljttt, ia which met \i ii killed, wo slwnini obtain froin it a conaiderable ijuaii'
tity of the material spoken of a* prolcid. A iid manr nuthora gn no fur ua In apeiik
of prohiplHam aa bt<ing purol;>' iinileld io nature; ihey reganl ibe liviiiK proto-
plkMn M proteid malehal, whtcli. in pitMJng from death to lift', box BMiinicd cer>
tain cbaraclrni and pmuinahiT bas been chantcfd to <-(>n>tnictioti, but itill i* pro-
Uid natteT; ihey eoaielimrs spmk of pnitoplmin ns "liviiij; jtroield " or " living
^bmaiD.'' It ia worthy of notice, liowrnT, ibnl even liuiple lormB of pmloplaem.
like that ci>ii«tittitloe toe bodv uf n white cor|>uitcle. forma of proto)>la«m which
we may fairlv conaider »» native protoplaKtn. when they can be obtained in auf-
fideui quiuilily for chemiral annlyni*, are found lo contain aome reprnwDtatirc*
of uirUihydrntra and futo iw well a* of proleidi^ We might. perbaM, even go aa
Imt M Io My, ibat in all forma of liTinKprotoptaam. the proteid baalala fonad npon
aonlyxia to bare tome cafbohydmte and aome kind of fat aanocijued with il. Fnr-
ther, not only don the normal food, which it eTentiially built up into prolaplaam,
eoaabtof all Ibrec dnun. but. na wo hnvc tern >n Ihc lecliona on nuiritioti, i>ro>
loplasm mivea rixe hy melobollam to nieiubem of the aanie three claaw* ; anil, an
far aa we know at pnaeot, carbohedniiea and fais, when formed in the bodv ont
of proteid fiHxI. art) $0 formed by tlic tucvaty of living protoplnam. by aome lii-ing
Uaaue. Hence there is at least loiiie leaiwn for tliinkinjc it probable tfaal ihemQle-
eule of protoplasm, if wo may lue >ucb u pliraw. is fur muru complex than a laolfr-
culvof proleld mnller, that it eouiaini in itM-lf raldoce, »o to a|iMik, not only of
profeid, but aUo of carbohydrate and fatty material.
Be ihla aa il may, for no dntpnntic i^tatefneDt can nt prraenl be mode, whein ws
examine the varioun lijuiie* uiid lluidi uf the aniuial liutly from a chemical iioint
of view we find pmeiil in dillVrent idacea. or at dilferent times, several rarielir*
and drriiativea of the ihiec chief gIomio. wc find many forms of pruleicb, and
bodi*4 cIuMly allied lo pnitrida, in the forma of mucin, gelatin, etc.; many wie-
ties of fata; and Mveml kiiidi of carliohydraten.
Wo find, moreover, muny ulbrr brHtin which we mar rexnid aa atagoi in Iho
coMtmciivc or de*lructive metaboltwn iif boili native ana differentia led prolo*
plasm, and which are important not no much from the ijuantity in which they
occur in the animnl liody at aor one time aa from their llirowing light nn tite
nalor* of animal mctaboliaiu ; these are aurh bodies as urea, other organic crya-
talline bodif^. and (be extnwtiTes in general.
In ibe following pagta ih* cbemical featurm of iba more important of ibeae
nrioo* aubaiaDcca which arc known to occar in the animal body will be brfeflj
eOBiJdwed. aiH-b cbaraciera only Iteing deicribed ■« pOMeaa or promiae to pow«aa
lOia APPENDIX.
phfwaloxioal intorat. The pbysiologiul fonclicin ottny oubtUnc*' miial (]<'[iMiI
ullima(«r]r on ita molecalnr (inclndinK itn chi'iaicali tiKlurp; nml Iti'tujth kt pr*ciil
our cliemical knuwled^je at tb« coagtlluciiu or aa auiiuul inklv ^'im lu biit little
infi^bt into tbeir pliysiolo^k-al [iroperciw. it cannot be doubts tbai «ticrb cliiai-
cal inf»rnifttion a* in altninablo i» n nvtvvury preliminary to all |>by<iiot«gl(Bl
Hludf.
PBOTEIDS.
Then fbrm th« iirlocipa] aolldB of the mu«oular, n«Trou«, iind f landular tiauo.
of the wraoi of tjlood, of tteroLi* flnidi, aimI of Ivmph. In a hviilitiv condition,
■weat, tMra, bile nnd urine cnnliiin mere tr«ciw, i^aoy, of pruicida, Tii«ir genual
peroeotfl^e compiMtlioii inajr he Iak(>a as
0. H. N. C 6t
From S().9 CD I&2 61^ Of.
to 2S.& to 7.3 to 17.0 10 fi4^ 1o i (I
(li(ippe-8i]rl<r.t^
Then HxunM Brv obUiiKwl rn>iu n wiuidenlton nf nunieroiu aatlyuM, rilglii - 't
■lie rarlena itaulu Imliii Iminitrrial, irliarv (h* jiurtly uribn (ubtUinc* ofna'. tl
li«doDn<talv listennliioi,
111 adilitiun 111 tlin nl'iif AinMiliioiitciinitpida l«a*««in iitiiilimi a *iir<< iridiA
In Ilic uuf of e|u nlbumlti iha ;irlu<-lpaj cuiiiitltnvnuof til* »>li irx vlit- i I'ain u4
|iiitaMi<tm. Ihi^ lnlUr icrMilyvxuHHint tho ronuor in aiii>-iunu Tl>r in: ihtlMfrf
ciJliim (IkI lulnuiiim. in ■•>iritiiUfitii>ii irilli phrupboru-. iiilpliiirtc. i>u<i I'ldxaal
icr^ iuiliII 4fuan1iliDd of {u1t.>lillri, n>a^n«iir>iEil aii<l ir i' ti r i.'-, t< r' r> T^nl
U aLi^ft A ImoF of >L|jrfi,' Th<* a*H '>f frnifil-aJhuiMi Ir
liUt Iht- ruli rtf Il.n jifuTui.ii of iriutcle *iiiilaiu> nn i-v ■ ■ ii ;i,ii ,.i: il.i
titlnrvol IheptinuccUon <>rilic iipii niih tliu nnjiniil it all II a iiia(t<f >■! olxt'driiy. UMnafi"*
|](Mtiiitltjl>in j« KiH (j^ l^nv* iiii »Jh on icntV^I-
IVolddn u met wllli in Ibr animal body are all amufplioiw ; rodic bit folu
«>iHiie intnlublo in wnter. nnil iiU ntc, for ibf mnit part, iluolublo in altnliolt
«lbcr: (licyart.' all lolubic in iinirii; ntidiariO nlkatica, bat in lirC'>aiin);di*«Jtid
modtly undi-r|;i> dee'^iuiiOBition. Tiieir Buluiioun i>iiiu«iu it Irn-baml^ iiiiaviii
ncttoo on the ^lanc >if pulAri/.iliuii, the amount depending on vAriau* cirtus-
stanoos. and boioji. <rith one rxccpiiin, vit., pejitonca, cliangcd by beatiii);.
'CiyilaU iaio »ho«* coniprii><i'<ii rvrtaic imiUiit (tapieiiiallj globnllii)* cleiiwiiit ••(et mn
liAt tiDoa obaarvad (n tha *euil> of iiihiit pUnu ; ai y»l (h«y liat-* nni beam Mit.ila>rf aaf.
Ilolaullr luUM ar In r^tiinliti-M liiriv ct'ioufli Ut firrmil •ny'mnla aimljaia la b* mait
A mvlliol of iiolatloit In ciDantilv aail r«i'cyalaltulnf tliuM •uMuuiVf liu. Ivowenr.'lMM
liiillcalwd.aad it •ctma probable tbat analyaiaof llieae inar load to lUlafvatinf lilf.iiinallwi
uu tha aulyMl of tlia cvnililuUnu and ODmwnalioii* of |inil««d*>
Tie prcacDC« of pratt-ida may he drtprininod by Ihe folloiting Icets :
I. Hrat«) with ilranx nitric acid, Ibcy or their Miliiliuna turn y«llaw, ami thb
nilor I*, on tfaendditiuD uf ammonia, or cauHlic ftoda or pr>la*h, cliangvd tvadw]'
urwngtr hue. (.Xanthoproteic reaction,)
i. With Millon'H rcatcent they givr. when prraent in milTlctfnl <|UBntiiy, a fw
cipttato. wbii;h turn* rod mi hrniins. It tbi-v nrr 'inly prexini in lra<'M, no pn-
dpilalir ii ublnincil, but merely a red colotsiion of the w>luli<in.
3. ir niiied with Home coneenlraied solution of sodic bydrale. aod oti<> or two
dtotwi of u Nolutioi) of oupric aulpbalo, n violet rulor I* obtained, wbiob dMpaoi
ia tint on boiling.
Tbe above Mtie tu del«ct the Mmalleat trace* of all prDl«>bl». Tbe two fallovbl
tcatii may b« uoed irhen ibere i« Diora (ban a tnoe iirvwiit, but do nut bold to
erory kind of prot«id.
4. Render Ui<i fluiil xtronEly acid irith noetic Of other acid, and aild a few Anf
of a anlution of fcrtucyauiue of putaJHiuoi ; a precipitate atiiina thfi prwBBM •(
protddl.
■ "" ■' "i -■■M.B.ta.
■ ■ wi.
■ |i|f!i^-. Jipiim I [.■,ir,. ii.Hi. ->. I.. IM. IIX. MNTtt. AXIt
K. ICentler lli« lluid, a* befoK, lUoagl v mM witti &<.«t!f ncli], add nn <v|iisl
TuIuiBc of ■ cotiL-etiiritWd M>1utlan of todtc »iilpluii«, aixl boil. A pfNipiUbi t>
Ibnnod if proutiik i* itraMot.
ThlaUMrwctiiiii imvful.not nnlrou ■cixunlofitHZBflnMs.butBUBbMBUM thartamDlB
iumI proluM oo <l'<>'iii[i>mii<>ii »f AilMr budiM whlob iiidj tw iimtnlj aiul liaac* aftar fllln-
liDB Uie Mne lluM nikf lis [iiril>«r *nAlrwd fer other ■abMaboM. Adilitlo^iat mctlwd* of
frvaui|*<n)ii1iini Train |iiVtMil> aivi acMulatinC with w^tlo u-iiluiil tfiiliiil- nvoiJinsMljr
•soMOf th* >»•! '. |irepl|>IUIioii h^f exoMofalathol ; lu IIm Ullcr oub thn tolulioo tiiuil M
nvalnl or fititilly t^i'l lloppt'ScjleH iMoninwiiil* (he cmpln^inenl nf a atuntod •olntlon
Af fttobly |in>i-t|iitBt«il hnie lijdnM, fn MmUc xml : thi* it *i1iIkI Ut (Im Mlutlali, »Iiil m
botllag til* wlioie Kf ihApratoliU *re pmdpiUleJ m v*II u the rarrio hIi. Um latlor ■* a bMio
•(•taM. Ilni<t*'» Riaelind oF reuiovmii ifi* t*«t tnu<(* or pnit«iil» fAini f Ijincvii mliidim* U
*Uii iif iiH |n« g]ycoa*n>. I'tTCi|ikalfu>i of ttio IjuI tninv uf jirulciili b)r ruauu of li/dntod
«ii<l« nf kail ala twiliiig tcnijwnlijrr' inii}- im nlio rtaplgjrwi.
Prolcid* na; be cniiveDiently divided into cIhmw.
Ci.AMt I. ^/t'rc .l/AitmiiK.
4
Mcrabem of ihia clnM, a* Ihpir name iinjilir*, nccur in a nnturai coaditiOD in
nnimal t)«UM aad fluid*. Ther ar« soluble in wnter. ara iiot pn-ci pi luted by rery
dituic aL-lda, br ciirboiiaiai of the alkalica, or by amlium chlunile. Tlicy an
voa^tilaied Ity heating in aoluiioii to a U'lnjwrature of about 7"' C. If dritd at
40* ('., the rtviilting mana I* 'if n I'nlc yellow color. CMily friiililp, laalitlcw, inodor-
OUa and nalublc.
t. BcB'Alburola.
t'ornia In aiiueuus Milution a aetitral. tranaiiareiil, yHlowbh fluid. From tbia
it la prmipiuted l>y excwaof aining alcubol. If lb« al«ubol be rapidly rcmoTcd
tbe]>rKi|iitoU) may b« readily rcdlaaalTwl In witlvr, ir«iibjeoUMi tuleiietbicr action
■ coaculalion occ-'um, and ln« albumin is then no InnRcr iliui aolublc. KtruQg
acid*, eapeclally nitric ntid, cauic a coagulation ximilnr to that produced by hcAl
or br the nmloogrd action of ak-obol ; ibo albumin bcooaio profoundly chaniced
by llic Hdlon of tfae ui'id itnd due* not diaaolve upon rciuuval of ib« acid. Mer-
curic chloride, argentic nitriiie. and lead acetate, (>reciiiii.->te the albumin, funuing
inaahibtc comiHiund* of Tariable cnmpoution with it : ih« prooipituila may b«
rMHOvtd by roeHDn of aulpharoltod hydrogen and (he albumin agaip obtained,
a|tii«r«iilly unalietrd. in M^lutioiu
ntning iir«tic acid in excoM ic'*^ "^ pr«cl|>llKl«. but when the aolutlan U cod-
cetitrat^l the iillxiincn M iranaJToriiied intua transparent jelly. A similar jelly la
[inxluuW when atioTie cauettc potash i* added In a i-oneentrated sniulioa of ent*
aDiuntiu. In Wh ibne caaea the Mibelancc w profoundly allwcd beeOBing, In
the (loe COM, uoid : in the other, allculi-nlbumiu.
'Hie apedflc rotalofy iiiwer uf c^-ftlbumin in nqueoua aolHtlon la, for yellow
lichl. — ^.6^ Kydtochloric acid, added until the reaction h stroaxty acid, ia-
creaaM Ihii rotation to —37.*'. The fnrmnlioo nf the gelatinnus conpuund with
i-aiutic puiiuh ia at firat accompaniiK) with an increaie, but Ihia la followed bj n
decreate of nitatiun.
tWp-ifiii-'ii, White of hca'*-cf g b broken up with aciaaon Into small piece*,
dilnml with an c>]ual bulk of water, and ilie intilure shaken aUnnply in n llaak
till quite frothy ; on atandinic. the fonm riaca to the top. and carrica all the Sbrea
In wliow roeabworlc the ultiumin wiu c^uloincd. The fluid from which the foam
liMbeeo remond, la atraiiied, and trMtedcarefullv with dilute acelio acid aahmg
M uj ptecipilate it fanned ; the preeipitate b then filtered off, nnd the ftllrato
Kfker ncu trail jEitinn coiicentraled at 40" to il« original bulk.
Serum - nib umtn .
Tbitfonnof albuniiit rcaemblM, to a ptat extent, (he one previously dwcrlbed.
The fullowing may suHIm as dlalingulahlog fratorea:
]. The taeeillc rotation of eerum-albumtn b —S^; llut of egg-albumin ia
— ^."i'. bolli mensarfl for yellow light.
«d. I
(it>(M.,ft«7, ■ IMknalMar, btiMhr. r, ini|nM>'CI«n->M, tl. iinntftail.
i
lOlS
APPEXOtX.
i. Swum -albumin k uot i-auBuliied by bcii)g ilukra uji with rtbvt, ria;-kn»-
BUD is.
.1. Scmm-iilbuniiii in not vtty rptidily iirrcipiiiird by *ui ng byilruchlorlc idd.
uihI ouch picd[>itii[e >l> doc* occur in r«m<lily rcdUiwlTcil on futlhcr addilioii urilw
aeid ; the exact jvrtnn of lh<M l*ro restu'tM boldi gbod for vfg-a1bun>iB.
4. Pndjtllated or coagulated «crum-albiiiDiu is r«a«ljl t folDbrc, ryx-albuBiio il
with difiicult}- fKiliibIc, in Rliv.nii; nitric a«id.
5. fr^-slbumin, if injf^'lcd nubcalcDcoutly or into o vein, a p[i vara uDaltntd la
tli« uHdc ;' BLTum -album in uniilarly injecied doM not tbiu iMTinallj |i«a* ooi by
tli« kidney.
Sernnt -album in is found not only in Mnoii-wrum, but b1»i> in hnifb. bath thai
CODtained in tbc projief' lymphatic cfaannvli nod Ibal dittuHcd in the ti»ue*; is
c)ivl«. milk, Iran ludn lion H n:id loiitiy puiliulu|;ii-ul fluiila.
It V* ibis furm in wbicb Hlbuuiiu gt'cvrally ajipeHTn in the urinp.
In addition to (lie bIwvd. ticL«r*r' Lun ilocriUr'l two rIOKly relalid lulW, Ui abiifk )>•
SlTW Ihn iiniDiK juinilliiimiii nixl iiwUlbuiniD. Thv fint lie oMalnvd Itoia irtanali I'^ff*; m
■IkaliD* loluiioiii ■» raninikaUa for bclDg ven tojij. Il Hvaii ikniUAil •li««jHTiU«la4/
ba erolaiih it HilT'^naMiiibly iiKvmpMilTon Ihjm lliMe. Haeitia* giiva a* htaOMpnUta^
0. 3i.S. B. n.9, .V. i:.H. C. £13, D. I.t jiw mdL ll twoiu to ba ametatol wliL H'Xm W;
like Elyw^Qi oijiaUe of being DDbireit«) itklo a •utrilnuuc ei<ini tL« r>' -^
MslaJbiiiiiiii, fuUDcl in • ilrnniiMil fluid, tvnmbl** tliv iirfci-'iiiiK.fcvl ii ' i tf
hydfochlohc add.ciT by antic add aiid bTRH-yaiilde afiioUiainBi ^ it ia j>rr('>|t 1111*4. w »*
wagulalad. by alcobol , ita aolDtion I* (nirorly oupiIilM r-n boiling.
Albumins are gCDerally fniiiitl nisncJAled wiih rntiill but deGtiilc anuniiHa'
•■]!ae mulicr A. Scbmidi' hoi* (hut lliry niuy br frrrd rrom tlicae by dlalyM,
and that ibcy urc then not cooi^ulutod <m bulling. From ibia It mlgbt be Infmrri
Ibat the albuutiu uud ihe aulme uiaiters wiie peculiarly telateo, anil tliai il>
latter pluycd «onie specini pari diirin); the coRgulalibu of lh« tanatr by hm
Sehmitll'a oborrvntion^, however, have not been corictanively c<MTo1xint«d by Mb-
■ecjueut olim-rvfrn.
('labs II. Derive^t Aft/imiHt tAtbumiiuitety,
I. Ac Id -albumin.
When a native albumiD iri suliition, such ad aeruni-albumin. ia ireHied far Mae
little lime with a dilnle acid, »ucli as hydrochloric, the pmperlie* lK«(>me enllrdy
changed. The mnat mntkrii channe* are: (I) itiut ihe inlutiun ia no loogtf
eosgnlated b> hrnt; [2) that when the lolution in carefully nruirnliied the wbula
of Ine proleiu ia thrown down ai a precipitate; in uiber wi'irdH. ibe MTuinalbumia
which was soluble lu water, or ai least In a neutral fluid onliitnintc only a anufl
quantity of neutral taltn, hnii become conrerted intoanitwlanire invilublu in waw
or in ainiilar neutral lluid>. The body intii which aeruni' album in thud becann
cooveried by ibeM^ioo of an acid ia aiJitkeii ofas ncK/-aU«Mii>. Iia characleriKii
fealure* ari- that it ia iuHoluble In distilled water, and in Dcntral Mline MilutinM
anch na thote of sodic chloride, Ibnl ll ia readily wluble in dilute iicidH or dihnr
■ Ikalica, nnd that lis MilDCions in acid* or alknliea are not conKiilatcd by boilin;
When iiua|i«ndcd, in Ihe undiiunlTrd alntc. in water, and healed to Tt'" C.. il bt-
eoiBM ooafulaied. uud ia then uudialint'nisbable frfia coagulated neruni- albumin,
or iDdetd troni anv olher form orcxisgulnteil proteid. It is evident that tbr im^
atance when iu sotulion in n dilute acid is in a dillertnt rondition frmii that n
which it ia wbcn precipitated liT mill rail ration. If a iinaiilily nf aeruni- or r^
albumin be treatiiil >rilh diluli- liydrochlnrie acid, it will be found Ihat the me
vvrrion of the native iilbtimin into ucid-albumin ia gradual; a^|>eciuleu lieaitpl
to 7fl^ < '. iminediately alter Ibe addition of the dilute acid, will ronf^ulwte alsal
a« uaual; and annUier apecioirn taken at the fame time olU ii^ve liardty auj
precipitate oa neutral iutioo. Some time later, the iuicrcnt dcjwndiDit on tb*
prof>onionof ibeacld to thealbumin, on tcoiperpture. and on other circunutaKe^
the coagulHtioD will b*l«a«,aad Ibeneulralintion precipitate « III be (WBaldrrabl'
■nnko^nreb.up-ii
r. IVlliol aaal.. IM-K
• aBn-4(rClMan.ui'i <
^o, :
'^dc rAilioiniiiuncBraieltailMr: slBlxluiaan.jlJtb
. ll., *. lav
* kurV) Jtiehtv. s). iin»i, a i
thioiin doK'B oil ncuirnliuitiiin.
If fiDdf-chofiptd miuole, from which the aoTublc albumins bare br«ii rMOOved
br repMUd waahiDg, be treated Tor tame iltiie witli ilitute iO:i jrer cent.) Iiydio-
cfalonc add, the ircaicr put nf tlio muscle is diMohed. The Iranoparvnt acid
filtrate containa ■liiriti' ((imiititr d( pmtHd inntrrinl in n form wtitcli, in ita gen-
eral cbaracten at Icuat, ugnm iritb acid-utbumiii. The ncid anlutiun ol' the i>r<>-
leid i> not ci)«eiilaled by boIKu^, but the irhol« of ibe proteid Is pn-ci pi titled on
ncuir»li»ition; and the preclpiuie. Insoluble In neutral *odic i-hlriridr (oluitoni.
in rendity diwolvrd by rvrn dilQUi nridti nr nlkntie". The proleid thim obtained
from mUKcIc hiia bci-u mlled n/nlonin, bill wc bnvr at pnvriil no iiitisfiiclory I«4t
Ui diniinsuitb the acid-albumin [or iTDtoiiin) prepared from inuHcle from ihnt
prepared from egg' or serum-albumin. When coaurulaled albumin or oUicr
cosBulaled protfid or fibrin >■ dinolved in xtrong adda, acid -albumin ii rormed ;
andwhen fibrin or any ulfacr prolrid in acicd upon bj fcaatrk juice, twid-albuBiin
la QDeofihi^ firetprodui-ta; aud ihcne iicid-albamins cuinot be dbtingulabed from
aeld-albuiiiiii (>re|>Hr«d from musile or native albumin. Though nydrocltlorie
acid i> prrbnpR tlir mn»t eoovcnirnl Rcid for fornilng acid-albunin. Other add*
may sUo he ii*ed lor the purpOMo of tirrpnritiic it. Acid-albomin in toluble nol
tonly In dilute alkolle*, but alao io dilute Kirutioni of alkaline urbonatfa; tla
voluiion* in thtee are not coagulated by holUng,
If (odic phosphate in exrera i* added to an acid solutioo of acid-albumin, tbc
acid-albumin n predpilated; thii also occun on adding iodic acetate or pho*-
phate.
Ah upecial te«lt of acld-albumln niar be glren: 1. I'nitial coa|^latloD of its
•oiniton in lime-nairr on boiling. 3. FurUier precipitation of ibe Mine folntioo
after boiling, on Ibe adililion of caldc chloride, magneuc aulphate, or aodk
chloride.
JDlaaolved In very dllul« hydrochloric add, add-albumin (ayatonln) prepared
ftom nusdepoaetaMaaBpCviDclnvo-rotatonrpowerof— 72* for yellow light, tbU
bving independent of the concent rail on.' On hcMing ibe solution in a closed
Ttaael in n wiiirr-lialh, the roUmrj- [lower rim-* to — 818°.
The body kDonn n* panipeptuue. which mnkrn ita appeann^e during the peptic
digeaiioB of protf^id* in closely allied to the ■ubnUuiccii juol dncribed.
3. Alksll-albumln.
If aeruni- or egg-albumin or wnahrd nuede be trealed nllh dilute alkali litftead
of with diliile acid, ihe i>roleid und«rgo«a a cbiiiige ijiille fiimilar to that which
waa bmugbc about by tne acid. The alkaline •oluiinn, when the rlungo lua
become cainplc4e, ia no longer coagulated by he«t. the protcid i* wholly preclpl-
taied on neutralliailou. and the prrcipitatr. insoluble in wnler >ud in nculral aodio
chlorine •olulloni, is readily Mluhle in ililutt- acid* or alkallei. Indeed in a gen-
eral way it may be (Hid iha't aild-ailininin and alkali-albamin are nothing more
tluin aolutioDn of ihe name ■ubalnnce in dilute .icids and alkalim mpe«lively.
When ihe precipitate tiblained by (be nruiraliiationof n noliitiuu of acld-Blbamln
in dilute add la dbaolicd ioadituK! alkali, il mar be eonudered to become a) kali-
albumin; and coniertely when [lie ptecipnie ol'lained from an alkali-albumin
•olutinn i* diivfdrrd in dilute mid, it may \«- regarded »% add-Blbumin.
It ia utalrrl' ana chararteriiilii' ri'iulinn ni Ibia muiliDrd or derived albutntn that
it ianot precipitatrd when ila alkaline lolutioun mm neutraliced ia theprcwBCWof
alkaline phoiiphates; aolmlona of acid-albumin, on the contrary, areaaid to b«
prrdpltaled on neutral I sal ion to ih* preaence of alkaline ph<i*pbale>. and ibU
difference i* coniidered to bo a diatincnifbiiig feature of the two prateid*. But
doubt ba> been caat on Ibn BlateiueoL
Alkali-albumin may be prepared by Ibe action BOt only of dllole alkalie*; bat
alio of alrong cauMtit! alkalies on native albusilna an well a.1 on coaguUteil a1ba>
min and other protdda. The jelly {-rodoced by the ntiion of ouMic potaab oji
whiteofepg, ipukenof in Clam 1., 1. la alkali-albumin; the aimilar jelly produortl
bj alroag aceltc add i* acld-ulbamin. One of the moat productive nethoda of
& bj alroag
■ Umipe^cTttV. Bilti fliTi. ralb. Cium. AliaL. BiL IV. (L«t), % VA.
• nep|i»-i«ii»«. ibc <-it. s IV-
•WjiVa. rHipR'tArcb.,tl-l. lU •tr(n,!> MI,
^
1020
API>BKUIX.
obUloioK ul kali ' kl lilt m in U that inlroducwl bjr Li«l>erkEihii.' and conftiua In vlilitif
■ slrone idutiuu of c:iii*lie f^tiub lo fiurifiM white of ecg until Uie *borv-inw
lionod jelly It oblain^d. Tliin ii tb«ii l-iii Into tmsll pteom. and iliait-Md uabl
i)iiil« whlt«. Tbe Iuidjm Hre i^bon dionolved bv hratine on lh« wnt«r-hatli, tad tbt
alknli-nlbiimin prccipitatod bv (lie cnTodil niMiliiin of acetic itcid.
Both alkali* and acid-albuniiii me u-iih ililKculty prucifiitaud by alrahol
tliMralltHlinoor ncidflolulian*. Tbo imutniJixatlon {ifevipilalea.boNeTtr,''"
ooagulatwl under the pra]oa;ged action of alcofaol.
Til* bfldr "prutciu,"JM.>ribed lijr Muld«r,k|>fwan,iftl*xi(Uat >1I,U W eUaal^ miaM«l
witb tlkli hody. All *iil>*n)uaiit olwervvr* hnv«, liumvM. bll«J u> oMilnii lii* tivwk.
The rotatory power of alkali-albumin ntriea according ft lia XMirce ; tbuavluB
prepared by alroag cauMltc tiotiuli Iroin aeram-albainln, the rotation rtMi tnm
— 06* (that of aoruiii-nlbiicnin) lo — .Sij° ; for yellow ligbl. Similarly pmand
from <gK-albuiDin. il riM» from —3^-5° to — 17° ; and if from conjnilalM wfatia «f
^g. it nm to 68.8°. Hence the exUtence of rariooa form of ancKll-albtuoin b
probable.
In additinn to ilie methoi* glToD itburB,»ll(ali-«JbiijBln lay b« >1m r—dily «hiaia»l kf
■haklaji milk witli Mmnii otutlif tmla •ululi<in niiil •ili«r. roiBOving tho •ihvrlal (biIbUiw.
ginvipilating llir rviuniDiui; lliiU wIlli tcaiif ui-id and wiuhlug tlic pncijitLale wilb mtim.
oold alculii-l and Filler
Tbe niml natiMfnctorT method of regarding acid- and nlkali-Blbumin u to coo-
Mder ihuiu iw rccpuctivcly ucid iind iilknii compouDdi of tlie DPUtmliKalioo pre-
cipitate. W« have reoMMi'to tliiTik tliutitheii tlie precipitate l«diHaolved In eltbcr
nn acid nr an alkali. U doea ei)[«r iuiu txinibl nation witb llicm. The aeulrallu-
tiuu prcicii>iiate i« in itnelf neither acid- nor alkali*albuaiin. but nay bvcoot
cither, upon Kululion in the ro*pPctire rengcnt.
Ii il pmbkblt llui Mvtr*) <l«rir(ot nltiumin* riin,< Jil!«rinE uvwIiDit io tba jimuiUl tma
whioh (hc^flrv lanaei or (nuibly *L-ui>rdiuK Is lb* uiodc of iWr |mpualhia.BiiJ itulnd
<if lli*M> may ax id ill Hi mrr* Ulii* Tmm»nf M4d- aad idkiilJ-iilliulHla; but Ui* wliala MilfM
requlr«> taitimt iiiivalij^lion.
Aoid-albumin, prepared bv the dirri'i nclion ofililuie ooido (in natire albumin.
or on Eaiucle-iubiiiaiii-». coiilaiiiii auli'liur, as tihowu l>y the brown ooloratiou wUdi
appeaiawheii ibi- prei-ipltateEa bt«ted wItJi caustic |>olasU in the prvaenoeof biife
lead acetate. Alknli-albumin, at all eventi aa prepared by the aolioit of elroui
vuuBtit- poliub oriioda, di»C« not contain any >ulpbur; and the acidalbumin. pre-
jured by tile ■■ilulit>u in an acid of the neotralitUion precipitate froiu Mich as
alkHll-alkuuiin aoluiiou, i« Hlinllarly tree from Bulphur.
*. Ouala.
TbU it the wdl-knoirn protoid eKiMinic in milk. Whco freed from (at, and ia
the moiot condition, il ii n irbitu. friabte. npn^ue li'uly. In miMi of it> react *
il corteepoflda clMcly iritbalkali-albuBiin; tbiu it isreadily uilublelndilatei
and alkatiei, and i§ re-iiteclpilated mi iientralli^tliin ; if, however, itotoMlo |
|>liit» in prenent^ oa la the caM in milk, the eoiutton uin»t be Mrongly arid bdbft
any precipitate b obtained.
Vartoiu rMcHsot 1i*t« al diffareiil lttn««ti«aa OMijtiwd Mim»>Ii> » diiiii>';ii:<t>it.( ii ttat
(liBcloMir alliiMl body olkati-itlt-uiniu. Law rurMrrbci h*>' ^4
•o tauvh dnnbt on (mm diUvrcuoH Hint the idoniiiy nr mxi .Ji'
ull'uuun rauit (Utl bo left mi opsa iiumIIoo, th« dltc'uMiou vl •n.i.it Yri>L.iij na nm <» |UA
her*.
OMBin.aiounirrlDg la loilk, hat had Mtfral melioat aatribnl W i(. M^aroicteriMlei ImI
Ihea low their inporuuiM on oMuideiliii that milk containt. tn Hl<lllian to tmada. oitm
•«hllAaow. wii'li ■• |i il-uiifl phMphit«. anl a iiaoilwr of bodiM which yield M*da by brntMi
Uttsn- Tha prMHoce of pitouio pli«i 'i .' 'i < - ub >«paai«l mlluauo* oa Ibe rawiUMitaf
mwm. Id 11m entire aMeaoa of tVi* . .^.id lo Ibe •lUjIUii qoaslitiai, m aian ft-
bjii'C aiihydnda.gUet a pfeoipiul*; l> i Ji it pceiani. <\irb>nu^ aiiliydrida. k>>«* an
tmapflala, and ontie aeid only ouo whqii Uia mIuUoii !• atld tnioa (ha preauKH of b««a«<>
aii.l iijt rrocn that of acid paloaio jihoapbate.'
> FtaMtodorff^ Anaalen, M. luirt. ■. It*.
* Kamir. nanriAteh.. M ivN.ii>C«u&'l*i.
* Au* JCUK l-Hitb. d. t*yiul. Cbtm.. IM. fl. M.
OnSUICAL BASIS OF TU£ AXIUAL BODY.
10:11
VThtn i>np*TtA IVom mitk hy ms^iriic *ii1t>b>rp (Me> bflnw), tnri by eibrr
(ram fau, aoi) ijignolvcd in witttT, cnarin powcMwii ■ iprdlic mlulory pnwrr of
-^80' §0T yrlluw ligln ; iu Olluie iilkallnu solutioiis. ot — Tti^; in vlTOUg alkkline
•ohitiow, of— !'!' : in ililiile hyitrdcliluTiu acid. o( — SI".
Caacia h*» bcrn wturtcd to occur in muadv. in t^rnu* iluiil*, and in blood-aerum
iM-rum-ca*rin). In many eawa it ban probiiUly brrn courouDdcd wilb gUibaliD*
»te Clan III.); but blood-aeram and iDUncli'-pliiKinii iiodoubti-dlr coutain an
aJkall-albuDtin in addition lo whatever globulin may be [TeMnl, ^ut Ihe u«ual
dnubt exiats as to ihoidcntityof tbi«wiihliuecnaciii. Its prewoco mity bp*howii
bj adilinti dllulti ncetic jirid to blao<I'ii[>riim wbirh ba« bcra ffrcd from globalln
bf a cntrt-nl nf cnrtionic atihvdnilc ; a dialiocl prt'cipilale i« (lirown down. A
uibaooce diniilaT to i-uavia fiaa also bi-en dmcrlbed aa exbtiiig iu uwuialed
moBcle and in ibe nrntcplnntn of iicrvc-cclln.
/Vrporo/fon, [>iliit« milk nitb fevrrni (10 to 1.') litnr* il* bulk of water, add
dilnU anUcacid till a prectpilale begin to appear, ibrn para a current of carbonic
anhydride, fitter, and w«ali Ibe precipluie wltb auier. alcobol and «tber: tlie
complete removal of thu fat carried down with the caitcin preaenia some difflcu)-
tiea. Ma^Mtc rulphnle addiil t" ■nturnlioii uUn prrcipitatri vaacio from milk;
■be precipitalo ai thus 1orme<l i* rradily lulublc on Ihe addition of water.
Clabh III. OMhA'm
BnidM the native atbumlna then are u miralier of native proield* irbtch difli'r
from Ibealbnmins in not being aotoblc in <li(tilled water: ihc.v nrnl for their
aolntion tbe prewncc of an appreciable, ihoiieh it may be » rnnnll, cjnuntiiy nf n
nentnl aaline body, nuch aa aadic chloride. Tbuii lliejr rcHemble tliv ii I bum inn in
In not being aolubie in diilHIed water, but dilfrr from tbein in being aoluble in
dilute sodic clilorido or other neiitrat talino aoluiiona. Their general chnractera
may be «<ated a* follow* :
Tbn are insoluble in water, soluble in dilute (1 per cent.) lolutiona ofaodic
chloride ; tbey aie alio ooluble in dilute ncid« and nlkalie*. being changed on aolu-
tioo into acid- and ftlka]l*albumin rwpectivelT unlcM Ihe ncid« and alkallea are
ciceetllnaly dilute. The utiiraiion with soliil Hodic chloride of ibcir aoluliuae Ut
dilutt aodic chloride, prccipitalm mnt mcinben of thia vlaM.
1. Olobulin {Criftiallin\.
If the cryMnlline len* be rubbed up wttJi Hne *and, extracted with water and
filtered, the Ultnitc will be fouixl to contain at leiut iliree pntleidn. Dii paaalng a
rurreiil of carlioniu anhydride a mjiiom precipitate nccura; thin ia globulin,
Tho iddilinu nf ililiile iinitic uid (o Iha DIItbI* fiocn lit* ilobHliTi. tpvet ■ pioripilala nf
•Ikali>altRiruin !' nu-t ih* fillralD (rom llila, irhaatod. (IVM rt funltnr pr»cip)Ul«. iluo u Mniin-
In iia ceiwral reactions Bhtbalinonrrmjioiida almoct exactly with the nest necn-
hei« of inb daaa (panwlooulin and ftbnnogcn). but ha* nn power In form la pro-
Biote the formation of fibrin in tloida ooatftlniDg the almve-mentioned bodin, and
IMMCwea Ibe following special featarea: 1. AccurdingUiLehnianii. iiHoiji-ctiiited.
BCUtfltl Kilutlona become cloudy on heating to 7Jt°L',.aiid arc coc^'tilnted at Kf C.
3. It ia readily prccifiiiatcd on the additum of alcohol. .Vccording In lltrjipe-
Stjlvt. it ia not precipilalcd on atluration with vhIig chloride, iffeniblinit vitellia
in tiib mpML
Arcertlnt 1o Kiihna'inrl ElehwaM* a globalin with nroporlica Identical with (Ik«« Jwt
CiTaa ma)' » prwiptlatMl IVoni diliil* wrnini by lh« niitioii* ■'1-litiaa of aeell* ai4d, Thla
lady iitlalwl by Wayl'tn Iw th« mma a* funelAlxUio (ribrliinplalllii), lb* lallnr dlUbrlUg
bmn H niily hj n imull ■■linlxlora of Abrtn-fAniMat.
2. Parselobulln {FibriHopUftin).
£iition. IIlooil-MTUin in diluted tenfiild wlih waitr.and a britk cniml of
aofaydriile ta paaMil through it. The lltvi funned doudliMMtooa becama
»
• 1tartM*aMa
> i^ili <l rk;
' llrllMlti' (III ..._.. ,
CiNMi d tB«atif«4U. mm., U
s.n).
_.„. „_., Uarfla, vtn. a. I.
1022
AffENDIX.
a floi-culcQl |>rL'dpitat«, wlikb i» Hnally i]ull« xraniilar. and but »a4ily lin lapv
nttd br ilewintatiun an<] flluation; it dioald M washed on ttiD fllur irich valat
4.-r>ii Mining uirbonic iidil,
It baa usually bocii itatod tliut pnnucl'iltultn nay hr separated frAm Mruni by
Mtufation with wxliu chlunde. But Haiuaiant^u' bu abowo Ihat thu bunly ta
part true, a cuu side ruble \ionloa of tin- gluhulia remnlaing auprvcipltat«d. The
Mpariilion mtiy. howcv«r. bp cnmplctoly t-flV-ct*^ by MturatJoo with maga«io «nl-
fihalc. Whon deli'Tininpcl by Ihii method the amount of MraglobulJa in aanai i*
Tory ooa«idmible,aiiiriuii ting, iiiwime auw ucMitdlnx to HammanttMi, ta a* moA
a* iM5 per txaU {^eck<tlIl^d r>ti 100 i-.c. of aenim). Tlie quantity aeema to mj ia
illflereDt aiiimnt''. tlio precipitation bring much morv onmplAlo in aMum (Vdib ot-
biood tban in thNt from the blo'id uf home*.
From il* KolutiuD iu (Iiliit« nudic (chloride. paraf[lobiilin mar b« pre«ipttalo«l b^
a uurreoc of i;arbuiiii: anhydride or tbe addition of HweJi'ny'y Hitu'r (Inu thaa I
|)ro inille) acetic acid. If ih<.' acid is tirong. the pr«cipltat«id |>ri>t«id b«cua«i
iminediatoly chnn/rd into artil-nlbnrnin (Cloaa II., 1 1. In pan mter. frt* lh)n
osyji«n, paraslubuliD ii insolubli-, but <m nhakinff with air at paanni; a eurninl at
osygea. aolutiuu readily luket )»lace : fcum this it may be r«precipiint«d by a ear
rent of carbonic anhydnile. I W-/(Vir{« alkalies diaiolvetliifl body without chann;
If, howftvcr, thnatrrnKtbnf ihr .likHti be rained orva to I per ocnt. the panglobuia
i* chanjcod into alkali -album in (Uliun II., i).
According to Kilhae and A. .'^cUmidt tbe solmioaH of thin body iu wat«t coa-
talning oxyecQ or in ven dilute iilkaliiM are not cuaj^ulaled on heitiiux. Tb*
Hodio chluriBo soluliona (In, hon-cTcr, coagulate whon hoaMd to S&^-'lT C..' aai
if theKubulancc itaelfbe luiipanded iti water and beaud to 70° C>it ta eoAgulaled.
Althoueb insoluble in alcuhol, Its soliilluua are iritb difficulty precipitated by thit
reueot
Paraiclobulin occura not only in blood-«f'runi. but it in al«o found in wbiMcor-
piBclea, in the ■irooia of red corpuictcH (to noinr QXtrnt at 1r«M), in coonnlivf
tiMue. the oaraea. aqueous liuuor. lymph, chyle, and icruuR Huida.
Fnr llie umiirrtnnf of flobulln in urtu**M EdleAan* and SanaUn.
8. Fibrlno(reO'
The geueral reafrlioiiii of this body ure ideBttoal witli iboie of paraslobalia.
The niOBi luatked diircronco bctneen tlie two Is tli« point at which coacuIatMB a(
their solution* tnkca plocn. Hmnmanitcn^ hn* sbown tbat AbrlnoMD In a l->'i per
c60t. Dolutiun of nodjf chloride ooa^^tilulu* at from &2''-Ay C, WDereu, m stated
above. paraKlobuUo (&br{ooplitat!n) coaKulalen fint at from 68°-70° C. Thi>,
however. Is disputed by A. Schmidt, who bolda thai the subataiMc coagulaliitK at
Hf-Ai* (s not nbrinoKon, but ii tort of noacant fibrin. There la abu a iHail«l
difi«renco in the precipilabilily of tliv bodio* by rodic chloride^ (8c« below.)
Otli«r ditTerencet between tbe two mav be tbua enumerated: In precipli^w
fibrogen by a current of carbonic anhydride tbe containing fluid must be moA
noore strongly diluted, and Ihc gas mu<t paM for a much longer time. The pre-
cipitate thus obtainea diflen from that of pnraflnbulin in tliat it forma a viseeei
dvpuait. adhering more cloaely to the sidea and iiutlom of Ihc oantaining Teaiel;
there la alao no Sooculent stage prerlousto the visoous precipitate.
Pibcinogen occur* in blood, chyle, seroua fluids, and in variwu traniuilatioaa
The rdlationi of fibriROKcn and paraglobnlin U> the fotinatloD of flbrin hare ttMa
diM!UBcd in the text, p. 49.
PrejMration.* Salted plaama, oblaiaed by cetitrifugatlxing blood whriw coaguls-
lloa la prevented by the addition of a cortain proportion of magumic nulpbala, W
mi xcd with an aiual rolunie of a aatuntud (85.8* per cent, al 14* C.)' aoiiUioa «f
iHiilic chloride: liic iSbrinogen la ihua precipitated, while the paraglobulln renaJat
In aoluttou. Tbe adhering pluamn may be removed by waihiug with a aolntfaiD
■ l-BUfar* Afahlr. M. itii >wn>. & tu : B4. Xtlll. {IS»\ s. Si.
• tfamnanHcD, ep. dL
• CmuralbUU C nud. Wl» I>hr(. inOb S. 3V. Alv> Artk r. kiln. Jlod.. M. ta. 8. <W.
• Vlmlinii'i Arvlilr. r.; U ' iV^ • llMala IiSkanAlnBiiiai rirhsadUonr. M. it tct
' a« ItKianikiMcn, > ' r.-j. tea. Sal,. l/'iiMta, Bar. Itl. tdL x. Untl, p. H. AI*o fflaM'i
AmhJr. IM-iU. il!)7* - iiXskM. IIBHL&UL
' Bgntlal*, Abu. CUIa I'u,^ >., vol. fll. p. m.
A
CKKMICAI. IIA3IS OT TIIK A.VIUAl. HUUY.
1023
.. Mtdic chloride, and ibe libriBCUMO floally parified by being 4MTeral tiiUM il{»-
MolTcd Id Kud Kprpuipiuttd lij •ocUc eliluride.
There I* no proof tlint thp whol^ of tho mibntivncr tlin>wn Hftwn by cnrtionlo
uibydriflo (ram diluted blno'l-itrniin {■ librin<i|>liuitit' ; uiilc<<fl wc kiinir Ihnt K true
(Clotiolin dermd nf fibrin opt Aitic pmportioa miiy bv ptopArvd from aitiiiii.' WevI'
consider* Ifakl lli«re !■ onlr ouc dobuliii In iteruiii, which he diBrMcteriies br
the nameof "eeniDi-Klibulln," and rfijnrdK dbrinopja^iin a* a nitxiure of thi*
bi>dy with « pnilion of Hbrin-ffrnirni. Wc Icnow for certain (■» p. 41) that the
wl>i>lf of the fibtinufilnKtic- (iKcipiiali'. uicd In cuuic ibe cuRiculalion of « flbrioo-
B«auui dutd, does uot euiei- luto ihe caiiipcnltion of the fibrin produced : we alao
inow thai inch a |ir«cipitAlo may lo*e itn Hbrinoplaetlc (lowert without any
narked elinnit'' in il« (ti-n^-rnl rcnciiin*. It would wem adviwibl*, ihttrefore. to
(peak of ihr •tctuMil pnjiluccd by cnrbuQii.- iiiihydride in dilute Mrum. or by utu-
niioB with nodic chloride in undiluted serum, aa globulin, and to diitloKuUli it
■■ ttbrloopla^tic i^lobulin when it U able to five rise to tlbrin. F^brinojten Biml-
larly otieht be upoken of a« Rbrinogonou* globulin. The name eryatallin, mthor
than globulin, raiftht then be given to the *ub«I«nce oblainud from the cry*txllin<'
Ivne.
I. MyoelB.
Iliie u Ibe HiibttniM-e which rorma th« chief pn>t«Kl conaticiient of dead, rigid
miHCk; it* general proEiertio* and node of preparation hare been already
dMcribed nl p. 103. In tlie moist eotidltloa It furrna ii gelutinoiLi. elaatic. clottetl
luas; dried, 11 \» Tcrjr brittle, slightly transparent, and elaittic. From itsAolulion
in »odi<- chloride it W wrecipitated. either by eltrvme dilutjoa, or by Mtliratlon
wiib (be solid Halt. When prcclpilaled by dilution und nubmitted to the pro-
longed uclion of wat«r mya«in lotiea fu property of bein;; soluble In Hulution* of
■odic cbtoride.* The aodlc chloride iioludoti. If expotea to a rising' louiperature.
becomes milky at M° C. and givof a Hocculent precipltnte at IW C. Thia pr«-
cipilate ia. howoTcr, no longer myoiin, for it in innoluble in a 10 per cent. *odic
chloride aoluUon, and doe* not, until after many d&ya' digeation, yield tyntonin
on treatmenl with hydrochloric acid (0.1 per ceol.), It i4. ' '
pratdd (tee Clan y.}.
In fact, voagulaled
Uyoein la eioeeuvely snliible in dilat« acid« and allcalieH. Advantace may be
takwi of ita eolubility in the fonner to extract it from mu«cln.* Hut if (he
rcagenia are at all concentrated, myosin undergote in the nclof itolution a radical
ehaoge, becoming ia tlie one caae acid-nlbumTo or ayDloQiu. in the other allcall-
albumln (Claw IT.].
UIm Alirin. it nui In toin* nuM Henemf oM hydrogee dioxiile. and oxidiM guaiaenm wiUi
fceiMlton of ■ blua enlor.
A. VltaUin.
Aa obtained from the rolk oftf^, of which It is the chief proteld eonatltueni.
vitellin is a while granular body, latoluble in water, bill very soluble in dilute
Midic chloride •nluticma ; it aurpaaaea myoain in thin re*jiBct, for the noluiioQ may
lie eaiily filtered, [in coiwulatioti teaiMniture la higher than that of mjoaiii,
lying, aeivnliug lo Weyl.'Twiween TO" C. and HO* C. Saturation with aolid audic
chloride giree no precipitate; In tlii* i¥«pecC it dlflera Ctom moat olbee DManbera
of thi* cIm*. In _yn]k of egg rilellin i« nlwaya aawKtalod with, and pruliably
txiati ID oombination with, the peculiar complex body lecithin,
Uanl*. aB<l aller hiia lloppw-Sayler, bata abown UuU vMallln balorfi ih« traatinent ra^Mte
III free il rmin larithia pamema* ppDpartie* itntto diflbranl frem otlior pmUliLi.
A theory haa boeo adranoed that vitellio ia rMlljr a complex body like bcmo-
C* ibin, and on treatment with olcohi^ aplita np into eawubited proteld aad
ithin. When well puriSed It cenU^n* 0.76 per cent, auiphur, but no phoa-
pboroa. Dilute aci<lii or alkallee readily conTerl it lu ita uneot^^lated form into
nmeaabeiof Claw II.
• Kakn>>ao<IKI«li<rald.loc. eti. i tnl vlt
■ Wf7l /r^IBwIir. I Pbrilol. CUrm. n't 1. ni7'>. E. TT.
' [laiilkinkjr. Zi-ntehr t pk; 4at. Cbcm., M. t. a«li, a IM.
•(*.*b.
1024 Al'i'ESIIlX.
Fr?Tny ami Val^ncieimea' have described a aeriea of proleMa. viz., irhtbin. irhlhtdiD.ftr,
derived from fjah and ampliibia. Tbey appear M be either idemlcal irilh, or ctowljillicil
to, vJteLlin.
PrtparaUon. Yollc of pgg is treated nith succeagtve i^uantiliea of ether as long
88 this e^tlractB any yellow coloring matter; the residue isdiaeolved id modtratelT
stroag (10 per cent.) sodic chloride solution, and filtered. The filtrate on falling
into a lar^e escees of water la precipitated. In this xtate it is mixed with lecithin
and nucleiD, and in order to free il from these it was usuallj treated with slcoli''!.
Tbia, as above stated, entirety changes the rilellin into tt coagulated form. It
Beems probable that the separation of vitellin from the other bodies with which it
ie mixed in the yolk of egg may be effected by precipitating the aodic chloiiile
aoluiion by the addition ot excess of water ; the precipitate is then rediE>eolved in
10 per cent, solution of sodic chloride, and tbe process repeated as rapidlr as pos-
sible.
6. Olobln.
GLobin. staled by Prejer' to be the proleid reaidue of the complex body hvinoglobiD \itt
p. 457), ought probably to bo congidered aa an otillyiog member of this itIbm. It ia.howeKr.
not readily soluble eillier in dilute acide or sodic chloride aolutiona. It is ni<l lo be abto-
lutely free from n»li.
Class IV. FUmn.
Insoluble in water and dilute sodic chloride aolations; soluble, with difficnlu,
in dilute acids and alkalies, and more concentrated neutral saline Bolutions.
Fibrin, as ordinarily obtained, exhibits a filamentous structure, the component
threads possessing an elasticity much greater than that of any other known solid
proteid.
If nlliiwed to form gradually in large maesee, the fitamentflua Etructure is not si nitin-
able, and H rcwniMes in Ihi^ foriti pure India-rubber. Buch lumps of flbrin are fiplJf
of being p|dit iti uiiy direi'lion. aiid do definite arrangemeiil of parallel bundles of ]ibrH<^Ti
be made out.
At ordinary temperature fibrin is insoluble in water, being dissolved on I vat tety
high temperatures, and then undergoing a complete change in its characters, la
hydrtjcbioric solutions of 1-5 per cent, fibrin swells up and becomes transparent.
but is not dii'solvcd.' In this condition the mere removal of the acid by an excrta
of water, neutralization , or tbe addition of some salt, causes a return to the origi-
nal stRtc. If, however, tbe acid beallovred to act for many days at ordinary fem-
peraturci", or lor a few hours at 40°-00° C, solution takes place, and the reaulting
proteid i^ syntonin. In dilute alkalies and ammonia, fibrin is much more readil.r
soluble, thiiufrh in thiM cHse also ihe solution is greatly aided by wanning : the
resulting fluid (oniuiiiH no longer fibrin, but Bikali-alDumin. This property i:*
not disliniilv cliiiraeterihtic of fibrin, allhtiugh it diesolvei) peibaps miTc readily
in both dilute aciiis iind iilkuliea than do coagulated proieids. None of tfae>e
solutions cati be toHgulated on heating, wbiih is intelligible when it is remem-
bered that lliey no longer contain fibrin, but either acid- or alkali-album in. Id
flddhioii to tbe above, fibrin is soluble, though with difficulty and only after x
con.-<ideri)blc time, in 10 per cent, solutions of sodic chloride, potassic nitrate, or
sodic jiulpliati-. the solution being olt«n accompanied by putrefactive ehanp'.
TbcfC ?oluii<iiis iiiiiybe coBgulateil by a temperature of eO'C, and are precipitaitd
by dilution niib water ur saturation with solid aodic chloride; in fact, by the
action of ibe reutrul saline solutions tbe fibrin has become converted into a body
exiecdiiiKly like uiyocin or globulin.'
(Ill ii;iiiti<iii <if fibrin a residue or inorganic matter is always obtained ; iti^.
hoivf viT, c<in-.idefed timt Hiilpliur is the oiily one of these elements which entfrs
I'weiitiHlly iiilii its lonii or-itinn. In other respects fibrin corresponds entirely m
pener:il compcisilion "ilh tbe other proteids.
I ri.uic'i. li^'iid., T. \.s'i\iLi. yy. Ir^:>, .'li'i.
-' Vvil <.|i .ii.,S.7i, > Die Bluitryftalle (I'TIK 5. I'lV
• r. .ii.|.:. 1 1- -I.IM11..U in^LV. himi-n-t, inkt [.lnei> if ihi- fibrtn, a.« 1^ frenner.ilj- ll>e en*. .imiaitsV.v
" i.nui:,r, 1 1.1. n.n.t., T, Iwis. 1 1-7 II. [.. iSi.
\
cnsuicAL Basis uf tub xsiual jjout. 102S
8u«|it>iute(l in water and kotil«d (o TO" 0„ il toum iU olacticitf luid beco4DM
Opaqiip; it i* tbcn indinlinjpiUhnblv ftom (>tli«r COHgnlatcil pmloida.
A pamlur pn>p*rt]r or (hi* Imdr wniiiia yH bi Ixi iiiiintii>ni><l. tit.. il> (nwtr at ilMom*
, peung hjdn>SDii dtpxtda. Pi«u of (Ibrlu pUiwi In Ihii tlui-l. though themnWu uiular-
Kins *a<b>angc> tocoi braiina cnrciwl irilh bubbln itt oxygen: and puaiacum ii turnvd bliM
(twin i» pnMDM or hydntvti iKoiMa ur aaoalaad torpantlaa.
Pmaration. By vignmiuly utimng blood wiih ii Immile of litigm and thoi
wuhinv with wulcr until it U iiuili! wiiil«. IT requirMl uerfeolly pur« and colo^
lew It •faould be preimred from pla-cua free from curpiudw. If the blood, before
ahrrin;. be dllaird witli an ojiml biiik of ivattr. the culiiMyiuoDt wuliing of the
Abrin i» much fAoititntrd, and ii mnj reiidily be obtitincd ijulii- wbile. Adj* Hdb«-
fcnt fat» may be rcniorrd by clhrr.
Wbca globulio, luyuain. and llbriii are iwin|iared «wh with Uic other, il nlll be
■ecu thai ibey form A Mrlca in which myoela Is iatermedialc iietween globulin
Bod fibrin. Globulin ia «xceMivel; M)la&l« in even the mint dilute acid* Hud
alkitliea: Rbrin ia ■ImoKi in«-ilubIo in tbne; while myoain. thi>ui;b luoru wiluble
tban fibrin, ia Ima naluMu than globulin. Qlobulin again diaaolTW with the
CreBteal eoae in » rerv dilute wluiioo of eodic chloride. Myosin, on the other
MDil, dlnolvea wltli difficulty ; it Is much more Bolnble in alO per c«nt. than in
a I percent, dilution nf fodic chloride; and ctoo in a 10 per cent, tolulion tin-
Dijmiii eiD hardly be said to be diaudveil, ao vucid ia the rraultins fluid and witli
awh diSculty doM it (liter, fibrin again diaaulrts with grrat difflculiy Hud very
■lowly la even a lu per cent. Huluiinn of nudic chloride, and in a I jicr moU tolu>
tlon it la practically insoluble. Wh«n it iit rcmrmbercd that libna and niyoain
ar*. both of them, the nvulu of criii){iilntinn, their sirailarily in inlelli|nbl«,
Myoain ia in fact a wiiuewhal more iiiluble form of fibrin, di-pmited not in threada
or ttliirnenCa but in clunipa and niaaieB.
V Cli«» V. Owsrii/aiyrf Pmltid*.
Thew aro inaolublo in wntrr. dilute ncida and alkaliea. and neutnJ taline mIu-
tioAH of all sirenetha. In fact, they are really aoluble only in Atroas acids and
■Lroafc alkalit*, though prnloni^ed action of ttivn dilute nciiU and dkalles will
eflbct aoene eolution. especiHllv ut bi^h tern peratu tea. MarinK aolntioii in atrong
aclda a&d alkalltft a detttruotive decoinjioaitloii cakea ptaee, but >ome amount ^
acid- or alkali-albu^iia is nlwayi produced.
Vary little i* kDonm of the cbrmiinl i-hnractrrinira of ibis claw. Tliey ars
produced by beatiug loTCi'C.. nolution* of rjtg- or aeiym'albumin. globiiUna, aiia-
pended in water or dfiaalred in aalliie M>Iutluua ; by boiling for n ihort tiiuo fllwin
■uapendcd in water or dlMolved in saline aolutlona. or precijiitated acid- and
alkali ■albumin ■utpciided in wster. Tbe^ are readily converted at the (cnipera-
turc of llir body into pi-plonea. by the action of ganlric juice ia ao acid, or oi pao-
cr«alic juice !□ an alkaline medium.
All [iroceldN in itolutioas are precipitated by an cicea* of atrong alcohol. If the
prtcipitant be rapidly remoTcd they arc aenin «ohible in water, but If ili« nrecipi-
tate<l pr<itwda are >ubjrct<vl for aomo time to tbr action of the alcohol tuey are,
with the e^cceptiun of pcfitoun, coagulati^il and lu*« Ibcir wilubility. It appears,
however, thui t]ie urotctdii conlalnedlii th« Hietiruoe'gtaina uf plant* are excecd-
lugly rcfiittiint to this coagulating action of alcohol,'
It •a«aM warmly nerMMty M point «ui (li* didiEioliaii Id ilia u«taf lh« word "ooajrala-
Iku " ai applied in lilood- ce mtualo-iiliunui on ili« una luiid ami tu Ui« KUoa of bnu and
almliol npou protoida oa the otiicr. The itiflMwnoc ia olmoiu when it ia rauMwiWwd thai
Ib tlin flni (««a the eoacuUliuu ImJ* lo (ko fannatMU of Abriu (CIbm iv.t.ar ■ny«aia iCia*
iii.l.aii4 Uist UioM IhhIIw ni*y then fuMkor iw eoagulslcil liy iMatar akotwl ■• iliirfHwl
abev*.
Ci-Aat) Vt. P^toata.
Vtry aoluble in water, and not i>re«ipltal«<l fVutn tfaelr aqneou* aottitioiM by tlie
addition of acid* 'it alkaltea, or uy boiling. Iiuoluble in akohol, they are pre-
cipitated wUh didlrully by thi» reagent, and are ancbaaged iu the procwa; Uie;
i*»»\\i»^Kmm,t4 nijrdol., mL ML (i ]
06
i
1026
APFKNUIX.
difler from nil oiher |>n>lcid* in not Lriiij; CM^Uted by pralaagcd «9
skobul. Tbof are not precii>Uuivil by luprlc anlpbMv. ftme chlari4*, '
:ij>ttuivil tiy L'upric aniptiMv. Rnte chiarul*, or, n
in th« liMtencei to be iiieniionti] jiteMntly, bv {K)UHic ferrocyuniile. and .
acid. In thma point* tbr.v dilfrr fnat mo«l oilier proieids. On lb« oUicr I
prei-i|>itiitii>n i* caiwcd bv rblorino, iodino, tnnnin, mercuric chloride, uitrkharf
nipicurv aiid vilvn', and Wli uueiiiUs of lend ; mIho by bilo-Bdili in an Mcid aola-
lloii. Id c'lmmou witli ail pruleids. Ibeue bodiai poMsna •pecifio In-fo rolatorf
fMwer over polnriRcd light; but tbej ditTcr from all ollitr proteidii in itie fact Uiat
boilinK priKlui-n no rhanKc in lh<i ■nuiunt of rnlntion.
A Kilutitiu uf |)e|'i(iim, mixed iriib a alron); wiluiinn of cauatic potaab, gimkoa
the addition ><f a mrrr Inifr of (.-uprlc eulplinl^. a /liat color. An rxceM of tb«
cnprir uitl tiri» :i viold enlor. whtch doeji^rm in titic on boitine, in fact th» ordl-
niirj proteicrr«tcti(>ii. Other pmlrid* hiin|<ty givn (h«> viol«( color. But tbei
cburucltTiiitlL- feature of peptone* irt their tcliitivrly Krtat diflbailiillty, m i
which they aloue. ul all the pruteids. may bu aaid to poaanv. aince all otiMC J
of prol«4ds pass through membmiiea irith th« grealnt dUHctilly, If at all.
Tho diDHiiitrilitT nf peiilouM la, bowevor. ■baotaLsly amall •« cnm|ia«Td <ttlh Uial nf •.,.
lalliao tiodla**iichta*niftFoli1i)rid«; in llwlM>1ution«of pvpUmw may be fr<wl tmtm ^ui,
dlalyiti. a prwwH euiployal ni llialr j<n>panitloii.
Notwltbntandiiig their ]>rubabl« furaiaiioii in targe ()unntiti«* in ibe tfawtiacfa and
IntMlinc. to judge from ibe reaulla of aitiAclal ufgoilloti. a very Mnall oaantili
only caa be found in the euiitonta of theao organ*. Ttiry are probably i^Mofh
iiK (nun OH rririned. Another point of intvrcal in llirir reconvrn>>on iotii o|'
fornm of prolf ids, niiioe ihii must oei'Ur to n Kreat cxtQiit in the body. We \
liowever, nit yet ignorant of ihe manner in whlcb this reverse cbmii.'!- i> I'lTx,
fyodirlion. All proteida, wiUi the ext«f>lioD of lardaoeiii {-' -
poptonea (and other prodactaj on lrraiiii«nt irith acid giwtriv i ji|
creatic juito uhhiI readily at the Icmpcmliiie of tbe human body. IVptonnl
likewbe pniducod, in die ubaunceuf pe[Hin and trypain, by tbe action <•( •'"
and inodotntely strong acids ii[ ni«diuin temperatures. hIm by the acUoo of dli
water at high tcmneralures under pr«wiure. Fur Tarioua meUwda of prv)u
iin, see Maly,* Adnnikiewicc.' Henotnger,' and i^kelbario^*
peptfint
Iba vUuD fll
Ii ai-iiean poaible (o renlilain ordinary cdM^iilalile proUU* tnmi peptonM by
«lllier pruluQctil lioalint Uf I4(t°-1!0* C. ot of'Miy^tMig agoiiu.*
Ko diflirreooe in percrntage composilinn bctweeo p^itoaeaand tho proUM tna
whicb they are foruied lini. at pren-ni, been deflniiely etitablWbed,
We have u*^d the pbrate " [leploneA" in the ptaral nunber becauae ir* liaTe
rcitaoii l[> think llmt more than one kind of oeptono exwla. MeiuneT* UMcriM
ilirce peptonni. Qniniog them ropectiitely A- It- and C-pApUne. He di»iioguiihri
tlicm M follow! : A-|>ept(ii)e is precipitated from itaatpieoaa aolutioni by eoficM.
tinted nitric acid, and alao by [lolaMic ferrocyanide in tbe prcwuice of even wtak
aeello add. B-jieptoae ia not precipitated by concentrattd nitric ai-ld. nor wIU
lioUMtc ftrrocyanide (pre a precipitate iinleM a oamidcrable •iniintiiy of airotif
ac«4ic acid be added at the aanie time. C-neptooe la precipiiuted neillice by uitHe
acid nor by pol4nlc ferrocyanide and acetic acid, wbatcTttr l>e ibe strength of iho
acetic acid, tn place, however, of apeakingnf all ihraa m pei>tonea. It ia better ■•
consider C-pei>tone aa tho only real peptone, and Ibe A- ana ll peptone* ai> not
peptone* at all. Nevertbelwa'we have reason, from the reaearchea of Kflbae, Ur
apeak of more ihnn one peptone, vi^.. of u bemipeplone wbicb ia oapabl* BMiiT
ue AciioD of tryjxiln of being converted into loucin and tyrodo. and nf Uk aiili-
|tepl<>ne which reiiMii micb n de«i>mp<i»ition. Tbe natne ■oti))rptiine is riven li>
the latter on account of ibiii rerintance wblch It ulfers toward Irytuiu ; thin uaar
beat i peptone, given lo the former, aignlflea that this peptooe is (be twin or 0N>
tvlallve half of antipeplone.
We have iwen (p. 311) that when any proteid i> digtated wilb popaln, what n
1 pai,p.r -
• I^IbS-: ".i." M
•Iwln.s.. Ivt l^piinvn. I^rit. ISJN,
• I^*j^i'i Ai'ti , 1^1 iin .:•.•■.-(. t', I".'".
> t(«ii>liic<r. lor. n\. ; IlLiOnrl.itf. Zgluoli. t phydal. Ckam., fid. It. 11*1*1. 9. aW: rtkdtalBB.
tor. ell.
• ZrttMb (. rat. Uta. B4*.m.,iW.,a..alL.a.ilv
OBBUIOAL BABIB OF THE ANIUJIL BODY.
102T
I
e
nar pivllinlnnrilir call n by-product makwtta appenrnuee. TbU by-product which
bu mniiy riiM-tnblancw lo acid-albumin or ■vitwitn, appMrin^ im a neulrallutlon
pKrifiilah- wilublp in dilute luldii ami atkalim but inanliiblv tn dinlillcd water, ia
fanrrally <pK»kcn of no |«rapepto«i(>, Aoctirdiuc to Flakier' thi> ii«utraIiulioii
precipilalf in ei|i«c'iatly abuntunt if tlic i>epslu be pr«vlotulv miHliG«d by npo-
aur* to B ieinp«ratur« of Vi"" to W C. The petxJn thua modified la apobeo of bv
Pinkler aa " tonppnaln. ' Many autlmn rcmrd pa»|H<pt»n«. ayatonin. Kod aold-
alltumin iiK hfiag ihc nHmo thine. MpiwtiiT. howoT«r, inive tb« naiDe panpfptone
to B buily. irhit-h Deed out tuiA probalilv doM not make lla Kppearanc* during
BonaKl nuiural dlgmlan or during iirlilk-ial dlgeatlon with a thoroughly aclivo
Mpttto. bul wbii'b k« formed nhcii protciiU ari^ aubj«ct«(l to the HctloR of weak
nydrocbloric acid. ?iih<-r alone or in i.i>ninitnv wllh un impcrfrr.Cly .vUng ])ep«tD,
and which in cirlaio rhiiniclcn i« quilt? iiistilti-t rrum ordinury ■viilimin or acid-
albumin, [u diBliii[;uiMhing ft-aiurc that it caiinoi be chaii|^ intu peptone by
thi' action of eren the nxMl <.>iiortfeiii.' pvptin. though it i* readily «u ci>DT«rtMl
under the Influence of tryiwin ; othtrwlsr it vrry rlo*<-ly MHunblea ayiitonlD. VV'e
have here aii iedication that the niintilc chnnii.'U'ni br which we hiavo described
acid-albumin may be borne by budics liariiijr inaiked ailTercncM from each other.
The r«M«rclia of KQbae* biive thrown an lmi>onaat lleht ou thoie iliffemiocik
The fkutdamenlal notion of Kllbne'« view b that an ordinary native albuiuln at
Sbrin oonlAin* within itHcIf iwo rfoidiim, which he calU rrnpectivoly an aatl-
r«iilae and a homi'miduc. The renull of either mplic or Irvptir digution ia Ut
wfiit ap lli« albumin or llbrin. mid to prodooe on the pwrt oftlie unli-rc«idue anti-
peptone, and on the part of the hemi-realdne faemipeptone, the latter being di«-
tlagnlahed from the former by ita bcipg auMvptible of fbrther change by trypUc
dinMioti into leudn. tyrwn. etc. AntipepboiM lemaiiw aa antipeplone oven
wneo placed under the action of the moal powerful trypatn. providnd pntrofactivt
clianm do not In terrene.
Benra the ala^ of peptone (wbethcr unti- or liemi-l ia reached, there U no
iaicriBedlate atage correspond! nj; In the formation of Kyntonin, In both nonual
Miplic and trvpiic diKoticiti antipeptone in prccMlnl by an aotJ-albiimoMt, and
hcnitpcptnnc ny a hi-mi-iillmm'ike. Of tboM' the anti-albumoae iicloMly related
to nyntonin, aitd liiu hithcrt^i been regardtO a* ayulunin. The bemi-albumoa*
liM not been ao fre<iuenlly ubaerv^ ; it waa. however, boUled by Meiauifr; il ia
appar«nllr the body called by him A-peploiie. It pomeawa aeveral peculiar IW
tureii. If its solutions are hemed ih<.'y pariinlly coagulate at about 00*'-43°C.;
the precipitate ia soluble at abiiut TO" ('. and is re -precipitated ^ the temperature
■gain (alls. It s1«> yield* a precipitate with nitric acid and putii»ic fcrnicyanide
and this aliM) U iiuliibk- ai the hlgliM leuiperntiire. r«-preciiiitatiug on cooline- In
thMe reajxcis it cliwely rc^embtoa a proleid body obterriMi hv Bonce-.Tones in the
orin* of miromalnciB.' It approaehae myown in beinx rcailily koIuIiIo in a 10 jier
cent- aolution of mdic chloride.
If. hiiwerer, nibumiu bo dicealed wllh InaulSclcnl or with loiiierfectly active
Eiptin. or Hiinply with dilute Hydrochloric acid at W C, aull-albuiDoee b not
rm«<l. bill in iU place a body makni it* appearanco which KUhiM calls antl-
■Ibnmai*.' Its i-.hBrac.teriKtic properly in thiit it cannot bo oonvertad by pevtic
difcottion into peptone, ihnueh it can be to chiinged by iryptic digeation. 11 b
in fact the purapeptoiie of .Melwoer.
Il WMf pfvliapa ne adviuiblo, now that Meissner's pampeptone ia cleared up, lo
rcMW* Ui« name parapcptone for the initiiil producu nt both peptic and Iryptic
diK«*tion. and lo apaak of anti-albumoie and iienii-alburn'»r a* bcinx '^'^^h parm-
peptones. But in thia seuae perspcpione will be uu iotenuedlBio and not a col-
lateral product of dlgeetlon.
MeiMiier also desoribed a particularly jnaoluble form of hia parspeptone ai
i^jnpcptone, and another intermrdiHie pniiluet an nietapcfitaDe ; but further invas-
tuetiOD of both Ihiw; licidle*. an wdl a.i hi* llpeptuoe. )■ noceauuy. Under the
inilaence ofdilute bydr-x^bloric ncid, Buti-ulbitm»l« becomes chnoged iuto a body
irbkh Kfibnecallii auii-altiunild.and »lilch leemi idtnliciil nlih the very liiaolnble
inotrid described by SchiiUenberger as " fauDipnitein," and probably witb Mebe-
' PtuirVi .trcblT. ilT (ISWi. B i»
■ itelji ■ ihi-n Miminl al itaae liu H ytt b«a PiiMlifcul
IUI<lill>m. Ikl. I. Ilffi t, tin.
• aa albduiai* noM not be ranAniiKlRl wlUi an allmailnaie
VarluudL d. SuutWaL-OML V«fDla,
1028
APPKXDIX.
n«r'« dnpeptnue. Tlie Mm* bodr It produoed lU oac« in eompknv with i,
b«1oDgine lo Uio beini-grotip by the action of S to 5 por cent. siil|>burte i
native nlDdnim or 6lirJn. Tlio following teblm 'linw the relaiinDo mkI gamia of
tho bodicm vro hare jiwt dcacHbol. The teverat praducta (aatijwfiUiaA, vtc-) art
Kivcn ia duplicaie. oa the hy|Kith««b(wlikl), thoitj^b not proved, la |>n>babl«) that
tbe ubangM of dige»ili>ii are casenUally hydrolytic chnii|^.' atcompanM bf a
deduplication; thai, jiut a» a moloculvof o'lorch ■pliln up into at iMat iwa ttul^
cu1o« of dulrvae, or a* a Dioltculc of can«-augar «plil>> up tolo a molMala «r
dcatromj and a molecule nf lerulone, to a niolMuto or antialbuiunae. far Inaiuot,
■pliu up into two molevulen o{ aoUpepton«, and to oo. But tli« «bol« •ch«B* tt
or courae only prOTiaional.
i
DRCOMroniTiox or Pkorsioa bt Diosnios.
Atbonitn.
» .
i I Aniipopton*.
AiitlalliUBiiMaL HsmUlbuiiioM^
Antlp»|itan*. Swnipqitoiio. ll*iiitp*ptMia.
LtudD. Tytvia, L«ai^ii. lyroalB,
•l4k ate.
DBOnWPWITIOK BX AclDB.
1.
By OM fM- Mat. UCl at 4U" a
AlbHiBin.
'I
AsilalbuiiMtCi
I
AaUaHmnild.
HcnialbnmM^.
Jleniipepcon*.
UcmipoptoM.
3.
By IS par cant. BfiiU at 106° a
Album la.
Antlalbuniid.
HaaiaJbvDKMfL
ll*nilp»pta»«.
Ltuoln, Tjivaiat «(c
llaiuip>p*n»>]
I
LvurlB, Tjnnutl
Clik VII. Lardacri't, or Iht to-called AmfloU £MM<mw.
Tbe itubMaiii-« to nbiL-b th« above oaxue la applied U fbund u a patlKdMlal
deposit ill th*> spleen aiid liver, alao in numerouH otber or^na, aueh aa tbe bim-
Tesaela, kiilneys, lung*, vie.
It ia inMtut>lc in water, dilute ni'idn. nnd nlknlj*, and neutral •aliof aoIaiioBa
In oentotiroal oompoailiun it ia almud idculii;al iriib otber protdd*,' vtf. :
0.and8.
M.4
B.
7.0
N.
C.
The Bulpbut iu thia bodjr ezbta In the onldlwd atate, for boHine with cauiik
potaab glv<a no mljihide ol the alkali. Tbe above reaulta of analyua wuuld Uad
at ORG* 10 tbe raokins of Urdaci^in ax a proteid, iinil Ibta i* *tr(>ng1;r ftup|<orted In
Other Cuia. Dtrons ny(lnKbli>ric a^iid c«iivvrt« ii into acid -albumin, and wutir
alkalies Into alkali-alMinin. On tbe other hnrid. it cxhibila the followiojrmajM
dilTbreucea from other i>rot«ida : It wholly rtoiitB the acUoo of ordinary dicwb'*
Ituida ; it i* colored red, not yellow, by lo>lf ne, and *IoIet or pure blue by the JniM
) IlKinliiatr. to*. dL p. Ml
• V, AshmldL ADii. il. Cb«ni, a. Phano., Ud. ex. S. :U«. aad PtMxdob a. Xelrnt*. VtMbmt
AkMt. iu. nl. 8. W.
CHBHICAL BASIS OF TOK ANIMAL BODY.
1029
»
I
i
*
Mtioa of iodiiw and mlphnrir ■cid. Vcam tboo lant rcMli'ina it his dNlvcil one
of ItsBUBCa, "amjioid. ' tliouKh (hi* ii eviilevlly badly cbMen; fof bMooly does
it dlfl!^ from tti« HUieh gn^up la cum pu«i lion, bui by ni> nieajia c«n It b« ouii verted
iolo f>ii)csr: (bfci latter io one of ibi- iTiirUl tnU fnr n Inir mfitibcr of th« carliD-
hydrate |[rou|<. According to llriu^hl ' nnd t^^rnil,' lUiiHn-viDlct luirtLyl-aoiliuJ
coloni lardai-cnun tiiou« ruiy red, but lound liauie blue.
Tba sijon nwBiiMiiiiiI •Invv, ■« ti*iiig pro<liu«l by Iodine and mlplmrlc add. an mneh
dcantr aad lirifbter ■'heo Iho reagents arc applldl to Ih* puiilloi) Urrlireiii. Whmi Iba
ivsfBnU aia ■pplird l» (ha Finds anbttann) in iu iwirinal jnaiiioii iu (K* iiiauM, tlia oolun
oblalDad an aJwayi dark aud ihrty-lnoking-
Puriflcd lardaoein b readily loluble in moderatelv dilute ammonia, aod citn, by
•THioratina, be obtained from ililn Nnhitloti iti the form of loiicb. gotatiiiou' HaL'rw
and tompa; in thie fonn iljcive* focblv rracli una only wilh iodjiio. K the cxccaa
of ammonia ti nppllcd, the «olution becotnm ncutml, and u precipitated by
dilute addN.
PrrfMtraiioH. Tlie gland or olher ibaiie coiilaining Ihia body is cut D|i into
aniBll pi«cm, anil tw iituub oa poMible of ibp iurroiiiidinK tissue rt-mnvcd. Tho
pifvm are tbrn cktractMi ■cverni time* wltli wjiIvt and dilute alcohol, and if not
ibtM n-ndricd col<irl<i«, are rcpralt-dly Imilcil wltb alcohol oootnioiDg hydrochloric
ftctd. Thu rcaldue after thin operntion im dlgoaed ni W C„ with good artiflclal
gaairtc Juice in eiceaa. Everyihing, except tnrdaccin and xmAir qiiantitin of
mucin, nurl«in. kfi:ain, lotn'iber with uimi- pcttion of the olastic tissue, will thua
be diutolrcd and n-movcd.^ from Ihc liiilcr iinpuriliia it may ho separated by
d«cantatiun of ibe liiii'ly iiuntlvred Bubataace.
Tho chief products of Ihc decomposition of pi^lcld* are ammonia, car^nii:
anhydride, IniRtn, and lyiusin. t^ornrnl other bodiw, for the most part, like
levcto, amidalfd ucida, nucb us aspartic acid, glutamic aaid, etc.. have also been
obtained; alto by trjptic diKMlioii, hypaxaathin, and perhaps xanibin. Bui
orcabaa never yet W^n derived hy direct decoin position from proieid material, lUe
tWtenents to this olfert having been luiscd on errors. In spite of numenrai
rMiarcbea. we cannot at present state detlnitcly wliat is the real constitatioD of
a pmlrid, or in what manner th«n several r«alduniareoonlain<d in the undecotn-
puMMl Rututance, U is unnecemary In give here any of the formnlic, ncnriy all
emplri«al, which have been made loreprewot a proteld : they all f^iiv with e<|ual
exnclilude the percontofe compodtioD, but beyond ibis lliey are un trust worthy.
Uf tlie various attempts which nave beon made to nasign in nmlcids some deRniUi
molecular atruclure, none appt-'ni, at the present xtaffe of inlortnaliou, suffideiilly
reliable for general acceptance.
Among ibe most clsbomle Isluni in tbii dirKiloo may b« manUonsd (bw oJ UlMlwals
and HaMnoaa. In their Ant piitilKatian,* «Urtiug from the pnetal simikritr of lli» pr«-
■liMlB of daeompuailloD of tlio jirulaids aud cartwIiydraMa, (kay triail to •stablish ■ iMInlto
relstioa bslwcan Iha t«o clsaot of bodies. In this tbey were not aorneiafal. and ia tbnr
Mcond mmnli* Ui*y mine ki the oouelution Ihal the oartwliydrstea lake do part ia the
foriDslloD uf the prolflldL
OUier experinienM In the samo dlreelioo hare been laaJe by SthUtseabarxer.* Re ahows
lliat albDrain cso b« dscoinpoMil Into tartunle aahydriils and anunoaia. sad that lbs rslle
of tboe two i* Ike tmau> as tboegk nrea hwl beea the bodr oa whieb ha opceatad. Kfam
this h« ooiioladss Ibat *' tlis molenile of alfaum'n onnlsins tho |[muj>ii>ji tf urea and rein*-
■enls a cgauplsx urclde." Ia lik* aeaiad publloatlou' lio oouAnn* hli pnvivna rtaulls. «t«<nc
that the aaiaiaiiia> ofboiiie sahydride, and oxallo aeidi nrodwed by lbs dBaampowtioa of
pratald*, ars *o oonnoolsil i|uanliiativsly ■> la Iw eabsbis of darivatioa from vaiyias ptMwr-
tlo«* of ntoa and oaamidc Ha al» obtained from tba deoaupeuDon of pMtald* a aiuo
K»WB rsssdiss whioh oould he ftirmulsled at gi'I&K rise to all ttis sni>>Uie-l vidi ■»■! otlior
las qiokoB of atBVOL Tlius, suconhnc to hiia, ■Ibuiinu, built ugi as ■ minplsx uialdo,
deoouipws into soinioiiiB,imibanie.M(Bliej sad aeatle aelds, aad Ibl* altrDgeuoiM body { this
UMiboii %\ifnm in lb* oihor prodaetsof d««««np«altiea,*
I Wtm. ninl WorbcnHlif,. Xo. U, R Til
a (Voi^ Hsnd.. T liii. <■*»). a. law.
• XnhD* and KadMir. VIntiow's Arahli. Dd. ixxllL <IMU 1. «.
• Ana. d. dKOL tL fhinii.. Bd <IU. S SM. ■ lUO.. M. dxis. a IW.
• Geaipla»ltsBilufcT.sO|ISJU.p.Ba. Bull.4a1s8uo.CMn. axlll., 1«1, IM:.lU,>KMf,«Kulr.
Setiu.
■ UDmMsHsBdos.T. laxsl. r IKK UulL da Isteo. CMm. xst. liT
• 8se alift aelMMabHivf . Ann. do cksOL 01 <Se rays., T. KM. (IMI>> ft. atb
1080
APPIHDI]
Botubl* in HC1 0.1 pta cent, in ihe cold
It wilt b<- notlcfld tbKt in the Kon«r&) description or tli« rariooa prolwb i
tinclive rOAcliou tar Mch oould not l>c k<wb. but that varying •otnmlitW '
tbeohlsf m««nHat ourdUpoMl for iliniinguiahiiiK ih«tD. Tbc^ tna^ t>« i
Moofdlag to their Mlublliti«H In tlic fullowlog ubaW forui :
Solublt in iluHlled \etilfT —
Aqu«')U8 solution not coRgolated on boiling . , l\ptonf*.
Aqucoiu wolution con^lntH on boiling .... AAmtidma.
iMOtuhle I'ji dUtillrd water ;
Soluble in NaCl volution 1 per cent OtohvHa*.
Insoluble "
jAcH- atid J/faff-
Iiwoluble in IICI 0.) per cont. in tlio cold, bnt Mlubia 1 rl_;_
«l 60" a . . . . . . . . ,1 "*"'^
laaoluble lu UCI 0.1 \wr cent. &l IIO'C; aoluble in rtrong acldft.
Soluble in gMtric juice Qtag^ibOtdi
losolnble " " lianiaceiA.
'fliudl K clflMifirAtiOR i* however obviously a wholly artificial one. uiiehil
ilMp4uo' puriMiKM. bill in no wny iiluMntiniK tbe natiitnl rplalioiMof ibi? u-i
memben. Nor i* n diviaton into " native" and "derived " pmti-idii mui'li n
unlhtacliiiy. It ia true that we mnr ihun put tofetlter ■crum- nniI ciri;' nlbnu
with vitellin. myn«tn. nnd fibrin, oti tlie one hand; and iieptonra, coagulated |
telda, and arid- with :ill!«liKlbuniin, on t)ie oilier. Bnl in what light are we U
eoneider casein, icciiie tliat thoiiich » nntural product, it biu to many mffcnblaaMi
to alkali-albaminf Moreovrr ibi-- >yiitem of clKMification moat be luwIeHwliitk
would plnoeflbrlnoplutlcgloltiitiii'arid fibrinogen in the luune claw ■■ fibrin, lad
yet we can hardly gpeak oteitlii'r of the two fomier bodies a> derieetl protelda If
the view be true that wbrii fibrin is uonverled into pejHone the large iiioleralf of
tbo former ii split up, with nwumplion nf water, itito two ■mailer molrculn "rihe
latter, one belonging to the " anti " nnd the other to the ■' heini " groaj). we ■igto
speculate on n (loatible doiwitication at all proli-id* inlii hctRt'frniridK,
proteida. And holo-proteldi. Thus serum- luiif oi;);- album in, niyunin. and
would be undoubledly hnlo-]>roleidit, pt'ploriM either nnii- or hemi-prMieids, .
we should have to (fi'tin^iiinh I'r'iliahfy jn the heteiogeneoas grmiD of <'
albnioin* both anti-, heni-, and htil'iiirMrid memhen. it U pnatihfr. mar
tliat flhrinoplastic and Gbrinogennus cldliulin stid rnnrin may )ir natural heioi-
veBiBkt
d..iSH
id M^H
^ids,^H
anli-proiei«t.afld not bolo-proteidi^
on these |ioints.
But we have at pmvnl nu puaiiiw knuwledgt
NlTK0<IK.VOD« N05-CKyitTAI.LIXK ItODItK AMJRO Tft PROTKIDi.
Thse resetnhle tlie prolelde In many general jxtinlH, but exhibit ■motiic thrm
selvea mtich erealer dlfTerences than do Uie |>ri>l(>iihi. As reganls Uielr molociilsr
stmclure nothing Hatiafaetoryi* known. Tlieir pcrceatage wMnpoaltfau) apptoMliw
thst of the ]>roteidii, and like Ihcse they yield, under ^drolytic tnalRient. ~
i|uaiilitie)i of leucin and in oome ease* tyrown. They are all anior]ihoiu.
Muoln. {O, S&,T.V n, d-Mi. X, HSO, C, *8.'>».)'
The chnmcterifitic component of niuent. It« exact «im]KialUoa ia ant j*i
known, the fixurea Kivon above beinj{ merely an approiimstinn,
Ai occurring in toe normal condition it given lo the fluidn which contAin It ibr
well-known ropy eooalsleocy, and can be precipitated (r«m thsM by acetic acid,
alcohol, alum, and mineral acids: the latter, if ia esccHS, redtuiolte the on-
cipitate, but this u not the eiwe with acetic acid. In Its pt««I|iiiaied foro itb
inaoluble in water, but iwells up strongly in it, and thin dlTeet u tncrea;i«d by tht
freaeuce of many alkali anlts. Alkalies and alkaline earths dkaolve it midlly.
Is solutions do aot dlalyse: they give Ibe proteld reaetiomi with Milloo*
resgent and nitric acid, but not that wltli salplutte of copper, and are precipitaltd
by basic lead acetate only when neutral or faintly alkaline. According to Hichwald,'
I Bcbwslil. Aiia.ll. nhBti-B. riuirni., Kd.exxalv.&lMk
• Op. at
wb«o beaUxI vith dilut* miticnl adds, niiiciD yields iicid-i,IUiniiB, bdiI iuinck«r
body which in nuuiy of ilx |>rQ[icrlit« cloaelr memblr* a nugnt. inuniiich im it
rrducM itolulionii of cupric sulphate. Prolougtd bviliug witli Buiphuric acid
pvM icndn and abuut 7 per cetil. of lyroalo.
Ptrf/tirafion.' Oi-tjall or aii nqUGoiM cxlract of finclr-choppcd mibmaxillanr
fland lBaciilulHi(«l wiih at-pticacid; the pmHntCatcd mucin to tbrn wimIicI with
waUf, diuolvril in dilu[<i Rodic carbonate and nnallj pracipiuicd iritb ac«tic acid.
It tnaf iilao bu ubiniiit-il fnim iiuaila.*
Ohondriu. (0..tl.i)4, H.fl.TI, N, IS h7. C,4T.T4. S, OHO por cent. )»
Thia la uatially regarded aa forming th« iMiGnlinl part of the matrix of hjaline
GUtllage. and la contained in the inlonticei of Ibe fibfi* in dnilic mriilagc. A
■imitw aubatancc can be prepared from tbe cornea. Eloiltil wilb n-iii^r, it dia-
aoIrA alowljr, fonuinjc an oiwle>c«iit solutiun, wbloh ia precf)iiui<.il by Htrtic acid,
l«ad acetate, dilute mlaerul acid*, alum, and «alta ofHllTer and mpper: aii exce«
of the taat four reagent* r«disMi1vM the preciptale. SoliiiionH of thia body
g«lflllRir« on Maiiding, wen if vrry dilute: the inlid miiaa ia iaonluble in cold
water, teiulily tnlublr in hot water, alkalies, and ammoma.
Tbe aqneooii and iillculine nolulionii of choadrin powota a left-banded rotaijr
poiraron polariwd tight of^itlS.ft"; In presence of «xce«a of alkali Ibtobeconea
-603.0°, both measured for yellow li^hi.'
Itaeems. accnrdinKlo ihc; nbtctvation* of many, thalcbondria can, bv heating
vith hydrochloric acid, be i-onverlcd intu a body wIi'mc reaction* reacmble tboM
of aynUHiin, and anuthemubauinoe. which like' tin- simitiir product from mucin,
*o till reaeuiblea grape ^ugikr IhnI Jl reduces cupric twltii in alkaline »oltitlon :* it
npiicart, however, to i-onliiin iiilrocen. The cxiolence of cbomtrin b" a dMtincl
(uMancc bus bowever been denied* itii thcnupiioaition tlial it la in all i-iuici a mrrf
mixture of other boilien. It is itnted tliat a auHlanoe having all the reniti'irin of
the ao-cultrd chundrin, may iit any lime be produced by a mixture of mucin,
glutio, and inorgaotc aatts. The extreme ulmilarity in tlie reaL-tionit of chi>ndrin
and mncin point to a cloac rolationnhip l>ctwc>en the cini. The whole subject,
bnwcTor. reijiiiriv ni<irf einuolctc invt«tij(atinn. With alkulir* »r dilute Hilphuric
acid chondnn gives leucin, but no tyroitin or glycin. Whether chondrin exi*ts M
uicb in ciirtilnse i* uncertain : it iiecms probable that it does not. liuce ita exirao*
tioD from eartlldfe rer(i)ireH an amount of boiling with irater much greater than
that reijiiiHiie todissolva dried chondrin.
Prr/ioT'itinH. From cartjla(« by extracting with water, and precipitating witb
acetic acid.
OAlAtln or Olutin ' (O, 2«.SI. B, 7.16. «. l&aS: C, M.». 8, 0.60 per
onai.)
Thia la the aubatance which U yielded nheii coo nectlve-l lane flbr» arc heated
for several dHy» wiib very dJIuie acetic aeid, at a temperature of abinit 16° C, or
by tlie pnilonge-t iii'tion of water in a Papin'* digoler. The clantic olenienla of
t connective tiiuui' arc unatTected by the alicivc treatment,
Aanbtaine<l in this war glutin w when tirait'ln thin riuid,an|{difylDg on cooling
t<i the well-kodwn gelatinous form. M'htii dried It it a tnlorleM, ttvntparcnt,
briliie Ixidy. swelling up, but rrtnniniiig undiwnlved In cold water: heat tog, or
the addition of iraco* of acidn or nlknlim, rcndit.v elTccta its aolnlion. Whoa dla-
nlred in wnU-r it ponieaaea a lirvo rotary power of - ISO*, at W C. ; the additioa
of Mrong alkali or acetic acid re<luc(« tlila to -113°or-ll4*, both mcMUrcd for
yellow llcht.' Its soluiion* will notdtalyse.
Mercuric chloride nnd tannic add are the only two rcagenta whtcb yield
inaoUibli' precipiiutr* with thii body, tis presence prpvenla tfa<' action of Trom-
■ner** (Ugar-test, since it readily diMolves np thi; precipitated cuproiii oxido- Th«
proteid reacUooH of gtuilo are ao feeble that they are probably due merely to im-
I
< nchtiklil. op. (Mv aad Chcm. f-pntnlli. IM*. Xo. II. RaaMar Ano. d* rb«M. n. num., Bd.
eai.att iADdw«iiT,z*HK<b.r.|ihx>b>Ln>tK, B<i.T.na«iks.>n.
> LiUHtmbr. ]!t««ih. t phplnl llxin . &1. ti.il'Kx.fL-n,
• naweiiSkt. B4b. pttya. pMk. Olwai. AitiL ( .Inl.. Wn, 8. m
• )IonM<6a«ei?\'i^Mn'i, ?Mulbli.-tDm). V«t. Il«i4tlb«rt. U. I. (UHh DtL S.
T yMlntKoonfainuIal vtUi ttrnKaKtMatmUliffliiMa."
• Uopiw^eylu, U'lk •!. t>Or*. f*iIi. obMa. XaaL, « Ann., wit, B. 2a.
1082
pvritlM. Heated wilb Bulpburk ackl It yields amiitoHU, l«acla. tni glrcla, bai
BO tjrnMla.
Il nppMU» impniluililc Itinl itli'tinexUtii t«iu)yfoniital in mnnn-i' - i' flhra^
•intw ibiae tin not Bwell up in wnivr. nii<l nulv vii.'1il glutin alirr ' Inat-
■i«Dt will) builli>|; iinur; to which it niAir be'*dd«d luai wliil«|:liiiiri ]-i Aiinl mpon
hj trypBiri, tl)c t-niine>.-[iv^tiMU4> Hbm in Ui«ir Oklurdl i-ondiiion rraUt its Ktioa
(i«pp. %'>i}. Wlion vlutin ia Knhmitlrd for rowe lime ut ttic ncliuii of dllM*
hydnidiloric acid, at 38° C, nixl llio dungc ■■ brouKht about eren mon rawUh
hv the Hc^loo of p«p*iu, il lobta iu poner of cekiiniting and ia now dlAMlib
tlin^iigli noroiM metnbrancfl ; tbe name of getalin-iieploDe baa been given toUw
produoi tniu obtiiined.'
Elaatln {0,SO.&. H,T.<. N.lfl.T. C, 6G S pnrceoL)
Tbls ehBracUriBtic i-oiupniieDt of elaatic Dbrc* U leA on lb« removal ut alt ib>
gititiii. niudii, oic, fnini Mich tlHii«a as " llgameniam noclue." odraninf ic Wiuf
tnken of ti» not biinji nlirrccl when il is hcniod with uater, rvtn uoHvr i>teHura,
Itilh Nlr(iti)( BCL-lk lu'iil. or wilh dilutr nilcniir'K. When moiM il U ji'llnw and
clnHiic. but uD Jrt'iDf; tiecoincB britilv. It in lolublc in *ui>ii]c alkitlieB at boitlif
tMtiperalureit. and conceiilratcd bulphuric niid nitric addadiwolve ii eten lalW
cold; it i« al«o diwolved bv the action vf pfliiaTa juit-iv It Ii preciirttatc) fros
tinlutioTii' by laniilr Acid, but not by ihr luldi'tion nf ordinary aci<l*. Ndwhfc-
aUndinic Ihnl it clwclyappruiichoilhi' praloidtiinita pcKcnla)(v compoiitJan. tat
giTCa liittl^ucl altbou'gb feeble pruteid reactions, any very cliwf rc1atloB»Up
between the two appear* imptubable. atnce elaalin wben treated with sulphiuw
acid, yicldn leucin |S0-40 per ceiil.) only and no tyroaiii. ~
i lilKcr' hail obuincd a mmilnr btxiy from the abvtl t&embtane of anaka
Keratlc* (O. SaT-Ss 0. H, 0.4-T.a >', IS 3-17.7. C, W a-£3-S It, AT-U
ptr cunt.)
This body, [hotigh somewhat resemblinc ibe prolcidi in general oorapcNlliM,
dill'en) fniiii Uiem anil aUo rioin thi-prei^ethiig boaieaiowidvly iu other pmprtliM,
Ibnl il» dc»crii>l)i>n i* ptiiccd here for roiivenlence rather than anything 'W.
Hair, nniU, fratheni.hitrn.itnd epiilcruiicocalnconaiitfor themo*t part of keralin.
Ileatcl wilh wiilrr iu u diErmtrr iit IW* C. koratia i» partially diaaolred with ew
lulion uf HuIpburrttciJ hydr<i|;en: ihe wibition then jciTe* with aoatic acid and
fermcyanide of potauium il precipitate soluble In excew uf the acid. Pnlonnd
tralliiig with alkalies and scidt, even acetic, dtaaolvea keratin; Ibe alkaline mm-
tions cTolvc Hiilphurvtted hydrogen on treAlment wilh aclda. The eul|>hur ia
kemtiu il rviduntlv rcry Icinat'ly united In the nibiitance, and in all it* rcaelioaa
there iippean to be a wunt uf xiniiliiritT between keratin ami eitJier pioleldi;
mucin or gelaiin. The bio«i ci'niiuon uf itn pniducis uf deoompoKitioii are leode
ilVjierccnt) and lyrnMn (8.0 per c«nt.]. and aonie aajiartic acid^ no cljcta it
irmed. What in jcenenilly known iw keratin io probably nconipound bQ^,«Uek
ha* iKit yet been molved into it* cumponenta.
Kwitlil unJ Kuhiir' buvo deKnt>«l ■ now (nily to which, —aam It e««iir*aaa e«i<UlBMI
of uetrav iisiiie tboih of nerve* ami "f lli« eeniml nertuiu in<icin|. aiiil ia yat rinwity tSt^
liotl wit^ ordiiiarv horny liMna. iliay (tire ItM tiiiinn ••! ii<tin>konitib. It U mparol ia
quaiiUly (hiiD tbebrsin byeilraoliTiK Ihli tjBUe witli iI.viIkiI ■») Hlmr, aii-l hiI^hiIIki; IU
residue to the sdifln otftptia nnJ Itypin. The liiial nwi.luu ii ueiMO-kciMiB. aira •mnrtt
Id I& Iu te |<er o«L of lb* orisinal tl«u*.
Nuoleln. C„R«S,P,0^
Diftcovered by Mincbei* in the nuclei of nna cmrpiHclM and in (lie yellow mr-
jiuicle* of yolk «f rnK' fXhct obecrvcni Iiaw NUhMqntfitly obtained It from
ypMt. from Kcnicii, I'lum tlie nuclei uf the red bluud-AOfpiticIni of litnia and
amphibia, from liepmio cells, and il b tirobably pKaciil In all nuclei.
Wbt'U newly prepaied it la a oolorleM amorpboua Ixidy, eoluble li> n alifbl
extent in water, rvadilv aolnble in many alkaline MitutKuia; but lie aolubilttie*
•Iter on keeping. If added Kradually in tiiibcieai •(unntily to aaoluUonof camtle
■ pciftMlM«(,MtMli. f. pbr-tol (TiMin..Bd.a.<in*I.B.niL
• SnTft. muliwtL Cb«l> '-- l.-^ t,?\ -^ w.. ii,,r__-r,
• lMnII."Kan»IA]r
• X'trtiaod. XaHiUM^ <
» Mtd'ltan. UmwwiJi.. 1. , .. .,..-, 111.;, t, . ... .;;
■' •vA.-nnoc
Il ITLtt«l).^aldL
CURMICAI. BASIA OF T H IC ANIUAL BODY. 1033
ilkali It Drill neulnilixe* tli« HQlution and tlien rend«n ll*cld. Ilseenw Co piaur^*
all iDiliKlinrt »nt)i(i-|in)t«<ic tcaclinn, but glvM no rMi-tJon with Millon't llitid.
It fields prcvi pita ten with sctctbI mIi*, r. •/., linc chloridtt, nrKcntfc nUrate, and
cunric oilphaie.
/yrjiaraliort.' Since nucttlii is vttf r«ii*t«Dt to the kCtioii of pepsin, Jt mny Im
obtained fiofli ih«i granular n-oidui; coiwlalliw oblelly of uudci, which m.-i.-un ufter
dlKn>lii>K p«i» wiLh jicpsin. The iiicMt rvinarKabI* featarc of thin body U ita large
Cerceniaee uf jilioiiphorus, 9.6fl per cent. This pbonphoriM U nwdUjr separatod
r boiling witli Btroii)- bydnivblaric acid or caiiatioatkaliea; the aame occiira wllVB
Mlatioa* of nucl«ih are aciduUt«<l und allowed to itaiid.
OWlln- t;^n,N,0„'.
Altbuuch nut found lu a con*tilu<'Dt of any maTniiiulian tiMue, thii aubiiUnu
CMBpcwfl the chief part of ibe exo-akelelnn of dudt iorcn«bratea. It mty
pnibabljr be reearded aa tli« animal analngue of the cellaloM of planta, and ttom
tbh pniDtoriicw it poMeMca conatderable morpbologk-al Inlcimt. DoUi oolluloae
an<t cbilin njiprai to Tivld tomo form of augnr nbeii ircjiti-d <ritb *trong acida.
Whrn purified, (.'biiin U a white amorphuua bcxly , nlli'n rctHiiiiug the tfbape of
tb« llttue from which it baa been prepared. It i« in«oluble in all r<«K«nla eacapt
atimtg mineral acid«. the beat solvcnia beloK mlphurlc or hydroculoric acim.
Tbe iinmvtliatr addition of water to the«« ooluUon* r«precipltate« the vbiliii in an
unaltered form ; but the prolonged action of itul|iburtc acid cau*r* a decomtioal-
tlon rAnlltDg, uccording to mme iilscrTcrc, in the formation »r an amoruliotu
fermenlablc earbubydrate ; nutl when bydroi^bloric acid in used an amldatcd car-
bofardnte ia obtained to which the name of glyt-ottamin* baa be«n given.
/VaM?rj(i«n.* The cleaned exonkcletoii of a lobnter i« thorou)tb1y cxtmcied
with dilute liydruchlorie avid and thru with cauntir aoda. To purify it l^natly it
ia Hibinitted to prolonged boilin); witti a Hotution of potawic permuugamite.
CARBOHYDRATES.
Oartain m^mbcra oaljr of thia ctaaa occur in lb« human bodj ; of them lb« neat
imputlaot and widopread uo thn*c known hh glycogen and tM twoauganitgTkpe'
■agar <>r di-xlruii- Igluctmc), with wliich diabdic ■uciir iieMna to be idOBticar SDil
nsltuae. Next tu ibeite L-ouiea milk-sutcBr. luiwit la another body of this cltM,
•lIlHMigfa it dlfTera lu many inipciriaut |>oiuta fruiu the preceding two.
8nfu* tn oRon miuiilikrvd to be puljtlioiiic aloohala. SannJ of tbeiu itand In peculiar
relation I" inannit, and may be «>nveri«<l inln that ■abttanc* by Itie u'lion uf toiinia
aBalcaRt.*
1. DaxtroBO^Oni-o-tugitr]. C.n„O(+H,0.
Oocura in the cont«Bta of the sUmentarr canal to n variable extent denendcnt
on the nature of tha food taken. U in alio a normal conittiturnt of blood, clirle.
and lymph. Ounceming it* prcaenoo in the liver, aee p. 'iT-l. The amniotic fluid
alao cunlalna thia body. Bile in the morputi condition >t free froBi ougar, ao alwi u
urine, llmugh thi« point haa giren riae to great dittpute.' Tlie dUeue diabetea ia
cb*raet«riMd by an picau ot dextroM in tba fluida and tiwucti of the IkhJt (w*
When pore, dextioae ia colorlena aod eryatalliua from iu ai]ueaua aolnUon In
•lg-ald«d tablea or prtoma, often agrio™'^'*''*' iniowartylamp*. The crystaU will
diMolve in their o»d irdglit of oolil water, requiring, however, aume time for ib*
proeem: they are vetf readily aoluUe in hot water. Dextraae ia wKnewbat ipar-
tngly aoluble in nlcihul, and cryatalliuK from anbTdraiu alcohol tn priauii free
from waUr of vryatallixolioo ; it i* marearer iiwolubia iu cihcf.
■ ate komh, znuoh. t nhnini- ctimi , Dd. ni. ntiff, SL m : to. nawh B. M) j vlLllaa). B.T.
"yUMWoli. aterd Nurldii u liini ■'[■HufiMirml." Blrw»a ■ Un.
*htiATtum.7i»-rh. I |<i>r>ini 'luon. ifl tl. ii*7«).a. ai
• LiMCTlm*. I'ffi cit.. M. IV. imjl, !^ XK.
• Ba(wii)i. Areii. t A»i. u. it>n>»i- Jaiux. in«, & an.
*l^qiw«ii9n,liiiw»T»f.wliiniiw>»t«i»i««rt»ucaofia«areeewnlaalaiheaywallioJyh>wnM
bam conhundM io(«llwir uudv ibe eamnnn nama td AunvM tm ilaeoaa n*? bt onwilital at
MMM an opts «n&
• UnuMMiia, Ann. d. Cbvm. n. PhMU., M. mill. B. laa
1 Sm»mt». tMrMa»«MHMU«ut.laL.lL tM
J
AVVKnOlT.
Tlie ftMhljr prapnnd «ild «qaaMl* ■olution or tliD cry*!*!* tMMMaM « daKUo^rutalor; pM*
»r-rt04°rurf#tlnir lialil- Thi*. qakblr oil lia*ilB(,nHt*H«wI]r on Maailiug. rklti (" -it',
Ki which paiul il nioMliii oonaUat.
ItaxtnMB resdilf fntin* Mni;»und" witli *oiiU lad nanj MtU; the laUrr u>» vary ■Ilu•ila^
doeonipailfii.iD nkpidty i>nliiiii|[ va IiMling llivos. Wh«a lU laatolllc oaminaiid* »ra iimm-
pMtd, Iho docompoaltlwi I*. In auutj cmo, •iin>in|«niad br Uia pranpiintin* ol l]i« mMUh
«■ If: taivm, itiild. meraurjr, blefnuUi. Ckuilin iitkiiliN rMilj' lifMupsM ihMD, ■« aim Aim
Dextrose in nwillljr nod coranlfftety iirvcipilAUxl by lead koeUU and aniiiKmii.
Ad (mpiirtnal [»rniiert;r of "i'n body in iu |>ower of undergoing Iwwmitow.
Of tbeae the two princlpiLl ar«: I. AktrMie. This u ptx>dtiM<(l ia «qu«atu aoln-
Itonn of doxtrose, undvr ilie inllueDce of y««Mt. Tbo dfConpOMlioa b ibe folio*'
inc.- (',n„l), = SaH,a ^ 2C0,. yicidinB (*lhrl) nlcoboi »ml c»rbonic «ahTdti*.
Olber ali-nliDU of the acetic Mcrie* uc feond in tisce*, w bIm ar« glyoefln, ■uc
L-inlc add. and pniliably mnnv other bodiw. Tbe fennenUlion is most adi** H
about 23° C. Below S*' C. or'nbore -tA" C. It almost cntlrclr ce«<«>. If the mc-
cbarin* iolutioa cooiainH inoru than lli por cenC of sugar it will not all h« ik«at>-
piHwd, a« exccM of alcohol stops the reaction. 2, iMflic. This ucoura la U*
pfta«i)«e of decompwRg nitrogonoui mH(l«-r, twpccially of caarin. and is probahlf
th« rMult of the action of a spvcifio (erment ' The firvt stage Is tb« pitidunka
of lactic ucid, C',H„0, = 2CHtOr Iu tbe aeond butjric nvld is fon)i«>d wKk
cirolutionof liydri>geuaodf»fboolcanh);drlde:2C,H,0, = C,H,a, ■■ 2<X', - illr
Tlio nbova ch'aiiK««. lh« Hrst of nhich is probabiy iindi^r^oovi by nuxar to a am-
siderablo oxtcnl in thi^ intestine, nre moat aeti*e at Xi' C. ; the preaeiice of alka-
line ciirl)iiimt«ii il itt>c> favorably. It is, moreover, aisential that the lactic icil
should bi' neuLrulizeil na fiLal tia it in forui«J. otherwbe tbe pres«aoe of ibi* fn*
add alopt the proceiu.
Tbo preparation detection, and catimatioo nf dextroaa ant so fnlly gifen in
rarioun Imoka that Ibcy need not be detailed h«re.
3. Maltose. C„U„0„ • H,0.
Tbi« form ofiugarwss Hrsl described by Dubmofaut' m a product of tbe acHea
of matt extract on starch. It* cxist«iico was for a \on% time doubted aatil
O^nlUran* repeat^ and (^«tilirmtnl the previous DsperiinuntA. Aocordiiw lo bia
It etratalllcea in fine atrit^ulnr crystals, posaeMeu a spedfio rotatory pow«r of -r IU*
and a reducing power nrhii-h looulv one-tliird as great as that of dextiiMr. ll
•eens probable t^at thin in Ihe chWf «iigar obtainod by Uie action nut only if
diaataae butorptrniin and nitnrToatic rerinenl upnn ■inrch and {irrhnps alao a|M«
glyotwen ;> althougli suini- ilcxiruae may st the Mime time be formed. MuMului
and Oniber.' have ahoarn that mullose may alio be formed b^ tbe action ni dOatt
nilphuric acid on atarcli. and that it is rspable of undergoing alcobolio ttrmn
tation. ~
/Wpiitnliini, Sec Miimuliu and Oruber {loe. eH~],
S. Mtllc-susar. C„H„0„ f 11,0.
AIm> known as taebme. It bi found in milk, and b characterislk of tbl> ««er^
tioo. It is said, however, lo oc>cur abnomially in tbe urine of lylnc'tn w<M»m.*
It yields, whoo pure, bard. colorleM crystals, belooeiug to tbe rhombic ayilsa
(fournidttd prisms). It i* loan soluble in watv'r than dextrose, roiuiriii)^ fur «1a-
tlou six times its weight of cold, but only tvn |>arts of bnilinj;. water ; it is tatMj
insoluble in alcohol mid ether. It is folly precipitated from ila aolntiona by IM
addition of lead aeetalo and aoimouia.
Wh«n frMhtr <li*»lvc<l. itsiu|u«ou> ntulion pMsasses ■ (piMift« ittxtm- rotatory |Hvar rf
i-VAJ'tDr sr>dluin iichl; ihit <liiniii>i>li<M, •lowlj nn Stan dill (. rs|ii<lly on iBltinr, anUI a
nslly iBiiiiuDs cooMkQt at -^UA'. Tha sniiHinl of rolaliou Is Inikjisa'lDat of tft Bsaa»
Inlion or Uh •utulmn.
LaciDMi aaltss i«aili1y with basts, fnrxninx unslalila ■viiiKintitl*: tittm Its meUUIe vm
pouiiilt Ikc neUl is pivcipltaled la Uia rvluvod slAt* i-ii la^lin)-: II toliioea wppw •slwst
raadlly u d«itn*a tnl (o a less exWal, ris., in the ratio of :<■ t loll,
■ Llsur. rMk Hm. Trans., vol. ft* lf% v. ts. abo QiMn. Jaatn. of Uteraa SiL, ML s*ui lUOh.
p. in.
' Ana. t^lDLPhnCDixl. llUTi.nlTH. • Jooro. Cliiiai.Soa..SM.% *•!. X.(aB& h OK
• Mutculusa. v.Weitiia. 2#ltKh f plu«lol,ClimD.. M.fl. fls3«>.il«al
' Zf iwttr. b phnhd. cImd., m. II. mn). ti. m. • UoiMMer, twd.. Bd. L (wrri. «>, m.
J
CaBHlCAI, BASIS OP THE ANIMAL BODY.
toss
ctuM ii Kenutlljr tUted to udinit of no Jirrct klcoholic fcrnvnUtinD ; this
miy. hwTATer, MHDetlinffl ba Iui)ui;«<] bv ibe pruliitipMJ ociion of fetuc Bjr bolt-
tti^ witli cliliile Riinornl acids \nctnne In convorlod into giilacUHtv. wbicli readily
nndrrKu ■lo»h4ilic ftTmntilalioa Aiid poMtcm n grcAtcr rolACriry power tbun IncKMif.
It fiia; b* ramarkod h«r« lliki Ihoiijih ifoUttd Uotu* la inospaM* of ilinot kloaholle br-
iMitollon. Billk ilHlf m*f be Ibrmanteii; BoftbdoC ru naabU la thi* ilipwt »lcnhr*lic
ArnMuUtlaD lo ilFtM't uiy inl«na«dial« >h»ng* of Um laoteM into %WT Mlier bnnonuUo
LaebMe i*. hoire*er, ttimtty cnpab1i> of iinderKning the luetic mid butyric fer-
mnitBlloiw; tli« circamsc&ncM&nd iiroducia itn? tbe name m in ihei.-nneof il«xlraM
{•w above). The nction b gcnernliy prcxlacllve of n coUaE^rul small iiiuntiiy of
aloobol.
Lactnoe m lbu» ili>tinj^i«hed tmai dvxtroM by itn mnaller toliibility in irater.
Inaolubllity in aloliul. urytuliin* form, loii«r cupric oxide reducing power, and
its incapability of uailenrning direct alcoholk- fermentation.
fiftparali/tn. After the removal of the casein and other prfteidn of tbe milk.
th* oiolber licfnor i* <ivaporal«d U> tbn cryntallixing point ; the cryalaU ar« putiBrd
hj rvpoatcd crjrttallitation from warm irat«r.
»4. iDOalt C,H„0, + 2Ht*^-
Tfals lubciauce oocnn but ajiariiigly iu tlie human body ; it woa found ori^nally
bgr Soberer' in the miiacle* of the heart. Cloetia abowed its prewnce in ihe
kiBp. kidneyv, iiplccii, and liver,' ami Milller in the brain.* It occttt*. aim, in
diaMtic urine and in that of '* BriKhl'* dlaviuie," and ia found in abuudauce In
the vegetable Icingdom.
Pure iuoslt fornia large efflnreeceot cryBlalii (rhombic lablen); in ulcroacoiiic
I pf«|iaratioDalt ianeualtj'otilained In inftcd liini]ittorfliie cr7>bil». Easily Holubla
In water, it la inmluble in alcohol and rthor. It nowcMo* n" action on |iolariii«l
Illtbt. and doc* not reduce aolutiona of metallic aalta.
It adniita of no direct hIchIiuIIc, but ia capable of undergoin); the lactic, fer-
mentation ; according to Hllger,' the acid furuied ia aarcolactic. It ia unaltered
bj beating with dilute nilTieral acida.
IlrtparatioH. It niny be precipitated from il4 aolutioim br the iictlrin of 6a»ie
IavI aMialc and BoiiDonin: tbe lead it then removed by anlphurcttcd hydrofreu,
and tbe inoait precipitated with exc«M of alcobol.
Ai a apecial teat I.Schprer'*) ninv be montioned the production of u bright violet
cnlor by care (III evaporation to dryneaa on platinum bil, with a liltle ammonia
and calcjum chloride.
h. Dextrin. (',H,gOj
Uy boiling atntcb-past« with dilute acid*, or by the action of feraionta, the
•tareh >■ converted inia an iaomeric body, to whicb. from it* action on polarixcd
light, the name dextrine baa been gireii. It insoluble iu water, but in precipi-
taud by Htciibol. Itdoea not nnderco alcoholic fecmeiitation until after it liaa
b«tD chaiigeit into drxtroM, nor can it reduce metallic aalta. It yielda a rcddnh
port-wioc cnlur u'itb iodine, which diuipnear* on warming and doc* not return nn
ooolinp. rurlhfi aciiun of acida ur of rennenla cunrcrtii dextrin into dextriUe.
Dextrin ia preveai in tbe conientB of the alimentary caual after a me«l containing
atarch, and ban alao benn found lu tbe blood.
Tliere m not tbe Icaat doubt that aevcral modtflcationa of dextrlu exiat, and
may be oblaiaed by the action of acJda and fermenta on atarch. Of tli«M two of
tlic bwi known are tboao dncribcd by Bruckc' under tho name pferTtlirodextrin
and aebroodexiriu. tbr fijrmer giving a red color with iodine, tbe latter not yirld-
Eng any color at all. Eryibrodextrm ma/ be readily convcricd lulo a augat by
tb« action of frrnienta, and thiu ia not found iw a product of the oomplvle action
of ptyalin on atarch. Aehroudextrin, on the other baud, ia not thu* coDwrted by
fennenlH. and tberefure rcmninB in aoluttou, toj^tber with the Mi)(ar fi>rnie<l by the
action i>r utTallii on alarch. Achroudeitrin may b« cuiivetled into doxtrMe bj
boUiiig wllb dibite hydrochloric acid.
I
I
Ank.4,CtVHa. IL l-hartn. IM. lazl1L&
IHd.. Bi. eU.. a IW.
atOm. d. Viva AL*4.. tK-i, W. Al*h.
• II. .
Aln. Vortamacm, 2 AnL .
10S8
APPENDIX.
a. aiyooe«a. c,n„Ot.
Bcloan to the stnrch division of carbnhrdratw. Oi^rovcrtu) by Dcraon) la th»
liv«r Mid other orjcani (•rn p. oiai.
Olyconn !•, when pure, un nmorvluKt* powder, colorI«M Miwl Ujttel«M, Nadl]/
tolukl«In water, IiuoIubl« lu aleobol and euier. IUMiu«u«uaoltiUoB i* eencnDj.
though not slways. nirongly opaletcent. but cootaiiis an particlM Tiubip Biov-
MoiHcally: the nimlwrtinci- in much iv^uc-od by th<i MtMctte* of fr<<» alkali**.
The tame nolulion pounuca, accotclii))t to lloppU'i'cyri.-r, h rcry atmng d*XU^
roUtory pow«r, about thrw tiinM «a great u that uf dcxUuM;' it dUnltM
hydraliii uupric oxiil^; but tbU Is not TMUoeil on bollitig. _
Ry lb(- aciinn of ijlliile tnineral acids (eicepi nitric) il ts partially onnrvi^MH
into a fiinii nf niitcnr i-nry clotrty m*mblinj(, thoiijcb probably diffrrini; ■ninewl^^H
from tmc cIcxlriHr, and ihe intiir conversion is aliw rwdily edecled by the act^T
of amylolriii: l'«riiivnt«. Tlir nuj^r into wbich the g1y«30gca of tb« \frtt U oaU-
rally coiiveited after il«alh <Hi'e 11. i'7'l) ikjipeflrs to be true ilestrt**;* ao alao Ibt
•agar of diabeioa. Tlifi roxiili i.f iho aciion n{ Jia^iaM. or ulivary or MocrwUlc
ferment, lino n Klycojt«n in, hnncTer, iic«:uniiag to MudcuIiu auu v. alarioit.'a
mixlurruf ai-hriiTiclrxtrin unci nialtate; the quantity of dextroafi niiUEtng lla a^
peamrice at the wime time being very ainall.
OpnlMcetil itoluUonti of );lyi-OKi-'(i UHiiully become clear on the addition nft«ujli(
alkali ; VintNubgan tied Hic'tl ' liavo shown ihiit ihi« is aooonpanicO on botliDcbf
a change which i-onvcrtu a portion of Ih? slycoKm into a sabntancv to wbieh Uirj
give ibe name of .'■Klyi.'oKeti-drxlriiiP. (KuUiif' bad prevtoualy d«*cribrd a bodj
to which he gave the tiniae (clyro^jeu-dexiriri. That deacrlbed by Viatachgau aM
IMttI dilTins slightly frnm Kdbnc's body, hcnctr the nan>c. According to itat
authora one-fifth of the KlvcuKcn in at tbci name limo diangcl ititu Home olbrr, tt
prwent UDdetermined. i>ulwUint-e. Nuriuiil lead acetate Kicca a cloiidintw. iti*
basic salt a predpitikte. in fluluiioni of glvcoKen.
As l«eti> for ibis body mflv bi> used the Tormacion of a |Mi(l-wiue color with
iodine; this disHppMrs nn narmin^, but relurna on cooling. Tb« sama colof i*
produced by the ncCioD of iixliuc on licslrin, but this doca not f«appMr on osol-
ing after it* diMppearanco bv nnnning.
Pffpnraliaa ^ •il^.virn. 'riie follovtiog ia UrOcke'a* metfaod. The filt«r«d M
simply simlnediiecoaiuu of perfectly froth llv«r or other glycogenic Uaane i*.
when rnld. Treated alternately witli dilute bydroeblorlc add kimI n aolution nf ibi
double iodide of pntnmium and mercurf' as long aa any precipitate occurs. Is
lh« preaencr of free hydrochloric acid Ibo douole iotlidr piccipitatea iitotM
malteni su comptFtelv iia to render tbeir loparation by Gltrslimi eiuy. Ts* ■•»
teidslieing (hu« ^oc rid of, the glycogen is precipitated' from the filtrate byadditc
alcohol to ihf «)(tent of between <!<'> and 7t) per cent. Too much atoohnl (sub*
avoided, mdcc other substance ho well lu fflycoBcn are thereby precipltaUJ.
The glycuKrn v. now washed with iilcbol, nnit of t;<) and then of SQ porcruL,
HiWnnrd with ether, aud finully with abiulute alcobul. Il is then dried om
sulphuric add.
Tunioln. (C,H„Oj)^
Ttiia budr ia regarded by nany observer* as identical with the true cellaloienf
plants, while others hare iiHcribeil tu it properlim dilfering from Iboae of cvlluhoe
sniDcientlj to Justify its receiviniEH distinct Dsnie. !l appears to be more re>i#kni
to rhr action of chemical rca)[cnt« tbiiii plant cellulose.
tl (untlilulci ihc chief part iif Ibr inteifutnent of the asctdia or (unlcala. is
prepared fnim tliiM source it i*. when pure, quite while, and usually retains ill*
shu)-* of the liosue. It Is uiiai-tnl upon by any reagent except strong odd! m4
alktilies. luul by tlie action of tlie forner it yields some form of sugar.
■ »r>i:iiii.niutvt^Anh..iM ixi*. (um), 8. n.
' l-noflafs Ann.. »l Kli. |is»i. & Ul, aiul aalr. om). B. Ml
• atiMUu. c
Ai-h Kai« IML. Bd. ajdr. (Mft
• PMnfaAicK
• UU>b.it.
^%'
pbrMoL '
- 4. WViJ
>>'ii, B-l il.(tffin.8.4M.
■: IM.
I- lalfl. |ISI», U. Ahlb.
' TMi Di*T l» piv]«id t>y i^ccuullaUat r^taaao kOIilp irtlli mMTMiric «Ii1b*1iI» and ilMMmwl
rnsbcd p(Mi[Suw lusliiM tnluuon of pMaitfa Mblv ■* fanfMitcannaiiesMks takaooiTJ
foollsf, Mnw aBosni oT snaiSiaw oecnrs, which dui tie
At aw.
■a
(omi olT: tbc BltiMt u ih^ i
CUGMICAL BASIS OF THE AXIUAL BUOY.
1087
FATS. TUEIR DEBIVATIVES ASD ALLIES.
Thb Accrto Acid Skrih.
0«n«nit fornitiU, C>1I,*0, (ini>ni>l>Mic).
This, which i* nnp i>r tb« mott complete homologous wi'w of organic cliem-
itUj, rua» piirallrl lo the •eric* of moiiKlonnr alcnhota. Thii< rormic add corro-
■pood* lo Diolhvl nkubol, aatilie Moid to rthjl (uriliuarv) •k'ohul, and hi on. Tbc
Mveral Kcida may hv regarded M beinK derived rron tbeir respective alcuhul* bj
umple oxidaiion ; ihuo elbyl alcobol yielda l^ oxidation acetic acid C.H.O -I-
O, — C,il,0, + H,0. Tli« viiriouH members diHer in composition by cll,. and
t£e botiing-pointK ri>e aticciwivrly by about 19° C. Similar rclalioan hold gn<Ml
with r«f«rd to tbeir mrlliiig points and sprcific grnvitie*. Tbe acid pnipcitica
arrntrotigeiil in ibosiMrhere u baa ibe least viilue. Tlie luwi-at uierubcn u( tbe
aeries are volatile iiouids. acting sh powerful acid*; ibMc auccewively become
lea* and Iras fluid, ana the highest menibere arc cjlnrlem mlid*. cluely resembling
Um) neutral Tata in outward appeamnce. t'oDKCulivo acids of the arrie* pns«ni
but very sinaJ) diflTerencea of chenuiuil mid phvuical pro|>er(ies. hriici! the ditGoulty
of Mfwrating them ; tills Is fkirtber lncreaw>il in the auitnal body by tbe Tact that
•ZMllj Ihiwo Hcid* which present tbe Krcuirut ximilHrititH usually occnr together.
The free acids %n- found only in Hmall and very variable qiinniitics ia various
piarta of tbe body : their di-rivuiives, on the otfaer band, form miut im[>ortant coa-
elltuenis of thebuman frame, and will be oooaldBted flirtJier ou,
Formio aold. cno.OQ.
When pure is a sttonKly cormeive. ftimintc fluid, with powerful irritating odor,
(olidifyinf; at IJ° (!., bnilina at IfHi" 0.. and capable of boinji mixed in M propor-
liona with water und alcihiil. It biui bocn obtained frum varioiu parts of llie
body, such *» tbe oplecn. ihymuN, pancreas, uuadea, brain, and blood; in tlM
latter its presence idht beduc to the action of ncids on tbe hieoioglobln. Aoooctd-
ioK to some natbors.' it occnrs atwi in urine.
Heated with sulphuric ncid it yields carbonic oxide and water; with cauetlc
pOUali it icivn hydrogen and oxalic acid.
Aoetlo acid. C,H,O.UH.
Ja di»tin([ui<bcd by itit characteristic odor; iu bollinc point ia 117* C; it
•olldlllea St &■, and la fluid at all tenipentturcs above IS* C. It Is soluble iu all
prop'ininDs in alcohol and water.
It occur* in the stomach as the result of fenncntaiire chnnKts in Ibe food, aad
ia frequently present in diabetic urine. In otfaer organs and itaids it exbt* ooljr
In minute trace*.
With ferric ebturlila il y)el<la a bbod-nd lolntioii, deeoloriaad b^ hydracblorie add. (It
diSm in Ihi* lad reaction (n>m mliibonyaoiilK of Iroa ) lleaisi ■iih aloihot and Hljihofk
acM. Um rtiBrar<«<>tic uflur of aeetle ether l*ob<aln«iL It does tint ndnoa illver nitrate.
Propionlo aotd. C.H.o.OH.
This acid cl'Kwly rcHcnihIes the preceding one. It posscHes a very Miur laate
ud pangcot o>Iar; it is soluble in water, boils nt Ml" C, and may be »«|i«raud
bum its aqueous uilutinii by excess oC calcic chluridc-
It occurs in small i{unntitin in twcat, in the cvateiila of the siomacb, and ia
dlab«lic urine when undeisoing fermcntalion. It ia similarly ptodneed. mixed,
bowever, with other products, during alcoholic fementation, or bv the decom-
position of gltcerin. It pnrtially rediMMsiilTcr nitratoewlution on boiling.
Butyric iwsid. C,ll,o.ou.
Ao oily uolorlcas li<)Uid, with an odor of raacid butter. anluUe iu water, aleo>
hoi, and ether, boiling at 1112* C. Catdc chloride Mparatea It IVrnn lu aqueona
aolution,
I BnllflnskT. tioi'po«Df k
rktrnTaiK.. lui. TfiL p. ool
tar's Med. obeiD. Ulilhslluuc. II*A & ft ua TliiHlklMin. Joun, at lt«
1038
APPBifDIZ.
Found ill meat, tlic conivnu of (he Ur^e intcstino. feoM, And in nrlnr. Il
occurs in trmvo in many oUirr fluid*, ftod m |>l«nliruliy obtained wl>»n dub«tie
urine i* mixod willi pun-ilcrrU chnllt and ki-pt at n temperature of 3fi*C> Il eiWi
oa a uvutral fat iu auibII (|UUiLltieit in aiilk.
ThiH 1« ib« iiriiMiipal jnoduci of tbo tecond stage of Uctic fenaentatloo. (6«e ,
dexiroie.)
Tolsrlonlo ooid. C^n.O.on.
An oily liquid, nf iinnetrflting ndnr and buMiinK taDtr; »olut>lr in SO fWtaaf ]
wilier al !?'('.; readily wiluUi- in iilc<ibol ami i-ihcr. Buil* at 1TA*('. PnaoHh '
in Tree and combfoed Votid. a I'eeble rigbl'banded rotation at the |tUn» af ftikr-
imiou.
Il ia found in Ibe lolid ex«<?mrnifi. und is fonnfvt r<?adily by ibe decoBfwdtlci^
tbrongb |iu|[cfa<'lion, nf impure Irucin, ■mmonU bi-int; nl ihr uunr tilli««««l««d;
bcnoe it* occurrence In urine wkea llint lluid contain* Icuoin. bj> in raina of acOM
ntropby of tbe li«-er.
Cttproio add.
Cnprylio "
Ottprlo (Kiiiiu) acid.
C,H„O0H.
c. n„o,ou.
L„H„«.OH.
Time tbre« occur togetbvr (a> fat/>) In butler, and arc contained In
proporlloiift in the fecM froiii a tnt-at diol. Tbe &ntt in an oily fluid, s
wiluble in walor ; tb« olhcre iir« stilifU and •cnrcoly aolublo in wntvr ; ibey u*
•olublc in all proportion in nlchol and olhrr. Tb«y tiuy be prc|>arcd fron
butler, and lepuratRd by tbo varying aolubtlitiM of tbdr barium aalta.
C,
„H„O.OU.
Iiatiro»t«arlo aoid.
MyrisUo " C„D„0.OU.
TbeH o«;ur u ii«atntl fata in (pomiaceti. In butter, and otb«r hU.
wtnl no points of Jot«niM.
n«yi
PnJmltlo sold.
Stearlo
CV^d.un.
TbMs are aolid. colorlem when pure, taatelea*. odoritn cryitatline bodloi, ibt
former meliinit at 03' C. Ibe Intter at lld.ST. In water ibey arv aui(« itMolubk:
palmitic add i* more readily unlublc in cold aloiihul (ban ■teoiic ; bolb ar« ntiHij
diMolTcd by hut alcuhul, ether ur chloroform. tilsciaJ acetic acid diMolTw Umb
In large quamitv, ilie solution being amUied by wanning. Tlwy radii v Dm
foapo wltb ibe alkaJiea, oIm nJth manr other melal*. Tbe varying solnblllUMW
ibeir bniium (alta afford the mean* nt iicpnriitin;; tbem when mixed ; ' tlila n^
nlau be applied bo many othem of the biKner mnubcni of (his scriea.
ThcwacJdcin eombiiiadoii wilb glycerin (nee below), togi-tber with tbeanala-
gOBa oonpouad of oleic acid, form uie principal coi)8ti(uenis of buinao fat. At
Ml(a of calcium they occur in (lie fecea and in "adlpocire." and pnibablyii
ebyle. blood, and Herouii fluidii, ox wttiii of «odiuRi. Tboy nrv found in the,'
•tale in decom|io*ing pan. tuid in <!ajM.'<fUB dapo*i(a of tulwrculoaia.
Tk* (rsiMenae ot mariarie Add, intvriii«dial« to Iho abaTa lwi>, it ftol now ailinlU«rl,i
nun lleinU' bsiibowa thM It ti redly a mlxlura of palmitic and tUiie aeld. Uupiwl
Bolil pMMMta the anoiBaloiu melting pwat at ftV.S" 0. A nunara of A" faaU Naktle and M J
of palmtUo aiidi wald at SO.S".
Acim OP THK Olbic {Acktlio) Sbbibs. H(CiiIly»-JO,(moaobuic].
Many addi of ttiia aerlea oeour ■« glycerin compoumU in rariona (ats, Tiief]
M* Tory niMtable and readily abawb oxygen wbi-n eip(i«ie<l u> tlut Mir. Tbt|
higher iooB9b*TB ftna deeompOMd on aKetnpting to dladl ibetn. Thoir ohM
|ieailiar property ia that of wing converted by iracM of NO, into *olid, «Uibk
netainetic acid* capable of being dtaiillcxl. Tb«y bear an imereaUng rdat
BeOM. Anml. da nj*. u. cliciii.. Bd loilL h. Im.
■OtLI
CUBHICAL BASIS OF THE A.N'IMAL HODY.
1039
I
\
»
lb* acbb of Um> acclic *tntn, Itroithin^ up whta hemed with ututic iMlnitb Imo
■Ofttic acid iui<l «i>iiH- iiilitT ini-Diliprof the kbiiki m-nra. than :
nC„H„O,-r-JKH0 = KC,l[,0, + KCwQ^O, + D,.
01«io add. CuUuU.utl.
Thin i* the only arid of tb« Mrin which !• nhyiiolo^oBlI/ fanporUDt ll ia
fi>uD<l uniled with gtyceriu in all the IkU of the nuDUUi bodj.
Wh«ii pui^ it >». Bt otdituuy tempera 111 res, ■ ntlorlne, Morlnw, ts*lel«M, ally
liqnid, MrutiryinK at 4° C. to a CT7aiallin« uih*, Infoliiblo in wilcr, il Id soluble
in alcohol and pthcr. It cnnnnl he diiitilli'd without drccmpocition. It readily
fiMiit> witli putnssiiim and Midium ■nap*, which arc mlukle in water; it* coui-
|>nU[id> nilh inMl ulher baitea are insolubte. Il uinv be didtiogultbvd tnm tlie
hckU uf Ihe ai'etic aerlca by ila reaclion wilh N0| anu by ibe ch&iigM it uudcrgoea
wlieti fxpuMcd to the air.
Tbk Neittkai. Fath.
Thrw may be coniidcrcd lut elhent faniipd by icplnciiiK the cxirbnngcablc ntora*
of faydnigeii iii ibi^ triat/jinic akiiliut (•tyceriii (»M> below) by the acid radicle* of
the aci>tk- and il«ie ncritv. Since tbere are three •ucb excbangesble atomiof
hydrr«en tii glycerin, it in powiblo in fiirin three claMCi of tlM«e eihera; only
thoae. noircTrr. whi<'b l>cl«n|[ to the third cIom occur m nntiirni conHiituents of
the buraaii body; thune of ilic lint and aocoud are of throrvllcal impurliineo
odIt.
Tliey poMeM certain general cliaract^rtstlcu. Inaoluble In wat«r and cold alco-
hol, they are readily imluhle in hoi alcohol, eiher, chloroform, etc. : (hey alM>
dinolve one another. Thi^- are nrniral bodies, colorlen* and luielm when pure,
are not capable of being distilled witboitl underKoioi; deeompoiiilion, and yield as
B reaull of IbU deconipoaitioo solid and liquid hydrocarbonii, water, hliy odds,
ami a peculiar body, acrolein, [Glycerin cootaina the eleinenls of one moleuule
of acrolein and twn'molrciilceof waWr).
They pouew no acii'>n on poloriied light.
Tlirr niiiy rrndily be dccompoocd into };l>'<^'<'> '"■'l their rcapcctire fatty ocldi
by Ihr Bcliun of eaiiitic alkali<« or of auperheulcd «lcaiti.
PalmitlB iTri-p.laiitfn). ,(!*|P|i, }0p
The foUowiug reaction for the fofmalfon of this £at b typical for all the othen:
Olyowtn. Pntmllle adil. Pnlmlllii.
Jmiiin in allgbtly ooiubie in cold alcohol, readily to In hoi alcohol, or in
ether: when pure it crystalllies in One needles; if mixed with Ucaiin. il gea-
erolly formii shapelesa liinnp*, nhhnngh the miilure mnT nt titnen oMOnie a cry*-
tatline fotm, and wa« then rt^nrded a* a diMinct btidy. niuuely, maripirin. It
ptHBCMea three diflero-nl melting pointa, acoordiiw h> the provioua tCRi|i«raturei
toMhich it bus been aubjected. It •olldifle* la all caeca at 4A* C.
/Vipom'iun. Piuiu palm oil. by remoTlng the free palmitic acid with alcohol
and oryitallising repeatedly from olber.
St««rbi (Tri-itcario).
*^Yy»}o.
This is Ibe hardmt nnd Icnot fiiiiMe of the ordinat; fats of the body, ia also the
ImoI aoluble. and hi-iK« i» the lintt tu cry>talliic out frtiai sulutiooa of the raised
fatji. It crytialliiM usually iu nqunre tablca. It preoenla peculiorltka In lu
fiiiinK iiointa niiiiilnr to choee of palinilin.
l^fpnralwn. From mutton suet, il* Mparation ftom palmitin and olein being
effected by rcpeiiteil cri-sinlliuition from etlMr, •l«arin beii>K the leaat eoluble.
Oleln (TrUoleln).
<^*-5TrM«-
[t ill obtained wilb dilBcuUv In tb« iiufe alat«. uid U tli«a fluid nl m
teingwruiuFM. Il Im mora soluble th»n [lie two providing ones. It rrsdilj
fon oxidation wh«n pxpoMxl to tlic air, nnd !■ uonTcrl^ by inrrr iracM of
into a Nolid iiiomocic fat. Olein yields, on iItjt diittilUtioa. n I'lmrnrirnitic mA.
ihe idbiicic. und it mpunilled with inuvli KK«tor difficulty tbuo arc palmilia u4
■tcarln.
Prtjuirriiir/n. From nliv* oil, either bj ooUrk to U° C. Biid praning out Ui»
olrin tliat tcnininn l^iiiH, nr b.r dimolvinjc in mlcoDol «Bd cooliof. *rh«a thf «lcln
rviBiiiiu in noluiion wbile the other fmu crvslnllitie out.
Qlyoarln. ^^f^'IOr
ThU priodpiil cuu!<tiluenl of tbe nculml f&l« may, m above (Utied, be toatgi
upon a» a irinlomio ak-ohol.
When pur<i. glycerin in n vincid. colorlcow liifuid, of a well-koown aweet fate
It i« koluble in wntor and nlcnhol in nil pnt|iartinnii, inioluble in etber. Esf>w4
to Tcry low teniperaliiri.' it becomca alriimt solid; it may bi- dia>lil]<Hl in d«)
vcaielB without aecomiKisliioii, betireeti ■SlT,''--'m'' C.
[l di»solvea the HltcallM and alkaline «<arihii. hIho many oxld**, uicb h thaw if
lead and copper ; many of the fnity acidx are aUo soluble in glycerin.
It nnnewni no rutatorr power or nolaiixL>d liKbt.
It \» eaaily recogniiea by ita ready sulubiiity in water and alcubol, ite LdmIii-
billtv iD ether, lut «weet ta«ie. and ili reacliou with boM*. The prodottka if
acrolein n alio characteriHtic of glycerin :
Cjafi^ — 3H,0 = C.H.O (Acrolein). |
I'rrjjaration, By laponificAtion oftlie variocw oilt and (at*. It U alao (bniicil in
Ninsll qunnlitiM during tbe nicokollc reimeiitatliin of eucar.'
Soap*. Theae may be formed by the iwtion of cnuMic allEnlic* on fata. Tk
Erocew consikia in a >ubtitilution of tlie alkali far tlie radicle nf glynoin. iW
titer combining with the elemeui« of water to form glycerin. Tliua :
I
TrfMcarin.
*''i'ffr'*}°'+«"i
airwnu
FaneitatJc JiiIm can *p1lt up Ail* Into jljoiinu •nd trr* bttr an<lf,aa4 tk« Uto likn***!
tobocapablsoriBponilS-inglhcH 1iMyaB\Ai. Thb amouutor «Mpt Ibrwieil la thaallnuivT
taoal It. boir«T«r, Bmall atiil uiiim|nKiiiil.
Acids of tue Glyoolic Sskigs. \
KunnlDg parallel to tbe mouaiutnic aloobola (OH,ii-,0) U tbe Mrlea of
diulomfo auonola or glycols (0>H.4,0,;. Thu« corresponding to ethyl akobqlh
Aa front the Bumatamie alcobob. H fhm dw
tlie diatomic alcohol, elhyl-gWcol.
glycil*. atuda may be derived by oxtdntioB ; from tbe lattev (glycAh),
iwu lerim of acidii can be oblaineil. known reapeotlvely aa the glycolic and
•eriw. 'llie lir»t Mage of ojtiduiou of the gtyeol girai a mevber uf the gl]
Mvlea. Thiu :
EUiri-|lTO)l. aijMlleavld.
C,a,U, J- O, => 0,n,O, -r- 11,0, or mora geBerally
C.U^ ,,0, -t- O, = C-H^O, + B,0.
By further oxidatloo a ueoiber of the glycotlo terka can be «Nirert«d
member of the oxali« uriea. Thiu ;
atjcuUc add. Oxalic mU.
^t^fii + 0,* C,ll/>. + 11,0, Of motagaotraar
CUlyU, 4- O, -OU,— ,0. r U,0.
TIm acJd* of the glycolic terim are diatomic but monobaalc : Iwt tboae uf tha
oxalie aerin are diatomic and diabaajc.
■ hiHui, Altn. A. QuiQi. u. I'lurm., VA. ctL S. »&
CHRMICAL BAEIIS UP THE ANI»,V1. BOllV.
IiHl
I
r
Th« followliif inblp may ht f it^n li> show tlio gtoonil rvlalMxialiip at kIcoIiaIi
■ntl acids:
■MblK
H*ih>t (CH.)
Bihri (C.H.)
Pfopyl (CH,)
Bniyl iC.H.k
AlmlHl.
c B,[or[)
c,ii,(onj
C.H^OB)
A«it.
atjttit.
icUL
Artin.
Formic, i
HrnO, 1
Cubniilo.
H.CO,
aO,ii,o,
IK'.II.O,
Oxvbuttrk.
ca.H,o,
A<-Hio.
m-.n.o.
Pmiilonk
Ouiyrtc.
HC.11,0,
Rtlivl-Elvcnl.
Pfoi.vt-cWpo^l.
f.rt.ioii:,
f.il.lOU),
o'lniiii.
M ■Iodic.
U.C.HO.
auo-lalc-
H,C.H/.,
ObvcoLio Acid SBKin.
IaoUo odd- C,H,Or
Next to rnrtxinir hckI, t1i« moat important mmbtr of this aertv*, m IWr us
B phy*i'ilo|C}' b I'Mncmircl. i« lactic acid.
L«ctic iit-i<l exiala in four Uomeric modiflcaiioua. but of ibew oaly three Iia^d
b««n fouiiil id die liutnaa boilj. TliMe'lbtee nU form afrnpy, colorlem fluid),
soluble in all pro|>rjrlinDK in water, nlcobol, snil pltitr, Thr; |kii<i<«is an inlt-nstly
sodr laMc, and a slTinji ncjil rrnction. Whrii liruli-d in iii>liiii<in Ibry arc par-
»luilly illstillcd uver in the tvcupiiig vapor. Tlivy funii salts with iiicialii, of which
*tboM: wllh the alkalies luc very iN>lubl«Biid erytulll^e wiih •iilUiulty. Thr »!•
clucii aud dnc salt* are of the )(r«atMt importance, u* will bo Rvoit iatei ou.
1. Bthytiden« t&otic scid. ThU I* the oHlnarjr form of the acid, bblAined an
the charscterttiic product of ibe well-known " lactic fcrraeninlion." It wcurt in
the eoiileiitu of the fttomacb aad ioieHlinca. Afconlinu to Ilviti(i>' it ta fouiMl aho
la anaelcs, sod according to Osclivldlcn' in the ttHnfC'ionir cell* of the gray aub-
stMica of tbs brsin. In many db<*M* it i> found in ntinr, and exiiiti to a large
atnouBl in this excretion after poiKuninK by photjiboraa.'
Il tuAy liu |n)i*rT>l I>y ihr nnxnil iii*Ihu|j ot iIawIv oiHlKlBt lb* torwificwJiiij icty««l
or by aMjDf np nioDichlvrina'vi prMiioaLo dckI triili nioiil irlvrr ojiliie. In ubtAtuIng it Frcrni
Ike proilail* i-f Ui'lic frni-vuTnlinu, t)ii> i-r<»li> •'f liiir brlat* nr* piin(ii<d 1>y M vstsl ttytlaMi-
aaiioai^ aiul III* aeid lllxmliil frt-in tli* «>in|wuDJ by il>o ni^lioii of iulphur«ll«l byilNii^.
-■ Ethylene- 1 kCtie ketd. This acid is found accompMnjing the next to be
daacribrd; in [Ij<- watery exinurt of mtiaclea. From ibW It la seinraird by inking
adrautagc ut the <liiri'rf<iil Kilubllities in alcohol of the zinc taliii of the two ncida.
It aeenu probable, however, that it bun not yot been prepared in the |>are slate by
this metlMMl.
Wi*limi«t Anl nhuinvd Uii* a<id by iiMlias hydroxy cya«i4« ol ethyltoo with aqwcoM
iolBlioa* of ili« allmlln
The BtiiM ohnrvcr luaa'l II alK> In laaoy palbaloglMl OoUl
S. Sareoludo aoid. Tbi» acid baa not yet been [>mciired syntbettcally. As
Its name iini>lie». it I* thai fnrni of th« acid which chiefly occun In muscle». and
h*DC* axi«la in lance auaiiliticK in I.eibie'* '' extract of meat." It in often found
kIso in pnlholngicrHl HuidH. Thi* in the only acid of the Hcrini nhivh poifwca
MV power of rotating ihe plane of polarined light; it is otbcrwlM Indiatincuiab-
ableiroTa the preceding elhylidenr-lnclic acid, and la generally rciHwseiited oy Uie
nme formula. Tlie Irce ncid ha* desiro-, the anhydride livio- rotatory acUoD.
TIm specific roiatiiin for the tine salt In solutfofi i* — 7.0d* for yellow light.
The tine and calcium talla for aarcolactic acid sio nan soluble, both in mter
I Aim. •) <. tunn. a. I'litnn.. IM. ftitl 9. KR.
' nuxi->-i Anhir. Bd tW. fina-tiig. iTi.
■ Mgliiiliicn inO lltvs. I'pbtr M«l« rhMiihomiitniilaB.
• Ann. il i7i<M II lluinn,. llil.essttll.B.a.
t'faM. Oniialb.. iMk f- an.
1012
APPBNinx.
aikI klc»hol. Ibnn tlioveofctliylideni'-lAclic acid, but I«miu iban Ibmeof filifl^
lactic ■uid. kdiI the Mme njilbi of ethylene lactic kM cootAiu morr iiaufi
cr7*ullluili>n ibau tboM of ibe oilier iiro.
Ilnnti' hu uodipirtd the alioie acliU li> Uie laoiitiaainu atp»\>U of «xihui( Iu iMbnr
ICinlraiTytla luiiil, Ih* Tuuitb la thU Hirlca of lutle iDiib, It i!i«litt|iii*lukl hf lb* B4ta*( <l
IU ilcuMnpMiliOin on hoaliaj^. It u n«T«r ouod m a naaMltumt of aalotal badlaL
OxAUc Acid Ssriih.
Oinllo add. H,C,o,,
Id ili« fret MaU thU arid <lo» out occur Id the buraan bod;. ('«totc oxahb^ !
howevMT. iflB not unfreqnent constlti)««il of urine, und eotem into the ooa)po*iti<a
of aiany urinnrir cnlciiM, Ibc «a>oall«d mulbonr caleulua o^nilatiag almM
entirrlv of it. It may occur in feoe«,and in thv ^U-bladd«r, lbou|[h thUbnr*!}
obarTTi'd.
Ah ordiuariljr pccipltatcd ftom solallnns of calcic ulta br ammoalc nialok,
calcic oxalate in i|uiu< aninriihowt. bit in iiriiiar; dejiMlta It BamiiMa a MtiHi;
cbaractcrialtc cryaullino form. viz.. tlint of reclAngalnr odobvdnt. In •ocnv tatm
it preacolH tbe anoinalwia forrna of niiindud lumiM, dumb-bell*, or aquar* colamai
with pyramidal otidit It ii insululile in wabtr, alcohol, and Mbpr. atw) in anoi'inu
and ac«ticacid. Mineral ncidsdlaM>lv« this nail r««illly. as aJxi to a imallcr e ti*»i
do fliilacionii of Rodic phonphate or urate. All the above character iatica tittrt vt
detect ibi* salt; ita micmacopical appearance, boitcver, ia ^nerally ut miNt fw
for this |KiqH>te.
The pure acid ia prepareil either bv oxidizing sugar wUb nlutc Add, ordrtiM-,
pueing ll|;neou* limue with ciuMic alkali*. .
Sucoinlc add. H,C,U,0,.
Thin in Ihc third acid of Uie osallt' iterfea. being leperatMl from nxalic a<-id bt
tlM! Intvrincdiulf maloulc acid. 1I,C,H,0,, It occur* in the iplecn, the Ihymiu
and tliyroid huditH, bydrooophallc and hydrocele Buida.
AeeBrtliaj to Mbihdct a»d Stiepanl* It li (buBd u ■ anrnul cantUliwal nt iirin*. TV* •
cael«alad 1^ Salkixrtlli.'and al« \>j rmi 9pej*t. It H«nu prabaUc, hnwwiar, llkM •t*r*
trinM aed nmaal«d linnan eoutain (ucelulc aeid, ami ihli laiwr fa— ii iinohaii^nl iiib &■
iiriuB, tlwl it may lli» tw onMAJunally ]>mviil in Ihii exrratictt,
Succinic acid cry^talliM* in Isrtre rhombic tahlea, also al lim«i in the furm ^
Iftrgepritmn; tbcy'nrc xiluMe in ■'> partaof mid water, and 2,2 of boiliofr, ■licliili
•olablo in aki>hol. aitd ulmunl iniuluble in rther. The rrtMaU tneli at iW C..
ud boit at 2-'ti>* C. being hi tbe miuic time deconpowd iiiio the anhydridt lad
water. Tbe alkali salts of this acid ar« Milubte in water, inifiliihle in alcohol a^
elber.
Prfparttlioa. Apert from the avfithetic aielhoda, it maj readily be obtiUnei kf
the frmicntntion i>f calclu malnte, acetic aoid being produced aimtillaneoualy,
lu pn-MDce \» revuetiiKd by the tnlcroscnpic cxamina^ou of it* cryataU, ■»!
lla characterlalic reaction with nnrmnl lend acetate. With this It girt!* a pftdid*
latccatiil^aolublcin exccmof the precipitant, but vnming down again <in wannu{
and ahiking.*
OHOLKifTltlllit. (CmH^O.)
TbU la tbe only alcohol wfalcli occurs in ibe human body in Hat ftt%
(The trlatomic aleohol givcvtin in almotit always found combined na in the fiu;]
and cetyl-aleobol, or »lbal. U obtained only from •pormaceti.) It in awhile t-t^i
lallins bodj, crTaUlliilng in fine necdtea frvun itaMlatioB in ether. chl<>rt>ronB,i
' WifU(i-tii». OK (41. AIn Ad«,4.CIimb,I|. riMrm,. Ihteldte.)! M.oUrflLftl
■ I iii.-(v<)c.1 ubRd- Eiinuh.41. niinni''*- llniiixi-r. two.
■ I'llDni'i ArrMr, Od, U. ^IMH, H. W, 404 M. Iv. iih:)). & M.
■ |-<.rllinlwrraniciiUnMeMMBHa.»p.clu<utd Uvlniai awl MIS'. K<*iw1ir. t rat MbI ^U
twnml : tr<im iu bnl alcoliolic xolulionii U i« i)(!poiiit«<I oa cooling in rliotnbic
ubie*. When driird il luelu ul l^^i" C, urn) disiiU iu cloacO vaueh ni ;ti>0*C'. It
I quil« Imolubic in water and cold dcoliol ; Bolable in •olutJona of bile miIu.
^ eoluUon* ordiolntenn pomeM a li^fl-lianilvd mUton Klioa on polariiced light,
of — SS' for Tflliiw Ufcbt, thin being iniicpendoot of (vinoentmtioD ■nil of the
niilure of lh« Milvvnt.
tlenteil wiib airuag tuliiliurlc AL'id it yieliis n hydrocafbou ; vllb concentnied
nitric il itivm choleileric acid and itthot prodiict«. It ie capable of uniting with
■i-idn and forming compounil otberx.
Cbolcsterin occun in small iiiiantitits in tfae blood and many tinauea, and ia
pKwnt in abundance in ibe wtiitc luntier of tbecerebro-tplnal uxin and in nerves.
It la a coaaiaot conatitueitt of bile, forming frequently nvarlj the whole maaa of
•onie call-Rlon«e. It is found in many pathological fluid*, hydrodv, tbo fluid of
ovariaTryatc, etc.
IVtparation. From nll-sconea by aimply citractioD witb boiling alcohol, and
aeail with ak-obolic putaah to free frriin extraneous inalter.
Aa tMl« for (his Bub«Eance may be given: With roncentratod eulpburic acid
and a little indine n violet vnlor ■■ obtained, cbnniiinit throush sreoo to red or
bine. TliiK U anplicablo to ilic mU'ronfopic erviitalii. AAer oioaolTiiig in chloro*
'sriB a blood-red aoluiion i» formed uu tbe addition of an e(|^ua] volume of cooceai-
nled anlpburie acid ; this aolution if exposed to the air in an open disb turo*
Ant, ^nwR. asd Rnally yellow ; the sulphuric acid ondrr the chloroform haa a
Krcpu IluorcMcnce. After cvapciration tu drynow vith nitric add, tlic reaidue
turpi red on treating with ammonia.
TU* body II lUwrlfaHl ber* tailier for Ilia wke iircanTaDlonce than l^m II* pnaMslai aMy
cldM rclatkoiikip ta Ibe wbitjiiiau I in mediately pracvding.
K OQMPI.KX NtTltCHtRKOI/H i-'lTH.
" LMdthin. C„U^NPo,.
Occtin widely ipread throughout tlio body. Blood, bile, and aerou* fluid con*
tain it in amalt quaniitleii. while it ia a couapicuoiis oomponent of the brain,
netvn, yolk of egg. bcmivii, pu*. white blood-corpuaclea, and the electrical orgaiia
of the ray.
When pure it ia a colarIni«.»lightlycTj>ta11in««ub«tauce. which can lie kneaded,
but often crumble* during the procew. It ia readily aoluule in cold, exceedingly
•o Id hot alcohol ; eilivr diMulvea It freely tltoogb in \vm i]uantitiea, ua aUo do
cbloroforn). fata, benzol, i-ur)>on dl»ul|>hlJo. etc. It b often obtained from ila
alcoholic a'tlutioD by evaporation, in the form of oily drop*. It »wella up io water
and in thi* (late yields a flocuulcnt procipilale with lodiuin chloride.
]..«!iUiin ifDoailvdeoompoaed; not only doai tbiidecniniHiiiiiiuu act in at 70* &.
but the i>'iluiii>ns,lf merely allowed to aland nt the ordinary lemiwratnre, auquite
aa add r«actloD, and tbe tulmtanci^ is deciiniiicHeij. .^dda and alkallea, wf courae,
«Abct thia much more rapi'Ily. If heated with baryta water it is eoBipl<4ely
dMOinpoaeil, tlie produrta bring m-unn. glycerin ■phoapbimc acid, and Iwrio
atMiato. This may be thu* nipruienleil :
r
c„iVNPo,*8n,o-stvi^o, + c.ii,po, + c;n„so^
When treated in an ethereal *olutifln with dilute aulphuric add. It Is merely
eplil up into nenrin and diMeHTyl-glrccrin'pbmpbdric arid. Hetica, Diakonoir
regslda lecithin a* IhudintHirTl-glyrerin pbmiihaleof OFUrin, two alMttaof hydra*
■an in the glycrriD-ph'MplioHo aviii beinjc n-plnce<l by tbr radicle of Stearic acid.
TtaMwaia also that tliere iirwbnbly extft otlieranalogoiu compoonda Io which the
radiclca of oleic and palmitic acid* lake p«n.
Prtfiaralioti. Usually frooi the rolk of egg. where it occura in union with
vilclho. It* ianlation ■■ com plicated, and the reader i* referred to lloppe^iaylar.'
■ Ham mylW* Ucd. HiPtD. I'liUHudi., IMt ILOMTi, axn: Hell. UL aMl. & «■ CwlMlk
f. d. MmL mm. I IIW>. M. I. T R. >.
* iMXVm. i-iuHiiicu.. iMi n. yxmy, tm.
KH-i
AITBNIUX.
aiyoarfn-phosptaorio acid- i-,H,ru,
Occtira lu a procIiMt i>r ihv ilnromprftlilim of leciibin. an<l benro U liiiii
thoM timuoa uid (tnids in whidi Dib IsUor la I'rMotil : in Ifiikhtraiia lti« ivin* W
■vd to conlain thU ■iib*t>ncc. It hiu not boon ufabLinoJ in ditt uiIhI furm. It bit
Iwen proiluccil ityi)ili<.-ttcuIlT hr braUng glyccrio aiitl gUciiti |>ho>pbi<>i(- itclil . it)
may be Tey:ur(liMl aa furiiioil \sy tbe uninB of ofi« molecuk uf glyoerin wjib w
|iko«{iliuric acid, with elimlnaiinii or one tDoln!iil« ol «nt«r. It in a iiitHiii«|
itflMilta with liariuni sDil caJutucn nrc iiualublo in Htcohol, Mlublc in n>1i) -
Soluti«D« or il« nlt« arc [>f«cipilatc<l by Iciail noetolc.
Prota«on. l<-",«ir„N,PO„T)
A cryiitalllne body containiug nitr«f«o and pbMpborati, obtained hr LkIift^' '
rrotn tlie brain tiibHtiincc ami rcgnnlM by hliii lU (u j>rinci|>al con*'-itii<-Di. Tia <
rcMAfchca uf ll(>p|i^K<^lcr anil Diakoiiov icoil«d tn abciw ibat I'mUff •■> •» {
iDMdj a mixture nl lecilbtn and c«refarin, A rvitrlilion of I.iirlireicb ■ esnrt-
lueotB ban. buwever. led QamgtiF and HIankenbotu* to confirm ib« tititb oflxij
result*. I'roTafciii] appeura to wparxtc out frutu warn alcohul ua pwlual cuullBfl
in tbp (turn of very "mall (i««dl<«. otitn arntngvd in ^root'*- ■< ^ aliftbUy •iii*k«^
in cold, more Milublc in hot alcohol and ctltcr. It U lluwIubW in ciilnr. but varib
Up and foruii a gdatiuuus uiaw. It tuelt* at 2D()° C, and funnn a bruotn q^rufj
auld.
/Wjyiriiiioa. Fladr diTidcd bruin ouUlaove. frtcd ttwa blood and cnniiKtii*
Umuo, U digMlod at tS' C with alouhul !S.^ per tuot.) as long ■« Ut* alMMl
cxlTBCta anrthing frooi it. TboprotasoD Khich M-pntnits out frunktht fltiraM ll|
well waabed viib e(h«r to wrt rid of all chi-tntcrin and ulher boilln aotvUt I
ether, and Anally |>utined by repeated crytullijsatloo from warm alooltol.
Nourln (Cliolii.). C,1I„N0,.
Dincovcrcd by Si f« lew* in pig'a-gnll. then in oi-^hII. It d(>« imvI •■CL-ur Uj
ihe free stale except an a product of the drcompi-iiion uf lecithin. It u a enli
lew Huid. of uily cuixJslence, poMmut* a «ining nlknliiie rravtiou, and forsw «Ml
aeiiln vvry deli'iucaccut uilis. The nulla with bydmcbloric acid and dio ehl«(1d«|
of tiliiiinuiii and gold «re ihv miiHt ini|>oriaDt.
hcnrin in n oiimt uohUblc body, ineec bcaiinK of iu nqueoiu aoluilon isAda
In tjilit it up into itlycol, triuietbyliunin nnd cibylrnc oxide.
PrtpnrQtion. Krom yolk uf c^ig. fur ihiii nee Diakoaow.*
W«rt<* bu otMiuixl >l *ynlliall(Mlly. Ilr*t liy llio wtlon of clcol )iy>ir< -
iMlkylatoiii, and liicn by ifa&l ot nilirkbc nxiik and <rMar on ll>« mih'
aboes. McfUiar rlUi llin m^ife nf iu dKViDiiiailinn. loial to Uw Mw tbn-
r*s»nlnl m iriowthyl-oxyctbyl-aiBinoiilBn Iiy4ral«k »iCIf^i(OIltO,OIt.
C„n„NO,(T).
la found in the axia-cylinder of nervea. In ]>iia cornonclea, and lareeljr in
brain. In former timm many name* Here eiven to the «ubMance wheo b «a
impure itate, r, </„ cerebric acid, cerrbrole, etc. W. JlUIIcf* fintt prepared iiia
Hie pure fonu. and couairueied the above formula from bi* anidvaii ; the mean 4
Ibeae ia O, 15.M i H.ll.i; S, 4.5; O. «S«. Great doubto are.'howrver. thwHi*
npon it» parity by the reM«rchea of later obwrvert. Accordlni; to Liebreich ' and
Diakonow.* it is a gluooaldr.*
Cerebrlii is a llghl. oolorlea*. exceedinRlr hygroMtipic povder. which owelU
BtrOBgly in water, slowly In the cuM, rnpidljr an beuing. When heiiied to N**
it tum* brown, and at a woiewhat higher temperstare mcltti, bubble* up,
linally buniH away. It ia inwdnble in cold alcobid or ether ; wana alcobol
- : UI.I >nura. uT rh)>l.4, wL n. fUOW. k IU
lul- eilvlO. £. W.
'(K
V irtMIOn. m.
'All- _ , . AMI.
> atttk (. isilixl, A>«' IMITK
■ fMlnlti. r <l luBt.v ... T
• e*»aW(itc«lw8l9>£. ......:. : vIiTiM. rlMB.. Bd. 11. nS?l|i. & XR-
:l
NITK<VJKNOUS MRTAB0UTE8.
TiiB Vhbk Gbopf, AutPRs, Avn SiJitMK BontKS.
CUKMtCU. BASIS or THK ANIUAI. B01
•oItm it Mniljr. llut«d vrilb dilatr mineral ncitt*. ccrabrin jiekb • Mgu^llkt
bod*, puMMsint; l^ft-handed rotation, but inoapuble of f«nnciilati<i(i.
B /*ipitratiou. For iIiUam VV. MOller.'
' The cUi«r CAiuUtu«tit of nufmal uriiw in m kin mall* anit none trtb«r Aiiimnli ;
th« uriiM of birdtt aho coDUiiimi nmiill aiiiuuiil. N<irmiil blood, KraOH fluids,
^Jjmph, uid the litvr, all cmttain the samv liod^ in tnioc*. It Ja nii4 Touad in thv
^■BiiHclcs •* a normal coiwiiiupiii, liut may tnaku ita apjiriirHnvv lh«ro aiidor ucrtain
^KfAthological M>ndilioaa.
Wbcn |iiir« ii tryHialllMa frnm n nmcvutrnti'il tuilution iu ilie form of long, tbia,
slitWrinf niwdlM. If depottled aluwlr fnnii •liliilc «i>lutione, Ibo form !■ tliat of
Imir-KidM priuiiiwilh pyramidnl PiidH; lliMC arraliraf* antiyilrmin. It puotFioeB
a aomowbHl bitter coolinp taitr, likp uiltpclrr. It U rrndilf soluble in •nUvt uud
alnltvl. Ui«»nlution« boinff n«utral. In anliydroiu etlMr It U Iniuitiibtr. The
CfiUU may be heated to ISCi'C. witb'ml being decompowd ; at n higher tero-
peraturf ihry are tint liiiiiefled Atid ilien decoiii|>w«. .Imvlng no rwidue. llM[«d
irilb MrDntl acidi (ir nlknliiw, cleroniiinititinn eriauea. thp final prodnctn being rar*
bttftte anhydride and ummonia. The larae d«cuinpaaiiion tnay al»u occur na
Ui« rettult of the action of a HpeciGc ferment oo urea In an aqueoiu aolution.'
Nitrou» odd at once decompowi It Inlo carbonic anbvilride and free nitrogoa.
It readily form* compound* witli acids and basaa; of thaw the rollowiog an of
ImportMice:
Nitrate of tiroa. (NII,),caU>'Or
CrfBlatliiea in nU-tilded i>r rliombic tablea. lnaolubl« In elh«r and Qltrlv acid,
•oluble in water, aliiililly toluble iu alcohol.
Oxalate of area. [(NH,>,CO],.H,C,0. 4- H,0.
Often crrstalliijea in long thin pritiiu, l>ni under the microscope i« obtained in
a fortn eloMty rcavmbliog the nitrate: It In alightly aoiuble in wat«r. low so in
alcohol.
With mercuric nitmlt! urea yieldi three Mtta, ooaiainlnfc. rcapectlvely. (aar,
three, and tW'> e^iiiiatt-ntn of mrrcuric oxide Ui one of iirefl. The flnl i« tlie pr»-
clpitale formed in I.iebig'e oiian illative detemii nation of nrea, and may bo rvpra-
•Mit«d by Iba formula; 3N,H,GO.11k(N0^»1IkU. Tbo ucavt conttituiioa of
tbcM aalto baa not yet lieen ueMrminea.
/Vr/mrafi'm. Ammonic Mil|>iiHte and pulawc cyaitat« are mixed tucMber In
a(|ueuu« »o1utioD. and the mixture ia evaporated to drynea*. The resldiie, vben
extracted irllh hIiiioIii[« alctiliol. yields urea. Fri>ni urine, either by eTajMiniliiiK
to drynm, buvini; prcrinunlr pr<<ripiuted thp urlni- wiLli nncmal aixl bwir lead
acetate in ■uccnuion ami removed the lend by lulphurcllvd hydrogen, and then
exiractiDf with aleoliiii; or wocenlralin); only i« a ayran, and then fortnine the
nltr«le oi urea; lbi« ia wwhed with fiuri nitric acid ana decompoMNl witli Miio
carbonate.
Iletection in tolDtiooa In addition to the m[cr<k«i-opic apnearaDce of Ihe
rry^laU i>V>tiiiiirit on (-v:ipurati»n. tlie nitrate and oxahitr *hi>uln be formed and
cxumlued. .Vnolbci piirt nhnuld girr a precipitate with nirrcurie nitrate, iu ibe
abivnce u( vlic chloride, but not in tlie prMen«e of this luKt unit iu exceu. A
ibini portidu is treateil irllh nilrir add containing iiitniu* nimc; if nrea la
present, nllmgen and carlmnic nnhydride will be obtained. To a fourth part
niuifi acid in exceM and u link- mercury are adiled, and the iniitute li wnrmed.
' Oil, (II.
• Miseiilw. rHnctf I AralilT. M. xU. (MTCk. %■ 311. Ail»cl>. JMMeh. t. pbrMol- rim*.. Sd. T. (tan).
J
IMA
APPKNtllX.
la prtMOfVof nr«a « niAjricM mixture of ^Ror* (N *i>d iX>,> ia ^t«ii «£ A 4
pnnion bWpi in«}t«d forMutne time, dimolT«<l in WHicr. nml ctipric Milphaicud
<aiutie tods aiv »diit4 : ■ n^ or vitilct color, ilui* to bitirel, it ilervlopM.
Asantitative detarminfttioti. For iliis codic Hicdsl mann*! mihc U ohi-
•nlted.' U nill aitlBcelMT* to point out tliat tli« detaniiinitioD k ta*A» aiifeir
with a Milittion of mercuric nitralv nf konwn ilrrnxlli lLi«bi|() ; bjr i
ibe aiVB by meana of «odi« hv|iobroai)i« into niimjteo. tuubouic anbvd
waiM^. anil mcaauriiigilio oilrocen (Knop) [N,H,00 -r SNaBK> = 3NHBr-
■f 211,0 4 N,J. or by lifaUng uto un-a wilh cauUlc baryta in a nealvil lul
urM Ming deicnninn) by ibi> ircj^ht of biiric carbonate fonurd I Itun*«« |.
Urea in g«ii«rally coimiiliMFil tri bo bd nraklc of carbonic aciil, >. r.. autaaaUt
Tbt aaiul« nf an add b formed vrbcn wuivr U r«mor«d from tbe ammuniuai nti
nf ibo acid ; if the aud be dibaalc and two uiolecalea of watrr be rnBorvi]. ibt
MBit k often trpokon of hi n dJamlde. Tbm if from ammnaic carbiwata (NH,',
OOhtwo molcculn '<( water, il),U, be remored, cirbooio acid bvioj a dihMc
acH, the result U uroa ; than :
(SH,),(X).-2H,0-(irH,>,C0,
vUeli uaf be wrjtteo dtlier aooording to tlw anirooaia ty|ie aa
COl
COl
Br a»
^{su'.
iw<> atouk <»f amidoceti (NH.) being Mib«tiloled fur two atooM of bydmnl
(HOI.
TbU connection betwron carbonic ncid and iirrs ti sboirti by thit fad dul
ammonic I'aiboaalc maybe tbrmed out of urra by hydration, a* when tim a
aubjMlcd ii> ilip ipeciGc ferment mentioned abuie. Re^'Ied. ttien. a> a diuddi
of carbonic acid, urea may be itpokea of a> carbamide. Hut ibc tlio.n-tlral dnf
■ralioa nf itrca I'luru uinmunic carlmoate by debvdratioa cannot he mllael n
imciii-e, kberciu urra can rmdily be foTtncd iVniu ammooic rarl>aniaie, aad,
Ktillie )>■ iuclined to recard it, ncfl as Ibc diuiuidc of carhiinic acid, but IfllHH
aHiid* <>r carhuuic acid. Ammooium cnrtiikmutc. <-'0,Njtl, Mtmn ll,4t.^^^^|
urea. t-X>, >',, H^ — wblcli. if carbamic iieid be written tu tXt. <)H. NH,. m^^^H
wrlllan aa CO, NH,. NH,. 'rw alom of Hinid-ijEen bciuj^ fiibatituleil fur onr atmH
„f 1.. ,i,,,...i ,m,i not two. iia vhra lh« uiWtaaca is rrgardrd an derived tttAu at- ^
b-. t>r«-biier:> exjicritiiciits indloote a ready dcrivaliim uf urrn fm»
■uiL. .... _j.rtMaiale. lie liai obtained urea by the elecir>ily>iii of aaulutiou -i
tbu lalt with rapidly alturiuling current*, Ibiia removine tbe eletnenta '-f «aur
frum tiM oaibamate by auch alternating nrocemea of oxidation and retlactirm n
may be tuppuetd tn take place in the body. Tbe mution i« expreaaed aa M-
I. XHct>oxn.-(.o-iNir,.<'ouKU,— u,o.
2 Sil.COONU, , H,= NII,CO-NH,^ H,U.
ft'aaklya' aod Gamgte.* bowofpr, Hnee urea, wbco heated viih a laije exoM
of wnaiir iierwiuiKaDMte, ^ive* nd' all it* nilroKen in a fttv >tntr. ux] not In lb*
'al'Ii'*-) f-Mm of niiric ncid. iw do all oilier nmideo, conclude that it b ool an
■f that it ■■ isomeric only and not Identical with carbunide.
' itaui to remember tiiat urea i» also laomeric with ammooto qyanait,
. „ and indeed iraa llrM formed ariiAcUlly fay Wfifalei (18X8) tivn
..' three tMmefic eompnuodi'. auimonium
'Mill other in »ach a way that urea may i
-■ii<>( ii<>*." .>j"tii' 'Ilium cyanate or from ammonie carbamate, mxi mar nui. luij
...,,, .^, AiwlTw<)all»mKtULaat.TO.&»l
. nil. il. p. 'A
4
CnSVTCAL BASIS OT TRB A:flVAT. HOliT,
eaae be converted Into unmonic corlioiiiii^.* Now aren in n much moro
■bU body than ■oimonic L-]ranat«. mod In tin.' Imittrortiiftiluii of the Utter Iniu
tbr foTmrr tnrrgj is aot free : and it u northy o( iioiice ilint, ihongli tht ]>TewDc«
of BUl|ib('cvimi<i(a in tbe mlivm prntmlily indlcatM llii- cxi'tnnce of cyanic rr«iduot
In Ibe body, tlie Dlirocenouii product* of tiio ilccom|jnaiiiDn of pmUad* bdong
ehirflj to Ibe cIbm of amides, cyanogeii eonipuundH beinfr rare among tfaem.
Pflfl^r* lis* railed attention M the ereat molfcalar eoNgy of lb* CTmnogeo Mm-
ftMnit. and liiu :>u|c^e>tp>H that th« ninctioiul mMaboliran of jirotopuaiB 07 wblch
eiMCgy in Mtl Ctce luity be cumpared to tbe converaion of tbe eooTROttc nostabl*
(gruiegen conipouudH into the lew enemtta and mure liable amidea. In olber
worda, ammonium cyauato Is a type of livuif , and urea of dead nltroeeD, and tbe
QOnT«nnoci of the former into the latter i» an imago of the esacnltal cbange whicJi
talm place when a liring protrid di«a.
fW/tmut Hnoi. The hydtngca aloni of urea imb tv replaMil by alMihoI and aelil ndlelai.
Tharaulu ar» AjRipaiiurl <ir*Bii orunlitno when Ih* hyilruc«« it n<|ilBaH bv an icid rulicja.
Many if them tta mltcil nirldt. liurf ihn liy^lniflpti tit-m ibc amide gnnp, if »m all r*plaMd
a* ■bat*, (mil br r*|ilar«l by mdal. Tliu), tlie futntitutioii of oxalyl (oxalit aciili ^vai
IKintHOfe add,
fCO
y,\ H. nrC«, Nil^S.CAs
at iiMtntjl ilanroDle acid), dial uric arid, rx). Xlli, XAIIiOii of maaoxalyl imwoaatle
Miidl.allman, Ct). K)I^X.<',f<r TKmp boliea arc iDltratlBg a* l«ing aUsotrlaiaad liytlM
anitcial oaldalKiii of uric acwl. (f«« Wleir.)
XSt\o add C.H.N.U,,
Tbe chi«f cooatilueoc of the urine ia birda and re|rtl1ca; tt occur* ouly apar*
iacly in ihia excretion in man and mo«t mammalia. It ia normally oreaeBt in ifae
■plMD. and traces of it have bran found in (he tungti. mivcin of the heart, paii-
cnaa, brain, arid liver, ('ritiury and iriinl calrull iil\en cotuiat iarirely at Ihia
body or ila aiill*. In K""'- lu-cuinululiona of urir acid Mlta may occur in ratinna
parta nf the body, forming the «o-cnJIed goulv coucrctiima.
It is when pure n colorlcM, (Tryalalliiie powder, laateleu. and without odor The
cfTaUlline form U very variable, hut UKUally tends tnnard that of rhombic lablua.*
Wbeo impiure it cryalnlli/m readily, but then ]iiiwmura ayellnwiah or brovni»b
eolor. In water it ia very inaolubl'e (1 in H,OOU or 15.000 of cold water); ctlier
and alcdiol do not diaaolve it apfwechtbly. On tbe other hand, aulpburic acid
lakes ii np wiihoot ilecomfiosllioa, and li ta alito readily soluble in many aalla ot
the allcalits, is in the nikatics Ihenuclvea. AmnoDia, however, scaiMly dtmolves It.
84IU of uric tcid. of Ibew the moat ImporUnt are the add iiisiea of sodium,
potaNiium. and aniiiionium, Tbe sodium salt crT*>alIiz«ii in nanj diStfoot forms.
tbwe aot bciiiK chatai'icrlKtic, aince tbey are alm<at the same (br Ibe corroapoad-
iDt «>miMMiD(b of ibe other two baac*. It i> veir Inaoluble ia c«ld water (I in
lloOor 1300), moreaulubte in hot i| in VJA). It it ibe princlp>al coosUluent of
MTeiml forms of orinary scdimrni. and compoMa a lurj^r paii of many calculi ;
the eicremont ofanakea contain* It largely. The potnaaium rracmblD* the awlium
aalt verv cliwely. a* ahM> doea the compound with ammimlum; tbe latter occura
generally In the ardltnenl IVoni alkaline urine.
PrmiratuM. Caually from guano or snahe'a excrement. From guano by boll-
log vltb caustic i>n|a«h <1 part alkali to W of water) aa long as ammonia ta
vrolTed. In the fillnilc a precipitate of acid urate of potamium ia formed by
pataing a curmii of carbonic unbrdride ; tbia aalt i* then wathed. diwolvcd in a
eaiHlIc pdiaib. and decomiicned by carefully pouring its lululion Into an eiceaa
of hydiochloiic acid.
■ Tta iDllniriM IllcntniK ti iMCRattoa ta CMinertlan *lin l>» quealtua at \tit eyaaXt a* aaMi
~ ID dC una— lirMhul ^ Bh. li. k. t.tinall d. WliK'LflMs- Hlf "' '"'' <-^' >•■'' ' •^i"'«t..
LflUA «. KalwfaiD Zt.f Rk-l. KI i.il*;i'.R»X Mimk 7i
_. J K Mlkotnkl: Oiuniur.d. Mfl HI*.. in>. N.>. W; |(«>
R IM )WUch r fftvlfj <i^in.. IVI. I. llfTI), *t 1. U.JJ); UJ. Iv. ;.. .- ... .•_ ,-. l.i.— .l*-
Urv kk*.. I .■\y i'ti'.'].ibt ttii a*ni.g,i.
1 l'll*i[iT'" Ar.-1-jv, 11.1 1 [-l.'ii. *. MI.
• 8(aLuaw:>ii«ii^ K. Ii UaAnuiDiAllaadirllaniardlaente. WIen, Ua
1048
APPENDIX.
Tbe pratcoeeof uric acid Ik raconiied by iho Tollawlng tmU: Th* m
haviiW bMn oximiDwl niiemieapiflMljr. » portion u arnpgrdlvil mrrt'uttf tu
Don with Mie or iwn <ln>pa of nitric Mid. Tlte rcsidaewtll. \( atie add U firMvni.
l>e nf ■ red euhr. ir|iii-ti on (ti« addition of uhhobU tunu tu ourolt?. Thai li ih«
muivxi'le Imi. itiid cje|ieii<U on lh« prMBBW of slloian and nllnxantln (» thi
residue, ^'nllilf ' ha* EJven ■ dolical* rttwtion Tnr utic acid. Tti<^ •uhttuK* b
diaMlvcxI in itodio carbonate and dfi^pcd tin |>*pcT moiilonod «ii)i ■ aiUrr mIl
If urie acid be proanl a brown >uiu ii fririuc'). 4fu« to tlio rndiiL'tinn >*{ \\\r ci^
bonoM or silrer. An alkaline Motntlon it urii- acid can, like dext/nw, r*diM*
ciipric KUlphAlo, willt |i(«cipitaliua of ilic caprniia oside.
Uric acid naiais vorv la^ly lb« action or on>a »tinn% acida and alkitlln, >i>
lillnttng iu lliii rwl a innrki^ diflemictf from am. It mii^t ibrrcforr pi-rhi)*
be su[ipo4ud tliai urea residues do not jirKsiit in urio acad ; o«verttaeleM bjr 90-
dation uric acid doss five nsa not only t-> ordinary utm, but alsu. and al Ih*
same limn, to the compound urcM (iir«idM) Bpnken ofabove. Thus, by oxldMigB
irlth acid*.
nrl« acid. AtlMCSD. Vraa.
CjH.N.o, 1. a,o t- o — OtS,u,o, + cs,n.o.
Now alloxan, at waa atatod abr>v«, ii n Rnmpoiin') um. vti.. mmosalyl-vns,
and by hydration can b« ooavertcd intu aivsoxalic acid nnd urea, thua:
Alloxan. Afa^intiR. rros.
and by tlie aciloo of chlorine uric acid can be Hpllt U|i dir«oUy into a moleculsif
nwsozalic add and two mi»It«ulos of aroa :
i
Uric uid. Mtaeialip sciH. UrM.
UiH.N.O, + 0', + <B,0 — C,H,0» + XCN,ll,OC + 'JUCI.
i
By uxidation with alkalien, nric acid is 0(Hivert«d into allantoin and caiboab
add, '
Oric adit. AllsnloiD.
C,H,K,0, + H,0 + O - t.li.N.U, -f CO. ;
and ntluntuiii, by bydnttiun. beuomes allanturic or lanlanuric acid nnd nrw,
AlUbtoln. UrM. Allantiirlo and.
c,ii,N.u. + 11,0 =cn,N,o 4. c.u.x.o.
Now allsalurlc acid \* a compound area, with a residue or glyox^lle add. Rr
Other oxidations of uric acid, imr.ibanic ndd (oxulyl-uroa). osalnric acid (wbitft
is bydraled parabanic add), sud diuturic ndd (tartronvl-urcsil arc oliiuinrd. It
fiui, all tfaeac dccorapoalliomi of a m'>tecute ni uric adtl lead to ibe i>r<idiii-t)nt» tt
ur«a und of a carbon aoUf of aom« kind or other. The reluiion <>/ uric acid u
ureL as illustmted by Ibe above rt>Hclioit>i, is brought very pmoiinenily into nr«
by tbe arnlheds of urir uoid which hiu' r<-ooRtl^ boon fvtUtftarA,' It ts obtalnei
by nmplyltMing loKclhrr £lyci)i:iiiv(auiido-aoeiio acid) and urea at n t<>iBp«nt»»
or S00^-33iy C The converse fonnutioo ol' glycocine from uric ut-id nitb iW
•t mill taneoas prod notion of amntoiiis and carbonic anhydride ha» lieeii k.n'>wD far
aoma time. Since in this Iatl4>r reaction th* nRnnooU and <-jirbonic anbydn^
are in the proporlion* in which they would be obtained fr-im ryanii- or ryanarir
add. uric ndd bn« been rennJed >■ Ixiilt u|) fri>m midui-n of cyaniiric acid and
lllycin, iuHi aa bippnric a«id In furmed from Klycin and bciir'iic add. It wa* ak»
u'one time snppoaed that urie acid might Im regardod as urironyl cynnamidt.
If the osidi'oce of some cyanogen residue is thna ansumod in tlt« raolMola 1
urie add. then it must be euppotwd that before urea can be obtained from it >
Ann. d. Cbi-m 11. rhaxB.. M. e» :^ SI
* UnttaniMkl, Sa >l tloMiili. CbaN.OtMilU Jahrg 1>«1.S. sn«.
I
I
I
• SMV.KnleilM.X*M>*fe.r BM.M.RlH.(l«7).a.«l fthiMtr. MiMfc. C ttr^lL
> i*hib. ln,um. VM. M Xilll i* 41.
• OUMiplxr. a. IM}>rtMk. At»a. I*CK llnft ■. Sl m.
M.
nnlncutiir rlikO|ti> iitkr* |>Uct by which ■ iMrtiati at Unuil nf thp niunx^i ■■' tlic
uric BL-iil u oinvciicJ into the Mtn« cundilion u lfa« rod of the nltmgen. vli.,
iiitrj ih« anld« state.
If ibu b« M, aiiiM the mi^tabnlMiD nr the animal* in which iiric B«id rci>lacM
urm cinnot bir au|i|><a((l In W ftindiunciiUlly dilTrredl from dial v( tin- urt*-
nmdacinf aninub, wi- muy iafet that Ibe iiiiUi-i'ileiit nl Uilti uric iicid anil urtra
In lh« regrtaalve ntetulHiliuii uf (inii«ldii U, its we Auggvalv'l uImtc, a \>oAj c«ttt»iii-
ing Maw at lout of ita tiiirug«n in thci form af c^anogcii.'
KreaOB. C,ll,N,Or
Occura aa a cnnslant cnatilituant of tho jaicm or mnncte', llioogh poMlbly It
may be foniiAd during the procoM of rxtmcti'm by the hydration of noatinin.
KraaUti >• not a normal conalituont of urine, but it ii uiiil lu uccnr in trnco in
Mt«ral flulib of the liody. Wh«ii found iu iirine ii> j>ra>eac« 1* pmbablj ilue to
tbi- conrenloa of krcatlnln. a coitsuim ciioMtitui-nl of uiiop. Inio Icrpattn during
il* (■xlriwlinn. »inco Dcvwaijinw' lia* ihnwn that llio mar* layniUy thr uppamtion
in rlTrL-tnl. lb« Irwi in thv iiuanlity uf krnatin nbtainvd, and Ibc gruil«r tlio naiount
uf krratinin.
In thi' anhydrouH form it is wbito and i>pa(iiic, but cryitallizca with one mole-
rale of water in coIorteM, trantpar^nl rhombic |'ri»m>. It pnweaMB a antncwlial
bitter taiiie, in solubk in cnlil. exlrcmrly onlnlilp in hut wuler, ia lesa (ohiblc in
abuiluic than in diliitr ak^ibol, and in Nnlublr in rlhcr.
It is a wij Wfuk boiif?, tcnrcvW noutnilixlug tin- wenkenl aoida. It forms vryn-
lalline compound* with Bulpburfit, hydrochloric, and nitric acid».
flftpamtion. From eatraci nf maaclo by ]>rccipiCotinK rompletcly with baric
lead M-etntc, and cryHt.illiEinR nut the krentin, mixed wilh krcntinin. Fmm thia
latier it it lepiirnird br the formation of the xinc-aalt of kreatinin. krcatin not
readily yielding a nimilar com]>oUDd.
Knaliu DMy be eanTonad Into kraaliala uuiUr the lalluonco of udiU, iha ifBuifbriaialiM
beiu|C oiie of naiplu •tvliyilnliun,
Kreatiii may be decompuaed iotu vtrcoain (niethyl-glydn) and tirea:
C,H,N,0,- H,0= U,H,NO,TCn,S,0;
il nay be formed iiyntheliailiy* by the acliao of aar«otiln and oyanamUIe :
CH,XU,tCD.N,=C.U,N,0.
San^n [aBlydnin wlilchoneatoinorbydrngen haabce«i replaced by lb* akolMtl
radirlo mMhyl, tbui>:
Cllyei- 5?g." } O beeow- ^.«4';.'* } 0 i
ltk« f lycio, (areonin ha* nut been found in free etate in llie boAj.
KiwAtinin. L',H,N,U.
TbU, vfliicb i* aSmply a ilchydratcd form of kreatla. nccura Bormally a» a
•tant cdOMltuent of urine and of tnuwic extract. It cryatidllren In color]
•bintng prtMnf. poMtenaitig a alrong alkaline taele and rcat'tion. It U rei '
aolublein culd water (1 in 11.*'). nlpwj in alcohal, but i> icanvV ■nlublo in ciIiot,
It acta nil a iiowerful baae. formioK with acids and mil* cumpouitd* vhicli
cryaialliM* well Of these the ii>o«l important !■ the anil with sine chb>rlda
(C,H-N,0|,Xa t-l,. It \» f'truied when a concentrated aolution of the chlorhle b
added !•> a not t<x> dilute solnlUm of fcrealiuin. Since the coiufixind in very little
afdnbl*' in alcohol it U better lo ow- alcnbolic rather than aipidHK wilniionn. Il
i*ri->tallJMa in warty lump* comiMMed of nfgrc(al«d niaattai uf ptiauia or fine
tieedlcK.
frqtamiion. EJtlier by the action of iiclda on krentln, or l>i>m human ur^ne by
M "••"■a"
i
iMoO APrE-vDix.
COnc« Dinting an i] precipiUtiDi; with lead kcetit«; in the filtrate from thisaMCoad
prectpititt« U caiued b^ the adilition of mercuric chloride, and consists of ■ com-
pound of thin Halt wiih kreatinin. The mercury is removed by suiphurestei
nydroEcn. anil the kresiiuin pariSed by the format iou of the zinc salt and vssIub^
with skohol.
Kreiiliijia-ii[i<> clikiriilr rnHT '« rnnverte-l inlo ktvslin, by Ihe Bcti->n uf hydnlcd nxiittl
]«ail 'm iti ^ppiljnif acjueinin ^'luticm.
Allantoin. C.H.S.Oj.
The characiemtic cnnstitueBt of the allantoic fluid of the fcBtaa; it occuitaho
in the uriae i>f animalg for a short period after their birth. Traces of it aresoae-
times detecu>d in tbia excretion at a later date.
It cryntallizeH in »mall, xhining, colorlesH prisms, which are tasteless and odor-
less. Tliey ure soluble io l&l parts of cold, more soluble in hot water, insoluble
in cold alcohol and ether, soluble in hot alcohol. Carbonate« of the alkalies <^\t-
solve them, and compounds may be formed of allantoin with metals but not iritli
acidfl.
Allant'iin, an already stated, p. 1048, is one of the products of the oiidilioa of
uric acid, and by further osidation gives rise to urea.
/W/iirii/iiin. This is best carried out by the careful oxidation of uric tai,
either by meiins of potassic permaoganate or ferrocyanide, or by plumbic oiidc
Hypoxantbln or Sarbdn C,H,S,0.
Is a normal constituent of muscles, occurring also in the spleen, lirer, tnil
medulla of boneK. In leukiemia It appears in the blood and urine. Itcijiul-
li^es ill fine iieedlea which are soluble in 300 parts of cold, more soluble in hoi
water, insoluble in alcohol, soluble in acidsand alkalies. It forma crystalline coii-
[HiundH with acids and baaea. It is precipitated by basic acetate of lead, the [«■
ci]>itat<; being soluble in a solution of the normal acetate. Its preparation &aa
in use Ic-f.t tract ilepenrls on its precipitation first by basic acetate of lead, and lh«
by an iiinnioiiiatiil no!ution of silver nitrate after the removal of kreatin.
iliilli hyintxjiiklhit) HOil Mie nc.\t IhhIv, vjinlliin. cnn al^^ bf oblainecl from proteidf bv lit
HI''! f |"ilr.TJi''liv.> cliiiiiKei'. "f "•mIit «t lnjiliiic luiiijwralurp. of dilute !iydrufhliiriiri>il
III.:; [-ir 1 .■lit.) lit l() ' I'., mill liy {\ii- atlioii of Ruslric Biiil fiHUc-reatio fermeiiU.' Ciiiueiniti
tj»?i Miilirril ;i if'-iiUiiv difW-ri-Uf- lk'IW4-*>ii tiliriii Hiid e^e-<ilbiiiiiin vrhen 3i>bmjtted [<* tboEi^^^
]iriH>^K.H>ri : III. fiiiil.4 i}ijii ilif^ liitler d^>r^ iii>t vifM iiy]*iixanItLiii when tnrat^l witb boilsEC
K-iilcr, 11-11 h ililiiiii Ly.lripi'lil.iri.- iii^id. nr I'ni'lrii- tVrrneiil, irliile lhi> former does. Ei!g-ilbum:o
nil till' I it 1 1', r fill III I yi,iM:> liyjHiXLDilhiii by tlio ■I'lion uf ]>Biirrealic furment in alki^liuriwluUi'o
biiT inil ,.-.1 midili' ji?! liliriii liin-s.
Xanthin. CiH.N,",.
Firnt liiHi'overt'd in n nrinarv I'alciilus, and called xnnthic oxide. More recentl.r
it hn.1 lit'i'n f'liunil as n normal, though scanty, constituent of uHoe, muscia, wt
sevnnil orpln^<, hiicIi as the liver, spleen, thymus, etc.
Wli<-n prci'ipitaliil by ('<'<>lin); from its hot. saturated, aqueous solution it Ub
in ivliiti' llocks. but if the solution be allowed to evaporate slowly it is obtained in
:<mal) soili'H. When pure it i:t a colorless powder, very insoluble in water, m]ii>i-
in^; iriOi) time:* ibi hulk for solution at IW V. Insoluble in alcohol and etfaer, n
readily ili^solves in ililulo aciil:< and alkalies, forming crystal lizable compoan<b.
ny]io\:intliin liy oxidation becomes xanthin. ISoth these bodies, as well uii'
foUiiwint;, (luiiniii :inil c;irnin, arc evidently closely allied to uric acid ; indwi
uric avid hv ilic :u'li'in of :<oiliuin-am^ilgani may be converted into a mlitiiKi''^
x.iiilhiii ;iiiil liy|iii\;iiilhiii.
/'•••l-i-'iliiiii, h is obiaiiicd fr^im urine and the aqueous eitract of muscle b.f
prMci':i.-> siniihir tn tli:i1 for hypoxaiithiu, and is then separated from the l«11«r br
ilio iu-li'Hi i<( dilute hyilruchliiric acid ; this separation depends on thedifl*r»i
Kohibili til's of Ihc liyiirocblorlili's of the tn*o bodies. For further infonnalioBiM
Ni'Uli:ilii'r ami Vo>;i4.-'
7'*-l''7" . S. >'>•}. Kthvm!. Ituu£. liisft. Iferiiitl"^
KifSM.-!. /^'U-oh. f. ptiy^i^«L v'bi'm . BtL. v. .IW". |
: ll.i;:i .Vn.i >-^- IM i -i. I—'. - -■■■. .V^Hi iln- liiitaiiirv .|Ui>le>] aleveun hjrpoxuuUa.
' J ,
..' 'I- ,'.
■-,-h: ■■ 1
'> ~
llVlil
"1
li
• h:::-
.-I '1. -'. '!
r, .■! Il;i
Si
11 }-:
' ^
1'-'
.liilirj
\';^ 1"
- .1
.-...1
1. Mill).
s
nil
I
CHBHICAt. HAfllS or TIIK AMUAI. BOUT. lOAI
OUBlB. C,H.N,Or
DisooTfred by Wei4«l' In exinci of niMt. or whidi ft oiaiiilutM nbuut on«
pereeot.
It ciyntnllizM in t«hil« miiwoi crnnpoMd or very sirtall. irregular cry«M]» ; il ia
■olgUc wllh dilEcuhj in cold, mnro ciuily Milublc in tint nalrr, iDMiluhli- in nloci-
hol and Gtti«r. lu ai)uvoun Holutiim in not |>rvcii>i[iiitMl liv nonniil Irail novtate,
iMit ia b^ tbe baaic acetate of tlijg metal. It unltM with adda and mIM forming
cijatallin* coHipoanHii.
jyeporatwti. In found In the prrripitttp cnuard in Mlract <if laeal by baalo
acetate of l«d.*
Tlii< liol; psaiNH* sn iatara>liii(r*Uiluii tf> liTi-maiitliin, inlo whioli II iiMjr W<DnvDrtal
by til* Mfioa eilbar of nilrii' beuI or. fltll boitnr. V>f lircFiiiii*.
Ouanio. (^tItK,n.
Fintot>tn!n4Hlfrrimgunno. but recentljrobserredaaocourrlngin atnall ituntiliM
in tb« pancTPiiH, livrr. and muoclo pxtrncl.
Illaa wfali«aniorpliou* nowdvr, iiuoliibl* in water, alcohol, other, asd ammonU.
It vnittt* nilfa acldo, slkahen, and anlta tu Tonn cryalalliubtc conpouuda.
Prfpamlion, From giiiuio by boiling MicceMlvely with milk nf llm» and caoalic
•oda. i>redpituiliig wttb acetic acid, and puriiying by lolutinn in hydrocblwk
acid and firecipitntion by ammonin.
Goanin mav, by tbe anion nf nitmua a«id, be coorcrtnl into xauttiln. By oxi-
dation i( can be made to rield iiHuciimlly guonidlne and parabanlc acid, accom-
panied, IiiiwcTtir. by »inaU<tuaiilitir* ofureift. xaiiihln, and oxalic acid. Ca|innics
Ua4 giv«a oevrml rratiion* chanirteriRtic "f thin body.*
Ita sefiaration frum liypmiuiibin mid xanthin drpcniU im it* inauluUlitj In
water and behavior with liydiocbloric uuid.
Kynurenlc aold. f,TI„N,0,+ SH,t>.
Found In ihi? urine of dogD. and Orat deaoribed br Lleblg.* Wlien pure il ^^*-
talliT«« in brilliant irbite needle*. IniolubJe lu cola, HtubTe In bot alcidiol. Tbp
only Mit or thin body nhirh cry»lnltiies welt la that formed with barium. For
preparation and oibcr parti ciil an xr Liobig.*SchtiltXMi, and Sebmiedcberg.'
Olyoin. C,U,(NH,|O(0H). Attn oikd Hlyooedl and Glycocioe.
Dow not nccur iu a Tree ulate in the human Imly, but ontera into the eoBlpad*
tloR of many important aubttaneo, t. a . hippuric and bile iidds. It crystalliM
large, colorlea*. hard rboinbohedra, wblcli are ea^itv soluble in water, inaolnbla in
cold, ■liKhlly aoluble in hot alcohol, iaeoluble in etlier. Il po-wcow an acid reac-
ttion, but a ««r«t tutte. It baa aUo the pmpertr of uniting wilb both acida and
baaea, to form crystal t i «ib le oompouodii. lu thii'il eshibita ita amide nature, and
that It b an amide Ih rendered evident frwin the meihoda of Ita Byntbetlc vrepam-
tton: tboR oiono-chlar accllcacid and ammonia giveglycin and amnontuctiloride:
^dn,CI 0^ -^ 3Nn, = C,II,(Xn,lO(0II) - NH.CI. it » amido^cedc add.
Healed with caiucic biir^'ia it yieldn amnionla and mothylamine.
J^rprtratioa. Pruui slutiu by thr uclioo of acid* or alkallte; from hippnric add
by decoffl]>jain|c it will! hvdroehloric at-id ai a iMillug lempeniUTe ana rraWTiBi;
by precipitation the ftimiiltaneou>ly formed bonioic acid.
Taurtn, C^,NO^
In addition I'l enterine into the cninp<»ition of taurocholic acid (•«« p. IDTifi)
laurin i* found in trnim in the juiots of muKcle and in the lunipi.
It cnrstalUcua in cuIoHmb. regular ■ii-aided pritnu) ; thete are reulily Miluble In
water, leea ao In alcohol. Tlie aolollann are neutral. It Ida very Mable coinpoand.
neiaCinc l«mpenitnrc« of lee* Iban 240* C. : il >• not acted on by dilute alkaliea
■ad acta*, even whrn boiled with tfaem. It ia nut prccipilnied by melnlliu lalt*.
• Ana. <t. t-bnn. u. phUK, 51 rtviii ji ais. ■ ^^ »vii]ci.o(. tit.
• ItHtti. t phnlnl. <lMm.. IM, tv. nWDI. « va
• Aaa. 4) ClMtni u. Ptena.. IU Ixxnl »i. ir.. *i>rt M n\a.h.*H.
•0|i,eU. • Anr. ri n>eui iL.naflB, Bd.tUlT.&Ufi.
\
Tuiriti ii ■Dida-i<Hlii'>nir «i:i<j ; bbiI tnar t>e ■yntlirllmll)- )>rp()*i«<l rruni Urtk»
onlc (fllhjrl-aulphurtc) acid by the ftcUuu ot •aiiuoiiia ; ikus :
^"•} so. + SH.= ':;fH|}so,+n,o.
Prtyata&m. An i> proilitct nt tbir dcooiniiMlUwi of bile, and h [KtfiGvd bj
ths mnnr!u)[ iiiiy iriicm ul' bile aci<J« by ninoti at lend aeeUbt, And Uihii
wccwitcIt cr.VBintlixing fruui naCer.
Loucin. r.HuNO^
la •mii (if the prindpol prodaci* of ili* iIci'>>iDpri>iii»n nl niiroeenouA innurr,
eJlhrrTinilrrtli<<indacnoeor|iatrcfHi;tii>ii or ornlruagiwiiUuixl nllcall<m. Iiiiccun
bowevcr. nonanWy In lb« pnnvrivi, iplncii, tliyiniu, iliyrnld. nlirarr ulandji, li««,
Mv.. and in oDe or the uruduem of the irypiic (pancrt«tlc) digoiign of pTtAmit;
iaai.-iiieiilr<-iphyoriheIirer it U preHcnlin ilieurliui in laii{«<|unnt(iy in c«aipi>;
iriib trriMiii.
As liaunUy oliUiint-d in an impure A>nn il iTyHUUitea in n>iii>d«d lump*. wUd
an often ootlecied t'lK^ibtr and ■nnietinie^ eihlbii rudiAlIni; iitri»(iim. ^Vka
purr, h (<-irm% vttf thin, white, slitieriiiz. [!iit i-ry^taln. TlieM are eauUr aolnUi
;n Imt nnti-r, Iwn so in cold waler and iilcoliol. iiMotublc in «lbw. Tlief »«d tttj
lu tile luucli. and iin wilboiit *iiiell and ta*te. Acid* and alknllea <ll*sn1ve ibMi
readily, nnd crjrtallizablc compuundu ate liinncd.
OaN/Ully boalMl to 170° C. It jiiibliiiiM. bnl nt a hlgbar l«nporaiure >■•!■-'
ing amyUmiu, nrbouk nnby<ln<l«. and Kninioiiia. In Ihe prewnof of pn.
BiaUor >^ tplilii u|> inlii v»lvrir wiil aD<l aBBiniiik.
..ni)
I
I
4
Leuein ia ainido-caproie acid, and may be rejircaented thin:
''l^i
J
prtpitnidon. Prnni horn «havinp by boitinK witli nuiphurie acid, nvumlldif
with uiiryla and *<-|>*iatiaK fti>m tyroain by f<ucei»iirv vryslalliKitl'in, See ab«
Kilbne,' nbo prcpon-s it by ihc itclion of pnnertiatic furineni (trv[i«in) on pi»-
teida.
Scberer baa given tbo following tut for ieucin. The aiiapecled anbatanct I*
evaporated oarefully In dryntw with nitric acid ; the rcaidoe. if il \* leudn. will
)(« almuat tnuNparcnt and turn yellow or bmwu uu ibe addition of cnustlc aa^L
If titix Iw Hcaln rery carefully Joncenintied with tlie alkali, an oily drop ti ob-
uiiieol, whiuo t« quile characteriitlic nf thlariiliataiice. I^eni'in, if mrt uw tBipuni
may be cwily reojigniiici by tin Miblimlnjc on boiitg htatcd ; a cbaractfriatio coliff
of amrlaniin in at ihe mmtf time evolved.
I
m
Asparagine. C,ll,N,Uy
!■ noi found m a caiinliiaenl of the animal body, 1>ut appears li> l>e fomim
the d«oani|M«ilfoii of proteidit, notnbly iliirin|c Ihe gornimntive cliiin|[4« i>f tbit
proteida In leguminous aeeil*.' It ia a cryalultine body, and mbva Unled wild
aclda or alluiUca ia rradily oinveit^ into lupantc acid.
Asportio <iw atjiaraginif) ooid. C,H,NC,
Tbu acid haa boen obtained in amnll qunniitia among the priiHuclB of tlie i,
creatio digeation nf fibrin' and vegetable gtutiii.* altbouKb tir.t mrcarring at.
conultueni of any uiiimal tisaue or aovrelion. It i*. on the other hand. T
normally In plains. n'>tablv in beet-augar molaue*. Il ariaea also aa a i
produd of tlie action of alkalie* and other reageniii uu both vegetable and i
proteidjs and nf acidn on KeUtine.' Il thu* |-MNe«ea OOTMhlerable inltr.^ ia
reapcct of il> relation to tbe protcidH. Il eryMtallixoa in rhomliic |>riiiKit which.
< Vln4Kiw'*ATeMT. IM mil fl IW
> UiHlatnbwh. V'--.. I
■llmldclendlu. >^
• V, Kutfffeui, 2i)ii-. ...
• HorhKaMvU. Hub. <). i. Akaj ii, \'< \f.. v, xu, x^D. 2 JMh. Jnal H«<L
■leralL. Jalug- vQ- imV. !• lU*
»
»r* l«H ■p«rin|[lT nolgble in coM irniw or Rtcobol, mdWj idIiiIiIp in buillii^
»»t*r. Ita kciil •ulutiun* are dexlro-mtatory. iu alkaline tn-Tu-niiali>rr luitl
rednoe Fcltliii):'* Buid. It furaw « cliaraoUrialU- rMdilr i-ryaulliubl« com-
pound with i-u|>[jer. Nltiuiiit acid conrerU it iolo malic acid,
Olatamiolo nold. C.U^'O,.
Tbf circuniiitaiicQ* and c»n<litiiinii under vbicli tht* body ocnn are iii pneral
the ooaie lu fuf ihe aapariiL- hcIO. un<\ lii'ni-e uh a product of Dmicid ileL-omimaiti'in
It ocqalroi Mim« ini|ion>inc(i. It hnn ii^>t. Iiowever. a* yet Iteeo obuined \iy th<*
action of poacraatic fivment* on protcidii. and iii thin it diflVr* from tlie prM.'«diut,'
body.
It erjilallixra in rlinrabic iGirahedrn or octahcdrn ; l< nnt very aulnbli; in coUI.
bat re«ditv tolubic in )iotw»i«r: UiKoluble In alfohul and eib^r. In iK'id itwlu-
tloDB poBBCM H Btrong dexiro-roiatorr fiower, and it retliiL-M Pebliiig'a fluid.
Orstio. C,U,NSO,.
la the chief cttiuitiincnt i>f n rarply occuTring urinary calculus in kko nml don.
It maf mlao occur iu renal concreiioiut and in Rravel, and in uccosionully foand in
nrine.
Frotn calculi it i* obtained, by eximirlion witb tiintnonia, as colorlHS ilX'tiidcil
tabic* lit rlionibnlicdm, which arr iiriilr«l iind LMlclnw. It in iniKilublo in i>at«r,
alcohol, and elhiT. wiluble in ommouia and tb« other alkniiiw, nnd aliw iii minenU
adds. The Out tlmt tliia body la one of Iho Csir ajntalliiie iiubsUni.-ea, ucvurring
pbyaiotogii-iillv. which contain mlphnr, renden its deteoiloo very «sMy, Apart
Rom lu inHoliibility in wntor, etc., it yii>ldK with cauhUc p'>tMU and talts of
either silver or Ind. a brawn culuratioi) due to tbe proMOCO of lli« sulphidca of
tline nietal*.
AecoolioKta Dswacaiiil iiauitfi*:' cyiiiii l>*iiii<l<>-Mil|>ho-tivrurloaad,aiiil II* AirmuU ii
CilUroOi — pjrnino boing Inciic ueM aiiaui mo sl>>iii* of bylroijiiii.
Tub Akouatio (iKKres,
Bensoto ncld. nC.UtU,.
Thia t* i»it found iw a normal coiuUturjil of the body, but Owm its pmenoe I&
urine to the fcrmentutive decompueiilion uf blppuric add, wtMreby glydn Hiid
benxuio oi-id are furni«d :
tlippiiric acii). Glvrin, Oouiiila uid.
C,H,(C,II,0)SO, 1 H,0 = C,U,NO,+ C,HjO,
The lublinted acid ia generally cmUilliMd in fine needle*, which am li)rbt luid
gliateiiint; : niiy odor they poMeM iii not dtie to the acid, hui to on eHenttiil oil.
witb which llicy nre mixed. ^SHicn prcdiiitatcd fWm solution the cryBtallin«i foTra
b always indiitinct. Thi> arid is nolubic in SOU patis cold or %'' parts of U>iling
walrr. but is ca*ily xilublc in alcohol ur etlier. It *ublim«* readily s( IVi" C. ; It
also puuMC* off in ihc ra['on urinin^ fniln Ibi heutc^ a'llutiuaii.
Premmli/m. Either as abuxe ftftm liipiiuric acid by femientiuion, by boiling
lite blppuric acid wiih acids or alkalies, or bjr aublimattoii fruoi gum-benMln.
Tyroolit. CVl„.\u,.
Q«Dorally accampiinin Icucin, and ia perhapa found normally io small ^uan-
tilifs !o tlie pancreas nnd apleen. It is also mually ublaiocd in larxe qnnntitiiii
by tbe deoooipositlon of proteld tnstlcr, eiiber by puircfoolioa or the outioti vf
adds.
Tbe rvaaaiDlMaof Riutiif^awaliy' rmdar it prnhotila ilial lyraain ilaat ael ocmr aonnallv
in aay pod of Iha bnnian anaaiam, azetFa as a prodeiS *t paaaaalli) lUitaaUsn.
All attempts to synthetlse tjrosin «et« for sofne lime fhiitlma, although eci-
denoe wna »liiaiii<yl MilHcieut to Indicate the probable exiai«nee iit ib molecule at
< Juiim. oi \B»\. and pkv«oi.. ytn„ imi u. ita.
> AiDlilr, (, lalli. Aaal, bL suvL S. 1. ZdlMb. t asol. Ckav.. BJ. *. K. Mt.
1054
APPEXDII.
some aiviaMic (ph«nyl) radicle.* More nvcntlr (he •jnlhMb bM bne*
formod.* «itd «c BOW hBTo pvery nwon Tur rcguiliiiK tyni»ii> u p«n-fa;ilrat,
pheiiyl-(i Kluniite. This »i-uth<Miii u wiMl lu tliai of uric' acid, referred Ui aburt,
w of oonal<I«rftblo iin|>(iriiiii<.-tf, since llie idotc defioitc tlie linoirledi^ wkich w
poAMwed of Ibe true molocultu' etrnctiirc of tlir prnducU of pti>tri<l deeoaiiNMi'
tiMi, tbe mora mwxi i> tlior« Ibr ripcctini; Ibut Uiv Kynlbmia of ■ prouM fbtU
may be realiiable in the not Tcrr raiitMe luture.
TTrodn cryiUllizM in exceedfiigly Ane n««<llea wliicb are ii<mallr oollvctwl Inlt
fMllicry DiaiiMs. Tbr cry^uls tire itnow-whtte. Uslelmh Mid odorlMa, klnart io-
Mluble in. cold vttxcr. ivjulily notiible in liot water, Bvid*. and Kikaliw, iiMoloWi
in Hlcohoi and elber. If errnUiUixcd fmm an alkaline *oluUon lyrtnin oAv
anHimo tlie form of rotetue oooiputii'd of line needle* arraaeed nAiAielf.
TyrMin doe> net ■ubllme by bi^aiiiiL'. but u deooinpmed With an odor of pbml
and nilrobenxol. On boiling wiib Millon'* reageal it give* a reaction alnM
ldenli(«l wiUi. but much more marked tban, that for nrotuid* (ilnirmaa'a tM^
If tvroun ii irculcd on a watch-gtaaa trilb one or Iwo dro|M of wUoiif Milpbtnt
acid, then diluted with a little water, oeaimlixed with calcic oaxboflate, and tW
•olutioo lillcrul. a cbaracivristic violet color t* oblained on llie adiilthm al a iuf
of acid-freo firric chloride (Piria'« leit).
Pftfuiralion. Ily nienn* liniilar lo tlio*n nnplorci) for Icacin. llieBCftanUkarf
the tiro ili'pcoitinp on their widely diircrinc sulul>itiiirs. Accmlinu to Kdkac't
iDciliod,* large qusutitlca are eusily ohtaiueJ aa ibe rMull of pancreatic iligeriiaa.
Hlpptirio sold. C.n.NO, Or BMiRoyl-Blycin. C,II,(C,IV))N<»,.
la found in comudemble ^unntitic* in the urine of herbinira. and aWi. ihMlk
to a mitch amalirr amount, in the urine of man. It ia (braied In the body bf Ui
nni'in with lii-liydmti'm nf i;lycii] and hpiiKoic iicid.
CrvolaJlind from a Hatoraced iiqucuiu iiulution It K<Hume« the form of Am
oeedlee: if IVom a mofQ dilute tiolutir>Ti. white. Miiiittnii«)>iti<>ni f'tui-*id<d friwi
arc oblained. Tliwe when pure arc i>dor!cnt. nilh a itomcwhat bitter lavte. Thij
tun ■'iltibic io 000 p>irta of cold natrr, rcadilv luiluble in bolliriK water, mdUJ
eolifblo in alcohol, lew (o in ether. All the ■oinlion* rcddeo lilmua.
Hippuric acid is monobaiic, and fortiii Mita which are readily aoluble In waur
(except the Iron salu); from Ihtec, ifSuBuffictenily concenlraied aoliithxi*, cxnw
of hydrochloric acid iireoipilAlea the acid In fine needlw. When hcRtnl viih
coiiCADtnted mineral acid* it !■ rewlvcd into bentoic acid and clycin. Tlir us'
dcooBipnoitioD occon in pretence of putrefying bodiw. tflrong nitric acid prv>j
duc«* an odor of ollrobenxol.
Pr^tmition. /WsA> urine of boraee or oow« U lrf«ted with milk of liiM^
order (o form calcic hippuratc and thus prevent the dconmpuutioo of ibr hljifm
acid. filtered. and the liltratc eraporatcd toaamull bulk; the hippuric acid i*lka
precipitated by nddinc an cxceM of hydrochloric acid: the acid ii then jwriM
by aeveral crystal lli.t I Ions from boiling water.
VVhen hentcfl in a *miiU iiitie, bipivuric acid given aenbliinate of betiMtc artd
and ammonic bcriKonto. accoDipnnied by an odor like tliat of new bay, while oUf,
rod dropi are obterrcd in the tube. Tbia it very characleriotic and diatln|[abW
it from neoxoic acid.
Ptwnytlo (e*r*.iiM) aeld, or/»A»iiaJ. (',0,«.
ThI* body la undoubtedly oblatneil m th<t mult of the putrefective ittau
tlonofprotdidR, notably in putrcfnciivc pKncrcuiiciligeationL* It may be obi
from (he dtrtillate of mch dij^mtivc mixtum. It in aUo fmind in the cmilenlK
the alloirnlary canal Utidvr the same ciioditioiii which kI"' >>»<-■ t'> indol. Whl
M uecnrring a jwrtioo of it may be obtained from the fecea, while Ibc rea4 i
pears in the unite.'
■ Diittti. aicm. i>ntn)h.. laca.fl.MaB: tStK&Tn: ISn&NM. Ilnfwr, IMH. imu, & laa
■una n. Kaiiit.-fit n.vi , irfrt8.«aft
• RrleniuDiv' ' .<r. >l. ilcoMft. Cham- itmUl, Mum. xr.fOKXS. IML
• Oj- rti .•
• SUkOHlkl. Ikl. :i. ImilJCll. CIlBUI I I . . - .1
t-ia. Kfi. 4. 4«4U(4. rtiau. (lo«l
«». ate alio tVnlnlli. (. iL mttl Wi-
ll. Iinm.a. Ml.
eilEMtCAL BASIS or TUE ASIUM. UOUY. lOdfi
Bulif latky ■ wn iba urin* of nan)' Miimal*. or cor* uiil hnm* ■Imjri^ conUiiu • Mb-
itouo* iiiMlubIa In ilmliol. Nnd not praetplUad bjr load a<«l»i« and knuiiMtlii. wlil«b ky Urn
•rtHMi uf iltlal* mlnvrBl aoid* p've* aarboUr avid The •am* ■eld afii>lii>i to Ibc hodir
•sUnulty ■■r iaUrailly aUi pawM ialo (ha uriur.' BiiniUrlj bWHDl <(Vtl<), «h«B lAhW
JniA tlio ■kiniafU, appt^ari ■■ a catbfllip acid In Ilia tirinc-*
Til* pure acid crntslluM in lonji;, colorlmi prinBiitIv omJIm; ihcy meli «t
Vt' C. and boil nt Iw 0. It m rMdily aolubk in alcohol an<) ctlitr, BlifliUjr
Hiluhir ill vraict f^l part in 30}. In toort caa«* it net* Ha n Rcak acid, running
cryaUiUinL' Htilia wiili the alkalitt. With nitric aclil {ljI»IcU pimcncid. IukiIu-
tioni reduce aflTcr mnd mercury wiltM.
I^HjMmlion. Bv the dir di*tillulinn o( RalicUic add. rIdo from tlie acid pro-
diU't* of the ilUtillalion of coal. It ii obtaiacd in Ihc laal portinn* of thr dintll-
lalp when preparint; iiidul. and la Mpanit«tl l>y furmiui; u comiHiimd H-ilb briHiiinc
q,UBr,iV
TltK ttlLK 8KKIKS.
OboUdlo (or eAi>f<«) Mid. U.CIIhII«Oj^ U,0.
Occur* in Iracnt in the Bmall iniMtine. in larger quantili«a in lhi> contfiBla of
th* luff inloitinv. nud llie ttctt, uf uii-n, oow>. mid il<^. In ict«ia», the tirin*
often conMina trace* of tliis acid. Bat it* i<rini-ip>l inirrwi liea in lu being the
aUrtlng point for tbp varinua bile acida (■•« lirlour), The pure acid may be amor-
pfaoua or crjnttnlline, in the lattrr rate crfoalli^inf; fr«in hot Rlci>hi>lic M>lutJoiM in
tOlraJtednt. Theae otjiUit are iuwilublr in wnlcr an'l vlber. In tbr nmorpboM
form, it li aomewbat aotuble In viati-r and (-tlicr. Ilcnicd to -^UOp C, il u cos-
verted Into mttt and dyilyun (C„il„(J,).
Tbia a<:Kl pcMiaata. in the anhvdroua coiidilinn, n niicdflc rolatnrjr power of
4- 00* for the yellow light; when it crvntalliMa with II,(). ihr mtaiion ti + S3*
The roiMor^ power of the alkali nalta I* alwaya loa thnii the nlHire, and wbon ia
■olutloo in aleoliol ilio rotniinn i« independent of the i-oni-eotrtuion. For the
alcoholic anlalion of the nodiutn utlt ibe rotation ia 4- 31.-1'.
/yrj'iraiiftH. Ily the deconipmition of bilencida by mean* of acida, alkalio*, or
ftrmenlativechanice*.
Bajcr* ha**xainiii«il iba Ul* bci-t* atrial oad froiH liuiuan lillQ.aDilhaiprrjairv] fnot iJiau
•hoUlte aild. To ibii )in BMignt Iha Ibnaiila CttTTi^i. If ikU ba •>, ih»a diutalK a^id of
biunaa bils vouM mm b' b* a body Miili*ly diflktvnt ftow Uial ohtaiMed beta ax blla, and
analyml bj Stndior. lUyar** ramlla, bowwrar, roifuira fcrtbar araCmaUoo.
IVifiiio/er't UtI*
Thi* well-known toit for bile add* dcpenda on the rea<Tlioa of cholalic acid in
Kreaence of aagsr and «nlphuric acid. If to n (o1iiiit>ii >if the aoid a little lagar
e addod and then nuhihiiric acid, keepinfi the trni|irf>lure below, but not mucb
below 'ff ('-. a beautiful rcddiah purple is obtained. If diluted with alcohol thla
•nimiiin givea a chamcteriMic a|)eclrani with two abmirplion luod*, one betweeo
D and E, neare*! lo K. the otlivr cloae to F on the red aide of V.
The reaction i« mnrh inipoded bv the preaemv of colorinf matter* ; morcAver
pcoirid* and other liodies eaaily deconijioted by aulphurie acid. Much at amyl-
klcohol and oleic, t;lve u niaillar reaull ; ihe colurint; matter iiroilunHl from iMM
bodlea di>» [i<>i, hifuevvr. ^iie the abanrplion bands deacribed above.*
Olyoocholic iMjld c,u„No,.
Thta body waa firat obtained in the erratalline fom and dnaerlbed by Unelin
(1828), who ^ve it the name of "cbolie " acid.
To aToiJ eunfuaiuu It la u>iw b«t lo ■■• tlta Imib ■■ eba4lo ' a* a •^uiiaym b* " aftDlall*,'
Dafnarmy, who f nl ( I938j ibacrlbtd Ilia elioUlia and a* a pmlort of Ilia ilaeDaipoaJUoa of
• Uopp^dnlar. Uol. rbein. l'nK««i«h.ll«nit. ID«7i».3t
• AlSln, ^«ii»JaatlL .1. riiani '
__^ . inn.. M. uiir-^-ltl. SaltowiU. r«*(*r> Artklr. Bd, T,(Un-ni
• IkfenlUpn IL Xatiujii, Hclobim <L ina^lM* VMrntm-it Anbl* ilMI), tUA 10. S. aW.
• 7eIi^'I> r i*>T^tt Ctnm., M n il«nbTMlLjM.
' t^liitiV>.n.-r, Ankalni il. riKK K lliarm . M lit irHli<>.««.
■ • KaninliM liilOrauUan mi (hUi »■" t- . - - -. i r .,< j(«J,. i^v ;i. fiit tii n. ur>.
■ ScAultB. aaa. O, i^htai- ii. rbanii., |i. vnaloni.nlinlul. I nurnHdi..
■ infa.t9n.&n. A4uBkW«tra.riv<
APPK^IDIT.
Ula ai-i-b'. Iiarltiii |;h-«ii it tlio t»in» of ahollo mtU. TW tuiiw rhoUllr t>
iinre it IjiiliublBi IliD Dialhoil t/f vliirli (be blla Wlldi U« ijillt ii|>, vlL. t^ I
■ItaUi.
This in thi-
ipal bile acid of os-gnU ; H w alin pnywnt
' jii iif ciurnirora. In icicrut
thr lillr of 1
n 1
' ]>nnc)|
[ilHcrreil '.
coiibuD Unoes nf (bin atiO.
tt i!rj*ullisn in nne. glintc-tiiue ne«<llM. TlieM u« aUgliily nohiMe In oaU
water, readily m in hm wnlor. Bn^falocihol. but Initolable in eilicr. Tltcj pomm
a t>ill«r unci ypt airci-t tiuir, and il »tn>nv acid r««Ctti>n.
Tbe sails or tliia acid niu readily ■nluble in nkirr and crjutiillttc mcIL Tb
mIu, an nell AS ihc rr«« acid, etm rifilit-baaded puiariiailun aniounlinc k
-f 29.0° for tlic ncid. and ■+- 25.7' Tur thu todium uli. Iioih meaMittyl ht yJltv
lijrbu
Oilyoncbolic acid i« n caoaptiuud of glycin and choUltc a«id ; Uiu* :
rhelnlir •ad. Ol»i>io. (ll»por|ioli> »pi4.
C„II,0, + C.SHji) — 11,0 = <\,l'o>'W*-
Prvlm^ed boiling irith riitnte mincml uilitie«f*«>lli!iilk>ll« JprampnMa etvmrlMUxM
Intoilyp-'i IF'.' -liilnlicMM; ifiliMolvnH In mnceiitmlM inil|>t
cIddwIi" ' . llio rsnoTal of xrie laoliw'lla of wal*^ ywl
The b»ri r' ihi* l»i BTv-l it iBX-liiblc In nlrr, wtiioli
ebolcnlc iivi-l fijn*ue* DMTly ilio mwio qiMllli) rotalocy powor x xivwicbutiu liui-l.
TViTiaroKoit. From ox-gall by <rvapomti'>n to a <yrup, dt-coloriiiog with rdIbiI
eharcoal. cxtraviio^ witli Mrong uli^bul. uml |)n.-cit>ilntinK by « Itrgo eirtw ij
cth«r. It« »e]Mratii>a frvm Murochulic ai-id dci>«iitlB on Ita iirecipitMluo b; a*-
mal lend oLvlalc, taurocholic acid not being i>reci(>iuicd by tLis rengent.
TBurocholio oold. e^n^NSo^
(h^curx iklxi in Dx-i;all, but b found Mpocially plvntiliil in banian bil« and Ibt
uf cnmivoru, (loUUy of tlio dOK'
It cryiilutliti-B nit'li difliculty in veir fine nccdlts, which arti tfxctvdiaglj il» ,
UoueAoent. When dried il !a mi aniucfiliiniH pi)wd«r, with iiuru ktt(«r i
(olublv ill water and alcobol, insuluble in ether. Alt ita ^iiIIa ore iinluh)* Inwala. '
and are precipitated by basic lead acclntc in ibe |>r«aenM> of fre« sinmitnlA. 71*
•odiiiin lutt diuiilved in alcohol bu mipecific natatory power of + If 4.5*'; if 4^
•olvcd ill water tlii* routioD ii loaa. And in thin reaped il rcMtnblM i^lyondtallE
acid.
This acid if fur more unstable tlian ibe precediog one. belii^ decomnoant il
boiled with water. The products of decompoeition are tauriu nuil cbolalic atniL
Taumcbolic acid is a <xiHip<iund -if taurin and cholnlic acid; thus:
Cliololk aci'l, Taurin. Taiiiwbolic Mid.
<''ii''wy» + C,II,NO^— 0/J — C,HuNO^-
iVijMtrafim. From the bile "f dog* by a pmoM* aiinllar to ibU for g\yt
add. It b nparaleil froin Irsceti of ibis lalier and IVoin cholalle kcid oy \
Uaioa with basic iMd acetate and ammnnla.'
1(1 1.B Piouxjcn.
It ia found cbidly in llie frwh bile of man And otmivnni, to which It ^i
cbaraoteriatlo dark gnlden-red color. It frequently ci>ii«titu|ea a oonaiderable ]
of MNDC kinds of gnll-stODes. not, however, a* ftM bilimbin. I>ut hr a compoai
with e«rlhy malter, cbieRy chalk : the gall-stones of oicJi and pig* iiTltM eoDUtn '
40 p«-r cent, of Ihi* coraponni).' These are tfaerefhre the best natenal fmm whkb
to prepare bilinibin.
PraftamtioH. The ft" H -*(■>■■ e* are trOAleil with «lronE ncdie or dilate hy^r»-
chlnrle Add, tn sepArate the eartbf niau«t, and the rcHdoe ia thuruuKliiy washed j
■ ll(kv.TtiMK.)f(>4..obiin.i;iiwn..Bil.l.3.M(ii
• Xalr. aitilNr. d. WKn. Itad.. M. Irtt.. I«H, II. AMk. V^M- IIII.
CHKUICAL UA&IS UP THK A.VIJIAL SOUT.
1067
ralcT nud alcolH)) tnd dried. From Diiit nnldue ihe ]jn}Iong«(I autlon of hot
cbloroform cxirecta ifae btliiubln. wbiclt may Mtber be obUuacd in tb« smorpboiM
form bv ivrpci|))tntion«rith ftU-i>Iii>l nrilatmlutioa in cblorofbrni, ofm wcII-Hcfimd
cri[irUli> li>' itit iiliiR rT*)inraiioTi of ihe chlororonn nolulUin.
T1i<> DxHt uminl fonn of tlii' crymulii is thAt of rbombic prisinii ; ibe^r At« reudily
•oluble in cbtoroforai and nlkatino solutions only.
By irrainierii irtlb oxidinn^ n;tcnt*, tiu<-b as niirou* acid, bilirubin takn np
oxjgra and brcomto biliriTdin. thn rnlor nl tbo nunc lime chan^ng Ui i;re«D.
Tbt MMaibk oxidation doM iml end here, and if contiuui'd a ncrie* »( prudiiL-ta
an owalnfd. each with a cbaracttnMjc color, n» in tbe wctl-known Omelln'a teat.'
Of thme only ihe Hnnl product of thf oxidation ban been oblainetl in aatale of
f DlHrient purity lo nmbV niiy dpfinit* ■tntruicnla to be made of it« chariLct«riatic».*
fcThi* i« ibc btidy kn<iwi] a.i Ctioli-ielin (>ce below).
Biliverdln. C„U,.N,0,.'
nb iRoiluct of the oxidation of blliroldn gire* tli« cbaractt^riiilic color to Iho
le ot neriiivunt and Id biliary vomlu. It oocnr* also probably at limen in the
•iilnr of jnundk-r and in tlie piftntcntary loaiter of the placenta. It lo fonnd, m
oecan in imcc* only, in gall tttonp*.
tWfamlioa. An impiirr prodmU t« obtained br prccipilnlitif; ordinary hrr-
blvorou* bile irilb baric chlorid»^, ivashini; ihc jiiccipilali- with wiiter and alcohol
and dccoDiiKMlng it with hydroi-hloric acid. The blliverdin tbu> obtained U
waabfd with elber and diHtolvcd in alcohol. From ita anluUon In the latter it i*
obtained M an nmorpboun green pnn-der by nlow evaporation. I'uro bJlivridin M
b^t prepared )>y the slow oxidation in Ibe air of bilirubin, diMMilTcd in dilute
oumicwida.
Ii da«« not crvBlalllce. and »■ liinnlnble in ether or chloroform ; readily ivilublu
in alcohol. When nxidixed it f.iv<r* the oarnD play of cotura mj> dOM hiljniblti,
with the furniation of tlio vatne final and inleniietliale ptiidnela.
Neither thi> body nor bilirubin girea any cbanicierlNtic nimorptioii haodn.
Tfacre Mema now no reaaon for douhtiiiK tbal ibe bile jilgmenta are derived
nltimately from ibe cotorinp- matter of the blood.
Vircbon ha* dracrihed' tbe gradual cbangoa In old blood<clots, a« of cerchral
beiiiorrbifp', which lead to Ibe jirc^cnt'e of the iw called bwmaiiudin cry«tala.
Though tlieae ba*e not been obtained in HUtScient <iuantili«« to enable their com-
poailion to b« finally tlxed l>y a obemical analyslo.' tiill tbe Identity of tbeir cits*
tolllne rorm with that of bilirubin, and the fuel that they both give (he aame play
' of Golota when oij.lixoil. an In itmelin'* ie»t. jimtify tbe awumption that hinaia-
(oidin and hilirubiii are identical. " Moreover, the balance of expen'nirntal evi-
dence diiliuiity (upporL) the view tbut a Uberatlov from the corpusclen of lb*
coluiing matter of tbe blood In tbe bloodveawla by aii Injedlon of cbloroform,
water, etc., teada generally to tbe appearance of btle-piKinoniM In the urine-* Tbe
uccarrence of bilirubin crynlBlii In the urine him frcijuentlir been obaerrcd after
tbe operation of tninKliwioti of b1<>od in man. The ebemica] paaudbilily of the
Ooa*eralon of bietiioi;1iiliiu into hilirerdin b readily aeen by n coaip«rii»on of tbe
Ibrmul* of hamiatiu (M-e p. iSi) and billrutitn. The (urmer baa. accordinu to
IIoppe-8#yWr,* the coini'oiiiion Indicalfl by Ihe furmuln 2(CHll,N,Fe(\) while
that of hillrubm i* ('liHi.NjO,. Allbough ibo convel«ion \uu not an yel been
directly elfectcd, the tollnniiig facta are ■ifiniftcant. If bilirabin i* trenlitl with
fodiUBi amalcam theaubatauce known as hvdroblllrubin (see Wlow) it obtained,
ir biemaiin Cidiaaulved in caustic aoda ana treated with mKlinni anialviui ur in
hydrochloric acid vilution with xiiic dud, a nubHtiuicc In ohtaincil whii')) i» now
rceopii(cd aa identical with bydrobilirubin.* Tbi* I* Ibe most direct cbetnical
evMeace of the relation of the coloring tuMtera of the blood and bile.
■ ItodnaonunitainellD. Dtn VrnUuuni. ISH.&n.
• Rendu* unH Cuni'lHtt. l-Uuvti-i An-Ii.. lU. It. iiniL & «7.
• ilal7.«Ub.<l- WItii- Akwl.. Al lii.|lK4i,lll. AIMb.
• Aran. 1 paih. Jtiwi^ l»l >, » 3-a.
• BetiD, AnD.d.Cbem.u. rtiftnn,. Od. ciM, 8. Wl
> TMdiaaoir. niVMr-> Anh . I^ IxilKO, 8, «L Sm atn Bd. a lUm. *. 9)1.
• nvtfOlOKlKbe CTwmle. isn. a. m^
• ksfn^nln. Ucd -cUfflD. rnMfeuai. B«n Iv. IBIL & 901. Bcr. d^dtnixti. <b*m. Q«mU-. *V.
07
AfTKyVlT.
Thin *ulMlaiioeiNobUineA lulbe final pTwIuct of Ibi- '" '' ' i'
rabio ur liiUi-erdlii. It ir hv^l I'reinrod liy M-iinu u|>^<
■dil In praMdconf aI<^i1iii1 ; tlic vniioas colon iif (Iciieliu ■ >^.:l' i'--i' -•■ ■
ud the fin*] redilUli-vpllow i-aliiiion, ir p<iur«>l into wni«r. j i«l<lii ii pf
nholftcliii. It U not vrvKlnlliao >nd U loliiblr in ■luoliol. rthrr. and ivi.
Wlion iVntit)- jiTcpiued It tMUiu to gift) ma unci-rlsin nlnurption )iAn<t if ex
in an <jn<l Hitutlon. On thh acvouul oomc obipntnr line brco l«l lo n-i_
Bt lilcnlicHl wilh tiT<)rotii1inibin |urt>ti<lin|. Tb«r« ia. Itotrercr, no doiiU Hut thn
aiv <)uit^ distinct biHtiisi.'
HydroblUrubln. CaH^N.O,.
Tbifl brtdy nM firet dnuiribcd bv Maly* iw mulling TroiB the acttoB of Miliiua
■Kialgam nn an nlkslint! I'llulion nf bilirubin. Wbca Ihv rMrtion l« cam^ru
the tolullan i* pntcipiLaiiMl wiib liydrocbloric acid, the pr«cipital« dixolvM ia
aminaiiia. again {yrvcipltatrd by ncid, and Uie sub*ui>u« thui finally ohuin4«l U
wwbcd ailii water. It ie rcadilv wilublv in aicobot. Itm w In etber. Iia aikaUar
4olntion* an- rcllov, and titi-a* turn pink on UioadditMn of acid. Dotli ilsicU
find allcnlinr mlutiuun, tbc hitter eapecrallj i>n tlir ■ddition of a tev dropi uf
chloride of xini-, jcive a i-barainoriillc abaorption band bciveeu b and F.* In lb*
coloni of Ita alkaline and acid !in1uttnn« and ibe greeulab ttuoraacenci' of it* aaaao-
nUcal aolution in tlic addition of chloride of dnc and in ilRabaort>iioo •iwclnia
bjrdnbilirubiii nhown itn vlme relatiuu to urobilin (>c« bclmr), vriib which iBdM4
it ia now oTuJilertHi to bo identical. It la aUo ideutical with a bodjr luiMd *bc-
cobtlhi' wblcb hK<l jirevioiisly been dMcrlbed. as a product of the alteration of ihr
bile- pi gin en t* in the ntioioota/v canal occurriiijf in fecm. There ia do diScnkj
in MeiiiK liow llii> chani;i- (hydroeenation) can be brou](bt about in the inteadK
ffnre it t> known tbai n iroiialderable <|iiautit}r of hjdroKoo tnay mak<' it* appav-
ante by feriii«niiitlvoj>ri>ceucfl In the ioteotine. aud in ita niuceot itale mi^t
raadlly timdiire the niinpte change which ia known to occur when bilirubiD '
convened into brdrobilirabin.
I
PioHcicnt OP Uuxs.
Our knowjmlge of theae bodies U at prcwnt limited and impvriaM.
probabt}' tbe;r are numerous, but only t<n> appear HUlScienily well chancUriifd
to deserve mention here.
Urobilin. CJlJSfi,.
An itatvl above thia is now rvitartleit an Identical with brdroblllrubED. Il i
liret df«cribml bv JaflV^a* n well i:hnnirtrrit«d normal urinary pigment ud lO '
identity with hyarobiliruhio inibie<|oi>nllT deti-rininc<l.*
Normul urine conlaitin only amnll iiuaotitieH oi' urobilin but iherr ia prcatM a
•nbttance (chroninKon) which under the influeiK-e uf acxi*. with abnorvtiM ^f
oiygun. yield* urobilin. The urinr of lever frequently i-ontaiiiit a con*ld«nbl*
aoaount of Hrliial ortibilin ai aueh.
The jiroiiertiiM dracribed above for hydroblllnibio an identical with Ottmrnt
nrobilin. Iih prei>«r«l<on from urino la Mmewhat difficult, and for thia
apeclsl nianunl tniu't beconnuUud."
> MMy. AtilL •■ WdaAkaa.Bl.lrtXItMliIAMIi. Pltr.. iind M. Itx. IMt: «AUlli . AhH-
am Ihjnilu* mA liaBpbrtl. Ute. clt
> ]|ifFiul«* mwl OnuMell. loe. liL Slukt1i.Ouiii>Ib. td. ued. Wlw., X.s itilnTJi. a.m, I
•Ma]y.CautKil>.tdBed.vnu..No -ii<i«fi(.a.ci. UttonMon.l-nLMn Anh iiiuiikmI
KtH. ■- 1
•Vnnmlvr. >1. mnl. Win.. .\o.»Ltm. AniHl. d. Cheifi., Ud. el(IU(U»l,&TT
' \-leranll, Z^l ■ - h r n,.i . M. U (IKSva toi,
•Vnuulriiii i.inlb.td nvl.WIM_Xa.XI*'!.
' VKronl:. i' : •trinKlaUDalna. *tK. TnUncm. fCC-A U.
' Wnlrall. ' ■i.tMBLS.-JU. Vlfeliow-*Ai«k.,M.altU<t«9l.am
*Hal]r.ADii J i.Ua>. u. rhariu.-lkt clHll (ICrA. H ff.
- VI.I.- .SViitxiiU't and VofrL lUrnansIpD. rI. Mil (IM)), & *!.
CHEMICAL UAiilS OT TITK ANIMAL BOOT. 1059
lb VniSitni lu be the •ulMlam-c which k>v« to tlw otine of rbcummti*m it*
dwactcriilic color. Wrjr littl« n koowii i>r {ta ch«iatcal |tra|>«Kl<a.' It npfwam
10 bv in ■morpltoim ivJiIirIi body nith uii nci<l raactlon, nlowlr soluble in water,
»lcobi>l, Htiil rlher. Whrti tmttrd with cAUntlc nlksli i[ turn* grm-n. Urine TOO*
lAlnioK this bixlr tnlttii on ■ i'haractf-ri>lic rcddiith-yelliy* color on the addition of
riinc«ntiiite<l h/droclilorlo acid.
Ikudldiuio ocoitiilan that ooruul urlno Mntolu aalf oaa pIsiMHI, wlilch he <alU
■rodirona.* Ma); i* iiidioi^ v- minnl thi> u the ntiia m iirobiiin.* Mora rvcvnil;
nmdldiaiu bu oplislil lib furirivr viuwi.*
TuK l5i>tao Sbkibs.
iQdloKD- C.U,NSO,.
A bodv WKi IiiiiK uffo dniL'rihrrl' uoccurrinjc in th<' urinp iin<l iiwnRt nf men and
otlier aninwlii wliiuh yiFlcieJ by tlie action of ncida the hliie colcirinjc inaltL-r indigo
Hsoiieuf lh<* prodiiclAof ilsdecauipmitioTi. 8ehuiik cun«itler«(l thia Bubstiuoe tu
Im» idenliral with the indicaa known to occur ioiev^ral planlo (Indlgnfera, (lalla).
Ho|i(ie Se.vler.' on the nthrr hand, having r^ard to the urcater cuuie with wbicb
(he indican from planbi undrrgoea deconpoaition, reKnrdcd Ihrm n* mtat probaUv
difliBrttat ■ubstanom. Banniaoa haa «bo«D' that Ibe two are really different, and
hM eoollrmed hia earlier ntulemeRtK in n more recent publication.' AccordiiiE In
bioi. Ibe indicnn obtained fmni urine is not a ^lueotide (m alao noppe-i^efTer)
and yirldi ititiihuric acid by the action of bydrochtorir acid. lie aaiiirni to it
the furuiulu C,H,V.O,SO,.0)I. and regard* It ju induxyUulpbaric acid. The acid
Itaelf 19 Dor ret known In the free stale, but i< vleid* atabfe aalta aueh aa tbat of
potamluin. C'jI.X.SI) K. It on-ur« largely in the urine as the rwult of the prea-
<iitt nf indol in the Hiimeiitary mnal. In thi< nay llaumann aod Hrie^fri' wore
eiiablrd to obtikin laritc <|uanlitieB hr i;<v'')|! indul to a doK> I''or it> proparatiaa
ibeir original paper mint be cunaiilleil.
Wben treated in aqueoua Koluiion with hydrochloric acid iu presence of oxygen
it yields indigo blue
rc,H,NSO,K + O, = SC,H,XO + SKUSO,.
It ia always otimaied in urine by conrersion inU) indigo blue.
iDdiso. C,H,NO.
It ia firiiipd, a> ulated above, fmni indican. and giro rM to the bluiah color
•omelimn 'ibierved in sweat and urine.
It may. by *low fi>r[tiitti'>n from indican. be obtained in line crvstals ; these are
inaotuble in walor. •lijchtly aotiible, with a faint violol color, in alcihrtt and ether.
Obloroform aUo diat'ilves ibeni to a aliaht t-ttent. Indigi) h loluble in itrone
Milphuric acid. I'urinioK at tbe tttme time two compuaiiaa with tbi* acid : Lheie
■re Miluble in water. It poweaees ajture blue color; when presied with a hard
body a red'tlnh copper clored mark M left, and tbo orytlats oshihit the *ainocolof
If seen in retlocie*! licht,
Tbe luliible cmnpnundii witli ■ulpbiiric acid give an ahionitioQ htkinl in the
niiectram which lies cIum t» the D line and to the rtd side ol^ It. This may be
iMOd to dried iadi)*o.
Treated with rediicinit a^ont«, indigo i« dae^loriMNl, beioK reduced to ioiligo-
while. The latter coniaina two at'tma more hydrofreo than Indigo.
< Hclki'* AretUviIifli. mnvxi, ^ mi i BiiLM«d.n . N.B.M* W1. |BSI, p.lM-
• Mal». Ann rt Chi'm u nuit™ , I'l.' oi liTt. * M-
• JmrB.CIwui s>eL,H« ;.ToiiiIni'<:i;.pfiiM.«il. _
*i«hllilk. rtilL »u. val. X. K n.xiT,p.2X: XT. pp. A 117. m nKOLOBMnlb.. ISM, 8,10;
Uar,a«TllBML» W. Il<>pi'«-ite;l>r, Anh. tpalli. AuauBl.iiTU. am. Jsdk, l-AU^M-i Arab ,
Bd.hl(t89m,atM
• Oiadb. d. vxh Chan. Anal . VjI iTfiniii, 8. m.
'ntnnr^Aitik., M. alUiixTit), s nil. ZalMCkr. r. pliT«laL Cham.. M. lOVn-ln. & a),
• ZsMHh. t ttjial, Clifn . h>l. Ill nont. a. »*.
*tilmai.tttr*iA.nem.nn iuaWf.iW aMal*envr.d,a«uMb ClMm.a«silUall<Uin.
•D.U01U1S, nM:asdaUI <UWj.«.«M.
loeo
APPENDIX.
Indol. C,n,N.
To ibh body (li« gpecifli: cwlnr or lh« fecn >« Pflrtlr dor. It to obuioMl I
liiinl prnduci of Ibc reduction of indigo, mnl mi»i> bjr ih* dinlilUtiun i>f
natlcr with nuirtie alhnlim.'
Il olirn <iccun among the product* of (h« ai-tion of paDereatk- renDeni ob piu-
teldt; its jimence in eiich chbm appean. hovevei', to b« due, not to th« acUoD nf
the trymin. but to ■ *imultnncoiis piitrvfaction iindrr ilio inlluriu-c of baeUhi,
etc' If tbo puncrratii: diftmlion be carried on in the ytcrvncr of mIk^Iic add,
indol doM not tnak« lla appi-aTPiice. Iitdol b a cr^aiafliiM body, aolttble In ball*
Ine wat«r^ alcohol, and cturr. Il paait» ov«r in ilie U^am «h«n Its aijutem.
HnTntloD IB boiled, Il le chnrnclcii/cd by tho Ibllnnine reaction". ,\ »ui\i ii%
Sine-wood moinlcncd with hydrochloric add i> colored briftht rtiniMin oiira
ippcd into a solution iif indul. It* alcoholic iioliiiion luriin red «hen traM4 '
Willi nltrouH ncld. aud !■■ aquvoiis iiulutioii gIvM a copiotiH ttd pieclpiiBt« wU ,
the Mnie rcsgtnL It also jieldA a chars ctrristic crT^talliDo oonpotiMl witlij
picric aciil.
Skntol. C^,N(T).
NolJc<^ by HricRrt' a* onr ol'lhr producl* of pntrefnctivr chan|[<-ii in the until
inieatiot. I?«crrtun* had previously dmciiWd a tiniilar *ulBtaoi.-c an eriBing frMi
th« putrefaction nf Hlbumln.
Skatol ia crjMallinc and c/intaiua nitrogen; iL ia mor* aolable in wain dian
indol. and doc* not pxi- rii>p to any r«d coloration with nitrou* acid.
Skatul readily pa»M into tbc uriue when it uccora in tlio alimentary canal. a«d
then KiveB a viulet-red reuction with Mroog hydrochloric acid.
T. Kenckl' prepares this (iibatance by the jiutrrfactlun of a niliture of Knrif '
diTJded pnncrena and Biiiiclc siibKlHiicc. After the addition of M-elir add th* i
Biaxt iadiriillcd, when the skatol rendilr piiuc over. Frcni ihi- dinlilUt* Il m
precipitated by piciic Mcid, and the predpitate when attain Oi>>iiltrit oiihanii
gives olF pure alialo), which a>ay be finally puriSexl by cryatalliaBidin.
I KaLno. B«r il dtuiKh- cbcni. (looU- Mil. Jlimi, B, JBO.
• KUbiir. VsfliDiKl- HcliLIIi. I>uliiil>ltl.-iiit4. Vcr., ii.9., Bi). i„ BR. 1. BsrtMild. tNrvUolm (kiw
0,^.ii^ti„n ivT. n-jj*.
|W^. rlicm UvhI)., J*)iFi; i ilNTTi. I*. 1(97.
■ I -III iPtitrrrBfildii ilu I'ulliuniiiit, llaiioin. I'M.
■L...,,,.,L. ;. il, luiiiJ.WlB., JKW.JLM1I.
INDEX.
ABSORPTION", from olimenlurv ramil,
4111
uf ilittuxilile siilh>tancei< imd wat«r, 4'26
of fnw, 4'2a
An.-nninicKln[ii>n, «9S
nierhunisni of, 8i)8
Atrtd -albumin, KVi
formntinn of, :in
AciiL a.H|>arlic lUo2
lieiiznic, lOii:)
Imlvric, 1037
cniiVi.', 103H
mpmic, IftiS ■
ra|)rylic, HUB
(■lir>rii', lOVi
«?lii_vlene-luctip, 1041
ethvliiltne- lattic, 1041
fiirniic, IC^tT
f(]iiliiminic. 10<'>3
Bl.v('erin-j)li(w|ihc)ric, ll)4-!
Klyc<K'h<iIio, low
hypiiuric, III.Vl
kvnurenir. 1051
lactic I Ml
laiim-slL-uric, lO^tS
mvrwtic, 103S
oli-ip, iO:ti>
nxnlic, 1042
inlmilio, lOikt
|ilicnrlii', 10">4
liniprinnic. 10:iT
wircDlni-tii'. 1041
Mlt-nric, IIWM
Hiiifinii'. 1042
taiiriH'linlif, 1056
vnU'riniiif. lOHM
Ai'fluKlic a|i|)iiratii», 1)4((
Adlpiiw tissiif. Iil4
AHiTonr iiii|>iil*<, 1172, K55
Afler-iiiindes iii'ipitivi', 1)24
jjiwilivc, !'24
Al™nlr>in. Hl'Ml
Alliuniin, iK-iil, HIIK
nlkiili. Kll'.)
ilcrivol, 1<ll4
pffU, 1017
nalive, lOlT
Htnim, il>l7
Albimiiniiiir' jrliirnis, 1S2T
ohmiKU-i ill, 'liiriii;; Myrctiim, .141
Alimcnturv c-^iiiu1. tmiwiis iiR'niUraiiit of,
aiT
mii-ioiiljir c'lal <•(, ^117
utrurlilrf rif, :117
HiibniLiiiiu-' •-■oat of, ttlS
] A I kali -III bum in, 101!
i Allantoic, 'jy:(
Alvi'gniat'K [lump, 454
Alveoli of lunftN 433
blriicture of, 433
I Amblvopiu, 843
Aniipfia, cliami'icrislicH of, 33
Ani'i'biiid movementH, liiT
Anelettrotimils, 131
AnimnI binly, (-hemicnl baneii of, 1015
Animal heat, 041
diHlribution of, IHl
pniduclion ()f, 045
rejliibition bv variationii in loto^
644
wjLirrts of, 641
lem|>erutnre of bodv, 043
Apha;<in, H23
complete, H27
|>artii(l, )127
Apnii'n, -iW
Aqueons buniur, Stl3
Area for muHtiention, R14
l>nMlnction of voice, 814
Kpecfb, 823
HwalloirinK, 814
Arteries ehanjn^ in calibre nf, 2(t5
stniiiiirt' r)f, 178
Arterioles, 178
Awiiew, 417
AB|iBrogine, 1052
A«i)hyiiiB, 402
A.'tiKniali^m. itDd
Atropine, lu'llnn cif. im heart, 262
im pnpil. 905
Aiidilory jiulicmeniH. U52
Anlomnlie action^ 171
I ALiti>nintiHin, irrvgillar, 870
r PfKular. 870
I Axis^linder, 121
■ Ax isH.'y Under proee.*es, 186, 80"
BILE, Ml
acidi. ;i.>3
I formation of, 508
action nf, on food, 353
I antiseplie ijnalilies cif, 'S-'A
' charaelen of, ;t51
I riini[«>sition nf, :lol
I formation nf ninHtitlientx of. 51)5
inHnence on )icpli<' ili){L>f<tir>n, 39'i
I piKlnenli of, :)52, 5il5, 105ti
', Cmulin's te^l for, 352
, rcwriilinn of, 3(ii>
^^^loSl^^^^^^^^^^^^iNDex^^^^^^^^^^^^^^B
l!i)iiwlt>.:i>3
HUiwt. roll- of i-tniiliiii>iti. iV9 ■
ttvw Tor, 3&S
n-lilioiuxf "tii, ti ill. (.Vi ■
l>rtll■lIk«^M'^ x-a
rvsjiinli-r^ it, t'll M
*n-n!<io:i of, ^11
"'11''.*' '"' '" <~onlr»rtU* S
miiriibiii, 3f>2, UM
•iii~>. 1 1 : ■
liilitvnliii, '^''i. Wu
nf li^or. M-J 1
Itiiim-iiliir visiijii. tC!ll
vnvoU. (ii)>llbrr, ITS ^^J
lllndili'i, iiiiix'k-- nt, ■•i*
Knin. J^H
-irorl'm' '•(, ''iti
>ltH-iin- of •I)(iit of mltllMi andW^I
Illiiiilim, Mi
Bt«v.>n4 ^
kill) <|ilnal nirri. Imttiliniir amafl
blind ■w.>T, tiiiiiitc ui> •< vm
inrtiii'iir. - 1-
IIUxkI. 41
nicnilifaiic*
mrlBink- lu'i.l in. -MMI
rRHIllar ilrr.irii.-« »•< ,m> <t, r-j
cliiuiK"- ill •[iiiuiUlv Mr, "iOl
aiti-riiii' of, illMrllnlion nnri rhaiwM
rllcniiiql (i.>iii|H>«iliiiii i>f, Ul)
S--!
ritillinu i>f, U
IiUmhI HIIiIiIt ■•{, «t
>ici Itmu ■>f Inm-iimilni:. w9
biifly luil in, -U
vuUMM of, 'rj
bulb vi. Tin;
MqitUFli-* in, SS
6bta> nj, 74s
i'ni>Miiiii'tiiiini III, 4'J
ifiitral itniy iiiniii>r nt, 74>
(-nii'l "r iiciilrnl -ulwoil, i-*
i^Hn'l>rii"-|>iidl tliiiil, '*>^l
rtTvrl \'f "-Kliiiiii clilori'li? nil, -H
i'hiLiirtr< ii* L'TiM' iiiAltrr of, Tl^f
i-llii'i <<r ii'iii|H'niiiirr i>n, -17
(Tniniiwiinil flhr^v of, 7I<1
lilirin in, 41
i"iin(i-1)'-n- •-! ^itiv am) h IiIiu a||^l
filirin rrnurn) in. ''>(>
^H
lllirliiiiuv'ii in. Ill
nirjii'"! ci-nivulntB, 77'-' ^^|
in ftrit'rit^ ■'•!(
iliiiiilriKMniiM, 77;! V
in vfiiii, :^•i
t^rar mMU'r cK, TTS 1
ii-iiffili -ir tiiw of, '1:1
n>r|ni> •(rtiiliiin (if, "idi ^^^M
IKinnilt-liiiliu in, -Ml
liliri'x, 7TH ^^^H
i>)iuniiiii- iii. 44
(-mninl m-rvAuf, 73<l ^^^|
nii'iclilv of, i:i
oiiibminii', 731 ^^^|
■rnini in. A'l
lilirv <if. 774 ^^^1
■enini-aliiiuiiiri jn, -M
fiiiu-iinio of, i*ll5 V
color of. <'>4
tnu-ili* iii»l (iiDmlr niirlH tt. TM 1
briulxiKw ••r. line ii> rrllLHli')!! iif
eny iimlUT uf, 74H I
hiiii>I>-;;ii3l fcalillts nf. TW 1
lifrlii iliroiifrli n.riiiiM-lt^ .'iil
vrniiiix mill iinvriiil. f )',!
itiliTiiinlixir Krnr ninUor nf, 7K ■
OiliipixiliKii III*, I'-V
luneiliiiliiiHl lll>n->»f, 77>'t M
lIUM'Ik 4-Vi
nnlun.' unil rdolkint of lllr t^^U
iiK*ili"il>-f>ri>htalnln)[. ViS
iiiu-li'l iif, 7lM ^^H
llirlnlitili. -tiM
tilliari iiiii'k-ii4 of, 743 ^^^M
fi-niiim of, I'H
o)fiii- iliiilimiiis 7K ^^^M
lliplhnll iif •'•lilalllinx, 4lH
(HwiiTiot liiiiiilliH of. '(to ^^^1
IlHIIllll, i'U
]n»U'rii<r r>.>niri<' 1/. 777 ^^^|
iTTalsU i>r. 4114
|)yriiriii<lui Imrl <if, 77o ^^^H
llntiiiiutol-lli, 4''iT
i|jlniii'tiiilf fiiiii-liiiiii ol. SUV ^^^B
oliuraiioP' "f. 4-'>T
pltui-liin- iif. 7-'(l ■
nii-ih'Hl of uliriiitiinx, ihT
r>iii*rfii'inl pnir mailer of, T62 ^^U
(fNvtnwojiic rintum of. 4SH
■>ii]H'rwir ■■! •f'liH.n' ilmi-Kili«ii,|^^|
iiiriclitw iif iTTHlnli, 4.'i7
lnii!l^ fdiiii ix,r|K>n iimidrii^tf^^^H
■iii>ihK<ii'>i;l«l>lTi, 41V4
^^H
IMflllwIc-* <>f. S'[
vi>nuiiH iimnKvavni iif, tIKS ^^^|
>vi-nii!;v miiiilirr iil', in liiiiiinii
inkllri tif. Hi i ^^^^H
lil™«t, 57 ^
ItliK'ii'* iiaivnliitkin, B2i1 j^^^^l
rliiinu-b'ri'lirs of. '>t
Urundiin. I.-Ct ^^^M
iwl. •inirtupF 'rf. ■"■«
■Itiu-iiitv 43-> ^^^^H
lurni<>fllo)>ln in, o4l
Broiii4ii<>lc^ Vii ^^^^H
wliiii'. 1,^7
Nlnu-iiin- «r, 4% ^^^H
vlln-l of Oi-liL-M'iit BrrntioD on, 468
Oiiiilii<'h. rotuiinu- uf, (Wtt ^^^1
Itlntvlrti. U»
^^H
]iliH|Uns W
1
jireiKinv 1'*'^
pALAltAIC liran, BCtioti of, nn laoH
■iirvn from rnmllil of nifalill. IXP
quMiiily of. It> iIlMrDiiiiiiin in I>'1<. 71
('*l4riiiMTm^ fi39 ■
1 X U E X ,
u>m
('n|iilliiL'i('S ITt'i
limilaliiiii in, I'.il', '.'S'i
I'niiii lui-sfiiti'r.' Ill' «iiiin':i-|ii}ri '"li
lin->siirf (if IiIiiihI in, I'.HJ
%i/i' "f, 177
-liiinuiliiiii'iir .siiisis if^ll
Sinn-inn- nf, 17(1
vi'-Hi-ls rif niiwli-, 'Mi
CiirlKilivilmtt'^, ]ll'.>, \m:i
CurlHinii' ai-i<l I'xIiiLtttI, 4'itl. -170
(jinliiM- (■•■nlniftiim, fi-iiliiri-.-. nf, ■.'-"ill
I'lirvfs, disi-ii!wiiiii lit'. 'JIM
^'1 II nils. :l:.1)
fniiii <]nf!'H nlnniiicli, i!21
iiiiliiil«>, Ul 1
plNisi-^ <liiniliim iif. '2'li
tU-ilH', fLNiliircs lit', i'll
Caniin, lu'il
Ciwiii, :ll.'j, l(l3i
Cdl lnHfy, Hi;!
Cells fill I'll III' I in I. I7tt
i-]iilln-lii>iil. 17U
piMi't, -1711
livniiiiiiliil. no:!
i'villr.il iiiTVciiiB .■•y<ii'ni, ViL-iiiiiiiliir fiinr-
liims 111', "27li
CercU'lUir Inu-I. iiiiirw iif, 7'Ki
(.Vri-K'lliiiii, iiir|>iis ill'] II lit 1 1 til 111'. 77l!
fiintrniiis iif, ■ili.'i, Mli7
(jrjiv iiijilUT iif, WM)
lii'iol.ip- >}{. W2
Mi|nTiiir |Hilii'k' of. 7WI
t'ervliriil <tmviiliiti(>ns iir«i> nf, "Hi
'•f iIkmIok. mi
iHirli'X, NIK!
lii-iiii-[ili<Ti-K, [ilu'inmiciiii c^liiliiliK! I>y
:iniiiiiils ilf|iriviii of. 7S4-7'.HI
"[K'tiitiiiii.-i, tiiiir tiiki'ii nji liv, .■i7"i
Con-liriii, \-i': 11144
(.'erfliriw-iiiiiiil lliiii], ■'•7'.l
tllTVl-N llil
(Vri'liriitii. miller siirfmi' nr li:iii<' nf, lllll
Cliiiiitn-?' nf liv ill): lis-uc, l>4
4 'li!iiivt-iiii iiiiil l>l^(f^^i liii'iiiiititi-liiiuirlrr,
■2\r2
( 'ht-lllii-iil i-ll;lilj-l-^ 111)
('lifviif-f^lciki-^ niiiiirniidii, 41il
( hiliii, Id^C!
( 'li>>U*(iTiii, li.'i. ■.i.y2, IDl-Z
Oiiili-ii-liii. lll-'h-i
I '111 mil rill, 10:11
('lliinlii rvilivii. '«'>'.>
tviii|iiitii iicrvt-, imiiti'i' :inil iirliiui nf.
■ :i:t-i
t'lmniiil iimt, "■•■h
4 'limniiilif iiln-rnilimi. 1117
Clivli'. I'liiinii iiTs nf, 4HI
Chyiiii-, ;is!i
Cilijirv tiinvcnii'iit. |.'"i4
rirciiliiriciii, niUM-. nf irt^'Klilnr livurt-licat.
■2'.a
i'iminisiiiii<i.>?< I li-tt-rti lining i-liuriii'lrr nf
tln»-, hi:!
Hli-el- nf !ili-ii!i(>l nil, "Jllo
I'Hi'i-l- •'{ I''"kI nil. :i;is
fii'Iiil, ll)IHI
< iron till inn, liyilniiilir |iriiiri|il(> nf, Ili:>
iilHiif-mt' nf i.'K<'n*isc nii, lillli
CiMi's, (liHiiniiin, Slk'i
( lurki-. 111] u mils of, (IS.'i
i'oiilsnf iirlf'rU'f. 1711
of vi-iiij, IMl
I'lii-lili'ii. 1142
(nlil, Hi'iisiilionK nf, KTi]
Ciilnr-bliniliiess 1'2:!
sonsjuiims, I'lis
Co Inst run I, li^,"!
( 'oJiijHiKjtinik nf liiK', '-i^tl
III IiIihhI, lill
nf gimtric jiiiiv, IMK*
nf milk, li'.>4
of |jfrx|>innioii, .'iiil)
of «.^ivl^ :Wi
of Hiiirviiiu IkhIv, (>:iK
of iIk' iinitiinl IhkIv, IJ2X
of iirini', '>'2!^
Cnniiii'livi' liiwui', 174
Ciinsliml I'lirn-iit, lU-linii nf, l:!S
I 'onlnictilc iinwli-^ HHi
tisswH. 7'.;, 141'
( 'ontnirlion, lirfiikin;;. V2',>
i(lio-iiiiis('iiliir, 141
iiiukiiiK, l:i'.>
li'timii-. 74
Hiivi' of, 5l7, l-'i^
Cnnniiimltil niovtiiu-iitK, nincliiiiiTv nf, 7!KI-
7ini
CnnI, Miiniil, 'i7o
volitintiul i»i|iiilws ill, KVl
Cortuii, .SS7
Cnniti uninionii, ,sti7
Cnr|>nni ;n'niriitulu, 772
Cnr[n>ni i|ii:nlriin'iiiiii:i, ftimtioiis of, Mi7
I oqiiH iiniiilii, 1^2
liiti'iiiii. "N.^
Ciir(iii.-«'li's. 1 1 •limit ivc iis.-ii(', 174
ntiiiili(.'r nf. in liiimun lilnoil, '>7
■if IiIihhI, 5-1
( oMi-x. .Trc'litiil, S(i:l, N7I)
!li-(nlo«y ni; SIKl-MW
ji--yi-|iii-:il |.rni-t\-**s in, S71
Cnrii, nrj;:in nf, 114:!
I nrtinil iin-iis, rcmnviil nf. >21
motor ri'fioii. J-lO-Sl-'i
< 'nini|>s, miiH'tilar, l-'k!
(■ryMiiliiiu' lens Mi2
Cnrrcnts nnistunl. 7")
olit-lmtcinic. 1:12
iiiiliiiT(l, 7,'"i
nf ai'tion, Mli
nf t^.'^l, llli
( iirvi-s nf i-ii'liM-.irdino |irc*iirf. 217
CnliimiiiiH M'liKut ionH, K.'il
Cyatin, llJ."i:l
|\A>LKLI.S Iminrv, 7.'>
I' lifiitli. 101:1
m'finiiioii. ;!'<:i
l>0)-liililion. :[7ri
Ih-|irf«nr iiiTvi', 2711
l)i-rniis, .V)2
1064
lVt'('lr>[iDirnt "f cmbrvo, 090
I'f (.luci-nla, SM
|)v\lmi. UW,
l>riirDM'. :hh, laia
Diiiboll!-. oH.'>
I^DgTMnmnlii.- cvf, HUt
ItUpliraicni jv, ii'(v>|iinti>iT uiiWkln U4
Diet, tiotmnl. lifil
Tiii>)ilii.iiti»Mi< n(. (bIT
Dtltitfiiif ill h9|nniii.,n in wxo^ 4H
Ilixi^li-ili. ;U)1
l>ilntntii'ii vf iirtcni«, IN."!
l>i"|<triL' ll|•|■l1mlll^ inijWTrrcijoiii- in, Wfc*.
Ea(i'li.'iiii>iii>H '4114
Di«fliiirK'- iif t-niTity, N7i>
r>iiiri-tiiA tVIA
i'ivixinii^ [if oir, fttn
*if 'jiiiial (vini. 677
thilnii.-Itcyiiioiiil hey. T7
DurlH i>f iiiHnininrv Kt!iliiJ>, li'M
Di-.i.i..™, 441,+>(f
EAR, WiV
fiflllcii nCiH-J
■liviilriii I if. |i:^
KiwlJicliiiin liiU'. Uth
liitvraitl. -T lalivrjnili. Ml
UKiiiiii' niiiliiiiriiu piicrniH. «t1
nifHiliriuii- iif |{i-i»n(<r. llj.'t
mi-iiil-nini iyiii|«viii. mil
iwniliniiiniui lotivrlntk. M>'i
oivaiii.r ('•■rti, ',11:1
ni»n>ii> lulitriiilli of, IH7
■nwlili- <if. HIT
|i[rrlKin|di. 'Mt
|>lii-*|o].,i!i,';i| nunininv ,)f, |1S9
*Iii|Hiti<in niilwlc ••(. 9^»
Itniior lymjBiii rhwIp of, SI"
Eili-renl iiu) III !•(>«, *I72
Eliuiiii. Itn-.'
P.lwlrinil rhunxv^ 118
"limiili, .H
KlninNK'^ nnn-tHiUriiMlilv, ll:i
Q(i4n>ii>iiic furrmt\ Vi'i
Tariaiir-Ti. 1^ irrilithililv (hiring, l-ll
KlfiJU-Htnri- aiin^ tneiumt* uf ■liilinl. H^'
Rlcvvnlli norvii, 7'ilJ
Kmlwyii, tiiirritiim iif, siil7
Knd-|>liiii^ VIA
KiierBV, 'liH-haifv .if. «7IJ
f^lMTnliiun- of. fa*
iiiK^iiii: (rf, AX!
of inn'liaiiiinl mirk. IHO
EniD|iili- |>li(>n'riiivnrL INM
K|■t■l■■nlti^ A.'i.1
Kpjulnltit. Hit!)
Kpili-|.tiforiii i»nv<iUiiin>, Kls, ««:i
K|Ntl.fiiuni of ilir .tmiA, fanrtwno of 341)
PAt>tnrliiiui iiiln'. !Hn
Kiir«i'ni, JS7
JCxtiiiliiiiiiii of aijuauiiK rapoT, WO
of rmrlmiir arid, <S0, 470
of mgnnii- luallcn, 461
1SDK3.
ElllalMKin. r«ll«ii of. 1471
Ksiilnitiuii, «■(
HfnnKiii (iiv, iia«
(■llUrr rrnlinl of. K^il
•tiuk-niiiiiiuiH-, !i9l
i-mntrlnifijc, "UT, MW
Ix'tioHiul tviltni of, 81*7
hvjii-niwliri|iir. dUt
luivbiiniMiifi •■/. liilT
■iic>VFmMit4 of. ;i;in
niavk> or, li-tl
niyiij.jiv KSW
fliKnii-iit ivll. .rf. nail'
CAdAI. i.,n-,v T.'hI
I loiirw of, 7.'»)
Fill li>i>i nil |iil,r^ i(;s.T
l'|nr:iJiMT>iin, S4I
I'uU, •-i>ui|>lcx nllr<>i[i-nnu>, 1ii|:(
ri'Diiutloii of. (ll»
naliin- ..f. ill iiilljinv liMiir, fi|t|
I (Ifiiiral, liii^i
I llicir •trriralh^ iukI alliif^ UKIT
rfii-f. :ill.'j
[j-iiinlc jnmi-ratiTv niiiatniliH. W2
1' ilir»-Mitli-reul, 15U
cfli-n-nl, |.)ll
■ifn«.rv. |.)U
luMM-iifKlrii-lar, :r7(l
I TiHiililnior. 27(1
nujrur of, 'S'i
J'll'Hii. m H.niiiig of liUni. 1 1
lil'riiifim-ii. 1)1^2
1-ic-fc'. -j.ririK ttinnniuclrr, a^O
I'illd ni-n,.. 7'Vi
lininriMn of. 7^0
I'lmulctiiT, :RJI
K.iCiil dn-iiliiit«n, I(m«)
, F'ltlidc irf hiiJr. .VV. ^
Fwid. u-tioii of Iillo aiul ■•nnernOlr Jnbvl
(■mnll iiiiiiiiiit-. :i!il
dmiitti'^ ..f. ill )ri2v in|4^iiic SM
in luimili, :w«
in •<iii;ill ill till Itii', .'tUI
{|| •(oainclL, ^1U
vftvbof fniiT iind rarlH»IiwInH>,i
•■r pn'lntin uis GSlt
wJi ai^nSll
-•llll^^ :tii|
I'lnnHi lit 3(tJ
ro'iriti jiiTiv. 7'>l»
KoiMi l>i-iui>-|>litni«, 1142 ^^
riuicii.uH.1 iiriiviiv. inttimiiT oC nn ^mJ
iniriili.-ii>><t»,144
G.\ T,KX. it'inn of. Xtj
<Jall-(rinCN iV^
; (ianfflin, ^ii|inthDlir, lilA
(nuiirilnnic tio-irlt. lAJ
thuiw in liIfMiil. (;ii
I
iitutr'ic nml inlnliinl morcmMiu, niTvoiu
DKrfciiniuiii of. :tS'l
dlfiatian, HrrnitwiMwtM iillwtinK. :<i:i
drrtiRMtmcoi NtI<rCTi^t^ tcmixTn-
lnM\ :1I4
KCtion n(, on milk. :ll-'i
nil [itiiifiiK :mh
rhiiiiionl olinntt'lcn nf. '.¥1'
fcirinuliiin or Irrr ite'ui ••(. 'M>
nuiiirr lit iliv u'lion ■•f. 't[4
nx-rvliiin I'f. :t)T
U«lMui, \74. 1031
Otoanilvp <in(*ni^ fenul«v Wt
ShTximl nnnuimv uf, M2
. >i'
Krtivmt iiLtlurc iif. HiH
nf lEninnrr. ^>7i
>if l.k'lHckiiliii, :i7'i
(llJMOIl'!! calMilp, olUl
<iloUn. WH
(iUiiDiTuli iif Iriilni-y, fll7
(ilalll". mirninmK ■>(, ITl
wilk-L1iri|C <rf, *j7.
liliitiii. KKIl
niv.i'riii. VHO
lllvdii, HIM
((iTmiii'ti, iin>. ut:i. milt
■■iKnu-lm 'if, STIl
iviiivi-t>i»ii at, )bi>i Aiiinr Inr llivr. 473
in nmx'li'. 'M
ill jtLirvni^ A^4
ii*-* nf. ill tivcr, oSI
(imi'lin'' lr<it fur ink; 3A2
lioll, (i>l<i>iiri<' <it, li>*lt,
tinnlinn r<illioli-. 1)^7
liny inallrr. Iflll
C iTOTp'" bnliiTc, 7 A
(luunin. I<ir>l
limlili-n'' niniiiiiiKurv, "10
tii»nsl<>rr liiilU, U"m
iniicHU'i iiiriiitirnnc. |ihTiiioIi>g!cBl anM-
omv <ii, IL'iil
H.EM \t \-n 'M ^^•HK ..r «Jow,rr«. S!«
ll'rml■ll^llllllm■•l<'^, Vnlkiiiiiiiri'p, AN)
flmnatai-lioiiirti-r. OixitYnu iiul Likrivl'*.
a)ii
himntiii, 'J64 ,
Ilnntii. 4tA '
IlapiiKiiclobin, 4^7 i
In r*d I)lwKlH«nni*'I«^ M
Ikin.AM
ttruiiax. ivnalUinii of, HAO
Heart. 'JOK
^ -ImM, uigmoniatina nr. Sill
■ flovOTnuMsit uf, hv tbo iwrviMU *r»-
■ tnn. ^1
H n-lk-i ifiliiliilinn o(^ S&i
H <«nliac tyfli: oj, 2U7
H koimmI and iwnbniir, 31'>
K viMiige of form of, SOU
I
Utwrt, ilumiim nf tlir •c*«f>l phwo* at die
ranllftr t^yi-lp. '"H
viiiliHiiriliiu; (iroMire, 'Jt3
•ueibnita ot dclcrmining,
:!U
llr-lxmixl nf. 212
Bngiia of, ^-Vt
ulnltif^T nf. 'i'll
mnliui- miwuliU' liaoe, $11
nifijiniirv oT nutmmwIJMi, 34J
illl)iill>r [if, :!11
iiillumuw rqpilallnjc lient of, 3(1.1
lUKln tvtMt mnirring in vi-nlridf dtir-
inft * bMU, i2»
in0Uiv« (itvnutv. nsiiuvand nam at
nrrvo <)f. 2Kt
iionuul lirut iif. 'Jtxt
iinuli<i!> nf, 'i47
ik-vi'l'tgiiurill i>f, S45
iwyiiUliun uf kill of, -J-lfl
Mcwul (ounil nf, 'iVi
Mundoof. :zl:!
iniiiiniarv uf »vifiil» om^iiinilnK ■ (imt,
vnlvBi i>f, '207
vviilririiliir nvrtiili", S*W
vjxihli' iii'avTiK'oU of, l£l)6
wurk <l'iiii- liy "i'ii
IICUI. •t'IIMIli<-ll> of, S.M
lldiiihotii'* iirniiietii)i-"i for iH|ii(ilixIng ili«
make niiij liriak kliookis m|
|lI)llk<~>Hlii|iv. IHMI
llFiiiiiiml>lvi>|iiu.S'l-l
lli-uiinnriiiui. t-IH
llr'rikijiiii>|niji. K|3
llLiriii'piii. Sirl
Hfni.-'- .Ixnilli. ISS. IM
llik-riinit-m. ittl
lIi|>|)iiniiD|al Inbiilc. H4!l
Ilipgiiirii^ nriil. r'liniuiliiiii iif. 54S
llipt-'Iiiriiulin, IDT
lloni'lcr, |i:l-l
llaniui) bnun, noaiiimclatiuv of ill* (orflwD
of. H34
OVUSI, W47
Ilvnioid RiMiibrsno of rrr, HW
llvdniUllruhli). lUOH '
lly(«rii<i'a, 4S7
Ityimnlic (-liuaw, SSI
I ly I H i)[kHUil npi^'p, 7-ID
(••iiiw ■•f. 74U
llri>oiamliin. K/iO
IMAfiK fnrnialion of, KM
liii)wir<T-i -jiifwli. Km .
liupnTifiuiiiiin. twi
liniiiilM'. ni'miiit, 124
Iwlknn, IiKVj
Indifo, lajli
Inilai SSU. IIMd
IndiKtiAU will. 7T-71»
InlliUBinalion. iihcinjiiiiein of, SHI
InfmuUIialB c>f lll^|.■^ -133
tfniiiirc nt, iH
^^^IW^^^^^^^^^^^^rjCDBX^^^^^^^^^^^^^^H
InhiUtl-iii, fiwliiri*' »r, 2^1
].rrljiih-(ii^ Iwilrry. TA 1
••t iiy'if^ hcon In ^linulnliDB ut 'rvtctif
lA-Ud-cvlhirsiia, I'liarsTfrrUod t^ ilMnM
m-rvi-, :;.'H
of whil«> ci>rj fl
InliilHlcin' iivrvi'*, I'l
l.lfr. |>liaK*>('1. J
Iniijtvii. I-I'
l.itniniciiliim mil If-. 1. ■ ^^^|
IniMil. WXi
l.iiiili, iiHiitsiidil* cf, SSt ^^^M
lii»|>initi>'ii. 444
^^H
liiTi'rtiinliil miii'lcf- in nv)i!ral)iiR. 41T
hill! aijiilliirins i'tl ^^^M
Int<'rrilirillLir>iiUlftiiix>, I<'l
'illHIK -ViN ^^^1
Itiualliiiil iti((i^li(i|i, :|91
•iriii'liiri' ^^^1
Iiit(T>lIiic, cluxil- "C ''SMI
(ilUfllll' lIllMlIc, -JllO ^^H
Innfiv 3T4
IviBpliBlin I'r, '>7Z ^^H
kfiivkl fiMliinw of, 374
tirrvivi of. ■%H>'i ^^^1
niiiti'-ili mtriiibnini! of. 3Qfl
■iTiiLiiiTi- i>r. 'Ml ^^^M
miiJL 3lVi
(^•k'k ^^H
-IMvlul fraltirtw'if, :iiL*>
l.tiltnK Ih'Iv, ttiHli' of, 13 ^^^1
iilniL-liirv (if, 3<lil
LiijmI, iiilliiciMv "^ t:lV ^^H
IriisSMa
I^oIkv ft tiniin, 732 ^^H
lrT)ulin(I»ii,(IJT
l.'ibiitn' (if luiitE>, 433 ^^^1
Lunii, lOi^:! ^^M
I.nilinL-'H kTiiuifni|>li. ISO ^^H
].\<t)[l'S .in-iii1imti(\ S91
V Jniindicv. -'itl^
■liiiiriii^r. :MI ^^^1
Luiiirs 4:11 ^^H
JuiiT, ipintric 3<iT
^H
ull'in oil nrulcEil*, :tmi
■niUcinl. !»»
HU'ln-imii-. W ^^H
'■r. 4M' ^^H
rnnij'witioii of, 308
|wiii-miiii', Xi*
funcii{in cf, 4^1 ^^^1
infuiuiiiiiiiii nt. i:a ^^M
nviiiiii iiti fixxl-^iilli^ Xn'i
lyniphnlio- <>i, 437 ^^^|
DFrrt" of, 4;t« ^^B
pmHuiT <-xt'iuil in limiililng. tin ■
yATELWTUiCrUNl'S; 1»1
Mnirliin' ••{ mniiiDiiiliaii, 433 1
IjiwplMiic kUiuIis sinuiiiiv of, ¥A m
KMifj", ■■II
tTMmi, :u*i; ^^B
Ktrri'in. jivnuuiiU of, SI5
i[liinii-nili cif, .".17
ru*«K •inii.-iii[v '4. l&i ^^^M
Ljinplinil<>, KfiKin of, :ny!( ^^H
I4riii)ili.(ii|'il1»rii'>s 3»T
Mnll'ichr. {ivnimic) nf. alS
i'liHr»i-t<-i> cif, 41 W
MiU]'i|{hi<ui miMil«v 'MA
>'bi-lliii'Hl i"nill|"> i-J' '■ ■ ' '""
iii-rvi-- ..r, .'St
IkhUI^ ■Itiii-liiT' Us
riinicriirv i-f. >'i|l
iiiicnM'oiiit.- dui.
lulilil) iirintrvri, .M2
uiovcmMiu of, 411
viM.il]nr nr ran t;v limit* of. 'iSl
viiMi-itilnior ni-ri'<i of. o^i
vnwi-tiiri'-irii'tor iirrvo of. AM
UACIMCTII' iiiti-nu|<h!r, SI; \tt
viwiiivl'>r luvrhnniHniH of, fiilD
Kim—wlir. fl^:^
t
Krtntiii. Ml', tl)4V
i"-^ 1 .H
Kmiiiiiin. t<>-l'.i
WIIH1I.WW ^H
K} Iiic-Knil'li, IVO
({wmnlOMM, BW ^^H
l.ii(1wI|r'>S 11)0
iwUcka. 084
lunirs ■JtoiRinM. VH
v»> ilcd'n-ii-, IWt
I AfTEAIJ*. :Wfi
Li Lninliiu' in a liunlrntd laiu^ )*V:!
rmirilln K-niiiLrilt^v V^l
Mnli.iihl ! Mto
Idpdanin. t<KJS
r.5III
lutO inuvlinc, ^74
Ualln-r, :
iiKivcmtmb «( SKt
Hatiiniarv jilnnils ilA)
Larjrni. KIM
I'liiiiiliv- rxy-iirHnc !n «23
ruiilii)!**- i>f. W»
nlniiiiii ti' I •ton, ItC
nvilv i>f, U70
•tvmlliiii >"
miwdi* of. 070
nlrtii-lim'. i>'.'<> ^^^t
I*1)yi4i>l(i|j;liiil Bnniuni7 of, MS
Muioiiicur. nii'ii'iity, l^V ^^^^
Twnt oinU, 1170
vnioiK ly(^ lUi ^^H
Ijili-nl i*ric"), ^7
Uan-v'n inuiljoiir with iBnllor aMintL SIM
l.«<-illiin, \-i'\ 1143
•
"■"■" J
11 .*■
. J
IN'DEX,
1067
HRxiniiini manoiuentiT «f (ioltz anil Guule,
214
Maiwell'H niflhod of fiwing oilor mnm-
M«'hniii)-mH, dioptric, N!'4
iir uiuiniiiiiHlHliiin, >-'JS
locomotor, !li7
Medulla, IT2
McDibrann lynipnnl, Wi
propria, !7)) i
Memhrant nf KeiHHiitr, '.W-i
Menstniaiion, US"
Mercurial gnu-piimp, LikIwi^'h. ini ',
Metalailisni. ^nerul reiihircx of, tl-~i2
intiiience o( nerves on, KST
MetabolileM, nilro^noiiH, KMii I
Mflaniere, npural, 1110 i
Methipiiiogliilnn, 4(i4 j
Movnerl'd rommiwure, "40
Mii-roorgaiiismis atlioil of In iilinienlan'
canal. SM
Micturition, 54K
invoUmtarj-, 550
nervoiiw niechanlwrn of, -3411
vutunlnrv, '>M '
Middle ear, tHI)
Milk, human, ti23 '
cotupoiiiiion of, t!'24
constilucntH of, tiXi
quantity sccrelcil, Ii24 1
recretion of, (ii) i
miear, l(l'i4
Jlillon » reaReni, 4.1
Morse key, 77
ilotor area, tliaracteriHticn of, CtW
for left, arm, unci fact in ninn.
uviiion and relative extent iif,
Movenienln of heart, 20ti
of limbx, »-2V
-killed, WO
Mucin, ■M-2, I(K«)
Mncoun eliind, i'han)!:e» in during ^ec^etion,
34-J
Rlands, Si')
MuH'H- volitantei", 'JW
MuK'le and nerve, decree of irritabiliiv of,
1411
experiment" wirli, 7S
lilienomena of, 7)i
eart; 00
eliemislrv of, 104
(imlniclions, simple, H2
i-nrrent,", li;!
ctirve, M
double, !10
from tlic gantrociieniiii.'' of ibe
froe, f-i
ninBle indticlion ebork repented '
rn|.idlv, yi,ii:;
i-lowiy, 111
dead, 10> '
dnrinK coniriidion, clian^'^ lakintr .
Jlnee in, !M
-plate" of, yii
energy of, 147
Mnwle fibre*, elemonlary, 94
microscopic I'liangeis 102
-nerve preimralion, 7->, IHO
u* a mucbine, I'l-i
plasma, 10«
sarloritis of frop, U7
size and form of, 140
slnictiire of, Vi
>iibi.tnnn', striated, HI)
under ptitarized ligbt, 10-{
Mnscniar and nervous acliun, nature of, 146
irretobilily, 7;(
contraction, single, 74
simple, 74
fibre cells from human arteries 160
structure of, 9!l
irritability, 74
sense, 9*Hi
stibstanre, striated, It-i
lisj.»c, plain, lAI
iinstrialeil, 149
Myoglolmlin, 107
Myogriiph. pendulum, 8li
sprinjr, Mii
Myosin, 100, 1023
Slyosinogen. 107
NASAL ri>s«i, rifrht, '.(-14
foMP, 9.')3
plivKiulogii-al anutimiy of, tl.').')
Xerve and muscle, electric currents of, 114
cells, Kroupint; of, fiM2
of spinal i^ord. KiT
variations in, tWti
(.entri', I IS
cbemistry of. 12.'i
eighth or auditory, "-A
idevenlb or splnid :ic\'es?ory, 7.'kO
endings in striatetl muscular Kbrea, 123
tibre, niediillated. ILO
fibres in retina, conmi'tions of, 81H
fifth or triceminal, 7.>)
fourth or lr<H'hlcar. 7-V.I
ninth or glosso-pharvngual, 750
roots, cimneclions of, 1191
seventh or fiiiial, 7-")4
sixili or nlnhuens, 7o4
structure of, UK
tenth or vngus, 7.".0
tetanixuliim of, V27
Ihinl or oeubi-moior, 7-"i9
iwclflh or hypoglo!*al, 74S
tuh(.s, hunmn, I'M
Nerves, ilej^i^nenition of, 141
electric iiirrcnts in, 127
Servotis itiipulse, I'-IH
ebangen in nerve during passage
of, UK
meusuremenl of velocity of, 88
svsttni, ceniriil, Itlil
tissues, general fealureti of, 159
Xeurin, 12ij, 1044
Xeurcikeruliii, 122, 12(1
Nicol prism, 102
XilrogeuoHs melidsdisni, KVi
^^^^^^^OTW^^^^^^^^^^^^INDSX^^^^^^^^^^^^^^B
^^^m Nod» of Rniiricr, 119
Pnwiri' la MplIUri«s IVi ^^^M
^^^^^ Nn<-leiii,
Prmlirct* nf iliscMinn. (viiirHF l«k«<i M^^l
^)^H
^^^^^^^ ACTIPITAI. rvifioii, |ii«niinmc« of nu-
^^^H \J cinir wlU ill. Wi
In |>r«ivi>)< <::i
bjr mg^r. 4ti
^^^H ttculiir iinvini. tUS
\n tiairr luul mI*s 4!ff
^^^H <F<I('II1B.
PrvmlAIf eUiwI. II>4-1'
^^^H Ov«ipliiii;iii. ^'I<>
Pmuxon, Ii\ Km
ppotflil Ikh1)<-. chuMien «f ilw nxm k<
^^^^1 •niii-i'mi-nN nf, !IT1>
^^^H BIU*rli«
IKiriiuii. :iiW
^^^^P ocn'riu* «■ i|)lt- or. XSl
I'rutcids I'il"
dimllimlloii i<f. StS
^^H Olotii.
cnivri' ,■ ! !■ "■■
^^^H UlfiiKli>ry luiiiMiK iiit^ibniov. i^l* ot, M4
ikti"!!! ■ li* iJiLtt-Mion, IOS>
^^^^H viiMliiins
I^Tiur^l i.'i>.i""<ii'>n vf, 14
xan(biii>roI(Mo tcA fvr, 1>S
^^^^P On<yitrrji|>li.
^^^^H
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^^^^B Ojiiic ntijinlioii. Ml
^^^B ihnlaiTllK
I'in-rliiral [•Mcv*<i~i, liiimliuii nf, .174
^^^^K Urgonic luiiili-r, i-ilmtaiiim of, 4%l
lli'iiiaitin. X.H. ir,\ H
^^^^f Otiiiuiiior
Pllltl t%S HIlJlVIl, '•'*II ^^^H
^^^^ irf ri-|>riH)iii'iioii, Wl
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kntiKTMic, ZM ^^H
^^^^^H
iiii-[>i<h1- ••{ rx-ivinlliiK, 'JSO ^^^^
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^^^^B cliiuiii-i ill. iliiriiia «wnlt"n. :((<)
Itvi-p^ 3111
^^^H i.r <!•>£. >n-ilon of, :i:;i)
i^ll•■■-l^lU'inx rnxri no urlllk'ul ihinIcL 91
^^^H iir rnU>iT,
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^^^B ■•Iniriiin-
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^^^^^^^^ PUnrrvuiio jiiirv, Il'>4
PilW-nnvc, ailiiiT>»lli', V*li*F> oC "iW
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I'll nihri'i of, olnog lh« BTMtW (IK
^^^^^^^g nil r(>iHl-*IutK>. ^Vi
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^^^^B HicrFiiiis tif, 'iy.l
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^^^H Pa|illlir. rimiinvnllRbe', V^^
fcjiiiirri <.r, 39u ^^^1
^^^^H fillucifi)riii.
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^^^1 Panutloliiilin. IIIJI
|.nHliopilir, ^9 ^^H
^^^^1 In iht' rlollinjc <if IiI'khI. W
v*l'»-ii>- iif. 'JXt ^^H
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Pnpll, mi.vv.T.cnu of, VOI ^^^H
^^^^H l'Bni|H-f>l'>iii>.
nvM'i-i ri.iFniinii. 110:2 ^^^H
^^^^H Ptmiiiil (Utiil. ni-miit* nm*llltiu<in of, !t3>
I^ukini". «<|]> ..f. MM ^^B
Piiiklni-'- riuun-*. IM'», Oin ^^H
^^^H
^^^^H IViiiliiliim inyivrapli. H.^
Pvlnriv cUiii)-, ;l£t ^^B
^^^^1 [V|>i--^iii>ii>i fiHHl, 3i0
rvniiii'liil iilU >"3 ^
^^B IV-flnn.'. 1(i:U
pVlVtin, 'M'l
^^H IVpiin. 1
I'vrifiiriii )caii;;l).uiknrrT«4«1l, atnirtulB
^^^^1 pFrMn>'!i'n\ |<^ivn1 i*n>i1illltr and tartlle.
\nn
^^^B
1(.W. sil»
^^^B m-Klitifil.
^^^H Ppri^Mldr lU'ivi^iciils 3iH
^^^^1 PtTiTwi-uUr IvDiphatki^ ifK4
AUAIlHUiEMISAL Iwdi<^ 772
\J (Juaiililv lit •(|iii<ou* ra]H>r rdufa
4oO
of hlnod, 71
mudo of aMlnalliui. 71
^^^^B Pi>r"{>jm(t'<ri,
^^^^P pFllfnhnfrr'i tril ^r We alU, VrS
^^^^B rpvcr'' iinii'lwr^ lOI
^^^^1 Pliv«H|(|[iniiii-, BrlloD ci 'fO pipll, WU
^^^H Piliiilan- txicty. ttlft
^^^^1 I'liumiap, iu rlntling of Mooit, 49
of inrbotiir nriil cil.al»L 4G<>
<if iircanlc nuilarrefiliBlol, 451
Qnnlliv of nanJ, 9tH ^^
^^^^1 Foil- Vnti'lii.
^^^H
^^^^B Kniv matin' of.
nAXVIKR-Snoa*. IIU ^^B
ll IU|<iililT of nrcaUlioa, ISW ■
^^^^B I^ulrri«^ mliaiui of ■inaal roni, 677
^^^H Pmuirvs srtiiriii], I3e
.,_. J
r
-■
IN'DEX.
1069
Bapidily of rimilation in veinH, 203
of i'lotting of hlood, 431
Readion of urine, H'lS
Bectiim, nrurture of, 3T5
Bed-blind, ^23
blood-corpiiseles, 54
counting of, 08
a|>])ornlii« for tlie, -18
diameler of, o4
ili8inlcgralicin of, 50
number cjf, 57
slmpe of, 55
clnu'liire of, 51
iiludv of, 54
ninrrow in bonv, u Kuurce of red cor-
pviM'les, 00
Bi^llex uciionii, KiK
nalurc of, I(i8
of i^iiinn! mrd, ill
inliiliition, 2>5T
fiefnirtion, MI4
KcGiilulion of heal, <J-I4
BciN-nvr, niinilinine of, 8-W
Rpniok'N ^iiii^lion, '2H
Kennin, ;il(>
Keprodiii'iiiin, VUl
coqius liileuni, 1*88
of nienwtmalion, iliHti
of pregnancy, WHS
develiipmeni of embryo, Wl
allanloip, '.Hit
pliiK'nia, W'>
ir^mtnlaliiin of ovnni in,
vari'iilar aren, Wi
(imnlian follii'lf in, ;if>T
imi'tefiiialion of oiide, VM
mtnpfriiHliiiii, it!-"
iirpnns of, !>2
Fallopiuii xui-eK. iit;i
mall', '.'Mi
ovuricK, 'Jt:i
vagina, 1182
ovnie in, !ll 0
ovum in, UW~
siicrDiatozoa in, ^1(1
Hi'Kpitution, 431
abforptiiin iifosvgcn in, 4(iT
apmia in, 410
iipporHliL!! for tuliinK ttacing>i, 442
an|)liysia in, -!41, 4'.''2
plicnnnu'na of, 4'.r3
carbonic acid eihaled, nmouni of, 450
chanKcn of air in. Hi)
Cheyni-f'^lokc^, 4{U
complioitniul air in, 43!)
dyspnii'a in, 441, 487
cHccl of lirtalhinK foreign gate*, 484
of chanfiKi m atniOKpheric pres-
sure, 41)5
of mutculsr ejerci>i' on, 489
elhalalion of aiiiieotm vapor, 451)
of csrlHinic acid, 4>JU, 470
of orgunic maltens 451
expiraiion, 448
! Bet^piraiion, expired air, impurities of, 451
' nuliire of, 450
tcmpemmre of, 449
' facial and laryngeul, 44!)
pnphic rcfordw of niovementu of, 441
influence of vagu.i nerveK on, 478
inspiration, 444
labored inepiralion, 447
niodilieil movemcnlB of, 509
I cougliing, 509
cryinfT, 510
hiwimph, ij09
I laughing, 510
I !<igliinK, 50l>
I >tibl>ing, S09
i-nteiing, 509
1 yawning, .'lOy
movement" "f diapbragm, 44o
of expiration, 448
of inFjiiraiion, 444
milnle^ of, 475
nervous nii-chanipm of, 475
nnniber of, 441
relation" of riTpiralory Kysltm (o Tji;-
cular and oilier systems, 4110
residtial air in, 439
rills in. action of. 4411
I funi-lion iif. No
Miitionary air in, 4^18
lidiil nir in, 43.S
vipible movtniinis in, 444
Ee^pirlllclly lenlre. 47(i
cban>;e" in bicr d, 4*il
in tlic lip^iies, 471
nndnlalii ns ^■<'^
Bete mu^o^lnu, ■"j-i^t
Keiiciiliim of fpltiii, 5U1
Betina, ^i.0
connective lipfue of, t-'Jl
inner Mirl'iu-c of, 8(^2
niugnitled verlical nclir.n of, >■'. "J
of man, icd and lone fri ui, ^91
pbdlo-ilicmislry of, 1)11
Bibs, action of, in retpiralion, 44G
liinctirn [ f, in rc^piraticn, 443
Bigor mortis, 1tl4
Bitler-Valli law, 141
Bods of (oni, '.143
Kolamlo, piibj-lantia gelatinofa of, fi> 1
ml ctilc of, 74")
CAI EAL nerves of dog, lfi2
l" Valine malterf in blood, 69
f-alivB, ;Hi2
action of, on starch, 303
cbaracIerN of mind, 306
of parotid, 307
of publingual, 307
of tnbmaiillnr}', 307
chemical eharactertt of, 302
composition of, 302
nature uf amyloliiic action of, 305
secretion of, by lueanu of chorda tim-
pani neri'e, 333
Salivary g I amis, 324
^^^^^^lOTf^^^^^^^^^^^tNBEX^^^^^^^^^^^^H
^^^^^^^B }|Sslivirj' flsiiil's Bi-iicnl >)nii-lurp of, SH
SiiwU. H-nHiiiAiw of. M7 ^^1
^^^^^^1 n<<tTiiii4 •i]|i|itr of, Si^
S'.n|». I»I0 ^H
Siiliiiiry follii-liw. luS ■
^ Sall>> t>r
^^^V }<ii»i>>nni. i-flniln;^-of, M(l
.Sinuiii- ftnil i>|<Uiicliolc B«rv«i^ IHI I
■ Sfirlin, 1ii:.<i
Si'inint'rrins, vc41uw mmi of, (M*.' 1
SnxlnnTiitmiil. iM.1
SoiijiiL '-'(H ■
rrrtilmli, 'M:*
[au-h »r. Vf^ ^^^
Srliviiicr'* t'\|ii*riDii.'i)l. iliajcnuii iiC ^^^
miu1il> or. V4'( ^H
RotlDili ■>f ili« linin, 313 ^H
Siililvtnm, fJiijil i-r. S-"*
i<iiintic oprtr nf iiiiui, 1 lU
S|M>ci£r );niiilT irf nrims A34
nl' nililiii, iM-rrip-ilhfv froin, !!!■■
S]>wtniHii|>ii' iuiii1v>i> ■■( hirllxiicliJilK 4)
Stk'rolif C'Bt, WW
Si^-wli. v:4
HttbiiccDiL-* kIiuuIvi ""(l
Btvn> tor, KSH
Kn-rrti'm, i;i-n'Tnl nniun-nf, !H-S
IliitHTfnI. 'Tili
ii.iKirc <■{ iiii- ni-i "f. ;Mr
(■BUiol W Iwllwr Jij^nn. R37
!4cttim-ii:iiiiim 'if miiiu, OTI
iiiick. ss:
S^iiiviixniinr cniinla '>13
S|H.-riiiii|iiUxi. WHi
Si-iiiitinl llilM. KH"
S]>)ii-riml nU>rnit(»n. VOU ^^
Sa-iiiinirrnoH riiliulMv BW
Hpli.Vi,-ni<>((nipli. t>iiil|;<raitlV 227 ^^H
Svii4iT!'>n, W-*
ilnn-y'-i. :r;7 ^^H
Scti'Aiii'ii-. iiii.liturv. W^
8i>iiuil iMvvmirt' nerve, iM ^^^M
■■iitiini'viii', "I'tl
n.nl 171 ^H
ili-linrllriil nl)<l fiiMiKi iif. I<IS
unu-r>>-Ul(Tal iMvnJimc M^H
of iiilil. S.M
:uu ■
.if .'(.I'.r. HI"
ii>iH-ni)in)t inwt> nf. AMI 1
priiiinrr. yaa
mill 'Ilia ill- itriloiH of, 734 I
oT lifiiriiiii. H.'ifl
cciilnl mnni i>f, ltT7, iMll M
.)( Ii.-i.t. J*".l
n-rvU'lliir Iran, XlK ^H
irf |ip»«lni, HftI
C"liimn> tii RiiHwIi, IHUI ^^H
llf VUK'II. "i'
of 1 Inrk. f.V. ^
nf in-'lt-. "(V)
llf (iflll, IWI
of KrlllllfniUKV, >*->l. VSi
niRinlcihviif rrflpx mnvvMnail
7 il-7»l
rvlnlinii III, t» «tiiniilu.\ fllS
ninm nf. (t7T
■I<-ci-n>liiiK inrO u^ (US
fi-iDiiiv llf. I'-iM
viMlnl. 93*1. IK)*
liMiins of. ITA
S!U*r. Tiiiii'iilsr. '.'■'•li
ici-Uiiit'iii-- MitHtMicp nf Btthd
Scnuorr imgiiil-vs WH
(iNl
p<iTrm" iM vilify -I'W
(rrav ii]»lirr. nxiiif nl. TWIi TifV
fliiltl. rlii-niliiil Hianu1«r nf, 410
-trui-uirp <)f. HTfe
Srniiii-ntbiiiiiiii hi rlnltini; of I4(kmI, 411
loiiKilmlinuU, cnnBi'iaMiM) tnrtti
ripruiii int'liHlinjt "f liI'miL ■Wl
70!i
tl<!i*i-iilli jivnv. 7'i4
|i>n< iif tfinp laf filidMal mavtn 1
Scic^ ilif1i-n.-iit^i? ill mjilmtiim, -144
7*-,
SllhV jX
n>Ilpi artinnxif, 711
iSiKth orn'Pi T-^l
inbiUllon i>r. T:tl
Sknii>l. I'>«l
lisir nvpiimJ frir, 731
Pkin. .Vil
n-liUU* tbv ami fonn r/, lUM
sl>u>(|>iii>ii liv, MI
rpik'iilNr fnrnitiinn, iVU
cBl>iif<ut rv-plraliMi, iVK)
liXiilllT nf mii-L-lo* tlirMuA tak
di-rnii* '>r triiv, 'i'>1
.>r :"■;•»
HildpniiiK 'VM
■ 1. i[7't
li«l»>. -v.:
ixiuiifoa oC 714
fnllii-lu' nf. •'liiT
«.! 1- '1 ,[!■ r,- of, ICT
tM'r'fiimunn, ninoanl uf, M9
rr ■
.>.TT><->iiiioii trf, .IW
H|ijn*1 'li- '-■ . 'I'liiiu. .,.. .,, ,-.itn HI, WH)
r- Hi of. *"fc*S
f.ni:.'!: ,, ltl.'t
h'l . Dimli^ Wl
[-.■■T.I . li'l
mailvn or, -"■'i"
SplcMi, OSH ^B
Hv«vlioa of ntMi, iDi-chiiniiini of, S62
l.l<Ki>lvc>i>U o^ fiUI ^B
rbcinlral i«ii<iiinrabi nC, 8M ^^B
•IRIi-liirr nf, ^>l
»<>t4i-i[lnnil^ 'AS
lrni|>hBtici •■{. 'I'A 1
..f. IWI
^[nl|>i).'llii<n mrpnK'Ica of, &PS 1
mitri'iiiiiiil- nf, ■'IlKI 1
1
^^^^^^ iifenf. **3
^^^^^^"
J
INDEX.
1071
Spleen jHilp, ~>'yl
nHloiqHwIe^ in, 0(1
rvliiiitiim i>r, ')\fl
-iriH'liire uf, -WK
ISpriiis n in Hum Her, Pick's, 22li
(*la|Hiiiii' [iiii»<.le, 'J4^
Stearin. IO;l!l
Sii'llale, iiiTve wUn Ili-'i
Slt'rni>-tiia-^li>iil in ninn, »ti-linn fmoi, !>->
Stiiiinli, I'liiinivUTs of, li(!j
Klimiilii'i. luilnn- iind iiinile nf n)i]iliniliiin
of, IMi
Siomai-li, :!i;'
iintnim [ivlori, .t^ll
puniiiif irl anils of, 'fM
jtnslriv jiiiiv in, ;W"
iiiDvcnitiilji of, ;tMO
nervdii'* siippiv of. :tH
]>_vliirii' Kliinilf, ^2^
-tMiflurc iif, :il'J
(t-mrjil relU in, H^'J
[•iirieliil i>r civdiiI cvIIh in, lt'£\
Slnicinn- nf alvccili uf liinps 43(i
of iintries, 17>>
of IjInddiT, 'i4li
I'f Imiiii, Tit
of bronchioles, liUl
of {.-upilluries, 17l>
of eilemul ear, "'fl'
of froit's innir, 4^1:!
of irrav ninller, (171*
of internal ear, 'M\
of kidnevs, .'ill
of liiryiii, SHW
of liver, 'ili'i
of niidille ear, 'J4D
nf nerve, HS
of newt's hiiiK, 4ti2
of spinal corn, (iTo
of spli«ii, oSS
of s(o»ia(4i, ;lll'
of tlivnnir^ tilo
of inicheii, 4W
of ureter, ■'>4ii
nf iilcriis. Its:;
of veins l"!!
of villi, :!7()
of white IiIoimI t'orpn^^'li's, <il
of white iii;il[er. ii77
Sniininvinurv (flund, -!'!:<
iHTvi'?. of "t:{2
Snivn- eiiti-rii'ii". itW
ii;i(nre ;inil aWion of, Ilo-I
SiiHori^ia rolls ylund'^ h%V)
Sn]ira-rrn;il ii'Klii-^ (il 1
i'ht-iui™l consiiiiiuiitH of, '>!:!
fiLnrtiims of, fii'2
-triiitiire of, til 1
Sweat -((I a ml ■^. '•■>'i
Sylvius aijiuiiiivt of, ~'.i'Z
Svntonin, Ulil
rA(TII.K ]MTiv|itions and judgments, 'Mi
Taiiiljour, Marey's, :ilti
I Taste, aw
drcum vallate imiiillte, fl'i"
tiliforiu pnpilhi', iliiT
fungiform pauitipe, IVi"
giulutory biilLs, itiiH
Kensntions of, M50
Tauriii, 10.>I
■ Teniiieradire, 043
eflect« of great cold on, O.jI
of jireal hem on, I'lM)
influence of fooil on. IS4l)
oil contrnetile tissues, 142
of miiscnlur action on, 046 .
o( time of day on, (j49
of IhhIv, «43
, of varioiis animals, U43
pyrexia, 11411
I sensationH of, Hoi
, Tensor tvtupnni nmsi-le of ear, 947
Tenth ner\-e, T-W
, Tertieles, SIS4
1 Tewta for bile, :«:?
I Tetanic coniraclions, 74, tH)
I Tetanus, iH)
I conlnu-tions in, liW
I Thalanii oiitioi, 770
. Thermal cliangeH, 111
iThemioiiile, 111
ThinI nerve, T")!*
I Thomeir diu'l, ;W7
' Thyniiis, fiinclions of, 61,1
atriicturf of, til.1
. Thyroid body, (i07
functions of, (fO!>
! slnK'lllrc of, (JOT
I Tidal nir, 4:jH
Tisflies, conlnictile, 72
■ Tongue, 9511
Tonic eon tract ion, lo4,H17
Trachea, ili-'i
stnLCture of, iH-'i ■
I Tracing from heart of cat, 210
, of resjiiralion, 443
' Tmnfiidation, phenomena of, 414
, Tniniie-l lering variations, HS4
Trii-emlnid nerve, 7"«
' Trochlear nerve, 7")!'
Tryiwin, 34li
Try]isinOf;cn, ^(47
Tubules, nriniferoiis, oli
Tunica albngiuea, 1*S4
exiiuiii, 17'.l
intirim, 171>
nitiiia, 17V
iTnni.'in, ll):ti;
Twelfth nerve, 74'J
Tvrifsin, lOO'l
UMBII.ICAl, vesicle, HO:;
I'rari, [Kiisoning by, 73
riw.' of IjIoikI pri-J-^uri' in animals
I un<ler, Soli
j L'rca, IW--.
I formation of, in liver, (HI2
I nitrate of, 11J4.>
1072
INDEX.
rvlsiiim* lo cytiniiecn vaaD|ia«iiMU COQ
•ynllMTii-i of, Dftl
L'rUM, Mrm-liini of, -Vlli
Vric •I'M. oiA, (lO.-., 1(H7
■nllK of, 1017
rriiwv W4
NbuiuiQul «inilitti(^iitn of, 'I'JS
iilbumiii, hri^
acidity iif. -'iSti
■iiiiiLial t'i, W£:
I1>III|UU4ll<i|l l>f, .'j28
fcnin'nlp ill. Ji'JT
at-iK-i'iil i'liiniru-r> of, 334
Til)'|>iiri(' mill in, hHi
iii'Tinmii' -"ll* in, "lii
ii.pn-iiiiro(!i'iioiw ii'tuiituiiiU ftf, 8S9
iK'Tiiinl iTinutnii' iiiiuiiliiirat* t>t, hit
liiBi'icnK ■•J", Urt*!
rciiciii^ii of, *>'2A
rclnlitiim of HH-tvl1oa of, w Ibntl mhI
itrink. -VU
•Crrvlioii nf. •li'!'
xlcimFmlnr, M8
•jifcilii' itmvity al, tt'iA
urea ill, iVJ-l
nrii; oriil in, A*J->
uTiiniim u(^ in, 321V
rriiiift-ruiK iiihiili^v 'iM
rnibiliTi. Il).'>'*
^^•<'^' ill rill, XfXiii
Uterus WS3
o* uteri, V82
ftniniirr ..f, W2
tllrUlM .rf Mir. tfVi
\rA(iIN,\. mi
Vof^up ni-Tvi". 751)
Vii1luiuiiin'> hn'niiiitiviiiMiwti'r, 200
V'illr«> of ht-iirl. )»:;
VnriotH forniK of Kiimuli, 75
VoHiJar nminKcuii-'nl i>f tfai; kiilix^, ti'll
IT1lvhHnT-M^ ITIt
Vac difi'irii", K'l
Vo»o-v€iU!"irioi.ii librw, ruorw irf. 27:1
arrviw, I 111
-«lilali>r lilirvr^ ii'iinv of, £7-1
■Rtiitxr iiiliiiiw, 'I'Vt
etiitti. oT, 27 A
VrioB, IM
IiIikhI j.rtwnro In. IS6
orviiUli'Mi in. Tn|>idily*<>f, JOS
miin f>r, I si
oln>ii('ii,T of, 184
tirurtui* I'f. ISl
TRMimoliir ncrrm oJ, 2ft6
wIDi valvMcUwd, ir'2
opening
Viil«ciQr uf blood m uterins IW
ill (^pillarin, SOS
«f ill* jiulw VBve, 333
Vmoiw {>iib^2S0
riiMiM^ 883
Vcwnil» f caiinaJai, l'S>
Vllinlinff liminit-furk with lhv|itTt> i
»7
ViiTordl hniMUftianwfer, MM
Villi, chanrtot* «f, 3l»
rtruaiur of, 370
ViMiii, 908
biniHulM-.tiSO
rtt;>i>n of ili-iiiii.*i. VM
VwudM>l,».t7
Initiulim, origin u(, IHIU
vt llillMMV. DM
iif •!»?. SOS
i.l^r-.lHlii>.90n
(nini-iiK-iii^ •irf'tilinoiiiin i4, VSt
iwiri-iiii-iii-. '■^''
(imlnv.-! 'if. tffi
)>ar|>tc, I'l't
•ramliniw, kAi. WM
irliili%li|:i
v,.|t..n^, ttl.!
Vhplliii. 1(121
ViinviiiH tiwly, fVS
Vocal itinli, U7U
lll('VI-ln<*lll> »f. V'S'i
Vnlilioiiut lniii<il->-> in ilie ninl, iCS
Voliinimry mot' |< ' tin. «ii
111 I II ilip lontr
I I III
Vwnilinv, U\
WATEltY vniH.r, nlinlallun •< «lt
M'nic, i-rinlriHlion. US
WblK blood-corviiH'In. '>!
rlM^nlcal i-SBRiiniittH
iiii^niiliwi (if. M, '.SM
■imaiimu nf, 114
niiLlri i>f, frl
Diuiilrr "f, fit
iriri;e>>i "', <)>i
|irui»>rtu>n of, lo ml,
fi(r of, 01
tffiiFliirv of, (II
iniiiif»niidiiiia itttv
«7
Wrfk <if, G0
Whitv uiniiiT. I£>. lOU
Wtlli^ i-in-Ic of. KM
n'nrlc diinv hy Uvn, SSa
XAXTIIIX. HCO
>kaiiilir(ii>n>icic tM for iiin>uUv<
YOrNO-llKLMHO!.T2 iheorr oT
ZISX, looe of, hS8
:!faia iK-Jliiada, WO*
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blc ■ mluciianin ibe pricvof Thk ^Kin bi FofU Dollars per year, n that it it kw ,
bj br iWcbtiapcM M wfll u llio bnl Inrfo w*«kl7 Journal pnUkbed in AmtHca,
By in«BiM of The MzdicaI. fi'Eird rv«qr |ib]raicUn b now abla al s nilniaititn i
lAliwurabisoirnrK«ipt<irtbeekrlMm ami mart oatborilatlve iafomaiion on all mi
bBA BkoTHBM <
THE MEDICAL NEWS— Continued.
of ifll«r«tt to lli« gnal nmliiBl worlil. Tli* (crainat writvn, Wadwr* aoi finctlL
e( Ibe dajr fiirnUh nrigionl utidc*, otiniciit l«ctiir«i Nail tmUst tia pnctlal ail*
tbs UUM mdJioita In liadin); bmplinl* art i>tamptlj r«]i«n«tl ; • onodMiMd Mn»W74
pngm H glnancd mcli vMk ham m lox^c eii liaii)[* llM, CDn]|«iB!ng Ibt b«
at boms luiil nbrntil; a *pcdal dcparttncnt ii aisigDeil lo ilalrai-U rcqujrii^ Ml tra^
tDUDI for projwr jirMiinUtiMi ; adllorUI article* an Mnind from trHtan abU la
iiwtnictiTel/ wiib<)ucBllo(uar Ili«ila7;lwi>^*r«e»rvru11/ rerivwHl ; momj pmomllip
•r« t«pftMi)t«il li^r tli« |>llli >lon«; rcfiilu cnrrctpaodenc* i» fumuilwd b;r gsndaiim Is
poaitiun to know nil otviirrmcea at itnpiittanM la ll>« dlHrlrl nirrrnitulla]! Iinpwtul
medical ccalm, nod minor mallere of tntercel an groupvd lacli week iiDd«r nen IMk
ETorrllilng b yrMtatod vitli sitcli bn^vltj na » compatibU with clcannw, ami ia lU
mnst aiinclit e BaBnrr. In a word Tub Msucal Sm b a criip, fnah, wmIcI; m«»
pajicr and na mch Mtnpin a wgll-niailud spli««« of tuafiilMM, dbtiact uul ci]iu|>»
uMiuirj (o tliQ hImI montlily nwrKuiiic, TuK Ahxhicax JovrhaL or tmx Mkdku
tJramci-nt.
The American Joui^nal of tijB Medical ^ciencej
Published Monthly, at Four Dollars Per Atmwm
&lenuiioaila>eventrHMKocid7faT(li491)wilhaMiiraiKworincT«aatdiiMAiln«k. Eacw^
■gxl by Uia gpiplrtllc tPdoneiawt otth< prefewlon, m inJicrttd by a growtli in iUwbMrl^
Um t lot of fiAj p«T cent. usM iu appeanuioa M • Doonlhl J at a reduced pricp, UiiiM in diatgi
will tpare no nffi>rt to Riaintaia iu place n* ilia l««dv of latdlcal pwlodLcttl UlanMK
Bang th* medium chuwu hj (tt« twal mindi of ll>« proEmion diirioK the pMlwtMf
jMn for the pracnlalion o^ their ablM pa|icni, Tiik Ami3ii'<ik Jocbxai. baa mII
Mnwd tht inaiM aoeotdtd it hj nii uoquoitjaaed atitlmrii;— " rnun ibb □!« alose, ««*
ftU otlior puUimtinn* of lh« prcn> for ihelAnt Gft^fcan d«slniT*d, it wooU bapnMbbM
npaodDoo Ui« )[ri«l maji^ilxr n( ihv ml contriliutioiu uf (he world to medial toMM*
during that pvriod." Original Articlei^ Reriew* and Pn>grcn, the tbrea nain dcpanmnai
ialo which thaconUnUorTiiK JovnNAi. arvdiTidod, will b« fotinliopMHWMUt Ki«*ttr
ial«rwt lluii III t)i« ful. Tlii> liri^htaM talent on both lidM of 111* Atllatio b cnliilid b
ila behalf ami luiclliiri will bo*pai«d lo moke The Joukxaldiocc than ev«r wvcilij d
ba poidiioD a* the rvi-tv*vnUilva of tlio bl|[liwt fcrm of mmlical ihooghi.
i
COMMUTATION RATE.
Takoi to^nber. Tue Julkml 4n<l New* form a pcculiarlj uefiil mnUa
■lul affiml UiarrcaJen itic nminuice Uiat nnitiioff cif ralue In Uvn praffrcM of m
DatierathBllfacaMMtmiioii. To lead evee; reader 10 pcoee tbb peraoaally tba '
muiaUoo rale faoi been placed at the viceediogljr low flgare of 9T,60.
SPECIAL OFFERS.
The MsniCAi. Niw» VouriyH Ltn invent piic«, 11,30. *e« ooxt r^c^} or!
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paf ing (obKriben to ciilivr ur boifa of lh»o perimGoab fur 79 ceou apices ; or JoanaL
Hn^ VUUOK Urt and Ycat^ Book, 96JA.
SabmHtrt mm iMaai, tU lAc dip ^ «anl mtmM, eM& (unrr* /ar Thk Joitwiai. i »
OManHy), andljvr TiiK Skit* (eaa cmiaBf),/rt4 Ay noi/. Ay rMtJftn^ TVs Otmit lor Tai
l9*T1ie MlHt mode of readtuim b bj bank ohedt or poMal maiuij order, dnwa »
tfac'jntcrofthainiilenigoed; when thsevamaotaooeeriUe, ramitUDOcaror wteeripuai
maj- t«Miiit ni ihe riik cif th« piili)UIi«>ri \>j f'innirding in rr^iMtral \eitm. AdJrna.
LEA BROTHERS & CO.. 706 & 708 Saasom Slnet Philadelphia.
THE XSDICAL ySWS riftlTING LIST FOR J899
Im ^ubli>lie<l it) four nrlcn, Wtvkly ^liiti'd fur 30 palicuu] ; Miiutbly (uwlnlcd, for 190
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fvtiial (uaibMd, fur 60 tulirnt* wmkl^ )ier rear). The OC^wtient )'cr(>r[u&l cotuirta
du* uiil 17(1 pan* of tyMrt*! lilankt. Rm4i MvU U id «n* iirali*t«hiin«(l bock, lexher-
boiiDil, wUh puck(!(, (WDcll, niMivr, eosiMr litl>lct ami nlhMcf-Kale. Price, ckcb, t1^.
SPECIAL COMBINATIOHS WITH VISITING UST.
Thk Amkk. Joiriiv.ti. Iff/ niih Vl-jtisu Li>T(t1.2&X"'VEAH-Booic (f l.SO), fot ^TS
TlIK MkWC*!. NkUtOU - .1 .. n .. 4^75
The JoiRXAL Asi> Nrw* iJTSOi " " " SjSG
The JdiKKAL, N»:«>. Vi.-itjs.. Lt-t anJ Yluit-BiXiX (•« (Mpi 17), , 8g60
TIili lUi ■• ill ihu eouM Im iIhIiviI. Ii eon- 1 tM of dliMWH wraond *J|itiiLbet1i»ll7. (Irloi
Hints (■•(Wnoiliil at uwful InOlItnUIOD, Mact i UDdsr Mrnh ■ lUI ot Ul* pnuniaaDI drUK* •(•-
ally lar Mn^rffvnu and KlvM food Mblacf ■(•)••■ . I|l<>r*'l ta ili* inatmwit. whtn ordarad,* ll»dr
MM ih*r«fi*iitl«.— OnuWiai jV*(«Hmv. I i(ffrr*D<wThucnUI*ii*rlfl-if> itfnralf*«L Tiila
ll It • muur^lcet. Bmae- of Ilia ft«uu«a ua i it a baWra iiaoullu la Uil* Vblltas lJH.-^p»tm
paolttf M "Ttif Madlctl Sntt VtaKInK LUI." nan wu ffanon, Daoambar.
ooMbV <'>■ Than[>tMil'^ Ttli1<-. i>r<]«rad trnm Pr. | For ooara Amv* and ala^ona II It not •iirrwa
T. IiMidar Kpnnlon'* bonk, ohlr b poolalna tlia aWa— OMMMt OawHf, Nnvtmbtr.
ry/ii; itEi>icAL utews phyhicianh' mixiBR.
OODtabiiae 300 page* of floe liaen " lei^«r " Mp«r, ruled ao thiU all the accoiinta of t,
lufa piutic* uajr b« cQDV«ni*oll7 kepi in it, rillirt hf ungtc or <li>4l>lc entry, &>r a long
pfrioi Slrmglr bonnil in Icatliw, iril)i i-Iulh <itlM, iimI villi « |uil«iil lleiil'U liaefc,
wliick p«nniU it to lie perfectly IIbI wheu upciiol ai any i>U(«. Price, f4jXi.
BASTSHOBJfE, UENRT, A. M., M. D., LL. D^
tMt^ PnftttBr of ayfimt tn iw IMnrtUf of i^uu^mnt^
A Conspeotua of the UedJoal Soienoes : Cnntaialmt Haodboeltaod A:
PhjtMnfr, ( bc<iiiiiiry, Miilvria Minlita. I'rM^icr 'if Mnlicine, Hur^et? ud 0)»t«lri».
BmdihI editioo. llioriniulily rcTJatU and Kidly imurotod. Id oao lai|a mjti l^o.
nluadof lli2» pa)t«,wiili 477 illuMiBiioiu. Ctotli, HS&; laatbar, fthOO.
l^a ol^aet ci l^lt i^uiutJ It 1^ arTorJ a coj>iva-
taa< varlt ot nftraDoa tu itudanu auri^K >>'•> >"t'f
momaaiaal iliair commaod «hk|p ib tlcvmitno*
open Badlaal lannraa. 1( it ■ faioratila ilea lliit
tt >M %■«• (MOil nanaMtrr. lo • thort apaoa <4
IndatUT aad aaticr at lit able adllor.— AMloa
Jtahfol and anuafVinn< SanL 1, IK«.
W« tan mj with Iha Mrt(Ui>< irtidi IbaJ II It Iha
Dati wetk at Uia klB4 oMIi *tHA «• ar* as-
. . --...-- - - -- qualntad. tl ambudlM la a eoodantad torn all
UMa, lo Itma a naw aad carallilly r«ibad vdiUoD. racant eoatrlbuUont la anttlctJ fnnlloiat. and I*
Tka illnaUBltoDt an nryDUmtmit aad saUM-l lh*r*A>T«aiafUtlDa»t7liiin|inwminnarllirouCli-
■lijr oImc, aad tach pan taamt to bar* raaal*«4 ', oat our oanony. baddat twlnjt a4inua)4)' ailtfTai
HadaaabMaot aiMDiloa. Wa oao eoaotlt* taatt ■ to tha oaa of atuiltnu at ne-Dela*. TiM bOc< la
a *of% (a ka utatal, doi anly lo ttodanM, bnl U ■ UUitullr aad ablr aiooiUad^-CtarlMiM ITidiatf
pfw uuoaan aa voir. It raRtot* oradll oibd Iha I JWiiol, April. IM,
ITEILL, JOHN, .V. D., atul 8XITH, F. O., M. I>.,
Uitt Sr^tm la l*t frttui. HitrptlnL Prof, i^ lAi t-<ttil%lM tf JfsL Ih Ud Vmw. af nnaa.
Ad An&ljrtioftl Conit)eocbam of the Various Bronohea of Uedlo*!
BolOnce, fiw lh« useandeiamiimtiunaf SlodMU. A new tdiliua, reriaed aod iinpri>Tal,
InoMlargeniyiJianiaToltiBiaof g74pf B^»iUia74 wnodooti. Clr4li, $4; Itathar, H-^S.
I
I
LUDLOW, J, L., M. D.,
OtjulUtij r^u>u»r> It, U( P*iftidtipMa B>tp>Ui, dc.
A Manual of Bxamiiiatioos opoa KvMimaj, Plijikilciar, t$arfl«ry, Pnctkw et
Hcdldnc, Obn«iri<i^ Materia Mtdlr*. Cb«a[ilMi7,PliunM(7Uid'ni»n|i«ull(a. To wliicb
ii added a Medli»J Formulary. Tlli^lellilioD,tllonwfhlJnTiaa(LBn■)K^eallTeaUr)(cd. In
OHO ISmo. vnlutneof SIS pago, with S70 illuMMttaUL Ckdli, |3l2A ; lentlicr, f3.T6.
T)i« arranecincnt of thit Toluue IB Ui« form nt quouunt aad umier render* il <wp»-
dally tiiltatila fcir llie nSt<e«xaailaatio<iofHud«fila,aBillurtliawpr*iiarlivfnricnidiiallaci.
SOBLYy, RICHARD ».. M. V.
A DioiiODarj- of the Terms Used in Hedtome and the Collateral
ftclBQCes. K-^vi-i-il. with ii'imcnxM *•!>.( jti<.>Il^ t<y Ihaac Uavts M- l>., Intc «litor at
The Aaieriuii Juuraiil uf ibc Matkn] Scicoce*. ta one Urga royal IZumx Tobiaia of SSO
doul4a-«oliiiuiiod paitca. CIollk,tl^; leaili«r, (2.00.
tl l> vb* kMt took at daSalUra* <ra bate, aaa e««ht alnn *• k* aaoo On WtfTt V^.-aamUm
Iftifcxu g-d Ja'iwal Jttrwal.
Bbotsxu k Oo.'a Pdb:
THE STANDARD.
THK
DaiionAL UleDidRi dmionmY
INCtUPINO
£i^Ssli, French, Qernau. ftaSan and Latin Teckmcal Terms used in Mg
the Ccdataral Sciences, and a Series of Tables of Useful Data.
BY
Jaho % Billing?, I^.D., LLD., Ediq. Bud HQfV^ D.C.U ^^
iltmitr ef IJu .fjtiinwt Ae»t¥'X o/Siifua, &irfm D.S:.A^IU.
WITH TUB COLbABOKATION OF
Pmu. TT. o. AlWATKK.
a. M. ttrKXElT, M,I>..
W.T.COi;.tCiI.MAN.M.Di,
JAMRA JJ- FLINT, ». D-
J. II. KII>I>KK. M. K
WILLUM LKE, M.U.
R. LOHIM, }I.Li,
WA)«niv<;roN >iikTTUtir&ttB.
C-S XI^'T, M.ll,
H. C.TAUItUW.M.Ii^
Id two very hundaomo royal octavo volumea oonuiaiag 1874 pacM,
with two colorod plates.
ttr rotunr-CVDM, $«! lA-alA-r, ST 1 Naif M-^tfm, Ar»rM«l IW««, O.AO. ArM*
bjf 8u6*cp4j*tUrH only* ffjiroltfiriijfrtjf^irit uji^f(n1ff*fia ^44^f««CA« ^aMIjJfc— ,
Th» MiUbh«im han gml pI«Miii« in pnamting to tb< ptoftMloo a n«« ptMlmt 1
'linildiii olctlonuj •mbrsciiiK Id oa» alphabtt ftU cumni urmt nacd in emy dcfan* I
nicnt «f medicine in the li>e irrMt lingukgn wiulilultng modern medioj lilenUon. '
For ihp TMt nnil mnipWx Inbor inTnlTcd in surli on iinilcrtakinK DO one tifitcr (|ial^
fled lluin Dr. Itillinjjpi I'jiiUI hair brrn evlvelwl. Ilu liaa jilannxl th« wtirfc, cfaoco tb
inoal nccontplulied men to imist liim in «|i«rinl dcpartnicat*, and pctsanalljr miptt^mA I
and ouuibiiiml ilieir wurk into a conajslirnt luiil luiiforin wlioto.
6p«ciii1 rare hii l>Mn tukni M rviiilfr tli# ilcGnlllnoa clear, sharp nnil um
TI1C7 ore gireti in English, nith ijaoii^uin in French, Uetnian aad IIbIuid of lit* 1
iupottanl Ronla In Englinh nn>1 I^ulln.
Bepnleil HH > dii'tionury, ihercforv, t1ii» lUmlnnl wndc «i]>fili«B the ph;—
■urgeoo uiil tiiecinlUi with all iafuriiiation ouncvming mnlitiil wnnli, Hmple a/iil
pound, found in KnttHiili, Ki'inji rairccl k]>cIHnE, rimr, iharji ilcfinilloiis and actirr
Don, niul furtliennure it tniiblw hint to n>iiHiili forvii:" ""rk* and to Utnlentawl ibe bm*
and inccnuing number of furrign Konla iik>1 in nmliiral Engtiili. It m eapr^allrfan
in (iliiiius ooRiptbiiTiK two, llinio or more nurvls uxd in qicciil wnaw in tha tatiaa
(l«i>iuiiijriiu uf iii«ilicine.
Tliu Hork in, huwoTvr, br more than a ditiieiuiT, and poiukr^ <•' '•— ■■-mrvc/i
mcvcloiHnliii, lui it kitc* in lis body a larss amoiint 01 raluaUo iIhti- .i..| cfafBi-j
oil informntion, mid Kf^tipa In It* tablnt. in a tomlen8><l and omt' riu, k la
anuMal of impurlant iliita wliicb will be connitlinl ilall^ t^ all In avtitc juratiic^
The auniilctcncB of (he work ii luuilu eridenl £7 ill* (act that it drflnc* M>Hj
wjmnte woiui and ]ihnu4i>.
The type ha* been nia«t cnrefully Miected for Inhlnen dikI doaraen, auil avefTtluiy j
beco cfooe to ncure ewe, rapidiiy and dnrabUitjr in i»a.
thr** tnr»l«ts taBllDen'a) Iubdhcm ahltn M*
rlchaii In mtdlnl huniikn. To iwl.l u »• hb-
fulnrt* n± « truta of rrC*r*iiOf .<im* Tt<aiT-
ublva u* BliOD. AnoUiK (vaiurv <rf ili>W",a m
ilm Mciicai'y ol It* drAnltInn*. an nt whMb twi*
bna eliackc-t by caDiparUon allli luwv •<>■
*Uu4uil votlu Is Ida cR*cTt9l lincuiiaa> II *
with, A|«rlfnni} lAf i ' ■i-'r<aoMn*"r
llfin whlDli may b* K " xudretaa
dleUOD*ty.unBli>»nbi . - . n k un>Mr kwv
lorswllnwllljpnnjr no; ivrlio^fal inailMlai
«f in<F laur^tHef ninlarn Tinfiai 1 1 Tlm<
aoi bn two auiuldna aa U> Hia araiM lala* ■
(UUwMc4ilil>ill(«lnn*ryM«l<a4iof|
•ooaRvall t«t* aad cM>dl(lanaor«a
BoJWr at m* haTebcaaaUatotaat^ n*|
liaabeco
tu aoora M eoa wblob wtll al nnfr ■4llt(y lliv
■tii'lvnl BDil ntMlalliliamiiilivnHnlaoniiainfJ'
leai i-iwidilaaar. ciMrand eenivmMa>lTe ilHI
Din>iiii*4r ir«nl« *hoii)d fbm the prim* IWalur* ol
any dJrilanary. an-l In iMi on* ihf talpl aim
a*>cn*bib*i"cli* ibrriai-l ■■■BHIoallon ai)>l<h«
dUIWrvut niHDiBic* or l*rn> la lua ia nicilltlaa
aod laa ooiraiaral *r]tfn4<p» lu laoguaga a* Ian* ai
■■MBpMIUaatih InrldttT' Tn* almoal br*il(*
aaderadbancM ha<* l*»akF|ii IdtIr*. Tbenotfe
laniaarkatil*, uw. lor lU IUIowk. The aaumara-
(WnaaBil ■DMirltHini uadar aash <rent haadlDs
are airthtnslx oomplaia, a* recuiUMIkt Ui» Kor
llMIMngu* anil LM luisuic* cMeBy aniiiloTail
br Bnl>>fal sad madarn apleacK li l> ImiiuiBfMa
(a do }a>tloa l« Uia dlBlUrtiarf VL**>y iwual IIIu*
iraHan. tl fraMctt tn th* InstViVi naAM
Ikoreuahlr wl*niinc cnoda ot *«iaM\u<
•ocabularir andvAaii an aeeatM>a^tM _
•f refrrmca la coanitUtii «<»\u in %b.i
Vi iwAm iiVVMnMauud.aBdlnrMpMloCcQRiidaiaaSBM
(mAt «M«na\YAifAM>KwV— Thi Lv^tw. \nMA,v««i.VWa
OBAT, UEKRY, F. Jt, S.,
LKlurrr (n A*nl<ii:iij nl SI. Ott-t*'i BnpiHII, tri^lwi,
Aaitomy, Deacriptive and Surgicsl. Edlud by T. PtcKBnnio PtcE,
P. R. C. 8.J Siirfctiit til ■ml L«ctur«t on Aiutumij M 81. Ocorga's HoapiUl, Lomlon,
Eiamiuvr in Annii'iiiT, Kojnt Cullcm ol SurKcoim vl Eoslanil. A now Ameriou from
tb« clcvsnlli cii)ar]gRl anil iiopmioa IjondoD etiiiinn, ihoroaglilj rvvitoil Bnil ra^lMd
hr WiuUAM W. Kexx, M. D., frofcMiir o( ^iir^ir? In lli« JctTtinwD Medknl ColUge «f
Philulelpbik. Tu wliicli la adilcJ (he •cmml ADivrican frotu the Intat Koeliah dlllion «f
Lanpmahiw, 1Iu>iual axdSi-iuiicauI'T Lrrimt 1Ioli>k!i(, F.B.C.S. In ona lunfrU
OttnTti viiliiin* of lOOSnM^ witb iis.'> \aifca mnd tUU>rmt« «n|inTin)pi on mod. PlrlM of
•ditKin in U*di! CloUi,|$; leather, (7; half RuMis, (7.60. I'riceof edition id colon
(wcbtlow): Cloth, 97.2S; lauher,9S.2.S; iialf Kuw«,$&7S.
Tliitiiaiilt ooTvnsBionaxlMuIed ranfcwvif nibj«ct*lhui la ciutomnrr in the onllnu^
text-boolu, gifioK nut unly lilt! delaiU Dewnarjr fur ll>« «tu<l«at, but >l» t'hc apiilicatiaD of
tlicav ilelAiU to Ilia iinurlicc of medicine nnJ luner;. It thiw firm* both a fiiiile for th«
Ifanier nnil an iulminlil« wotk ot ntmnacm for ibt ticllee pr*(t!tl(in«r. The eagTavinM
foriD • ifirciul Tenture or tli« work^ many oT them heln^ the siw ot nature, iwarly ul
orig>iu].and baTins the nunia of Ihe vaiioiu parti pnntnl on the bodyof thecut, in
0tie*orflKureinrr*reri>iici> wilh itMorlntlona at th* foot. In iKi* clilkin a amr ilcpanur*
naa hem tsben hj the Imio uf the work wllh the nrlerioi, Tcins ami nerrea diitinKuiahed
bv tllt&n'iit oolorn. The cnpn(ingi thiis form • conipUtc aiv) iiploiiUiil arrini, «hl^ will
Sc*iIt kwul theilnilenc in forming ■<^tnl^ iilraof AiuUm;, naa will alfiiwrre to r«0«tib
• iiiein->ry nf llit«e wlm uinr linil in tht exiiceni'ita uf pnMicv the neccHitj of recatl*
iDg ihe ilctniU of the JiBeciiog-ruum. Conihining, aa it d(«a, a tmnnlele Alloa of
Analuniy willi a tlioroiigli ircuuao on ayvleinalic, doiripliir* nn<l aj'pile-l AiiKtotuT, J
the work will lie foiind of gttM Mrrire Ui all phnicians whu r«*tTe ttiideula in Ihelr I
ofiic«t, rcliering ^Mh preoeptor and pupil of inuca labor in laying the grouwlwurk of ■ *
thorodgh medical education.
Fnrtheconvtnleneeof tiMae who pioffr rmt la pay the alishl inirmM in cml neT«al-
toteil by the uae <it <xAe>n, the rolume la published alio in bloidk alone, end maintained
is ihia uyle at iho price of former odilion*. DoiHriihitandinK ita lancolr inrwaaad hm.
Liinilnuirti, Wtrluai anil Su'^ttal.hj ih«iliitliitciiihlie<l An:l^|lIlipl. Mr. Liilher Iloldea,
ha* been np|i«iiil<d la Ihe pm«iil eilitiuu oa ii wia to the prerioiis one. Thu work giree
Id a clear, conderued and (ynteraniic way all the information by which the pncUtioncr can
determlDe from llie exirrnnl mirfnre of Ihv lx'\y ihe poeition of Internal (dft*. TTiue
aonplete^ Ibe work will furnUb all [he OBiitonoe that can be rcoderei] by typo and
DhMfatiofi in ■nntmntcal itndy.
^iamf*lpaputflJ«(>rkonanat4mretv<iTTH(40,
It n •ufllElvnt 10 ujr t>r tl lliai ililx tJIiIod. itiaak*
•o let AmFflmn adltnf, riirpu-*B all ultiar *4t-
Uam—Jo^.-fi'uA"*^. ii'i. Ah'i, |)«o ni.ian,
A wwfi whIaTi for morfr than twnly feanha*
kad Uia lMai>r alluhrr ttii-lioaliiioD anaconiy
thronallinui tba dvlUBatl laortil ^omrs 10 haoil la
•aell MBUIJP ol •laoutlnn und vfaney of l>il
and llliuMUon a* morn ihan 'o mall* eoul ih*
lut* prMnlia of Tha pru>|wi?iui. It woala <ia in-
itoA dlfflKiili (■> noma ■ fiviun itliania Iha pr»-
•at kta9t\f*ti #H]iik>r] "I *ir%y E^'iiitil Im mandod
or Miimd. aD'l 11 ornl- n« rmpti*! lo t*r Itiat
Iha roraJ work liddvdDcdrormaDyyvar* tneixna
M hofd tha Irtt plaM amoDK aixuoinleal leit-
bonkiL Tlin wcrrk It tiuMlihad atlh tiark aad
eoloKii jilLEf-f. It Iva marrel of t»Tk'mak[nc,—
A^tfiffi ['t-t^litui'iit u'ul ytiifi^ Jbi- Jl, lua.
Oraj'n A-iAl^irny L« (bo Rinti m ui^ ' l^fan I work
iipoo aaaieniy whlah tiM attr hr*D iioMiinvd M
ih« Snallah ft any oUtar laacuaaa.— OiwMaa^l
AaUWb«oknawD«tDlhapuiolia»r h* U r«-
ealTlas Dia baatworkes aaatom* thai s rulillahad
InanyraaRiiacai.— IVgiaujr*l.JVv</tfy.liw. IMT.
Oray'i •ian>l*rJ jtmtotvv ban IXBU Ba4 kUI ba
for yttrt thi iriMaiN for ■iiidaate. Ths book
ntMBonEy 1
■■'m-JMMI /VfH 1/ ITaWm Bmt TmK J»e.
on'/ 10 l>* viamlnad^ls a* peclaiUy uadar<
WM."
t Auo nm iialx xxrAUAT^—
BOLDEy^ LUTUSBf F. Jt, C. A,
awywn lo 51. BarlManu^i enf lit AaHlVUf gwplali. laweaa.
Landmarka, Kvdical aad SurKJCSl. Kemnd American &oa tbe iMeat twriwit
Engliih olili-'n. with aihlill»n« by \V. \\ . Kkkn, M. D,. Pr(if«««a<r uf ArtiMlc Aaalomr lit
Iha I'entu. Aowleiay of Fine Art*. la on« liimu. votiiSM uf 143 |<eeea. Cloth, |l.oO.
DVNOLISOy, ROBLEW M,D,,
Ltit PrMtttjr ^f u,u'^r,n,f Mi.ltti'u in tin Jtfttmtt Utdit^ QiUiff nil^OflM,
UEDICAL LBXtCON: A Diotlooarr of Hedloal Soleiio6 : CvotAiniDc
■ cnnnip riplnnulionof the tarioiu liubjocto aoJ Tifruuuf Analoniy, I'bytiolonr, {"aUiiM-
ncy. Hyf;^iFi»', Tlicraiwiitiai, Pliamiuolagf, PharmairT. Suri;ery, OlBtetrka, Medloal Jori^
pr'^ulen'ce an.1 DealiMry, Nottoee ofCllmaie and of Miuenl Walent, Furmilv hr Ofldaai,
Einpiriinlaiul Keleiic I*repanllcaii. wllJitlie AceanluailM and £lyinol(«T of ttie TWnM,
aiul the Krrodi iiul other ^Twoyne*, eo aa (oooiutituUa French lawellaaBii Ki^bb
Uedivl Leiicon. Edited by RictiAKti J. DvxaLnox, Si. D. In one vety larnand
lMiAa«e royal octavo Tolume uf 1 139 lavM. Cloth. tUO; leather, nieed baadi^ f^M;
wtrj liawleoaM Lalf Riurie, nuaed banda, 18.00.
It to* t>ie ran marli llial M MtWelr ha* no rl*et la Ike EacOak laacwHi* *t aauwawt
b4 exaal «f re*f«MK— XMrfM JfaaMal OaMla.
I
ALLSNf UAMRISOy, M. D.,
A System of Human Anstomy, InoIudiiiR lu Hediokl and Sorgioal
SelAtiOnS. t'ut tlw ii«e ul 1*1*1-111 i<>mn Rnd SliiilniU of Ik(«<llctiip. WlUi U> UArk
dactiw; fiifctkni on i\iAo\«gj. B.t E. U. Suaxbfkak^ U. D., U|ihlWiiKiha( U
iJm PkUaildphU HmpIuiI. C«iii[irnin{f 813 doabb-colimitiiid iiunno |>4«*. wiui M
iitii><nitiaM on 109 full page Iitli(^n|<liic plutM, niiua; of vliidi mr* la ouon *ad MI
•tignviM* In Ik* icic In *ii Scciioiw, cuh in n (xxifolkii. SnUoo I. Hamunn.
tjcdien u. Bones una foitrm. 8<vttnn III. Mr-tci.ra aud Pahis. tUMUon IT.
AxTSun, Vn.sa ahu LrwrHATiL-H. t<«ciion V. Ninron BrmH. 8«cUoa VL
OiKikm or Sknm, or Diuncrimc axd GtwiTo-UttiSAiiy Oroak*, EwoiTOUwr,
Dm-ELOI-MIVT, TSKATOI«OT, StTERriCUL ANATOKT. PoCT-MoaTnl BZAMUIATMIB^
AMi> Ge»KKAu AKu CuMK-Ai. Ivi-KXEi^ I*n(* {icf Section, I3.S0; Kteo Iouk) iatv*
Tnluiii«, clolli. $23.00; very handianie half Rluvdn, micctl band* Ubd gfxn Imck. K&A
FVr tait bf nJurriptiim only. Ap/iiy hi At PuliluluTt.
Tliar* U w hm*
l(Ulab« con*] -fliTAd Bsladyuf ftfifiUfil 4nMoiny
la Ha wIHmi >mi>w — • uiiiiannitii: vi^—niniloii o(
mcb «0Mi>inkBl fipu u tao >.< t|<|ri]til to tlir
Pfattloa c4 ntillolav a« ««1J aj of *iir|:4ty. Qui
Mibor !■ MDDito, acoiinac aud |jr«^ll(^ In li>
Malanaiita, and imctvad^ adi»ir&hly la Intuilng
•■ iDOnM IMO lk*MuilT»riihat 1iK*nara]|y cdo-
aldara4 • 4n tuhft*. Tlia ilniianniaDi •# Blitol-
■■■J la traatod ta> a DUatcrJj niauuaf. and Ui*
fmiai b UaraUad mar byaoa IhuiHiKhly Ikioll-
Itddih tL ThalltaalnUooaaramwlainiri (imi
<ar^ and ar* atmt^y upark.
' pracucal a)4<llvMloa or aBMnmlnJ pBUM
t4
ihr't'dirT-dB]' waoM nf Uia n>«4>ra( ftlelcNa,
tCoUKht kaftin an ao aall |«aa«Mad tor
afanllnit nncaHi. la IHI li«
oaanvilliaMlUiannliaMoata
•riwd antlVaUM Uia> ta man
*oo<*rDtii( aliich Uiay tnaf na>«r ta^
Ku>rat praeUUMian
AiiDg o{ Hr|>riwd intlVaUc*
pCiDI
•Idaratlan. II I* a *urk vkleh la 4aaia*4 M la
Uia b«l of lu kUd l> aay iMUMga^-JMMK j
CLAnKB,W. B.fF,R,C.8. Jt LOCKWOOD,C. B., y,B,C.S.
The DlBseoUx-'i Manual. Inoocnoekel-aiM 12an.ToUuD0<if 8M pa^o^wUi
49 iUtMnUionai. Limp ciMli, red •dpai, $)jA H^Shii^aU Sm»vf llmiaU,fmftil.
l*aaar«Clarlc*aadLMkwondbata«TMMaalaak IntlmaU aiMMtUlos «Mi iMiaata aoaU kva I
■■MM '
thtl «ui hardly lierlnllnlaaaHBCllutald loika
dliaaMar. Ttiair purvoaa, which to '
arrlba tlia tail way Is dl^ay iha
bow 10 da-
analomtfl
Taa'abfurf
atrUDiBra^" li»b**nnillyaualnad. Thayaualto
a lu«ldl<y ol dcmnDnlialloD aad tiaprilc laraaaoia
at aii'itaaliiD. whlc)r only a loug imlntog lad
Etna. With aKft a itaMe a* tbl^ a
Ij aa otuaellra a naamaeiary ar 1V*
AndM <laaliH>v<>aina aarlail. an Madaai . __
iMIUMdaMlyandalMnrblacly lotaaaMad la IM
■Mdy«( laatomy^Aw Orfaaaa JftMaaf aadA^
fItiljKtnM, Affll, UM.
HIJiST, BASTONC, M.D., A PIEBSOL, GEO. A., Jf.Xfc
Pn-funr tf 4biUln»H On Puiiril^ /HfaiMr ../ A •■alt'v m-J ft»t»iwfcm ai
4r fgnHj<fi<a. ' ■*■ (,'—nfiii> ■■/• fma^liiiit^
Homsn ICoiUtrOBtttoB. Hagnitiofnt fcUo, mnuiuing ahnit ISO MgMof HB^j
lUuMfflUd with ra^vinfiL and nanjr plialcignpluc pbtc* froaii nature. In fMir pan^ ]
pric«, «acb, fS. LimitGil cdllioa, for m» ty aDbKriiiuan only. AddrwM tii« pHbUthmJ
TREVM8, FBEHEBICK, E. B. C. 8.,
fif^v*- l**mimtlT«i^ Off AtHttty'ii, at^ A *mJ«< ^'faF jw at lAa £ctadm tfa^yalaL
Surgical Appliod Anatooiy. In «■• podtM-aiw ISmo. votiuue of 640
Witb ai (llnurUioB. Litnj) dolh, ni\ «(|||W, ti.W. 8<« SCwfniH' 5cr><a qT JW«wiij
P"Be 3L ^^
BELLAMY, Bl>WARt>, F. B. C. 8.,
Tho StudoDt's Guide to SnrcioBl Anatomy: Being a DoKripUoa of Vbi I
BOH ImpenaM Kiinioal Kwlawof tlwHumaa Body, ukI JnltwUd aa an lolradtietba M j
opentirefiurgcfy. lDo«ielXni<xTCfaiBHofSOOp<ig«s^viili MilkutnUeoa. CiMk.C'"
WJLSOy, EBAS.VUS. /'. R. 8.
A System of Humim Anatomy, Gsnonl and Spcckl. EdiiBl
QoBBScMT, M. D., Pratascn- of GnunI aixT aurglMl Anatooiy In tli« HkUoI
Okio, In ooe large and hmdionie oclaTo Tolum* of flltt pogei, wUb SS7 lUu
Cloth, KOO; l«.tbpt, »S.OO.
CLELANV, JOMN, X, D., JT. «. 8.,
JVtfwairnf .lMi>«a^aa<rt»Mife(iyi« Quin*! tWfa^ Oa>i>«.
A Dirootory ftor the Dineotion of tha Hamaa Body. Ii
Tolnma of 1*8 |iacta. (loth, 11.29.
HARTSaoityra bakdboor op axatout i
AVP PHY810LOG1. SaMod adHlM, nalawL
la «■• m al Umo. ttitnua «f «U) ta«ia. <aVk n>
WOMMIM. Cl«lh,tl.».
nORtrBlfftBPECiAI.AKATOlIT AKD
our, _ nckUi adlUi^ aXMiMHly rwrkMd
mfTMrl
ibMMlal. [DM«oo(arontnMaaa( MiTne^
wWlil
I
fMBt JOJSWUr, M. />., LL. D.,
Hedloal Phjraltn. A Texi-book far !5liiilei>C>i and PncthiiMer* oT MmUcEm. Is
MWi>ci>iu TL-liinii-of 734 |»);n, villi 376 voadcuu,mnaU_*ori|[iaal. CloUhMi
PROM THE PRBPACB.
TTi* hct thkl B Knowlwlo* of PliyiiiH is iiiiliaiiwiiable U % ihocoogh oadonUnilliiK ti
Medicine lias nut Ivtrii m. fully n-nliu.il in th U oouciI(;)t •« in Eiin>fi«,wMr*lli**iIiiilrahlft
woriu d Dtsplai* snd (inijcl, of IbibcrtM-m mnd of ounwoui G«tiaaa writm OMkitilute a
Icwwb ti •ilucniioiwl IJMnttiira Ui wlilch w« on Atm no nKnUlel. A full B]i|>r«cimlloa
of ll)l> lh« author tniK* will be cuffiviMit jiinillalmi lor plidjis in bixik IWm th* nb-
Maiuia of fab ledum on Ihii deparuiiont of •nonce, deltTcrwl diiriiur maa* vmuv >l tbe
DBivMaiU of il>« City of Ke« York.
Braadly siieKktiic, lliia wark ainu to iinpan h knowlcdg* of the raUll««w ciiilLiif
bMveeo Plinis uiil MciIiciDe in their UMI lUtc »f d#r«loi>uivnI, and lu l-idIhhI.t in lli«
piinnit of lab object «bNt«v«r eniwrienm llie aiilhor lioa gainnl during a l<.<n|t livrind trf
mcbiog (lib fjwclal bnodi uf applinl vcieiicb
Wtlla all enllf *■«—«< Bhytlclm. nitl >Kr*ail>si ; No maa In Amarli-a «*• btuar fItUd Ihaa Xtt.
•kBairMu or phjiilr* f* df*lt>hln (nt llin rnM<l> Pn|wr for Iti* laali li* uodenank, and h* hM f^
«■! lUiteol, nnl^ Ihn-* w^iiiiiHir *iiKii|t<l to ih* (MM Ih* ■Isdtiil Ka< prarlltloDar of aitdl«ln*
NMftlBS Vi l^>■ vdifiWT itililKU «au ba hilly wllb ■ ntuine M ono* r— <»lil» and llMniU(h.
• M Ih* ditllctililr* ■eeauaUnl by ■uidaOM Rtm la ihg ■iii>toa( <rtio haaaom* kDowUdptoT
atumcl l'i> •tii<l)t nf ll>*>i> mibjaou wlltinitl |.1i]->laa tlil> bsok la nHltel, w H •hooii Mm tu
■l>pli'4i.rinii In Iha prnlWolria iliai b« hu ttioMo.
Lir. Urk|itr. wuivid Iwrobar, kn«* iratl Uiailin-
cuUlBatabauirvuaUnJ la brlDgliig hi* •w^*i?t
nlltiln th* |i**p of (h* aranc* ((udanl, nod (hat
b* ba* lUHwAdvd lo mfl |>roT<* oB*e mora thai
Ih* loui la sTlia Ibr aad eiftmlna aiuiliwil* I* Iba
OD* oho ha* tariKht uid l> tMchliaa iliam. The
b<0k U mil prinlad and full* IlliwIinWil.and tn
*T»nt war d***nai fraunil iweomlUvB.— n*
VinlrcnJ ValifiilJ'eNnui^Jnly.UMu
a hDOKlatqia M •llh'sr idi)r*l«a or rhamidrr.
Ttiaaa ar* npeclally C*I1 by lb* laochcr of pby*!'
Ifl*. how*i#r, tmpoaalbls for him I* Imparl a
kooalvdga (4 IJio main ttcu uf bi* BQl|]ac« and
•itebllah lh*n by raauint and »|iarlni*aul dem'
vnMnlloo, aad at ih* aama llinv uodtrtaka I*
•MVh ■« MM) Ih* prld'-lii'** <r
■■a. INuM th* daairaBilliy.
aially, for •om* »ueh iroru
•awh at MHo Uicprld'-lii'rof rri*tnl>'ry or phyi
daalraBlllir, a* mar aay "">
>rlt aa th* vraHDi on*.
BOBEBTHON, J. McGBEOOR, St. A., M, B.,
FtlVKioIogloal PbysiCB. In ooe 12>iiu. vuIium of 537 patica, wilh 319 illiutn-
liotiiL Unii> riolh, SIOOl S«« StiuUnU' Stnf ^ ^anuaU, pnme 31.
Th* ItU* af thia wcrk lulDclnDlly rip1*lfl> th» mxnt*. It irKI b* fouad of ffwtt rmta» t* Ite
■alura of iaa iwntaadL
STot Iha atadanl ot iD*dlrln*, u aiitillarr lo
lait-boaktnphyBlDlop.Bndltvouldbrpaniao-
r naaCUl ■•aculd* la Ua lahunury aipcrl-
tinullUanar. It la a <ar«Aill7 prapand took uf
■ - .^^_ ,„j a* ai' *
Tttinnet, wooia* and Meanln, and *• auob <■•
h**r(ltT faoMamaad ll>— Jmr
Xadlaal ^uaMan^DM. «, U*t.
DJXTOX, jomr a, jf. j>^
• rinftawi mill I' ' fit (i ' ji II iki QiJfjj* nf P^Hrtaw an* Om'tm u, Vae frrk.
DooMuM of tho ClroulstioD of tho Blood. A KblatT of PhTnolo^nl
OnJnluii and DisniTery in rtgnnl l(> l)ie Circiilatloo of Ilia Blood. In on* hawliwDt
ISbio. Toliimo ol 293 page*. Cluth, 1^.
Vr. Paltaa'*«nrk ltth*(rult of ibad'^prMoaruh
•C a enlMfad ntad. aad laibebuay prvmionx it
aainMIUI tab* a lourea a( InaUvciltni. ti wilt
laapli* Mm irll)> a (Mllng orrnillluila aad a<tiiilr-
atlan tor thota ploditlng «nrk*n of ald-a Uma^
who laid III* (eundailan of ih*maininwnll«<n>pl*
ef raarilcal (niaao* aa lts<fa tlaod* Xa* Otoau
waa
In \h' pn»trw of phnlolosloal aludy do Ibnl
u oC ftr*auF momanl, no
nana mora potnplatfly
rvialuUooliad Ui* UiaiMiaa ol uaehan. Uua Ih*
di^n**rT of th* elniulailan of tho UooaL TbM
■iplalaa the •itraonllnaiy IqImm* It baa to alt
HMillFal hlalottuu. Tbo (oliUD* bafMs na la taa
uf iliF« c>r four ahieh hai* b*«a wrliMD wKMn a
f** T>ar> by JloafteMipbnlcaaa*. Illalnaaiaral
raapeou (bo noiit rompMa. Tba tDlaotk ttMHsfe
amall )a lUa. la ose of 1^ n«al MvdlHbl* e**-
IrfbiilloDatromaiiAmarloaapaMlotoa^kalbMoiT
thai baa afvaand^Vad. « *r«. Xv, Pact *, IM.
BELL
LL. F. JEFFREY, M, A.,
tnfiuo' 1/ ClMparafiH A anliiy al Klmi^t OiO*s; Laodan.
CompArUlTe Fhysiolon UuIAdUoiqf. In me t2mn. volumaofUl pafai^
trallona. Iiis(iclolh, 12.00. )ietSbidti»U Stria af MaiKiai*,f*t^»\.
with S3B illurtralloiia.
TtkanaaaaJ i*p«sCinlaaDily aiitidnnt'ibook^ ll lb* b*M •ark Id aiMaaea ta Iha Eoaluk
<l*ar and •tnsla la laDatuwo aiiJ uiaoiemaDL laaaiuca to Maw lo lb*_baB>l* of Ma uadlflal
It b> W4ll Hut aliaDdwiiIy inu>ir>i*.l. aoll* raad- x ^HtM^-HfiM M
■H* aad iDlanailnf. (in th* vh':lo •« an
I HtUto-CtHnryuM JovMf. Mm.
ELLIS, GEORGE VINEBf
Aamttw Prafmtar uf ^-wIiMy •« I'tlvtnUlf CUI«M Iial'*-
Dfunoiutrtttioiu of Anatomy. ' Beinc a finida to iba Raowl«d(« of the
Htunan Bodf bjr IKMactioa. Froiu the eighth and nrtiad Lnndoo eilitlno. la oim r«n
kUKtaccuocUto Minna of 710 laffo, «ilh 349 illmtnUooa. doth, M-^S; laathar, t&M.
BOBERT8, JOBK B, A. M., m7j>,,
Tbo Compoad of Aoaiomy. For uae in tha <ll»acHng^w and In jircfiailog
Jbr anuiiinaiiotM, f a 'joa tSmo. roluoM of 194 (•««•. Uiuf i^MCktlb wwa.
i
8 LtA BllOTBtBS & CO.'B PUBUOATIOII»— PhfBlOloer, Vb
CUAPMAJft MENBT C, M. D..
A TrcatiM on Human FhyittoIogT. In om tuuHbrnne otum iii1«m %I
926 I>agM> villi 00% fiD« cnipiivinBt^ t'loili, ^VO ; IcaUiirr, IlkSO.
11 iv|-rManU •nr ftil>y Uib *iImI»( >ui> "t\ fuih*r,>a4 ll»Ull*rwKI i*< •alctulOMml .
phyilology, TM t<r(HTal<i"irk tiua>p*i:UlT>iiir | inFtniallan to u k4nili*U* b«oh a< taArtuM—
Mtba •ludpnl >Dd {'■'■'tlilont* k> dffoi^'l »><>(*
lo 1^ t*'*^''-^! *p|(tlp«lLoD nf vrll^kDOKn Imlbi
vhlrk (tin adtanr* »! •cioDO* liU (l*cn Id Ihs
profrotloQ tnih)«(lv|«rtmaDt, nhirb may bomii-
■IdrtHl Ih* (snDdallniKif nUtiaiil m#dli'liii> —But-
fii/u jr*lifalaHl8iHfbnl ^MrwILDrc.ltxl.
Mitwrs vlileb haV* ■ pnctlcul iHarlniiea Iha
prvclka of Di^iKclnoAn lurkdljr *i pr*H>*(l^ l*rh-
DtM) (Mllrr* tn k"'" <» mtnulo dmll^ rlRbn
tMa dlr*«>ODi «■ iiiiKd fbi ibt BuldkDc* of du-
dtau le Uie laliomAry. In e»ty ntpftrl tlia
vork tUlfila [U uracnti*, «h*lb*r v ■ rCiniplMa
W<«llw (»[ th* »[iiii«nt or fir lli# pnr>>''>uii for
tb* fwiD«r II I) fo romplrU ihat K# aecd Iwh no
.Vn'IA OmrtUwi Mtt^i JoirvH, X«t. IWI
Tb* votk nnali'r rc^mVMl* iMal
■ludrnlMKl i«*MIUoot'r. VlwilvmeM
lij- ilt> ptoamBln phjralclu of l*4*v
lullan of phjtlolu* tn pncUcal KitnimiDn, ud
ihltwoTk lta4Hlde4linT(*M*BBDllDitil>MMl
mar oltira mtk* In Ui* nurk*)- ll vUl iiiJ^f
Ink* p1«o« WDonf th* Boat raliuhU >»ii tokfc
Jfiifwl Ajt, Kat. », UR.
ll la Iha rrodMUoD of Ml knlltM «*lltliM4«llk
hl> Hork. and atria to laiidra uu4*ala BllkM i»
UiDiliuin aklnlo biiaa
•nd JVnn, Ku*. It, Un.
I
j>jXToy, jojry c, at. j>.,
A 0^«atiBe on Haman Physiologr. DadgMd fcr the luc oT Swdeoto mi
Pnctitbmera of MfdldM. Seventh tdllUin, tli'^ixxit-ltlj mvitd ud rrwrilMm. lom
Tei7 hftDilaooDe ocUto Tolumeof 732p*Bc>i"'i(l>''^^-l'<'""<i'iil*ncraTliinon«Md. CktK
>fi.OO: iHihcr, Sfl.O0.
Fn.m rhs nt-t ai'iicaranra of Iha l>oafc II ha* hai* natar baao la a*; doabi aa to UaaMrtMf
tiavn a (ati^rirr. iitnE " **<l to tha BiKbnr** aDith,— .V, f. Madlrol Jawvoi; OaL Utt,
iWDoirn (A an I'lal U'lc^rr a> lo in* cbaim of Piofawot DMon't nalMBMni aM 4aBarta4bi
almplioliy Bliti tIMh. at a *rll*r. h» dviiyi ipT>i*eUwJ work bw ku faaaai Un Migi ii
■UDi^ortlji III inii*iBllint Bt*n InlrlraN *ubJ*rL>- '-'■■- ■ - • ......... ..
ll Riitirt hr (railfjinii To him In otBan* li>r fi*
qiionpf wtih aMcli hit *ork.«rlll*n (Or riu-laol*
and itrat'tLIJtiB^r*. [a quotad by othar vrllara ou
phfiuliifa- Tbla tiv'i aiuau lit •atiia. and. In
gfU cn«M'ii», lt» oriKloalHv, ll nof n**d( no
■nul. anl i>( ■|>pr<ib*tion. hnHrtti, (or Iha tbnn-
■nnda who haTv tiutllcd It in lu ?ark{iuj aditlon*
ipT>i*aUwJ work bw ku |._ . _ . ._
wliU-l. ilouuld ta mi*«»« la lb* onttuaj
Th* «Dtk It amlDoally *•• for tha aiadtal |n»
IIIIoDar.ilDro llmabunaalfBUjroflkoaahnMat
of pbyvlolocr wlilrh hara a itlreM baarlac as At
■llajjninl* and Iwialmaut af dl*aa*«. T%a WMb K
onewhieh «• eaA blshlr manwiaad i« all lar
CMd*r*.-i>abNajMnattf JfaiMalfMaaii,
r^^^
FOSTER, MICHAEL, M. />., ^. JJ. «.,
i\tlnfuT et r^vituii/vif atui /''eI/i.ui uf TVinity Ctit%is, OiHA'idfi, AflaiU.
Text-Booli of Fhyiiiologj. N«w (ruiinh) and cnbrBcd Ammon fna lb*
£flk and rovicpil Enctiih edition, with nnln nnil additioni^ In one tuadaama ucUto tvt
KM of about 000 |M|;«^ witli olicnit 800 IltiismlloM. Beaif lao/fvitay*.
A RKVIBW OF THE FIFTH ENOLIBH EDITION IS APPKNDBD.
II b ittlicblfiil 14 nixui 1 1'—V "lilcTi ili-r>t«a I Ham, and hia odmxIb* ara mi liittrct^ awij aid
only nni)uallA*d prmlti'. m^Ik ••vit. I>itiiirlir(.>r* I d'lK'ii'mi'.l nn t«lty aad intld ilttalW. IUtw*
D>. Ill* In all C'lwou an Idtal iv>i-Ik>o)i. tt'lili ■
f^niplata. Acf^racaand dvtaHod kiioMlddjt* of l;la
Butijavl, tlie aiiilioi haa aiirciwdsd in (ilrlni; a
tboroujibi)' «nna*riihra aod phl!i»<-'f>l)l0 arctitint
tf 111* tpiaupc \ •tudrni't ai'.-nilon li hapl
IkKVChoul Aitd OD tlio iraai aiid lalli-m 4^ut»-
t«lty«
iri i< 'atawbolawaan jBKlllad liaar
III lit oBly (harauahly mot l*«i > iil
.ll ., inihaRoftUb buu«c*.aMtMin
l> i.ii>liaj.l>' ttrf baal laii-boa* la nw liaOMB
m
POWEIU HENRY, M. B., F. Jt. C. 8.*
fiia>Hncr n ptiptieleg^ Banal Otttrntif Surgttiuof ITnuIand.
HumUQ Ph7>lolOg7. Second edition. Id ime handKimc jracket^st I2auu««)-
niM of 60» |.[>, 'itli as ilTowwllciin. aott.tl.lWL SmSiuitMt' Stritttf Mlammalt, |i,fl.
8IM0N, W., Ph. J>., M. J>.,
nVtHOr (rf Ctnntfrti onJ Tunfolon m iXt OMm tit npiiHani airf Swytpw, aalriBi*!. ad
~ ' r (gf CAwntlry m Ui J(aryl«i4 Otaaft 1/ Aormary.
Manual of Chemlatry. A Guide to Lertum and LaboraWrr nark for BccioMn
la Cbmintrr. A Ti'itti'tik. i,|HviBl1y ni1aptc<t (ot Siiidcnl* of Pbann«n nnd Mtdiclw
New (•ocDnil) nllllon. In »iie fl?o. vi>). of V,^ tips with 44 woodcuta nodT oolond {ilM
UIiuiralijifE 56 of llie mist imjiortaiit chemical IMU. Qolli, 9^iA.
In thlatiaak Iha aiilhor liaa andaainra^ lo maal
IfaawBOtaof (horiudanlofmedlolnaor pbaima^y
ba raaara in hit ohamlcal nndlaa, aad ha hat ■iic-
oaadM in pratoniliiit hl**ubj«Tt*oo1*irl]rUiat no
Ona who rnlly wHhat to ai*j)UTre a fnlr IrnovHdM*
ef rliamlalry ran bU Co do to «IIh ih* help (4 (bit
nvk. The Uttttt aaeilon of iha book ItnalunUIy
Ibal ittTOIad lo Iha mnililiiiallan <d iha rarhon
eattpooad*, or organla ehamlMrr. JIn d-vllrnl
ftaliira It Ih^
(honni ihe ' .
ohrmlfal mi-
nf Iha alkalDi<l>
■ «ui^ ia (
■ I Ll.i. ■jriii.ij
pan iiMtlagof phytlolof InU rbanitCfyUia
on uiahvl* of Iha iiilaa «ill ba CMa4 *«I9
eaJ, ana wall •nliadioitaaaradaatilisimi '
of inadielna.— TAc JMKalJtMarA May «■
Wdhler'a Outlines of Orgaalc Chemistry. Edited farf Firno. Tnmlalal
I7 lu RkM8SE,M.D^ I'b.D. Idoqc 12aic>. vuliuueur&Mpagt*. ClaUi,t3.
LBHH«N>'«MA:njAL OF CHEKICAI. miTK- Ciftpeymt'SrillZB IMATOIITnSDSBASV
i£yS^U )1JS' "^^VJ^S^ "^ "" ***"•'. hwwoa buHHUe Umm V HaaLTa aa* Dte
by llasn Poata. '-■>»- "t»%n «Aiiw.*. >
In ana oMaio t^Aiub*-
\W»ft. YKfl^lH. OlMb.VmWk.
htK Beotiibm k Oo.'b POBLiCATioM— ChemUtiy,
rRANKLAyD,E.,D. C.L.,F.R.S.,AJAPPf F.B.,r.I. C,
If AMitt, Itmiam,
XDOrcAnfo Cbontebrr. In one hDndiom* octavo folnma cf 0TT f»gt» wllb £1
Icuu arxl J plalM. Ctolh, fS.TA ; laathfr, H' S.
wDodcuti
Tills wwk (Aoairi iiiMrMd* Mh*r aarka nf »■ i
«■■» la III* imJIOiI cMUgM. 111! OMUIniT bMUr
•daplwd this Bfir vork DpM cbanlrtnr.vldi whleli
«•■■* KqiiniDWd, M ImpHt ihnE «rakr uit (U!l
ImowledB* cf Ui* Htano* ahMi itudnnu nrnwH-
Iglua ■bould k«TiL PliriliTlkiKWlioffvlitiaKbalr
<h*nl«l litMSleiln H tab ID J iho Urn—. ■uUlU
4ow«n MMUdyihT* work. Tht •iMerlpiloni anH
dVOWfuMMteDi m (Ddl* n plilo Uui in»n U
no dUKcultr In UDiler>UD(l]D| Ihvm^OMoiiwii
jr<>r>«t .VtM^ J««ufT, Ittt.
Thti •iMllaol trwtlM «(ll mi IUI M Uk* It*
ptii« gu una of iTi« xry b*M oa Uia autltcl cf
which It tmtiA W* )dT* bMn nmoh piMMd
vfib lh« rAfnjir»ii«nrW* fend liictd nua»4r Id
■hiKli lbs dim'-ulii" qt eh«mi>>l kvtaiIMn Ha.
D«a«iir]ftiur« riArt tf*«aclav«4 u|ibrUi»«rli«1
II ibuwi iin >iar)r pap tbM lUa tnblvmi.
r*4-lrtlni( (ft* oMrurlil*' of lhl> KiaiiP* taiTJ
of MMnpivhandOD ha* 1"n( uJ tiio(*>'full7 1
HSMaiT Iha aitrDilMl ot I'la t\iOuin.—M»hntt
and&rf tMt JBdwrUr, OctoUr 11, IML
jPOiryBS, GEORGE, Ph. D.
A. Manual of Klemenmry Ohemlstrr; Thcorctka) utA Fnetinl. En
badlring Watti' PkyticuIaii'lItutryanieCYitmuln. Now Ali]cr;aui,rron>til»t<««lftk B(l||IUb^
•dluoa. In oaa Iuk* roj^l I'lbno. toIuid* of 1061 fiftt, wflh 168 UloMntlou on wood
tad « oobrvd iilaln. Clotli, flTS; leather. t3.S5.
fWnw" fVnutni hw ii*«n ■ lUQilitnl tait { work » ena ef lb* t*ry baal lait-booka «p
upon rh^rnl'try fof manf jnar*.
* " ' ham Ilia kn<I phfili'lAna
•■MTanar* la tlil> Muniry auU Id Eaicfuiil. Aa
■ba aslanca baa adiwioad hjr ilia RiakiBR of a«w
Jticct«rt w. Iha wofk ha* haon r<»l>''l ao a* is
kaar U atin«at cf Uia tlln*(, II baa tl*a11lT
nalDMInad l«i poaitlop aa a Millxwk wltb majl'
olaludanU. InUilawnrkara ict<iI*iI FiiIIt: E)<ai,
uiAlMMa alfttad t^ tn-r* fvnii in rli-mlntl
MItMnpM baaUb and dlaea».alc..1i ofth* inwl
iMpecMI klad, aod abouM ba nunlllai In trtTf
"' ' pnwUUonar Wa oao oommoDd Iha
lamotlU ebamlttnr •itaal.^O'HIaMn Jfaf. JirtiM.OM.11,1
or all Uia worka on vtiamlal^ Int^ndad lot Um 4
aaa oC inadlaal aludmU^ Fi>iaa«a' C^tmuttry '_
parhap* Uia mual vldalr aai-L lu ponulailiy h
baaad uddd It* avntlan'W, Tlil* la*t adlUoa cdl^
tain* all of lbs naaarlal found la tha nnitou ,
and 11 laalaaaiirb)b>dlv(>»«^»lom«r WM0 '
Pkmtal and /«rinnie Ctinitlrt. Ail gf (ba mat-i
wr 1> hnu|b( lo Ilia praaaal ataadpcdot ef <ibMnl*'l
raJ Imawladsa. Wa may aafaljr pndlal IW UiMJ
■iiik a«on(lnaaiuworUiefMtiaaad k*or 11 vBlof*
amoDS madlcal alDdrola.— Vim Oiaaaa Jlfamtal
I ntfAvBleal Jnumol, March, IHL
ATTFIELD, JOILN, M. A., Ph, D.. P. I. C, P. R. 8., Etc
Chomintry, Gonoral. Uodicsl and Ph&rmBoaatioBl; Includioc th* ChM^
iMiT cif the U. 8. l'l]iiriTiiii-i>|xrin. A M.iii(inl of ihe Oncrml frinciple* of I In nnifO,
■M their AfiplicBiioii tn Molidnc nml I'banii.-icT. A no* Atnoricui, tmta Utr twilftli
Bof tUh nlitioii, apNinllr rKviMvl ly iha AuiIi't ftir Amfrlcit. In on* hoBilaama rojftl
UDO.T«Uiin«ol7tt2iiit[WS withlUt illuttratioiui. Cloth, tS.TS; leather, t3-^SS.
AUflald'i Ohamlatiy la Iha moat p«p<itar honk
MIHHiB atadaala 01 iDadlolDaand nhainacy, Thli
popularlly hat a (ood, aobauoilal laalt. It raaia
nioD mat iDarlli. Aitnald'awork comblaaala Ilia
bapplaal manB>r a <i*Bi kiikbIiIod of Uia tbaniy
Otanamtairy wllb (ba iriFllcal ■pptlcailin ofihia
fcDOwladna lo tlia afat^day daalliiM o' Iba pby-
ateiaa aad pbarmuliL HIidlKarDimal liahowD
est only lo what ha pula IdM bla work, bul alao In
whal hfl iMTaa oal. Kla bnok la snclaely what
lbaUUaelalDu6>rll> Tba admlnblaamnaiaant
Of Um Mxt >e«fcla( a rvadar ia ^al a pwd ld*a of
olMnlaUT wllhmt Iha old cf •ipanineDli^ and
a[ain li )■ aanod InbAralorT ti'da, and Knal^
roDtAlaa auona maatof wvirarnAcad iDfortnatli. _
thai 11«lllalwayaaarT«MabaQiIybivk ot nttt-
anor. Ba doaa DM allow any uauUlliAl-ia kiiawl-
t4f la altn Iota hia bonk; hi* Idiir y*ara •<(
riparlaQca ntTo prcducad a v4Tk vbicb t* both
•elautlBfl aad practical. hikI uliirli altuu ool
afarylhlns In tha naiilra cf a ivupariluily, and
IherrtD I1«* iba Mi-nl nf lla aunWHk ThU laai
adltlOD aliowa Ihrt mark* ot Iha liMal pMCtaaa
madalneliainliKy and ebamical Uaetilac.— A*
Orlfoiu JfadicaJ aad A«vial Jotmtal, Km. Ml
BLOXAM, CHARLES L.,
Fntfitwr ur <.^^rvl^tfTy ■■ Kutifi OMtft, Ijrtidoit,
ChQcaiBtry, Inorganio and Ori^aaiQ. Nvw Amerion from ilie fifth Loo*
doa cdilkiQ, iburougliljr rcriK-i ui>t mucn improTciL la one vctj' bontkaUM octaro
vdUMof 1k7 ftgm, with -i-i-i ilhwiniiioEw. aalli,$i.OO; Imlher, (aoa
ftwnKBlftnW «■ CO thla -I'mUM anik
*««l Mfaraaoua, IlOllknwIUrly In ac<>j>e aud
alD fton IliM of AtlflalJ. and la lu ■«• b atjually
aondoMtclani. Iia<l"[>u Iba mini diml nialh*
la •latlnglhr iirlncIplaa.hypHlHata tod bti«
Ot (ha aelaoM. lulabHuaR* It an Ian* aodlncld,
aod It* ■rrauaoiaal of niallar ao lo^lcai la ao-
Bea Ihia (h* atudaot naitr baa unaflan lo
a Ihat chamlaUy la a hard atndy. Haeh
il* foU ID aaparUnaaiAl Uloitratloiu of
1 pAnvlplaa and ptiaoonaDa, and Iha
»afCODd<ielinB(boaaaiparfnwn(*. Tbabook
mala lalaa Iha pOalHoa It haaalwija bald atUtaW
Iha liaat maouala nt aaneral cbanlalry (a iha Mtr
IM> lanKUaaa.— i>rtrbi( tatett. Fab. IWt,
Wa know d bo liwaxt>a en cbamMiy whioh
«fintatBa M niub innlnl laSanaaUaa In (h&,
■ana Duaabtr ef pMaa. Tlw bask Can bo ri
adaoud BDl only lolho DOoMof Uioaawho c.
a Mhrably eoniplalo cwiiraw of ahanditiT, tal i
•0 Iha naaib of Uioaa who ilaatra oot* a nu
kwirtaiteaorthoaubjatt. Illabalhai '
taitbooE, and a uaonil btok of rafanai
Madiral i-d Airjwaf J^ivhI, J vaa 1^ tWL
GSEEyE, WZLLIAM H., M, J>.»
Dmmutratvr tf OtmtU&y ta IM Jtfadiasl O^ptrMaal tf (M Paliwa% of i^ililnala.
AKanualorUedloalChemlsbT. Fur tbeDMoTSiadntii BMadnpeaiBov
nan's Utilitnl Cbemulrj. In one lano. Tvlutue t/ SIO pagMt wtUi 74 lUiu. CIolli, tUS,
tllaanoeUa maniia] of thra* biiadrad pMaa. 1 Ibo tveofotliaa of otapenada daa lo paUioloctral
■lilMaa •uallani aummary of Iba »hI naUwda ooadWona. Tha 4aMaUM of poUoo* U Irtalad
SfHBjMBiibalfqiadaaailaalMaotlkabadr.hotii with aanUval Minsao tor iba^purwa of ,tto aau-
Af a* atlwafif «/ UaOr aormal eoDMiliioal nd , daMM f«tMUMNnr-V*teK It. •i(Q«Bk 1
IiEA BaoTDEM Jt Co.'b PuBLtoATiotrs — Cb^nilAtry.
JtSMSSN; IBA» M. J>„ Ph, J>„
fiimlplea of ThsoreUoal Obomlatry, with nwcU nrfemm lo ik* CuHtiUh
1 of CbnalcalOiBpoUDdA. New (third) Bod UiorDUgbtjrroviMdadlttoB.
■OHM rujrnl liam. TolunM of 3III pagw.
Tkl( Kork of Dr. Bvlun ■■ Uw *«]r IciVtMnk
Mcil'd, KBd Ui> ia«<lkBl >liicl*ol i>lu> hu K ■(
M* ODsm* (oilaiM la uuwk, aw, U ba elMiao.
Bkk* almHIt biDlUu wllh Miy bnui(>h ot oham-
lalTT ahleh )iMB«r ilatln lo puiHi*. 11 ■vuld be
dinfouli Injtad to fluil • mots lucid, full, kod M
Uwiana(lnt«roifipvi aaiilJcvUun of ilio plillo*-
•pbr OttfirlDltliy. UiMi Ifie lx>i>li bcfut* ii>. mid
Vtt racoaunaii(S it to th* rmrnfiil ^bd Iini«ni»J
Innnahaod-
«IUllBUMeOl<»llt«b«l1ll«Hf*< MtMM^tf
(iosl Jnnal, Jwinwy, UM.
llt*khMlthfDli(ca«kH**iHa4anud Iv
■ third (dlllPBbftuchaboek •■ Ikta. Tlitoi*^
tlon to IftTEH Ihao til* l»i hy aba*! «*nnif «••
Haaj, and much ef li ha* bVMi mrtftfa. IM«
btinglag ti tiitir abr*Ml af the IMhi laraaaWb
Unoa.— S, " " '
r. jr««M( /owmI, Dvc ■!, IMI.
CHARL^H, T. CRAN8TOVN, M. J)., F. C. 8., Jf. A,
^brmtrly jHl. /Vof. mt Daawut. qf Olfnii'I'v wl C'AfMltal Fl^la, Qwva't f'trirr. J
Tbo Elements of Pbystologlcal snd FarhologiatI OhtmiKOT* -&
flnnillnok (»r MedicaJ StudeoU aAil Pniriiiicnim. OiDUuniiig s |tm«fml KiraiM of
h' lit rill- .01, {''inhIi- nnd Diipialinn, and the CliPDiulry of tli* TiMtiM, OtxMu, t^ttMliomaai
Kicr«<iuiu iif ilip B<«ly in IImIiIi kikI in niMiim. TogMh«T with ln« mctbod* fcr pt^
pBciag «r aciKraluig Uwu cbivT inniilitiiciiU, u «l» Tor thtit aTnmlMtyiai !■ daHil, ud
MoiiUtiMBfUafauaofmnwaickleDnraeofiiMnictieaferctudraU. In on* h»i»JM«D» ottew
volniM of 468 pa^M^ with >8 irondcuu and 1 colored plttt«. CtaUi. |3J0.
Dr. Chwlaa In l\i\tj Impriiaid wllti Die Imnor. I nowadara. Dr. Ch^a* haa lUaoMi ■■r> Man
tanaa and (invUall raoHi a( lila lublHl, wid IiB I to tlia oludUallen at nrlaafT Walatlaft B« aam
tiHtrHiadUinaeompBlaatanilloalrurllTa R»a- thli lakh oiiioh drlall. lad nt la • fnMlatf ••«
wm. W* caDiipl (vominaiid a bKtrr book Ihmn lnt»niRllil« nuumer. la Ate^ tM (aliM* lM> MM
tliaprr*«-iii. In bf4, U Allaacap la madical (fii.
bODtn, and Uiki l> a liilag •liivh ema ranly b* (aid
IntMIlRlMi ^ _ __.
hla look alih maoy naaUcBl Ma«a, ■,
■rd, bv»ln>>ar)i\ IBM.
nOFFMAJf]!ftt^.tA.M.,rit.l>.» A POWES, F.B., Ph.D.,
AUftnual of Chetnlcikl Analjii^-Mkrolicd uUm Exanufutianof M«£dMl
Cbanlcab unl ilicir PrriNitstioiu. Bdiw ft (rtiliU Rirllic Dolrrmiiuuionor thatr IdKlUf
■nd Quality, nnd fur tha Daurtloa of IdiimtIUw unci Ailntipnlkoia, For lh« vm m
Phamadit*, PbTriciaiu^ DnioiMs and tlanntacWring C1ieBiL'>ia. Mid PharaaceirtitKl wd
ibdfaal StudoMa. Third edition, «iuir«l j rewritttn and miidi cabrnd. la oa* *«t
haodioiDeoclaTUTotiiineof m ptfita, with 179 lUncrlratiocu. Cloth, tuS.
Wa wDOalulai* tha aiiihor on Ui* apiaaniBn
it IhaUiltdadllloaariliUwDTk, pabtMiadforllia
nl Uma Id ihia (miiBlnr aUi, II laadiwlrablaan.l
tha iDlinmialioa )l nndvnaliH to wppl/ la both
•ilaorin and tmMwDnhy, Tba aalacOoa of pn>
■[nine U>a puflU' of iha nhalan.
" oil
I
tfOrdaunBlDiBC I
eaacf *hlqh HlnaU UooIUatud Ihadaaarip-
llou eT ttaam ulnituUtly air*l°lt- Ua»«m,(|>
aE<4pll<inAjJy fiM frani IjpDf raphtc«l arra^
hai> aa btKllailea In rvaommandlu ii u T
■ho ar« vDffMKd bIiiim in tha laaaiuSMarBoti
iMiInNPf iiadlcln*] «baBilCftlL— .£•■«
caatKoiiAiwwlland Trtmuiittamn, UB.
nl In tha urAirniaatfa of a«cb iaal;
> elianiDWrlaUe* ara iirajirra^ la tha n*
A,vhlrihvavBn baafUly f aPaanaaanJ aa i
CLOWES, FBAJfK, X>. 8c,, Ltmaon,
An SIem«atar7 Treatise oq PrAOttoal Chet&uti7 and QualUatini
lOOrganlO Anal]^. .Sivcinlly ■daiilcd for OM in the Ijdnnaarica of Retook mII
OoUogca and bf Bagiimcn. Third AiMrieattfroai tholDunhaad lariHil Engllab ndlltaibl
In DIM ISmo. «oUiraa of !IS7 v^gt*^ with tA illitalntiaiw. Cloth, 92,iO.
Thla arark ba* loaa bcvn a hvorlM vlth Uboi»
lary lBam«to» an a«Duni at lla vMcaiBlle plaa,
<*rrrlac Uia Madaol (Mr *•</ t\tf bom tbaalnplaal
Dnaallan* td phHnloJ aiialjrpu. U Iha mora ranni-
dUa bfablama- fiaium «Im ■* (ommMidaMc
Mr* tha r*«ularii7 and ar*lan d*aiau<l*d of l)i«
RALPE, CHARLES lt.,~M7^.,
Aitubnt Pt^tvutn at tU Ionian HotpUoi.
Cliniost Chomutry. In ooo imrkaf !i« l^oiu. tdIiko* of 314
UlDiinliont, Llin]> dotJi, red <dg«^ f 1.60.
Till* U aaa of Iba moM ta*im<HlTe tltLla work*
■hat aa l>*ia nial wlUi In a lunt Udib, The aLiilioc
i> a phjalctae aud uh/dutoilal, «> mr'A >• a rlirm
■•i,<«n>*iiu>aU> Uia li« ' '
neol, lalUn; ~
ftsaw.af
aludiml
ThoMcl
•dlUOB,
Irthmorr ■uldafoiiha-wdaMoriMtMala* .
leal aoalrite^irw ^ar* JVaAmJ jMn«L 0« % '
utm.
F. R, C. P.,
I'Tf'
•mH
■■cue auu un/Huiogiin, «> iCii a< a riirni'
*i|U>aU]i Uia book l> Boqaallllailly inaa-
Una lb* nhyalalaa loal ■bal ba mtgM to
r i£a apprieautiiw «t ebamlrt^ la toadl-
Sm Srwfml/ Sma of V-tnttaU, paga U.
elua. Dr. ttalfc la IbWBUBhIy ao^ualaMd «U Ika
laUKt oDutrllialhii)* 10 hk af'l^DI<>. asd li ta atM
i*rrt*hlna to And Uw anbjaoi <iaaH wUli ao «mmv
and Blmp^,7M laanohoManl I
, modam tDlanUfle nMteda
I JiHard, PatmiT 1 IMi.
inl hatwoM "U **
and apwC »»«■■
CLASSEN, ALEXAITDRR,
Elementary QuttntitatiTe Analysis. TmulNtol. with notaa and aildilji
EuQAit F.SuiTii, Pli. Ii„ Avittani Profmor of Cbemiatr; in the Town* Bci«Dtific
Ui^T«n)lr of P«t>in. Id one IZmo. volume of 334 imgtm, wilh S6 illiu. Cloth, fS^W.
It la |i(«b*faly iba bulBiaBUklof •D*I(irnilar7| aad (h«a *dreR<-lniialhaanaJ7itl*of BlMankwi
ona ■
OMOi* aitaM. iMomiub aa Ua m«tti«4k •>» Mm I anah pN>4Mt»aaat* nial *IUi la upHad (toM*
baL It laMtM kn«SMnD)«*,«MitntMiC\ii%<inih\«j, \\ \* «b\b«!ag«awaMa fcMk fcr aaadaMak
ala«la dMOfOtlaaUMW, nnm*4 tor »>;■»»»««».', AaWi*ir|^»a«to.J"-—H '»ii*nr».Oa^Mi
LiA ItnorHuui Jt Oo.'8 PnBLiOAXKWB— Ptukmi., Mat. Med.* Tharap. IT
I
llAHE, HOBART AMOBY, B. Sc„ 3f, !>.,
IVni'no' nf Vnfiril JTHiAa Ao/I nn-tpii<. fxi ■■> It* J-frrrt^ J(>rfiA>r n^llffo "f PidnnltlpiMi
.y4ftJ^-v'f n^ (Vi.i...i...n fi.'H* AfTu«->i' r*« f;-.>f«i.v6«i«i nM-*>nr.,>ai ^ titn
A Toxt-Book of Friu)tiO*l Thorapoutio*; Wiib l-jit<i>cul IMcmM* to tli«
Apriintloti of R^ninJia) TkCtuMir** lo IXkvnM unil iImit ECniiiioTmrat upcni ■ ItMlooal
bM& Wilh ■pc'UI cliat<ii-n ti,v Dim. O. E. De 8cniri;isin^ Rehcarp Mariim,
J. DoWAlUt RitevM ami Baiiton C. lliKn'. N«v (3d| loJ rcrianl rdilinn. In one
biBlaWM ortaTO viiliiciie <•{ ^M) |>*|c». I.'ldb, (:{.T-V; Icnibcr, $4.TA. J<y« rHwfjr.
Tkl* *wk liM NKWlrvit Ih* rar* illitlaAIInD niliar iHliirM at Uil> pfMilcal'* Iwl^l lr*alM»
■omOK MwJItwl <'aft»<>if r»»»ililnK> iMnndrtliltai I irhlrh «lll mak* raOnni* >a It npfmlaai t»4
•li modili* ntUr ll> llrM k^petiwio*. W* sola { ■•rotlubtB, an Ui* wnu«*n«n( of (UIm «< iliuc*
■anoDBths IspvrtaalK** ftoluf** ckancterlMDi , >n'l iIImbch In ■IpliBbeilcd criIh, ■BConllBff <a
tl>a>M00ilnllllan,ul4lilani>lluhrai»Uiian«Hi< , Uxlr l\niili>)i niK»Mi Ifct lairadMllm a( Ih*
tks iha nmvdlM r>v*aUr ■<l>l*d to (ha Mki>H* pro pant loi.' <Jthe Rrllhli riinrniwic-piiilk: aifAw
Hadkai Ih* mdlioa «l •mpI'VlWt IM MM rnrai I llM at 4Tnf* altlclnal mad -ranfflrtna). la ad<ltlK«
Uin UH or Buiu*u>iau Id Ihu usalounl ■>( looBmik- I b> ili* cfaanl ladei, a hpIodi uiiiI aiiiiuttAfy
IM alalia Had oIIIhI ■fflrcllont. Uioy ii*» pr»- In^m tftdl»WMMi 'r«iiwJW«hM>M«apf laiii
MrlHloni kat« a)«« bMD lOMrtvd lo IllaitnW wMi'h wiW nndttt Iha ooDtaaU vtdljr ftiwIMfc
iba Mil tnodu <tt aiiplTtai r«ina4l**. abmik I —TV JVonm' iIjk Jiilr In, inL
HARE, nORART AMORY, B. Sc,7~M. D., Editor.
A 8fat«[u of Practical Therapentios ; By AmiriRui an<l I'r^rflKn Autlt<>n.
In A Mrira of ranuilniliiNW bjr MTviitj-vercn ciaiii«nl phjiuiaus. In three uncc wUto
tDlotDBa of abcul 1000 pag« Mch, wilh Uliwlmlkina. For lalt by toAtrripiBu* mtf.
BBuyroy, t. lauder^m.d,. d-Sc, e.r.s., f.r.c^p.,
A Tflxt-Book of PbarmaoolosTi Tberapontioi uia Htttaria Hodio*;
bwhidiutlM riurnucj, ih« Flivuuloctnl Aeiian »Bd th* TliarapMUu) Uwaof Dnin.
ThM «duioii. OoUfo, 1S0& pa«««. &ti Uluairaiiow. Qotk, ^Uft; iMlbar. 9A.NI.
Ro mrda of pralM an Ii**dt4 tr<r IM* «aili. Iter i mad* la tarlona dliMIHiia* la Ik* an «r thawpaw-
II hai alnadr (IH-Iifn lor ilatUlii fnrmar ■^iUoni. | Una. lod II vowvlandiiuiufnllBd Ih IU IliotoiiChiy
ll •*» by unaolmout «otia«>l plB(«d amoni Uia tpltnlillBpriaiilaliaaMthamodPtof ira* awloo.
fDrvamI l»«li> on Uu miI^ImI (tot pabllahad la Ho on* wnn olaha* in b» lutlv ii» M th« flmfi ti
laj liTfitri. an It 1^ I Vttirr Itli fcn rrni *ai1 iiiirtl»4 r IkU nclrntw taa mati U atcki^l tha atady of Dr.
(ka a«M nlahlj II la ifiprvoUlad. Th* praaani i BruMon't ■ork. TW lodaia* ara ai«allau(. aail
•AUoa coaMtoa tnueh naw mallar, (ha 1am linn [ aiM bdI a tlilla la (hs pranltral faliM of th* baak.
W ablch hw baao H«aMlta(»d bjr (ha a>t>ao<wt| — Jtah«l ffmnt May », MK
MAISCir, JOONM., Phar. D.,
A KftDUal of Organlo BCMerla Hodioa; Being a tiuid* Lo Maierii Madica of
Uic VcgeUble mk) AninuU Kin^laiai. For (bo ii*o of Sluilniu, DnMgpM«i VhttwMiSalM
■nd Phjitciaiu. N'a« f 4tli) ailltiDii, llionHtghlv r«TWI. In on* IwaiMcinM n^*! ISmo.
of J13»fiBKW.<itlliZ5i)Uliuiu«Li>ana. ClotK.IS.U^
wlaaw
'h>N*iiua', nnt
^ . Ialartalf4 la malaria nadlcia,
>lai*!h1.»l»iiu»1, -
to lu faurtli odu:
For Iha Amarioaa pi ^
■hanitkahlrhirfll tlf hlmlhaptmaafj kiHl
MMlalfcaaMlHl*a;,H>rU7bwaiM*ll>*l>it l>
in*C naelM, and IVaa tram aonaeanaiy nMi '- r.
t.ua
nnt uublUhad U ua and dob
ilaa, la aa lodUpanMNa book.
■ phanna«aa(leal ttndaat ll I*
Bd pamr kanuM of (ha piimarKM (lliiMnlkiia, I'lali
■kini brloc hull wnith knoalac ImnMlaialr b» j UKL
tuta hki Bjaa. Tlial UaaannlUporraaaalDUila
rwHCI UM raplil auMoulaa af sdUion* it Iha haM
aiMmt*. Iibitafanruabaaksf (IkaJliDarliaa
atDdani bno anlil^ at MalMh'n Mfvnl biiii4rt>l
;<»r«nial Mu4«Ma. TVa anancemanl at lU n*"
tfaif ihDin Iha praatlfal vtoAaef of Iha baalL
Mululi'* •TMam of olaatlAcMlaa It aaqr aadMOi-
it alianatra.— W »n»« imHm l« tonutt OWMJ.
PARRJSn, EDWARD,
LaU fVaAu>rc/ Uj VuBrnml Prmhftf t^rmn^ 1^ t\t PltUuMplit* QMm^ ^^rmart^
A TreaUaa od PhArmaay: Dad^icd ■• a Tut-book for tfa* Sradnit, hkI m ■
Oniilc (or Ibo I'hjriiicino and Phannaraiitiit. Wilh mnay Fonuala ukI Pnacrlplkm.
Pinb •lliiuo, tlioToiidilT nriiiol, \yj Thumab S. Wubakd, Plt.O. In mm handaonM
ocUTO TohuQC oT 1093 Jftget, wUk 856 UlortnOJoBa. CliiUi,t&Oa;lNlber, 16.00.
oda of aomUaalloD ara «««**nitii, oaa tMmi la
laan thia vorH aoi of lb* IIM of UiaU itArka «f
(vCifVDOa. Tte ttmttWf MapUUf ar. alia bum
almin h* >b a maaaara hb ovn pkatVaanltL *ui
Had It ladl«aan«ablaL Xaim.«M MMtaal Jhm,
HarehV^ML
Xo IkonniBb-flalDBpbannaolM «in lUI lo
hlmwlf et to naaful a pilda M praelloa, and do
ph)i>i(^UB who praparlr ■nunaiM Ilia talnaof an
anrunia kBoirlBdcB of Oia rrmaillal afaala om-
pl«X*d bv hlo Iq dall* praMloe, an lar la Ikair
■latlMlltjr,Mm|M>ndlltr aad niaaiallMUtaiDalk-
BVBXAJrX, J>r. L.,
l\tfmivr of nytwlm i> lU Painriffy tf 2WwA.
Sxparimeotal Pharmaoolosy. A IlaniUirak of Ustboda far Pwaralalm tk»
Phjrtiologiol Actiona of Dnict. 'nuMlotad, with tlia Antbor'a paraWo^ and will.
WtunalTa addltlona, bv Rosanr M bamc Siuth, 31 D- DamwiwnUoc of Fhmlolan in lbs
Unittnitj of Pmio^rlmila. 12nia, IM pasta, with 33 iJliutntiona Clolb, 91-Ml
STIXlA ALFRED, M. D., LL. l>.p
Tharspoatic* KOd Haurla UediOL A STMoualio Tnatiiaaii ika Acikm and
Data of Uedkinal AxcdIk, indudinf tbalr Daacriptioo and Bidorr. FonitU adlUom
fmtimd Mill! faUmit. In ttr luprt and hilHlTWiT TntliTTi TrftniMl.Trti^iVTi^TT.'^''^*' '
OoUi, 910.00: /wibor, |12.0(f.
BTZLL^, A,, M. J>., LL. J>., Jb MAISCH, J. Jf., Phar. D^
#t lit P'-irviay
. _ . MloftMT^iOTJfraj-
Hh <if iri>tMH oaJaf OtMMt JTtMMH
The National Dispensatory. |
COMTAHtlMO THEKMWRAL HISTORT. CM£M/ST/tT. PUMRHACr. ACTIONS AMD VS£S »F
MSOICIt/ES. IKCLaDIMG THOSE RECOGKIZED IM THE PHAHMACOfailS OF THE
OHfTEO STATES, OHCA T BKITAIH AHO QtHmAHT. WITH UaMEROVS
ttEFCSEHCES TO THE FREHICH CODEX.
Fourth tditioin rrvLMtl, aii'l (v>vcniiKll» new ISriliili IliruuiMopfcia. In oat mtf-
nificetit impcri&l ocMto rolume of I'M nigot, trilli 31 1 tlabonle wiKravln^ Mr*
Is clotli, f::^^ : Imtlicr, rabed bmtwU, S8.00. \»Thu mrli vUl bi ftrmuiul nU AUrM
Baidy BfferertCf TKumk-Utlir ta^tx fyrt\M in adddim la tltt print m amy ily/t v/ iMmf
dUcnory Kits >*<»<<«I dua •ilsnUoa.— i
1
It l> ■llh mii'^h F'l»<"'i' ihitl »'p ("Drill pdlilon
of lhk*inuEiiinocDi Hf'fL l>r#t«lid(l. TliBuifhtn*
•nd pablblian litis mx'ii Ci f«l proad cj lliln,
Um movt i>oinpffl»>r(i"lT», rlthnnu and aorurBU , , . .
work or (he kind <•*' rniiu-l tn ini* rniiatrr; ll fhavQiuchupirclBiiciiiDttliliKiMlonkHUnll
I"- tui four ■dlltina wMhln ifc* foopi
aft llaaml Im^ir. Nnr. 1»T.
V> itilnk U ■ Bmilii fm Rmaraii
rlahnrua uid uvuiaH . |irohMluinrDi»llfinr idJ ihalW i4iuinMT>H>a
> itilnk l( ■ Bmilii fm Rmaraiitlallini Udl It*
M DO w«DdBr IMt II lit> Ixoomo ih* (landuil vi- tui four ■dlltina wMhln ifc* fooparMmlr WW
Iharlty tot holli IJi* in»dl(«] ud phannii'-BiiKr*) n«r[ad at •lahi r*u>. Ths mUitrT «Uh vlilr* li
pnlVHilfni, ui-1 thai ftnir fdttinna han h>#n r*- d«*l*ar* orV immal a naiei* iIiM .tlik^riKi
' lo ilifitlj tite MnrUAI and Ii>pr«a4^nc I'TijrtlalaD nor Uia IdiarmHlM oaA do i>»lli<«l Ito
* " " " l«
JnlrtJ lo aniitlr
•ciiajid ilDC* li> llnl afiptvianca lu IfTu. Tii«
•nlira fttiJ liM hrm kp>u9 ut^i and Dia laHfju*
I ■rlliitt rrrl'ad in wvcrdaniw w(lli ll>* lalml
I daTalDpmanl* rciatilmi iha aitrlbutat and than-
pauliual acU-iii of •iiiifi. Thcrcmnllraaf r*««il
pimMt
Ilia iiliarniHlM oaA do kIIIki
UlMI lalt-bDolo ue ltirm.MM«tall]rtli-<>«lluil>n
vuacfTbnto a*d mupcahapalv* B* IfrUofiau n*
l-onk la 1nFt*4Ti>arn*dlMblabMh uihaanllian
and loltietmuir
Ma/»t.
rilihan.-JCfw IVl Xa4ui*l,f<
COHEN, SOLOMOy 80LIS. M. ».,
A Handbook of Applied Therupouticii. Bdng m Bia^r of rrincirtah
A])pllc«bl« ■ti'l nn RifH^Mitinn of MnltCHit. KiuplnrM In iho UMingCIuttit of lli« fftd.
Id oDe lOTSC ISmu. volumi^, wilh illiiitlntioD>, Preparing.
TABQUHARSOy, ItOBERT, M. D., F. B. C. P., LL. Z>^
f^Hfitf rv uM ,\frirma XrAttn At Si. X"-y'* Ba*t*iiAl MBAfAt 3fJntit^ £d444K
A Qulde to Therapeatics and Materia Modloa. N«ir ((mitII)) AiMriow,
tram llic (uiinL Kngllxli *-1iliaa. KiilniKuI iixl aiht'lnl lu llir V. S. FliAroMmfKela. Bj
PiuKK WooiMiURy, M. !>-, l'r(>r«aor of Materia Mcdim and TberapMlic* UM lliunl
UoUdue In ^i Mpdla>-Cliknir)|i<al Collq* of Pbiliuliilpliiik. In net h»»«bmn> ISma
I
Ilnu>(ennrilTt*i*||Ud*d>athanoa* modaro
work CI Ita kind, tl u mndw. tm compli'tp.
OoikUI&Ing an aofAUnt <tf hU ivfri^cTin tt<ai hava
m |d*n lo iha llrlrlnh Bfid t'olird Stnt— I'iiarnii
copafftB.aa well a!«ron*ldf<rtiiji all non^nkokal bvl
lni|M>rlAala*«drD|f*,U brodniC' In fAf^l a mlDlaliara
dlap*Di*(tiiT^A>iyl< Mtlifil Jouiital, Juaa. lUt.
An ttp^wif acfrVtlta fv^tnra U an arranfO
nam ))]r irhlch Ihrt-bralcl "■!''■ I and ihatapouiNal
•rllnn* et rarlnn* nifivdi** ir* alwwa la iwiM
MluniD*. Tnia alda «ra«tly H «H» XHalla* m1
iKlUlalaa atuilir. Tte AMaMcas ailltoi ha* •»
largad Iha waii w u la fnalia II l*<luita all lb*
raiiHdlM aod |ii«Miratlan* fa Iha V. 9. ■'lunB»
mfala Tbe bMk ItaniMI aaluahl* adillilna M
tha lk>lariraMlHanQ(hto(no*llfniK«1wii MtfvA
—Amwtimm I'rasNUsMT ■«< JTaa^ Sot. •, Um.
EltES, BOBERT T., M, J>.y
JiilMfH J-rufrjiir ;f CTiviral liidlciM is ifnmrd I'-lr(riifv, JMImI OtfrrlmfM.
A Toxt-Book of ThorapeuUo wad Malflria Uedloa. ImmlMt Inrilw ,
the of Slii'lriio nn-i Pmrliiion^r*. OcMto; M4 para. Clolh, 98.60: I«m1i«-. $A.att.
Tb«pr«a(»lonrk>'«intda*llni'dIi>lali*apronl-'a'>inni*Iid Uia book aad caaBWuial* tlia Mtbw
acnt tlacaaaamt-biiDkotithainibJMIiiir ohlati
n iraata. li pDwaaaaa all (ha naiwlnla which wa
aiiact la a twoV t4 tu kind, anab aa niaaltaaata,
olaiaan, a ^dlalon* rlaivllldallen. and a raA*aii-
alilii Jiiiii ll inaiiiliiii All lhf> nawMi drufa
Ot fromla* an uiawd OL Tha vllolpal indM al
Uia««d VDI b* fanud nrt atttut. Vit hrarilljr
on liaTinc ntMnvad ao aood a an*.— if. T. Uttttt
Aw-'-iL Frh. IK IML
t'r, Edaa' limk rafirvMnta b^'.ltc Ihan »UT aUw
book tha pneilMl IbanpaniUw nt Ika praaaai
day. ThabMklaaLhanuiAlrpraMint oua. Tba
■■lawlOiMlnn nf raniadl** na* tabraooa (a l»*k
ihanpniilqatlon.— nnnaacnIMat Jlra.JaB.UA
BBUCE, J. MITCHELL, 3L D., P. R. C. P., ■
fHviiManaad J.aNur(rfn JTiUcwi jrnbM*>l nirnpnlntM lluirt^ »a«i 8—piltl, l^id».
KatorisKadiOBBndTherapeutlos. .^n IntrodiKiioa toIUiUniiJ TrauiiMnt.
FmtUi edition. Unia,Ulpicei^ CUh, 11.50. Stu SlmJmliStntnf Uuny^ fift tU
eMZPPITfl. BOBEBT EO LE8FIELD, Jf. D.
A UDlVOrwl Formulary, conuiulnK IIm Melluidt ot Vetparlag uirf AdMilal^
tfrincOffidtlBlukdcAkeT VftirinfA. "tVe vWA« tAa^uAvtV^i^Uaaa aul PtuutnaMM-
In oM ocUto Tduiaq c!
. • a\i a* WuATaSioKa 0«iu.VJ«> ■,'w«i&«i ,V.ifc.
■eUiBEH, llEXEAGE, M. !>.,
Practical Pathology and Morbid Histology. In nao rcrr liandaoioo ocUta
volume or 31 4 {i9Hi^. wilhGiiillii<.ifmiIi>nk,iucMll7pliobP)Eraplilc ClutbiV^TIS. Jiutrtady.
t»r. ait^M^ ■I'ltMlahAl r«t>uUil'>a »• ■ invtor I (ha (natul pAalMa >»liu (o tM labont'iTr
Th* Koltioi'i OKlboila at ll*nl*ali>|
.•■tlcl|«(* ■ vDik of icr*a> tvu*, and la inli ■«
•ra Ddl illiapfmlnwd. Tho obifiMT* ila»M<l la
ini* Mlbjui an modati if camplatanai*^ pon-
~* ' tn (uon a in»na«r m U nodM iham *>(
■MkM. Th« aoibori OKlboila or tukni*ali>a>
•acUtn'ODIUnic au-l •talnlDR at* Nlnn In iltuli.
■ ■" fall *
■nd ar* ■eooniMiilad ^
_, -Juabia fDrmutv fur
iltdint Xmt, Jmiy t. WM.
m./iEyy, nicROhAS, m.d., Ph.D.,
^L^ Surgical Bacteriology. !f«w (icoonil) nlUlon. In on* batubmia ocUn'
^HMBpagw, "iili i:i lilHUv, iif niilck IOiir«a>lor«d,snd D eo^nflng^ Cloili,93. JuCrWy.
> „_.i .. __.._........ .__j,^„j ^111^ . ji^i |j„ 1,1^ |((»ratBr» ef a ■nlt]r«, lh<i- •••Ina
y trUliiic aiiKh I'mr. W* fardlalhr aomnitDd ihii )onk <"
aJi |ih7Mclaaii<la*Irauj4CliaM4Df fa» vli/^ mn*!
«D IwrHllatlODs — QonaiMli ImiccMliMt, Hn
.11. IMl.
4
A Mty ihoniiRh and ■il>i>u>i{n t*Tl*«n( tnr-
TVbl iHrrviin (4 that varir.t bu-irrlnWy trUlinc
lo tllnnrf' flaoh Mok- »■ llili an nr IDcaleula-
Ma bBBatl W Iha ^aurat iirullllunvr. n> thmy
■Vtog b««K» bim Id aiunilaiutJaiidDiJiiilyiniia-
I
GREEN, T, nEKRTfM. D.,
£«hrn on nil Aui>vv t-*! JfKTbid ^naitoMf at aaiiive»«a ■d«pUa( Jfidtcal fttaal, £ondaa.
Pathology and Uorbid Anatomy. Xn (aixth) American from iha acTsnth
r**Ueil_EngUili cdiiinn. Ocuvo, nS9 nnv""!) 107 tAjtrant^ Clolb, fS.T&
I
I
Th* PaUiclon tnl M>>r'><il Anatnmv
[*4lrva« Iptao nil liauw= hj mfnibvr* of th* in*ill-
4Blt«a<>Ml<>Dtnii*r'l anf r<>mm'oiUili-ia. Tli>n
laaMivalT an UHalllnm phjiialwi iaymhnr itlm
ha* sot IM irarti to hl> ItWaiy, br II 1> aimo-i an
Mmilal, In tv\ II 1* kMMr iMapu»l in thr ni-nu
of iI»n*ra'stMilt1pii«nlhan>nT wnrK "fi halt lad
«iiri whii?6 «B ara a/'qoalDWd. tiis work* of
Oarmau anilien opos palbulugi', shlrii hiia ba>a
iranKltlcI IDU English, an loo atatruaa for tti*
physician. Dr. Cr*«(t'a vork nrHla^lj mafia LU
■IrtiM. Tha eau ■xJilhli ilia anpranucv* of
|i*ilialq[1o*l Mruelona )u(t aa inrr ara aMa
throush Uia mknaeopa, Tha ftfi u»l It !•
(•B>rKTi}r smplojivd ■- a taithriHc ^ mid iial •lu
'idrnca thai *# kAt* aot ff
UanauM Ittdimt ,VfiH,Onk
dMiU li an
murh In It* hror,
iimL
' bOt ff^tk'Q Ida
PA YNE, JOSEPH J-"., M. />., f. iJ, C. P.,
^nii<7r riHUi'inf /'.tyiinnn oiul Litiff ua nihali-jtca! Analonu. SC. Thomas JIMpHiC tmdkib
A Manual or Ooneral Pathology. I><~ii;nol w >ii liitrt'ilm-il'm lo ihp Pnc-
tiM of Mi'4idi)r. (U'lnVDtif 52 1 |>iigT\ Killi 1&2 illun. nud ii mlorrct pinlc. Ckilli, (3.6l).
Krkn«jr]^, b« a (la^hff and tiaminor. th« tiaol ' Ml fjfcri^r* lb (}>'■•* dIVAua* now *l1h reaanna^ta
rvflalnl)' aK'r|1i«4 10 pMllOlieDall? nlPmbH. lo
1^14 ii0|partmriii h«.haah«*ATKy l^iMaadaaplK*!*,
Dni no If Id a datnlpHM inaaoaT, bal In llMMi'h-
alqua at InieallRaUM. Tha iaoandll, Bl*lna
matiibti at t«MW«b,l*aloB« wcirth ina ivM of IM
book, aaTara! Ilfnaa ovar, to aivrT atudaoi of
pWhaiav^-JLIa«uW«taMlAirg.^r„Jts.'ML
Bfr'Li <if miTill4.>aJ hiuiivuL*, Lh« AfjLhur Jia> In 1h*
W'lrl* tifikrv If iiFeidFKd for ITiPlf v>^'-k«l il-i>
vhat wv doEioi rkpp]Ea^* Cj^ -Ar Sih« ho4i Inlrttduiv
tt«D lo pHTaT [.iilTiriloiEy Ihal wi hava r*l '■'
amiDfid. A dnjiAfiuFp «f hii-h vat airihor lia*
taken U Iha i^rraLar aliablton |4Jd U> ih* rwinfr
UOD Of dlaaaaf. aad iBor* aapaalallr lo Uia vtlnlngi'
COATS, JOSEPH, M. />., r. r. p. s.,
FaUolugui lo IM Obusoit WuIt^ JnXxiui'y.
A Treatia* on Pathology. In nno nn luiulanRw ocuto rolam* of 839 puw,
^Ui S» bMmlfoUlliulrMUiiWL in.>Ili,S.l.iO; l«al)i«r, (6.00.
M«U*al (tudaota aa ir*ll m* phTxIciann. who i manoat. Iha ehaniM tram ft naimal eoDdlUOB
i»«lf« aiaork fnr iludr or raffrfoca, ihti ir<«u I effaoiad In tiraeiunt by diaaaaa. aad polnia oiil
llta aoUacla In tha lutou* daparlmani* In a trry ' (ha rharvMrli-Uoa of ntrlana morbid anoela^
Iboroa^b manoar. hiir "iiiiaui ptviitHy^ wLI] <>•?. wi ihai th*r t*a ha aoally ratwfniaad- BOK oat
lalnlT |lfa IM- onr il» (■[•(•rin'v u> an^ <rlii> Umli^il i^ morhld aiatuwf, II aiplalni iHltf hna
which «a ar* atqnaloMd. II a»u (onb iha muat Iha funciMuaef arcaaiaradlMnrkad MaboDtaul
NMDl dMoTtrlw, eiblblu, In aa loMnilioK : eondliluoi— Oncinnan J/atiml .VHn,do(. Un.
tscnlFF.n, EDWAJin a., f. r. s.
Tha EBSentlala of Ulatology.
91 illiuusLioiM. Ctoib, S2.2&
In ana octaTO toIium of Hi |ias«k, wlili
XLBTK, E., M. D., P. It, 5.,
Jnml Uiwur 9% Owol JwK. a>l n««. H lA* Mad. AtM< ^T SL 5arfh>la>M>'( 0*ap
Elemenia of Hlatologr. Founb «iUUan, In ca« Itgao, rulunc o( 3T« warn,
with IB1 illoi. Ump cloth, »i!t5. flto A^MI/ 6Mn ^ JfMa^ pat(« SI.
I
I
vnnniiKAD^ rntcncAL r«rnot,oor. ai
HftftBal fur 8uid(«(4 m4 PrartlUonan. la «na
hMMIlal «aU«* Tidania of W paa*V *■<■> U*
PRprni-s SmtOIOAL PATHOI.iMV. la oa«
tat(ii»M«ia IMM Triooa of Ml p^M, vuh (I
llliMWMlMa. LMpcioih, radad||M,tt.«a Sm
1
14
LlA BaOTHEKS k Co.'b PcBLroATion
ofMeft.
iVi/. >r '** JVihMpIh an« :Pr«K><t cf ITtA oM </ CMl JVtA xi JIM.'<n< JToipNitl HadiMI 0>U«^ .<. f,
A Tro«tiw on the Prinol^dM umI Pra«Uoe of Medlolne. Itatsanl to
tiw uwof HludMU ami Prailiti<'n*Tit ol MmIIHii*. tint ifiith) p<lilii«i, IbunnMhl; !•■
vlaed Mid Kwrlunli; iKp Author, ii»iit«l hj Vi'ri.u*M II. Wcij.ii. M. ]>.. Vrtimmed
Pultolcigf, JohiM Ho|tkina tlnirrmiv, Balliiuorc. und ALvniti Fuiwt, Jk., U.IX,LI. U,
ProfMnrof PI)]rsiolo|n, BcIWvim' U<»)-1uI Mwlicsl I>i1I«e«, N. Y. Inona vHjr I
ocUvo volume of UriopagBs with illuBlrntkiua. Cloth, ^SU; t«iber, 90,60.
Ko Uittuok oa tk* ^ndplH utd |>nHlo» of < In «l(jr, Insn, illlics.
ma^lelii* iiH ■*•( Oial liiUili rsUBlry mlUi Mi^ TUtit't Ftm**/*. IT*
UOD*n u ilic vork «( PrafMwr Fllal, In all Ih*
fn*4lc*J e',nrtff uf M« VnllHl 8IM« II Ik Ux fit-
nnIM 'ark uuin PnoUt*; ukd, an n ka» fUtod
b*t>t» la •IIU'IIDC 10 It, lb*n ■■ BO oUin in*4lfal
wcffe (hniMn I* » MMnlljIniod in tha lu»nr
ita af phi*ki*i>ii In onri ttt»t aad UrrlMnr
of IM* •■«««> turr tha iMk Itel vmin »ai« tikdy
«ob*la>nil lBUHolfir*«(a»*ifkMBi«kirhMh«r
or •* •om* mrmt tnnii.*
nuk* UtU iiMfiMi m *
MrBtiitonit4* • ■ laal ftw rarwwal nkaviMMuM
it b ihB tMUnomy •■•» nr oUian. Ab i^lib
UiMi iit»m Uu( *i>>7 OMiMilBnWa ihakfii tan
bMD ni«di>UiUi»«liilt •dlUnn. Tli* VMk MV<>^
dovModl* bv f«|tBnl»4 H fklilv r*|>raHrBU^ Ite
prH«nl*lal* aflb* kIodm uT ■•hIiMm.mI m
ri»mlag (ha iWw* ol uma Kho Miwpllty w
■halj ioMIm Uw prwinl nuaa at wMi— «( Ka4'
ttnl wt.-.«H*iMM jr**ml J? MM, bfC UN.
BRI8TOWE, JOHN SYKIi, M. /)., Z-i. J>., JT. It, ft.,
gmur Pl^niuiii la and titlvtr m Xe'<i--\t ir SI. rx*wH' Uuf^lo^ lonJiM.
A TtmUm on the Bolonoe and PraoUoo of Uedloine. Seventh ed»-
lion. ln«M]ugaiicUfOToludMariaae|Mgc& Ckith,$e.SO;h*tbM-,$TJQ. J^rmlt,
■nnrxTirkiMtnnlwtlir wllhwhlah M«a4l-
tlant "f i-'>1i n»k* Uotr ipponinM l>
Milkln . la 11* •intlUdo* and •■l<i«.
TMi, 1' rif tho Diinwran* rli)* for ih"
b<r«r i>( UiD ■Luilsut nUck hu* hMO pal KirlK
■tMln ll>B tltWnn VMM ilDM BrWoiN^*'H*dl'
tfiie" nr>t iipi>nr*4. Kor cu II t>* MI4 U>M u»
■aUtst lilBiH-U hH IWII*d
10 kt«p M<
■>>iMl of wltnnclait know'nlK udaoui la Ikal
laik muM rniT*, Ho )nn| tn iharo I* *ha«n MMh
•nra t»t *LrviiNHpa«Uott (n t)i* tnctOKinn ot all
navBatlar IhMlHwiacAd ll)" iwi nf prliMiai, •»
tama wtll Ml* iMrtI i*lala Ilia taior trhloh It hat
■hiat* mat Pnr il l» ■ wart ilkal la Inlltcvft
MabTa IbuaOuiun, •v>|M^all^ •etttltf and pnc-
Meal, oaMxInlnc lh« malcrad ««p»H>»t< of k
iihytlilBa dho haa Fvwr eWm lotwi
an lAlbDrti;, and KipnaaJ la ani^la vhaui »
imPU 111* [Viir<lilimartaiMH<hakihaMiliMii. k«
Mia ms «; iliai iMabeok t>u <.t<i*irL..i iM-aaia
whklimaandsai>t»d,*»d •■ iiaaaAif
■III lone wMlana i" auf >■ mrtmnf
tn»b tillUa** lor Uia ailt»: ' * aBnina
faaaiallaa vt tnadlnd alUdaDUs.— TM AtaiW,jrf
TsLuM
Dr. BrttU"*'- "jw riin'fli if^itiu. af^Mn M
III Hraniii ' iHaiia^
In irhtah i ' in t*-
mandalkn - . i id Bvir
•a XBiiafc Wie iihiI <><>iiii.<in iiu.t >.»r'ilrfinb
tacikiL-9i«M JtMMat Jbiiniat,Sa»<Mntw R. ■«
OAMTSHORIfK, It EN BY, M. !>.» Z£. />.,
tttlttf pratrntar tf.Hva"^ >* t^ C«*Mraii» »f ftaiqli aiiii,
BaMoHalt of the Principles and Praotloe of Modioine. A RmAo*
for HtiMaiM anJ PnclitioBan. Fillh »litit<n, DiorcaiKhlj r«TlM>l NmI t«writt(n. In <••
n>r>l I'teo. vo)llID«<>r««»t««:«^«lth HlilliMinitkiM. aMb,nLTA; hi>inKaiD<l,|lJli
niinin Uw oranpaM •< aM MC*« tl IraMa «(Uu
y af andWlBt, »ovnl l«il
iunaiAlo|[y,and|>riir*
wiOO|>rt, o|4ilJiartno*>
aamiiM^Knaoloir.aadaMCWratiwiapjtni
ifw. rhara ■• a irnadmni amonat <draninnailoa
EmHunaiAlo|[y,andpliir«te*l dlacnealannaBdlaw
7afeaooi«, oiitilliarnwec^, atc\gananl IkM^
•«ralala<dln Ihla mvk.aM M b on* D( lb* bail
t4 Na kted UiM *e lian Maa^Qlam* J(a«M
JbvM'. Nil. iiM.
An liull>|»>n«Uilat>iflk> Ko*ork**«r*iMhM*d
a bpllar aiTnii^r of a<t<iaj praoUaa) Inalmeni Ikaa
ihia aaa: *»d prukaWy noi on* srlio la air 4*
hail a hollar <Mp«t1llli«r Ihu IH. Ha '
«andsn*liM •" 'Ha ilava of »'i . r. ■. .. < .
Intaa inti'i. 7h* nutnoMU-
ran Oaaflll In •(odoBl* *f
UaU, •■ l>i* Baa»« Bvinab, i
■opanxls lb* Mil-l>«nk>of niM mm
tM ilioy *f« Iba iDivl raluaMa la
innuii Ki a*« Ma alaikr* Uia atHila IllantiiL .
JI«*H>.an-llhaa***i<*laiMainalMH>BL— I
3la4int .fMrwtoad giaii^r. April. IHL
MMrtMW
■nliii^lll
wbl Clip
BETNOLDS, J. BUSSELL, M, D.,
iVa/iaHri^IM AvinplHiMd^-arlmaf IMiriMik tl*MmM<t OB«f«, t'iiiAi>
A System of Uedlolne. With nut^t and a>Ulilocu hj IIkxht HAsranonA
A. M., M. b.,liite Crofnaoro^HnictieiiiUM UnivarMtr uf PeDn^lvuiK. In lliiv* Hi*
«acl hMiibeimc ocUvo voliunei^ ocauiidtig 90H (loobl»«nliimMd p«niL wlUi Xti Uhaui-
UoiuL Price par Tatunw. clotli, 19.00; sliMU, VAXM): nn hMiibolM halT KlMalJ^ niiol huA^
«e.M. Per M, cMli, |1A.00; leather. fl9.oa AU »>^ ty nioripMn.
STILhiC, ALFRED, M, -»., XX» D.,
rv,/ubi' JSH»ii<ur>f Ihi n»ryand rVwiiAai;' Jrnl.<i.Wfl< Ci-aetl MM. It I** tfutr. 1^
Choiora: lunrtgin, H(Mur]r,(^aiuatiim,B7iU|itiMU,LedMi^ Pwi-entiuo and Tran
nent. In ono hanlaniiie 12niD, volune of 163 |Mga^wtth«dlMt. CloUi.fl.SSi.
WATSOX, SIB THOMA8* M. Z>.,
ia(* i^paiNaa In OiMtaty lo llw Qutm.
Xeoturaa on the Friaoiplea and Fraotioe of Phnio. A tmm AiMriM
fnia lli« fifth EiikVmVi «Auiw. ¥Aiur&,'«\.vi&«>Uii.i<Hw, a».l IW illiainukiMt hj Hsnt
H*Kn.iioR!«it, A.M„M.tl.,\iW VTOh#«»«<*^l»amt\ft WcV:^'(>\iTnh<.'iQCI%taqrlnaifc
rSsornns k Oo.'» I'vauoAi
Flar aah bg 8%^»eripiioH Onty.
B A System of Practical Medicine.
^B BY A UERtCA N A UTUORS.
H Edited BY WILLIAM PEPPER, M. D., LL.D.,
^V piioTo«r Airn
piiOTo«r Airn rHoniRMR or thx msORY amd rRAcnci or mhiicivz axd or
c>.rHicAi. MUiiraiiB IK THB osiivKiuuTv or rB»nTt.vA)iiA,
A«lueil bj Loots tta ASS, M. P., Clinical rnr«nr of the DImmm of Cblldiw ill tbt
HMjiiul uf lb* UnivsnUf of PtiumjlnalM,
Tht eamfUK mrk, in /at columa. emiaininy 5673 pnyE^ vilk 19S iV^MtrnttMU, ii mw m^y.
I Mm ;Mr rolumt, dolA, 9^^; Uaiittr, $6 ; Ao^ Kama, rattd bamlt and span iod, 97.
In ll.is erenl work Aincrian tneiliciiw n fur (ha fim time r«ll«ct«d hj iU worthiMt
tencbcrt, Diul prnvnlcil in the Cull ilcvalopmmt of Um ptmOieal tuilil* which b ila pn-
feiKirut cIiniMcifirittici. Tliu lucntl aiil« iiitn — fruin IM BmI mwI us WmI, tttm tba
Koith ■nil the Suiitli. rriim &U the iironiinMit cvoln* of cduCBliotv aiul fann all tho
hMfiiUJa wliich nflbra ipecud opponunilia for aludr sikI prvotioe— hare mtilal to
fMWKU* riralrj in bring Mc«th«r lliM Tut wgranM of •MCdMuod «xi«rin]c«t.
Tbi dUiiiipiioiiMl vdllor hu ao ^ipoitunMo th« ynuk that lo mcIi nutlinr hn* bMs
■aiODcd Ui« Bulyeci wlijcb he ia jKHTnlinrl; 6tl«I to ducUM^ >iiil in wbkh hit tiem
wUrtw >ccc|ilod u the lattst rijirtwiun of KicntiUc and pnctioal kDOwlcdtc- The
practltioaar will lli«i«fur« Gad tlioic vulniuai ■ nim|>let«, mllioritUl** and un&lTlng woi)c
of nftnnee, lowlildi li* nii; at all timea bini with full L'^ntiiitr of tadliK what be neali
!■ Its BKat rcocnt aitieet, whethsr ha aaoka Inronuniion on the gmenl pnodplMor uadl-
cln«, or uiiniil« jtiilflnnra in ih* irMtment of «ii«dal dbawM. Bo wid« U lli« kii\» t>( tho
work thai, villi the vxcvplian uf uxuiv'dtij aod maUfin aUicllr kuigical, it uubrMW the
wbotaikiniLiinnf nivdirin', indwling lliadapartnicnia (or which (h* pli/aiciaii b aooulnnicd
lo rtljr uB q>MJ>l trNiiii>««s (ucli ai diiewaa Ol womfs asd chlldi«n, of tba gaiilta-iitlnary
ocnua^of ihvakitifOf ihenerrM^hyginw and MiubuT acienca^ and awtical ophihAliiwlop
■M Mologj. Moreover, wlhora nava itunUd Ui« rantuilaa which tha; haro (aattA man
«fid«nt in th« trwtmMit of il>« nrlotu aAMlona. It mar Ikua be iral; rcganM a* a
CoarLXTx LtSKABY or Pkaohoal MxDinxE, and the general [inililiniiiii |iiiMiMliijt It
■Bj fael aacaro that ba wUI r«|«iira liltla aim in Iho daily touad of tuofaaioDal dullaa.
Id apila of CTerj aSbrt lo eondMioe thv rtutt amount of |iractical informittan Utr-
niabod, il liaa Wn iropoNible lo pnaeot it io lea thnn 6 JArga ocU*o vulunia^ anUaiaing
aboal WOO btautiAilty printad yugat, and rmbodTUig Iba mailer of about 16 ofdiaaiy
octtToa. Iltustraiiona ara lalraltiotil wh«n9T«v re(|tiMil« to alucliUla tlia taxi.
A dOaiird proipfelmi nil be tent lo any adJrat an appliealiim to iht paUitktn.
phjvliitaDi nliu If* acaualaMvl lalili all Uw Twta-
IIh ot ollmaui tn (ha Dntwd Su<*^ ib« cliararMa
at ta* iMI, tb« maanvni and auMom* ot in* 1*0-
fla, fto, 11 11 iieculIailT «ilayl*<l to Uic waala
or AnMrteas i^rsctlUoowa ot mfillolB*. and It
jaavf la ua Ihai «f«f)r odd uf ihdni (tuuM d**tra
l*h«Ta IL ll tia* li»n uulr rallc'l ■ •■ fntttft*!*
Lnmrr at ITapUoal Hedlolna* aa-l Ilia (iiaaral
■neUUODar vtll mioir* lliUe alfa U kl* rouad
fiarrh.UH.
Barh of ih« tnlumM U pmfdHl «tth a Beat
ooplflui lBd*i. ud laa w«rk alutathv r f««ml(M
lo twoaa ■hmti WW add moeb lo Ui* madldal
Innrauin at tha praoMl oaalurr, and nSoO g»M
mdll BpoD Uia Mholanlilp and t>rv(lnal arunaa
ofll* aulaofa.— nf {«J« taaM, 0<4. 1, tnt.
Til* lHlle«Dr proud aalbbMloo iriiii wbieb tha
Amarlrui pntBHlOB taaa UiM, Ila twatanMUTa
■rtUm «f pr*ftU«*l mailliilBe liauad io Iha Medi-
cal awl^b ItiM JuallDMl tir Ui* rkuatw ot Uia
wak. TTie aollra cwl« ot Ih* ajPMMk I* In kaap-
laa with lb* bMt ihonalMa at itaa leatora and M-
lowan o( our bona aohoed ot marta*. and Uif
eeiMnaUoa «d ifca aolwiHa awdr oTi
Thai* IwA tolunixB brlUR Hi!* admlraLlo wAfk
M a Pin*, and (iiJIy aiuiun ilio IiIkIi nUindard
tvMlwd Of Iha aarUM toloinri: m l»<o oaly
tharctora f« ociia the anlc^um f^roDoubcaJ ui<iu
Iban. W* would ■■rmly TuDnrMuIaM Uiaadllor
aad hia obllaibomjDrv *i \>i,p mnaluBloD ot (hair
lalMinon* latk on iha a'tmiimMn mannarln wbloh,
(tMa kmlo la*l,lh*]r liai* iwrformed Itialr (aTonl
AoUaa. Th«y h»*a (uawadeit In urDdueloc a
wark wliloh Mil] Iook ramala a ttwidatd work of
lattraoe*, to ■hioh pntoillloaan alU lot^ tot
MdnM, aad auihof* will laaan Itar Iwiu.
rroM a tUaniT polot «4 Tlaa, tha WW* I* alUioat
an* MriMU bmnlah, aad In nw|>aM ot produMlan,
llhMlWhMuUfUl flnlab ihu Ani«ni«B> *!■■>>
■Ita Uialr aorka^— JWIiln'g^ Jtciiral JnrwW, Jan.
fi?.'
• • Tha ttmusi dliUnetlToly Amorleu work on
Iha prw-im rr| mnltoln*, tni, la<laad, Iha aopar-
IhItv ndji-oilx vnuld Dot ba Inai-pniprlala Kara
avaa all iithar |>ro4iiPlloDa phKad la cunipUtHm.
aa aiamliiail'io ut lli* Ata Tolnina* >• ■uflloleol
to aoBtlucB uod of Ui* nunltu4a of Itio anta^
pUri aod ot Ui* (iiixvaa wEliih ha* auaaAid Ila
Iniainianl.— TW Jtidwal ^t", Julf 31, UK
ThlB huaa nilafna formi a Aulnsolo** lo the
MMiraUm ot madlolD* whtah In aa ■borl a Dma
baiKinao hlabaplatalo madloal UUtataia, and
hta 4«B* aaen nadll u> the nntaailaD la ihli
aountir. AnionR tha laantv-lhre* eaniithulon
at* Iha nani» ot Iha laadlaa oaureloglM la
itmarloa, and mcHlof ihavotk fn ih* rolama li af
Iha blghi-t ordar.— JlHloa JUadHnt and JIHMal
Axoin', Juir 11, lan,
Va oAoaldat II osa of Iha craadail worka ta
Pnciteal Madlolo* In Iha KngOah laacuaaa. til*
a aark cf ahira tha proJ>Hlon of IMacooiltrT can
Mat prnuA W/fMaa alcfivltalj bf Aa»rMan
lb* praMleal •ppllMaloa 0( aaaM and oaparlaMa-
U) kooDladKa h> Iha Ifaalnaoi of banao mat-
a.li<u. R.akai ovary on* of na shata [a tha prtde
itiai ^a• aalooiBOd Pr. raaaor'a lahoia. Bhiatid
ct Iha protutir thai waaitoa Ih* raadaaa of iha
llannaa aahoaL IM anMaa alaaa <mb* <
aaldafcr*lllhill*Tataafal». Ill
of aroaalBaa brain', anil la
_ ._. aain*
la tha ouitonia
_, . . oualiod l&nashoul
Mr oiueb *4 Of alurdr Indaaaadane* ot Uiouahl
and orlalnaUIr thai l> a nallaaal tkaraalarMla.
Y*« aovhara la lb«r* laefe ot MUd* af Iha raoM
■dtanoadrtmafthadat.-Xo'W.OvoUMHiA.
10
hMA Bnornsas A Co.*s PcBLRunom— CttnicKl M«d.,«ta.
VOTHERCILL, J. M., M. D., Edltt., If- R. C. !>., Xond.,
The Pr«ctitioner's Handbook of TreatmQnt; Or, Tbt IVindpIo orTh«r»>
peuUci. Xtr<r|llilnl)tilitiofi. Id mic t^to. vi>l. of (141 ]iiib«il CUiUi, {.S-TA^ lcaUi«r,Kl)L
m* it k iwj.flul tMtli. ir IDMvbatBM*
thl>>( •• "m*dIalB* oiadii 'ur," ihia u Uw •«% W
MclcwpltabthtorMiin.— I'a. Na<. Jhaia, JnM.'K.
■n(t>tnui|tM*M(Mfly ■iiirmml. uhAiImUh
MuiUdI iM *««Da ifnvililbuar. |i»>lui^ ana it
T* b»t* • 4«Mrlf4lon of Ihs nonnil plinMttsl-
«>l pTtefM of wi oritB ut4 i^ ilr* maihedii of
IrMUnaDl of U* morbl-l fan>IIL*<M hnii|lit
liwMliar la ■ idnitl* etttittt, ao<l ih* nniUM
b*i*MiiihBliMvl«(tyMM>4,nt<>aiHnil1 M|ire>a
■ iiaii •■>a**DMnM to n>M> UHwhtAil bui bHy
pCyilclaoi. ThalirKiWal nJsaof Uia (nluBWb
Bml^ IncMuad hj ilia latnJuttMi or maaf
prMVriHIon*- TXU in* proftoloa a|4«*cla)a*
Ihat lh« aalkiT ku ■DH*rUk*ii aa Importanl watfc
Bjil hM ueonMHhHl U ■• ihoon iw iho diioiand
r IbU IhM adlUM^ ff. r. Mtri. /Mr, Jane 1 l,-n.
. _- ,-- "•'. ■■aihaa* i _ ..
W* do nul tno* a niura raMahl*^ fiteltimi «M
iiHflil wutk (• ika ifBaimvDi of diiaaan UMa ilw
on* «• han an* b(4lu* iit.— /^KtM JMuaJ .M
Airpiial JlHmitf . 0«utor, Itsl.
VAXJGUAy, VICTOR C.,Ph,JK M.D.,
and XOn\ FREDEHICK G.. M. D.
Ftomalnes and ZiOucomataeB, or Putre&otlve and Phyaioloctcal
Alkllloids. Kcsnliiioa. In ono bindwaia l^nu. voJunwoflOOiiagM. Ckith,R.3S.
Andy tAow/jt.
FINLA YSOX, JAJIES, AT. th, EdUor,
/*«)■•«>" a-^ f.wfiir*' M dwHUiil JMlniM ia (M CthUftw n'«<(rn Stflrmmrt, lU.
' Clinical Hauuol for the Stady of Hedioal Casob. Witli Cha{4ta
bT Prot tiftinlner on lll« nijviogiwiiiijr of UfaWP; VraS. Slcptictiaoii on IMmmh itf
tha FcoiBla Otigui*; Dr. Rnbartcoci on InMmltjr; Dr. Ucniaelf oo Phralcal UafaMb;
Dr. CmU oa LtfrnfMcci^ md Pua(-UonMn Eiuumitkouv and b* tha Editor OH Cm*'
uUdg, Fuullj WMrj tiA Qtm^/Unatot DiaorJer in iha Varjoua SfManw. N«« editiciK
In oiw ISma. vdume ef 682 paffcs «3tfa I&3 illmtnUioa*. Cloth, 9150.
Tba infiHatbb tannol t-nt nlMioia IAaMO»a4
•dUlan ef iliJo nrf laluaU* work o( Plnla^MW
Kl Mr oatUbaraton. Tha *l*a vt Iha bnnh Iih
M IncwH* BOil (ha nuBhit of IliuiinUmi
Marly dooblad. ThaiDaDaarla«liichlIia*ub}*ci
U Uaalad !■ a fn<M pnciictl cna. SymfiKaai
alona and ihatr fHanoailo iDdintiUm* r»rin tfia
baaliot dItOBUWn. Th«lailait<l*iD«rlMtl]paad
fDllT <Im n><lh«li of aianlnaUou aad tha Brno-
clo«l<M M ba drara ftarn Uw pbj afcal ttKaa.—
n* Xaftcnl XOHL April ti. Mm.
Wa ai* filaaaad la t»* aaacood adMloo or llita
adnlnMa M«k. II ■■ aaaantbuly a pravUwl
InMUaoa awdtal dlwaaaU, In vMAannttii
ud tfaMMB o( dlMsiI U «wMI> BkUnad, Ml
Uialr t»1all«« alcntlWuM la Um diawaM tfa^
tlDuilavbleiiUM'raoawpotatadaM. PMniMr
■jrMhaala IM ainJtM oM aoMmMv dMMWM
iriadlHHMirltlKabifbtiabaatodaaK TWbat
haa ba eoBiiniHnr, nni In It Itkaly hi ban la Mai
H hMura aalUaBt oialnuin ita fqasaal itaiidild m
ataailaacia. Tka (aoaral piaetUtonar vUl iM
■Mny pracdeal bIMa la U> r*cH. vbna a Mnkl
Mudvaf ibamik laUl ht* bin (rooi nuay ••■■
la dlacnoaU^taMrvaal Maiim-CttntmUti JM^
M^ JauoatT, UK.
BROAl>BEirT, W. H., Jf. D., F. R. C. P.,
nyilnan la aad Utiynr » ATaAr Im nt SI. JVarp'i fftaiHMf, ivadm,
Th« PulM. In cm* I&dq. toIuum Di 812 jafM. Clotli, 11.73.
teal JfdmKiti^ |Mg«8I.
Tbia Ititia baob pn>««bly repraaiBU Ibe b«a(
Celkal UiaBRht au ltil> uitil*cl lu Iba Eailiab
I '"■
SttSmimafam-
aiuiai. A wRMt InUrpraiallan <i((ha tida*L
ttllli lia altnoat InKnila niadin«Mlni]ii. tirciisbl
aMHit by tiotwl uaHmlWil bodily Ta(taik>B>. tan
oolyW adilarad by*>parlaa«a, ai>4,
Uwatd »uai>iia<Uil*B«alL •ouilni <■
•anloa Umb l£lt broMtuN nn Iha
puiah— n< J
Haptamhai. UM.
■■?«, ^ ■■ wm
•UlilniKlIf haaTiM**
aluM aTttl
laliM*^
BABEUSBOX, 8. O-t M. J>.,
a—or fli>am— «» aad tola ft. — Mtfrim <■< f»««lfc« «f Ifad. 1 e^« BfpUat, Im^-k
On tho Diaesfles of the Abdomen; CtonpTiabiK tbote i^f ib« Stonudi, mi
oUirrMrtiioribeAluuentar7C(aAl,ai>0]ibwM,0KoaB,lDl«aU(ie*UHireritoa«unL !)KMd
Ani**leaD fmn tbtnl cnlamd And rcvlaM Eu4U> wlitMD. In cna hAodsoaw ocMR
Toliim«orftMFac«i,iTilhllliuinlIoD«. C]otb,CLSO.
Tbia TllMMa IrcatlH nn •llHwei of Iba alonacb
a*d aUiMDaB Will ba li>ua-l a i-yclopiailU tit Infar-
WaWn. ayilamlleaWy ajiaiiga-J. <ni ulliliHamar
Iba alunaMair InM, frum ifta maolb U Uia
iwwm. A t*ii pNforitM <4 aneh thaplar it
4atal*d(oay(n>iau»,paibote«y,a«i4lhen|>tuU<».
Tba i<nnn[ *dlil«a la filler Uiaa liirinar onaa U
!• Iha limat, ■Dd nabl»i II ■ minns at inlaraN tl
ry a*M t4 madMiaa Ml
Alt
U>* prw^li' ' r
•iircarTi
aiasFlaiHl
■imutt,
U In
ha*a ^
aiMiy panlcuIaM.aBdbuhaanlbacDiic^
Ud amaadad by Iba aititiof, tlatrii.
(ara ba«a bara addad. biinRinf iha * <
laad '
Af«i,un.
<uf l.-t-.K-h — -V. i, ijffit..
TAXXER, TTIO^fJS UAHKKH, ^t. J).
AHanualof CUmcutlUcdicmo und PbjBicnl Diagnosis. Third AsMriau
train Iha Mocaid Luadra Mlitiua. Rvrianl .irnl fnlartMl ^j Tii.xtiKr Fox, M. IX
I[iuiMMMlllSiiw.TOlunMa(S62pgttt,<'i>^ ilJiii(tnili->ni. Clodi, ll.Sa
LIUnrREfl O.*) THR STVDT OP PKVKR. By r LA Ii
A. KcMH, M. IX. M. R. I. A. !■ on* Ocuio .^ :
nituma of 9* tap*- c««ti,iuo. I _„„
A TREATISE OSrtVSK *j &»««. O. \.W\ Tlw™»
. RLI.OW rCVBR, «iau.<af»4li
. , I'Mbatac*"*'. Blli'^'Ct'a] aat
^BaWlMi. laiwel. t-
ftuOM^P^-Rr^eiie, Electr., Piact.
BAMTHOLOW, ROBERTS, A. JU., M. />., LL, £>.,
/V>f. 0/ JMana JfaAca and Ovwrol TXnpnina w M> >(ri>^>tn ifai. CbIL 1/ miK,
H»dioBl BleotrlcitT. A Practkal Ti
10 MoliiriiM nod Hunctj. N~*w (thlnl) tdilloa.
SOS pWM, Willi 110 illiiBiniiDiu. C'loili, tlfiO.
on tho Ai>plicaticqi» of EUvuldt;
In CMM nrj bmdMHue ocuvo rolune of
TiM kM Ibti ihl* <N>rk hH muilieil <i< iMnI xll
tlOd ID •li r**"! hikI UiM tl liv b«a kcut hillr
mtn«M aUh ih> incrvMing uM ud kduwInditB a(
•l*MrVtiy.di><niir»lnM« iHolklm Mb* ntntldoiwl
• HWtlMl inAilH «f ttla4 Mill V l« 111* pnifewloo.
To* mUMT •dil*d talbe iimcntttUllMicnibMCH
UiB mtitl r*o*Dihl*h^o«t lu c4*ulrL4?h] infttnttmi,
Tha iUuitniloni w* ■liim-Uul *a-l fivar. uxl ilia
■«k oaniilliilu ktull. a1a*r ui-1 courut inaniuil
■•ll ailiiiiiMl to Ui* DHiU <4 bnlh t«ailvD« uhl
pnmuloomt^Thi JVafMsl A«n, Mar II, IMT.
Y£0. I. BVBSEY, M. D., F, R. C P.,
Frtfutcr if Otnttal Tlk'n^aM) I" A'»i/i OWi>)PS ^wlen, ai^d /^yaMwi M JITof't Qh'^tm
ffowilal,
Food in Health kod Diseaav- In on* ISma mlunia of AM pag**. CLmli, 13.
Ste ^r>M 0/ (.Ii'niiiuf Maxtiai*, page 31.
LtTfo •nriillr* In ii-;>>mpiw<lanns*wl)rklllhM
Uis (.rvililonvrnquliMio knavou ih* hii)*cI i-r
dill Tlia >ork U diiidwl inia In |«iu— (uaJ In
tiMlili and (dad In du>»a. Dr. T«a tin* K»l)i*r*d
■VBtlhT (mm ml inartf r* an Inmaaw amuunt of
lutAil tatBTinailon nkicin a comiiarMliilj unaU I JnwiuU, Fab. 1, tWO.
•rnn^M*, aod ha hu arruiBMl aad ilftalad bl*
niatetlali with (kill fat ihauM tit lb« iinMiUoDar.
W* hA*D Hldom laen a iKwk whivh mora that-
DU«blT nallaaa Ika nldacl br wbUli N n* vrltlaa
(ban thli lltUa work «f Dr. TMu— J>riilM« JIabtal
h
BICIiARDSOA', B. W., M.D., LL. J>., F.R,S.,
Pr«T«ltiTO HcdiOinO. Iii(ni«ocUraroliuncol739 pa^cm. Qulb, M; IcklllM IS.
Dr. KlnhudMn:! bai luoiiipjod In |inKlu«lD|t a
VOrk «blc«hl*al*TaladlBcouE<B(TiLoi],cfijDprBbaa.
■It* In (mp*. lelantlllc lii«haraiit*(,>x<ManaUela
Wnuit*ni*nt, and vblcli li wrKlva loa«laar,mD>
Claaand i(*auiii miuioar. Uanlnoai iba happr
feanlt* oi'iinH'UnglhaplUi of what i> liu<i«u nu
fta aobjvt. mnd n1 iifatlng II In a fDo>i nimpln.
tnuilltviii and (irHdMl lOrm. l'h»r* i> nerhan
DO ttmLlar i<T}fk vfliwn for ih* nafraf piibrio
tbatronlaha* iiu'-'ltaconipIflL*,rp]lablaand Inttnio-
iliaiwllection of data upon Uin •tI»aM> ecmaiaa
li>lb*noa,ibalr9rlclnfeucau»a*.aad tlia in#«»ur«a
tor thalr pr*>*nllaa. Tn* d*i«rlpllnn> nrdtaraaaa
a» alaar, aba*l* and •oholarl;; ilia dlaomsliinaf
itie<)uB>ilaDa(iI Itaua la eoiDBrvheoilTa, maaWrU
and (ully abrnait allh lb* lalaal and bvii knoitt
Mt|>»a iha •iibjnci, and tha prvnnU** inrawina
•rlittcd ar* BiMDraM, tinllFlt and F*UaUa.-.71ta
dmwicait Javmaliiflii JMimJ SeiMiu, April, U»L
TffE I'EAB-BOOK OF TREATMENT FOR 189t.
A Comprehensive nnd Critical Beview Tor Praotitioaora of Uedi-
Oind> In onv Viiw: viiliiiii' iif Is/i |»>km. <.:lotli, tl/i'). Jiulrtailf.
,*a For ijicoinl inmiiiiituuuiiJi iiiili j'enodiisia ec« pafiO 1 aoil S.
Tba ptvirni li-Tii-, iho >«T>niti. tiu U*n In*' tani'yloraeomineHdlinth*f»*aaBH«*ama,i»hleh
eraaaa Jby ffi'>fBlbi*n runir rmu'Irt-l and IVfljpaKVP.
iMst tba <ir]t;iEiaJ |>Uii njf rha book li nol ■iurtil.
It «ltlL tvmaJni n ixik^L*!! fpiioiiid ^t ilio ?iiL«f
arUri*^ t>f lJi» pjMl i-*4r— nrtlH^lB* K^CtkHod and
■plCiimU4d by a iilaff 4'if coDlrLbul4ft OL^cti lA'
eludM. many ur Ltia tvit-knocn and ablrvl apa-
n ooaitdfrr In t>* aspcdor to anf «f lia prtrtaota-
aor*: mora pmstlrd. mora ciB*n>llr UMful ta
iTanilllaDan Ili**a«ilonBl«diMr*haTa«oaJiD*4
thalr aiiaailon lo tulfllllagih*ah|>oto(lha booli^
ailiMOM— Ih* imTliUaa at u •pltaow tt tha
.._..._. _ . , matt pracHoal and uaafOlntlhaiDcdlealarllel**,
OJaJliU, and Ihcw i^aatUiiian liara wlMly kapt ia i vl til* pwl r«ar, Ibr the UM or «■■ ith>»* prv.
misd litat Ui* 1 tar llwk I* lot praoUUeaan^ and , lira doa* not aUaw lima IW U>* aindy <■! a laig*
Uwy tar* «It*o obai asuial praollM raqulraa— nnmbMot hDmaaod CxalBO RiadliW'joDrDala.—
omJuI, w«rGab1e iDformaUoo. W* h«t* DobMl-l n» IfMui Jtifitmt tf MUmA Selmft, Hay, Iwl.
THE YEAR- BOOKS of IRE A TMBNTfor *8€, *8 7 and 'itO
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aCBRETBER, JOSEPH, M. J>.
A. Mftnual of Treatment by Massage and Uothodioal XOBOlA Bz>
•rolM. Tmugliilpl I'v VValtkk Ue»i>klih>h, M. D., of Sen Yufk. In ou handmns
OMBto raliUDc i-r STJ {ages "i*^ '^^ ^ae cugravinifa. Clidli, f^lTS,
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AMDORCAT VBMRit. Thlri AnariCM •dl'
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SMITH OX CONSUMPTlOHillaKartraad Rama-
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LA ROCnX 11.N P.N-EVM09aA. I toL «T«.«f MP
tut**- (.loWitMa
WILMAUS O!) PULMOiCAftT CO!ISVIiPTIO:ii
11* NauiTL Varlallaa and TraaUDaot. Wllkaa
•oatpalaofs** than— « (aaa* 10 *«*«>bIU> Ub
dntUlM. taoQ*tT«.*al.at»in. OaliLMai
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OPCLIMCAL UeUIUlNK. UBlaiaGulda to
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I)*V1.T Cl.IXlCAI, I.XCTI.<ni« 0,1 VARIOUS
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RAHLO«~a KAXVaL OP TIIS PRAOTICK OP
MKDIcidK With aiHIttca* tp a P. Oioi^
M. D. I T«l. Iio., pp. MB. aolh,|UCL
CBAMBEBtl-MAia'ALOP Dm AHKRBOIMRH
TK HBALTK AHti MlrKHUM. In ooa haad
t Mttro nHuBB of m pp. t^ou, ti.n
fnttua' nf Un Pnnapla a»< Prttiu if MtUmt m iMIaw flwpMal Mt^mA CUtft, H. f.
A Uanaal of Atuooltatioa «iid ParauMioo; Of iha Phpleal IMnfi^iil
Di««wc* oftW Luii)D Hod IJnui, anJ of Tbondc AMoriuo. Ncrw (fifth) edkte
Elihcd I7 Juut* C. wllaocL M. D., Lecturer <m Pbjvlcil Diaipx*'" '^i ■'•<> J'lhn*
Mfxllcml ColUn, Plillnilftlplti>. In OM kaMlMme njrml l&ao. ruliiina vt 274 |Mg«i, vU
12 i[lu>ilraili)as. autb.fl.TG.
Tkl* ttUla bMk Uireash tu twImh edition* bu I onf^tam of ppof. niBl^ iDTHllnUoMi. htM*-
~ ' doDO moro u> adfium th* «cImio* of 4Mto n li •x(>*Uhi. Ii* tain* t* oIimi
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proistlr doDO moro to adfium th*
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dM*Tt»U<« upon UiB niUM*, aod no* la lu flnb
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DoMlly «Hh which ftvrlous •dtlWaa vera ««
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«andM »>)• of axfTHalon. J'WUw
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BT TBS BAUB ABTBQB.
A Practical Trsstiae on the Phjsioal Ezploratlon of the ObeM tad
ths DUjpoflu of DiaAuea Afltetiag the Baapirator; Organs. Scamd atd
nrbfd «illi]i}(i. tnofi* luiiuWin*<>oftavo*nIiiiii*of Ml pagva. Clotb, $4JM).
FbUiiiris: It* Horbid Anatomy, EtfologTi Brraptonuttlo Evonu and
OompLioatioDs, Fatalitj and Prosooaia. Trotttxtant and Phyatoal DlMt-
noaia; In ■Mrimor CIIaImI Btudl«a. Tiiom ocUTo«aluiiMof44Spac«& Clotfa, fSA
TrMtmi
Jn oM oouvD
A Practical Trontiso on th» BiaKnoaia, Pathology and
IHseaaea of the Heart. Scoood Mrianiuul mUiskI adidnD.
of 6S0 pagM, wUli a plaU. Clolh. H
BauTS OQ CoDserratJve Kedldne and Kindred Toploa.
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A Praotiool Ovide to Dlseaaee of the Throat and Koee, tnclodlng
AMOolated Ailbotlotu of the Bar. New (thini) and enlMg«d «dition. In oaa
inivrtal ooUitd Taluate af 731 pagw, «iili IN illualnitioiv in oulor, and SSUi anKtariiiy
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n* UilH odlUoBcf Mr. Lraaoi Snnni*'> in-
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oliaoRi tiv beta randwHl d**lr*lil* byttie ad-
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SELLER, CJML, M. !>.,
A HandbixA of Dtagnosla and Treatment of Disoaaea of the ThMai,
note and Kaito-Fharynx. Third edition. Inocia Iwadmne njal ISmo. raJnaa
el UTS |»>lt<>, willi 101 lUiiMnLli<4u and 2 colortil plalw, CUdli, flZSi.
Frw ni#dl«al wrNan nrfBM thi* author bi I «f Mplo* and molhodL Tha laek aaaarvo* ■ IhV
abulir M mak* bM maaaJna parfMilT hImt n a »*la.o*P»qlallTMaonaaiaira< pfOMouio W"
hw »«rd^ and tn dbcrlmtuAoa la MjMUea>,ba<b | «v> VidiMf Jianal a "^ ^ - - '
£srs£
AltfltlM.
COHEN, J. SOLI8, M, I>.,
LtttnivmiM'v^iatatitfmtd tUnn—tf Uu Wril id Otat ■•> tkt Ji/«v*a ttvlHmt CkOwa
Piaeaaea of the Throat and Naaal Paaaagca. A (toiiUioilia [Kapxafanl
Tr«aUncnlnf AffacliaMaflll«Pll■l7»,<fiK>pll•gD*.Tr•a■H^L>ilvnsllml Nana. 1\M
•dllhm, tli<>r<"i|[bl7revlBei1 and r«wrili««, tritli a larKvanoiMroi naw llluttnaJaa. la
OM T«ry buubonia octavo Tolniaa. Prtfiirimy.
0B088, B, B., BT.D., I.L.I>., MJ.C.L. Oxtm., LL.D. CMiMk
A Practic%\ Tre&viM otkTotc\cD.'&<ia.\«AV&.Vb& k^ts-^uaaQse. Ibom
ans ft Cd.'s
BOSS, JAJIES, M.Tt., F.R.C.r% l.L.l>^
A Bftodbook on Diso«*os of the STorvons Syatom. la oiia ocuto
nJuiiie<i( T:&i [oign^with Ittlillutinuiun*. Cloili, HAO; lottW, $<U0.
Til* kocfc bWon UJ ii MiUII«d la Ux hlfliMl
WWiHwiUtai II !• p«lnMi>tIa<, mUdiiSe kad
•■(■•Alkc'y «<>iiipnh*iwtit*. — iT<u Fv* JMant
JlMnaJ, J air ID. IM».
ThamUinr luu ranilarwl > riuI Mrrlo* to ilm
vntamiaa tv eondnnttiiK ■»<> *"•» voliHaa ttiv
aMiHl Am* farulnlns to niaivlagjr ud D*r>-
I itflTIT- W DtlllCmMKi It III* prXHtl tMtltk
Kli* hw (iMMBiled loplfluclnK ■■aikMooM
t *nd Hwiloal y*i kImUAc, oUhuui •DUriiiK
IBM Ika •IbmiHloD «f Ihcariau, er bardaDlof liw
Btoilwlih «oui«l qnwtlanii.— nw^h ITaiiMliMd
ThU •dmlnbla mrk I* lalandBd ibc MiKbdto of
■ttllcln* an-1 (or niph owlloail aira u !■■•* no ilm*
tet iPDKiliy iieaUMu. lu ili* pr*unt laaUDM ID*
ilul; CI aiiMiElas Iha ib-t •(«• flf ni*i««BJ u iK*
•IllHwl Af Ih* naDiat. and ct ahtlitctiiR Iha de-
xriptlMi «f Ita •Dnnrrrii vpHto <4 namiu tfl>-
bHU |w>rn>ni<*'l «llti(in|(<iliiraklll, wid
Uw imill !■ ■ DtiBeix uid pltllaH|>lil«l ■olA* lo
Uia d«fanin*n( ol ncdlala* n( *lileli u mala.
Dr. KOH holda Mieft • hith mIbdUHc (oalltoD Uial
anr vrlUntt* wlilflii Imm hia nma ara naliirallj
•i|«ol*l to han Uiafmpran nf a pcwMfnl lalal-
1«(?L In anrr put ihl> handlmik m«fla iIm
liltthail t<mH,aiid will nodauU ba Imid of Iho
KTiaial raloatothemiJeniaa wli a* to Uia pnt^
SUuiwr.— Sttnkiity* JV«AmI Jearnat, Jan. Ufl.
MAMJLTON, ALLAN McLA2fB, M. J>.,
Au-kdi^ Fliymtbmal in B—p%IM/ur ^MiplHm uid pBrta^tia, a^ttfTi Umd. I*, t
HorvouKDiaeMM; Tbi-irDamptionuHlTreaUarot Heooiiiledi(km,thDrcinBblr
■vriMd ml rewritten. In oni>ocL>Tu voluiDeofARS |*gf^ with 72 iUiMtntiow. CloUutl-
Whan UwAnO'liO'iii nl\h\-ti~^ IhhiIi •I'lirani'l
«9 wtfl% 11 aur tnipliaii'3 ehd^rHtruauL aad tha
■Nml aitllhin •n^aDCHl"I^ >|«T*flauan nl Ilia
took anil lu auihoc u a hIo ruido lo •todaala of
Daa of tlia baai aa<I nvoM
iuroli>BloalJ(inmjklii, ilraii^lua
rriBraotorliad IhU IwoK a> Ih* b*M of lu klod la
an]' langiiac*. ■hl'h I* a haBdwoi* i« iwiamanl
trtMn an *lal(*d MUM*. Tb* Im^vrtvaWMB !• Ite
DBv adillon. aod th* aadlUoaa (0 II. VlU lliaaiy II*
parabaa* aiwD Iv Ihoaa who pnaaM Dm old.—
A\\mltt <md Xmmlagut, Ipril, lib.
•UaWal aaiirolaur.
WlUoMotKaillaliMi ^
TUKEt DAXTEL HACKf M. !>.,
nioMrationa of the InflottiUM of the Ulnd upon the Body Id Health
•ad Dlseoao. DMWud to eludiUlo Uic Action of lbs ttnaffioaUaa. New nlition.
Tbnrouit h I <r rpv bed MM rewritten. lnuii«8*o.«(il.oft(IT |i[>„«iili9a>l. plats. (Jloih,fX
Ii la ImpciaalMa (opHHOthaao iDUrcallna chap-
Mr* wlUKiiiibalsfffoiiTlaooil of lha aulhor'kpar.
fcrt •lD»riIj',lmp»nlalH]r, anil Ihoroiigli mantal
n>«> It. Tnli* haa aihlblwl lha raquMla
iBOOBl Of aclantlllD addraiB oo all ootMloai^ Ud
•ha «or* luirinaio lha pliBaummatko mora Analy
ha* h* ad)i*ra<l to a phTrloloKiaal and rational
tn^lhod of iMprprMMloD.
' daduciion. ih<
OaUad t^ an onll^
uWjgU,
taOHj ha* rMhUmodnr
■oianwaoiMllnWcHltniidnniBln In |NynholHT.
nrailaiialir Otwidanao to rharlaUna and •mnlrlcab
Thla bnak, wnll MSMlnd and aril wrfMan. Mnit
eonnoad Haalf te arm UioiiiMftI uvdaraiand'
|08.-.V*H r^rk Mtdktl AwimI, topUDbar <. IMt.
^'
GRA r, LA yDoy carter, 31. i>..
A tactical Treatise oa Dlaessee of the Renroiu Ssratem. Prtforing.
CLOvsToy, xaoMAa &, jc z>., f. r. c. p., l. r. c. s,,
ladHrn- ua JKaMal iMM*a« ia (ka IMf*^ 4f WkAmj*.
CUoioal XiMturM on MoaUl DiMuee. With an AnpcDdii, oDoiAialiiK u>
Abatractof tbf HutntMoTlli* Di^twl Sutw mnd of ihc Sovoral fiuiM asil Tcrrilorie* t«-
kting to the CuMwlr of tfa* Iimdc Bjr Cbarlb P. Foiaum, M. b., AMbunt Prefwnr
of Uanlal PiMwea. M«d. Dtp. c/ Harrud L'ni*. In nnehaodaonwocUTOTolusiiiorMI
MM*, with «ifht llUioctaphlc JiUlw, four of wliidi arc beautifiillj cotorail. Ctuth, fi
tk* pfanlilonar aanll m th* Mndanlwlll •»
«*f( lha fUa, tawtldal waclilnt of lha Mlhur aan
fcrvard Map in lha tllonliira of loaaoliy. Ii L*
I«fraahtn( io And ■ phjraiolan of W, t^nutlBD'a
llif c**"*! pracUMontr In inUlBB hiaa loa diM
UMl> and jndJcaUnii the iramnaBL if itlillyh
maojr vliatvta and deutntul «*»• et^BMBUl -11^
To Ik* Ataarlcan raadar Hr. rataaaa'a A^
pnriii »M* fliaallr to Ih* nlat at Iho ««rk, and
wlllin>k*ll*4«afrMa*ktdMM to mry llhiarv.
-^■■■^aaa J^rtjlnjil JauiM, Julj.lih.
•RHriano* and hMt Muurtion tl*lnK th* twd-
tiilt oo«M a»on rtleti hli aiparlosoi.- ha* baan
ead*d aiuf hi* mauiit* Jad(manl aaiahtWitd.
;h alfnlral obaariaiinn* cannot hiil b* upalSil lo
|0r-Dr. FoIiKuo'ii Aitlrael tatf aito b* obtninod ncpanUlj in ono octavo Toluaw of
IOS|ncM Ciolb, 11.60.
SAVAGE, GEOBGJB H., Jf. D.,
LMwrr oa U.^ial DftittmalOi^t Bttpttti, Lillian.
Inmuciity nod Allied ireuroeea. 1'rBAi<»] uid Clinical.
la MM ISnwb ToL
rfl»ltiuM.»ltli ISltliM. CIoUlRXM. BM&Tica^aMiMf JfuMwl^MNSt.
■ farafUllraalaelad.andMtracudalrai
Aa a h'aa'dbook, a fsld* lo lha pni'lllliinar and
■ ■ ■ ftiH
o» ...
•hanMorlatIo eltaniaM, Uia Illiiainiitta run are
aMdaot, lha bonk fulltli an admlnbl* piirpiM*.
Tha Maov knn* of Intanlljr am dMcrlbnl alia
•UnODli
I* oiaiyahor* BivamnL |>T.an<
ua Km >rllt*« an auollani manvaJ ^« !»• jmr-
uitonotaadatadani —Am", Jtv, efIntm.,Apr.Jt.
In
PZ.J ITAIR, W. S., .V. IK, F. R. C. P.
The Syatematio Treatmont of Nerre ProetratloD aod Hysteria.
on* handiDfue anwU Ittno. voliuiM of 97 pogw. Clotli.llJW.
kl-*»l>l^>fll>ON IIORAKITT AKDn*TUUT-IJOiagrtTLI!ll(UI.UI»EilVATt(MniO« TVVC-
MK^T. I.of4UMaM lha TrHtMtol, IMIsall TIOKAL XBkVOUS DBORUBWI. '
as(f/.*CxLa'fn>anar»UMM. lB0MTCt*hUi«.\ AlMIMan UMMn. \u oiK^iBte
I #M»* opiatw rafntna. I Wwn* at Mh T««M. CteNb-Vkia.
£OA£j;r.s, JOHJir B., Jf. ».,
Tb9 PriDOiplM ADd Prootioo of Uodera Surgory, For th« w ofWlirfiw
and PnetUliMMr* <>r M*cllcliw niki Kurj^ery. In<uie rtvjr h»»iliDiBc ticuTo TolacMcf TN
[■««, vitti 60t illtutmioiu. Cloili, HM; lMlb«r, $V50. JuM rendf.
InlUmnck tb* villi ir hM VadMiorVd to cl>«
lo I'm ko'*'*''^ "< "i'"!!''*"'*'' ''"n *'■■ doctruM*
■■A yfiifv^urM ot aiMlarn Hur(aty. ><• >•« mkt*
(I ■ vorli 4h^/IwI Mor* f«p«cUMj lf> ir*n i>rv^W«
(hail to <>» UlMOi; o( •UF^fir. IIl< ftUD tu*>
«ip*tl*iio* liifi a^wd tnuiy nttijM'l* fHl«r**la
th« ooik. It «ODtoJiw miiar ptaelWal nttnW la
illii|CM»)>, vhWIk mdiv il Ih* >D<f» talusM* lo
tlu pnMUUbnar; viJ l)M artUaiMliMlon abMi
p«mi>4M lb* irliiJ* mk, los*l>in irtik 11*
litntMiiXr. POtMOM Id vahi* M ■ ituilvnr*
■nasiial. Tti* Um (hM Uil* vorK Is amiiiMiitr
r<-IV>l nuuM b* loo flrooitlT ■mytmiiil. It
mo<<r&, icil K> fl( tMi4il<m I* thM g»ii«i«l>y
imBMil wd Konh thai kVvrit IWI* oppanMlV
(grMcawlon. Itwlil h* ItMinc, It !• tUn tiA
««aalM,rMt>ilL Thn MOk tt (BtliM lo ■ plM*
In nudarn *«r|lnl tlMmlani, — At»nl* tt Bnws
Ju ■■•(.
Thtawerk It •T(47r'>R>'>nii*ntlt«nrM*alH
fftatnl MirRv^y, bh^ wiIT <i'«i*iilrii- nawt «lu*
tMOnib* rBM|HJtuii by (ba bnfwalxs. I> ten
llMnHiilhIt' \nMotl <il>ar*oUr, Uw MklMUm
Mini >flbi«r*}*ilstii'nt. M* 0<«<>iu4aUMiM
WtoM WIIA IhOM ol ina l*»illM |ii»nlllwiinl
lb*ail,**dlW llisnlura l. riilljr npIaMlthtia
taoort dactrlBH »•! ni*Ui^< uf pnaUM ii<a<
pr^wnl dv- '■>* ([•'^•■'^< nrivinmaal IM
ikli rala, lad (ha «ii' .laatn M HI
ctaaaad pra/^ilcal laa' <il<lv|
Iftldt aBllratr alia*!" . i-rla^t
bta maamm la whMi i<- in- iriua ln4
ilMMnaai ar U* mrk.-jr*Kral AatM, t^ V,
U»I.
AHBHVRST, JOltX, Jr., M. i>.,
iljrln. /V,i/. «f Surftrjf ami QM. Arfiry la FJaif. v/ Pn«L, Air^aM fa lAa iVaaA. ifa^M
The Prlncfplea aad Praottoe of Stirgery. NVw (liah) ediUoo. calaqiii
■ad tboroughlr tvtHKl. In one Urge uwl haiiilwni: ocUtd rolume of 1 H4 MM •lit
64S UliMtruioiM. ClMh.fflj UMlier,$7.
A camolaW and i
I aMallaat mtk an ■uiytiy.
Il t» esty nacHiwy la aiamlua ii la *aa al miM
||q uio»llaD» and r*at m'rM nithi't M laiVboak
. (h lti« atnlani at a Enlda tor Ihr (ananl pnicll-
■ ttftfjar 11 f-itly ooDtidan Id datait orary iur^cal
I Injury aed dlwaia (u alikli th* b«dy i> [labia, and
'rttrty adv^^ la *'irir*ry vnrlb n^liif la la ba
bund In lu proparplt-n. Il U iiaqumionably ilia
uiaad ntnai mniHTti^ «FD(Eta Tfiluniaoa ■iiri^rr.
. I Itia RnglMi lnnK<ia«a. and '■■' ' '■■•> radlrn
|)inlcontliiiifldap|fr*tTiatian al: : 'luttly
tbnand — j#>BiA(Fii lyitti-turi
"rtla It ona o(l(ta fn —
Um many «all-ka«n i
II PBinl-hM In ■ ■-' '
ooaiipFTbtealva da*ciij-..<.r.
tin noil itcnarally anii-luy<'
tiirglval alTvoln^. with a i
prUi«t|d*< an ufalfli l^ —
Mafd Tbv aelln >a r 1
■od • uamkcr «f &-
ft ^-
'•:tara*d wM *t
K***nl •arrt-
i-cT a dMT •»
iHaariti
jnriwaw*
Iran
Ibal Bnally anbi
QaWaantl Lm»ttt~
CbfUt, LXn. I*. IHi.
. •; 1 nruM •••
raliilly rn> <
HIM la«i>iia«
I ef II
baMk-
irnriTT, iionr.iri. m. r. c. s., etc.
Hituual of Hodorn Surgurjr. TwelUi odltloii, ibomighl^ revtatd brSru-
Lnr ttuvi>. M i;, !1.S^ !■'. K.(:.8.. In one Sra.voluiM or 9tt5 iia«M, villi 3T$Ulu«»
tleni. CUii, M; I..>ih«r.t0.
ItlaoHaDtlftlly a Ti«<ir heat, TairrlU''n from ba>
SIbMu M aud, Tha aditw lua broucbt hi* ir«tk
BMlKa lairM data, and oeoity aiory tubject on
nfah Iha alud*nl aii'l [trar(Uli-D*r mfwid <li>*li*
ft>OA■llll a *urf loal V4if(]i7i«. ha> fjiind lo plar-*
ra. Tha Toluaw rinw aiili alixil larniy ■<■■•>•
«f fannu-'M (oTcrtnit a brotr:) raac* vl pracltfal
[Uaranuilea. Tha iciidanl will And thai Iha naa
tniltl It lo thbt B*aaniil4n what iha old CAa ma*
to Iha fermar, aad n» hlcher pndH d»hI b*
m»ett4»t to a>r rnluma -XariM &ra«ai MtitiatI
Jmmml, QMobar, 1»T.
l>nlU'* Sii()p>ry hai baan an axpaailtncly h^
IM vork In ih* prattHiMa, li i> *iaia-l ikaiawV
raplaa bar* bHn Mid la Ennlaucl. aiiila la Ha
t:Biwdae*i«>LaTar •inoaluMrwi i>tu*.iitn*l
■ip*d at a taiUbenk la a Tary larw *>iaat I
lilt Ih* l4M mr to Ihl) «Mntry ii om t> '
■ppRclalad ihat acopy waa IimimI by UwO
awal (AaachtuTcaoo. Tha poiwni adlUta. •
II ha* iNaawiia ItaUrH r^-uliar tn (ha wart a
Anal, (•mbn^lat all ranni dtMnrartoa la ■MiWIJi
an.l 11 tally up 10 lbs llmea— ClMMaM MU
.VflH, Saptaatbar, UBT
OANTf FREDERICK JAMJB8, F. R. C. B,,
Smiar Air^vi'* lo lU AiyaJ Ft— B<ttpUtl, £anA>a.
The Studflnt'fl SargtTf. A KaHnm in Pom. Inotw aiinuv ocbinfoUnt
of mS pagc^. with 159 ongnTingL doth, tS.'&.
OBOSS, S. D., M. J>.» tL. !>., D. C. L, Ojetnu, LL. !>•
Cantab.,
*MnhM /■'•few ^ JiWyry M U4 Jtftrum tltdmU OMf ^ I^MtadalpMa.
A Srstem of Surgery : PacbolcifUal, lHi«niAlc, Tlitnii'mtle mil 0|Mmllt«.
Sixth nliiUin, thurwiehl;r rvviwil and ^itatl? imPTOTcd, In tB<7 Urge uhI MMUhlll
Erinud imperial ocUto roliieae* inntBining 23n pgei^ illuilraUd Sij 1933 •cynutlfb
IroBUt/ faoeiBd in laalber, rabrd Ivmln, lift.
BALL, CHABLES Ji., M. Ch., J>ub., F. R. C. 8., IC,
fi*rpt(ii aad rbat**' altfir P. /hn't Il*tp4i-il, l>^U».
Piaouca of the Rectum and Anua. In one iSiuu. Tntunv at 4IT pn.
with M oil*, iin.1 4 a.\i>nd |iUlw <:k.lh. gJ.SA. Sea&rwarf CHmmU JCMMatt H.
OLBNEY, r. P., M. D.,
Orthopcedio 8uTtraTS< TDitV\M<meiA Vvgst.ti.wniniand8nid«fiia.
Inumhaa'-
kTiOKS — 8urr«tT-
21
MBICHSEN, JOHN B,^ V. B. S., F. R. C. &,
The 8oi«aoe ftnd Art of Surfrory ; Beinir ■ TmilM on Riir(i<«l tDJiiriM^ Vi»-
«H« ukI OpcrMJoM. From the ciitliili nod cnliir|[«(I Eiigltili (ditknu Iii Iwu large and
faMUtifiil ocUvD TotiunM of 3316 |uig«», lllualrited wtib 9M eafcrwTiiigt on wood.
Cloih, SU; iMihw. niwd baD(bi,911.
An of Sarf*r]P~ •>■ on* of Iha I
toolulo in* Eoc'lin Ikngiiag*,!
•4lllaB*Blf toeflrn* out unTMoi ot^nton.
HJ» KnM clwHur* In csnlwly eammanillDiE It U
MI nwUrI-r>MM**oa( -Vw^ April II, InU
For Buay TMn tVU filual' irnrk lui bMD
(niri* bv H«l*rMtM At u*«li*r> ilti> prlaeleU
Mti-Mik •>• fur(«7 for mtdlcal »iuJwii^ «bil»
■Tba Seine* MBd t thmuRh liantlallOD* IdIo lb* iMdlsc «ODIlDaDUl
ti**t (Dialol Uit- \ lugiiiigr* li tnar b* nlil U fnI4* III* aarglaikl
•,•04 >hl**<fhth .Horiinaior III* clrllliad vorH. Xo •lealfise*
l«t)i oMnton. W* of Ui* finnar •dHlon IiM li**n droivHl and ao
of Ui* Rirmar •dHlon lia* li**n ActiffrA and ao
dliearM), d**lM or hnvn>rpm«n( which haa
marli«4 (h* pracioaa o( ■iinrrr dutinc the laai
dwad* hk> bHB omluad. TUa IlluMratloaa an
miuty and riMutad in tha Uahaal aljl* of art.
— J>><.i»iif> JfefKal gfti. Pab. K UU
BRTAlfT, THOMAS, F. R. C. 8.,
airanm a^ tjttmmm Surgrry ni Ihv'' BatptM, Lamiam.
Tb* Fr«Ctico of Surgery, h'otinh AnMdaui ftcco tb* ronth ud nrlMd Eiw-
licit tdilloii. In one larfte murvfiy liinJaoine itnporial octaro TatUBM ef lOUpa^w^wlUi
727 illiMtnLiooi. Cliilb, |IUCt; l«uth«r. fiM.
Tha hMitlb odltiOD of (hla mrk I* fUllT abraaat
or lk« UuM. Tha aulhoc baadlaa hla anhtaeta
«tUi IhM dapwa o( JuHgniaal and aklll ithlali Im
•nalovd tvTnn "■ t«u>Bt (oil and Tarl*d •>'
parte***. Tha ■•raarnl odilinn ti a Ihotalllb t*-
tWoaoflhwa BhltA prtccd*d Ji. wlih mu«h i.rw
malU* addad. 11L> dmlnn l> » k'^'''"! and
IHlwl, and hi* axplaaatlon* an u lucid, aa to
AllM laa mwh anoai lli* niiharl ncdfr of (otl-
fcaaka tor Ih* nadleal Undant, Almoii •rar;
(oplo la lursar; I* proaoDWd lu inch a fbrm aa %a
asablf Ih* bu>T prantilcnat lo r«*1*> aarmMaM
In *<r*rr-da)' yrvUaa In a thnrf (ImtL Ko Um* la
lUl II ' " " '
iMlwIui uhIvu Uivorla* or •upfftuoiu (Wlilasa.
In than, Iha Kork H amlnanUr «Mar, loclral and
praoUoal.— OU«^ XaAnl J^nal b«I fcawawr.
ArrLI, Mat,
ThUbvali li aiaantlallr irhal U rafporlaloba,
•li- a manual for Ih* frnrttH at aurfcry. It I*
p*^u I iMty vM I niin] briboMudaoln hw matal
l-raMliMaar— Tka MttMit /tnt, AasaM U, U«.
TREVES, FREDERICK, f. R. C. S.,
/fnHlimn Pnfutar 'l lit Sa]/nt tUlm Vf SurywiU af £a^taM.
A VftDUal of Surgory. In Trvsliaea hf Vanoiin .tattion. In ihtv ISmo.
TolumeL oontaiainK 1S66 tinges, wilb S)3 en^raringa. Price |i«f toIiiri*, cinili, ti. S««
Slititm/ »ria of UatumU, p««e 31.
W* liBT* (i»rt Iha O]>lai0D> of ihlriT lh(*a , _ _ .. . .
aqUwn. la aa *aoyei«[aBHlc term fcr tanjr and I Jxta ar* aloari pnoltd lo (iir»-h*aTT tfp*. 'o
rndr rftoroaM. Tha thrao lal'imi" vn>hra«* I
•Torr nrlalj atunlaal UhcilDiit llktiy m I>« |
aol «llli, Iha i«r>cnptia *» ahmi and I'lihjr, and I
] Iha aaJiani lalaU and tha baclnalsia << iirw inb-
k-naa
ihal a pamn majr And ahalarar Infainulon ha
may ti* in dh»i '>f ai a tnomtol'* iiaoor.— C>a>
nnmitii £o'i"i-'Ti>-ir, au|ubi tl, IBM.
I
I
I
I
P«
HOLMES, TIMOTHY, M. A.,
Sar^rmm'4 Ltitt^f v l*.'fn 'I ■^- ('inf^t fTi ij T. T i *ill.
ASysUmofSurgory; Thooratical and Pr«otloaL !>' TREATISES BY
VARIOCS AL'TKOK.'^, A3iKHi<\tK cniin.jt, TiionoroBtY RtniCli aho iiK-CDiTm
br Joii.s II. pACXARit, M. I)., Siiryeon tu lli« BpiHopa] md fh. Jtaapli'a Hoiqiitali^
Pliila<l('1)<hia,ai*iit«il I17 a mqia of Uiirtr-lhree of Uiangat siDJntnl Amwlcu MitgcotM.
la tbn* laiYO imporinl cvuvo Tolumva CDnUininf 3)3> ilonU»eMunuiail pun^ wUh
ftrS iUiHtntlofU on iro-l miil 13 Ulhognuihle |ilatM, beautiAiH; colond. nie«
■M, dMb, tlX.0O: twlber, (lil.OO, &M mJf fry n£*i»^ii>«n.
WliARTOX, HEyRY R., M. O.,
OwnwIraWr ifSur^mt atd ttitii'ir pa Ai^iMt /haauu g^ CUUra* la IM r/« r, qf f IH.
ICiDor SormiT and Bandaglog. la cue rvrj huidaoim* l^no. voIubw of
4n (MRe*. wilh l(n cngrsrtnsi, nianr twing pliotogmpbic Clolb, t3.00. Jurf rtmdf.
MAJS8II, HOWARD, F. R. C. S.,
S*w .Aauloal JiVf*Hi lo m^ L^<irf n> ^aafo** al JK. AorUalMMIi'a Moi^iial, LMdaa.
DiMUa* of th« Joints. In one IXmo. rulume of 468 |Mg«^ willi <H wnodcau
md a mliiraJ ]>1aIp. CIoUi.»2.00. 8ee&ri«iorClt»iiaat JfaawiUmi^tSl.
BUTLiy, HENRY T., F. R. C. JS.,
4uMlaal AarjtM la St. SvliMMHtr'l Btapiltl, loflAn
DiaetAM of the ToiikuO< In mwi I2in". roliuiM of 4M t^gt*. »lth 8 tolortd
pUuB aw) 3 vooduiita. CloUi, tU9. 8— &rw a/ OiaMo' JfoMui/j, |Mge SI.
TREVES. FREDERICK, F. R.~c7s7,
amrfmn t* ait Uilt^tr mt Swintl Ol l/tmien Htmptlal.
Intaatlnal Obetruotloa. In on* iMckM-alse iSroo. volana oT S3Z pt(•^ ritb M
Ulnttrui'iiM. Limp ckxli, blue odgei, t^OO. Hee Smm 0/ CUmieal Uamialt, juft 31.
GOULD, A. PEARCE. M. 8., M. It., J. if, C. A,
Jtauiail Utrftfm la ittUlnu /(.••j-.Tnl.
Zlemonts of Surgioal DiagnosU. In oat pndcM-aba IZmo. toJimm of fi89
pagCi. Clolb, 12.00. See Sladi%u£rit, af UaiuuU, i«gt 31.
rtuiim pRi9R.-n-Lf!a
■UMMiy. MiMd
axd r»Acno« or
3J«n Hani. M. D. ta
thiMiiiM rioui.nn.
■ruLJunspjuffofpLnorspBaERr. PDunit
Amamw Awn ifa* ittM Bdtabnnh •dlUoa. la
HILLKR-S PIUmCB OF eVMSBT. PMUlk
•oL«i(«nw..«VftL«M«taMnU«Mk. OM^]
22 Lka BKOTHBoa & Co.'s PuBLiOATiONa — SurE«>T> Fnc, Oiiduc
SMJTU, STEPUEX, M. It.,
Tha Prlnclptes and I^aouoe or Opanttive BurBary. V«« (Mooml) ul
Uuirviflf^tv rcvi>«l nlitiuii. In one fvxy liiuiiliKiiiic oulaTo raUine of SM pMn» wtlh
10' I'M. C'loUi. tl DO; l«Ui«r, (i.au
Ho viinPT- '■d UiB lataal tiiMtiauMiM hMva ••
t<>'L ft-r 1Ji4 >1(icj«i. - L.^a I
Mid «p#-» c» M )v>^)
nuni^*<-tt;j)iftnBti;.... .- ; ... ,.._.-.>iir .._ ^.
this tvlama vflriuDS liplil ■ bkim Ma*tnmB
plac*. aad Mldom «lirila ntMlMT. oo nuOM lia
aniuual lh« hUmlcmmiH IM pav** in tMit. n
cam|«M(MB,KXMlant i-rlBV »'ni*TiM* T
Umu, wtd Ma(iun> lu dwMMltr tnuata
aeWf.kll OSMIiIm M PMBRMBd M.— >«h4 J
Mil Atrfltal Jovwr. Majr 10. UK.
Tl. ud ivtr 'kliutil* book U OBDQr
Ihd ^4'^- --.-. ^jh'U17 vtfrkAOD mocldm Dpvrmtm
•Ufynry yrt pucluhvd. Il* aiiltiot aad puMUhar
har* •uuail uo ptlBi Iv niklto 11 aa (lu m iHulUe
■•IdraUand Uialralliirta ha>a ^no liap«lUoii
pronitn«niamonBih«r«oanl mrlra tuiliEiidaparu
maul of tmttett. Tha hoali 1> a MmiWDdtain foi
Ilia cnnlArn •(ifKHA. Tha praaMil, tha only rwieJ
•dlitnn •ii>"D in*, iikmdu muj ehuiSM Inua
Ihp iiriflnal rnkiiLial. Tho Toluaia I* (naoh an
l»iKfd.kDcl Ihrtrxt Urn t>Mn UiorougblT r**(M>l,
•oaatORlta ihQiDDn impmrHl ae\aoJt In laap-
• Mrmi
noLMEs, TuroTsy. m. a.,
A Treatiss OQ Surgery ; Its Prmoiplea and Praottaa. K«w AnwtiM
ftauUic fifth Bogliib vJiti"ii. oJiird tn-T. Picxsuna Piux. F. RCH. In dm mm
TcJuliM of MT toeos «il)i 4^ illiutrntloiw. CSotb, 9S-00; lonthor, ti.OCH
Td tba jranopr iD*inte<n of Uia prolMrian nnd br Ik* ganinl |>raMlilanBr. (asDhlnc ihrnaitliB
'■■■'■" thai nr* aH«urr lol* lioftwn fur Oi» isaMM
pruaeeutluD at lAa phjrilf his'* oann, tnp
MoUwn 'Mt nsqiulnlad wllh tho beak Mid Ka
marlu. *b tahs pleuura In rwantmandlDf It a* >
9Ht%'*ty DDin(i1«ea^ thoroKxh,
lll(Uinl*d, iDiHlara, n voiii
niMU for Ui« aoiiaan ipMlnllit. adwinMelr conclie
w*l1-i>rltt*n, hilly
oolhlnK (hal In Dnrpraarai k.
•ml LiiiMf*. unHlnnttfle or iBvjn
attJ,«H Jiwytal, July, un.
as"
S
inol *Tnid tiklnt tJibtb* oC UaM «
•lansuihatarara, ha Ba*4* nil ite atjl*
nir*. Iloi «kit bHiar waMiMn nn la
^W
HAMOJLTON, FRANK A, Jf: D., LL. D..
A FrsotioBl TrootiMi od Fraoturas aod Dialocatioiis.
lioD, reTi>«<l Aotl cdltLs) b; Stepiieji Nmitii, A. M,, M. !>,, Prufiwur of < i^rj
in llw Univenilrol iboClly of N«v Yoilc In one <FcrT linxnlaauw ovUvu twluiu* «i
pagM,wItlt SOT illuMMion*. CloOi, 9M0; 1«m)i«i, (u.V.
Il hM rdHliail tha hIjhMt andarwnantihitn
■*(k U|4u a dDpaxtmanl c4 tumrt ou pcuUiljr
raeMm. Il I* ii«i>d aa n mhi-Vji^V in eirry Ffix.ti
*ni «Mi«M of Uui eonnirv , ' :
ten boo enlled upODM |>ri r
WhM mora nn la Mid In -.
UtanbNn Htd Willi iniih ii.t: ii It duuhiiu; if
aov *nf)tlc*l <•<»>( l>u np|M>r*d diitMa Iha lual
tialf o*»luiy wlifeii toon ocmplaiely Hilcd Uia
»laaa to whlah tl vM da*tsn*<L Aj Dt. Saillh
nya, lU (reM martla npp»at moat ronipleupBiIy
N lu dltar, I'jfi.'t-o. an J yalcoinpnihaiifliTaaiala-
maalofi'.'. ' lUii rin-lcnlcaBailailnbla
(asI'lK'^k I I'ld c-;](>n. ancNn ltaiir*allh
Of ellnKsi ' '<>"'■ a-Infiii ll(i> iti* dally
AManltln* I'f jhr»oiUl'«npF. VraoiLirFt and
OBloadiDtia *ra liijitrHK wlilcii lbs gan'ral im.'il-
j Uonar, tn hia tbrnnmltt a* ■ an rvnn*i>*
Up«A I* iraM. They Itm m paM of
ha cnonnl *Tnid tik' ~
' iiqilaiaU
la •enir*. _. _ _
-ok (ban n irorb IhM la d^Vni'i! atoliMiMf n
' ifoatlnf rm^mraa ab^ 'H- - .- *a-l (»**
qn^nUy ooauina IiiM Inf. i^ pWa ••
|l<iaCB,taF Uia nianw*ai>'i i . . . •oitrana*
(hfti b liktiy to baakai mu. U wah MbM
Tliaooiinttyl* <UI*i*lth nUlrwInMMi.^ .
ivnaa wliatv aotliUat* uv Ci^tiuilty aaMin*
nnd towhlck innanl prb , i>ad iw *
llnnUMd lurMti. ate . ilaUi all
ourwl W« aSdWItr (.' - ..n i
•kcvid t« la UM Uhnrr -I fK'ry j.nat
OwwaMtl JMlait JltH, P^Wunr/, foL
BTIM80N, LEWIS A., B. A., M. D.,
PkM/Iy of Uvr. ^ ai» qT .Y. r., CbrropowCiu jTonter qT !<• Aoala dt Cturmr^M t'^
A Uanool of Operativo Burgory. ^ew (Komil) editkm. In ooa *«7 hi^
MiiM mral ISiDO. T«luBia or ACS p«)(«n, with 342 iltiiatnliotift, ClMh, $S.AO.
Thnra ii nhw* raom tat ■ ipxid botk, to thai
*h<la Maa* ir«rk« on ppmita •urvry maal W
oouliUrM (uwraunDi, (hat nr lir. IMn»on hai
Mil li»n«n. TliB aiiilibc kno«i Ida dllD^ult an
•t oon^mMlfoD. Thiu Uia manual Mrna ■■ a
w«rtl cC ntfannia. and at Iha ••ma Uma w a
hnndy auiJa. II iraEhnt iihal U firaCuMa, Iha
•Up* nr a|<>nitlaniL In thia vdltion Dr. NJmMn
bMaovihiiO l»JltaM the chauit« lla( bnvabann
By the eamo Author,
ATreaUM on Fraoturos and Dislocatiooa. InlwoluuulaanMMUNMt
iMiM». Vol. I, FKAI.-TUIIIS, •'JH3 |KiK««. 3eil baaiitifiil illiwtnili>nu. Vol. II., Paucv
nov^ uo page*, Willi 103 illiutditiuiu. (^loplMe work, cloth, $5.50 i IcaLbar, (T.
Either Yoiilinp t«|
ennoud Id (warellT* tnathode asd mxitiiam It
SaMUlMpUe aral'nl.anil ha> >dilt>d aa im^b
mas* n»ir OHtMliMia and tulatlaaa M at
tupa sf olitM DpanllonaL Wa do »->i 4a>»>a
aiiAl Uii* maoaalalMva niaojr atfiaXoni iiialpi
KriUtti publleMloUB ol iha nama nlaa^ •lUl aaiv
llairo UiH II annialna muoh lliat t* mttnia ri ■»
lallno.— JlrtluA JfarAMrJowiMT, J^ (^ OR.
tponlolr, cinili. $3.()U ; Irailivr. HOO.
fUieaaoond ntema muki lb« ol DlBknMloo* •< It
Tha appanmneaoi
eemnlatlon otthaanthor'aonEloiil plan '-f iiniar
tnK ■ *nrii ablck aliiMld praaani tn Iba niftfal
Gunaar all Ihni It knoirii on tba otvnate ■ut^acti
nt Fraiiuiiaa and PltloraltoB*. The teluma on
Tnetana aaaumad al ooca Iha poaltlon of nii ihortlr
«n Iba lUtijM*. aad lia (MnpanloD on Diilooailant
vltl oodonMba alnllarty faeel>*il. Tha cliaInK
TOlunio ot l>r. Stlni*nn'a work ixhihlU IS^aurxBry
la taiiaht an
moataanlnant anrKasiUaorUia praoaait tii
lalolnalhararallaoCauoh *>i*aila)l rw
iniiai lor • long tlma be T*Bar<lad ■• a» —
on all aulilecia pMMlnlSf la dlalo^hiaa.
)iracilltlanararwrc*g7 vlll IWI U lananhHI*
lilm to hat* II for wnalart rvfaroaaa.
ifalical .V«H, Mar, Ml.
nCK, T. PlCKERIJJO» y. K. C. S.»
Jl«aoalo««*U*l«'«o*iS«vn.«*E«-a«i.i<.ll'3f>»»»,U— -. '
I
LxA Bkotbreb
BURIiTETT, CHARLBS H., A, M,, M. D.,
Tbo Ear, lU Anatomy, Physiology and DUunaas. A PnctioLl TrastiM
Air 111* "<• fil Mnlinl i^tiiiivnw und I'miliiiuarn. Sorotiil tdillon. In one hamlwDtiie
«tUTi} Titl<uu« uf 6H0 pHg«, witli lOT lUiWtratkiiM. CloUi.HDO; UotliM, l&JXL
WanoUiTltlipliiuursihfiippMnaM*^ aHenod e>rrl*d out. and fnii(4i a** mailar iMtA. Dt.
•dIUM) of (hu T«lii*bla woik, Wh*n H dnt eMna Baroxi'i aaiH must Iw t*ur4M ■■ k vny nlOB-
•U n «M MotpMd by iba f«ii|itMi)an *■ ona uf Ma «at(IbuUc« id aunTnircT, SM miy oa
ik* MHiduil vDik* OB fn«lam aiinl lursarT In ascvnnt of lt> fomprat— »lTi< »—. inn >«rain« H
tt* Kagnilt lancnuni Mil la hit Hvood triltlnn MoMua Uis mnli* oT Uia (araAil panonal obiai M-
Dt. Bimaii lua ftillT maloBaliiad hia rapulaUan, tlon and a>*«rlan(*or)>il>*inlB*BluinlaDni*ab
Jar Um beak I* taplau alih nliulOa lohumalioo —LvidM lamal, Fak. n, IHft.
•M4 •oaBHCloiu. Tba raiWIaa ha* b«*D cvarnM*
BBBBT, GEOB6B A., MI. B., f. R. C. B,, JSd.,
Dlaeaaea of the Bye. A PtmiImI TtmiIm fnr Siuilvnu of oplitlialaioJonj. In
one octaTo Toluine of 483 pages *i>b 144 Uliuintioiii^ U of which are ImutlfUllj
cotond. aotli. |T^0.
Thia navaal ooadlduM tar ttmt wnaBC apnthal'
Balo^ralaiiidanldaitnilgBadloMparatirollairal
In oiiaiavlar kiid <ha plan Usall adkand to, Wa
k»»Maa toMltily (Uack by tl.e rut loal Uaia
tn lb* MlMtiaa or ulial !• aueniw whleh pai^
■Maa Iha book- Tha aiilhor taana lo iiaTo tha
■Doammon IWcultjr of *lnln| hia miHan m •
whole aad lalilDR tha wliaDi polDUaBd noloini-
ni>lD| hit rvadur— preiuniably a Bludaul aoil ■
nodco— «l(h a man of daUJi* irll^ no kty l<> Uioli
iirinit«rilnc, Li I" HpparaDl f hat lb« Ultratiu* af
••ch ■ubJoK hai hna (nno onr tn ■ tan iImt-
odO naanar- Tlis tet itiai ha na ariMM •
Olstnii iniillin tm (ipgiiiiian and oot an aaiTalO-
H>dte hM ahnji b««a piaaoDi alih iha •Dllior.
Tha namtar and alo»llanc*ar thaoolorad llto*-
inilona In Uia i«i* doaarto mora than a paMlw
ooUo*.— ilnkina af OrUMmvlatt. B*pk 1M».
NBTTLE8HIP, EDWABBf F. R, C. S.,
QpUMMf AifW" a( A. TteMi- Uotptftl, Lmdim. Surfam 10 14« JlDiral totdm (JTooqleMa
OrUtalia<fV>M}nML
IHaeaaea of tho KyO. New (foiinh) Americ«n fium the nrth F.ngliih cdilicu,
thoroiurfitjr revisoJ. Willi » Hi]|>|>l«tii«Dl on lh« Ivtm-tion of VajIot BHtidiMM, hj Wa<-
LUM THOXiiOlc, M. !>,, fnirnoKC t,f OjihthilitioUf^ in iLe Jefleraoii Mmlical <JuU«^
In tmv ISino. Tolumvur oOU iioroi, vith 161 i1lii>lralkai>, Mlection* from Snellen'* icrt-
ITpas Bod fanDolBi^ uid a colored plaW. Clotli, f2.0U.
Thit la a *«lt-kaowa and a raluibla varlc ti
vaa prliiiarfl)> iataodod for Ilia u>* <if •lodanu,
aad *«ppll*» i^Hr naadii admlnbly. bui It if a«
••alWI inr tha prarlltlODar. or IDdMd oion to. li
Maa aot pmuapua itae larRt wiiaaui«f ncondlla
ksoirtodi* 10 ba pKMnl which Mom* u> ba aa-
mmmd tn mm* of our Inrcar workiv u sot Udlou*
(ram vrar-<<oo«la«n*nv and yst eorara iho mora
Impartant patt* ef eltnlcal oi'lithalBolofj. A
■applamaol I* madotolho prasont adltlonoo tbo
prwtlcal xiunlnaiinn vt nlltttA acaplo)^ ai Is
«alor-bJ1ad»t^aod aoQi«no«a4^r Tlilian and heof^
lag. ThIa l> ••■> ■illlan, and oeatalaa feod
*"n«*loni br ihOH irho uf b« culled ■■ "
maMo (ooh •ualnaUooa.—Jnv
Awaal, IMMUber )*, IMO.
r*rk MMmt
JT7LBR, HENRY E>, F. R. C &,
A Handbook of Opbthahnlo Soieaoe and PraeUoo. Handtuma »to. ml-
uiuror460t«igea^wii)i IZ^ voedoiUy 117 cokircd plnM, Ml«ctuiiu fiau TeM-irpo* of
JM^r and Bn*Uni, and IIutmgrMi's Oolo^bliiidiMM TtM. Clotli, fiM ; Uatlicr, f^M.
II pnMOM l« Ika atiidont ooDOla* daatrlptlniu
•o-l i/[>L»l lUuMntloM o[ all UnpottoOlofwtflM-
Uoo^ iteeod In JoiieinaltlOD. aa ai la ba RrMiitd
M ■ aUnea. nayuail a doiibi ii !• iha boat lime-
irlat haadtook of Dj-ltthalmlc •clBiii->«i,lohbaa
•T*r appeared. Tlwo.mxt U illli Uiler, ihree
llluWtaih>n*ar«nearl]F>llorlatnaI. IV" h«»* ««•
BMlacd IhUaMJta wvlk >M (Kal tarr, and It
nproMhla Iho mmnoolj aeea|iicd ilaaa <4 a4'
na^Hl ophthalmolMlite. We ean tnoai haartlljr
UUoMni
PH«« *9tW Infill UfMUl/ ■■^£V|<1VU IJOW* Vfl HV
: ophthalmolMlite. We ean fno« heartily
ad Ihla book fo all madloal Modanbt. pna-
n and eiwdallaM.— Oaiirofl t/mnt, Jan. It.
yORRIS, WSt. r^ M. />., and
A Text<Book of Opbtbolmology.
Wllb UltHlratMHH, Pitformg,
OLTFEBt CHAS. A., M. D.
In OM ocuto volnna ef almui 500 pagi^
CARTER, R. BRVnEXELT^
.F. K. r.
' Smrfay «r SI. CoriH*! Whj" '
Ophth^mlo Surgery. I« ^
ooloT-lillndneHi t«t, tnt-lviia nu'! < '
Strim t( Clmitml Mtmmai; page 31.
& FROST, W.ADAMSy
H., F. R, C. «.,
Jh"! QfUkalmt Strt—K ami Jaitil UiL
oa OpV Sv . Si. Oml^'t Hop., t*A«M.
viliima of fiSft ptvai,*<('> f*' "•'oduili^
1 |..'i,ilic of fanmOa. Cloili, 1225. Sm
VRLLSDXTHKBVK. Id one oouto nlome.
LIUUNOI ADD HOOK'S IIA»UT IWOK OF
UPBTBAUtlC BUaaSRT. for Iha aaeof fnc-
UUaaera. Hoeoad edlUoi. Is oso oolato nl-
ona at m fiase*. «nh M lllua. t^oth.ttH.
I.aU-|IO!( on IXJUKIU to TBK ET^ ORBIT
AMI BTKUIm' Tbetr Immadtdaaod laneM
BBacta. Ia<iaa<ielai«mlaa»*e(«Hp«v»a,w)th
M IllwIfWi-aa. Oath, KM.
i^.
J
ROBERTS. SIR WtLUAMf Jtf. D.,
A PrftotioBl TrstttiM od Urlaair a&d Booal DlsaAses, inoLtuliiu IT»
any Deposita- Foiinb AmciiaiB Adbi tb« routtfa Lnndoa ciliti'in. In tnauai-
Moi* ocUvu rolunM ot OM IMfM. with 81 itltiMntloni. CInlli, tS..'-'<
IlinBybaiald*Ob«<h*fc*ttlM>»k laprlninslha
•UtM«> of «)ilDh tl W«M&— n> JHV4«H./*flrnal
if bU M«iU»>' Sctfuo, iiL UM.
TbB iraonlkf TkliMsnil tiiUik <f (ha book at* Im
■ mawMr* ttTt*t4 trom H* nMliiW malBtanuK*
nf •<tiaiult*4i>rwUc>lahHV*ar. Itk ta un-
rinllM •luMdlMs o( arviTthinR which riMn
AtatUy or iDdlMMlv •» th« dI>c■l«<^ l««(o«l«
tad iwlnwu «C umtf iliMili^ ■nJ pmmhm
mm*lal«a*»ii ttol fn
voMi- IB lU a««Miiil i-l
•!• >a orr Mr
nt tlliiwi
>f JfiHftaif*' JfaUni < 1<U
TiM **l»a of ihti irv.. —(Li at
phn4(kao ia ^tity poi' i* '^v
•attntlfil. Thcl U U I~.., -r — -.. .siMllM
pr*MBl ka«ltdc*l*kfHitn««M;iiB«ra«aMi
upon Dr. Robam, «h« hiM a irMa npUMMsBt
ini*r pf»uii««*r.-ir«tw>t AMe>4<>Mra.t»
S7 the Samo Author.
Diot and Clgeatlon. IncmelSno. volauMofSTOpp. C1otli,91^. Imim^
PVRVr, CaARLF.8 IF., jif. i>., Chicago.
Brtgbt'a DisoftBo and Alllod AfToottoiu of tbe Kidneys. In oo* «■>
niliitu« ul U8S lugtM, nilli illminiliuiB. ^.I'-lli, |^
The aAlant nt IM> ««ck UtP'tUrnlih >*)r>tw< ' (hon ■■■«• (ha UimMc^ IhoW ui4 ir« ■!■■««»■<
MM, aneilal'aDd omwIm dMorlpIlM) «f Hie letOR mats ftiHr UUa ttair ■■i-.r ■ranb^HUi
nUJMMST and truatmaul of Ui* <t>l«f Orniale un irvUinaH Uia mIMr ia naninjlailt ■»■!
Alw— a»t< Uw kldbtr MBrtataJ wiUi ■Ibunlou- JiUatlac c)*w uf sNitnlUIw. mnA mtttm mi
rtfc nthlob lAall tr|tfwan( (ha moM raofal ad> uai. anal (•x^'bgoK* nanally do, tM uaJupanM
tMtaa IP ont knowladw <ia (lm« Hiitjaela 1" and "" "" "
lUi Moidbo ut Ilia oCjoM U a iib 4M«clr4U>n of
IM bo^. Tha aotk t> ■ uh^iI ««*, Ki'^BE '■> a 1
Uai. ahal (
IW«a «hkh ara all lanratlaot (a Uia cmmM (■»
UUsaar^TM JTawAMbr MidUml dtrMHH tM
MORRIS, HEXRY, M. B., F. B. C. S.,
Sorgloal DlseaadB of the Kldaer. In om ISbw. vo)iim« of 664 pap^ 'iA V
WMilcut*, kod 6 (ulurrJ flnUs. Lim]) doth, flSS. Sm &rlM «< Cltimoa/ "— rri'f. p^ ft.
. .4 (fw««w«r
l>a KloalJSla
la IKIa luauua) ■• hara a dMteM mlOMta M
•uixtial ilWnlUTa. ahloli RlTa* laforfnalloa aM
vlMmharaWbamaiaHh In a alalia vork. flueh
■ took ■» dl-lln«(l]r rHulr*d,>a4 Mr. tUitU
hat rtrj tlllcanlljr m4 MSt fartMOMd th* la)k
h« (oak la taa4. Ii
bcniKf rvtwanoa, U4
IMoaafa la Harioh of ■" 1
■•(iMMitand UMflhrom'j-iiDt'-crarli* anki__
Dm
^
LVCAS, CLEMENT, M. B., B. 8., E. R. C. 8.,
S«Mr^Mi*laa(K>vfini la ihmft thtpiml, lo'^t^
DlaoBMM of tbo UrMhrft. la en* lano. roiuaw. Prtparit^.
tf CViiMnJ Xmvoit, jmg* 4.
TBOMESOy, SIB HESBT,
LeotuTM on DiMUoa of the uiinvr vrgmns. Baeood Amerieaa froM iIm
tUnl EokUMi adiikiu. In on* Sto. toIium of 4Kt pp., «iih S& UlHamUov. CloUi, yttt
B7 the Same Author.
On the Fatholoff; and Treatment of Stricture of the Urethim ud
ITrinary PiatuUo. From tha Uiinl EuliiJi ediuon. In ooa ocIbvq Toloaa «f W
ptgw, wilh 47 «iu and 3 platM. Clixli, $3.40.
THE AMERTCAX SYSTEM OF DEXTISTRT.
In Troutisca by Various Authors. IJIiinl hj Wn.Drit r. I.rmi, M. D.
D. l>. S^ Pntrntur of l*ranh«tlc beotiMn, M&teri* HwUa uil Tb«nipmtl« ia lfc«
l^niu^lfUkLa ODtl«g« of DmIaI Burgerr. In three t«I7 IuumImjm dcUto tuluBa o*
laiDlnK S10U me*, with I8SS illiiMntilotii anil 9 liill.paitB plawa. Par roliima, ek«h, K;
lealbtr, ITj half Uorooco, gilt Utp, Pi. Tbe «xiiu|JcU) wiwli !■ mib r«aify. AraMtly
M&Mn'jnIiMi a<Jy.
Aian auPjvloMilJa of ItantlXiT II bM no hi. ;.teuMaul*M,lu niark nn opAr* la liw bkMT 44
nrlut. U •hoiir.t t:rm a pan of nary danUal'* At^OMtr. LwMtMi alll i* ratUOmt mU M m
■•4 of tl— 4h«* mnii. 1( U iar« to ba iiiulufT
n) naada to noi ttlm aM lit^ hH
ick, Bhlla M* arbfrMloa ai toaa
Hrlut. U
llbraiy. a* Ui* InlorlnaUM II untatD* It of Uia
craataal ralua I* all auM»d In Uia pcaallM at
&aUui7^.1>"^Ma JowTDwiL jSH., Sapl. IMl
A Cand MMMn. bif anough aB4 fsod aMoch
■nl i.ts4tM»* anaugh (or a monilBianl (vklrh
■aM»n*l Mill I
pr«<*4i
•hat (te oludanl
la lb* ii|b( mck, ahlla nia arbfrMloa ai toaa
•ill r-<^*a iMakuUMa baaaM Awa IL Otpfc
givplK Jjw Mi; Jan. Un.
COi'fiUiAJr^ ^., L. R. C. P.t E. R. C. S., Eaoam. £. J>. &,
S-tar ttml. Am. and X4M. «■ Btnl. Avg. n4 % XarUotaMiri JTsf^ and !*• ItoX. Jbtf, ^.rfn
A MkDual of Dental Surgery and Pathology. Thnmi^ilj nrlatA aal
adMited 10 ih« lue of AuwrKaa Stiukau, bj TuoxM cTttnLLvriOKM, U. A., M. Ii,
D.D.a,Pn<f.ar PlijualoKrinllMPIilhaaiphlaDvftUlCbllM*. latn* kauboMoeun
ToluDw ot 41s pam. allb 381 UliaOntlona. CIoUi, tSJSS.
II ibAiiM ** (a IM p«na»taa ol ararr intU- a «*n. II lb* nionB* fM Imo aona n( iv *
tM**f In Ihb CMtnUT. Tfca nan *»toie4 la Brai - • ■^...-
and tacoB'l danilil-n and IrrticnlanUM Ip iha par.
nanaai totih I* IWIl; *ortb ia> prlOK ta mi,
prlr«*bouUnMt«ct«tldKa4\avaRV«ataiavu«
_j n(iw I
•MM uandM^ IMV4>aaka(!Milil baoaMfMU
aatb paMli'Mlaaa a* 1*1*, mnah a«*d vunU nadl
-SHUin /iv^al Jht-A, Har.lMa.
Gul<l« 10 tfi*l* lAacnoMa aafl tn^mtiM. \« \ «■»..»».
SA BaOTREBS Jt Co.'8
6BOSS, SAMUEL W.» A. M., M. 7>.,X£. 2>.;
P'ofaiv' ttfim'niiMm^aarttr^iu^itf OuttnUlK^r^ ••> (A> ^xfirwi ««iui*( CbUtf* *f rute.
A Prsctloal Treatise oo Impotenco, 8t«rilitT, sod Allied XHeorden
of the Halo Sexual OrguiB. >r» ruii) tidUkiii, uioiauebW revbtd bt K. R.
&mtoi«. M. 1*., I'lof. uC DivoKaof lh< licuiwUrlmiy (hs>ca mm M Venereal Dii
H. Y. rt«i Gra.1. Med. ScbooL In oop ver; huxlnaie ocuvo voIium cf ]<& (ofli^
with laiUiwiniMiix. C3Mh,tl30.
Tbn* •AUtanBor \'mtntor<iKit»' nltakitUiak
hsi* Mao vihkuiiwl. lUd ■UII tba daOMCxJ ■•
DBaiipptltd. Ifr. SiiirKlii fc*> (VTlcvd and addwl
to Ida r<*<^n>* *<'>*Mn^ and Iho aavanaappam
Eaat« complaw and men nloabla than btltonL
Faa> lmr«fMMBud|iaaar*liyml>UD4M*ioo<l>uii-
JacU ar* tnai'd— iiniMlaaM, Marlllly, ■pamauir-
ftuaa, aad pmtulnrrniaa. Tha (sok I* a nnmli'il
ona and in adiUiian m iha •alaouflo anii lary In-
MraMIn^ dtMiualnDaonalMogT.tTnipianiihale .
than ara llD*a ef (Naunant laid duau itiat any
fTa«(UI«i*r na fallaw and whlvh tiaia mal vlui
niMfaa la lit* hand* of aulDor and adilor,— Jftdi-
crt JtMvnf, Fab. IM»] ,
IthaiMMi tha aim el Uia aulhor lampplj to n
«oni|iaa< fnrm, pra^llral ami ■irlc*(ljr krivaita-j
l&ibnnatlonr*p«ola[lj hdnikitiii (n ih* wanunf 1^4
^AalVl pfMlllwnar la r«iCBftlioaclap<4orroniDi4>n
aad (Tnt* •llorim T^* mrk co>lnJ»< ftwf
mtaj (uru In n(»td la IW HXaaJ dtHTdm «(
naai, «f iiia iiiml ioWr*>lia|[ ghancut. Wa nun-
masd iha ttttif of It la tnij f*o(H*lonal man,
and aaixalallr u thoaa •uivDd lo lha (M*ral
pnetloBot niadl«ID«^CIa.3M. JTMn. Jan. IM1.
Tha oark brfura ua ha> Ixeenia a rtandnrd Uil>
Irsok on lha ^ul^^cra ot vrhlcli H traaiL In lija
t>rFMi>l adliiqii ih* aiiiliot'a vorh hia M«D«tQ'
■Idanbljr auKmanlal by tx. Starfla, wliciaama-
IrthuUun* and Tlnn ar« Is ba taan •taiyaltfra.
Thay contain Rianj' nioaMa annt^Ulan* maa ara
Ih* fnill n( a rip* aipaftoiaa irklnh aaDMil IxK
pnhanca lha OFlKinal MiL Tlia i-ntpvlnn ti
uulck 10 appraoiMa tueelnel irtalk*M ohlrh an
rtill vi<1 eompltta, aora aapaaiatly wban tba
4>iiiiinr> ai* kaoTD lo ba •ronh* ot ratpaM hi«
wnnjonof.-^ l«alaJf«landAr-J«''.F«b.>l,
N
TAYLOn, R. W.f A. M., Jf. />.,
JViff. ^ Vtinnal and An Oitauii tn r*t (Mii'i'iitu n/ rnwal.
The Psthologjr and Treatmant of Venereal DUeaaea. Indmllng i)i«
NMjl(««r Ncont iflTcatigatioiw upoo th« xilijcct. Itrio^ Uiv uxih edtUan of BunuMad
and l^lor. Entirclf r^wrillfn by Pr. TojIot. Ijuhp Stq. Toliini^ thtmt 800 MM^
wtLhkMullSOenKnTiiigii, shwvII m iiuini-r»i8 <'lir<im»'lill)ugni|ilu. /n ocliR' pnpapalt«it,
A ooiioe iif lh« iirerioiu edition is n|>))ciirJ»J.
II (a ■ aplaodld racord o( liooait tabor, hm* uipod lha mbjacu nf wLleh II (raata. Inl alto aba
ruaanh, fual eomparliun, nnfiil •rruitn)' and 1 nMcd hu neaqual Id otliirun^aa Itar Iti «Mar,
«TlalDal *tHrlani!*,«hlah Hill al'ay I* h>lil a< I i-ompiahauilta and cnnural k«iidll» or lla
• hiKberMllloAiiianMinnieillul lllrntiira TM- inrm".— dni. Jour, if (*< Jfaf. IM*w«i. MB. UM.
la Dol only tha ben vork Lu tho Eu«llkh iBD^u^ct I
CULVER, E. M.. 31.1>.. and JIAYDE\. J. R.. M. D.
HdnAnlMn BoijUtal, X f,
A Uanual of Veoereal DiseaAes. In one 13ina.T«luaia<ir>baat £50i«gai,
with illittlniiii>u>. It"3iiii ih'-rr!ii,
CORXIL^ r., " "
/V^B.or III Ib> >Vul(u uTiTaMM s/ JWt.^ a-f fkw^naa to (^ I«>«<a< itdvaial.
Syphilid, it« Horbld Atiotom;, DlagnoBia and Treatment. R])«dBl1r
nTiMd V ilic Aiitlior. nii<l lnin-Iiii<^< villi iimim. and addlilaiM Iit J. Hkmrv C. Siusi,
U. D., DpDKiiulralni or I'alhuk'Kit.'nl Ili'Inki^-T in lli« Unit, of IV, knd J. W1I.I.UN
WBtTC, M. t>, Lociuror on Veiirr«il I>iic«6c«, I'niT. of Pa. lo one fautdMBDe ocuvo
voliimr of Ml vagtK, wJtli S4 T«rr l>«iiull(<il lll<iHn>ii<MUi CUdli. 9374.
Tha aoaunx, lha hlitnlofT, Ih* rainolntr and , paniial wllhoul tba iMltng UmI hi* nap *t Ika
Ut* pllnlnaJ Ifatam «t (jrphllla ara rppr*HnW4 In I wUa and Important aiibjairt f« whtph H irMt* I*
•bit work In ihalr twtt, moM praeUtsI and mol a (tronicei and lumr on*.— 7>a JDaaioa iVain.
lOMMcUia turui. ui J uo oaa will rlM flrom lla i IHwr, Jim. lUX
CKkf nf Oiaif ■''nfrMl linrntmrnt, Haa-
aoih.
HVTCniysOKf JOKATHAy, F. JJ. S., F. R. C. A,
Qv*««-rfi'«j Hu'^tnn to rhc Lomlcn Hotjiital.
BTpbllla. In imp ISmo. voluiiip uf ^Ai [Higea, with S diromcr-lilliognptii.
$3.36. lit* Stria of ainital Man<atU,ygtA\.
ThotawIiD ban aaan mut af lb* dUaan and ' an.! ynintBl ohaarHtlen. bulef hb patianMaad
IboM nha hBH fall Ih* raal 4Moulll>tor dli^niv ■"•dTiliy Jii uhlan »>*-•* el bla «•<■■ aad haap-
■!• aod malni-ni «ll) moat bl|hlT appraclua Ibe , li>( <h*m ID a (<«m amlahla (or >ueh aKMllaDl
fK-i< asd ■i>i|i[f-'ti'<Di ahlch abonnd lu ihua iixia' ha haa put IMmlninlhli*alaB«.— Iwndm
fnifa. Ii la ■ ■'inliy and Taluahia tacotd, not 1 JliAai Aieard.ilM.ll: HVI.
onl^ of >l^. Hui4'hiLx*on'« Tary Urfa aaparianga i ^^
GROSS, S. />.» X. D., LL. Z>., D. C. L., etc.
A Practical Trontiso on the Diseaaett. Injurtoa and Halformations
of the Vrinaij Bladder, tbe Prostate Gland and Iho Urethra. Tliinl
edilion, (fanroughfj revianl bj Hami*CI. W, Uro«i, M. L>. In oai octavo tulnina of 6'i
|«g«a, with 170 illiiamiiiorni. CVdli. iiJM.
CULLBRIER, A., A BVMSTEAD. F. J., Jf.O., LL.D„
Jhrawif-IM AtpU^du JMi Lf,up'«f,,-f IW 0U. CUL Mm nadAvf.X T.
An Atlaa of Venoreal Dleeaaea. TmiHaMd aad aditad by faBuiAii J. Bum-
m-EAD.M.Ii. laeaaiii4MriBl 4(i>. *oluiiwom8pagai,ilniiU»«oluinn^wltli 36 plxe^
onnUioioK about IM flfarai^ baasllAiUT aolamd, ananj of them the aiae of life. Strongly
bound inclotJi,|17.00. A ipaduasof tliaplataaam) Icitaam b7 mail, <inrai;<alpletfSScU.
H"!kPJH?"l"»*'">''P<'*'-^«*T»oior«; FORMS ~or LOCAL pisRjsK kmxmw
i-BM UWrUKSaOir SrPBlUS15D SOMe.TluS. 1aoo««ioiol-ot-M«acik OMav.«Jk
.*8 POBLWATKW*— VMM
TAYLOR, ROBERT W„ A.M., M.I*.,
^rwan IV |«< /lafHrlau^ «/ r*i*>m( Md Wi* Uubmi i/ ikt Aiw fart V«*aM;
A CUnloal Atlas of Veaereal «iid 8kln DkMMU: Indudbig
Procwaui nml Tnuliumit. In cigbt Urg« lolio pwti, BicaMiriiii[ H X 18 IndM,
eonafKtinx ^ ixnaufuUj nilorod jilnta wllh S13 &(iiT«k *») 431 pagM ol uxt «ttb I
CBgnvii>g& Coinplvt* H«ik jW rwu/y Prio« per put, fSJHX Bwnd in one tnloM*,
half Hiusik, |£7 ; lialf Ttirkej Murocoo, f£4 Fvr Wt iy «it6wT»i<ti>ii n/y. ^p«ciai«
pbU» Moa on receipt of ID cent*. A full prwnKlm Mnt to my addrw* on ap|'lloi(K«L
TbceoniiMIaB of thU naaanioaul vorh l> > dMM (o • noUes K Uili hlM. U'l •'.il* «a
■ahMtcf asiuTUBlMlaD dm odIj (o Ih* aiiUot b*n BMIitu but h*Im titr Mis k4u>lnlUa alli^
■of paUUIikn, bnl lo Ih* pn<r<M*IOD u lug*; llttiu(b*adil la>*Mn(aUan ihain toMmUiM
ladvMltlf MUisUUcrlhx ItdlNoUyitiMiila M vkmaifd by ili* nwrlti o> tk* wark ~
k wDoittrfallr eleor •iikxHIod of ■ nntwMily nttfirftnl .V««t, Dw 14. IHa.
dinnilt linnch d( nidlEio* Good lltaraMra ha* tl woul-lli* larillu uh KunUaiilf*' annld |
Klnari h*nil> vnli nod ait ■Itli hIaUjr Mttuho- (rlmioualy aoAUCb mutst lo <ba i*«-t>i Ui* antt
r> rwulu for bntb. Than an dlAplhar lU talao nf llUa C'fninl ^iLw Tlila Ailw la i ' -
hturaa.nianycif vhlcliara life tlMianifrtprHSDi
IM blulioat Mrfntlon «( lAo ohtomD'lill)^-
Kimphtn>rl,aDdwMtaTad th(au|bout Ihauitua
HiDuin«nibU •atnrlosih (JiiImb pronoitlaa o(
Iheaa lltiinnlloaa an from lb* aalncm^ dkd
•ollacUau, oiillii od Iho Mbai htbd ilia baat
allaaaa of iha wiirld baa* bran dmn U|»>a fur
■k* meat l^pln*] and niroaiaifnl j.i'iuim at tha
naajr dlStnm Irpa* ot leoeraa' Bod aMIn d)*
MMl Va think m nu ■» irltbaui unduo
•iwnnuiao UiM 111* rvprndueiliiDs Iwih In oilui
■nd Ta blark and whlM*! itttr »Tmn*t litvnrlaljly
•uv<««riil. Tha Icit l> pracilml. Ion at that*-
paDllc«J ■UKS'*l^'iii'. ^'1 tliu Hlulcal >ac«udU of
diaaais ara c]*u and Inclaln. Pr. TajlBf U,
habpii)-,ui*iinlCi«ntanthiirtt|'tnhDUidapaffnt*iitfr,
and «• find at aonnuxiDpnra thai Ih* lw<> dliti-
tODJB k4 itit, workuuajatt an e^iuU aclanKAii aud
tlMnrr merit. Wa haioMrtady f««>Bdili«llinJla
OMBV)*!* •Tan Ihaa an MiHbmjt eiatai •( nlM^
l*i!Iuroi, fer In noaaaeaUcH av laviAal Mwn
U II al all |.n>baUa IhM aB dF tba dtMwaa hWM
r»['na«ata< vonld ba taaa. Il la aiaa mm* aa^
tloaablaM Ite mitftU at aMdasU Itaa Mtaafr
•a«a Bpan ollalcal Ncuaraa, Cw »dm at na
aluilanUKha all on i«a»ia aaaJa Ui (ba laakn
hall naonM ■•« Iba avb^avt aa wli aa Um (■■■
tludpnlDanciamlaafhaaalnja u»ILfb«fenipalll^
'Kmii'ik Conipulo) iha t«>l la a laMiiMr.B k
rnorfp AAiUfHciory In BiaetQ4va and ftilaaaa Itm
h* Koiild t« Ittoljp t(-t4tn tf lafiacvaaraaaiili
IndaM, ikia dUaa la latataalda w ■M/*»
anj praoUitoMr. fat II ««rtt»i ih* *)w ' . _
■•hnfolan Ut Kkaka dlaffaaala aif a alTn Mv <(
•klB mulftdailno b* compartiil Ilia «a«a •■
111" piflorp In tha ^Uat, viMra irlll ka mmt Mat
Iha tan nt dlaxiwaia,|«UKil<iC]F.Md MiMaUai
«o UaalaaaL— rirtfinia ifadtcS Jra<KM».M«.M»
ipii..
HYDE, J. SEVIlfS, A. M., M. I>.,
/VitrsK- ef DrrmtUbin and Vtitv,at JMataw) m Ku"> Mtditnl CbltaBa, Otiiitt,
A Pnctionl Tre&ttse on DIbobbm of Ute Skin. For lU
Pnctilicmon. N«w (»cojutl) wliiiou. In vat hudMine ocUra Tvlnmc
wilhSiMloml |i1litisjUldS6licaiiufilland^abotsMllli»ir<l>(>ni. n.iili. t-l .'
Wa <an hBtriilx fr-rrifnt^nd it. not imTy aa an
litlnlnUa UiMnok Tnr l>-a>:h«i ami nlii-lvnl, but
tlla tJaarand <4iopr«h«n*laa rulaa fat dl«a»«Lla,
•oand and Indapendani doctrtoaa tn paiholMty,
and IW mlDuit aad fuiHaloua dlmUlona lat Ilia
teaalaaanl of df'-i— •■- t mosl tail'lkaoty and
•Qn)Iil*Ca|ir«pl1<'' ' lifiihjrtlL'Iaa— .liaci'
mnJlfrmiUi^th- lihl July, lua.
A uarral itluauai ...-^i i>.liiii of Itrnx u ginn.
Tha il»»crliillt» i>nnnm4 at thli wnth ar* plain
and 'ifLlf nndvrntaoil. and abore all va f«rv
•ecuraiii. Tlia ihnrapaullcal fiafl li ahandautly
aupplled aiih aicallanl raaoininnndallon*. Tha
eoLura pan I* wall dona, TJ>a Ta'Uf ^f Iha laofk
praalttianrrt la Erval baciaiiaa otUia aaoallaniw
af Iha dparrlpllona, ilia ■uanatftvaaaa of Ula
■dflCk and ihaeorracluaiaurihadaiallaaad tha
prlnclploa «f Uierapanllc* Irripratwd npon tha
(•adar.— FVolaix JVal. VvatMK May. tMfc
MMor8tud«M>an
Inmcof 0711 pn^H
LI T.n i.-.ii,r, i&jdn
Tlia
r.Tlnli^al i ' .'
in dlcil'>ii. aTiii •ui>i'>
TIiB iba^tat un ^aaral n-mpM'! <
no olaarar daa
aod OOTUMUtlra laaloD* •< ■■n- -arfi lah
In It* ■ay :
pilniarT ao
be iiiel vtik laywhara.
la and ihcfapauUfa am al*« >on»
Thaa* tm aaaatat diana
« al*« >on»]r tt aaaaM
mdr- IM. Hr<l* haa abovB hhoaal) a i
hanaltaraadMof lteliMa(HI*raUira.*nlkaa»
oerpamtad iDta hla btok all Iba bcai Kiliai aliM
Uia laat vaara have braucM toiUu Tha ttauilr
IlDnaaDd fbrmtiiHtr* )il**u In boib «u«itaiwH
matrlo arauni* r>il and tllnatrailana af*«*S
and etdand piMaa of far* «aaaa land adiPibM
auraMionm, AllosHherUlaa anxli aiMU^aiiad
to Iha na>d*o( acananj mttDIUtanari •■4 aa mm
will maka a mlalaka tn purchaalnB IL— JMM^
Prwef ll'aaUn A'<* IWtJEna. UM ^1
FOX, T., M.J>., r.R.C.r.,awlFOX, T.C*,B,A^M*B.OJt^
n»*<flai> la («( SaoaifaMX riir ^Itia £afaaafL l^nnnftr Oi»aaia»rfa*<M«li«i ^H
(Aiirwiuv (Wf'ja i/aapiUI, Zo^ba. Kwifanoiir WtipJit taiMkK ^f
An Bpitome of Skin DiMRSM. With Pormuls. For SuiikoM aad n»
Utknen. Tliitii ulition, rcviMd and *tilait(0(l. In one li;mo.r<il.uf338|i|i. CUKfA
Ilia Ihlnt edlilua ut Ihln ponianlMil handtmnU ' ntannal (alia upon UiBtaUa fcrlaaiBBi ra*a«aa«.
Salla(brBollMouii<i;'"i''*rt'lplooanda>|auiiilc>ti , ItaBtnbatwtloal afrangamaal UtailaMi MiMa*^
artiieli It haa iimiccinne. The anancFmeni nf rkln tot all one haa hi koow lalh* aama of Iba dlaaaaa^
loalphabtticalctfiiar.ahlch lalbcmalhbd aad haia «« li> daacrtpabiu and UiaanmpiMM
npailoB adnplad la ihla onrk, brruRiaa ■ i in>aCni<'nt al hand and taadr ku iBatant aM<H»
adranlasa Is tha aludant Th* hsok ta UoD. 1'hr pratani adilkB \tm ba*B *vr* oarataOf
af elaaaUli .
toalUaa adranlac* _
ana Khli^h w* ou aliwiBlj rMciDmend, DOI oolr
taaCndanta bol alaoiopneililoDarawba raqulra a
OMBpandliHia aummari' at Iha aa«aant tUia
iannaulon^-antut UmUtnt .
Sratant •■UIMb hm baaa rwrf oar>ta>>T
a niimbat tt art dtaaaaaa an d*
Mrtbad, *hlla nioat of Iha nnat BdillU"aa ta
dwRial ih>nr*ullca Sad inofiUe«i, and Uie fa
OMBpandliHia aummari' at Iha aa«aant tUia of, _ ^
tarmaulon^-antut ifadMnl JsMwil, JulyLIH^ laixallha andnf thabookhaal _ _ _
WaeardWIr m>ommand l'oa> Xp^awi'toUita* lantmaiitrd.— Tbi JfaltioJ Aiaa. DOMmbar,
vhaaa Ume ta llaMcd and whu vUh a liandy |
id Uie tatma
■DM4*fal^
nnbar.Hn
m
WJLSOy, ERASMUS, F.R.S, _
The Scudeat'* Book of Cutaneous Uodioino nnd DisoMoa of the Skts.
In OM huuboin* miaU ocUvo votiinie of &35 |iae«*- Cloth. i-<M\
niLtlRR'S lUXDdOOK or ftK\« \«a6KSOM, \ aataAWM. Xjou^cuu.'ntanxtqt IttHe^
faraadeoU an4 I'rwlllttMWta. MonA VwdV \ -Ma^i^tUm. <>>.V.^-^
J
SjMA BuiTBnu k Co/fl PuBUOATiOKa — Dis. or Woaicnj
The Arnerican Systems of Gynecology and Obstetrics.
S^'HtuiuH ol' Uyniioology and Obmoini?!.. iii Ti-cbIim>b by Amorican
Authors. QyaMalogjoliKtIiiT.MAT-riiEwD. MjkXK. A.M^M.D^rnnrawtofObctslna
■ul UrnMoloxr la Ui« Jt«ti^ul l.Vimrtiiimt of t)i4' (jDiTtniij of BuflUo; Mild OlwCM-
rhs Mii«l \ijr Bastox (.Vmkk Ilinn', M, D,, A«ociai« I'rnfasMr ot Obttetric* in ih«
(jnlfvnllv ol rmwjrlvnniti, I'hUaiielplita. In fMir rery handaoiii* ocUro TotiiiUMi, tea-
talnloK :^>M vngM, 1092 cnftntvingii nnd 8 plates. Cosiiplol* irorh ••■ rtadf. IVr vol-
une: 7:iolli, iS-OO: tcMlicr. t'iM<; h»\t Ruw!», t^.Ott Pitr mU hf tuAKn^um onlf.
iMrat lA* P^M^t. f'tll ilnoriiitivr pirrulor fnv on applinliou.
LIST or CONTIIIDUTORS,
irit.l.lAM K n^KliB, H. V,
ROOKKT HATIKC, M I'.,
I UMCKL. C. I»t ■<KT, M. I),
JAUKH C CitMKiCl>K ». Iv,
I HKNHV C. CuiK. A Jl_ M. D,
RlXVAIIU C. liiVlB, Ml'-
iG K. !•■ ti(niWKIMIZ.ll t'.,
E. C. iJllI.'l K^ » I' M P,.
B WrK. FV
liKxH'-h I .SS,M- P,
UEMIT J u .. :.s. A M.,N. Dh,
WILLIAM 0^>1>ELU A. M . M. [■..
BllRKKT H. OKASI'lX, A. M, M. D.,
1 ri,,
■ II.
i- ■ ■■ . I..M.B-
\. mZv- IX,
KIiWAbi) W. JKNKb. M P . LU tl.
_jiliuat ■(•nltcmiacC lli)> founirx li> il>m d*'
paruaenuonltDpKdBaslOQ- Kw? hooT1trl^^la^p^P'
I•M■■ iB«Basn>|>h uf»B hbaparlkl Mpla. mi^i
•TnyWIaa id Ihr tnj dC IiUMit. Ui»rT. mBlhoJu.
■d4 mnlti If nrnvowil lo our nttlHl nwH. A> ■
•«rt alt*n»n!l rah no m, ll Mtl tw (Mind rMnnk*-
MylUII knd iBiiriictlts In •t*ry dLnrUoo W
tDqatrj.— Tm r>MMn<OaHtl',H*|<Minb*T, IW
Than (kD Iw tiut lllilo ilniibl u.u Uil* •mtfc will
•sdlhaMHic (>iat wlUi lb^ |iror»«lon llut Iih
la>D aroimlfd U lh« "fiyawm ot HsdIelDa by
Amtniui Aatlion," aud ih* "8T*(*m ot Of om«1-
XbyAnivrlcaii AuiViorH.** Onvlt»iitliM«loka<Ftt
I loAay <if ihii rolunir. I'>t r>u iiiM Jail and
marlMil pfalw mi]>t<> ml-ti>k*a (or flailaty. Tba
f0»noll>n. Enjittinann, M»nln, lllnl,Jii|sard
•■4 B«a*« ara lawmpanbljr l«]rODd anjUIDa ihM
(■Bbafouad la ubaWtrieal HAdU. C^rtttnrriba
EdHor mar b* coofiKlulawd tar laif ag mirJa aueb
a viH wlBMlnn nl hi* matrlbuwn.— JMnml qf
UUiHrwar Jlfal«*l.lMtM«i^,Snl.fiMI.
la em kAIm of Uia "eyMaBi ot rraallcal H«dl>
•iMlv Aaitrkan A<iibon,*<i« mada <ha follciii-
iBg aUMniabd— "It I* • wntk ot vblPh Ih* pIO-
HOWARD A. XRLLT. K !>.,
I'HAKI.Ktt lUIIKIII.I. I.KK.N.Pw
WILLIAM T. t.rSK.N. I'.LL. D,
J. )IE\LiItlE LL'lTlJ, Jl f>,
JCATTflKW !■ »IA^ V A. M, M- D,
H Nf.u'Kl.L HAwnS. r. ft. B., fl. 0-
KICIIAIU' IV HAl'KT, M. U.,
V. l>. PALMKR, H. D..
HOSUEM, P*hK, M, tl,.
THE-'l'lirl r> CAIlVtIt, M. IX. LU Di,
K. A K I't-MlueK, M. K, i.L U.
THAIH'Kl-S A- BIUMT. A. M, M. IK,
J. C REEVE, M-D-
A. li. Kuc-KWKLUA «.>!. D..
ALSXAKUKR J r CKKKK. ». IK,
J. LEirtBBHITH. » II,
STBPHEK SMITH. U.I>^
It. aTA.SORL'HI HtlTT>>!(,M-D>LI^IX.
T. OAlLLAKli TItUMAIL M. !>., U. D,
ELT VAK PE WARKKR, M. I'.
W. OILL WTLIE, «, D.
faul-iti In (liU roiiKtrT nk (tal prnutt. VrMlaa
•xrliuliafi by AniarlPaa i^iytlalwu ob* •>* a»
«iiUDl*d slili all iha rvlaitM it allinaw M lb*
Coliad 8ma». ih* obanswr of lb* •oil, lb* mao-
oara aMd rmioma (4 ik* p**fil*,al&. It li iHmil-
latljF adafitnl hi lbs iraAU nf ADMrlcaa pni'iMaD-
•»«( in«>l(PlB*,aBdI>HainatuB»Ui*l xarjruia
at IbeiD HiNitil dadn (o hai* IL~ Ktary w-rd
lhu> axiiniiMd In rMwd Ia Ui* •'AnarHMi ^a-
lam at I'neU'-al McJlela*" la aiqJIoabto (a Ota
■■jnarm ot uroBnlou by Amarloa Autbora.'
II, Ilka tba Dtnar. luu baon vtlupn •loluMraly
KAmoili'aD fhyalGlaaivtMBnaoquaiDMd ■llli
lb* HiarvunaUc* at Asaf lean peaiJa. alia ■!•
■oil ■•toinied in n-Kkii lu iba paaullwllloa of
Amari«an iriiaian, Uirir maimtra. <«»b>ma. tmsiw
ot IWInc aW. A* rr»r]i I'racUtiac iihyilalan la
oaJlad Dpon lo Ifaal UtHuai ot frmal**, and a*
(bay (ciDvltlDM a fla-> lu ahlcb Ilia Aaillr phy-
■Irtan muat fli* alMouon. anil CMitioi uu> otat
Id a apaelallM, «■ da BM kno* (4 a aotli to any
dapan»af>( of nadlclBe lba< n ikould MaitaoKly
taeaniM»Dd raadkal oiao eanoMly uutvliaalua.—
Oacpnali Wtd. VaaiJuly.ltn.
THOMAS, T. GAILLARI), and MVND±, PAVL F.,
M. »., LL. n., Jl. D.f
firrftutrif DUmtm tf f—^ •< <«• CM1<«< /Vi^biid- ^ Oyaiaabn In li* X(« rorl
APraotioml Trcntfno on tbe Dlseaass of WemoD. New (tUth) edttUm,
UMnMghlT wvbcil in'l tpwrhlon by Dh. Mtxdk. In one laixo nnd baiidioBia pctaTo
volume of aboiitMN) (AM*, vitfa om- 300 illiHUBllaaa. Stad^ tl^ilf.
Tnt. TiioKAlf ■Unifatd weak En the origfawl lest ami Iiy tu (nnclatinna inlo rthtt
bjiSiitgai la aTMyiftieiw rcRotdcd w tbe b«t wpiawiMaliTW of Uic peculioj ly AniaricaB
■d«K*orKrDecni<^. Ill iliia nrinon tiMjofnt aiiihon bat* owWrtakaa M nrtMm ib»
fnctlcal nturr nf ibe wnrk wLkli hu rmil«r«d il ao uwflil (anadcnta anf pli7«kiaal,
■dcncvorKmecn^iy. Iii iliia nrinon llMjofnt aiiihon bat* nadtrtakan M i
fnctlcal mturr nf ibe wnrk wLkli hu nf>il«r«d il ao uwflil (anadcnta aail .
and alia to bring it alimU wilb tli« btat ^yDetotoffial vi«w« anil pneticc of Ibo' prcacal
^j. He ihorni^ rliarnrl^r of ibt TOTiBon mn l-o iimWritixiil fniio lb* fart Ihal t4 Iba
itluHnUDiii nno litimlTttl and KTCnIT-lti-o an.' ii*w.
EDIS, ARTHUR IF., Jf. !>., ZotmI.^ f.B^CJ*.. M.Jt.C,8^
J>n>f 'ifrirtiii rHimialll In ViiKInai Ifniitfil Iilfl r>)iiriai tl ffilWl^ f|lla>ai HaiBlllit
The Dittoaaoa of Women. Indndinic U>*ir Patbokcr, OauHUon, tiiiiin*iiiin
DiatpioMi aifcl TraitmcDL A Mnniuil Gw Stiidmu aad PnctltMiMa. In one liBBilaaraa
ocuvo Toliiinii ot 5Tii riu{«i, wilh 148 UliBtnUlmK. Uolb, fSJW; ImUivt, HOD.
Tba tfamliJ qoallilH olilrt: ara ooniidtnMnia
■ra iboNniRbaaaa lo ootarlni t^• «hula ipwiiid.
plnaraau ut daiwrlHIo" "'■ midlwnaM oi rtaia
DMM. AooUiaT nurlied Iraliu* Ot iba bosk U
Iha *«lanilio palil to tba ilatalla t4 any mlDOt
avflnl opprallon* aad fasotdaiaa, it. tar
teaSBoa. Ib'a oib ot Mata BM««llon «f laaobaa,
~ a*ar lolaeUaiiih TMaa
•nont tb
maot, ud
tba noiv «aaiBwB mMbada «t traat.
maot,' aad ya< Ta<7 Uttta t> aald abaiu ibalD la
many of (M taibtoaka. Tba booh la ooa to to
wwnl* ratonif dad tafai^aily lo aladaau aad
praenilafian, vho oaail a aiocIh to) «i«i.
plalar«wa^otUia*bala>ali}fVL SpceitlMMoab
wUt a»d many uaahl blow lo Ha ,
X*£ and Svy. Jaom, MaMb ^ INML
26
Lu BftorHUB & Co.'a Pdbuoatiomb— IHb. ol' Wobmoi, UMwt}.
£JCjir£T, THOMAS ADDJS, JH. D., LL* 1>.,
The PriDolplw and Praotloe of GTiwoolocr i FortlMtiMoifattidMUi
Pnetitioaeno/ Ueilldae. Nn> (third) edlilan, tboroufhlj rvrlaad. In uo* l«rn«id f
butdaome octara volume ct 660 jNgoL witfa 160 Ulortntiuoi. Clulb,|6{ UallurJ
*tr7 haadacnw hitU BumU, niMd bandi, f&M.
W* m hi doubt whHhcr i« mnirwalue ths , Iba prlrllMI* *>>■■ «ShM Umoi •! rMadaa ife*
■uUiar nor* than III* prefmioa ii|>«n ih* *pp*M- tlBnwt>lpmpitM«(iA«wun<H. Hm wnwlANi
aiMof iha Uilntt-llUoDirf ml* w*ll-kBoirniir«'k. l«l ^tV«M *B< oaaWtooMMMntH ir* iki&lhal
nM]PUw.MII<lwa,t)>* Ilf^l<inie(p»fli<n'-i'n( fI>|ltHna4 talyhli MlvMn*! ■iiwrUn
«M «b«lkHCOI»piiniinu>]r4bt[uitulUioil i < < <i<-Mi>n tp nfnnagi IM ill lint »»M* <
■* • laid Md raefMnk) ap*rmi"r, uu.L » : <ela*l (tlMM «Ml UL— ii4M* . ~
dtlDIW) M miMh ■UcaUOO InWa Kiicrui'. < , UUP U^ U&
MIibiwUm pnlWinu vtll DdiSilJio ftpiin^itle.
> of inv
XJJT, LAWSOy, F.Jt. C.8.,
Prtfttmr of Onarctuav »> Qkmh'i CMhtM, JIf iii|>i| lot* JVirtiwI xf Ik* A-MM 0*«r
DtMMOB of Women and Abdomioal flurgor. ta two mr luMHtMot
octATO TolutMs. Voliune I., 6M pag«^ tfZ «iigni*ingi iM 3 pUlM^ Cloth, f3. <V<w
Norfy. Voliiiii* II., prtparing,
Th* alw «( 1^ »rk doM aoi IndloM* lb* r*(tt-
I V »yii»Mir> <rf ■ tKil teok, Mid 7*1 B*wlr tniT-
UilO(*tdlHMe lanatBlos to Ih* tiiHom arai*>
■BMrnaftbir canM4*nllte. Tho dwOtpUoncf
dWMni «aD4ltlsu I* axondlnilr diMr, mJ th*
tnaUn^Dl, ni(<dlcitt«rrariH'«I.M nry MlUMletj.
MiMti or tbs Mil If atnarin-i'*- '
rau*, wblcli (dd niacin*'
|^• tuM**>*d fi»a* ft ir*>i
B^M llial r» KTHMttcliid ' 1 .
tut la Mtr* Ilia wnrk la UibU
(AtMrM V«"l'. M>r<ib. Uaa
,> -., (lU
libnnaa.— na
DAVENPORT, F. H., M. D.,
DiMasM of Woman, a Kimual of Non-8ui-gioal OyoKcolosT- D*-
dciwd MpaclAlly for ths Um of SlodMila unI G«imc*I Prtctltlociare. In on* SaatlwM
ISIDO. tdume M 317 [«ge», wilb lOS illiMnUkog. Cluth, fl.SO.
Wa ajpwa «fiii iti* aaaf M*l9«*t* obnta a» ' S**«IMac !• of ih* im*o*(, (raaban aad noai
tlo*>aonBrat*adIao(harJ<nirDalieoD| " ~ ' ------ '-■ -
Dr. ittnuforl an Ilia MJOaa whlol
■lulnad. lis hu irMd u «ttla a book (or lb* author
ffiidtnt and ((orral praeilitoot* Hhkli vonld
leoDcrMuUillai; , uwiuMt, •« niu«h m that w« hara weonnnwidtd
whloh ha haa lllaow dowaf K]roaMla«y aUVMuUi. TbalUw
adTlMi ' "
Mil ihem >uii sliai Ihar oURht to liao* wllbaal
dIttiacUBB ihair altaDllan allh alM«f onaplla-
llan* for ahlrh iber fould bara ao (MMalbla ua*.
In IhU ha b— bun am inanity lutVutOL Thar*
b &M *f*ti a p4ni|raph of UMlaia BMMc.
Mt Is llM wwr el uwMieani Iim all
baea BnKtIeallr M*te4 br htaacll; asil ■■cb
■nthou racalia* onl^ n BMiah aoamMMaUoa ai ba
kaa fanod Itac It daawTaa. W« w« aara lUi
ihaM «o4 aaalhlifa nlll onnMafiil lltr 11 • laraa
tala—OMda IMtml llMord, Da«. UW.
MATf CBAHLES H., M. D.,
A Haaual of thoDlaeaaM of WomSD. Balm * amOm »bi1 truanutki txf»-
rition of the Uiaorr anJ practica of gjoeoJIiigy. Kmr (3>l) editioo, «lttM 17 L 8. lum,
iL D., AHaodinK OjoMnlwIit at iha Hiirlatn Hotpitol. N. Y. In mm ttmoi. voluna «
SOO pagw, iritli 31 illuainttOK Clotli, tL.TS. J«H raady.
r«pMI]r bal bu aet ika Una la •nuaali Ur«ar
iMtkik Wa ara niiub MnMk vKk Iha raMlaaaa
Bkd owianiaoea vlUi wblsh «*a an rvhr m aar
«uHaM ruBiaload In ikit Tolnna CiaraAitly aoB-
Mtad indaiaa and anfta lllMimlaaa abwaarMll
Uie •orK. TblBMaawl «iu bajMikU tuMii Ma
SUfMMa Tarr aaililiaMorUr.— fka nyittHM 1
urytM^JBaa'HML ^^
Tbla la a manual t4 cKnatfolocf in a very cQiH
4>wa4 fonn, ^uil iFim Cii-i ilitl • >SMnd adllta*
kaabaanotlfxl (i>r iiiili'--ii*> ihut 11 hu oMtirilb
a (il*«nbla m.<>i.ilon. It I> lut*n>lfd.llM»uihor
talla mt. lA Bid lb* MOdanl who aflat bavbac car*-
Ibilv p*rua*dlarKar oock* daaira* to rarlao Iha
•nl4at<,andliaa4d*th>u Unayba u*>M 10 iba
finoiUloaar i>h« wlihaa to latraab kit atanoijr
DVSCAy, J. ytATruEWSt m.d., ll. d., f. r. s. e.,
OUnicoI LecturtiM on tho Dia«aMa of Women; l)«tir«t«il in BaiM B«r>
ibolMuvir'i ItcMpilal. Inona liuiilMnM oclaTD voliune uf IT&pogM. Cloth, |I.S(X
Ttaay ar* in 'tDiy oay •vnlir of Ihsir Butbvr:
la>t*ad, ■■ k«k ups« than a* amonK Iba aaaat
nloable tt hi* canuflnUoBiL . _
mallan ofcrtal lotaraal 10 Ihaaananl prvlli
Thay va all uaoo
n of Krtal lotaraal 10 Ihaaananl prvllUonar.
oribiimd*Bl«ltbMJb]*ola ihaiara na^ua
riila, Bda«uiit*4r baadlad la Vtm laKt-luaka ;
of itiaoii abu* baariBK upon inpln> Dial ara t
IftMaaof Bl lannli la iiirii •"(k<. yi l>ar 1
of iBdlrCtMUlj inai mer da»m
IMl— X r. JTadwW .bHniaJ, H«r<^ I
HOIKliE OK IX8IMH MtCVLlARTO WOMKK.
lBt>*dla« iilaptieainantaoriba UwfiM. iiaoand
•Lilian, tVTitad and aalarMd In oaa baaau-
Inllr ..r."i»i r.-( = m ..,iiim. .,f 111 t>a««^ wlik
rUi. .■.-..:.,.., i.-l'JlXK .IM»
&t;iK>i:itV. lu rcltcauua iv Um Proeaw «<
Fanuiillen. A Daw anJ aalarBtd adUlon, thor-
OUaUy nti»ti* tn Itia liithnr. Wtih adqllUtM
bfV. V. Sutoo, H. t> ,Vi«(awM «tUM«WM.
alf..tatha UBlntm Madleal rvdlaga af pbd^
datvMa In ana Ibth and tkAn4*dnia liar«ri^
oMBto nlaoia of nil i«t|«i. ami n« full paa*
rliUa*wi4« «om1«<ii* ' '. I'l" i--ii.«n(alaka«I»
all aurly aootiaauiir -iiraasly biaaJ
In laubwr. vlth fai^. .
W»rrit UKCTt^'Ilt;'! < _ MRBAHM Vt
WOUEX Third Ainaili'iA i/voa Ih* third La*-
■liaadllkia. In una oMara Toliuna of M1M«&
C'lol^. t*.'y. laMltar, HTl.
Lba Bkqtuxks a Co.'S Pdbucations — Midwifery.
PASmr. THEOPHILV8, M. D., LL. J>.,
The Soienoo ftud Art of Obttetrioe. Nev (2il) dlittoa. lo on* hudmn*
TlMMieaad*dltliiaotltiliiin]rlII>rullr uptotbaiabMnrkllurMoneu ta tnikd kooik ■bkli Ik
MHntdaUor adnDDMiwal ct ()i* obaUtrle aiL 1 te BompnhanidH nod j*Keamf*Bi wx) |iriHil<>*L
Tb* wMlwr hM *oc<>**ilfd *u*«<dlii(>]r aptl Ib ' In ■tn^h iw>p>i^ Ici>h»«ImM|'bhii bank »rihv
flni mritt Tn* Inalngiil ofth* (UliiMUilmi
tothor hM *oc<>**ilfd *u*«<dlii(>]r aptl Ib
iBMtvoiulDK nSK msltcr •Uhniil wpurvDIljr Id-
MMMhK LbBsiM of kl> oork oi iDUrfviluR with »■) lATas lo ilia v««— Uia laillrliliiallitM u(k
IbaMnoDlliaua ud EntrsoriUlltaiHy '"'"'riiti- T<raciji.-iil trvtiK. • ikllnil ebaWirMtiU. • siM*
ifii - - . . . . — . . _
Uao. II* I* ftf Itlfctuniw In hU d*ii'r)|>t'BOi< of
(Midllloos. ud pKii** hiTnMlt tn (nl* f«*tw«4ii
■ckalif ksd ■ UMtM. bnly tn tb* nsg* of
ililnk*ruid * rl» •cbatu'.— JV*dual Sio"i.)Ut.
IT, inl.
PLATFJJlt, IF. 8., M. »., P. », C. P.,
^ t/tnar of OftiJiing Jf ilnwM m AW' OiKiji, Lon^n,Ut.
A Treatise on tho Scionoo vnd Prvctico of Hidnriforr. Ne« (GAb)
AmxHiiin. fnjriilli»t»vv(ith Kniiliah rdiiioii, Edilol. "illi u.iililiuii-, liv Robert P. H*b-
mxi, M. X>. la on* \>am\if>%n« [k-ioto votiimc of M4 |m{[». wiUt 207 tngnvingi aod &
pUto. aolh, 14.00; Icmhvr, £3.00.
TruTy * wnndcrful tio'>>L: ad ppllom* of all ib ' linn frtini th* ni'tniaai cT tfonwitlun to ih* II nM
•IMriMl lisri*l>d|ir. full, elMT add morlir. lo , u( eomr'oM latuluUoo liM hwl tha •ulhnt'a
Ihlnrta r**'< *< ha* R-a'had HTrD vdltlwik 11 . t>iUI*nl allBDtloD. Tlia plaU* and lllualnlluDa,
UpMhapa IK* (n«ti inpularmrk of li* kindvnt nan-rvIlT ntuilttd.Ktli ivvh it>* rei*vti* at mid-
prill nl* J In Ibr pronHlon, BBilnaliis «l()i In* otforr. Tha milaraf tTilsbnok «tll hat* btifM*
— alriri]' and phynioIuKy of ihacrmuaeeauwrDed, ' him ih* »ry IMHland t«>t uf olsuulc pnOlMy
MUilst !■ l*n iioafltMa that Ui* prastuta) ae- 1 and alaa of all <h* nolni'ldsnl IrouMa* mODaoUd
«MOli*ar ihnnid kfiQ*. It loma that •nrv tbarawltb.— SSuMm JVoditivMr, Uao. un.
•Ivkbla phralolDBloal or ptthologlcal taodI> I
Kiya, A. F. A., M. D.,
IV'Jutur r/ f3t*lilritt mil AuMwaqf Vmmkm Ot MttiaU Dtparitnml of a< ai—iWan r/a4*(r>
•uy, H'oiAinfliH. A (l,«idto(ki Danwkly of r>niimt.<H.
A Uknual of ObKtetriua. Ne« (ronrlh) edition. la mm i*rj handiDaie 12mo.
Tolutnw of 4:i'j |iH|(tK Willi UO illiiMTAiiniu^ Cloth, (2.&0.
K>r. Rliig. In Th« pr*<fjK<* ta (h* Drat tdltlon af I (1)* daapflnrl'in or labnr, hnlh normal aiid abnor-
IhUmuiuI. in'j<l<»ily ilalH Ihat "Hi purpota I* , mkl, thai Ur. KlDt Laa> hlalwM. Har* hit alTt*
10 buDlali a (Vud KiifU-ndwork la th* iiiijaat al i ta au (unnUa, anJ iha Uliuttaiiuna ara an itnM,
th* ttaclBotoi nf hiaotawirlc nuiliaa." lu pur- (hat ih* **ri**ilrraeould nollulli<>r*ci*tt*aolM(
foa* laatMlxdi H iMU firniih ■ i^wd (toaDd- eoDMplIno of Ubor, til MnpttFillnia and lr*a^
oaritlo ihe (ludafll aim '^arftuUj laada ir: and mant. Ot Iha 111 lUualralioua li mar ba aatti*
* " ~ ' >aij ihai <hay atliltkufraitianditiatlhaaaKtmHri
fUftliar. iba tiu^y pruriliLuniT thfiiiid b^i tc^-tn ihi
folam* i^raii** wrlrian tor »i4idaf]i>,** ii •-i^tt-
lala* mucli Ttluabtc Qlwwi/ta koowladi*. piin*
•f which la DM (bund In rjora alaboraia laii-
Moka. Tha chaiilan ou Iha anatumr ur ih*
fctnala ^namtlr* orgaua, in*ni*lnja1lon, frviiuda-
[■Mli^ ^uvia«««« Ilia*"', ■■■«iiri4*jaiiuM,
IND, th* atria or prtRnanrr, and iha dlaaaaaa of oal Htut, Dta. T, UN.
fMfMOey, an ail aiotlltDiand elcat; built la In
wnrk !■ aicaflani. Tha oam* or 1A* puhllaiiara
L* a auffl«ifDt |u*nnt*« thai Ih* aork U pia-
Hnied la an afliractlTa forni. and fpotn *«a*j
■laDdfMlDt *•«» noai haanlly lacununasd Ik*
buuk bslh loptartlilonaiBod •tudaol.— TM JfaM-
BAJeyss, ROBERT, M. />.. and FA2fCOVBT, M. 2>.,
Mw. u> iKt aawrol l^tft* JfMDs 1<<«A Obidfru Pla,,. ta SL T»»wm' Btty^ Lam*.
A Sjretem of Obatetiio Hadiotoe end Surgery, TbooreUcal uid CUn-
tokl. For Ilia Sitiilmt •ml llie Prartltic«i«r. Ti\» Nartiuii «n RinbrrttlacT br Prof. lliloM
aotE.t6: k-aih«T,|fl.
Mm«1wU.
TM Imniadlala puti»iaa(il>a work l> lo (litoiah
^haDdbo•k nf ohaU'irli' in^dioiBa and auraary
tiha IIM ol Ih* aiuilrni anil praiTi lUoiMr. ll It
Dot an a.t4K]i|Brsilon lu >*v of iha l^ak mat It li
Ika baal Inalti* la Ih* Rnglli-h laacuac* )-at
pubilahatl, and ihla irlll noi b* aatirprtia lo ilto«*
«te ar* Ki|iialoiFd allh Uw vork at th* aldar
Banw. Eiarr praclUlooar aha d«ttaa M ban
In one 8to. voliime of 873 ppL, with S31 illiulf>tii)Tu.
Ui* b**l olxWtrlral spfolona of th* (Ima In •
raadKjp aooaaalM* and^ooiidraaad fOfin. onihl lo
nvnaaopjaf tha bMH.— >/*unH' rt tkt imirta»n
Mtitaii iMDoaUaa, Juna It, ina.
Tha Authon har« ■■ It a laivkook wtikh l> la
ann a'a* qntl* mrlliy In Uka a ala^ haald* lb*
l*at irMtlaa* «C Dm »ano4^.TM Toik JMMal
Jutnal, itOj X UT-
F.
WiyCKICL,
A Comploto TreatiM on the Patbolosy asd Trvatmesl of CbildtMd,
For BUuUaU and PnditHjiwni. TrvotUial, wilir tha oiMMnl ot iJia Antbur, froa th*
■MxadGcnnaaeiliUui, l>T J. K.CfiAi>wicK, M. D. OcUfo «H pag«a. Clall^HOa.
ASaWKLL-a PIUCTICAL TRBATiaK OX TRK
DMCAtas rSCULIAK TO WOMIUt. ThlKl
AsMtae bom tba tkM and iwna«l Laadaa
•dWan. lDOBaan>.M.v^MDL CiMh. CUO.
PARSr ON SrnUrOTRRIMK PBnr.XANirr:
lia ClUIOil 1(liMf7, [ilacnoilr, l>ii)cn<»t> Bi,>l
Traanaaw. (Miati^mraiah aoiA,t>Ja
TAniBR ON PRBiiNAXCT. OMara. 4W |
nit and tlataa^. I* «Ma Ololk.HJi
OBHaOHILL on TBB prxaPKRAL rBVKB
AMD OTHES DUKUES PECULIAR TO «•>
MKN. Inonarra ad nrtMiaan. ao()i,|tJ>'.
WRI'MCSTHK NATt'HK.SI&HaAHPTUAT-
MSdT or CHILIiBBU PRVBB. la Oao Ira.
i«)ra*otMIH«aa. OMh.lUa
80
Lka BaoTBBiu L Oo.'b Pdblioai
.. Uta. ChllOu.
SMITH, J. LEWIS, M. D,,
amuM fvyMc *t OuMMi tf c»MUf»w <■ a* ttm^m BffMi juttuu cwi«t«, Jr. r.
A Troatiae on the DiM«*M of luALnay uul Obildhood. Sew (mtmuIO
•dllkm. ibomuitiljr nriiwd uid r*«rlura. In oo* himbuiiv ortsvo volaaM of ttl
p^gM, witli Ql illiMntlon. Clolh.HW: l«Mli«r,>&«l, JtMrtadf.
IC*wy 4*p>rtDi*Pl (kaw* IhM tl !•« bwo Ilior-
«achl]r niwd. ■nil Umt aTMjadiniiuinlMBbHB
Uk'onrnnfMkdTiiBn M kniMiltdn tobrlnsd
<aair<*icir<ipio itiatimM WhBtiniikH(li**Mk
tt Itt. Smhh <4 at^a^al nl«» li Uia ■imdUod f«lil
MilMCucHlxand l^• nrtAil daiall of IrMUnml.
(* Id UBd(iaM*dlr ana a( ih* baM iNaUMii on
(klUtH'xIlHuMkWid us lati b«oh brsuubM
«r»4 tPiMa fur yonnif BtntlHtoXCT HhuangrpaiiaiiiL
— Aif-fu Jf*rl>«l ah) MiralAW rraniat. Jan. ml
Atn>»l]r In r»*lnn> sllilna* iha aniii of Dr.
tiniHIi nald puaili»B UB4H|fliwl al <t>r brad or lU
cteM. So bwk la any taiiiiuaf;a dduIiI ilwpuu
•UlttlW«Ult« lu praMTilaaaoK A lt>( of w<irk-
on dlaanaaacf ohllilraa, inada up In anyrvuMfj,
•OMldhafa Uil*irwKi>ilMhn'l,andfaiUa»|iair-
■ain «4 Uia snal nm/s-nly c( practKianan Ih*
lM*a■■d^a(«(rlplaI■■ vmhlhMonaalMi*.— diurl-
M* rniMMeawaid .Vw, Kay *^ <*•■ .
Nol««lk«laad[ua tha nuny amlMM nlama*
lit ■■■*• liroii [aaiiiM iMantly uo dt*— Jat of
<Ail4na, Uw watk (d Ifr, J, Lnt* BMUh
hold* ■ 6«*l •lata. Im (BMial *4IUaa* ka*
bran tftvouidkly OTliM. In fapm— d»w
ooUM taal IKtar n( Ui* i>lia|rt*n hara tiaaaj
llfaly TwrftlM^ Pull naUoa la Utm Of i "
raaml adiwio** Whi hat* haan nwJaL
aulbor MMM Is Ih* •••(»». Ul4 ■.Waa.afy ntii
kMTMiMllvrradMaiaiiBnaMk. IuUmm
of IniMKtllaa rtwMMd Uia WWII UMy pr
na Mintldand is Im«« 4aaU«4 fn lUa^ (
eondaDiMbHi ud Um aMlnalaa nr all Ma
i:.i.'>>fi.i 11^ tulsnia hat oM •■•d tamtarnd u
■'•.f- Maay illi»mi »M prath
iiai* noalrtd apMUl ahnpwiB.
Tfry laaMoal «»r. &iia«h) ran]
Muu loktii IliM (lia dLrafUi'b* lor tnaUBaal i
M HiUcntarand tiili. la Dooihtr aotk »»■
*■■ - •MUIUDllnl
iiu iif l"(anl&->
va pafucantrana li^^i. Ja ooo^rtar
oafifnl latucipltod" gltm 1a Iha A*
S«lana (Ml Iha anlfc-laJ ImiIIiu
uWrMl «<««< A-rW, Fall W(
LEISUMAy, WILLIAM, M. J>.,
ASyfltonioflCidwiflnT.IncludiiiKtIlo DiMMUwaof Prevuoiioyftad tiM
X^erperat 8tat«, Frmi-ili «iiiil«i. Ocuvo^
LAXDIS, HENRY G., A. M., M. !>.,
I^tfutartf Ofiilttrtti and lAi IhMiittiol Vtmm aiMaXMp Jfatinil Oullat, O
The Kanagemaot of IiBbor, and of tho C>yiQ«-ln Portod.
hMidMiM IZiiKi. ralniM of SM |HW«n. *llli S8 illuMnuiuoL Clcdh, $1.76,
Tha author hm dmitnad M plae* la Iba hi»d>
aftlMyaux prKUUcaara hoik la ahuk hf oui
■nd aae>— ry lnlbr»atluD In •> lulant A* fttr
aam oan kh. naUilu l> '<»'iii.,
aonnd. and lh« ^■■caduii' -
C>f>lr4/Man Mr OmwUAt.'
Thl« la • baoti ■■ on
Tlia auUiirf ROH nnch i»rts )<>
dMall* of llM naaaaamval ul
■aat-hoaki, and It as laaJaMn i
''■• adfloa it
■ CWM. .
>-,^.m.iaiaa,
r" Ul*
man
' I «> la
Maipl any asa wh.
tba bD*k t» tu
■uiyana • Umi
anil IM* amula^ '
lautk araryihUK
lyin^'la laoiu. ^<
lyinc'la laoiu. ^< •! If li !■ a* a
teaiTM II dftnL-. . iL> miun lu fm
«MMffe *tM«lo» lu Biwial— .Vm Orlaana
•alabl AiiMiialJawiiil, Uat. KM.
OITE-y, EDXVKn. M. B., E. E. C. &,
Surglaal Dlxeasea of Children. In mw 12bii>. toIuiiw of GV TWK ■*)>'■ *
Cbniiiii>lilliogniiikic plates and 86 winkIcuu. UuUi, K. 8m dvia of dMMit JVoaaa^
One In inim«dtu*4y alrafk an fvadlai Uik book
«lih ll> acrmbla ityla aad lh« aililvnoa llanaT'
■bar* t**«oue<lia naMloal fMnlUxtlyot A*
aMMr »II)> hl> mtlML TIm liaok nw ba
fccoMflty i*winm*»AM la toUi «*uil««(* (ad
prMilile*cn>. II I* fun of anuad lalniinatu^
plMMUly glTaa— 4aMii ^avfvy, Hay, ItM
STUDE^fTS' SERrsa or MAirVALS.
AtftMaoT FHlMaM«a«B]a.A>rllH>Ua*araiUik«l*a>id VnaUlltmn gf lladtM— Ud ttUaMT,
vmita bytoiiMaiTHDhennr Ei*i»l>*n,aad iMUad la vwkM-'MaUinafalaniaaalmMMpMHL
tkhty IIMHraMd aod ai a l«« prloo, Tta Mlmrlni: nWuraaa a» no« rMtdvi TuriaT jltmM if Ai"
Ootu/* AirTHM OH«»a4. m, ll«t.n>.>-» J-1I«AJmk«I /^AwM, Ki Raan^ W" U.«m ivd rbni-
KfiM (4Ui •dM(M),|IMi IVb>>'> Jfa<»a /-KinMw (A tsfliloii), |IM i>t Iauvmi^
Pima'i AiTpitvl fWALtoyp.lJ; aud Kitta** Mtmmunt ffHP>kfa IMhadiO' Hia lcll>wta«
lilnrnMi ripna'>n>rM>Mc««b>i-u. PoiMpafBlanoUmaMa tndaxaa UA^a^a.
SERIES OF CLiyiCAL MANUALS.
luacnuiRlDi ra(thl«ft*>(M II taaliaan UiadMlKU i^f ili'iutii|in«ratefnirM*th*
[■Ilanlan of aaUK«1lailT«BKiMiKnphion in>b«tlanl(
>talunaairllle«aMhi«kaatHil|iMa<aad«llII>aCra«l'
VMa. Tlia rallovlu miwnaaara nnit raanly; \'ta
Ik* ^dat^lLra: Ouna * Kain-a CMIA^mc 8Mi
■lia/t*Mu<a<i<,4a<M^ KS; Niihh on Iha *'
'jnalbi
•lpff«
•iMIVIIk
Matwm.
ajH>II*(*l-)n of aaUK«1lailT« BdMipaphi on in>b«tlanl eilnioid >uMpMa la'a«feM> M. . ,^ .
natalunaairlllMaMlo<*aat>M|iMa<aad«llII>aO««lrlItuiinw<jlnruhram»4lttqmaiiiiad«i^
i.n»; irmraiMMM 4»Ml*,aiL% luuoa
xAirtin.KX: irmraiMMM teMw.|K% iuu »a
JiiMi, IB; wax as dW^Ml XMSuM ^ CMHr*s K)
nm P>ra<« praMamaaWOMMWiaaZtli MmnM
(ha TWm, VOi Tutaaon /aiuiiaal Otafrinlfi^ |!l _...._
Tin foUualaK I* Ip MUra prrparMlou: Iadu oB /aum<j k^
hidai «« iMl MC*.
OOKI>ir« MACTir*L TBEaTIBE OS TBKI WEaTOXSOHl
DtmUBBaoPCBII.UBKN. &l>th adllloa.t*- OITA HVHTKM
n*^ Mid ■aimautod. laaoa tmuo mtaaaall Hbh tdIuibo "■'
mpaf«». Cl««li.tt.t»:lM«hM.«Uft. \
ind SiTjLaaoB Nwnduaad 4
nf tJu oraOf^ Win aapanu
■ TtD
I
Lka baumaa A Co.'8 Fubuvatioxs — Med. Jiuis.! Mlscol.
TIDY, CHARLES MBYMOTT, X. B., F. C. &,
AvIlHNr at C\muitra aftt of fWr^K J(«mnt ant rMu SmtllA at lit linftm fibt^ol, «lb
Loeal Mediolne. VoufMs II. Logitimaqr ■»<1 Pnernit*, P1WMIK7, Abof^
llod, Rapf, IihImuM EspoMU*, SedonVt BmIkUit, LJr* Birth, Itianticio*. AspliriU,
Droitnlng, HuiKine, StraifulaUoc^ SBSocaiioa. Making a tmrf hunlKme impcriB) do-
um voluine of 63» pngM. Cloth, •6.00; tcMiicr, fl.Oa
VoLPXK I. CoDlAiiitng 604 bnpcrlNJ ocuwi paM>, with two buiitiful mImmI
|>l»l«. aotlr, $«JN; Imihtr, rXO.
I
\
Th* mU«A9Um «priM«d «1tt Ih* ant tantm
al Ihln >arli I* tn do ■!■• kaarncd bf > ptniMl a(
Iha •HOB-I ToluoKi. W* ftod II fllikraciarlMd kj
ttia Mrina tula*!* of ilslall aod dImu-obw at ■>■
pravbiri vhlivh m hul orfTMif to hlchlf to «nn-
iii*n>l Id <iur fornxt d<rtlM,atut ■ltl«li raodarttaD
Taiiml-la ie Uin Diedleal fsrM. Tlia ci^ow
UMn of «■■«■ *|ipaDdt4 ID aMti dlitaloa of 111*
■iitiJMi muft hkia <<Mt Ui* auUKr a Hodlstona
■muUBi nflaiiof aod MaMtch, bill ihar OMiauiula
■'OS uf UiB niaiat •aluablo lutun* 01 iha bosk,
•apaelallj for rahiaao* la MadtoD-lacaJ Irttta.—
J awnnio Jwmif ^ Ut JVMlnl WauM, A|»ll, mi
TAYLOR, JIjFRED S., M, D.,
PeiBoiw in Bslation to Hodiool Jiuiaprudenco «nd Hodiein*. TUnI
AmericuL from tli* tbtrU bimI rariaed P^glMh oJitioii. In cine Uii* (KUto tcJuum of 7M
By the Same Author.
A Manual of UediOKl Jurutprudooco. RisliUi Anvrkaiifroni lb* Moth LdH'
den •ditkin, ibonniKfal; rvriatd ami r««TiU«a. Eiljtvil 07 Joiix J. ttuai^ M. D. In ooe
br^ octavo Toliiiiic
PEPPMS, AVOU8TU8 J., M. 8^ M. B,, F. JZ. C. 8,,
Vorwuia XAdldiu.
jWiM *f MimmU, Maw.
LEA, HENJRTC., LL. I>.
Obapiur.H from the RnliKioiia Htatoir ot Spitin.— Coosorshtp of the
PreM.— Uf-BtiOF. and lUummati.— The SademonudAs.— El Saoio Hi&o
de la Ouardia.— Brlands de Bardaxl. In om 12do. Toltme o( 5^2 ]>■«'"■
CUh, U..'iU. JuM rr.<Jy.
or a fraa-thlnkM. m* nodaraUon doptl*a* Uia
loill^iniMit of oona of lu lOtM. TSr [acta nai
Utf dncaiuaDU^«f whwhtlia vuiBttraod aotoltf
atiaaiaraUcnlafuillilDn.ara it»u|Oel In liitnlixwa
onler and pnduaa ea ih* nwlxan *nMt&i| lt>a
Tba width, da^ and HwrDncbaau or nHacti
whloh havo aaniiad lit. Laa a W|h Ibmfrmn pla»
aa lh« ablHl hlMctiia 1^ InqnlalMim ha* T*t
fbund ara bfra appllad toaancMOa-liaaaaof IMI
1 iuti)*el. w* lian odI* to aaf o(ibt> Tolnma
Afiar aMaatltalr r*B<ji(( tkt work o»« d-x* aot
hanrtrhaUMtUiaaiiibaclaaOUIialle.al'rMMUiil
«■■! iu(>)*el. w* lian ODi* to aaf o(ibl> TOlnma onler and pndiiaa ea ih* nwlMan *nMt*JI lt>a
nal li *nnhll7 c«nplaiB*Bl* tha •mIut'* aarllor nor* nnavrlVil Is thai tl tfami ika l*« dMlKaad.
Mdl« la •Ml*^<aU«*l hMonr. Ula aitaMlta Wkos aaaMtaal liia aUI* k* ia*Tanr varataal-
■•d mlBut* laarata, niMh of It bnai laadUad laaL that II I* tiaar, aotor and ar»dlia-no<olliH
naaiuatlpu la MailM^ appaan oa **•«; p*a*^— ~ - -
••ta^alifHity, Jan liaf.
r, aotor and oraolia- no <o lull
)aatla*lo avothwMoli r*a*>Ha{haklat>H( haao
aa tboMltMaoI iha wiMaraodav IIif aialkad«(
tha ■»«•» aehoel af bblDry.-AmM Q^HnM
d'SM*«r* M dt LttWalun, PtiU. J«a. IWl.
B7 tbe same Antbor.
SuponitfUOD and Faroe t Bmys on The Va«er of Law, The Wacer of
Battln, Tho Ordeal andTortur*^ Tllrd reviwl and Fnliirsol oJiiL-a. To one
ban-lwiiiiu rv/al 1 SmOi. Telnma of A6S {MgM. Ckilh, ILUX
' (t^ tha braadlh aad amiiT of kl* rorw^ha*
BOioni th« ntn *Dimaa •/kaewladc*. ilia |ir» lit
■ad laairarkMaof hliila**a»nilikMmUiir4 olUi
(li>R<alUMtBaaia«aaa<aia>laU«a,aa4 hla warn
■itvliinaat to Uia eauao af IfaadoM tmd la>all*«t-
oti procnw,— .V. r. TV16MW, Aoiuata^ tin.
Kr- t^a'i •niHniit blMOrltal noassrapka, at
tiiin 'lO' nn^ffni'Hl loipoitaM li hara |md«M4
Lb an »alaic*il form, kar* l>>a* klm ■ luUqu*
Eltloa amoM Baaiba aad Aaarloaa mr^tlan.
la dlMlocaMMlut kl* iMoadllo aad alllsMt
nlat kla fovar cf oshaaMlTa hMorlrai bkbIT'
B7 the Same Author.
StudiM in Church Hisiory. Tho Riw of tbo Temporal Power— Ben-
efit of ClergT— ExoommunioaiioD— The Early Church and SlaroiT. 8"^
and and reriaed edition. In one fata) ticUTo rakunrioTfllttrank. C1m1i.S£M.
Tha anOiM la nalMlaBDily a •ckotan lia lakaa
■pararyMffeaiiladittlk Ika Wadlac ibimo aad
tofoalionl toiho miBiiim daiall ■Ubavaalili
•tfkaMrlBdca aad laiMnulllr ot tntlmtM thai
•ompal ada'rattoD. Tha wi^at «( lafifnulon
eaaiprBusd lata Iba b«ok la ■ilnordlaaiy, aad
thBrraFinaollalkwaf awUMcHlBa a*d nbraaoa*
■aakai Uw mrk isrllf^lMfmluablBio Ik* lUMi*!
whu daalna aa aiktaMIn raila* fiWB w^laal
aaiinWB. In naaihar alniil* laliuno la Iba d««loa-
naatotlk* pr1n>MnobiuvhlnMad*tlliaoDiuoh
ilaamBBB aad with ao jMalla • pampllM of
Mnplai or owtlitlai ttn*^ JtMii Trmtlltr,
Ur. Laa ■■ /ttd4 /irttMt anaoc Aakartna
arbolar* la Ik* liMory «( Ik* Mlddla Aca*. aad,
lodMd. wo haow ot •« Biitofwan wrllor wbn hat
abon aoch 1 tu ttth, aaaaraiT and
^ anh, aoaaraiT and ataaa la
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