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THE 


JOURNAL OF ANATOMY AND PHYSIOLOGY 


NORMAL AND PATHOLOGICAL. 


CONDUCTED BY 


G. M. HUMPHRY, M.D., F.RB.S., 


PROFESSOR OF SURGERY, LATE PROFESSOR OF ANATOMY IN THE UNIVERSITY OF 
CAMBRIDGE ; 


SIR W. TURNER, M.B., LLD., F.RBS., 
PROFESSOR OF ANATOMY IN THE UNIVERSITY OF EDINBURGH ; 


AND 


J. G. M‘KENDRICK, M.D., F.B.S., 


PROFESSOR OF THE INSTITUTES OF MEDICINE IN THE UNIVERSITY OF GLASGOW. 


The Journal is intended chiefly for original contributions 
to AnaTomMy (Human and Comparative), PuysioLocy, and 
PATHOLOGY; it contains also CRITICAL and HISToRIcAL 
ANALYSES, Extracts, Reviews and Notices oF Books, &c. 

It appears Quarterly in October, January, April, and July. 
Price 6s. each part, or 20s. a year, post free 21s. paid in 
advance. 


Authors will receive 25 copies of their communications gratis. 


THE 


JOURNAL 


OF 


ANATOMY AND PHYSIOLOGY 
NORMAL AND PATHOLOGICAL. 


CONDUCTED BY 
G. M. HUMPHRY, M.D., F.RS., 


PROFESSOR OF SURGERY, LATE PROFESSOR OF ANATOMY IN THE UNIVERSITY OF CAMBRIDGE } 


SIR WILLIAM TURNER, M.B., LL.D., F.R.S., 


PROFESSOR OF ANATOMY IN THE UNIVERSITY OF EDINBURGH } 
AND 


J. G. MSKENDRICK, M.D., F.R.S., 


PROFESSOR OF THE INSTITUTES OF MEDICINE IN THE UNIVERSITY OF GLASGOW. 


VOL. XX. va 


q 
“4h 


WITH TWENTY-ONE PLATES AND SEVERAL WOODCUTS, 


WILLIAMS AND NORGATE, 


14 HENRIETTA STREET, COVENT GARDEN, LONDON; 
anp 20 SOUTH FREDERICK STREET, EDINBURGH. 


1886. 


EDINBURGH : 
PRINTED BY NEILL AND COMPANY. 


| gE 
| nes 

6 y 

V. 20 4 


CONTENTS. 


FIRST PART—OCTOBER 1885. 
PAGE 
THE ANATOMY OF THE MusciEs, LIGAMENTS, AND FAscL# OF THE ORBIT, 
INCLUDING AN ACCOUNT OF THE CAPSULE OF TENON, THE CHECK 
LIGAMENTS OF THE RECTI, AND THE SuSPENSORY LIGAMENTS OF 
fone. (by CB: Loc woon:. (Plate T:),...207.....-.. po ecnias stn fase os 1 


Two CASsgs OF AN ABNORMAL CORONARY ARTERY OF THE HEART ARISING 
FROM THE PULMONARY ARTERY: WITH SOME REMARKS UPON THE 
EFFECT OF THIS ANOMALY IN PRODUCING CirsoID DILATATION OF 
THE VESSELS. By H. St Jonn Brooxs, M.B. (Plate II.)................ 26 


A SECOND BuRSA CONNECTED WITH THE INSERTION OF THE BICEPS, AND ON 
SOME RARE ABNORMALITIES. By A. Warp Couns, M.R.C.S., 
ral OP LON die: sievose «xed Bit SAB AS acti an saat MRiSee S bhw o seid aaene tartan aes 30 


ABNORMALITIES OF THE LOBES OF THE HuMAN LuNc. By A. ERNEsT 
CRT PE IO Eyeed 8 LOOK aca cceceadantine s telledddccdeees Mchebeddoce shee carinestaeert ts. 34 


THE NATURE oF LIGAMENTS. (Part IV.) By J. Buanp Sutton, F.R.C.S. 
BLS yie0 ITS eget elle eet il es Remer is Rieee Meo Please a ber atta Rete 39 


uJ 


Virat Revations oF Micro-OrGANIsMs To TissUE ELEMENTS. By G. 
Sims WooDHEAD, M.D., and A. W. Haars, M.B............... se setansna eae 76 


TuE BLoop-FoRMING ORGANS AND BLoop-FORMATION : AN EXPERIMENTAL 
RESEARCH. By JoHN LockHAr?r Gipson, M.D. (Part L.)................ 100 


THE RELATIONSHIP OF UREA FORMATION TO BILE SECRETION: AN Ex- 
PERIMENTAL RESEARCH. By D. No&t-Paton, M.D., B.Sc. (Part I.).. 114 


THe INDEX OF THE PELVIC BRIM AS A Basis OF CLASSIFICATION. By 
Professor Sir WM. TurNER, M.B., LL.D., F.R.S. ....... Seaitt ses denecuce tee 125 


lv CONTENTS. 


PAGE 

Anatomy oF Sowrrsy’s Wuate. By Professor Sir WM. TURNER. 
PRLS EV. ) 2, yweas gese.ceasansen Sanvdswornie nchauabaesecseos' va cape case ese eee anaes 144 

INODIGES OF INEW BOOKS ....ccccestsctassens cceiwacee ve euesiecdieeend shee sean eee mnae eae 189 


SECOND PART—JANUARY 1886. 
MorrHOLoGy OF THE ARTERIAL Syst—EM IN Man. (Part I.) By Professor 
MiaGATITSMmR. ISAC MOD! WARN. c-. ccc ckecs+ ss: sssenencroereses sete eemeane 193 


ANATOMY OF THE SHOULDER AND Upper ARM oF THE MoE (Talpa 
europea). By R. AusSTIN FREEMAN. - (Plate V.) ........0.cc0ceseeseeneeeees 201 


REPRODUCTION OF THE CARAPAX IN TorTOISES. By Hans Gapow, Ph.D., 
PAPAS OHMUAN,) § (EUBUC Vi lo).ns sth on sieves conaGtebeseunreduare:=.cheten eres 220 


DEVELOPMENT AND DrcAy OF THE PIGMENT LAYER IN Brrps’ Eces. By 
AE CAN DRI IM. MCAT IO WEES MDs conn tccusen eee stes tos: ceee eer en eee 225 


CONNECTION OF THE Os ODONTOIDEUM WITH THE Bopy oF THE AXIS 
VERTEBRA. By Professor D. J. CUNNINGHAM, M.D. ............ceesees0eees 238 


SKELETON IN Grococcyx. By R. W. Suurenpr. (Plates VII., VIIL., 
Ba aia yeduendenk ts eb Ree Soe abi bes oh cetacean edn ee ge .. 244 


THE RELATIONSHIP OF UREA FoRMATION TO BILE SECRETION: AN Ex- 
PERIMENTAL REsEARcH. By D. Noiit-Paron, M.D., B.Sc. (Part II.) 267 


ReEcENT HisroLocicAL Mernops. By WitiiAM Hunter M.B. (Edin.), 
IME RGSS Sean bd ts eee en eed etl) SO, a or 307 


SacRAL INDEX IN VARrous Races oF MANKIND. By Professor Sir Wm. 
MORNER, MM. B., BBS, soccer pee a seeo. soph one xe eee sivas 0 ee 317 


BLoop-FoRMING ORGANS AND BLoop-FoRMATION: AN EXPERIMENTAL 
REsEARCH. (Part II.) By Joun LockHartr Gipson, M.D................ 342 


MALFORMATIONS OF PELVIS AND PELVIC ORGANS IN A Fatus. By Joun 
Patmes, L,R.C.P. Lond., MR.C.S. Eng, ..... Ass csasesessocose seen Ps ain! 


ANOMALOUS MUSCLE IN THE FRONT oF THE NECK IN A HuMAN SuBJECT— 
A STERNO-PETROSOPHARYNGEUS. By G. E. REnnip, B.A.,.............- 356 


CasE oF CoNGreNITAL Hyprrrrorpuy or THE Lac. (Diffuse Venous 
Neyus.) / By GriBerT BARDING, ML. B......s.5..stisescsccevnseadeteecouess tees 358 


MANDIBULAR DENTITION OF THE SuHREWws. By G. E. Dopson, M.A. 
Mena Dssscice set SUG ledeasonoues kere nents Marietta Meee Ee OSee A AES Bex 359 


oa 


CONTENTS. Vv 


THIRD PART—APRIL 1886. 
PAGE 


Action oF INFUSED BEVERAGES ON Peptic Digestion. By JAmeEs W. 
WRASER: MD Clee Big) MRC: Senos. oo srk. nscasn ext sasarcreimacacsss « 361 


Some VARIATIONS IN THE HUMAN SKELETON. By W. Arsuranor LANE, 
UN MRLEER aoe 00 oie adi bv'Ge on cles « Svs gatas tins gay bed othe weve tavonsgesteanioen 388 


Norte on A CAsE oF Bicrpirat Ris. By R. L. MAcDonne tt, M.D. ...... 405 


OsTEOLOGY oF CoNURUS CAROLINENSIS. By R. W. Susuretpt, M.D. 
MEM LES EN rena NS Me hecho stn cac taba ooh ns Seater Meats it ONG ott daly vane ost ohE Copieete asncna dae Se 407 


A Navaso Sxutt. By R. W. Sauretpt, M.D. (Plate XII.) ............... 426 


NorE oN THE Navaso InpIAN Skutt. By Professor Sir WtiiramM 
SURES OG Bier 6 ORS ie ie en rae aris a SAE iA a er ee 430 


ORIGIN OF CERTAIN CysTS—OVARIAN, VAGINAL, SACRAL, LINGUAL, AND 
TRACHEAL. By J. BLaAnp Surron, F.R.C.S. (Plate XIII.)............. 432 


THE Bioop-FoRMING ORGANS AND Btioop-ForMATION: AN EXPERI- 
MENTAL ResearcH. By Joun Lockuart Gipson, M.D. (Plate XIV.) 456 


INVESTIGATIONS IN THE RELATION BETWEEN CONVERGENCE AND AC- 
COMMODATION OF THE Eyrs. By Ernest E. Mappox, M.B. Edin.... 475 


A CoNTRIBUTION TO SPLENIC HistoLtocy. By Roperr Ropertson, M.D. 
FWieneTaerG (EXILE GO MAO) ocr sanice de ste vaisbianic Soeubecash tebe used ct code Ra aian eae ere eee 509 


SUPERNUMERARY LEG 1N A MALE F Roc (Rane palustris). By FREDERICK 
ReER MANNED). (PIAteeX WI.) cacnscsnssteas comncsnceacseteeeeenetemee. <3: 516 


THE NATURE OF THE RELATIONSHIP BETWEEN UREA FORMATION AND 
Bite SEcRETION. By D. No&u-Paron, M.D., B.Sc., F.R.S.E. ......... 520 


Hinp Limp or IcHTHYOSAURUS, AND ON THE MorpHoLOGY oF VERTE- 
BRATE Limps. By Professor D'Arcy W. THompson, B.A. ............... 532 


THE LUMBAR CURVE OF THE SPINAL COLUMN IN SEVERAL RACES OF 
Men. By Professor Sir Witt1AmM Turner, M.B., LL.D., F.R.S...... 536 


PREM ITOLNMT COATT IN ODIO Set sea c as cornca encore icciccda es sti Och econuaba sooneesinesooeeseseceus 544 


CONTENTS. 


FOURTH PART—JULY 1886. 
PAGE 


PuysloLocy oF THE HEART OF THE ALLIGATOR. By T. WrEsLEY MILLS, 
IASC DS cats sescnice cen siianieins Hodiiraaeedebue one Sara: soaeae venteaee oe eeEEED 


FUNCTIONS OF THE TonsiLs. By R. Hinasron Fox, M.D., M.R.C. P. sone 


INVESTIGATIONS IN THE RELATION BETWEEN CONVERGENCE AND AC- 
COMMODATION OF THE Evers. By Ernest E. Mappox, M.B., C.M. 
HOAs oe BOS eeepc Fe gsehe abe ae ap sea op BEE Tp oeitaeeoeree sphere tedmepieeve tne 565 


Srx Sprcrmens oF SpINA BIFIDA WITH Bony PROJECTIONS FROM THE 
BopiEs OF THE VERTEBRE® INTO THE VERTEBRAL CANAL. By Pro- 
fessor Humenry, F.R.S. (Plates XVII., XVIII.)............00-0cs00s its.) 


PENTADACTYLUS PES IN THE DorkKING FOWL, A VARIETY OF THE Gallus 
domesticus, WITH EsPpEcIAL REFERENCE '0 THE HALLUX. By JoHN 


ChON i240) A Pe? <8 OR SOS PERSE 3 a SM Wa 3 SOROS Since Jia ogbes eee 593 


VITALITY OF WILD ANIMALS UNDER Fire. By Brigade-Surgeon WILLIAM 
(CURRAN, ASML D): (Plate xu Xe) con. .cccessescaree op mene een SS peetee vseseee OOO 


ALIMENTARY CANAL AND PANCREAS OF Acipenser, Amia, AND Lepi- 
dosteus. By A. B; Macauztum, B.A. (Plate XX.) .4....,.80..gonueee . 604 


NEURAL SPINES OF THE CERVICAL VERTEBR& AS A RAcE-CHARACTER. 
By Professor D. J. CUNNINGHAM ..........056 sis apo s.ARIETY » coies scien pede en 


VARIATIONS IN THE NERVE SUPPLY OF THE FLEXOR Brevis POLLIcIS 
Muscitz. By H. St Joun Brooks, B.A. and M.B, (Dub.).............. 641 


MorrnHo.ioey oF THE INTRINSIC MUSCLES OF THE LITTLE FINGER, WITH 
SOME OBSERVATIONS ON THE ULNAR HEAD OF THE SHORT FLEXOR 
or THE Tuump. By H. Sr Joun Brooxs, B.A. and M.B. (Dub.) 


MEAD SRONUL, ) a,c; ocesivaaisceaseas Eteonear een sich e ac aaeeeantts ceed sasieces arse see beeen 
NATURE OF THE RELATIONSHIP OF UrvA Formation To BILE SECRETION, 

By D. Nozu-Paron, M.D., B.Sc. FURG.E.. ...cc.ccicvesecectes «se ahe eeeee Gow 
Bioop-ForMING ORGANS AND BiLoop-FoRMATION: AN EXPERIMENTAL 

Researcu. By Jonn Lockuartr Gipson, M.D. ..... diese sas ea eeaeae soqnen OMe 
ANATOMICO-PHYSIOLOGICAL NOTICES..........00.-+008 Reaieasticins ee chats eais soe eee Eee 692 


Hi ee gee - Ree ne ee A Meeecamnet 3 Fb Teg ie CS oka souadeasbhcspeyes cee tee 


Fournal of Anatomy and Phystologyp. 


THE MORPHOLOGY OF THE ARTERIAL SYSTEM IN 
MAN. (Partl.) By A. Macarister, M.A., M.D., F.BS., 
Professor of Anatomy, University of Cambridge. 


THE arrangement of the blood-vessels in the adult forms of the 
lowest, and in the embryos of the higher vertebrates, indicates 
that the history of the complicated vascular system of higher 
forms has been one of development from a simple and regular 
ancestral condition of metameric and inter-metameric vessels, 
through easily defined stages, to the more confused and irregular 
condition of the arterial system in the adults of higher forms. 
The blood-vessels participate in the metamerism of the 
vertebrate body, however that metamerism may have arisen; 
but as the vascular system arises in the course of the increasing 
division of labour of the organism for the transport of nutriment 
from one part to another, the chief stems are those which are 
dia- or inter-metameric. The simplest vertebrate vascular 
system probably consisted of a tubular arcade or half ring on 
the splanchnic wall of each segment on each side, joined to the 
corresponding arch in the preceding and succeeding segment by 
a dorsal and ventral inter-metameric trunk on each side of the 
median line. In the whole organism the vessels would thus 
form a double series, two long ventral trunks, two corresponding 
dorsal trunks, and the lateral uniting arches in each segment. 
One of the earliest changes which took place in this primitive 
system, was the fusion into single trunks of the longitudinal 
vessels. There are two primitive dorsal vessels in vertebrate 
embryos, and their fusion can be traced in the chick, beginning 
at the forty-second hour of incubation. This union commences 


behind the head and travels backwards rapidly, so that after the 
VOL. XX. N 


194 DR A. MACALISTER. 


fifth day there is but a single dorsal vessel for the middle and 
hinder part of the body—the dorsal aorta. 

In the region of the head and neck in mammals, the foremost 
euds of the two vessels remain permanently separate as the 
internal carotid arteries, while the part intervening on each side, 
between the united vessels forming the aorta and the separated 
parts forming the carotids, called on cach side Ductus Botali, 
becomes obliterated, except in cases, like those described by 
Harrison, Quain, and Flood, in which the persistence of the 
ductus and the obliteration of the transverse part of the third 
aortic arch causes the internal carotid to arise independently from 
the aorta. I sawone unilateral instance of thissome years since, in 
Dublin, which, like the other described case, was on the right side. 

There were also originally two ventral longitudinal vessels, 
but their union probably occurred even earlier than that of the- 
dorsal. This embryonic doubleness of the ventral stem has 
been regarded as a secondary cleavage, due to the presence 
of food yolk, but, I think, on insufficient evidence. 

The setting apart of one portion of the single ventral vessel to 
form the heart differentiates the pre- from the post-cardiac 
portions of the ventral vessel. The latter becomes venous, and 
we shall not at present trace its history farther. The former 
becomes the ventral aorta. 

The adult arterial system consists of the modified pre-cardiac 
ventral vessel, the dorsal trunk, the pre-cardiac lateral arcades, 
and the dorsal portions of the metameric vessels in the post- 
cardiac segments, together with new branches arising from these. 

As a consequence of the cardiac differentiation, the only places 
where complete metameric arcades remain are the pre-cardiac 
segments, where the ventral aorta is joined to the dorsal by 
vessels which become the lateral aortic or branchial arches. 

Behind the heart in higher vertebrates, a series of vessels 
extend from the dorsal aorta through the mesogastric fold, and 
end in the splanchnopleure; these correspond to the dorsal 
extremities of the post-cardiac lateral metameric arcades, but 
just as the pre-cardiac lateral vessels have become specialised by 
the development of the branchial clefts into respiratory arches, 
so these, very early in the phylum, have participated in the 
specialisation of the post-cardiac splanchnopleure and become 


THE MORPHOLOGY OF THE ARTERIAL SYSTEM IN MAN. 195 


visceral arteries. Like their anterior representatives, they 
become broken into a rete in the middle of their course, forming 
a capillary plexus, and from this their ventral continuation has 
become venous, ending in the sub-intestinal ventral vein, the 
specialised derivative of the post-cardiac ventral vessel. The 
modified remains of these vessels we know in human anatomy 
as bronchial, cesophageal, coeliac, mesenteric, and vitelline arteries, 
which, originally paired, have for the most part undergone, like 
the longitudinal trunk, a fusion into single median vessels. If 
these were completed they would stretch from the dorsal to the 
ventral vessel, but the causes of their displacement and of their 
being cut short are easily seen. 

On the development of the muscle plates, a parietal branch 
from the dorsal aorta arises in each metamere for its supply, and 
when the muscular wall of the somatopleure differentiates into 
dorso-lateral and ventro-lateral muscles, this vessel gives off 
corresponding dorso-lateral, and ventro-lateral branches. The 
former of these extends into the tissues arising from and below 
the laminze dorsales on each side, the latter extends ventrally 
into the somatopleure. 

These metameric parietal branches from the dorsal aorta are 
not necessarily similar in size and distribution, although serially 
homologous in origin, for, like all smaller vessels, they arise, for 
the convenience of the segment, to supply the needs of the 
developing parts; they also are not necessarily regular, their 
evolution being proportional to the sizes of the territories to be 
supplied, and to the activity of the metabolism therein. 

The metameric parietal branches are least modified in the 
region of the thorax, where they form the inter-costal arteries, 
also in the loins, where they form the lumbar vessels, each single 
trunk on each side giving off dorsal and ventral branches. 

The terminal branches of the dorso-lateral vessels form a 
continuous series of inter-segmental anastomoses along the 
closed edge of the dorsal fissure on each side, which becomes in 
the adult the posterior spinal artery, receiving supplies from the 
entire series of segmental vessels. A similar but late-forming 
and single median anastomosis forms along the floor of the dorsal 
groove, the anterior spinal artery. Free anastomosis along this 
region is necessary, as the spinal cord requires a constant, 


196 DR A. MACALISTER. 


uniform, and uninterrupted blood supply. The ventro-lateral 
branches likewise form along the medio-ventral line on each 
side a chain of medio-ventral anastomoses; these become, in the 
adult, the epigastric and internal mammary arteries. 

Coincidently with the development and consolidation of the 
separate parts and processes of the axial skeleton different systems 
of inter-metameric anastomoses arise. These vary in position 
and size according to the general principle which regulates all 
such anastomoses, viz., that whenever a primitive blood channel is 
so placed that its blood stream is liable to be interrupted, com- 
munications form between its branches and those of neighbour- 
ing trunks at the nearest spot where there is the minimum of 
pressure or displacement. In the dorsal region, owing to the 
development of the rib-basket and the consequent interseg- 
mental immobility, the parietal trunks are not lable to much 
compression, so, except at their ventral terminations, the 
anastomoses are small. 

Watching the development of bluod-vessels, we learn with what 
facility vessels form in mesoblast, and although I cannot speak 
positively, yet from what I have seen I think it probable that vessels 
have proceeded in development from the compound lacunar system 
to the tube; itis also probable that, historically, vascular space has 
at first arisen in the excavation of single plastides by diacelosis ; 
and when, in the course of descent, a trunk has become established, 
the process is then abbreviated as we find it in the chick, where 
in the formation of large trunk vessels the space arises by the 
liquefaction of the central cells of primitively solid cords. 

The largest inter-metameric anastomoses of the parietal vessels 
take place close to their primary divisions, the dorso-lateral 
trunks communicate by smaller or larger branches on the dorsal 
surface of the rib, between the rib-neck and the front of the 
transverse process; these ante-neural junctions are the chief 
regular communications, but others occur more irregularly and 
opistho-neurally. The chief radical anastomoses of the ventro- 
lateral branches is precostal. Each ventro-lateral artery gives 
off also a larger or smaller lateral branch close to the exit of the 
lateral spinal nerve, which is superficial in its distribution. 

Each dorso-lateral artery in the thoracic and lumbar region 
having communicated, although by very niinute vessels, with its 


THE MORPHOLOGY OF THE ARTERIAL SYSTEM IN MAN. 197 


preceding and succeeding segmental representative, divides into 
internal and external branches, the former being the lateral 
spinal, which ends in the inter-metameric anterior and posterior 
spinal anastomoses ; the latter ends in the dorso-lateral muscles. 
Each ventro-lateral branch in the thoracic and anterior four 
lumbar segments of man, pursues also a uniform course, having 
a very small precostal anastomosis and a fairly uniform 
segmental distribution. 

The parietal branch belonging to the lower cervical, lower 
lumbar, and upper sacral segments are much disturbed in their 
simplicity by the development of the limbs. As these arise by 
the consolidation of ventro-lateral appendages derived from 
several segments, so each limb primarily receives vessels as it 
receives nerves from several metameric trunks ; but, coincidently 
with the reduction of the basipterygium to a single rod we 
have a corresponding reduction of the main arteries to a single 
or at most a double trunk, which, in accordance with a well- 
known law of vascular distribution, runs along that side of the 
limb which is the least exposed to pressure, and least liable to 
alteration in length in the course of accustomed movement. 
The vessels which become enlarged to supply the limbs are the 
lateral branches of the ventro-lateral trunks; we have seen that 
there are such branches, although extremely minute in all 
segments, but they are specially large in such segments as have 
contributed to the formation of the limbs. 

The parietal branch of the last cervical segment is the trunk 
subclavian, arising from the dorsal aorta, above the point where 
fusion into a single trunk ceases, and as these constitute the 
chief blood supply of the fore-limbs they are commensurately 
large. Tach gives off as its postero-lateral branch the cervicalis 
profunda, and these, like their serial homologues, pass backwards 
and give off lateral spinal branches inwards, and muscular 
branches outward. 

Owing to the alteration in the dorsal aorta consequent on the 
retrocession of the heart, the trunk parietal artery for the first 
thoracic segment is obsolete at its root, but this is compensated 
for by the development of a large precostal anastomosis from 
the preceding trunk, which, on the principle that “to him that 
hath shall be given,” not only thus supplies the appended limb, 


198 DR A. MACALISTER. 


but by this channel forms the functional root of the first, 
sometimes also (by the enlarging of the second precostal 
intermetameric anastomosis), of the second intercostal arteries, 
the whole distributional system arising from this anastomotic 
root being named the superior intercostal artery. It is rare to 
find a direct union of this vessel with the aorta, but I have 
twice seen it to arise therefrom on the left side, and once 
from the bifurcation of the arteria innominata, that is from the 
right aorta. Tracing the subclavian trunk a little further 
forward, it terminates at the inner edge of the scalenus anticus 
muscle. Its posterior medio-ventral anastomotic branch passes 
backwards as the internal mammary, its lateral branch pierces 
the scaleni, the cervical equivalents of the intercostal muscles 
as the so-called second stage of the subclavian artery, which 
is thus morphologically differentiable from the first stage. 

In the more anterior cervical region, the vascular system of 
the component segments is necessarily much modified. The 
metameric arches are displaced backwards, and the portion of 
the dorsal aorta from which the parietal vessels should arise, 
the Ductus Botalii, is obliterated, hence the blood supply must 
be provided for by anastomosis. To this end a large retro-costal 
intersegmental anastomosis ascends from the root of the parietal 
vessel of the last cervical segment; this enters the interspace 
between the sixth and seventh cervical segment, and is, in the 
young foetus, placed at its origin at this level; but as the heart 
recedes and pulls all the cervical vessels downwards, its root 
becomes displaced ventrad of the seventh cervical rib (anterior 
root of the seventh cervical transverse process). This large 
anastomotic trunk ascends retro-costally but ante-neurally, giving 
off at each segment lateral spinal, and muscular branches pre- 
cisely in series with those of the trunk segments. This trunk 
vessel is the vertebral artery, the first stage of which is the 
portion due to cardiac displacement, the second stage being the 
regular intersegmental union. 

On reaching the first cervical segment, the continuous anasto- 
mosis is interrupted, so the whole vessel here divides into the 
two proper to the segment, a muscular outgoing, and a large 
lateral spinal ingoing, which, like its predecessors, turns inwards 
to the spinal cord, ventrad of the spinal nerve, which, owing to 


THE MORPHOLOGY OF THE ARTERIAL SYSTEM IN MAN. 199 


its large size, it overlaps; here it joins the two longitudinal 
anastomoses, the posterior or posterior spinal, which, from its 
disproportionate size, looks like a branch of this lateral spinal 
stem, and the anterior or median or basilar, which is the con- 
tinuation forwards of the anterior spinal system, the lozenge- 
shaped space from between the basilar and the anterior spinal 
being an island, similar to which I have thirteen times seen similar 
islands bounded by divisions and reunions of the basilar trunk. 

It is thus interesting to see how, in the course of the vertebral 
artery, we have so diverse morphological elements represented, 
the retrocostal, intersegmental anastomosis constituting its first 
and second stages, and the large lateral spinal branch of the 
first cervical segment constituting its third and fourth stages. 
Thus we have the complete system of durso-lateral branches for 
the cervical segments provided in the absence of a dorsal aorta. 

When we take into consideration the varieties of position and 
course of the vertebral artery in man, and its apparently 
discrepant relations in elasmobranch, teleost, reptile, and bird, 
we find that this view of its nature enables us, in the simplest 
manuer, to explain all its variations, and makes us understand 
why this system should be so prolific in varieties, 

The ventro-lateral parietal branches of the cervical segments 
are also united by a continuous precostal anastomosis, not far 
from where their origins had originally been from the now 
obliterated aortic trunk; this anastomosis constitutes the thyroid 
axis and cervicalis ascendens, from which the modified ventro- 
parietal branches start. These are chiefly remarkable as giving 
off lateral branches, which contribute their quota to the nutrition 
of the limbs, as the suprascapular and posterior scapular ; their 
parietal branches are small; the posterior branch of the former 
and the superficial cervical branch of the latter are the continued 
parietal branches of these two trunks. 

With the condensation of the anterior segments which takes 
place in the formation of the skull, all distinct vascular meta- 
merism is lost, and the anterior segmental arches become dis- 
placed backwards or obliterated. The common and external 
carotids are the continuations of the ventral aorta, while the 
root of the internal carotid is the altered relic of the third arch, 
and the ascending continuation of that vessel is the upper part 


200 THE MORPHOLOGY OF THE ARTERIAL SYSTEM IN MAN, 


of the dorsal aorta. The length of the carotids is secondary, as 
is the length of the neck upon which it depends. : 

In the region of the oral and pre-oral visceral clefts, the 
branches of the stem arteries are distributed in superficial and 
deep branches to the ventral parts of each inter-fissural tract of 
tissue; thus to the infra-hyoidean region we have, as superficial 
and deep branches, the superior thyroid and superior laryngeal 
arteries; to the supra-hyoidean we have the hyoidean artery, 
and more deeply the lingual; to the mandibular region we have 
superficially the facial, deeply the ascending palatine artery; to 
the maxillary lobe we have externally the transverse facial, inter- 
nally the internal maxillary, to the fronto-basal lobe we have the 
temporal. 

None of these branches are the vessels of visceral arches ; they 
are secondary medio-ventral anastomoses, uniting the widely 
separated representatives of the ventral aortic trunks. The only 
carotid branches which in any measure represent rudimental 
arcades are the occipital and posterior auricular arteries. 

The cervical dorsal aortic (internal carotid) has only rudi- 
mental branches in the neck, represented by the intercarotid 
ramuli. Its intracranial continuation gives off three lateral 
neural branches, the posterior, middle, and anterior cerebrals 
(the first originally being a carotid branch, its root being the so- 
called posterior communicating, but its anastomotic internal 
branch, which joins the median anastomosis, dilates so as to form 
its functional root). The ventro-lateral branches are reduced 
and modified as tympanic, vidian, receptacular, and ophthalmic 
branches. Upon all these branches the characters of the 
secondary alterations which the cranial segments have under- 


gone have thus impressed themselves, so as to obliterate all 
traces of primitive segmentation. 


THE ANATOMY OF THE SHOULDER AND UPPER 
ARM OF THE MOLE (Talpa Europea). By R. Austin 
FREEMAN. (PLATE V.) 


THE increasing interest in Morphological Science which has of 
late years become manifest among scientific men has led to the 
careful examination of the structure of a large number of 
individual forms, and to the publication of detailed descriptions 
of their anatomy. 

These contributions to Anatomical Science frequently contain 
matter which in itself is of considerable interest, and which, in 
the hands of the morphologist, forms the material for important 
generalisations. 

It is somewhat curious that amidst this great activity of 
anatomical research, one of the most specialised and aberrant, 
although at the same time most archaic, of our British mammals 
should have been passed over in comparative silence, and the 
more so as the difficulty of obtaining specimens for examination 
is so very slight. It is true that notices of the anatomy of the 
mole have from time to time appeared, but, as none of them have 
dealt in a detailed manner with the subject of this paper, it is 
unnecessary to enumerate them here.! 

The most obvious peculiarities in the anatomy of this animal 
are those which have relation to the immense development and 
singular displacement of the anterior limbs, and it is to the dis- 
cussion of these that I propose to devote the present paper; the 
forearm has already been treated at some length in the pages of 
this Jowrnal by Mr D’Arcy W. Thompson (vol. xviii. p. 406), and 
I shall therefore confine my remarks to the proximal portion 
of the limb. 

Following the usual order of anatomical description we first 
proceed to the consideration of the bones. 

Osteology.—The shoulder-girdle of the adult mole is composed 
of two bones, the scapula and the so-called clavicle. 

1 In Mr Dobson’s beautiful Monograph of the Insectivora a very excellent 


description is given of the nearly related form Condylura cristata, together with 
some notes upon Talpa ewropea. 


202 MR R. AUSTIN FREEMAN, 


The Scapula.—This is extremely narrow, and somewhat pris- 
matic in shape, its vertebral border being only one-sixth of the 
length of the axillary border; the mesoscapular spine, at no part 
prominent, is in its middle third almost obliterated; at its 
vertebral extremity a blunt, somewhat conical process (a) (fig. 1) 
is produced, the principal purpose of which appears to be to 
afford a surface for the attachment of the posterior division of 
the trapezius. At its glenoid extremity the spine expands into 
a broad and thick, but stunted acromion (0), at the extremity of 
which may be seen two well marked facets for the strong 
acromio clavicular ligament and acromial portion of the sub- 
clavius respectively. 

The supraspinous fossa, in the glenoid half of the bone, appears 
merely as a shallow groove on the anterior surface of the 
acromion, but in the vertebral half (c) it is of fair extent. The 
infraspinous fossa, of much less extent than the preceding, is 
almost absent in the middle third of the bone, being continuous 
with and almost indistinguishable from the axillary border. It 
exists as a broad but shallow grovve behind the acromion; and 
at its vertebral extremity it forms a deep fissure (d) which is 
overhung by the spine. 

The axillary border presents near the angle a large triangular 
somewhat rough surface limited internally by a sharp ridge; 
the surface affords origin to the large and powerful teres major. 

At the glenoid extremity of this border is a wide concave 
surface, on the inner or vertebral margin of which is a small 
tubercle (fig. 2, a) supporting a facet for the origin of the biceps. 
The outer or dorsal margin of this surface is formed by a slight 
ridge which, proceeding upwards, crosses obliquely the infra- 
spinous fossa and intersects the spine about its middle. This 
ridge limits anteriorly the extensive origin of the long head of the 
triceps, which occupies the whole of the surface above described. 

The anterior border at its glenoid extremity is almost indis- 
tinguishable from the supraspinous fossa with which it blends, 
but above it is thin and sharp excepting near the angle, where 
it widens out to form a somewhat oval concave surface for the 
insertion of the levator anguli scapule. 

The vertebral or suprascapular border is, in its anterior two- 
thirds, comparatively thin, and bears on its ventral aspect two 


ANATOMY OF THE SHOULDER AND UPPER ARM OF THE MOLE. 203 


small facets, the more anterior (fig. 2, b) of which, narrow and 
elongated, gives attachment to the anterior trapezius, whilst the 
posterior (fig. 2, c), which is oval and concave, marks the insertion 
of the serratus magnus. 

In its posterior third the vertebral border widens out greatly, 
forming a wide surface which affords attachment to the teres 
major, posterior trapezius, serratus magnus, and rhomboideus. 

The subscapular fossa is as such confined to the vertebral half 
of the bone, the ventral aspect of the glenoid half being occupied 
by a prominent ridge; this ridge (fig. 2, 2) supports the bicipital 
impression and extends to the margin of the glenoid cavity. 

The glenoid cavity is elongated and deeply concave, its long 
diameter being placed obliquely to the vertical plane of the bone, 
an arrangement which is rendered necessary by the position in 
which the humerus is carried. There is a smal] facet on what 
would be, if the glenoid cavity had its usual direction, the anterior 
margin ; this marks the scapular attachment of the glenohumeral 
ligament. There is no coracoid process, a fact which is suffi- 
ciently explained by the composite nature which Professor Parker 
ascribes to the so-called clavicle. 

The clavicle, or more properly speaking the coraco-clavicle, is 
a curious irregular little bone which articulates with the 
humerus and the presternum but not with the scapula. 

On its outer aspect is an oval or saddle-shaped surface 
(fig. 3, 2) for articulation with a corresponding surface on the 
humerus, and its inner aspect is occupied by a narrower concave, 
somewhat reniform surface (fig. 4, a) for articulation with the 
presternum. 

The anterior surface is deeply concave from side to side but 
convex vertically, especially at its upper part, where it turns 
over and merges into the posterior surface. The sides of the 
concavity are produced into ridges which afford attachment to 
muscles and the external ridge widens out below into a triangu- 
lar surface (fig. 4, 6) which gives origin to a portion of the 
deltoid. The posterior surface is limited above by an obliquely 
transverse, rounded ridge produced by the folding over, so to 
speak, of the superior margin; below this is a concave surface 
of quadrilateral form (fig. 3, b), in which some fibres of pectoralis 
minor are inserted. 


204 MR R. AUSTIN FREEMAN. 


The ventral surface is mainly occupied by a prominent 
obliquely truncated conical process (figs. 3 and 4, c), which is 
directed ventrally and inwards; on its inner aspect, near the 
free extremity, is a rough surface which gives attachment to 
the pectoralis minor, and at its base, at the junction of the 
anterior and ventral surfaces, is a minute foramen (fig 4, d), which 
Professor Parker believes to represent the coracoid fenestra, 

The Sternum.—The manubrium is of very large size and greatly 
produced anteriorly, its length being nearly equal to that of the 
mesosternum and xiphisternum together. It is considerably 
expanded, especially at its middle part, and onits ventral aspect 
is produced a very prominent ridge or keel, which when viewed 
in profile is seen to have somewhat the form of the lateral half 
of a coffin lid. The free edge of that portion of the keel which 
lies behind the angle or promontory is thin and sharp, excepting 
where it widens out to form part of the articular surface for the 
mesosternum ; in front of the promontory the edge is thick and 
rounded and expands anteriorly to support the articular surfaces 
for the coraco-clavicies. The promontory itself supports a 
tubercle which gives origin to a portion of the pectoralis 
major. 

The anterior surface, which looks somewhat dorsally, is 
triangular in shape, the apex pointing upwards and backwards ; 
it is convex both from side to side and from above downwards, 
and is traversed by a median longitudinal ridge. The convex 
surfaces on either side of this ridge form the articular surfaces 
for the coraco-clavicles, and the ridge itself gives attachment to 
some of the fibres of the anterior sternoclavicular ligament. 

Viewed from the dorsal aspect the manubrium is seen to be 
produced laterally into a pair of somewhat triangular ale (fig. 
6, a) behind which the bone is rather narrow; at the posterior 
end is a thickened mass (d) which carries the surface for articu- 
lation with the mesosternum. 

The space between the ale (that is, the anterior three-fifths of 
the dorsal surface) is occupied by a deep groove (c), which, com- 
mencing anteriorly in the middle line, deviates considerably, as 
it proceeds backwards, to the right side. In the floor of this 
groove, at about its middle, is a small foramen (d), which pierces 
the root of the left ala. 


ANATOMY OF THE SHOULDER AND UPPER ARM OF THE MOLE. 209 


The first rib articulates with the presternum, but there is no 
discernible facet for its reception. 

The mesosternum consists of four pieces, of which the two 
posterior are, in the adult, immovably ankylosed. 

The xiphisternum is narrow and pointed, and supports at its 
extremity a broad leaf-like cartilaginous expansion. 

The sternum is directly connected with eight ribs, of which 
the two posterior unite at their sternal ends and together are 
received into the space between the meso- and xiphisternum. 

The Humerus——This bone is extremely short, broad, and 
flattened, and the muscular impressions on it are mostly very 
pronounced. 

The upper part of the anterior! surface is occupied by a large 
concave rhomboidal surface (fig 7, a), which marks the insertion 
of the deltoid. At its lower angle is a rough triangular surface 
(fig. 7, b) for the insertion of a part of the complex pectoralis 
major, and at its inner angle is a deep notch (c) continued down- 
wards for a short distance into a shallow groove (d); this is the 
lower portion of the bicipital groove, 

The upper external border of this space is formed by a slight 
groove which separates it from a broad, smooth, convex surface 
which articulates with the coraco-clavicle. 

Below the bicipital notch is a very prominent outstanding 
process (fiy. 7, e) bearing two distinct impressions, for the teres 
major and latissimus dorsi respectively. 

At the base of the internal condyle jis a deep circular pit (/) 
which gives origin to the powerful ligamentous flexor sublimis 
digitorum ; the pronator ridge is thin, sharp and very prominent, 
and is produced above into a pointed process (g). There is a 
large supra-condylar canal, the lower opening of which (/) is on 
the anterior aspect of the bone. 

The external condyle is drawn out into a styliform process (7) 
of considerable length, which points upwards and slightly out- 
wards, and at its base is seen the almost hemispherical capitellum 
(j). At the summit of the bone is the superior opening of the 
bicipital canal (4). 


1 For the sake of convenience in description the bone is supposed to be placed 
as in the figures, although this is by no means its natural position in the animal 
(see movements of the limb). 


206 MR R. AUSTIN FREEMAN. 


Turning now to the posterior aspect of the bone we notice 
that the head (fig. 8, a) is very prominent and vertically 
elongated, that it looks directly backward, and that it is not 
placed at the superior extremity of the bone, but that, projecting 
considerably above it on its inner side, is a ridge (fig. 8, b) which 
represents the inner tuberosity. 

External to the head is the smooth articular surface for the 
coraco-clavicle (fig. 7,/, and fig. 8, ¢), upon which and close to 
the head is a small, shallow depression which marks the 
insertion of the supraspinatus ; this surface is limited inferiorly 
by a jagged irregular ridge (d@), at the outer end of which is a 
hook-like proress (e) bearing a facet for the infraspinatus. The 
articular surface above described represents the outer tuberosity. 

Close to the outer side of the head is a shallow groove which 
lodges the gleno-humeral ligament, and internal to the head is a 
deeper groove for the attachment of a thickened band of the 
capsular ligament. 

Internal to and somewhat above the head is a ridge (f) which 
overhangs the bicipital groove, and upon which is an impression 
for a part of the pectoralis major. 

The bicipital groove is very curiously modified, being, in its 
upper part, converted into a complete bony canal which com- 
mences at the summit of the bone (g), and passing downwards 
and inwards for about } in. opens out into a groove (fig. 8, h) 
which terminates at the notch before spoken of. 

Another very curious condition, which occurs in the upper 
part of the bone, is a large cavity (2), somewhat similar to that 
found in the humeri of birds. This cavity is conical in shape 
and excavates the upper portion of the bone so extensively as to 
convert it into a mere shell, the walls of which are comparatively 
thin and in some places quite translucent. 

The inner tuberosity bears two very distinct facets, which 
mark the insertions of the subscapularis. 

The olecranon fossa is very deep, but there is no intercondylar 
foramen ; at the apex of the fossa is a foramen (7) large enough to 
admit a very coarse bristle (in some bones there are two somewhat 
smaller ones), which opens directly into the medullary cavity. 

The trochlear surface (/) is placed mainly upon the posterior 
aspect of the bone. 


’ ANATOMY OF THE SHOULDER AND UPPER ARM OF THE MOLE. 207 


Ligaments.—lf Professor Parker’s views concerning the nature 
of the bone usually called clavicle be correct, we must regard the 
articulation between that bone and the humerus as part of the 
true shoulder-joint. It is, however to be observed that there 
are two separate synovial cavities, that the articular surfaces on 
the humerus are not continuous, and that the cavities of the 
joints are separated by ligamentous partitions. 

The ligaments of the scapulo-humeral articulation are—(1) 
Capsular, and (2) Gleno-humeral. 

1. The Capsular ligament presents nothing worthy of remark 
with the exception of a thickening of its substance on the inner 
side of the joint. 

2. The Gleno-humeral ligament is a stout, cord-like structure 
which passes from a small facet above the glenoid cavity to a 
small, shallow depression at the lower and posterior portion of 
the articular surface for the coraco-clavicle close to the head. 

In its course it lies within the capsule and is separated from 
the cavity of the joint by the synovial membrane only. 

In addition to these structures the tendons of the supra- 
spinatus and biceps exercise ligamentous functions. 

The tendon of the supraspinatus is broad and thick and is 
inserted into a smallimpression between the articular surfaces 
for the scapula and coraco-clavicle; it thus, becoming intimately 
adherent to the capsules of both joints, forms a strong partition 
between them. 

The influence of the tendon of the biceps upon the movements 
of the limb is fully discussed in the section of this paper which 
is devoted to the actions of the muscles, but it may be here 
stated that when the muscle is at rest, the tendon will pro- 
bably function as a ligament, steadying the humerus, holding 
it in apposition with the scapula and limiting its external 
rotation. 

The claviculo-humeral joint possesses a capsular ligament, the 
anterior portion of which becomes considerably thickened, being 
converted intv a broad sheet of strong vertical fibres, which are 
inserted into a groove at the anterior margin of the articular 
surface on the humerus. 

The acromio-clavicular ligament.—This very thick and strong 
band of fibres proceeds from the anterior of the two facets 


208 MR R. AUSTIN FREEMAN, 


on the acromion to a small flat surface on the posterior margin 
of the outer surface of the coraco-clavicle. At about its middle 
it receives the insertion of a part of the subclavius. 

The sterno-clavicular articulation.—The coraco-clavicle is bound 
to the presternum by two ligaments, one of which, placed on 
the anterior and dorsal aspect of the joint, is of considerable 
strength. Its fibres, which arise from the anterior margin of the 
internal surface of the coraco-clavicle, proceed, some of them to 
the head of the presteraum, others to the bone of the opposite 
side. It therefore agrees closely with the interclavicular liga- 
ment of human anatomy. 

The posterior sterno-clavicular ligament is a comparatively 
thin sheet of fibres which extends from the posterior margin of 
the articular surface of the coraco-clavicle to the corresponding 
margin of the articular surface of the presternum. There is no 
interarticular fibro-cartilage in this joint, but that structure is 
probably represented by the plate of cartilage which covers the 
articular surface of the coraco-clavicle, this being what remains 
of the so-called “fourth coracoid segment” of Parker, who 
regards it as constituting with its fellow “the moieties of a 
highly modified omosternum.”? 

The interscapular ligament.—This unique and extremely 
interesting structure consists of a thick tendinous cord passing 
from the base of one scapula to that of the other, its attachment 
being nearly opposite to the spine: it is continuous anteriorly 
with the ligamentum nuche; and some of the muscles which 
arise from that structure, as ¢.g., splenius, have also an origin 
from it. From its posterior surface in the middle line, a band 
of connective tissue passes backwards to the spine of the third 
or fourth dorsal vertebra, and appears to be a backward continua- 
tion of the nuchal ligament. The interscapular ligament affords 
attachment to several muscles—trapezius, teres major, serratus 
posticus superior and dorso-interscapularis, some of which it 
appears to have considerably modified. 

It becomes a matter of some difficulty to dotedivine what is 
the nature of this ligament, for, that it represents some structure 
which exists in other animals there can be little doubt. Some 
light seems to be thrown upon the subject by the following facts: 

1 Monograph on the Shoulder-girdle and Sternum, Ray Soc., 1868. 


ANATOMY OF THE SHOULDER AND UPPER ARM OF THE MOLE. 209 


—(a) the vertebral spines are almost completely absent from 
the third cervical to the tenth dorsal; (0) the interscapular 

ligament is in intimate relation with the lgamentum 
* nuche, as well as with some of the muscles which arise from 
it; (c) an ossification has long been known to exist in the latter 
structure in the cervical region; (d) certain muscles are pro- 
foundly modified by their relation to the interscapular ligament. 

These facts appear to bear upon the subject in the following 
manner: the abortion of the dorsal spines explains the existence 
of the ligamentam nuche, or rather its backward continuation, 
in the dorsal region, and the intimate relation of the ligamentum 
nuche, with the interscapular ligament, implicates the latter in 
the changes which have occurred. The abortion of the vertebral 
spines, together with the ossification in the nuchal ligament, 
seems to indicate that the latter structure may possibly consist, 
not merely of separated interspinous ligament, but also of the 
detached spinous elements of some of the vertebra, and this 
conjecture is rendered more probable by the circumstance that 
the so-called “nuchal style” commences abruptly immediately 
behind the prominent spine of the axis. The most important 
of the museular modilications above referred to occurs in con- 
nection with the serratus posticus superior. This muscle arises 
from the anterior surface of the interscapular ligament and 
proceeds to its usual insertion, whilst, arising from the rudi- 
mentary spines of the fourth and fifth dorsal vertebre, and from 
the backward prolongation of the ligamentum nuchz is a small 
triangular muscle, which I have called dorso-interscapularis,’ 
which is inserted into the outer two-thirds of the posterior 
surface of the interscapular ligament, and which is separated 
from its fellow near its insertion by a small triangular interspace 
which is filled by the ligamentum nuche. 

If it be assumed that the interscapular ligament has been 
produced by the metamorphosis of the spinous elements of some 
of the vertebre, or by an outgrowth of the ligamentum nuche, 
it would seem that the region in which the changes have occurred 
corresponds with the origin of the serratus posticus superior, and 


‘1 This muscle is called rhomboideus posticus by Dr Dobson, whose views con- 
cerning its morphological nature are somewhat different from those above given. 
Vide A Monograph of the Insectivora, part ii. 

VOL. XX. Oo 


210 MR R. AUSTIN FREEMAN. 


that the mass of changed tissue, extending outwards through 

the substance of the muscle, has cut off, so to speak, its postero- 
internal angle, which has persisted as a separate muscle in the 

position which it now occupies. 

Myo.tocy.—The muscles of the anterior limb are, as would be 
anticipated, extremely well developed, and present several 
interesting deviations from the conditions most commonly found 
in mammals. 

We shall first consider the muscles which pass from the trunk 
to the limb and limb girdle. 

Pectoralis Major.—This muscle is not only of very sit size, but 
is split up into a number of remarkably distinct fasciculi which 
may be conveniently distinguished by attaching numbers to them. 

The posterior superficial mass (P.M. 1, fig. 9) is of large size, and 
evidently corresponds with the sternal portion of human 
anatomy ; it arises the whole length of the sternum, behind the 
promontory of the manubrium, including the expanded xiphoid 
cartilage and from all the sternal ribs. From this origin its 
fibres converge towards the crest (fig. 8, b.) above the bicipital 
groove, where they are inserted without having become tendinous. 
As it approaches its insertion, however, this portion of the 
muscle becomes covered with a layer of glistening tendinous 
fibres. At its anterior border is placed a flattish, fusiform 
fasciculus (P.M. 2, fig. 9), which arises from the promontory of 
the presternum, and is inserted into the same portion of the 
humerus. Anterior to this, and separated from it by a small 
interspace, is a narrow fasciculus (P.M. 3) arising from the head 
of the manubrium and passing outwards to be inserted into the 
angle above the bicipital notch. In front of this is a somewhat 
large quadrilateral piece (P.M. 4), which has no sternal or costal 
origin but arises from a median sheet of connective tissue which 
is interposed between it and its fellow of the opposite side! It 
is inserted into the whole length of the outer bicipital ridge and 
into the triangular surface at the lower angle of the rhomboidal 


1 This condition of the pectoral muscles is of considerable interest in con- 
nection with the opinion expressed by Miss Lindsay in a paper read at the 
Zoological Society on June 16, 1885, that the sternal keel of carinate birds ‘‘is 
an outgrowth of the sternum of comparatively late phylogenetic date, and created 
for and by the attachment of the pectoral muscles,” 


ANATOMY OF THE SHOULDER AND UPPER ARM OF THE MOLE. 211 


space. The two muscles, when in action, must pull against one 
another, since they have no central bony attachment. They are 
probably homologous with the “clavicular portion” of human 
anatomy. 

Upon reflecting the sternal portion of the muscle (P. 1) there 
is seen, underlying it, a very distinct fasciculus (P. 5) which 
arises from the posterior three-fourths of the crest of the manu- 
brium and from the second rib. It is inserted into the posterior 
surface of the inner tuberosity, becoming covered, as it approaches 
its insertion, with a layer of tendinous fibres. Behind this is a 
somewhat larger mass (P. 6), which arises from the second 
and third sternal ribs and blends more or less with the 
first part (P. 1) at its insertion. A seventh fasciculus, 
small and narrow (P. 7), takes origin from the promontory 
of the presternum and is inserted into the angle above the 
bicipital notch a little above the insertion of the third fasciculus. 
Pectoralis minor is of medium size and arises from the anterior 
half of the keel of the presternum and by a small tendinous slip 
from the first sternal rib. It is inserted into the ventral surface 
of the coraco-clavicle. 

Subclavius.—This muscle, which is of large size, arises from the 
whole length of the dorsal surface of the presternum on either 
side of the median groove, as well as from the greater part of 
the anterior surface of the first rib; it fills up to a great extent 
the large space which intervenes between the first rib and the 
coraco-clavicle. Arriving at the anterior portion of this space, 
the muscle separates into two parts, each of which ends in a 
tendon. The more internal of these divisions proceeds to the 
outer third of the dorsal margin of the coraco-clavicle as well as 
to the adjacent part of the anterior surface, whilst the more 
internal of the two is inserted into the anterior surface of the 
short acromion and into the acromio-clavicular ligament. 

Trapezius—This consists of two thin and slender muscles 
which are entirely separate both in their origin and insertions ; 
the anterior arises from the outer two-thirds of the superior 
curved line of the occipital bone and is inserted into the dorsal 
lip of the vertebral border of the scapula in the supraspinous 
region. It lies at the side of the neck, separated from its fellow 
by the rhomboid, which overlaps it at its posterior end. The 


Aa bee MR R, AUSTIN FREEMAN. 


posterior segment, which is a thin ligulate muscle, arises from 
the last dorsal and all the lumbar spines, by tendinous fibres, | 
and is inserted into the conical process on the spine of the 
scapula. In close relation, externally, with this portion of the 
trapezius is a very thin and narrow ribbon of muscle, the dorso- 
orbicularis, which passes forwards to be inserted into the skin of 
the dorsal region just behind the posterior border of the orbicu- 
laris panniculi. 

Rhomboideus.—There is only one rhomboid muscle, and this is 
probably the representative of rhomboideus minor of man. It 
is a somewhat large muscle, arising from the ligamentum nuchee 
and from the curious little styliform bone which exists in this 
region. From this origin it passes backwards and outwards, and 
becoming somewhat twisted upon itself, is inserted into the 
vertebral border of the scapula in the supraspinous region. 

In the posterior part of its course it lies superficial to the 
trapezius, a circumstance which is due to the entire absence of 
the central portion of the latter muscle and the abnormal 
position of the scapula, to which allusion has been already 
made. 

Latissimus dorsi is of rather large size and consists of two 
portions, which are separated by a cellular interspace. The 
anterior portion, which is the smaller, arises from the lower five 
dorsal and first lumbar spines, and passing forwards soon comes 
to lie under cover of the posterior portion, with which it blends 
near its insertion. The posterior portion arises from the 
remaining lumbar spines, from the lumbar fascia, and from the 
fascia covering the external oblique; it is aponeurotic at its 
origin, but soon becomes fleshy, at the same time becoming 
rapidly narrower, as it winds round the thorax to reach the 
axilla. It terminates in a broad flat tendon, which unites to a 
great extent with that of teres major, and the united tendons are 
inserted into the prominent ridge below the bicipital groove. 
Although the tendons are almost entirely blended the facets on 
the bone for their insertion are quite distinct, the more anterior 
belonging to latissimus dorsi. 

Serratus magnus arises by seven fleshy digitations from seven 
ribs,—the second to the eighth—near their angles, and is inserted 
into the oval facet on the ventral lip of the vertebral border of 


ANATOMY OF THE SHOULDER AND UPPER ARM OF THE MOLE. 213 


the scapula, the posterior part of the border und the interscapular 
ligament. 

Levator anguli scapule, which is nearly as large as the serratus 
magnus, from which it is separated by a cellular interspace, arises 
from the transverse processes of the cervical vertebre from the 
third to the seventh inclusive, and is inserted into the oval sur- 
face at the vertebral end of the anterior border of the scapula. 

As the serratus posticus superior has, in the mole, an attach- 
ment to the shoulder girdle, it may, together with its concomitant, 
the dorso-interscapularis, be properly described here. 

Serratus posticus superior is a small narrow muscle which 
arises from the central portion of the ventral aspect of the inter- 
scapular ligament, and passes outwards to be inserted by two digi- 
tations into the third and fourth ribs near their vertebral ends. 

Dorso-interscapularis, the nature of which has been fully 
discussed under the heading of ligaments, is a small triangular 
muscle which arises from the spines of the fourth and fifth 
dorsal vertebre, and from the backward prolongation of the 
ligamentum nuche, and is inserted into the outer two-thirds of 
the posterior surface of the interscapular ligament. 

We now proceed to consider the muscles which pass from the 
shoulder girdle to the brachium. 

Deltoid.—This is of comparatively small size, and owing to 
this fact, to the large size of the great pectoral, and particularly 
to the singular position of the limb, lies entirely under cover of 
the quadrilateral fasciculus of the pectoralis major; it arises 
from the outer half of the anterior surface of the coraco-clavicle, 
and is inserted by fleshy fibres into the whole of the rhomboidal 
space on the front of the humerus and by tendinous fibres at its 
margins. It is thus seen that the deltoid has no scapular 
attachment. 

Teres major.—This muscle is of enormous proportions, being 
perhaps more hypertrophied than any of the arm muscles; it 
arises from the triangular rough surface at the posterior angle of 
the scapula, from the upper two-thirds of the axillary border, 
from the broad surface on the vertebral border by tendinous 
fibres, by a small additional slip from the anterior portion of the 
vertebral border and, lastly, from tke interscapular ligament. 
From this somewhat complicated origin, its fibres pass forwards, 


214 MR R. AUSTIN FREEMAN. 


forming a large fleshy belly, which tends to become more or less 
blended with surrounding muscles, and terminate in a strong, ” 
flat tendon, which unites largely with that of latissimus dorsi, 
and is with that tendon inserted into the ridge below the 
bicipital groove, the moiety of the tendon which belongs to teres 
major being inserted into the posterior of the two facets. 

Subscapularis is a somewhat small muscle, compared with the 
ereat teres major; it arises, as usual, from the whole of the 
subscapular fossa, as well as from the ridge on the ventral 
surface of the neck. The substance of the muscle is penetrated 
by two tendons upon which the fibres are, so to speak, gathered 
up, and which are inserted into two round concave facets at the 
summit of the inner tuberosity, immediately internal to the 
superior opening of the bicipital canal. Into the surface of bone 
which intervenes between the two facets, the muscle has a 
fleshy insertion. 

Supraspinatus is a somewhat small penniform muscle, which 
arises from the whole of the supraspinous fossa up to the 
margin of the glenoid cavity ; its fibres converge upon a strong 
flat tendon, the size of which is, in relation to the muscle to 
which it belongs, somewhat disproporticnately large, which, 
uniting to some extent with, and piercing, the capsule of the 
claviculo-humeral joint is inserted upon the surface of the 
humerus which articulates with the coraco-clavicle, lying in the 
cavity of the joint, and covered with the synovial membrane. 

This tendon also becomes adherent to the capsule of the true 
shoulder joint, forming, as already stated, a partition between 
that joint and the claviculo-humeral articulation. 

Infra-spinatus, of small size arises from the greater part of 
the infraspinous fossa, and also from the truncated, conical 
process at the vertebral end of the spine. It is inserted by a 
small tendon into a facet at the base of the uncinate process on 
the outer tuberosity of the humerus. 

Teres minor appears to be entirely absent. We now pass on 
to the consideration of those muscles which extend from the 
shoulder girdle to the forearm. 

Biceps brachii.—This muscle is perhaps more remarkably 
modified than any other in this region; it arises by a thin cord- 
like tendon from a small tubercle on the ventral surface of the 


ANATOMY OF THE SHOULDER AND UPPER ARM OF THE MOLE. 215 


scapula about +); of an inch from the margin of the glenoid cavity ; 
from this point the tendon passes forward and downward to 
reach the superior opening of the bicipital canal. The axis of 
this canal being as nearly as possible at right angles with the 
portion of tendon intervening between it and the scapula, the 
tendon becomes bent up, playing over a small trochlear surface 
at the posterior edge of the superior opening. Emerging from 
the canal, but lying in the deep bicipital groove, the tendon 
follows the same course until it reaches the bicipital notch on 
the inner side of the humerus, where it again changes its course, 
twisting sharply round the polished pulley-like margin of the 
bone and coming now to have a direction at right angles to the 
last. It now swells out into a fleshy belly of considerable 
size, and this becomes contracted into a strong flat tendon 
which is inserted in its normal position on the radius. It 
is thus seen that the tendon of the biceps lies in three 
different planes, each forming a right angle with either of the 
others. 

Triceps is of large size and has the following characters :—A 
large quadrilateral mass divided into two portions by a cellular 
interspace arises from the rough surface at the glenoid end of 
the axillary border of the scapula, and from the infraspinous 
fossa for about # of the length of the bone; another distinct 
mass arises from the posterior surface of the humerus below the 
bicipital groove, and a third arises from the outer and posterior 
surface of the humerus and from the conical cavity at its upper 
part. The muscle is inserted, mainly by fieshy fibres, into the 
enormous expansion of the olecranon. 

Brachialis anticus arises from the upper portion of the outer 
surface of the humerus below the rhomboidal space, and from 
the hooklike process on the outer tuberosity; its strong, flat 
tendon is inserted into a small depression in front of the coronoid 
process of the ulna. 

Anconevs.—This muscle consists of two parts separated by 
a small cellular interspace; the posterior portion is a round 
fusiform fasciculus arising from the tip of the styliform external 
condyle and inserted into the outer projection of the olecranon. 
The anterior portion, thin and fan-shaped, arises in common with 
the preceding and becoming somewhat aponeurotic as it passes 


216 MR R. AUSTIN FREEMAN. 


over the extensor muscles of the forearm, is inserted into blip 
prominent crest of the ulna. 

Epitrochleo-anconeus or anconeus internus, a comparatively 
large muscle of a triangular shape, arises from the posterior 
surface of the pronator ridge, and passing almost horizontally 
backwards, is inserted into a prominent process on the 
inner side of the olecranon. It lies superficial to the ulnar 
nerve. 

The present paper would be incomplete without a brief notice 
of the parts which are played in the economy of the mole by the 
numerous and important modifications of anatomical structure . 
which have been described. The movements of which the 
pectoral limb of the mole is capable, present extremely little 
variety, being almost entirely confined to a backward and 
forward movement like the oar of a boat or the arm of aswimmer, 
which latter it closely resembles, and consisting, like it, of a 
“stroke” in which the outspread hand is driven forcibly through 
a more or less dense and resisting medium, and a “ recovery” in 
which comparatively little resistance has to be overcome and 
therefore comparatively little force used. 

To the execution of the former of these movements the limb 
is specially adapted, and it is to the hypertrophy of the muscles 
by whose agency it is performed, that the structural modifica- 
tions are mainly due. This will be seen more clearly if we 
examine the movements in detail, and observe the manner in 
which the several muscles contribute to their production. Thus 
upon analyzing the first movement, the backward “stroke” of 
limb, we find that it is made up of the following parts: (a) 
flexion of the humerus upon the scapula; (6) extension of the 
forearm ; (c) rotation of the humerus around its longitudinal axis, 
and as a result of this; (d) flexion of the digits. 

(a) Flexion of the humerus upon the scapula is mainly effected 
by the long head of the Triceps, which, by virtue of its very 
extensive attachment to the scapula and the great development 
of the olecranon, acts at a great mechanical advantage, and its 
action is probably largely supplemented by the infraspinatus as 
well as by the teres major and the latissimus dorsi. 

(b) The principal action, however, of the two latter muscles is 
that of rotation of the humerus around its long axis, in which 


ANATOMY OF THE SHOULDER AND UPPER ARM OF THE MOLE. 217 


action they are powerfully assisted by the posterior segment of 
the great pectoral. 

This movement is of a twofold character, consisting of (1) a 
rotation of the humerus around a line drawn through the 
middle of the trochlear surface and the scapular head, and (2) a 
backward movement of the entire bone, carrying with it the 
scapula, upon the coraco-clavicle. The latter movement is 
obviously brought about entirely by the pectoralis major and 
latissimus dorsi, whilst the former is in a great measure due to 
the teres major. 

(c) The rotation of the humerus results in flexion of the digits 
in the manner described by Mr D’Arcy W. Thompson in his 
admirable paper on the subject (Jowrnal of Anatomy, vol. xxiii. 
p. 406). 

The teres major and latissimus dorsi have been above described 
as rotating the humerus outwards; it must be borne in mind 
that as this bone is inverted, these muscles come to lie on the 
outer aspect of the limb, a circumstance which, no doubt, led 
Professor Owen into the error of confuunding the teres major 
with the deltoid? : 

(d) Extension of the forearm is of course effected by the 
action of the triceps, 

In the forward movement of the limb the resistance to be 
overcome is very much less, whence the muscles which are called 
- into action are of less robust proportions. The movement is, of 
course, merely a reversal of the preceding, viz., extension of the 
humerus upon the scapula; flexion of the forearm; internal 
rotation of the humerus and extension of the digits. Extension 
of the humerus upon ‘the scapula is effected chiefly by the 
anterior segment of the great pectoral, the deltoid and the 
supraspinatus, which muscles, assisted by the biceps, as explained 
below, rotate the humerus inwards. The digits are extended by 
the strong extensor communis and the elbow is flexed by the 
brachialis anticus and biceps, the latter functioning also as a 
rotator of the humerus in a very curious and interesting manner. 
It has been pointed out (p. 215) that the long proximal tendon of 


1 “The deltoid, coextensive with the scapula, acts through its length with great 
power upon the well-developed humeral ridge.” —Anatomy of Vertebrates, vol. iii. 
pp. 17, 18. 


218 MR R. AUSTIN FREEMAN. 


this muscle runs a zigzag course somewhat like a capital Z, and 
that the middle portion lies in a bony canal in the upper part of 
the humerus, the canal passing almost horizontally through the 
bone in its transverse diameter. 

From this it clearly follows that when, by the contraction of 
the muscle, the tendon is put upon the stretch, it will tend to 
become straight, and the result of this will be an external (or in 
the inverted position of the humerus an internal) rotation of the 
bone and consequent relaxation of the flexor sublimis ligament 
perinitting of extension of the digits, 

Thus it will be seen that the singular condition of the biceps — 
comes to be of material use in the curiously modified muscular 
movements. 


EXPLANATION OF PLATE V 


Fig. 1, Left scapula, dorsal’aspect. a, mammillary process at vertebral 
end of spine; 0, acromion; ¢, supra-spinous fossa; d, infra-spinous fossa. 

Fig. 2. Left scapula, ventral aspect. «a, bicipital impression ; 8, 
facet for trapezius ; c, facet for serratus magnus; d, ventral ridge. 

Fig. 3. Left coraco-clavicle, posterior aspect. a, articular surface 
for humerus; 0, posterior surface; c, conical process on ventral 
surface. 

Fig. 4. Left coraco-clavicle, anterior aspect. a, articular surface for 
presternum; 6, surface for deltoid; c, conoid process; d, foramen 
(coracoid foramen, Parker.) 

Fig. 5. Presternum and coraco-clavicles, ventral aspect. a, alar 
expansion; 0, thickening at posterior extremity; c, carina; d, 
promontory. 

Fig. 6. Presternum and coraco-clavicles, dorsal aspect. a, ala; b, 
posterior end ; c, groove; d, foramen. 

Fig. 7. Left humerus, anterior aspect. a, rhomboidal space 
occupied by deltoid ; 6, surface for part of great pectoral; ¢, bicipital 
notch ; d, termination of bicipital groove; e, ridge for teres major and 
Jatissimus dorsi ; /, pit marking origin of flexor sublimis digitorum ; g, 
pronator ridge ; h, lower opening of supra-condylar canal ; 7, external 
condyle; 7, capitellum; %, superior opening of bicipital canal; 7, 
articular surface for coraco-clavicle, 

Fig. 8. Left humerus, posterior aspect. a, head; 0, inner tuberosity ; 
c, outer tuberosity (articular surface for coraco-clavicle) ; d, ridge on 
tuberosity ; e, uncinate process ; f, ridge for part of great pectoral ; g, 
superior opening of bicipital canal ; h, bicipital groove leading down 


. 


ANATOMY OF THE SHOULDER AND UPPER ARM OF THE MOLE. 219 


to bicipital notch; 7, cavity in outer tuberosity; 7, nutrient foramen; 
k, trochlear surface. 

Fig. 9. Dissection of the muscles of the pectoral and axillary 
regions. On the right side the superficial, on the left the deep 
musclesareshown. P.M. 1-5, pectoralis major; the sixth and seventh 
fasciculi have been removed, the former in order to show the sub- 
scapularis, and the latter to show the pectoralis minor. P.Mi., pectoralis 
minor ; 8, subscapularis ; T.M., teres major; Bi., biceps; P.R.T., prona- 
tor radii teres; F.C.R., flexor carpi radialis; D., deltoid; E.O., 
external oblique ; F.S.D., flexor sublimis digitorum (converted into a 
ligament) ; E.C.D., extensor communis digitorum ; Ra., radius, 

Fig. 10. Dissection of the brachium and fore-arm from the inner 
side. Sc., Scapula (the axillary border and subscapular fossa shown). 
Tr. 1., scapular head of biceps ; Tr. 2, outer head; Tr. 3, inner head ; 
C.L. capsular ligament; Bi. 1, tendon of biceps above bicipital canal. 
Bi. 2, tendon of biceps emerging from canal; Bi. 3, belly of biceps ; 
LT., inner tuberosity ; A.I., anconeus internus ; P.R.T., pronator radii 
teres ; F.C.R., flexor ‘carpi radialis; P.L., palmaris longus; F.C.U., 
flexor carpi ulnaris, F.S.D., flexor sublimus digitorum. 


ON THE REPRODUCTION OF THE CARAPAX IN 
TORTOISES. By Hans Gapow, Ph.D., M.A. Cantab., 
Strickland Curator and Lecturer on Advanced Morphology 
of Vertebrates, University, Cambridge. (PLATE VI.) 


AMONGST a number of land tortoises (Testudo graca) kept here 
were many specimens which, owing to the rough mode of 
packing generally gone through by these hardy reptiles, arrived 
in a very injured condition. In some cases the greater part of 
the carapax was broken through, most of the plates having been 
so much cracked or crushed that they were dislodged from their 
neighbouring parts, and were rendered easily movable. This 
refers not only to the epidermal scutes, the so-called tortoise- 
shell, but likewise to the underlying ossified dermal plates. 

All the tortoises have since been kept under the most favour- 
able natural conditions, whereupon the maimed specimens 
showed the following interesting changes :— 

The epidermal scutes fell off, followed, after some three to 
ten months, by the thick osseous plates, which were completely 
atrophied and then raised above the old surface, until the greater 
portion of the old carapax was bodily lifted up and was kept in 
connexion with the animal merely by the overlapping margins 
of some of the neighbouring uninjured scutes. Underneath this 
old armour, and separated from it by a space partly filled with 
decaying matter, comes a layer of new tortoise-shell. This 
layer is of typical structure, and even contains the usual patches 
of black and yellow pigment. In bad cases the whole renewed 
area is covered with a mass of horny large tubercles without any 
regular arrangement. Underneath this shell comes a soft, vascu- 
lair, and apparently sensitive layer, resembling the malpighian 
normally found between the tortoise-shell and the ossified cutis. 
Underneath this soft layer lies a more or less well ossified 
armour. In one of the most injured specimens about four 
neural and six costal plates of the dermal armour were bodily 
lifted off (fig. 1). The animal was then protected only by the 
remaining subcutaneous connective tissue, which although 
already considerably changed and thickened, was still so soft 


THE REPRODUCTION OF THE CARAPAX IN TORTOISES, 22ak 


that it was bulging in and out according to the expiration and 
inspiration of the lungs. On this soft cover there was already pro- 
duced some irregular and imperfect tortvise-shell. Microscopic 
examination in transverse stained sections shows the following 
structure (fig. 2A):—s.c., stratum corneum, partly peeling off; the 
lower layers are composed of more flattened cornified cells, 
followed by high cylindrical cells (M), which, like the typical 
malpighian cells of Chelonians, show granulous contents with a 
large nucleus and nucleolus; P, black, star-shaped pigment 
clusters ; 0.c., ossified cutis, z.¢., new dermal armour. Sections 
through less advanced portions show the thick cutis (fig. 2B, ¢) 
composed of the normal horizontal connective fibres and full of 
osteoblasts. Towards the peritoneal or inner side these are by 
far less numerous and the fibres are still irregularly felted, 
as in ordinary subcutaneous connective tissue. 

In another case the epiplastron had been severed from the 
neighbouring entoplastral and hyoplastral plates. The opposite 
margins are now quite smoothly rounded off, and the epiplastron is 
now connected with the rest of the plastron by a belt of skin of 0°5 
em. in width, consequently quite movable, but of course attached 
to the skin of the neck and the M. claviculo-plastro-humeralis. 
When this animal withdraws its head into the shell, the new 
jointed epiplastron shuts part of the opening and thus bears a 
close resemblance to the mechanism of the genus Pyvis. The 
belt of skin cannot in its structure be distinguished from that of 
the neck and legs. 

In order to investigate this process, I made the following 
experiments on TZestudo greca. The specimens are now in the 
museum of comparative anatomy and zoology in Cambridge. In 
November 1884 a ring of three-quarters of an inch in diameter was 
cut through the whole carapax into the soft subcutaneous tissue. 
The central disk, which had thus become a movable plug, 
remained intact. The walls of the ring and disk were cauterised 
in order to destroy the margins of the malpighian layer. The 
wound was for the first fortnight dressed antiseptically, but 
afterwards left to itself. The plug showed the inspiration and 
expiration for several weeks, and remained movable for at least 
three months. Now, in September 1885, examination shows the 
following result (fig. 3) :-— 


29 DR HANS GADOW. 


The walls of the ring R looked as if the wound had not been 
mended, but after slight pressure a ring of necrotic dermal bone, 
D, came off and showed the continuation of the malpighian 
layer M. Dead tortoise-shell, together with a disk of dermal 
armour could easily be removed from the top of the disk. The 
dermal part D was only half its original thickness, resorbed and 
necrotic on its deep surface. Below it, a layer of newly formed 
tortoise-shell, followed by malpighian stratum and then by 
dermal bone. 

On another part of the tortoise a portion of the horny layer, 
together with the malpighian stratum, was completely removed 
to the extent of 0°3 inch square (fig. 4). This area reached 
across the natural suture of two neighbouring dermal plates. 
The injured place kept its appearance, apparently without change, 
for ten months. Now, with some force a thin layer of dermal 
plate could be removed; it was quite necrotic, and separated 
from the rest by a stratum of horny epiderm, which latter was 
continuous with the uninjured tortoise-shell. It must, however, 
be noted that this regenerated shell does not reproduce the 
peculiar pattern of concentric squares and rings visible on the 
uninjured scales, because this pattern is intimately connected 
with the growth of the whole animal. 

To sum up, the injury done to the armour of tortoises is mended 
in this way that from the healthy margins of the malpighian layer 
the latter grows centripetally towards the injured area right into 
the osseous plate and thus causes the latter’s partial destruction, 
whereupon the superimposed shell and osseous plates to the extent 
of the area to be mended are cast off. Lastly, the remaining deep 
portions of the dermis make up for the loss by thickening, If 
the injury is very severe, as in the tortoise figured, where the 
dermal armour was cast off down to the soft cutaneous layers, 
the bulk of these produces cutis, which then undergoes the 
normal process of ossification, until at last a uew complete 
armour is formed. 

A superficial centripetal growth of the epiderm over the defect, 
as in mammals, does not take place in tortoises. The reason 
for this rather roundabout process seems to be that the epider- 
mal products (the tortoise-shell) are not only the phylogenetically 
older but also a more important protection than the much later 


THE REPRODUCTION OF THE CARAPAX IN TORTOISES. 225 


established dermal armour. However, we do not know of any 
non-pathological cases where bony or dermal structure lies 
superficially without being covered by epidermal products, with 
the single exception of the horns of deer, but these have to be 
periodically renewed. 

Considering the enormous extent to which our tortoises have 
mended their shells, there may be after all some truth in the 
often ridiculed story related of Malayan turtle-fishers, who, after 
the living creature has been slightly roasted over a coalfire, to 
render the tortoise-shell easily removable, set the tortured turtle 
free so that it may recover and yield them a new coating. 

I confess that at first sight the process of regeneration 
described above seemed to afford at last an instance of epidermal 
products being developed from the surface of cutaneous layers. 
One might suggest that through the development of the various 
skin-glands epiblastic elements had been conveyed deeply into 
the cutis and that these latent or dormant cells might under 
certain circumstances establish a malpighian layer. Such an 
assumption might, perhaps, hold in mammals, but Chelonians 
are, like most other reptiles, almost devoid of skin-glands. Or, 
if the mesvblast of the amniota is derived from both germinal 
layers, the upper portion of the mesoderm then being a descend- 
ant from, or perhaps a differentiation of, the eztoderm, we might 
assume, that the cutaneous layer retains in itself the potentiality 
for developing structures which normally are produced only by 
the ectoderm specialised in that direction. Such cases seem to 
occur. Professor Humphry kindly informs me that in cases of 
burns or other injuries, causing the destruction of large portions 
of the skin, the whole often considerable defect of epidermis and 
cutis will be mended by a regeneration from the cutis per swper- 
jiciem, provided there be any of the deeper layers of the cutis 
left. An analogous case is the reproduction of bark from the 
whole surface of the cambium laid open after the destruction of 
the old cortex. 


periipk THE REPRODUCTION OF THE CARAPAX IN TORTOISES. 


EXPLANATION OF PLATE VI. 


Fig. 1. Transverse section through carapax; #., epiderm; O.C, 
ossified cutis; D., cast off portion of old dermal armour; M., newly 
formed malpighian layer and new tortoise-shell ; «, see fig. 2. 

Fig. 2. Section through regenerated portion of carapax at point a, 
fic. 1. A. s.¢., stratum corneum; JZ, cylindrical cells; P., pigment 
clusters ; O.C., ossified cutis. B. c., thick cutis. 

Fig. 3. Transverse section; £&., the ring cut out; D., cast off portion 
of dermal armour on disk; D,, cast off portion of wall of ring; O.C., 
ossified cutis or dermal armour, partly renewed at 7; Z., cast off por- 
tion of old epiderm ; M., malpighian layer, newly formed. 

Fig. 4. Transverse section; D., cast off disk of dermal armour, 
below which the newly-formed tortoise-shell. 


OBSERVATIONS ON THE DEVELOPMENT AND THE 
DECAY OF THE PIGMENT LAYER ON BIRDS’ 
EGGS. By AexanpEr M. M‘Atpowr, M.D., Vice- 
President, North Staffordshire Naturalists’ Field Club. 


Many and various are the theories which have been brought 
forward by ornithologists to account for the coloration of birds’ 
eggs, but all have failed to explain why the majority are so 
richly and beautifully ornamentec. White eggs, deposited in 
dark situations, are accounted for, as also are those which 
simulate the colour of the soil or other material on which they 
are laid. All others, comprising by far the largest proportion of 
egos, have proved an enigma as yet unsolved. Theorising on 
this interesting subject, it occurred to my mind that if we were 
to reverse the usual order of study, examine first the pigmenta- 
tion of those eggs whose colours apparently bear no relation to 
their surroundings, and afterwards investigate the causes which 
might have brought about the loss of pigment in white eggs, and 
also the method by which protective mimicry had been produced, 
we might arrive at a more satisfactory and scientific conclusion. 
A review of other organic objects gave abundant reason for this 
order of investigation. White is one of the rarest hues in the 
organic world. When it occurs it is for some specific purpose, 
and its presence can always be easily accounted for. Thus, 
animals which live within the arctic circle are white like the snow 
of these regions, the perianth of many flowers is white to attract 
insects and ensure cross fertilisation, and most sea birds are 
white, with blue on the back, resembling the colours of the 
clouds, and rendering them less likely to frighten the fish upon 
which they prey. 

Much has been written about the variety of colours observed 
on eggs, but a glance at the solar spectrum will at once show 
the fallacy of this opinion. The only portions of the spectrum 
represented on the whole range of colours of birds’ eggs are a 
small band on the red (forty to forty-five), and another on the 
green (eighty-five to ninety-two). 

This seems the more striking when we consider the number of 

VOL. XX. P 


226 DR ALEXANDER M. M‘ALDOWIE. 


colours exhibited by the birds themselves, where every part of 
the spectrum is represented, as well as by other organic objects, 
as insects, flowers, &c. Only two colours of pigment are found 
on eggs, green and red—together with black, which is not a 
colour in the scientific sense of the term. It is a remarkable 
fact, and one which I have never seen noticed by writers on the 
subject, that all the rich hues and shades with which eggs are 
so lavishly ornamented, are merely intermediates between these 
three. ? 

I have arranged the following scale by an expansion of these 
three according to chromatic rules, and it will be seen that it 
embraces every shade of colour found in birds’ eggs. 


BLUE-GREEN Russe? or Orange Brown 
Green — _ Brown 

Olive-Green Grey-Brown 

Olive Brown-Grey 
Olive-Brown BLACK or Grey. 


The hematin of the red blood-corpuscles is the source of these 
pigments. It is taken up by the pigment cells where it under- 
goes metamorphosis into the green, brown, and black shades, and 
is then secreted by the particular follicles at the lower part of 
the uterus. “To understand how yellow, green, brown, and 
black pigments may be derived from the colouring matter of the 
blood,” writes Rindfleisch, “we must first glance at the physio- 
logical metamorphoses to which this substance is liable. The 
most important of these, and in some sense typical of all 
the rest, is its transformation into bile-pigment. The red 
corpuscles, as they grow old, part with their colouring matter to 
the serum; from this it is taken up by the liver-cells, which 
transform it into bile-pigment; as such it is ultimately excreted 
in the feces. Before it is thus removed, when retained in the 
gall bladder for any length of time, it undergoes further changes, 
passing through shades of yellow, green, brown, and black, which 
Shadeler terms respectively bilifuscin (C,,H,)N,O,), biliverdin 
(C;,H,)N,0,,), biliprasin (C,,H,,N,O,,) and bilihumin ; bilifuscin 
differing from bilirubin in containing two atoms more HO, 
biliverdin from bilifuscin in containing two atoms more O, 


> 


DEVELOPMENT AND DECAY OF PIGMENT LAYER ON BIRDS’ EGGS, 227 


biliprasin from biliverdin, again, by an access of 2HO, while 
bilihumin is a black, insoluble, very highly oxidised substance.” 

“The scale of colours enumerated above serves, as already 
stated, as a standard for the course of all other chromatases, 
whether physiological or pathological.” 

A careful study of these pigments will show the reason for 
this limited range of colour. If we examine the characteristics 
of the pigment layer on eggs, it is impossible to resist the con- 
clusion that its presence has reference to the sun’s rays. All 
organic objects which are liable to be exposed to the sun’s rays 
are protected by pigment of one colour or another. This is the 
chief use of pigment in the animal world. In many animals the 
colours are adapted or moditied for concealment, but their 
primary use is for protection from the glare of the sun. The 
delicate and tender ovum in particular requires protection. 
Professor Yung has shown the generating power of the sun’s 
rays on the ova of frogs. In 1881, I read a paper showing that 
the pigment of frog’s spawn was placed in the ovum itself to 
absorb the sun’s rays as well as to protect the organism.’ But 
the ovum of the more highly organised warm-blooded animals 
does not derive the stimulus required for its development from 
the rays of the sun but from the body of the parent. I think 
that this affords us sufficient reason for assuming that the use of 
the pigmented covering of the shell is to protect the sensitive 
ovum from being acted on by the sun’s rays. If we make two 
perforations in a piece of cardboard, cover one with a piece of 
pigmented egg shell (eg., a rook’s or blackbird’s), and the other 
with a white one (¢g., a pigeon’s or a woodpecker’s), and hold 
the cardboard up to the light, we see the great amount of pro- 
tection afforded by the pigmented shell. True, the same end 
might be gained by thickening the shell, but it would have to 
be increased several fold to afford an equal amount of protec- 
tion, and nature is never wasteful in material. Moreover, owing 
to the peculiar way in which the yolk is suspended by the 
chalazz, the germinal spot is always uppermost, and consequently 
exposed to the sun’s rays striking from above. 

Mr Salvin noticed in Guatemala that humming-birds were 
much more unwilling to leave their nests during very hot 

1 North Staffordshire Natural History Reports. 


228 DR ALEXANDER M. M‘ALDOWIE. 


weather, when the sun was shining brightly, than during cool, 
cloudy, or rainy weather.' 

I shall afterwards show that there is a direct relation between 
the amount of light to which the egg is exposed and the 
intensity of its pigmentation ; that direct exposure to the sun’s 
rays is necessary for pigment to acquire its full development ; 
and that eges deposited in sites protected from the sun’s rays 
gradually lose their pigment layer. 

If we examine these three shades from a purely chromatic 
standpoint, apart from any physiological views of their 
development, we shall see how admirably adapted they are for 
the purposes of protection and concealment. 

1, Green is the colour we would naturally expect to find used 
as a protection for eggs. We know by experience that it has a 
peculiar softening influence on light. When it is present, it is 
always as a ground tint, uniformly spread over the surface of 
the ego. Most eggs are more or less covered with spots, blotches, 
or streaks, but these markings are never formed of green pig- 
ment. Many markings appear green to the eye, but this 
appearance is produced by the green ground tint being seen 
through a thin layer of black or brown. The spots in that case 
are of a darker green than the ground, but never deeper or 
richer. The most common tint is a bluish-green. 

2. Red. If we see a reason for the use of green pigment as a 
protective layer on eggs, we can find an equally good one why 
red should be employed as the second or supplementary colour. 
In the chromatic circle all colours are arranged in pairs, and the 
colours in every pair are complementary to each other. When 
two colours are seen in juxtaposition, they mutually affect each 
other, both in colour and tone. A yellow object, for example, 
placed close to a blue one, will appear as if inclined to orange, 
while the blue object will seem to incline towards violet. But 
two complementary colours, such as red and green, do not modify 
one another’s contiguity. They merely enhance each other's 
characteristics. Further, Professor Church remarks that green 
and red have a relation to each other which is different from that 
of any other pair of colours. “That there is something very 
peculiar in the relation of green to red,” he adds, “may also be 

1 Ibis, 1864, p. 875. 


DEVELOPMENT AND DECAY OF PIGMENT LAYER ON BIRDS’ EGGS. 229 


concluded from the frequency with which these two colours are 
confounded by persons who suffer from colour blindness.” The 
exact tint, which seems to be the basis of all the reds and browns 
found on eggs is a russet or orange-brown, and, if we examine 
the chromatic circle of Maxwell, this tint will be seen to be the 
complementary colour of the bluish-green, which I have just 
remarked is the usual ground tint. Red pigment is usually 
arranged in the form of spots, streaks, or blotches. Occasionally 
it is used as a ground tint—as in the grouse tribe—but then it 
is usually more or less speckled. It rarely if ever exhibits the 
smoothness or evenness of the green. 

3. Black seems to be employed mostly as a tone giving 
neutral to enhance these two colours. Orange or red seen in 
juxtaposition with black is rendered rather lighter in tone and 
more luminous; green with black becomes more brilliant, but 
the black suffers in purity, and appears slightly tinged with a 
ruddy hue. The grey marking seen on many eggs, eg., the 
sparrow tribe, are produced by a light layer of black pigment. 

Next to their limited range the variation of the colours of 
egos in the same species is the most striking characteristic. In 
the egos of several species, as gulls, terns, and guillemots, the 
range of lives extends almost from one end of the scale to the 
other. Whata contrast with the colours of the birds themselves, 
where the colour of a few feathers in some instances constitute 
the difference between the species, or with lepidoptera where the 
pigmentation of a few scales determines the name of the insect. 
And not only do the eggs of birds of the same species differ, but 
those of the same individual vary almost to the same extent. 
Thus in a tern’s nest, containing four eggs, I found one of a 
pale green colour and another of a deep reddish-brown. The 
other two were of intermediate tints. In this case the green egg 
had been the first laid and the brown one last. 

Basing our theories on the development of pigments referred 
to above, which is generally accepted by physiologists, we would 
infer that green was the first colour which was developed in the 
egos of the early species of birds. The eggs of the species 
extant support this conclusion. Green is the most common and 
most widely distributed colour. Schmidt states: “The more 
stubbornly a character is transmitted, or, what amounts to the 


230 DR ALEXANDER M. M‘ALDOWIE. 


same, the greater the number of families, genera, and species 
over which a character is extended, the earlier did it appear in 
the ancestral stock.” It is also,as has been shown previously, 
the colour best adapted for protection from the stimulating 
influence of the sun’s rays, and there can be no doubt but that 
this is the primary use of pigment. Almost every egg which is 
laid in a situation where there is no need for concealment, but 
which is exposed to the sun’s rays-—e.y., rooks, herons—is of a 
green colour. 

Red or russet, and all the intermediate tints, appear to be 
developed chiefly for concealment, the different shades of brown 
showing in many instances perfect adaptation of the colour of 
the egg with that of its surroundings. This colour is chiefly 
' exhibited on eggs which are deposited on or near the ground, 
as the waders’, gallinaceous birds’, larks’, &c. Hewitson states: 
“We should scarcely expect to find the eggs of the crane so 
entirely different from those of all the other species which are 
most nearly allied to it in habit and in form. Whilst the eggs 
of all these species, with the exception of those of the spoon- 
bill, are either pure white or slightly tinted with colour, but 
always spotless, those of the crane are, on the contrary, richly 
coloured.” This difference may be completely accounted for by 
the above theory, as the crane habitually breeds on the ground, 
whilst the others choose elevated sites, Compare also the 
eggs of the three species of divers, laid on the margin of the 
freshwater lochs in the north, with those of their congeners the 
guillemots, deposited on cliffs. 

The coloration of the eggs of the Falconide are in some 
respects exceptional, depending on the nature of the food and 
other causes, which lie beyond the scope of the present paper. 

When I examined eges with reference to the amount of light 
to which they were exposed, I found that the ratio between the 
intensity of the pigmentation and the degree of exposure was 
very marked, and, indeed, almost startling. For example, the 
eggs of birds which breed early—as the thrush tribe, hedge- 
sparrow, &c.—have well-developed ground tints; whilst those of 
the later breeders—as the green finch, linnet, &c.—laid after the 
leaves are out, and therefore screened from direct sunlight are 
more faintly coloured. My own experience is that exposure or 


DEVELOPMENT AND DECAY OF PIGMENT LAYER ON BIRDS’ EGGS. 231 


shelter from the sun’s rays plays an important part in the 
selection of a site for nidification, by the parent bird. 

It may be laid down asan almost universal law in ornithology, 
that eggs which are deposited in situations exposed to the sun’s 
rays are much darker in colour than those laid in nests protected 
from direct sunlight. They almost invariably possess a well- 
developed ground tint. Some, eg., the heron and the hedge- 
sparrow, are spotless, but most show markings of some kind. 

Eggs laid in shaded nests, e.g., the yellow-hammer and the 
green finch, possess a faint ground tint, and the markings are 
usually smaller and lighter than in the preceding. 

Eggs laid in covered nests, as the tits’ and wrens’, usually 
present faint spots in a white ground. 

Eggs which are wholly excluded from light, as woodpeckers’ 
and kingfishers’, are almost invariably pure white. 

Not only does direct sunlight seem necessary for the pigment 
layer to acquire its full development, but there is also strong 
evidence that all white and faintly coloured eggs have undergone 
or are undergoing a process of decolorisation when the protec- 
tion afforded by the pigment layer is no longer required. This 
is in accordance with the laws of physiology. If any tissue or 
organ loses its function it will gradually waste, and finally 
disappear Moreover, it has been shown by evolutionists 
that the lost organ is apt to appear as a variation, or as a 
rudimentary and useless appendage: many eggs, therefore, show 
rudimentary pigmentation, and colour appears sometimes as a 
variation in eggs which are normally white. Eggs have been 
divided by Wallace into white and coloured, spotted and un- 
spotted ; there is, however, an unbroken series between white 
eggs and those which are highly pigmented and _ spotted. 
Further, the eggs of several species would have to be included 
at one time under the former category, and at another under 
the latter. For instance, the egg of the white-tailed eagle is 
usually white, but occasionally it presents well-defined markings. 
The egg of the puffin is as often colourless as it is pigmented. 
That of the whinchat is of a bright blue-green, sometimes spot- 
less, sometimes faintly speckled at the large end with rust 
colour. 

It has been stated that white and light coloured eggs are found 


Van DR ALEXANDER M. M‘ALDOWIE. 


in dark and sheltered situations because colour is not necessary to 
conceal them from observation. The fact has, however, been 
overlooked, that eggs laid in elevated sites, as gulls’, rooks’, 
herons’, &c., have well-developed colours, although they can be 
of no use for concealment. Want of the stimulus of the sun’s 
rays alone causes the colour to fade or disappear. 

When one (or one or two) species in a family, where the 
majority lay deeply pigmented eggs, lays either a white egg or 
one faintly coloured, we invariably find that it differs from the 
others in its mode of nidification, depositing its eggs in some 
place protected from the light, whilst its relations lay in exposed 
situations. In this case there is what may be termed a specific 
decolorisation, the loss of pigment affecting the individual 
species alone, and not extending to the other members of the 
family. 

When the whole of a genus or family lay white eggs, we find 
that either all, or the majority of the species, deposit their eggs 
in places protected from the light. In this case there appears 
to have been a generic decolorisation affecting the group as a 
whole, the pigment having become obsolete at a much earlier 
period than in the preceding case, probably before the differentia- 
tion of the family into the existing number of species. 


SPECIFIC DECOLORISATION, 


In examining the evidence of decolorisation of eggs, it will 
be necessary to enter somewhat into the details of the modifica- 
tion of several groups of birds to show the various ways in which 
the eggs are protected from the stimulating influence of sunlight. 
With reference to this, eggs may be divided into three classes 
viz. :—(1) those laid in holes or covered nests, (2) those which 
are covered by the parent bird with leaves, weeds, &c., and (3) 
those covered by an incrustation of calcareous matter. The eggs 
of nocturnal birds are white or faintly coloured but are included 
in the above. 

1. Eggs deposited in holes or covered nests. 

The egg of a woodpecker, or any bird where the whole of 
the family lay white eggs and are identical in their mode of 
nesting, affords no evidence of decolorisation. But when we 
find one species only in a family laying in a covered nest, and 


DEVELOPMENT AND DECAY OF PIGMENT LAYER ON BIRDS’ EGGS. 233 


its eges white or faintly coloured, whilst those of the other 
members are laid in open sites and are highly pigmented, we 
may fairly argue that the pigment covering has degenerated or 
vanished. Thus all the thrush family lay richly coloured eggs 
in open sites, whilst the dipper, a closely allied species, lays a 
white egg in a domed edifice. The egg of the black redstart, 
one of a genus where all the other members lay coloured eggs, 
is exceptionally pure and white. Again, the auk tribe is notable 
for laying bright coloured eggs on bare ledges on the cliffs ; 
whilst the puffin, one of the family which breeds in rabbit holes, 
lays an egg, described by Hewitson as “sometimes spotless, but 
more frequently marked with various tints of colour, but so very 
faint and indeterminate as to appear as though they were seen 
through the shell.” The wheatear is a good example of an egg 
which has undergone almost complete decolorisation. One of 
the most interesting cases is that of the Virginian quail. This 
is the only one of our gallinaceous birds which builds a dome- 
shaped nest, or, indeed, may be said to build any nest at all. 
Yarrell states that the eggs are white, although Hewitson figures 
one of a faint buffy tint, with minute spots. All the rest of 
this order lays eggs with a well-developed ground tint, and 
usually richly covered with dark-coloured markings. 

2. Eyys covered by the parent bird with leaves or other vegetable 
matter. 

There is no doubt but that a loss of pigment has resulted 
from this mode of concealment. It is interesting to speculate 
on the reasons which have led certain birds which breed on the 
ground to adopt this method of protection, whilst others trust to 
the colour of the eggs. Inthe grebe tribe it seems the only 
method possible. Protective coloration of the eggs would be a 
much less effective mode when they are deposited on the top of 
a large and prominent heap of decaying water plants; and it 
would be impossible for them, on account of their peculiar wings 
and legs, to escape from the nest unless it were close to the 
water’s edge. “It would seem that whatever they do must be 
done in the water,” writes Naumann; “they cannot even rise 
upon the wing without a preliminary rush over the surface of 
the lake; from dry land they cannot commence their flight.” 
The loss of pigment is complete in the grebes, as it is probable 


284 DR ALEXANDER M. M‘ALDOWIE. 


that the wet decaying vegetation shuts out the light more com- 
pletely than the loose dry materials employed by the ducks, 
pheasants, &c. 

In the duck tribe this method has probabiy been adopted 
because of the size and large number of eggs laid. Protective 
coloration could not have afforded sufficient concealment for 
eight or ten or even more largeeges. The amount of decoiorisa- 
tion varies in this tribe. In some instances, as the eider duck, 
the egg is of a pale asparagus green, in others it has only a very 
faint greenish hue, whilst in the shieldrake, which breeds in 
holes, the loss of pigment is complete; the egg being of a “ smooth 
shining white.” . 

In both the above groups the whole of the family adopt the 
same mode of concealment, but why should the eggs of the 
pheasants and partridge be correct, whilst those of the grouse 
are laid openly? The two former birds breed in sheltered woods 
and hedgerows, the latter on bleak and exposed moors. If the 
grouse covered its eggs with dead vegetable material it would 
soon be carried away by the strong winds which sweep over 
the moors. 

3. Hogs covered by an incrustation of calcareous matter. 

Only three British birds’ eggs are coated with this peculiar 
chalky substance, viz., the cormorant, the shag, and the gannet. 
The hard shell beneath is of a faint bluish-green colour in the 
two first-mentioned species ; in the last it is usually pure white, 
but sometimes tinged with blue-green. 


GENERIC DECOLORISATION. 


The fact that certain birds deposit white eggs in fully exposed 
situations has been pointed out as proofs that the coloration 
could have no reference to the exposure to light. But no notice 
has been taken, as far as I am aware, of the fact that all these 
instances occur in families where the majority of the species 
breed in holes or dark places. In this case decolorisation must 
have taken place at a much earlier period in the life-history of 
the family or genus than in the preceding instances. They are 
probably descended from some ancestor which bred in holes; 
and the change to open nests in a few members of the family 
taking place long after the colouring matter had disappeared 


DEVELOPMENT AND DECAY OF PIGMENT LAYER ON BIRDS’ EGGS, 235 


has not been followed by a restoration of the pigmentary cover- 
ing. For proofs that pigment had at one time existed in these 
groups, we must look to the eggs of allied species. They occur 
in the owls, the pigeons, and the petrels. 

“There is a strong and perfect similarity amongst the eggs of 
the different species of owls,” writes Hewitson, “ which we could 
scarcely expect to find in the eggs of birds which differ so much 
from each other in their mode of breeding. The eggs of those 
species which are deposited in the hollows of old trees and 
deserted ruins, and those which are found on the bare sod, and 
exposed to the broad light of day and the pelting storm, are 
alike without colour.” But the large majority of the species 
breed in dark places, and, being nocturnal in their habits, all 
have a tendency to avoid light. Now their nearest congeners, 
the harriers, which link their family with the Falconide, bear a 
close resemblance to them in many points, ¢.g., the loose and 
flocculent character of the feathers, and the circular arrange- 
ment of those about the face ; and the affinity on comparing the 
skeletons of each is most decided. In the coloration of the 
eggs also the resemblance exists, the eggs of the harriers being 
white, or sometimes a pale skim-milk colour, more rarely spotted 
and smeared with brown. 

All the Columbide, or pigeon tribe, lay two pure white oval 
egos. All, however, lay in crevices in rocks, hollow trees, 
deserted rabbit burrows, or dense thick trees or bushes. This, 
together with the fact that the nests of the arboreal members 
are crude platform-like structures, quite unlike those of any 
other bird, show that they are descended from an ancestor which 
bred in holes. One of the Australian ground pigeons is said to 
lay buff-coloured eggs. The nearest ally to this order, Pallas’ 
sand grouse, lays three oval eggs, similar to a pigeon’s, but 
coloured like a plover’s. 

The petrels are all more or less nocturnal in their habits, and 
all lay white eggs! The fulmar deposits its egg openly, on 
ledges on the cliffs, but the other members of the group lay in 
crevices in the rocks, under stones, or form burrows to the depth 


1 Since this was written Mr Bladen has called my attention to a mass or ring 
of very faint markings near the large end of the eggs of some of the petrels. 
These, which appear to have been hitherto overlooked by writers on the subject, 
are the best examples I have yet seen of rudimentary pigmentation. 


236 DR ALEXANDER M. M‘ALDOWIE. 


of two or three feet. Ali the rest of the Laride lay richly 
coloured eggs. ; 

It will be seen from the above that the decolorisation takes 
place in three ways. In some—swallows, wrens, tits, &c:-—the 
ground tint disappears first, leaving the egg more or less thickly 
marked with small light coloured spots; in others—starlings, 
little auks, &ec.—the markings vanish first; while in a third 
class—puffin, hen-harrier, &e.—both ground tint and spots appear 
in a rudimentary degree. 

In this paper reference has only been made to the eggs of 
British birds, not only because the fauna of these isles form a 
very complete and typical group, but because the views adduced 
are based solely on the study of thousands of specimens of 
British eggs in my own and other collections, and upon observa- 
tions made on the moors, in the woods, and by the seaside. 

Evidence has been brought forward to show that the pig- 
mentary coat on birds’ egys came into existence at a very early 
period in their life-history, and existed in the eggs of the 
progenitors of all the extant species. It has also been shown 
that the range of colours on birds’ eggs is very limited, but 
follows the usual course of pigmentary changes; that the pig- 
ment is unstable and variable, making the process of change and 
decolorisation a simple one; and that its primary use is for 
protection from the solar rays, but that it afterwards became 
modified for concealment. 

Lastly, it has been shown that eggs acquire a highly developed 
pigmentary layer, or lose their pigment entirely, according to 
whether they are exposed to the full glare of the sun or laid in 
situations inaccessible to its rays, and that the intermediate 
degrees of coloration are in direct ratio to the amount of light 
to which the eggs are exposed. 

The two causes which determine the coloration of eggs— 
protection from the sun’s rays and concealment from observation 
—act conjointly ; they are not antagonistic like natural selection 
and sexual selection. The limited range of colours shows that 
natural selection alone operates. Darwin states that, in regard 
to structures acquired through ordinary or natural selection, 
there is a limit to the amount of advantageous modification in 
relation to special ends; but in regard to structures acquired 


* 


_ DEVELOPMENT AND DECAY OF PIGMENT LAYER ON BIRDS’ EGGS. 237 


through sexual selection there is no definite limit; so that, as 
long as the proper variations arise, the work of sexual selection 
will go on. That the causes are different from those which 
produce the colours of the birds themselves, is shown by the fact 


that eggs from tropical regions do not surpass in brilliancy of 
tint those of more temperate climes. 


THE CONNECTION OF THE OS ODONTOIDEUM 
WITH THE BODY OF THE AXIS VERTEBRA. By 
D. J. Cunntncuam, M.D., Professor of Anatomy and 
Chirurgery in the University of Dublin. 


(This paper was read before the Biological Section of the British Association 
in Aberdeen, September 1885. ) 


THE point in connection with the os odontoideum which I have 
to bring under the notice of this section is a small one, but still it 
is not without some interest and also morphological significance. 

For some time past I have been engaged in an investigation 
into the curves of the spinal column of man and the apes, and 
in carrying out this work I have made mesial sections of a large 
number of {frozen human spines. Very soon my attention was 
attracted to a small lenticular-shaped plate of cartilage, which 
seemed in almost every case to be interposed between the os 
odontoideum and the body of the axis vertebra. On all sides 
it was surrounded by bone, so that it could only be brought into 
view by means of section. To obtain a proper conception of the 
significance of this cartilaginous plate, it will be necessary that 
we review some points in connection with the development of 
the axis vertebra, and its tooth-like process. 

The identity of the os odontoideum with the absent body of 
the atlas is, so fully and satisfactorily established, that it is 
unnecessary for me to do more than merely allude to it. 
Whilst this is the case, however, in man, the os odontoideum 
and the body of the axis in their early condition form one 
continuous cartilaginous mass, with no external marking by 
means of which the one can be distinguished from the other. 
Both are traversed by the notochord and the only evidence of 
their separate nature is to be found in a notochordal swelling or 
dilatation at their point of junction. 

The manner in which this cartilaginous mass is ossified is 


! Robin, Lvolution de la Notocorde, Paris, 1868. Miiller, Veber das vorkommen 
von esten der chorda dorsalis beim Menschen. nach} der Geburt, &c., &e. 
Zeitschrift fiir rat. med., N.F¥., Band_11. 


° 
CONNECTION OF OS ODONTOIDEUM WITH BODY OF AXIS VERTEBRA, 25 


very instructive. The body of the axis is converted into bone 
by one primary centre and an epiphysary lamina, which is 
formed upon its lower surface. The os odontoideum ossifies by 
two lateral primary centres, which soon fuse, and an apical 
epiphysary centre for the summit. The last undoubtedly 
represents the upper epiphysary plate of the atlas vertebra. 

The apparent differences, then, between the ossification of the 
os odontoideum and the centrum of the axis on the one hand, 
and that of the corresponding parts of the vertebree lower down 
on the other, resolve themselves into two. 

1. The absence of an epiphysary centre for the lower surface 
of the os odontoideum, and for the upper surface of the body of 
the axis vertebra. 

2. The double primary centre which appears for the ossifica- 
tion of the odontoid. 

Both of these discrepancies can be more or less easily 
explained. With regard to the first, Macalister! has described 
both of the absent epiphysary plates as being present in the axis 
vertebra uf the Balwnoptera rostrata, and he likewise states that in 
some cases they can also be detected in man. Further, 
Humphry has observed and figured one of the absent nuclei in 
the rabbit.? 

The second point of difference is more difficult to understand. 
It is true that many anatomists deny the double character of 
the odontoid centre; they see no material difference between 
this centre and that of the other vertebral bodies. Thus 
Gegenbaur, Henle; and Blandin give expression to this view, and 
Robin * insists strongly upon it. It is a conclusion, however, 
which we cannot adopt. On the other side we range Meckel, 
Luschka, Sappey, Cruveilhier, Quain, Macalister,* and several 


1 Macalister, Jour. Anat. and Physiology, vol. iii. p. 54. 

? Humphry, The Human Skeleton. 

3 Gegenbaur, Lehrbuch der Anatomie des Menschen, 1883. Henle, Hand- 
buch der Anatomie des Menschen; Erste -Abtheilung, Knochenlehre, 1871. 
_ Blandin, Nouveaux Elements d’ Anatomie, 1838, vol. i. p. 39. Robin, ‘‘Sur le 
développment de vertébres,” Jour. d Anat. et Phys., Paris, 1864. 

4 Meckel, Descriptive and Pathological Anatomy (translated by A. S. Doane), 
1839. Luschka, Die Anatomie des Menschen. Erster Band, p. 32, 1863. Sappey, 
Traité @ Anatomie, Tome premier, p. 312, 1876. Cruveilhier, Anatomie 
descriptive, 1871, vol. i. p. 78. Quain’s Anatomy, 9th edition. Macalister, loc. 
cit. 


240 PROFESSOR D. J. CUNNINGHAM. 


others, all of whom believe in the originally double condition of 
the odontoid ossific centre. In support of this I may bring 
under the notice of the section an extremely interesting specimen 
of an axis vertebra, from the Pathological Museum in Trinity 
College, Dublin, in which the os odontoideum is double. In other 
words there are two distinct odontoid processes springing from 
the upper surface of the body of the axis vertebra. Such a con- 
dition could only have been produced by the presence of two 
ossific centres which have failed to unite. 

Granting, then, the double centre for the odontoid, how can 
this deviation from what is generally considered to be the 
typical mode of ossification of a vertebral body be accounted for. 
Humphry has offered the best solution of the question. It is 
not uncommon to find one or more vertebral centra cleft into 
two lateral parts, clearly showing that they have been ossified 
from two centres. The very specimen to which I have alluded 
is an example in point, because not only is the odontoid bone 
cleft into two, but so also is the body of the axis. In view of 
abnormalities of this kind, Humphry has been led to suppose it 
possible for the ossification of a vertebral body to take place 
by two twin centres which fuse almost immediately after their 
appearance. 

For a considerable time the os odontoideum is separated from 
the body and pedicles of the axis vertebra by a layer of cartilage. 
It is essential that we should study the changes which this 
plate of cartilage undergoes. The accounts given of these by 
different authors are at utter variance with each other 

According to Henle,? Cruveilhier,? and Quain,? we are led to 
believe that the cartilage disappears at the third year, and that 
the osseous union of the odontoid and axis vertebra is then 
complete ; according to Sappey? and Robin? the union is accom- 
plished more slowly, and is not perfected uatil the sixth or 
seventh year; Humphry, on the other hand, delays the consoli- 
dation until puberty. 

The observations which I have made on this point would 
seem to indicate that it is not until old age is attained that 


' This specimen has been figured and described in its pathological aspects by 
Prof. Bennett in the Zrans. Path. Soc. Dublin, vol. vii. p. 117. 
2 Loc. cit. 


M 


- 


CONNECTION OF OS ODONTOIDEUM WITH BODY OF AXIS VERTEBRA. 241 


complete osseous union between the os odontoideum and the 
body of the axis takes place. 

In sixteen out of eighteen axis vertebrae which were obtained 
in the Trinity College practical anatomy room, without reference 
to age or sex, a small disk of cartilage was found on section, mark- 
ing the line of separation between the odontoid and the second 
vertebra. In dealing with dissecting room subjects, it is ex- 
tremely difficult in Dublin to obtain reliable information regard- 
ing the age of any particular individual. Three of the specimens 
examined I have been obliged to eliminate, as I was quite unable 
to arrive at any knowledge of the age of the subjects from which 
they were taken. Of the remaining fifteen (which include the 
two bones in which the cartilage is absent) I was only able to 
ascertain the precise age of two; whilst in the case of the others 
notes were made of the probable age of the subjects as they 
came in. 

The fifteen vertebree which remain may be divided into three 
groups according to their age. 

Group I.-—Axis vertebre obtained from subjects varying from 
twenty-four to fifty years of age——This group comprises six 
vertebree—two from females and four from males. The youngest 
specimen was taken from a girl of about twenty-four years old ; 
the others were obtained from subjects varying in age from forty 
to fifty. In all the members of this group the cartilaginous 
plate was present. In sagittal section its average length was 4 
mm., and its average thickness 2 mm. 

Group II.—Azis vertebra obtained frum subjects varying in age 
from fifty to sixty years——In this group there are only three 
specimens—all from females. In each the plate of cartilage is 
present, and its average dimensions are the same as in Group I. 

Group III.—Age ranging from sixty to seventy years—Six 
specimens—two males and four females. In four members of this 
group the cartilaginous dise was present, and in sagittal section 
its average dimensions were: length, 3 mm.; thickness, 1} mm. 
In two it was absent; one of these is marked as being taken 
from a subject seventy years old, whilst in the case of the other 
the age was put down as sixty-five. They were the oldest 
specimens examined. 


But how can these facts be reconciled with the statement 
VOL, XX. Q 


249 PROFESSOR D. J. CUNNINGHAM. 


made by some that the os odontoideum unites to the axis at the 
third year, and by others that the two bones are consolidated at 
the seventh year. Both of these assertions are, from one point 
of view, more or less correct. From the third to the fifth year 
the base of the odontoid joins on each side with the pedicles 
and the body of the axis, obliterating at these points the inter- 
vening cartilage. The middle portion of the cartilaginous plate, 
however, still remains and extends from the anterior to the 
posterior face of the bone. At the seventh year (often much 
later) consolidation takes place, first in front and afterwards 
behind, and the cartilage is thus entirely cut off from the surface. 
But the ossific union is restricted to the periphery ; the further 
process is extremely slow, and is apparently not completed until 
an advanced age is attained. Robin, therefore, is in error, in so 
far as he has mistaken peripheral consolidation for complete 
union. Cruveilhier, Henle, and Quain, with much less reason, 
mistake lateral consolidation for complete union. Humphry’s 
account of the ossification of this bone is more accurate. 
He defers complete consolidation until puberty, and he speaks 
of a space which remains in tke centre, which resembles 
the intervertebral spaces seen in section of the sacrum. If a 
dry bone be cut with a fine saw, this space can, as a rule, be 
made out; it is the cavity which contained the cartilaginous 
plate. 

In order to determine the nature of the disk of cartilage which 
persists so obstinately in the interior of the axis vertebra, I 
decalcified the specimen I obtained from the girl of twenty-four, 
and cut the odontoid process and the body of the axis in the 
longitudinal direction for microscopical examination. The 
cartilage was then seen to be of the hyaline type throughout 
its entire extent. In some of the sections faint traces of a 
feeble and sluggish ossific process might be detected around the 
margins of the cartilage. Not a vestige of the notochord could 
be discerned. Robin states that the notochordal swelling, at 
this point, disappears when ossification sets in, and that it is 
replaced by a small mass of fibro-cartilage which shades off 
on all sides into hyaline cartilage. Henle also refers to a 
fibro-cartilaginous layer in the very young bone. There is 
not the slightest appearance of this in the sections which I have 


> 


CONNECTION OF OS ODONTOIDEUM WITH BODY OF AXIS VERTEBRA. 243 


made of the adult bone. This is strange, seeing that fibro- 
cartilage is generally looked upon as the descendant of hyaline 
cartilage, and it is difficult to conceive a reverse development. 
The morphological significance attached to the obstinate 
persistence of this cartilaginous plate in the axis vertebra of 
man is very apparent :— 

1. The foreign character of the odontoid bone as an element 
of the axis vertebra is more forcibly insisted upon. 

2. It establishes a condition of the axis vertebra which is not 
so far removed from that of the Monotreme and Thylacine,! in 
which the union between the odontoid and axoid body is 
exceedingly late—if indeed it ever occurs at all. 

3. It establishes a more obvious basis for comparison with 
those reptiles in which the odontoid persists as a separate bone. 


wy 


1 Report on The Marsupials, by D. J. Cunningham, in the Reports of H.M.S. 
‘**Challenger,” Zoology, part xvi., 1881. 


THE SKELETON IN GEOCOCCYX. By R. W. SHuvFELpr, 
Med. Dept. U.S. Army ; Membr. Am. Ornithologists’ Union ; 
The Am. Soc. for Psychical Research ; Philosophical Soe. 
of Washington ; Cor. Membr. Soc. Italiana di Antropologia 
Etnologia e Psicologia Comparata, Florence, Italy. (PLATES 
VAG VITL; LX;) 


Wuen I came to North-Western New Mexico, early in the 
autumn of 1884, I expected to enter the very heart of the home 
of Geovoccyx californianus. Much to my surprise, however, after 
a residence of seven and more months in tbe country, not a 
single individual of this species has rewarded my many moun- 
tain and prairie excursions. Not to be foiled in securing 
specimens of its skeleton to complete my papers upon the 
osteology of the birds of this country, and more especially those 
peculiar to it, I sent a considerable number of letters to 
collectors in Texas, Arizona, and California. Many weeks of 
patient waiting brought me answers to only three of these, and 
all from the last named state. One said the bird was no longer 
to be found in his neighbourhood, while from my other two 
correspondents I secured each a skeleton, but both informed me 
that owing to the great demand on the part of collectors for the 
“Road Runner,” it was rapidly becoming a rare bird in localities 
where formerly it was very abundant. My last specimen, an 
old male, afforded me a perfect skeleton, and from it I made the 
drawings which illustrate this account, while the second bird, 
though not meeting my expectations so fully, still gave a 
skeleton, which has proved of great service to me, in the way of 
comparison and verifying points present in my type. 

The aim of this memoir is simply to give a full description of 
the skeleton of Geococcyx, and in it I will undertake but few 
comparisons. Some day in the future it is my hope to enter 
more extensively upon the entire anatomy of this form, so full 
of interest to all biologists. 

Of the Skull (Plate VIL. figs. 1, 2, 3, and 4).—In Geococcya we 
find the osseous superior mandible with a gently curved and 
rounded culmen, the curve increasing very modestly as it 


. 


a 


THE SKELETON IN GEOCOCCYX. 245 


approaches the apex. This part of the skull has a broad 
base, being both deep and wide in the rhinal region, while 
on all aspects it tapers gradually to the slightly decurved 
tip. Its buccal surface is flat, with cultrate edges some- 
what raised above the general plane behind. Posteriorly, 
this face is encroached upon by the palatines and maxillo- 
palatines. Turning to the lateral surfaces of this mandible 
(fig. 1) we find them for the most part to be slightly convex 
throughout their extent; the only exception to this being seen 
in the depressions which are found, one over each of the scale- 
like projections that close the hinder two-thirds of either nostril. 
These last mentioned openings are of a sub-elliptical outline, 
placed longitudinally nearer to the edge of the beak than its 
culmen, and just posterior to its middle. They do not directly, 
communicate with each other, but are external apertures, in this 
bird, of osseous tubes, one on either side, which are produced 
backwards nearly to the rhinal chamber, being encased in the 
loose osseous spongy mass that almost fills the otherwise 
hollow superior mandible of Geococcyzx. 

The cranio-facial hinge enjoys considerable mobility, and its 
position is clearly indicated by a transverse track. Mesially, 
this region is depressed, and may show the last sutural traces of 
the nasal processes of the premaxillary therein. Each nasal 
bone has been so completely met by the various surrounding 
elements, that, save its hinder margin, its boundaries are hard 
to define in the adult bird (figs. 1, 3). To the inner side of each, 
and just beyond the cranio-facial track, we observe on either 
side a small circular foramen, which is constant, and is to be 
found in like locality in Ceryle.1 All three sides of this osseous 
superior mandible is more or less marked by anastomosing vena- 
tions, and a few perforating foramina are always seen near its 
apex. 

A lacrymal in Geococeyx is an unusually large bone, though a 
light one, due to its very open cancellous structure within, and 
its being, perhaps, pneumatic besides. Superiorly, it articulates 
with the frontal and nasal, principally with the last on the lateral 
aspect, though it departs from it some time before reaching its 


1 “ Osteology of Ceryle Alcyon,” by the author, Jour. of Anat. and Physiology, 
April 1884, Plate XIV. figs. 1 and 2, nf. 


246 DR R. W. SHUFELDT. 


lowest point, where a slit-like interval is seen between the two 
bones. Below, its broad rounded margin is placed obliquely, its 
outer and at the same time posterior end, resting upon the upper 
side of the maxillary, while its inner and anterior end being 
elevated just above the superior surface of the corresponding 
palatine. 

The posterior aspect of the lacrymal is concave from above 
downwards, in conformity with the somewhat globular concavity 
of the orbit, while anteriorly it is correspondingly convex in the 
same direction. It lies in front of the broad, quadrilateral 
ethmoidal wing which overlaps it, the two forming a very 
complete partition between the orbit and rhinal chamber; the 
bone under consideration closing the outer third of the space. 

The ethmoidal wing, the form of which I have just given, is 
pierced above, immediately beneath the frontal bone, by two 
elliptical foramina, the inner one being the larger, and both 
being placed vertically. They probably transmit the olfactory 
nerve and vessels to the rhinal space. 

This “pars plana” has, like the lacrymal, also a somewhat 
cancellous internal structure, the plate being moderately thick. 
Its lower and outer margins are concave and smoothly rounded off. 

The expanded anterior extremity of a mazillary is immov- 
ably sandwiched in between the nasal above, and the posterior 
dentary process of the premaxillary beneath. Its rod-like 
extension behind forms about the anterior third of the very 
straight quadrato-jugal bar. The horizontally expanded end 
alluded to is quite ample, and may be perforated by numerous 
foramina. Its maxillo-palatine development will be described 
when speaking of the under side of the skull. 

The remainder of the quadrato-jugal bar becomes gradually 
larger and club-shaped as it nears the quadrate bone, to rather 
abruptly turn inwards as it reaches it, and is inserted in a 
vertical notch in the usual apophysis of that element, which 
projects directly outwards to meet it (fig. 3). 

With respect to the guadrate, we find that its orbital process 
is very broad and flat, being at the same time very short. The 
body of the bone is also broad, while its mastoidal apophysis is 
twisted in a way common to many other birds, and supports at 
its summit two articular heads with a distinct valley between 


° 
THE SKELETON IN GEOCOCCYX. 247 


them, At the inferior aspect of the mandibular foot, there are 
two condyles for articulation with the lower jaw. The inner and 
smaller of these is hemi-ellipsoidal in form, with its major axis 
in the same straight line that constitutes the longitudinal axis of 
the corresponding pterygoid. If this axis be produced the other 
way, itis found to be at right angles to the long axis of the other 
and larger facet of the mandibular foot of the quadrate. Rather 
a broad notch separates these two condyles from each other. 

The quadrate is a thoroughly pneumatic bone, and a large 
foramen is always found upon its posterior aspect half-way 
between the mastoidal head and the mandibular foot. 

Both the sphenotic and mastoid processes are well developed 
in this bird; they are of about an equal size, the first being 
directed downwards, and the last downwards and forwards. 
Between them, and carried well to the rear, is a sharply defined 
and rather deep crotaphyte fossa. It is separated from a like 
depression of the opposite side, by an interval of one and a 
half centimetres. These crotaphyte fosse are fully as well 
marked in (reococcyz as they are in many of the Laridea, 
and better than they are in some members of that group 
of birds, better for instance than they are in Chroicocephalus 
philadelphia. 

Owing to the great breadth of the frontals, the orbit is com- 
pletely sheltered above by an arching roof; the outer periphery 
of which is concave inwards, and bounded by a sharp edge. 
This orbital vault usually shows posteriorly a few perforating 
foramina. 

The rostrum of the sphenoid is pneumatic and rounded for its 
entire length beneath. It barely extends beyond the broad 
ethmoidal wings in front, and ascends but little as it proceeds 
in that direction. 

The inter-orbital septum is a thin partition of bone, which 
always possesses a considerable quadrilateral vacuity near its 
centre. This usually merges with the foramen for the exit of 
the optic nerves (fig. 1), while the small foramen for the exit of 
the oculi-motor remains distinct. 

As might be expected from what has already been said about 
the orbit, we find its hinder wall also very broad, and generally 
concave forwards. At its usual site a distinct, irregular foramen 


248 DR R. W. SHUFELDT. 


of some size is found for the exit of the olfactory nerve, and this 
branch passes forwards in the living bird in a shallow channel on 
the inter-orbital septum beneath the frontal, for its entire length, 
where these two elements are united. It leads to the inner 
and larger of the two foramina that were described above, as 
occurring over the pars plana. 

Before leaving this side view of the skull, it will be as well to 
notice the large, luniform sesamoid that occurs in the ligament 
that passes from the quadrato-jugal to the hinder border of the 
articular cup of the mandible. This sesamoid is present on 
both sides, and in all the skulls of Geococcyx that I have ever had 
the opportunity of examining. 

On the superior view of the skull we are to note the form of 
bony lamina that partially close in the external narial openings 
from behind; the position of the two small circular foramina 
beyond the cranio-facial hinge; and this fronto-lacrymal region 
generally. From this aspect, we also see the small foramina that 
pierce on either side the orbital roofs behind. Mesially, and 
between these latter, a shallow, longitudinal groove marks the 
cranial vault. Posterior to this again we find a smooth, globular, 
and ample parietal region. The crotaphyte fossze may likewise be 
discerned from this upper aspect, and a glimpse obtained of the 
supra-occipital prominence. It isalsouseful inshowing the manner 
in which the quadrato-jugals articulate with the quadrates. 

Viewing the skull of Geococcyx from beneath, we find, an- 
teriorly, the broad, flat surface, already spoken of, which forms 
the lower face of the superior mandible ( fig. 4). 

Following this back we come to an elongated median vacuity, 
that separates the anterior terminations of the maxillo-palatines. 
This aperture has irregular, jagged edges, and through it we may 
see some of the open, spongy bone tissue that partially fills the 
hinder portion of the core of the superior mandible. At the 
sides, the posterior processes of the dentary parts of the pre- 
maxillary overlap the maxillaries. They are long and triangular, 
with their apices to the rear. 

Returning to the mawillo-palatines we find them to be, upon 
this aspect of the skull, two very sizeable, elongated, subcylindri- 
cal masses, composed of an internal spongy tissue, but encased 
in an outer covering of an extremely thin layer of compact 


- 
THE SKELETON IN GEOCOCCYX. 249 


tissues. They lie parallel to each other, and to the median plane, 
nearly filling the interpalatine space. Anteriorly, they are 
separated by the vacuity already described, while behind their 
free and rounded extremities slightly diverge from each other, 
they being in contact in the median line for the middle thirds of 
their lengths (fig. 4). From their upper sides is developed a 
mass of open, spongy tissue; this is continuous with a similar 
structure that is found within the superior mandible; it reaches 
out on either side, to abut against the inner surfaces of the 
nasals; it joins the horizontal plates of the maxillaries, and 
finally supports a median vertical plate of bone that stands 
just beyond the rhinal chamber proper, this latter space being 
free from its encroachment, as it is from any development of the 
ethmoid behind, beyond its lateral wings. 

The anterior half of either palatine is quite a broad, flat, 
horizontal plate, the distal end of which indistinguishably fuses, 
and is directly continuous with the horizontal portion of the 
premaxillary. To its inner side alsv, in this locality, it com- 
pletely anchyloses with the corresponding maxillo-palatine (fig. 
4). For the most part, however, its inner and outer edges are 
free, not coming in contact by the inner one with the maxillo- 
palatine, though it is parallel to it, and separated by an ex- 
tremely narrow interval, while its outer one neither touches the 
lacrymal nor the maxillary, but occupies a plane inferior to both. 

The posterior half of a palatine also lies mainly in the 
horizontal plane, but its under surface is a concave one, and its 
upper correspondingly convex. Its outer free edge, directly 
continuous with the outer edge of the anterior half of the bone, 
sweeps by a gentle curve round the “ postero-external angle” 
of the palatine to its head.? 


1 Professor Huxley, in his ‘‘ Classification of Birds”’ (Proc. Zool. Soc., April 11, 1867, 
p- 444) says:—‘‘ In Geococcyx the principle of construction is quite the same [as it is 
in Cuculus canorus], but the postero-external angles of the palatines are distinctly 
indicated.” In the two excellent skulls of adult specimens of this bird before 
me, as well as those I examined in the collection of the Army Medical Museum 
at Washington, the postero-external angles are rounded off in the manner above 
described, and the presence of a process is in no way indicated whatever, and I 
must believe Professor Huxley had in hand, the time he penned the words I 
have quoted, an imperfect and perhaps broken skull of Geococcyx. (See also my 
remarks upon this point in my ‘‘ Osteology of Ceryle aleyon,” Jowr. of Anat. and 
Phys., April 1884, p. 289). 


250 DR R. W. SHUFELDT. 


The inner free edge of the bone extends from the head to the 
apex of a small pointed process in front. For nearly its entire 
length it is parallel to the corresponding edge of the palatine of 
the opposite side, from which it is separated by an interval of 
something like a millimetre or rather more. From this edge 


the surface curves outwards and back, forming the “ascending 


process” of the palatine. This terminates in another longitudinal 
straight margin, which is applied to the corresponding one of 
the opposite palatine, and both unite to form the usual groove 
at their upper aspects for the rostrum of the sphenoid. These 
latter opposed edges also extend from the palatine heads, likewise 
in contact mesially, to a common anterior process. This latter 
is nearly opposite the anterior end of the rostrum, and from its 
extremity in front projects a free, needle-like, and rudimentary 
vomer, of some four millimetres in length. It does not come in 
contact with the maxillo-palatines, but lies above the interval 
formed by their slightiy diverging posterior extremities, and is 
freely articulated with the palatines at the point from which it 
springs, and in the manner described. This diminutive vomer is 
equally well developed in both my specimens of Geococcyz. 

A pterygoid is a nearly straight and slender bone, and shows 
not the slightest evidence of the development on its shaft of an 
apopbysis, and indeed there is no necessity for such, as the basi- 
pterygoidal processes are entirely absent in this bird; and the 
pterygoids when im sitw oceupy a lower plane than the basi- 
temporal region, as well as being at some distance in front of it. 

These bones articulate with each other anteriorly and with 
the opposed palatines ; from this point they diverge at an angle 
of about 85°, each to meet the usual facet upon the correspond- 
ing quadrate at the base of the inner and smaller condyle ou 
that bone. 

The basi-temporal region is elevated above the prominent and 
raised boundaries of the auricular apertures; it is narrow and 
smooth and lies for the most part in the horizontal plane. In 
front it presents for our examination a thin tip of bone, arching 
over the common aperture of the Eustachian tubes. 

Beyond this it contracts to form the sphenoidal rostrum, a 
considerable portion of which is unoccupied before we reach 
the pterygoidal heads. This allows these bones not a little 


e 


THE SKELETON IN GEOCOCCYX. 251 


backward play in the recent specimen, an action which is quite 
possible from the more than ordinary mobility enjoyed on the 
part of the cranio-facial hinge. 

Either external auricular conch is a capacious fossa, well- 
defined by a raised and bounding thin wall of bone, with its free 
edge curled inall around. At the base of either of these fosse, we 
see strong osseous trabeculz converging to a point near the 
centre, to support the double concave facet for the mastoidal 
head of the quadrate. These stand between the Eustachian 
entrance and the passage to the middle ear. 

If the plane of the basis cranii be produced posteriorly, and 
the plane of the occiput and foramen magnum extended to meet 
it, we find the latter makes an angle with the first mentioned 
plane of about 48°, while the long axis of the fairly well- 
developed supraoccipital prominence would be perpendicular to 
it. Inform the foramen magnum is broadly cordate with its 
apex above; the occipital condyle at its lower margin is small, 
sessile, and hemispherical in outline, being so placed as to 
encroach upon the foraminal periphery for about one-third of 
the condylar arc. 

Points of interest within the brain-case are seen in the 
presence of a strongly marked longitudinal sinus, and the 
unusual thickness of the walls of the sella turcica; its fossa, 
though deep, being quite small, while at its base we find a 
double entrance for the carotids. 

As a whole, the skull of Geococeyx is a delicate and a very 
light structure for its size, air gaining thorough access to most 
of its parts. 

The mandible (figs. 1 and 2), seen from superior aspect, has 
the typical V-shaped form, with an extensive symphysis, which 
is scooped out longitudinally above. Either ramus is not deep 
in the vertical direction, while its upper and lower margins are 
prominent and rounded, the former, however, becoming sharp as 
it approaches the symphysis, which condition is sustained to the 
mandibular apex. 

The ramal vacuity is large and occupies its most usual site ; 
in outline it is an elongated ellipse, but its anterior third is 
encroached upon by a thin plate developed on the part of thie 
dentary element. An articular end is considerably concave 


252 DR R. W. SHUFELDT. 


above, and presents two facets for the condyles of the quadrate ; 
its inturned process is much tipped up, while the usual pneu- 
matie foramen is seen near its apex. Below, its convexity con- 
forms with the concavity of the articular excavation at its upper 
side, and its angle behind is obliquely truncate from above 
downwards in the forward direction. 

Beyond an articular end, on the superior ramal border, we 
find, on either side, the coronoid process but feebly developed, 
and single. 

When the osseous mandible is articulated im sitw with the 
remainder of the skull, its tip does not extend quite so far for- 
wards as does the apex of the superior osseous beak, a condition 
present in the skulls of most Coracomorphe and other groups. 

In the hyoidean apparatus (fig. 8) we find fully the anterior 
two-thirds of the glosso-hyal represented by a thin stripe of 
cartilage; while behind, where it ossifies in front, the usual 
median foramen is seen, having an elliptical outline. Posterior 
to this, on either side, the strongly marked cerato-hyals project 
outwards and backwards. 

First and second basi-branchials do not anchylose with each 
other, the former being short and thick, the latter about half 
as long again, and tipped off behind with cartilage. 

The elements of the thyro-hyals are long and slender, they 
likewise terminate in cartilaginous tips, and curve up Lehind the 
skull in the manner most usual among birds. 

Of the remainder of the Axial Skeleton-—The Vertebral Column. 
—This column presents us with eighteen movable vertebree before 
we arrive at the consolidated pelvic sacrum. This latter contains 
eleven more segments, thoroughly united together and firmly 
Joined to the iliac bones. Finally, we find five vertebrae and 
a large pygostyle in the skeleton of the tail of Geococcyz. 

In the cervical region we pass twelve vertebree before we come 
to the first one of the series that bears a pair of free ribs; the 
thirteenth and fourteenth both possessing these appendages, and 
in both they are well developed, though not reaching the sternum, 
through the intervention of costal ribs. The pair on the four- 
teenth vertebre has the epipleural processes fully as large as 
they are in the dorsal series; they are absent entirely, however, 
on the first pair of free ribs. 


THE SKELETON IN GEOCOCCYX. 253 


Returning to the atlas we find this segment rather delicately 
constructed, though assuming the form the bone usually offers 
among Passerine birds. Its neural arch is narrow antero-pos- 
teriorly, though the canal is capacious. A perforation is seen at 
the base of the articular cup for the occipital condyle, which 
cuts through the superior margin of this little concavity. The 
centrum is small and does not develop anything that might be 
called an hypapophysis. On the ais vertebra we note the 
presence of a low, tuberous, neural spine, occupying the entire 
central portion of the arch, while posteriorly on the under side 
of the centrum a feebly pronounced hypapophysis is seen. The 
odontoid apophysis is small and short as compared with other 
features of this vertebra, a fact no doubt due to the lack of 
depth in the atlas. At either side of the centrum we observe a 
delicate and vertical spicula of bone, which completely arches 
over the vertebral vessels, constituting the last remnants of the 
lateral canal at this extremity of the column. This condition is 
often met with among the Anatide in the axis vertebra of those 
birds. 

The postzygapophyses are directed backwards and outwards, 
and are very powerfully developed, more so than in any of the 
first nine or ten vertebre of this portion of the column. The 
facets they bear for articulation with the extremities of the 
prezygapophyses of the third segment are at their under side 
about the middle. 

On the third and fourth vertebre we also find a low neural 
spine placed at the centre of either bone, while the hypapophysis 
is becoming reduced in these segments, to disappear entirely in 
the fifth vertebra. These vertebre, as in so many of the class, 
have their zygapophysial processes joined by a spanning lamina 
of bone, which in either case, and on either side, is pierced near 
its middle by a small elliptical foramen, of the greater size in 
the fourth vertebra. 

The lateral canals occupy rather more than the anterior halves 
of the sides of the centra, and the processes that project from 
the under aspects of their free margins behind are short, and 
each is separated by a considerable interval from its fellow of 
the opposite side. This great inferior width of the cervical 
vertebra is a characteristic feature of these segments in Geococcyz, 


254 DR R. W. SHUFELDT. 


and is well-sustained throughout the series, until we come to 
the free rib-bearing ones, when a gradual contraction takes 
place as we pass into the dorsal region. But even here the 
segments are comparatively broader in their transverse diameters 
than we often find them. 

In the fifth vertebra the neural spine is placed further forwards 
on the bone, but is very small; it is absent in the sixth, or only 
faintly indicated, and it does not appear in the series again until 
we find it as a pronounced crest on the fifteenth segment. Some- 
times, however, a low, tuberous elevation marks its site in the 
few ultimate cervicals. 

Prezygapopbyses in the fifth vertebra stand almost directly 
outwards, while the postzygapophyses very prominently point to 
the rear. Little modification takes place in the former of these 
processes as we examine the succeeding vertebre, their general 
direction remaining about the same, but the articular facets they 
bear face more and more towards the median plane as we 
proceed backwards. With the postzygapophyses, however, the 
case is otherwise, for as we descend the cervical series we find 
these become gradually shorter and stouter, with a wider 
divergence, while their facets, from facing downwards and 
outwards, come to look almost directly downwards. 

We find strongly marked metapophyses surmounting the 
bases of the postzygapophyses in the sixth to the ninth cervical 
vertebree inclusive; after that they disappear, and are but feebly 
reproduced in the dorsals, where they occur on the superior 
aspects of the ends of the transverse processes. 

On the fifth cervical vertebra the lateral canal is at its 
forward part, appropriating about the anterior moiety of the 
entire centrum. Its outer wall may show a slight perforation, 
while the parapophyses which project from it behind are on 
either side a short and needle-like spine. As we pass down the 
series this perforation becomes larger and larger, until in the 
tenth vertebra it has broken through the hinder free margin of 
the lateral canal and disappeared, leaving in the segment only a 
shorter passage, and a deep concave notch indicating its site. 
Pari passu with this change, the parapophyses and pleura- 
pophyses pass through the usual evolution in that direction, to 
result in the perfect and free pair of ribs found in the thirteenth 


. 


THE SKELETON IN GEOCOCCYX, 255 


vertebra. Faint beginnings of a carotid canal are also seen in 
the fifth vertebra, in the presence of a shallow excavation at the 
anterior end of the under side of segment. This becomes better 
and better marked to include the teuth vertebra, where this 
canal is moderately well protected by lateral walls, but in none 
of the series does it become a closed passage as in some other 
birds, In the eleventh vertebra its place is taken by a strong, 
single, and median hypapophysis. 

This latter feature becomes faintly tricornuate in the twelfth 
vertebra ; markedly so in the next segment; the three prongs 
springing from a common pedicle in the fourteenth ; which pedicle 
is lengthened in the fifteenth ; still longer but without terminal 
prongs in the sixteenth vertebra; to be entirely absent in the 
succeeding segment, and the rest of the column. 

In the atlas the neural canal is capacious and transversely 
elliptical. From this vertebra it gradually changes its form and 
contracts in calibre, until in the fifth vertebra we tind it nearly 
cylindrical in shape, and much reduced in capacity. 

Passing down the series it gradually changes for a second 
time, so that in the eleventh vertebra it is again found 
to be large and transversely elliptical. This form it retains 
through the dorsal series, though once more reduced in calibre, 
In the tail vertebre it is at first triangular with apex above, to 
become a vertical slit as it enters the pygostyle. 

The fifteenth, sixteenth, seventeenth, and eighteenth vertebrae 
of the column in Geococcyx support ribs that meet to articulate 
with costal ribs below (Plate VIII. fig. 7). These ribs are broad 
above, but become more and more rod-like as they near their 
hemapophysial articulations. The first three pair of the series 
bear large epipleural processes, which are always anchylosed to 
the rib upon which they appear. These three also have costal 
ribs connecting them with the sternum ; this I believe to be the 
smallest number of the latter present in any living bird, «e., only 
three heemapophyses articulating with either costal border of the 
sternum. The last pair of ribs, or those coming from the 
eighteenth vertebra, never have epipleural processes, and their 
costal ribs do not reach the sternum (fig. 7). 

With respect to the four vertebrz that own these ribs, we find 
that they present all the characters of the dorsals as found 


256 DR R. W. SHUFELDT. 


among Aves generally. The neural spines are lofty and quadri- 
lateral in outline, each having its superior rim capped off 
with a vertically flattened tablet of bone. The diapophyses are 
rather broad, and project directly outwards from the sides of the 
vertebrae, having the ribs articulating with them and the centra 
in the usual way. Very close interlocking is evidenced among 
these four dorsal segments, and the post- and prezygapophyses 
are no longer than is necessary to afford the proper amount of 
surface for their respective articular facets. Anteriorly, these 
face upwards and inwards, precisely the reverse being the case 
with those found on the postzygapophyses. 

So far as we have examined the vertebral column, the articula- 
tion which obtains among the centra is upon the heterocelous 
plan, i.c., the anterior facet is concave from side to side, convex 
from above, downward, precisely the reverse condition being 
present in the posterior facet. All these vertebree, as well as 
both kinds of ribs, are eminently pneumatic, groups of foramina 
occurring at the usual sites in these bones. 

The Pelvis (Plate VIII. figs. 9, 10 and 11).—From its sagan 
unique form the pelvis of Geococcyxz has attracted the attention 
of a number of anatomists. Owen speaks of the ilium as 
forming behind “a prominent ridge in most birds, which 
generally overhangs the outer surface; in Geococcyx, to a remark- 
able extent, like a wide pent-house, producing a deep concavity 
in the outer and back part of the ilium, where it coalesces with 
the ischium.”! 

Marsh, in his classical work upon the Odontornithes, again 
calls attention to the same thing, and points out other particulars 
in connection with it, making admirable comparisons with the 
pelves of Reptilia, Tinamus, and other forms.” 

Strange to relate, the only other living American bird, so far 
as I have examined, that possesses a pelvis anything like the one 
we find in Geococcyx, is the common sorn rail (Porzana carolina). 


1 Anat. of Verts., vol. ii. p. 84, Lond. 1866. 

2 Marsh, O. C., Odontornithes, pp. 70-73, figs. 16-20, Washington Government 
Printing Office, 1880. There certainly can be nothing that advances our know- 
ledge of the exact origin of birds more certainly than the constant comparison of 
recent forms with the material paleontology has thus far been enabled to supply 
us,—not a great deal as yet. Professor Marsh never seems to allow such an 
opportunity to escape him. 


o 


THE SKELETON IN GEOCOCCYX. 257 


This bird not only has the ilia forming the peculiar outward- 
curling crests behind, but has also the propubis well marked, 


_ and identically the same style assumed by the anterior portions 


of the ilia, z.2.,a deeply concave inner margin, with the sacral 
crista mounting above it, and not coming in contact with the 
same. 

Viewing the pelvis of Geococcyx from above, we are to notice 
the condition just alluded to (fig. 10), as well as the raised, 
anterior emarginations of these ilia, with the processes that 
project from their middle points. As already hinted, the ilio- 
neural canals are here open grooves, and the neural crest of the 
sacrum stands between them as a lofty dividing wall, with 
much thickened superior border. This latter is distinctly 
marked for the entire length of the sacrum, otherwise the 
individualisation of the vertebre composing this part of tbe 
bone is not very distinct, as few foramina are to be found 
between their diapophyses, until we reach the last one, where 
regularly occurs a large pair, throwing the ultimate uro-sacral 
into bold relief (fig. 10). 

Owing to the spreading of the ilia, both laterally and behind, the 
superficies of the postacetabular, far exceeds the preacetabular, 
and these iliac bones are much tilted up posteriorly. 

Upon the lateral aspect of this pelvis (fig. 9) we not only gain 
a better view of the largely developed propubis, and the strangely 
formed hinder portion of the ilium, but we are also enabled to 
get a glimpse of the rather small subcircular ischiac foramen, 
with the reniform antitrochanter in front of it. This latter faces 
almost directly forwards, and only slightly downwards, and less 
so outwards. Beyond this again is the acetabulum, with the 
circular perforation at its base, the postero-superior are of which 

"merges with the periphery of the outer cotyloid ring at the base 
of the antitrochanter, while directly opposite this pvint the arcs 
of these two circles are far apart, and an excavation occupies the 
intervening space. This grows less, of course, as we proceed 
either way towards the base of the antitrochanter, where, as I 
have said, the inner and outer rings are tangent to each other. 

The elliptical obtwrator foramen occupies its usual position, 
and so close together are the post pubis and ischium that an 
exceedingly narrow strait leads from this vacuity into the 

VOL. XX. R 


258 DR R. W. SHUFELDT. 


obturator space, a long narrow interval between the last two 
mentioned bones. At the centre of the triangular area among 
these three apertures at the side of this pelvis, is found a group 
of small pneumatic foramina which assist in admitting the air 
into the substance of this light and thoroughly aerated bone. 

The Caudal Vertebre and Pygostyle-—As already stated above, 
the caudal vertebree are five in number (Plate IX. fig. 17). They 
are chiefly noted for their high and prominent neural spines ; 
the two loftiest being seen in the third and fourth vertebre. 
The diapophyses grow longer and more spreading as we proceed 
in the direction of the pygostyle, the last segment possessing them 
longer than any of the others. We find in the third caudal 
vertebra a small anchylosed chevron bone, which slightly over- 
laps the bone in front of it. This apophysis is very strongly 
developed in the last two vertebre, where it is also anchylosed 
to the centra, is bifid, and hooks well forward to overlap the 
preceding centrum in either case. Each one of these bones is 
pierced by pneumatic foramina in a number of places, as is also 
the terminal coccygeal vomer. 

This latter bone has an irregular oblong figure, with its 
posterior margin considerably thickened, the others being cultrate. 
The neural canal is continued into it for some little distance, 
its passage being denoted on the sides of the bone by a longi- 
tudinal smooth elevation, which gradually tapers away to the 
postero-superior angle. 

Of the Sternum and Pectoral Arch.—The sternum of Geococcya 
is a thoroughly pneumatic bone, but air does not gain access to 
any of the shoulder-girdle elements. 

In the case of the former, foramina are chiefly found in the 
concavities among the heemapophysial facets on the costal borders. 
A few scattered ones may be seen in the median line upon the 
dorsal surface. The number of these latter vary in different 
specimens. 

The “Road Runner” has a two-notched sternum, which gives 
rise to a pair of flaring xiphoidal processes on either side (figs. 
7, 16), Its carina is fairly well developed and moderately deep 
only. It extends the entire length of the bone, and is marked 
upon the upper side of its projecting carinal angle by a roughened 
facet for articulation with the hypocleidium of the furculum. - 


THE SKELETON IN GEOCOCCYX. 259 


Osseous welts are raised upon its sides to facilitate muscular 
attachment, and these in some specimens, extend on to the 
ventral aspect of the body (fig. 16). The inferior border of the 
keel is somewhat thickened. 

In front of the sternum a peg-like manubrium projects out, the 
lower margin of which is longitudinally marked by a sharpened 
crest (fig. 7). Below this, the perpendicular anterior border of 
the keel is vertically concave, and this inferior manubrial crest 
is carried into the excavation as a median raised line. 

Either costal border is very short, having but three facets 
upon it, and these are usually close together. In front of them, 
on either side, a prominent costal process is reared, constituting 
one of the most striking features in this part of the skeleton of 
Geococeys: (fig. 7). 

The thoracic aspect of the sternum is very much concave, the 
ventral side being correspondingly convex. Here on this latter 
we notice well-marked muscular lines, one on either side, com- 
mencing at the outer termination of a coracoidal groove, and run- 
ning backwards to a point about opposite the middle of the keel. 

The coracoidal grooves do not meet at the manubrial base in 
the median line, and each one is characterised as being a deep 
transverse notch, with upper and lower lips of projecting bone 
and extending laterally only so far as the inner or anterior 
limit of the base of the corresponding costal process. 

With respect to the pectoral arch, I find a coracoid to be, 
comparatively speaking, an unusually long bone ; its sternal or 
lower border extends beyond the facet proper, in order to fit into 
the coracoidal groove of the sternum. This end of the coracoid 
is not as much expanded as we find it in some birds, but, on the 
other hand, like many of the class, its outer angle is produced and 
bent upwards as a projecting process. 

The shaft is long and cylindrical, being marked down its 
posterior and lateral aspects by musculatr lines. 

At the superior, or really anterior extremity of this bone, we 
find several noteworthy and interesting characters (fig. 5). 

Its scapular process is very long, and compressed from side to 
side. This apophysis reaches forwards, and by its slightly 
dilated extremity articulates with a vertically concave notch in 
the lower part of the head of the corresponding clavicle. 


260 DR R. W. SHUFELDT. 


Another meeting between these two bones takes place 
above, and this is affected by the summit of the coracoid 
curving inwards towards the median plane, to articulate with a 
considerable facet found at the highest point of the clavicular 
head. 

These two articulations between the furculum and the coracoid 
completely close the tendinal canal, even without the assistance 
of the scapular behind, though this latter bone materially aids in 
increasing the actual length of this tendinal passage, by closing 
up the posterior gap. 

If we turn to the dotted outline of the es furculum, shown in 
figure 6, it will be noticed that it has a form about intermediate 
between the usual U and V shapes of the bone. Regarding it 
from a lateral aspect, as shown in the preceding figure, the actual 
form of one of its transversely compressed heads can be better 
appreciated, as well as its method of articulation with the other 
bones of the girdle (fig. 5). 

Below it is flattened in the antero-posterior dineatiagat and 
terminates in an elongated hypocleidium. This latter articulates 
when the arch is im situ with the carinal angle of the sternum, 
in the manner described in a foregoing paragraph. 

A scapula assists to form the glenoid cavity in the usual way, 
contributing about half the surface to that humeral socket. Its 
clavicular process reaches far forwards, to make an extensive 
articulation with the head of the furculum, when the bones are 
in the position they assume in life. It also rests further forwards 
upon the scapular process of the coracoid than is usually seen 
wmong birds (fig. 5). Sometimes we find the posterior third of 
the long, narrow blade of this bone bent down more abruptly 
than in the specimen I have figured, and its end is always 
rounded off, rather than being truncated, as is commonly the 
condition in Aves, 

At the outer and back part of the shoulder-joint in the adult 
Geococcyx, occurs usually a very minute sesamoid, known as the 
os humero scapulare, and I am led to believe that small sesamoids 
may yet be found in other of the tendons of the pectoral 
extremity in this region. 

Of the epenitiondin Skeleton— The Pectoral Limb (J late VIII. 
figs. 12-15).—Pneumaticity is extended only to the bone of the 


> 


THE SKELETON IN GEOCOCCYX. 261 


brachium in this limb, the hollow shafts of the other long bones 
being charged with medullary substance. 

The humeral shaft is much bowed, and in such a manner as 
to be convex along its radial border and concave upon the 
opposite side, which concavity is more apparent owing to the 
prominence of the ulnar crest, and the peculiar projection of the 
distal extremity in the continuity of this curve (figs. 12, 14). 

In form the shaft is nearly cylindrical, and almost entirely 
devoid of muscular lines. 

At the proximal end, a well-marked valley occurs between 
the ulnar crest and the spindleform humeral head. The former 
has barely any pneumatic fossa at its base, the circular foramen 
there found being nearly flush with the general surface of the 
bone. On the opposite aspect we find a short though prominent 
radial crest, which makes no pretence to extend its lamelliform 
plate down the shaft, as we often find to be the case in birds. 

The distal extremity of this bone presents for examination 
the usual oblique and ulnar tubercle, while, as already alluded 
to, the ulnar condyle of this end is much produced and very 
prominent (figs. 12, 14). 

The ancoueal aspect immediately above the trochlea is flat and 
smooth, the opposite side showing a broad, shallow groove 
for the guidance of tendons to the antibrachium. A fairly 
well-developed “ectocondyloid tubercle” is seen at its usual 
site, on the radial border of the shaft just above the oblique 
trochlea. 

Following the example of the humerus, we find the compara- 
tively short radius and ulna very much bowed along the 
continuity of their shafts. This gives rise to a broad spindle- 
shaped interosseous space, the two bones only coming in contact 
at their distal and proximal extremities, when articulated (fig. 
13). 

The radius is not nearly so much bent as the otber bone of 
the antibrachium, and presents nothing peculiar about it. On 
the other hand, the wlna, with its greatly curved shaft, its 
prominent row of secondary papille, and its well-developed 
olecranon, is quite a striking bone beside it. 

Composing the elements of the carpus, the two usual free 
segments are seen; of these the radiale has pretty much the 


262 DR R. W. SHUFELDT. 


same form as it assumes among birds generally, while the wnare 
takes on an entirely different shape. It does not develop the 
two limbs or processes that straddle the proximal extremity of 
the carpo-metacarpus, when the bones are in situ, as in the 
vast majority of the class, but is simply a bar of bone, with one 
end enlarged, and bearing at its summit an articular facet for 
the ulna. 

The carpo-metacarpus is chiefly interesting for its peculiarly 
formed mid-metacarpal. This is uncommonly broad at its 
proximal end, and curiously twisted, as it descends to anchylose 
with the lower end of the index metacarpal, or main shaft of 
this compound bone. So far as I have been enabled to discover, 
the phalanx of pollex-digit does not bear a terminal claw, and 
the bone has the usual form as seen in most Aves. Nothing of 
note distinguishes the two phalanges of the index digit, while the 
small phalanx of the last finger develops, at the middle point 
of its hinder margin, a curious little upturned spur (Plate VIII. 
fig. 13). 

Of the Pelvic Limb (Plate IX. figs. 18-27).—As in the pectoral 
extremity, the proximal long bone of this limb, the femur, is the 
only one in it that enjoys a pneumatic condition. The site of 
the foramen that admits the air to its hollow shaft is, however, 
quite unique, being upon the posterior aspect of the bone, 
between the trochanter and head, instead of on the anterior side, 
as usual, below the trochanter. 

This latter feature is not elevated above the articular surface 
at the summit, and the semi-globular head is, comparatively 
speakiny, rather small. A shallow excavation upon its upper 
side marks the usual point for the insertion of the round 
ligament. 

The sub-cylindrical shaft faintly showing the muscular lines, 
is considerably bent to the front, as shown in fig. 19. 

At its distal extremity in front, the rotular channel is well 
marked, the condylar ridges bounding it being about parallel to 
each other (fig. 18). 

The outer and larger condyle of the two is at the same time 
the lower, and the fibular cleft that marks its posterior aspect, 


is very wide and deeply sculpt, being rather more to the outer 
side than is usual (fig. 20). 


» 
THE SKELETON IN GEOCOCCYX. 263 


Above these condyles, behind, the popliteal fossa is but 
moderately excavated, and a straight transverse line bounding 
it below divides it from the general trochlear surface. 

We find in the next segment of this limb, the tibio-tarsus 
with a subcylindrical shaft below its fibular ridge, that is slightly 
bent so as to be in the vertical line, somewhat convex anteriorly 
(fig. 22). The bending here though is not nearly so great as we 
found it to be in the humerus and femur, or, to make the com- 
parison more exact, in the ulna. 

The cnemial crest of this leg-bone is but little raised above 
the undulating articular surface of its summit, while the pro- 
and ectocnemial ridges, that develop below it, are not peculiar 
(figs. 21-23). Their planes are at right angles to each other, 
that of the latter having its surface facing directly to the front. 
Neither is produced for any distance down the shaft of the 
bone, but terminates rather abruptly upon it; the pro-cnemial 
ridge at a point about opposite the superior end of the fibular 
ridge on the other side of the shaft (fig. 21). 

At the distal extremity of the tibio-tarsus the planes of the 
condyles are nearly parallel to each other, and these trochlear 
eminences are strikingly close together in Geococcyz. 

The intercondyloid fossa is deeply excavated in front, to 
become suddenly much shallower behind, as well as somewhat 
narrower. Upon lateral view, it will be seen that the general 
outline of either of the condyles is more circular than we 
usually find it in others of the class, where a reniform pattern 
prevails (fig. 22). 

Just above the condyles, on the anterior aspect, the vertical 
tendinal channel is spanned by the usual little, oblique bridge 
of bone, and this is supplemented in life by a longer ligamentous 
one placed in front of it. 

The fibula has a large head, which is produced backwards 
beyond its shaft. This latter makes a close ligamentous articu- 
lation with the fibular ridge of the tibio-tarsus, and at some 
little distance below it merges into its shaft, to become almost 
indistinguishably fused with it (figs. 21-23). 

A well-developed, subcordate patella, with its apex directed 
below, is found in the usual tendon in Geococcyx. 

It will be seen by referring to figures 24 and 26 that the 


264 DR R. W. SHUFELDT. 


tarso-metatarsus of the Road Runner is a longer bone perhaps, 
than we would be led to expect from the other long bones of 
this limb. . 

Its summit presents for examination the two concavities for 
the condyles of the tibio-tarsus, separated by the mid-tubercle. 
Behind this we find a short hypo-tarsus, showing two vertical 
grooves at its back, and two vertical perforations through it, as 
shown in fig. 26. 

The sides and front of this bone are flat, the latter for its 
proximal half being longitudinally grooved, deepest above, 
gradually becoming shallower as it descends. — Posteriorly it is 
likewise grooved in a somewhat similar way; but here the outer 
wall of the groove is raised as a sharp longitudinal crest, best 
marked at the middle third of the shaft, and gradually subsiding 
towards the extremities. 

At the distal end we note the three usual trochlee for the 
basal joints of the toes; figure 27, however, shows how, in 
this zygodactyle bird, the outer one of these is extended to the 
rear, in such a manner as to allow the fourth toe to articulate 
in that direction (figs. 26 and 27). 

Of these trochlez the middle one is much the largest and is 
placed the lowest down; it is the only one of the three that 
shows the distinct median groove. The trochlea for the fourth 
toe is much elevated, while the inner one holds about a mid- 
position in this respect. 

A well-developed accessory metatarsal, slung by ligament in 
the usual way, is found between the shaft and the basal joint of 
the hallux (fig. 26). The perforating foramen for the passage of 
the anterior tibial artery is small and inconspicuous, being at the 
same time quite low down on the shaft. 

The joints of these podal digits are harmoniously proportioned, 
both as regards size and comparative length, and beyond what 
can be easily studied in my drawing of them, and their typical 
zygodactylism, they offer nothing of particular note. 

Before reducing my specimens to skeletons, I failed to make 
any special examinations as to the condition of the ossifications 
of the columella auris in the adult Geococeyx ; I find, however, 
among other normal ossifications in this type, some twelve or 
thirteen sclerotal plates in either eye, overlapping each other in 


cee nda 


nihiar a gmeeblibaalipe aie a 


i 
| 
: 
} 
f 


THE SKELETON IN GEOCOCCYX. 265 


a somewhat irregular manner. The rings of the trachea also 
ossify as in other birds, as well as the plate of the superior 
larynx. Some of the tendons of the pelvic limb in old birds are 
also converted into bone, and small sesamoids may be found 
about the proximal extremities of the basal joints in the soles of 
the feet. 

During the preparation of this memoir several more Pacitic 
Coast specimens of this form have come into my hands; among 
others I am indebted to Mr W. Otto Emerson of Haywards, 
California, for the present of a very fine male bird. I also have 
a young one, just leaving the nest, but my further examination 
of this material reveals to me nothing in the skeleton of the 
adult tou be added to the above detailed description of my type 
specimen from Santa Barbara, California, from which I made all 
the drawings in the accompanying illustrations in the Plates. 


EXPLANATION OF PLATES VIL, VIII, IX. 
[The figures are all life size and from the same specimen. | 


Fig. 1. Left lateral view of skull and mandible of Geococcyx 


_ californianus, ad. ¢. 


Fig. 2. The mandible seen from above. 

Fig. 3. The skull from superior aspect ; mandible rernoved. 

Fig. 4. The skull seen upon under view ; mandible removed. 

Fig. 5. Right half of the pectoral arch seen from the inner side ; 
the coracoid, furculum, and scapula 7 situ. 

Fig. 6. Os furculum in dotted outline ; viewed from behind. 

Fig. 7. Left lateral view of dorsal vertebree, ribs and sternum; the 
bones all in their natural positions. 

Fig. 8. The hyoid arches seen upon superior view ; the thin glosso- 
hyal is dotted to represent cartilage. 

Fig. 9. Left lateral view of pelvis. 

Fig. 10. Superior aspect of pelvis. 

Fig. 11. Pelvis seen upon direct posterior view. 

Fig. 12. Right humerus, palmar aspect. 

Fig. 13. Palmar aspect of radius and ulna, and the bones of the 
carpus and manus; same limb. 

Fig. 14. Right humerus, anconeal aspect. 

Fig. 15. Right humerus, radial aspect. 


266 


Fig. 
Fig. 
Fig. 
Fig. 
Fig. 


Fig. 
Fig. 


Fig. 
Fig. 
Fig. 
Fig. 


THE SKELETON IN GEOQCOCCYX. 


The sternum seen upon its pectoral aspect. 


. Caudal vertebrae and pygostyle, seen from the right side. 
. Anterior view of left femur. - 
. Inner view of left femur. 


Posterior view of left femur. 


. Left tibio-tarsus and fibula, seen from in front. 
. Left tibio-tarsus and fibula, their outer aspects. 
. The summits of the left fibula and tibio-tarsus. 


Anterior view of the left tarso-metatarsus. 


. The summit of the left tarso-metatarsus. 
. Right lateral view of the skeleton of the left foot of 


Geococcyx, including the tarso-metatarsus. 
Fig. 27. The distal extremity of the left tarso-metatarsus, showing 
the trochlez as seen directly from beneath. 


ON THE RELATIONSHIP OF UREA FORMATION 
TO BILE SECRETION: AN EXPERIMENTAL 
RESEARCH. By D. Noé&t-Paton, M.D., BSc., Biol. 
Fellow of the Univ. of Edin. 


(From the Physiological Laboratory of the University of Edinburgh. ) 
(Continued from page 124.) 
Part Il.—EXPERIMENTAL. 


No direct series of experiments upon this subject has been 
made, but many well-known physiological facts strongly indicate 
the existence of such a relationship. In the first place, in the 
starving animal, the excretion of urea and of bile (Bidder and 
Schmidt) falls, and to the last days of life persists in small 
amounts. In the second place, after a meal both urea excre- 
tion and bile secretion rise, and reach their maximum some 
hours after food is taken. Lastly, the influence of different diets 
on the urea excretion on the one hand, and the bile formation 
on the other, is of interest: while proteids cause a marked 
increase in both, fats have no stimulating effect upon either 
process. 

Instead of following the lines suggested by these past 
observations, I have employed, as before mentioned, another, and 
I believe more satisfactory method—that of studying the 
influence on the urea excretion in dogs in a state of nitrogenous 
balance (stickstoffgleichgewicht) of certain drugs, which power- 
fully stimulate bile secretion. 

My earlier experiments were made upon men, but the results 
obtained, owing to the many disturbing causes which could not be 
excluded, were so unsatisfactory, that after two months’ work, 
this mode of experiment was abandoned, and the observations 
were repeated in dogs. 

For all the experiments here recorded, with the exception of 
two, dogs were employed. And I believe that the use of these 


268 DR D. NOEL-PATON. 


animals renders my results of more value, as it was upon 
dogs that Professor Rutherford conducted his series of experi- 
ments upon bile secretion; so that we may in every case safely 
conclude that under the administration of the drug the secre- 
tion of bile really was stimulated. 


Mode of Expervment. 


Large female dogs were selected, weighing about 13 kilos. 
Female dogs were chosen, because from their mode of micturition 
there is less chance of a loss of urine occurring; and, when 
catheterisation is required, it is more easily and safely performed 
than upon the male. 

The dog was kept in a cage of 40 inches square by 38 inches 
in height. - 

The floor was of smooth zinc, and sloped from all sides to a 
hole in the centre, under which was placed the vessel in which 
the urine was collected. 

Across one side of the cage was stretched a narrow board, on 
which the dog slept and fed, but upon which it rarely 
micturated. 

The floor of the cage was kept scrupulously clean, and was 
frequently washed with a solution of permanganate of potash. 
The feeces were always cleared away as soon after they were 
passed as possible. 

I am well aware that my method of collecting the urine allows 
of a certain loss, but that this is a fixed percentage quantity is 
shown by the very uniform daily excretion of urea which my 
analyses indicate. Using this method, I have been able, by 
carefully regulating the diet, to approach almost as near to a 
daily uniform excretion of urea as Salkowski, Virchow, Wolfsohn, 
or other observers who have employed catheterisation, or who 
have educated their dogs to micturate into a vessel held 
beneath them (Wolfsohn). In some of my experiments I have 
observed a tendency to a bi-diurnal type of urinary excretion : 
(see Exp. 1V.), a small amount of urine being passed on one day, 
followed next day by a more copious quantity. But if a mean — 
be taken for these two days, it corresponds very exactly with 
the normal daily excretion. 


_ 
RELATIONSHIP OF UREA FORMATION TO BILE SECRETION. 269 


_ The urine was collected at ten o’clock each morning ; the cage 
__ was then cleaned, and the dog fed. 

‘ The urine was measured, and set to filter. The specific gravity 
_ was taken, and the urea and uric acid at once estimated. 


Diet. 


The diet consisted of oatmeal porridge with milk. On this 
food the dogs remained healthy for long periods, even when 
confined to the circumscribed limits of their cage. This dict 
caused a firm well-formed fzecal dejection daily, a matter of no 
small importance, as the urine was thus kept free for admixture 
with feeces. 

The urine too was large in amount and of low specific gravity, 
so that the dilution usually required in observations on dogs’ 
urine was thus rendered unnecessary. 


Methods of Analysis. 


_ IL. Urea.—For the quantitative estimation of the urea I have 

_ employed the hypobromite method of Hiifner. 

In preferring this process to the more commonly adopted 
method of Liebig, I feel that I have exposed myself to adverse 
criticism. But a careful perusal of the very extensive and 
scattered literature upon the subject, and a series of experiments 
undertaken by myself, has fully convinced me that,’for such a 
research, this method has advantages over all others. It may 
be rendered more accurate than Liebig’s method, and it has this 
great advantage, that the presence in the urine of the drugs 
administered does not interfere with the results. 

The solutions employed were the following :— 

1st, Solution of sodic hydrate prepared by dissolving 230 
germs, of caustic soda in 500 c.cs. of water. 

2nd, Bromine. 

These were mixed in the propcrtion of 1 in 10, and the 
hypobromite solution was freshly prepared each morning. 

The apparatus of Dupré was adopted, and was first carefully 
tested with standard solution of urea, in order to determine the 
percentage loss of nitrogen. This was found to correspond 


Corrections were made for temperature and pressure accord 


270 DR D. NOEL-PATON. 
very closely to that observed by Leconte, Mehu, Foster, and 
Russel and West. 


ing to the formula given by Hiifner (Jowrnal of Prakt. Chemie, 
N. F., Bd. 1IL p. 11, 1871). 


V = volume of gas at 0° C. and 760 mm. 
V’ = the volume read off. 


B = barometric pressure in mms. 
W = tension of aqueous vapour at temp. at which 
decomposition occurs. 
T = temp. in °C, 
V'(B-W) 


The formula is V = 


760 (1 x 0-00366 7). 


Great care was taken to carry out the decomposition at 
a low temperature, and time was always allowed for the gas 
throughout the apparatus to become of a uniform temperature 
before and after the decomposition occurred. 


Il. Uric Acid.—For the estimation of the uric acid I have 
selected the method recently devised by Professor Haycraft, 
because it appears to me to combine, more than any other, the 
two qualities of accuracy and rapidity. I have tried Heintz’s, 
Cook’s, and Pavy’s, and have failed to get results so nearly 
accurate ; while Salkowski’s very admirable method is much too 
tedious for every day analysis. 

Through the kindness of Prof. Haycraft I have been enabled 
to use this process throughout all my experiments, although his 
paper on the subject has only recently appeared (Brit. Med. 
Journ., vol. ii. 1885, p. 1100). 


EXPERIMENTS. 


I. SALICYLATE OF SODA. 


According to Professor Rutherford’s experiments 62, 66, 
67, 69, 70a and 73a and 74a, salicylate of soda occupies 
perhaps the first position among the cholagogues in its influence 
on the bile secretion. Professor Rutherford says of it—“ Indeed 
this substance is a certain hepatic stimulant, never failing when 


b 


es 


RELATIONSHIP OF UREA FORMATION TO BILE SECRETION. 271 


placed in the duodenum to excite the liver within half an 
hour.” 

Only a few observations on the influence of salicylate of soda 
on the urinary constituents have as yet been recorded. 

In 1876 two papers on the subject appeared, one by Chr. 
Bohr bei Panum in Hospitals Tidende, 2, iii. p. 129, and the 
other by Solomon Wolfsohn (Ueber die Wirkuwng der Salicyl- 
stiure and des Salicylsawren Natron auf den Stoffwechsel). 

Bohr’s paper I have been unable to procure. But, from the 
account of his results given at p. 173 of vol. vi. of Hermann’s 
Handbuch der Physiologie, it appears that the dogs upon which 
his observations were made were allowed to drink what water 
they pleased, and he connects the rise in the urea with the 
increased imbibition of water, which is supposed to have increased 
the tissue metabolism. His results cannot therefore be considered 
of any value. 

Wolfsohn gives six experiments, two upon salicylic acid and 
four upon salicylate of soda. In five of his experiments, the 
drug was given by the mouth, and in one the salicylate of soda 
_ was hypodermically administered. In all his experiments he 
was able to demonstrate a well-marked increase in the total 
nitrogen and in the urea excreted, and this increase in his 
experiments was best marked on the days following the adminis- 
tration of the drug. 

Since he does not state the weight of the dog employed, 
and since the diets used by him differ from that employed 
by me, itis impossible to compare our results in detail. 

Carl Virchow (Ztsch. f. Physiol. Chem., vol. vi. p. 78) gives an 
experiment on the influence of salicylate of soda on the total 
nitrogen of the urine. This was performed in Professor 
Salkowski’s laboratory on a female dog, weighing between 22 
and 24 kilos., and fed upon 500 grms. of flesh, 75 germs. of 
bacon, and 200 grms. of water daily. The urine was collected by 
catheterisation. The nitrogenous excretion before the experi- 
ment was anything but constant; nevertheless his general 
results correspond closely with my own. 

Although the action of the benzoates upon the excretion of 
uric acid has, on account of their connection with the production 
of hippuric acid, attracted considerable attention, I am not aware 


Pi? DR D. NOEL-PATON, 


of any observations upon the influence of their allies, the 
salicylates, upon the urinary constituents. Sée, indeed, in a paper 
published in the Bul. de Acad. Med., 1877, p. 704, states that 
in gravel and gout the uric acid excreted is increased under the 
administration of salicylate of soda, while in health these drugs 
have no influence whatever either upon the urea or the uric acid. 
He states that in acute gout he has seen the uric acid rise 
from 0°30 grms. per 1000 c.es. to 3°10 grms. per 1000 ces. But, 
as Garrod has shown, a similar rise occurs in all cases of acute 
gout towards the termination of the attack ; so little value can 
be placed on Sée’s observations. All the more are they unworthy 
of attention, as he does not record the experiments upon which 
they are founded. 

The action of this drug, which is now so largely used in the 
treatment of rheumatism and gout, and which promises to 


become one of our most valuable remedies in diabetes, appeared 


to me to deserve special attention. I have accordingly devoted 
six experiments to its investigation, four of which I now 
record. 


Exp. 1. and II. 


For these two experiments a man, aged 56, was employed; his 
daily exercise was fixed, and his diet was daily, without any variation, 
the following :— 


Breakfast.—Bread, 4 oz; tea, 1 pint. 
Dinner.—Broth, 14 pints; beef, 4 oz. 
Supper.—Porridge, 14 pints; milk, 4 pint. 


He took little or no water—never more than a tea-cupful in the 
twenty-four hours. 

The daily excretion of urea never became very constant, but the 
influence of salicylate of soda is seen in the accompanying table and 
abstract, and in fig. 1 (p. 274.) 


(Exe, I. and IL. 


© 


RELATIONSHIP OF UREA FORMATION TO BILE SECRETION, 273 


. I. and II! 


Remarks. 


0°968| Weight of man=62°1 kilos. 

1°037 Diet — (as above). 

0-811 

| 0°837 

6 grms. sod salicylatis in 24 

1012 | 32°205 | 0°823 gh eat ‘106 grm. per kilo. of 
body weight. 

1018 | 34°901 | 0°874 Reaction with FeCl, 

1016 | 34°572 | 0°663 

1018 | 41°521 | 0°567 


1019 | 34°335 ” » 
1016 | 35°096 


1017 | 34°156 |0°720| Faint rr ” 
1013 | 33°398 | 0°701 
1013 |} 33°171 |0 800 | 


1014 vie 0-350 | ( 66 grms. sod salicyl. in 24 


2? 9) ”? 


1016 | 34-489 | |. iy | 


hours=0°106 grm. per kilo. 


; 921) of body weight. 
MOIS YN SA/982: | 0281 | | Reaction with FeCl, 
1016 | § 32°349 | 0-321 | es a 
1014 | ( 32°349 | 0° 441 | on fe | 
1013 | §30°590 | 0-257 | i ‘3 ) 
1015 | ( 30°590 | 0-206 ~~ = 


1013 | ( 27°860 | 17119 | 
1015 | 28°351 | 0°679 | 
1013 | 28922 


1014 | | 28-870 | 0-984. 


1016 | | 27-860 | 1-001 | | 


1013 | { 28°870 | 0°894 
1012 | 24°742 re | 


Average daily Excretion of Water and Urea under Salicylate of Soda:— 


| 
Before Exp. I 30°62 
During administration of drug, 2012 34°72 
Between Exp. I. and II., 2108 33°77 
During second administration of drug ug 2016 31°30 
After Exp. II., ‘ é : 2027 28°40 
— and after Exp. I. and II., i i 2003 29°51 
uring Saunas of drug in Exp. I. an 
Il. eee ee . | 2014 33-01 


Percentage ae im Water, .. -. : . practically unaltered. 
ay Bp Urea, . : : . +11°89e. 

1Jn this and all subsequent tables, the date refers to the day upon which the 
greater part of the urine was passed—not to the day on which it was collected. 
Thus the urine of 3°7°84, is the urine collected at 10 A.M. on the 4th July. 

VOL. XX. 8 


A “A119 "102d UIA UOTJOVAA BAVS OULM ‘a 4v WOATS “SULIS : 
1090 UIA worjover saves our ‘ . : Y 1 yo poe ue 
if 4 TM ‘p 4v WAAIS “SUIS 9.9 “Bpog Jo a9e[AOITVg Jopun ploy 94 Jo pue Baty JO WOTJOIOXY— ‘TT pue “I ‘dxq 


OT Ane 


OF 
suis 


RELATIONSHIP OF UREA FURMATION TO BILE SECRETION 275 . 


Average daily Excretion of Uric Acid in grms. under Salicylate 


0°868 0°732 


of Soda :-— 

Exp. I. 
Before. | With. | After. | Before and After. | With. 
0895 0-732 | 0-842 | 


Percentage change= —15%. 


Exp. II. 
Before. | With. | After. Before and After. | With. 
| = Aer a el i rend 
0-767 | | | 0-303 


| 0-303 | 0-981 | 0-849 
. 


Percentage change = — 64°3%. 


Haycraft’s method was used in these, as in all future experi- 
ments, for the determination of the uric acid; and, as the 
presence of this drug in the urine might have interfered with 
the accuracy of the process, this possible source of error was 
excluded by the following experiment :— 


Exp. A. 


The uric acid in a specimen of urine was by this method found to 
be 0:0221%. A large quantity of salicylate of soda was then added 
to another portion of the same urine, and upon testing the percentage 
of uric acid was found to be 0:°02287. 

Therefore, salicylate of soda has no effect in the accuracy of this 
process. 


Exp. IIT. 


On the 27th January, salicylate of soda was given in 2 grm. doses, 
made into pills with alittle mucilage, at 1 p.m, at 4 p.M., and at 6 P.M. 
There was no sickness, but next morning the dog appeared dull, though 
it took food greedily. 2 grms. were administered at 11 a.m. on the 
28th, but the dog appeared so ill that the drug was discontinue l. 
The effect of such large doses of salicylate of soda is shown in the 
table, abstract, and in fig. 2. The effect on the kidneys was marked. 
On the 29th the urine passed was of very high sp. gr., and contained 
a large amount of albumin, but no blood. ‘he albuminuria persisted 
for three days. The drug had absolutely no influence upon the 
bowels. 


276 DR D. NOKL-PATON. 


Ezp. Il. 
Urine | Uric 
matecnt hin sp. G.| Ure@in | acidin| Bowels. Remarks. 
grms. | 
c.cs. grms. 


20°1°85 |} 530 | 1013 7°260 0143 moved Weight of dog=13°6 kilos. 


21 550 | 1012 7°370 0°126 a8 Diet. : 

22 600 | 1013 7°380 0°132 ey Oatmeal as porridge 113 grms. 

23 610 | 1013 7°320 0°140 - Milk 320 c.cs. 

24 470 | 1011 7°442 0°126 | not moved 

26 | 600 | 1014 7°442 Rei moved 

26 465 | 1014 7°394 0125 | not moved ; ay 

27 240 | 1017 4°100 0°064 6 6 grms.) 8 een. sodee salicyl. in 22 

ours. 
28 OU | Seens ae na moved 2 grms.( =0°6 Fe. per kilo. weight 
of dog. 

29 1130 | 1026} 382°192 0:265 | not moved | Reaction of salicyl. with FeCls, 
marked albuminuria. 

30 340 | 1019 9°860 07102 A Reaction of salicyl. with FeCl 
trace of albuminuria. | 

31 800 | 1010 9°168 0°203 moved Reaction of salicyl. with FeCl; 
trace of albuminuria. 

1'2°85| 745 | 1009 8-418 0°223 | not moved 

2 urine lost or aa “4 

3 570 | 1010 7°353 aun moved 

4 550 | 1010 Tee 0°154 - | 

5 750 | 1013 8°222 0'192 an | 


Average Daily Exeretion of various Constituents under Salicylate 


of Soda :— 
| | Before | With After pipe | With Percentage 
Drug. | Drug. | Drug. After * || Change in. 
| 
Water, in c.cs. |) 546 502 653 599 502 - 161% 
Urea, in grms. |} 7°372 | 11°012 | 8-053 7°710 | 11:012 +42°77 
Uric Acid, ,, 0°132 0°126 0°189 0°160 0°126 — 21°37 


Exp. IV. 


This is in every way a satisfactory experiment, and fully confirms 
all the results obtained in Experiment III. The dose was more 
moderate, and the kidneys were not so markedly affected. The 
diminution in the amount of uric acid is very well shown. 

During this experiment the amount of urine daily passed varied 
considerably ; but on taking two days together and striking an average 
an almost constant daily excretion of urea was obtained. This bi- 
diurnal habit in regard to micturition was frequently well marked 
in this dog, but in no way interfered with the results of the experi- 
ments. 


RELATIONSHIP OF UREA FORMATION TO BILE SECRETION. 


Fic. 2 


grms. 
16 


=-j-- ® 


| 
Jan. 20 28 3U 
Exp. III,—Excretion of Urea and Uric Acid under Salicylate of Soda. 
ee given at a, and 2 grms. at e. 


bo 
“I 
io) 


DR D. NOEL-PATON. 


Exp. IV. 


Dat | Urine |. G Urea 
"eh ‘Sp. G.} 

2 yn e.cs. [>P | orms. 
14°3°85 | (563 | | 8150 
15 -, 563 8-150 
16 | (563 | | 87150 
17 (735 8°540 
18 1735 | 8°540 
19 $30 8-200 
20 (595 | 87538 
21 1 595 8°538 

| 

2 440 | | 12-320 
23 500 | 11°400 
24 (725 10-927 
25 Y725— 10°927 
26 830 8°300 
27 

28 (570 67945 
29 570 6°945 
30 645 9-550 
31 645 97550 


Uric Acid 
in grms. 


07136 
07136 
07136 
0°196 
0°196 
07166 
0°129 
07129 


0°075 


0-085 
07145 
07145 
0°107 


0°153 


Bowels. 


moved 
” 
not moved 


moved 
not moved 


” 
> 
moved 


not moved 


9) 
moved 


” 
not moved 


2 grms. » 0°16 grm. | 
per kilo. 
3 gTms. = > . O' 232m. 
Reaction with FeCl,. 
Slight “4 
) 4 - 
{ Average daily excretion of urea 
{ 


J 


Remarks. 


Weight of dog=12°7 kilos. 
Diet—Porridge of 113 grms. oatmeal 
Milk, 320 c.cms. 


2 grms. salicylate of soda=0°16 grm. | 
per kilo. 


=8°247 germs. 


The brackets indicate that the figures given represent the average for period 


grms. 
12 


11 


March 15 


Exp IV,—Excretion of Urea and Uric Acid under Salicylate of Soda. 2 er 
given at a, 2 grms. at e, and 3 grms. at c. 


20 


indicated, 


Fig. 3. 


Uric 
Acid 

germs. 
0°15 


0°05 


25 30 


RELATIONSHIP OF UREA FORMATION TO BILE SECRETION. 279 


Average Daily Excretion of various Constituents 
under Salicylate of Soda. 


| 
: Before 
Before | With | After , | eames 
| Drug. | Drug. | Drug. | jet With. | Change in. 
oe | 
Water, in c.cs. |} 691 482 | 695 693 482 — 80°47 
| Urea, in grms, 8353 | 11393 | 8-258 | 8305 | 11398 | +3717 | 
} 97 


| Uric Acid, 4 0161 | 0082 | 0137 | 0149 | 0-082 | -44: 


Results— With moderate doses of the drug, such as were given 
in Exp. I. and IL, no influence on the amount of water excreted 
could be determined ; with larger doses a well marked dimina- 
tion in the amount of water occurred, and in Exp. III. distinct 
albuminuria was induced, showing that this drug has really an 
irritating action upon the kidneys. This observation fully bears 
out the results of clinical experience of its use, especially in scarlet 
fever, where it is undoubtedly an extremely dangerous remedy. 

The urea was very markedly increased in spite of the diminu- 
tion in the water. 

The uric acid was very much diminished. This last is a point 
of great practical interest and importance, as it throws very 
direct light on the benefit derived from the use of the drug in 
goat. In the practice of M. Germain Sée it has entirely 
replaced the use of colchicum (Year Book of Treatment for 
1884, p. 81). 

How this diminution in the uric acid excretion is brought 
about it is difficult to understand. Obviously it is a diminished 
production and not merely a diminished excretion; for we 
have no great rise in the excretion upon discontinuing the 
drug. 

Either salicylate of soda must cause a more complete meta- 
bolism of those bodies which yield both uric acid and urea, 
whereby they are entirely converted into the latter substance, or 
what is more probable, the production of salicyluric acid must 
interfere with the formation of uric acid. 

To decide this point a quantitative examination of the uric 
acid and salicyluric acid under the influence of the drug would 
be necessary. Unfortunately, there is at present no method for 


280 DR D. NOEL-PATON. 


the accurate quantitative determination of the latter of these 
substances. 

Bertagnini (Annal. der Chem. und Pharmac., Feb. 1856) has - 
shown that this salicyluric acid is produced by the synthesis 
of salicylic acid with glycocol; while Horbaczewski states 
(Wiener Akad. Sitzb., II. Abth., 1885, Mai) that he has prepared 
uric acid synthetically from glycocol and urea, It is there- 
fore highly probable that salicylic acid, by uniting with glycocol 
to form salicyluric acid, prevents the production of uric acid. 
Dr Latham of Cambridge has already developed this view of 
the formation of uric acid in relationship to the treatment of 
gout by benzoates. 


IT. BENZOATE OF Sopa, 


Dr Rutherford, in Exp. 68 and 72a of his series, has 
demonstrated that benzoate of soda is a powerful hepatic 
stimulant. 

From the connection of benzoic acid and the benzoates with 
the production of hippuric acid, the influence of these drugs 
upon the urine has received a considerable amount of attention, 
though only two really careful scientific experiments have 
hitherto been made upon the action of the benzoates upon the 
nitrogen of the urine. 

Klitzinsky (Gist. Zitsch. f. pract. Heilk., Ba. iv. 8. 41, 1858) 
comes to the conclusion that no alteration in the nitrogenous 
matter of the urine oceurs. I have been unable to procure the 
original paper, which is referred to in Hermann’s Handbuch 
der Physiologie, vol. vi. p. 172. 

Meissner and Shepard, in their admirable paper “ Ueber das 
Enstehen der Hippursaiive im thier. Organismus,’ Hanover, 
1866, show that when hippuric acid appears in the urine 
after the administration of benzoic acid, the urea is not 
diminished, 

Salkowski (Zésch. f. Physiol. Chem., Bd. i. S. 45, 1877) has 
recorded two very careful experiments on the influence of 
benzoate of soda. The diet of the dog upon which these 
observations were conducted was carefully regulated, and a 
nitrogenous balance (stickstoffgleichgewicht) established. 


RELATIONSHIP OF UREA FORMATION TO BILE SECRETION. 281 


The following results were obtained :— 


|W. of Dog | Urine to Sp. G N. Bunsen’s | H.SO, as 


Date. Jin Kilos.| — c.cs. Method. | BaSo,. Remarks. 
16.7.75 19°62 400 1014°5 3°377 0°866 
17 : 1014°5 3°480 0°880 
18 19°65 ~ 1013°5 3°208 0-842 
*19 400 1028°5 4°865 1°332 5°12 grms. benzoic acid as 
soda salt. 
*20 19°62 400 1035°0 5°648 1°344 7°323 grms. benzoic acid as 
21 19°55 400 1014°5 3°976 0°736 soda salt. 
22 19°47 400 1014°5 3°132 0°884 
23 19°25 400 1015 3°440 0°840 
24 19°03 400 1015 3°568 0°850 
*25 19°05 425 1037 5°372 1°512 7°588 grms. benzoic acid as 
soda salt. 
*26 19°00 400 1038 5°652 1°206 7°527 grms. benzoic acid as 
27 18°83 - 400 1015 4°024 0°524 soda salt. 
28 18°73 410 ? 3°328 


Bunge and Schmiedeberg, in their paper on hippuric acid 
(Arch. f. Exp. Pathol., Bd. vi.), do not consider the influence of 
benzoates upon the nitrogen excreted. 

C. Virchow (loc. cit.) has made two experiments, under 
Salkowski’s direction, and finds a well-marked rise in the daily 
excretion of nitrogen under the influence of benzoic acid given 
as the soda salt. His results agree so entirely with those of 
Salkowski that it is unnecessary to reproduce them here. 

In regard to the action of benzoates upon the uric acid, it 
was long ago stated by Wohler and Keller (Ann. der Chem. u. 
Pharmac., xliii. 8. 108) that no decrease in the uric acid occurred. 

More recently, Garrod has investigated the subject, and has 
come to the conclusion that benzoates do very decidedly 
diminish the amount of uric acid excreted (“Lettsomian Lecture,” 
Brit. Med. Jour., 1883, vol. i.). His method of experiment 
was not altogether satisfactory. The urine of the 24 hours was 
not dealt with, and apparently no attempt was made to fix 
the diet. I here quote his last experiment (Exp. IV.), which 
gives results exactly similar to the other three, and will serve 
as an illustration of his method of procedure. 


I. Urine from 11 a.m. to 2 p.m., after 60 grs. of benzoate of soda: 
Urine = 5? oz. 
Uric acid = 0:17 grs. 

II. Urine from 11 a.m. to 2 p.m., after 120 grs. of benzoate of soda : 
Urine = 94 oz. 
Uric acid = 0°57 grs. 

III. Urine as above. No benzoate for 27 hours: 
Urine — 6 GZ 
Uric acid = 1:00 grs. 


282 DR D. NOEL-PATON. 


IV. Urine as above. No benzoate for 51 hours: 
Urine = 12.02. 
Uric acid = 1:25 grs. 

V. Urine as above. No benzoate for 75 hours : 
Urine = 10 oz 
Uric acid = 1-00 grs. 

Cook (Brit. Med. Jour., 1883, vol. ii. p. 9) opposes these con- 
clusions, and contends that the diminution observed by Garrod 
was due to the presence of benzoic acid in the urine preventing 
the crystallisation of the uric acid. 

Using the method of uric acid determination devised by 
himself, he gets the following results. The experiment was 
made upon a man with fixed diet :-— 


Date. Uric Acid. Remarks. 
May 14 13:0 grs. 
15 12°3 ,, 
16 13:0;,,4 
is ae 
18 13-2 ,, 
19 13°0 
20 on 500 
21 254 ae. 30 grs. benzoate of soda. 
22 13°7 9 60 9 ” 29 
23 14°0 9 40 ” ” By 
24 132 ;, 60 ,, 7 
25 A 


| 


The question of the influence of benzoates upon the uric acid 
must therefore be regarded as undecided, and as requiring 
further elucidation, 


Exp. V. 


The influence of benzoate of soda in the water, urea, and uric acid is 
shown in the accompanying table and abstract, and in fig. 4. 

The production of hippuric acid has already been so fully studied, 
that it was considered unnecessary to make any estimation of the 
amount produced. 

Its disturbing influence on the accuracy of the hypobromite method 
for the estimation of urea need not be considered, since Knop 
(Ber. d. konig. Sdchs, Gesell. d. Wissen., 1870, p. 17), and Hiifner 
(J. f. pract. Chemie N. F., Bad. iii. 1871, p. 18), have shown that 
none of the nitrogen of hippuric acid is evolved. 

The possible disturbing influence of hippuric and benzoic acid in 
the urine upon Haycraft’s method for uric acid had, however, to be 
investigated. 


> 


RELATIONSHIP OF UREA FORMATION TO BILE SECRETION. 283 


Hap. VA. 

Does the presence of benzoates in the urine influence the accuracy 
of Haycraft’s method of uric acid estimation ? 

Using Haycraft’s process, the urine of the 15th Feb. yielded 0-047 
of uric acid. 

To 25 c.cs. of this urine 0-2 grms. of benzoate of soda were added. 
The uric acid was again determined by the same method, and was 
found to amount to 0°039%. 


Exp. V8. 

Does the presence of hippuric acid vitiate the accuracy of Haycraft’s 
method ? 

To 25 c.cs. of the same urine, 0:2 grms, of hippuric acid was added, 
and the uric acid, determined as before, was found to be 0:041%. 

Therefore the presence of benzoates and of hippuric acid do not 
interfere with the accuracy of this process. 


On the 16th the urine contained a faint trace of albumin, and 
reduced Fehling’s solution very remarkably, giving on boiling a 
copious orange-coloured precipitate. So marked was this reduction, 
that I felt convinced that sugar was present and endeavoured to make 
a quantitative analysis. The amount of Fehling’s solution reduced 
indicated the presence of 4% of sugar. 

With Béttcher’s test the bismuth was only slightly reduced, giving a 
greyish colour. 

The fermentation test was also tried, and after several days gave no 
evolution of gas, although a check experiment with the same yeast 
yielded an abundance of carbonic acid. 

The reduction is due to the presence of a substance described by 
Salkowski in a short note in the Zeitsch. f. Physiol. Chem., Bd. iv. 
S. 135. Its exact composition has not been determined. 

The urine of the 17th gave a still more marked reduction of the 
cupric salt, but no evolution of carbonic acid occurred on the addition 
of yeast. 

On the 18th the reduction was decidedly less. 


Exp. V. 


{ 
Dates “apes 3p. G. Ureain | Urie acid Bowels. Remarks. 


cs, |°P- grms. | in grms. 


11.2.85| §745 | 1009] 6°910 0-230 |notmoved| Weight of dog=13°6 kilos. 
12 


(745 | 1009 | 6°910 0-230 | moved | Diet—Oatmeal 113 grms. 

| 13 $625 1011 6700 | 0°247 | not moved | Milk, 320 c.c. } 

14 (625 1010 | 6700 | 0°268 moved | | 

15 530 | 1013 | 6°095 0-212 not moved | 
16 650 | 1017 | 7°540 | 07130 moved 7-0 grms. benzoate of soda=0°51 


erm. per kilo. 


17 800 | 1023 | 11°680 0-216 | not moved| 7°5 grms. benzoate of soda=0°55 | 
grm. per kilo. | 

18 b> 210 1011 | 8°236 0-232 99 

19 / 700 | 1010! 7-140 | 0-210 ed 

20 |  Urin'e lost | | moved | 

21 | 500 | 1012| 5°600 | 0-263 | uvt moved | | 


22 760 1010 , 6°080 


a ae 


284 DR D, NOEL-PATON. 


germs 
11 
{Urie 
Acid 
‘rms. 
10 0°25 
9 0°20 
! 
A 
I 
8 1 0°15 


Feb. ne lo iT) 


Exp, V.—Excretion of,Urea and Uric Acidjund »1 jBen- 
zoate of Soda. 7 grms. given at a, and 7°5 germs. 
at e, 


Average Daily Excretion of various Constituents 
under Benzoate of Soda. 


3efore Percentage 
|| Before. | With. After. and With. CGhanae’ 
| After. ae | 
| 
| Water, in c.cs. |} 654 720 653 653°5 720 +10°00% 
| Urea, in grms. || 6°663 9°152 5840 6°251 9°152 + 46°47 
Uric Acid, ,, . 0236 0°195 0°236 0°236 0°195 —17°87 
| | 


Exp, V1. 


During the administration of the drug, which was given in the form 
of a pill made up with a little mucilage, the reducing power of the 
urine on Fehling’s solution was well marked. 

The influence of the drug is shown in the table and abstract, and 
in fig. 5, 


RELATIONSHIP OF UREA FORMATION TO BILE SECRETION. 285 


Exp. V1. 
ro, . and Uric 
Date. | Urine 10) Sp. G, Urea mM Acidin| 
by | | grms. germs, 
—_ a os ——_ ae ae ee 
23. 2.85, 900 | 1009, 87550 | 
24 (800 1010 | 8°550 | 
25 /(800 | 1011 | 8-335 | 
26 | | 675 1010 | 8-482 | 0°218 
27 |) 675 1011 | 8-482 | 0°218 | 
28 650 | 1022 | 11-700 | 0-186 | }7 
| | 
1.3. | 700 | 1015} 6160 | 0-132 | } = 
2 700 | 1024 | 13-020 | 0-132 ad 
= 590 1010 | 6°303 | 0°142 
4 590 1013 | 6°303 | 07142 
5 675 1012 | 8°140 | 07152 
6 675 1012 | 8°140 | 0-164 
7 710 1010 | 7°550 | 0°164 
8 710 1011 | 7°550 | 0:164 
Ig ie 1010 | 7:717 | 0°190 | 
10 742 1011 | 7°717 | 07190 
Wie, & 


ll 


10 


Feb. 


23 zZ3 
Exp. VI.—Excretion of Urea and Uri 
at a, 8 grms. 


Weight of dog=13'154 kilos. 
Diet as in previous experiments. 


‘5 grms. benzoate of soda=0°57 


Fs) 1U 
c Acid under Benzoate of Soda. 7 grms. given 
at b, and 7°5 grms, at c. 


Remarks. 


ee 
| 
| 


grms. benzoate of seda=0'53 
grm. per kilo. 
grms. benzoate of soda=0°6 
grm. per kilo. 


grm. per kilo. 


0.10 


286 DR D. NOEL-PATON. 


Average Daily Excretion of various Constituents under 
Benzoate of Soda. 


Before. | With. | After. | Before | With. | Percentage 


| and After. Change. 
i 

Water, inccs. . | 798 683| 664. 728 683 || -6°3% 

Urea, in grms. . || 8°425 | 10°293 | 7596 || 8-010 10'293 || +28°5% 

Uric Acid, ,, | 0-237 | 0133/0159 || 0-198 | 0-133|) —32°8% 


| 
| 


fesults.—My experiments merely confirm the observations of 
Salkowski and Virchow in regard to the urea excreted. 

They also show that in the dose given benzoate of soda has 
practically no influence upon the amount of water excreted. 

The point of chief interest is the clear demonstration afforded 
that benzoate of soda really dues diminish the uric acid excretion. 

At present [ cannot discuss the relationship of the hippuric 
acid produced to this diminution in the uric acid. It is a 
matter of great importance which I hope further to investigate. 


III. CoLcHicum. 


On the cholagogue action of colchicum Rutherford gives two 
experiments (Exp. 17 and 18), which show that this drug is 
undoubtedly an hepatic stimulant and a powerful hydrocathartic. 
The dose of the aqueous extract used in these experiments was 
large—60 grs. being given.? 

The influence of colchicum upcn the urinary secretion, and 
upon the urinary constituents has, from the connection of this 
drug with the treatment of gout, been much studied clinically. 

All the earlier observations on the subject, as those of 
Chelius (Lewins, Hd. Med. and Surgical Jour., 1841, p. 200), 
of Christison and Maclagan (Hd. Month. Jour. of Med. Sc., 3d 
series, vol, xvi. p. 24), are valueless, as the daily amount of 
urine is not taken into consideration. 

Boecker (Beitrage zur Heilkunde, Crefeld, 1849) concludes that 
colchicum has practically no effect on the renal secretion, but 
his experiments show too many fallacies to allow of his 
conclusion being accepted. 

Krahmer (Jour. f. Pract. Chemie, Dd. +1) gives the following 


1 In all probability the sample used was not freshly prepared. 


RELATIONSHIP OF UREA FORMATION TO BILE SECRETION. 287 


table, showing the effect of the administration of colchicam on 
the various ue Y the urine in grms :-— 


| 
| Dey. | Combus- 


'Amount.| Resi- rata Ash. Urea. | yar 
| : | > £ . 
due. nents. | 


| Mean of 62 observations 
| without the drug in usual | 
| condition, ‘ 2084°6 | 74°008 | 39°654 | 35°242 | 19°640 | 0'364 
Meanof 5 obser vations lur- 

| ing the use of tinct. col- 
_ chici seminum in doses of 
lto5drachms, . . | 1756°5 | 71°987 | 42°259 | 29°728 | 22°843 | 0°684 


Hammond (Proc. Philad. Acad. Nat. Sc., Dec. 1858, p. 18) 
gives a series of experiments upon the vegetable diuretics. He 
finds that, while digitalis, juniper, and squills increase the water 
excreted, colchicum alone increases the solid constituents. 
His experiments were conducted upon a healthy man, with slight 
precautions in regard to diet and exercise. The following table 
gives his results :— 


Urine. |Sp.G. | Total Solids. sabia Organic. 
Normal standard, . «|, 1280) | 1025 63°12 29°83 — 33°29 
Under colchicum, . ‘| 1556 | 1023 77°28 35°23 | 42-04 


S. R. Percy’s paper on the subject of the influence of colechicum 
upon the uric acid excretion (Amer. Med. Times, 1862) is absol- 
utely valueless, as the method adopted for estimating the uric 
acid—by observing the amount of precipitate—is exceedingly 
crude and fallacious. 

Garrod, in his admirable “Treatise on Gout and Rheumatic 
Gout,” has given the results of a tolerably full investigation into 
the action of colehicum upon the uric acid, and of one or two 
observations on its influence upon the urea. Two experiments 
upon healthy patients are given. 

In his first experiment the average amounts of water and of 
uric acid were as follows :-— 

For four days before administration of any drug— 


Water averaged 68°5 fl. oz. per diem. 
Uric acid 99 | 8°24 grays jy. 1155 


288 DR D. NOEL-PATON. 


For five days under colchicum the average was— 


Water 55°6 fl. oz. per diem. 
Uric acid Oe ie ae 
In his second experiment the averages for three days before 
the drug was given were— 


Water 
Uric acid 


37°7 fl. oz. 
5:03 grs. 


Averages for five days while drug was given-— 
Water = 25:2 fi. oz. 
Uric acid = 529 grs. 

From a more careful study of the details of these experiments 
we see that nothing can be deduced from them. The daily 
variations in the uric acid excreted were very great, and com- 
pletely vitiated the results. 

Garrod also gives six observations on the urine of gouty patients. 
Observations on the action of a drug in disease are of little 
value, and we therefore refrain from quoting these experiments. 

From his observations he is led to the following conclusions :— 

“1. There is no evidence that colchicum produces any of its 
effects upon the system by causing the kidneys to eliminate an 
increased quantity of uric acid; in fact, when the drug is 
continued for any length of time it appears to exert a contrary 
effect. 

“2, We cannot assert that colchicum has any effeet upon the 
excretion of urea or the other solid ingredients of the urine. 

“3. Colchicum does not act as a diuretic in all eases; on the 
contrary it often diminishes the quantity of urine, more especially 
when it produces a marked effect on the alimentary canal.” 

Certainly, in respect to the action of colchicum upon the 
urea and uric acid, Garrod has little ground upon which to 
base his conclusions. Painstaking and careful as are his experi- 
ments they are open to all the many fallacies of clinical obser- 
vation; while his use of the old method of Heintz for the 
estimation of uric acid renders even his analysis open to 
objection. ’ 

Obviously the question of the action of colchicum on the urea 
and uric acid of the urine must be regarded as totally unsettled 
and requiring careful experimental study. 


RELATIONSHIP OF UREA FORMATION TO BILE SECRETION. 289 


Exp. VII. 


In this experiment very large doses of the drug were given, and as 
is seen in the abstract a much smaller percentage increase of the urea 
occurred than in the next experiment. The marked purgation which 
was induced may have prevented a greater increase in the urea 
excretion. 

The influence of colchicum on the uric acid is not well illustrated in 
this experiment—though on the second day of the administration of 
the drug a well-marked rise occurred. In all probability the diarrhoea 
induced had something to do with the great fall which occurred on the 
third day. 


Exp. VII. 
| : : 
Urine eer Uric 
|. Date. | in Sp.G ae an | Acid in Bowels. Remarks. 
c.cs. erm: grms. | 
15.12.84; 620 | 1010 | 6°262 | 071041 moved Weight of dog=12°68 kilos. 
16 460 | 10J1 | 5°520 | 071381 Ac Diet as in previous experiment. 
17 565 | 1012 | 6°780 | 0°1911 not moved 
18 520 | 1013 | 7°800 | 0°1394 moved 0°5 grm. acetic extract of col- 
chicum=0°039 grm. per kilo. 
19 540 | 1014 | 8262 | 0°2106 | copioussoft semi-| 1°0 grm. acetic extract of col- 
fluid evacuation chicum=0'078 grm. per kilo. 
20 425 | 1016 | 8062 | 00858 copious soft 11 grm. acetic extract of col- 
evacuation chicum=0'086 grm. per kilo. 
21 ( 492 7-718 | 0:0800 | several watery 
evacuations 
22 | (492 7-718 not moved 
23 400 | 1013 | 6°560 moved 


Average daily Excretion of the various Constituents under 
the influence of Colchicum. 


| 
| Before | With | After | pe | With Percentage 
| Drug. | Drug. | Drug. | Renn Change in. 
ae te pee | 
| Water, in c.cs. || 548 495 461 Sees | = 1°67 
Urea, in grms. | 6187 8041 7332 6°759 8041 +1817 


The uric acid is not given in this table, as it was not erage on P the two days 
succeeding those on which colchicum was given. 


Exp. VIII. 


The influence of moderately large doses of the acetic extract of 
colchicum on the water, urea, and uric acid daily excreted is shown in 
the accompanying table ‘and abstract, and in fig. 6. 

In this experiment the chief point of interest, apart from the special 

VOL, XX. dt 


290 DR D. NOEL-PATON. 


action of colchicum, is the illustration it affords of the influence of 
purgation. On the 12th and 14th a copious soft motion followed the 
administration of the drug, and the urea was only slightly increased, 
while upon the 13th, when the bowels were not moved, a greater rise 
occurred. 


Exp. VIII. 


‘) Urine in | Urea in| Uric Acid | 
Date. | ¢cs, |5P-&) gyms. | in grms. | Bowels. Remarks. 
e 1.85} § 375 1013 | 5°045 0°095 moved Weight of dog=13'6 kilos. 
1 375 1013 | 5:045 0-095 59 Diet as in previous experi- 
0 (470 | 1012] 57140 0°124 not moved ment. 
11 1470 1011 | 57140 0°124 : = 
12 520 1014 | 7°644 0°122 . copious soft motion | 0°5 grm. acetic ext. col- 
chici = 0°037 per kilo. 
13 | 620 1015 | 10°726 0°167 not moved | 0-3 grm. acetic ext. col- 
| chici = 0:02 per kilo. 
14 | 610 1016 } 8°834 0°142 ~=copious re are 0°4 grm. acetic ext. col- 
15 | 475 1014 | 6°692 0°132 moved | chici = 0°029 per kilo. 
| 16 | $498 | 1013} 6°040 0-070 not moved | 
17 | (498 | 1012] 6°040 0°107 moved | 
118 | 550 | 1011} 5°940 0°107 not moved 
grms, 
10 
grms. 
. 0°15 
Uric 
8 Acid. 
0°10 
d 0°05 
6 
Urea 
5 
Jan. 10 15 


Exp. VIII.—Excretion of Urea and Uric Acid under 
Colchicum, 0°5 grm. of acetic extract given at a, 
0°3 grm, at e, and 0°4 grm, atc, 


RELATIONSHIP OF UREA FORMATION TO BILE SECRETION. 291 


Average daily Excretion of various Constituents under Colchicum. 


| 
: | Before 
} Before | With | After rr With. || Percentage 


Drug. | Drug. | Drug. After. Change in. | 
| | | 
; * 1] | | ; 

Water, in c.cs. | 422 583 | 505 463 583 +25°9% 
Urea, in grms. | 5092 | 9°068 | 6178 5°635 | 9068 +60°9% 
Uric Acid, ,, 0109 | 0-143 | 


0-102 || 0-105 | 0-143 +3614 


Exp. 1X. 


In this experiment a recently prepared sample of acetic extract of 
colchicum was used. This probably accounts for the great purgation 
induced by small doses. 


- Urea | Uric 
| Date. | ,Utine isp.a.| in | Acidin| Bowels. | Remarks, 
“peak | | grms.} grms. | 
| 
19.7.85 } 640 1009 6528 | 0-083 moved | Weight of dog=13-37 kilos. } 
| 20 580 | 1009 | 5916} 0-097 “P Diet as in former experiments. 
2 640 | 1009} 6-016 | 0-083 | not moved | 
| 92 700 1008 | 5°600 | 0°119 | moved—copi-! 0:2 grm. acet. ext. colchici (fresh), 
| ous soft i =0°014 grms. per kilo. 
1 23 740 1010 | 8314 | 0126 | moved—soft, | 0°2 grm. acet. ext. colchici (fresh)| 
| mucous i =0°014 germs. per kilo, 
evacuation | 
24 (645 1008 | 6°342 | 0°074 |moved—loose! 
25 1645 1009 | 6°342 | 0°077 | moved 
2 .# 480 1010 | 6°490 | 0-070 “; 
27! 480 1010 | 6-490 | 0-096 | af 


i=. 
z ~» e 


germs. 
8 


r 
' 


July 20 25 


Exp. IX.—Excretion of Urea and Uric Acid 
under Colchicum. 0°2 grm. acetic extract 
given at a, again at e, Very free purga- 
tion induced. 


292 


DR D. 


NOEL-PATON. 


Average daily Excretion of various Constituents under Colchicum. 


; Befo 
Before | With | After - 4 i With Percentage 
Drug. | Drug. | Drug. After Change in. 
Water, in c.cs. | 620 720 562 591 720 +21°8% 
Urea, in grms. 6 °225 6°957 6°417 6°321 6°957 +10% 
Uric Acid, ,, || 0°087 0°122 0°079 0°083 0°122 aS 47%, 
Exp. X. 
nine Urea | Uric 
Date. lated Sp. G in Acid in Bowels. Remarks, 
ger. grms. | germs. 
24.97.85 { 645 1008 | 6°342 | 0°074 moved Weight of dog=13'15 kilos. 
25 645 1009 | 6°342 | 0°077 7, Diet as in previous experiments. 
26 § 480 1010 | 6°490 0:070 24 
27 ( 480 1010 | 6°490 0°096 es 
28 Age ar ioe ” 
29 520 1010 | 5°772 0°087 ” | 
30 580 1009 | 5°563 | 07128 not moved | 0°15 grm. acet. ext. colchici=0°0114 | 
grm. per kilo. 
31 620 1009 | 4:960 | 07104 | moved—very | 0°20 grm. acet. ext. colchici=0°0147 
loose grm. per kilo. 
1 605 1011 | 6°836 | 0°109 | moved—very | 0°20 grm. acet. ext. colchici=0°0147 
loose grm. per kilo. 
2 530 1013 | 5°500 | 0°057 moved 
3 530 1007 |? 5°500 0°075 cs 
4 620 1009 | 5°300 0°091 “4 
5 620 1009 | 5°456 0°085 9 
Fig. 8 


Urea 
grms. 
6 


5 


July 


25 


30 


grms 
0°10 

Uric 
Acid. 


0°05 


5 


Exp. X.—Excretion of Urea and Uric Acid under Colchicum. 


at 


c. 


0°15 grm. of acetic extract given at a, 0:2 erm. at e,and again 


| 


RELATIONSHIP OF UREA FORMATION TO BILE SECRETION. 293 


Average Daily Excretion of the various Constituents under 


Colchicum. 
| ae ae 
|| Before | With | After || Before With 
| Drug. | Drug. Drug. |jand After. Pa 
1] 
Le | ee a 
ae | be 2 asd as bas | 
Water, in c.cs. : 548 | 571 606 557 5/1! | 
Urea, ingrms. . : 6°204 | 5°786 | 5°453 5828 5°786 
Uric Acid, ,, : . 0°081 | 01138 0°077 0°079 0°113 
Percentage change in 
Water, . ; ; . practically unaltered 
Urea, .. < . : 23 He 
Uric acid, ‘ ; . +43% 
Exp. XI 
| 
Urine s Uric 
| Date. | in [Sp.G yeaa Acidin| Bowels. Remarks. 
c.cs. > = “| grms. | 
26.7.85 | § 550 | 1009 | 67182 aaa moved Weight of dog=13°'25 kilos 
27 (550 | 1009 | 67182 aot - Diet as in previous experiments. 
28 { 620 | 1009 5°816 | 0°0621 = 
29 ( 620 | 1008 5°816 | 0°0621 
30 620 | 1010 5°332 | 0°0682 
31 ee 1010 5°454 | 0°0526 ; 
1.8 562 | 1010 5°454 | 0°0606 » 
2 620 | 1009 | 5-564 | 0-0806 x +s 
3 780 | 1009 | 7176 | 0°141 moved— | 02 grm. acet. ext. colchici=0°015 
rather loose | grm. per kilo. 
| 4 720 | 1009 | 6°523 | 07106 moved— 0°2 grm. acet. ext. colchici=0°015 
| | very loose grm. per kilo. 
| 5 } 700 | 1009 6°720 | 0°091 | notmoved 
Fig. 9 


grms. erms. 
7 0°10 
Uric 
Acid. 
Urea 
6 0°05 
July 26 30 5 


Exp. XI.—Excretion of Urea and Uric Acid under CoJchicum. 
0'2 grm. of acetic extract given at a, and again ate. 


294 DR D. NOEL-PATON. 


Average Daily Excretion of various Constituents under 


Colchicum. 
| 
| Before. With. | Percentage 
| Drug. Drug. | Change in. 
Water, inecs, , © «|| 577 735 «| «| 4878 
; Urea,ingrms. . ; : 5°7438 6°784 +18 
| Uric Acid, . : | 0°0646 0°112 | +73°3 


| 


Results—The above five experiments clearly show that in 
medium doses colchicum increases the water and urea excretion 
to a moderate amount, and the uric acid excretion to a much 
greater extent. Indeed the most striking feature of these experi- 
ments is the very marked increase in the uric acid production 
indicated by them, an increase which renders it difficult to 
explain the well-known beneficial action of this drug in gout. 
For, that an increased production and not merely an augmented 
excretion is indicated by these experiments is proved by the 
fact that after the administration of the drug is discontinued no 
great fall below the normal daily excretion occurs. 


LV. PERCHLORIDE OF MERCURY. 


Experiments 784, 788, 78c, and 78D of Professor Rutherford’s 
series clearly show that perchloride of mercury is a cholagogue 
of considerable power. 

In spite of the length of time during which mercury has, in 
different forms, been largely employed in medicine, I can only 
find one or two experiments on its action on the tissue meta- 
bolism (stofwechsel), as estimated by the amount of nitrogen 
excreted, 

Harvey (Brit. and For. Medico-Chir. Rev., vol. xxix. p. 515) 
records a series of experiments on the action of blue pill mass 
and perchloride of mercury upon the urea excretion in dogs, 

The dogs were kept in a cage, the urine being collected much 
as inmy own experiments. The diet consisted of “ paunch,” This 
is a form of dog’s meat sold in London, and consists, I believe, in 
the intestine, stomach, liver, &c., of sheep and oxen. 

In Exp. I. the dog got 6 oz. of “ paunch ” and 4 pint of water 


RELATIONSHIP OF UREA FORMATION TO BILE SECRETION. 295 


daily. For seven days before the administration of the drug the 
urine was daily collected and the urea estimated. 

During the next fourteen days the pil. hydrarg. was given in 
doses, commencing at 2} grs. and increased to 15 grs., so that 
in the fourteen days 100 grs. of the drug were taken. The 
following table gives the average excretion of water and urea 
before and under the administration of the drug. 


Before With 
Water ; 203°187 grms._. 212°39 grms. 
Sp. G. 1041 ; 1043 
Urea : 13 grms. 12°9 grms. 


In Exp. II. the dog got 12 oz. of “paunoch” and 4 pint 
of milk. For nine days no drug was given, then for nine days 
the liquor hydrargyri perchloridi of the pharmacopeeia in 
doses of one or two drachms. The larger dose produced 
diarrhcea. 

Average daily excretion before and with the drug :— 


Before With 
Water ; 393°687 grms._. 315°7 grms. 
Sp. G. ‘ 1026 : 1031 
Urea : 8515 grms. , 8:580 grms. 


During this experiment the water and urea excreted varied 
much from day to day. 

Exp. III. was similar to Exp. IL, but the dog was kept for 
twelve days without the drug, and for twenty-one days it had 
daily the liquor hydrargyri perchloridi in doses of from one to 
two drachms. The following results were obtained :-— 

Average daily excretion before and with the drug :— 


Before. With. 
Water ; 339°06 grms. : 342°77 grms. 
Sp. G. ; 1030 : 1024 
Urea : 15°9 grms. : 14°323 grms. 


During these experiments the dogs are said to have enjoyed 
perfect health. 

Unfortunately these experiments are not satisfactory. The 
diet given was much too liberal, and we find in consequence 
very large daily variations in the amounts of water and of urea 
excreted. Besides such a substance as “paunch” is not of 
sufticiently fixed a composition to render it suitable for such 
experiments, 

The only other observation of any value which I have been able 


296 DR D. NOEL-PATON. 


to find is by Hermann v. Boeck (Zeitsch. f. Biol., vol. v. p. 393). 
It is to be regretted that this most careful observation was not 


made upon a healthy man. Unfortunately a syphilitic case was 


selected, so that the results obtained must be accepted with 
great caution. 

The method of experiment was ingenious and admirable. 
The nitrogen contained in the diet was estimated and compared 
before and under the administration of the drug with the nitrogen 
excreted by the kidneys and in the feces. 

The diet was liberal, consisting of eggs, bread, milk, beef 
extract or soup, beer, butter, and salt. 

The patient was a man aged 44, who had been infected with 
syphilis three months before, and who suffered at the time from 


secondary syphilis, condylomata, &c. On the 10th of Oct. he 


was put upon a fixed diet, and from the 12th to the 15th the 
urea excretion was nearly constant. On the 15th mercurial 
inunctions were commenced. On the 20th salivation occurred, 
and on the 22nd the eighth and last inunction was given. 

Boeck gives a full table of the nitrogen in the diet, the amount 
of urine, the sp. gr., the urea, the nitrogen in the urine, and the 
nitrogen in the feces. 

The following table shows his results, so far as they concern 
the present question :— 


Date. Remarks. Nit. of Diet. | N. Excreted. Urea. 
10 16°1 12°4 26°7 
11 16°7 17°4 30°7 
12 ilyfal | 16°3 3l°3 | 
13 i (ay | 18°6 34°5 
14 173 17°5 32°0 
15 | Inunction daily till 22nd. 1/333 18°6 34°9 
| 16 175 | 18°8 35°0 
ita Rg 175%. | es 33-9 
18 Sone Diarrhea. 17°8 19°2 33°1 
19 17°5 18°5 35°4 
20 174 | ~~ 192 370 
21 17°8 18:0 35°5 / 
22 Diarrhea. 17°5 19°0 | 33'1 
23 17h 18°0 32°3 
24 177 17°9 33'3 
25 var. 179 32°8 


RELATIONSHIP OF UREA FORMATION TO BILE SECRETION. 297 


Before the inunctions—from the 12th to the 15th 


52°1 grms. of nitrogen were taken in | _ i) 
524 F- ‘a x ate ante OY, 


During the application period— 


193:2 grms. of nitrogen taken in | _ + 56% 
204°0 grms. of nitrogen excreted f ~ s 


I cannot agree with Boeck’s conclusion that, “Dieses Plus ist 
ganz unwesentlich und beruht zum Theil auf Fehlern der 
Methode, zum Theil vielleicht auf den Diarrhoén, die mehr 
stickstoffhaltige stoffe den Korper entfiihrten.” 


Exp. XI. 

The perchloride of mercury was given in pills made up with starch. 
Even with the dose given, 0°037 grms., which was large, absolutely no 
physiological effects were produced, and it is extremely probable that 
the drug was never absorbed, but that the pills passed through 
undigested. 


Eup. XI. 


The results of Experiment XI. were negative, probably, as before 
stated, on account of the form in which the drug was given. 

Experiment XII. was made on another dog weighing 13°14 kilos, 
which had been kept for more than a month upon a diet similar to 
that used in the former experiments. Before the drug was given it 
had been for more than a week excreting an almost equal amount of 
urea each day. The drug was given in the form of a pill, made up 
with a very small quantity of oatmeal and gum. During and follow- 
ing the administration of the drug no constitutional symptom could 
be detected. 


Kup. XII, 

Urine in| Urea in | Uric Acid 
Date. en ese G. grms, | in grms. Bowels. Remarks. | 
18.4.85 580 | 1010 | 67148 0°075 moved Weight of dog=13'14 kilos. +5 
19 650 | 1010 | 6°500 0084 - Diet as in previous experiments. | 
20 635 | 1010 | 6°500 “He aA 
21 635 | 1011 | 67500 0°102 o 
22 685 | 1009 | 6773 0-084 a 
23 685 | 1010 | 6°773 0°108 ~ 
24 605 | 1011 6°715 0°138 =" 
25 740 | 1010 | 7-400 07148 f 0°02 grm. perchloride of mercury | 

| in pill=0°0015 grm. per kilo. 
26 780 | 1009 | 6°552 07101 Pe 

| 
27 655 | 1011 | 7°598 0°085 P 0°04 grm. perchloride of mercury 
in pill=0°0030 grm. per kilo. 

28 810 | 1015 | 14°580 0°137 3 0°05 grm. perchloride of mercury 
29 745 | 1009 | 7:003 0°097 iy in pill=0°0038 grm. per kilo. 


30 630 | 1011 | 6°300 0°082 ” 


298 


DR D. NOEL-PATON. 


Average Daily Excretion of the various Constituents under 


Perchloride of Mercury. 


Before | With | After Before |yw34),_ || Percentage 
Drug. | Drug. | Drug. || and After. * | Change in. 

Water, in c.cs. , 639 745 | 630 634 745 +17% 
Urea, in grms, 6534 | 7°978 | 6300 6°408 |7°978 +24% 
Uric Acid, ,, - 0°098 0°107 | 0:082 0°090 |0°107 +18°8% 
Hap, XIII. 

rine : Uric 
Date. in (|Sp. G. Uisein Acid in} Bowels. Remarks. 
C.C8. o | grms, 

22.5.85 oy 1010 | 5°457 | 0°078 | moved | Weight of dog=13°37 kilos. 

23 517 | 1010} 65:457 | 0°073 4S Diet, as in previous experiment. 

24 550 | 1010 | 5170 0-088 ” 

25 560 | 1010} 6°600 0:089 _ 

26 700 | 1009 | 6°020 0-091 + 

27 690 | 1008 | 6°072 0°116 3 

28 620 | 1008 | 5:454 me :, 

29 705 | 1007 | 6°768 | 07118 s 0°0166 grm. HgCl, in iodide of potassium ; 

solution=0-'0012 grm. per kilo. 

30 600 { 1010 | 7°320 | 0088 A 0°033 grm. HgCl, in iodide of potassium 

solution = 0°0022 grm. per kilo. 

31 670 | 1010 | 7:236 | 0:088 0°066 grm. HgCl, in iodide of potassium 
1°6 600 | 1010} 5160 0°078 2 solution =0°0049 grm. per kilo. i 
2 ee 1008 | 6°310 0°073 + 
3 622 | 1009 | 6°310 0-094 a 
4 {605 | 1009 | 6°310 0°092 i 
5 1605 | 1008} 6310 | 0-092 x 

Fig. 10 
grms. 
6 
5 ) | | 
May 22 25 30 5 


Exp. XII.—Excretion of Urea under Perchloride‘of Mercury in Potassic 


Iodide. 


0°066 grm. atc. 


00166 grm perchloride given at a, 0°083 grm. ate, and 


° 


| 


RELATIONSHIP OF UREA FORMATION TO BILE SECRETION. 299 


Average Daily Excretion of the various Constituents under 
Perchloride of Mercury. 


Water, in c.cs. 
Urea, in grms. 
Uric Acid, ,, 


Eup. XIV. 
Urine 
Date. ino.cs. Sp. G. 
13.65.85} 670 | 1011 
14 630 } 1010 
15 640 | 1010 
16 510 | 1010 
17 560 | 1010 
18 590 | 1009 
19 560 | 1011 
20 675 | 1011 
21 420 | 1010 
22 { 517 | 1009 
23 517 | 1010 
24 550 | 1010 
25 560 | 1010 


Urea in 
grms. 


6°099 
67114 
6°144 
5°814 
5°936 
4°720 


6°216 
8100 


5°156 
5°457 
5°457 
5°170 
5°600 


| Before 
| Drug. | Drug. | Drug. 


5°604 
0°091 


Uric 


Acid in 


grms. 


0°0957 
0°0819 
0°0832 
0°0856 
0°0784 
0°099 
0°112 
0°087 
0°0705 
0°053 
0°094 


0°088 
0°089 


May 


io 


of Mercury. 
grm. atc. 


With | After 


| 


te) 
” 
” 


moved (soft) 


moved 


Pa) 
Exp. XIV.—Excretion of Urea and Uric Acid under Perchloride 


Diet of Oatmeal, 113°6 grms, 


Milk, 320 c.cs. 


0°05 grm. HgCl, in pills of 0°025 grm. 
each=0-0037 grm. per kilo. 

0°10 grm. HgCl, in pills of 0°050 grm. 
each=0°0074 grm. per kilo. 

0°10 grm. HgClein pills of 0°050 grm. 
each=0'0074 grm. per kilo. 


Before : Percentage 
| and After. With. Change in. 
649 609 | 606 649 +7°09% 
7°103 | 5°735 5°699 6°694 +21°5% 
0'098 | 0°084 || 0:°087 0°098 +12°6% 
| 
Bowels. Remarks. 
moved Weight of dog=13°37 kilos. 


orms. 
0°10 


Uric 
Acid, 


0°05 


005 grm. given at c, 0°10 grm. at e, and C10 


300 DR D. NOEL-PATON. 


Average Daily Excretion of the various Constituents under 
Perchloride of Mercury. 


Before | With} After Before With Percentage 
Drug. | Drug. | Drug. || and After. * || Change in. 
Water, in c.cs. . 582 608 | 513 547 608 || +11°15% 
Urea, in grms. . || 6°021 |6°345 | 5°346 5683 6°345 || +11°6% 
Uric Acid, ,, . || 0°085 |0°099 | 0:079 0°082 0-099 || +11% 
| 


Eup. XV. 


. Urea Uric 
Date. |.Utine Ign Gg] in |Acidin| Bowels. Remarks. 


TRICE grms. | grms. 
2.6.85 | (622 1008 | 6°310 0°088 moved Weight of dog=13'37 kilos. 
3 (622 1009 | 6°310 0°088 % Diet of Oatmeal, 113°4 grms. 
+ (605 1008 | 6.310 0°092 +S Milk, 320 c.c. 
5 1605 | 1009] 6310 | 0-092 = 
5 640 1008 | 6°016 0-083 op 
7 $652 1008 | 5°660 0°090 == 
8 (652 1009 5*660 a8 s 
9 670 1010 | 7°303 | 0170 5 0°06 grm. HgCl, in 0°5 c.c. satur- 
| ated solution of potass. iodidi 
in gelatine capsule 00044 grm. 
| per kilo. 
10 605 1013 | 7381 0°085 |moved—loose| 10°087 grm. HgCl, in 05 c.c. 
saturated solution of potass. 
| iodidi in gelatine capsule 0°0065 
grm. per kilo. 
11 710 | 1007} 6319 | 0:205 ¥ 0075 grm. HgCl, in 0°5 c.c. satur- 
ated solution of potass. iodidi 
| in gelatine capsule 0°0056 grm. 
per kilo. 
12 560 1010 | 6°720 ee moved 
13 575 1007 5°750 ae AA 
14 (552 1007 5°847 52 4 
15 552 1007 5847 0095 £0 


1 Urine contains a distinct trace of albumen, and the fermentation test shows the 
presence of a small quantity of sugar. 


25, 
grms. germs 


0°15 


Uric 
Acid, 
0°10 


0°05 


June 2 5 : 10 15 
Exp. XV.—Excretion of Urea and Uric Acid under Perchloride 
of Mercury in Solution of Potassic Iodide. 0°06 grm. given 

at a, 0°087 grm. at e, and 0°075 grm. atc. 


RELATIONSHIP OF UREA FORMATION TO BILE SECRETION. 301 


Average Daily Excretion of various Constituents under 
Perchloride of Mercury. 


| I 
| Before | With) After || Before | wii, | Pereentagel 
| Drug. | Drug. | Drug. || and After. * | Change in 


Water, ine.cs. I, 619 | 636 | 568|| 593 636 || +7:2% 
Urea, in grms. . || 59995 |6°930 | 5°798 || 5°896 6°930 | +17°5% 
Uric Acid, ,, . | 0-088 |0-158 | 0-095) 0-091 | 0158 | +687 

| | | | 


Results—Both the perchloride of mercury and the iodide 
cause a distinct rise in the water, urea, and uric acid excre- 
tion. My reason for using the iodide will be explained in 
a future paper, while considering the nature of the relationship 
between the urea and bile-forming functions. 


V. EUVONYMIN. 


Experiments 27 and 28 of Professor Rutherford’s series show 
that euonymin is a powerful hepatic stimulant. 

In regard to its action on the composition of the urine I can 
find only one observation, by Cook (Brit. Med. Journal, vol. i. 
1883, p. 1060). 

To a man kept on fixed diet, one grain of the drug was 
administered on an empty stomach on the days marked with an 
asterisk. 


j | 
Date. | Urine. | Sp. G. | Urea. Uric Acid. | 
Si oscee ita | 
| | | : 
Noy. 1* | 38 oz. | 1020 | 360 grs. 11°7 grs. 

oF ee $4, 1022 Lo. eee 14:3 ,, 

get eget es b> 1020. <4. sae: 17°4 ,, 

4 Ms 1020 |= OS Bay ae 1253. 5; 


5 43 ,, 1020 340 ,; 12°5 


He concludes from this very insufficient experiment that 
while ecuonymin exerts no influence upon the urea excretion it 
increases the excretion of uric acid. 

Practically nothing is known of the action of the drug upon 
the urinary constituents. 


302 DR D. NOEL-PATON. 


Exp. XVI. 


The dog had been upon a fixed diet from October 8, and had lost — 
3:17 kilos weight. It was, however, healthy and strong. 

Before November 3 the variations in the daily urea excretion were 
considerable, but the average from October 28 to November 3 was 5:4 
grms. per diem. From November 3 the daily excretion became more 
constant, and on the 7th 0°5 grms. of green euonymin, procured through 
Duncan & Flockhart from Kieth & Co., was administered in the 
form of a pill made up with a drop or two of alcohol, and next day 
0°75 grms. were given. 

The accompanying table, abstract, and fig. 13 show the influence of 
the drug upon the urine. 


Exp. XVI. and XVII. 


| Urine Fane oe | UEC, 
; Date. in (Sp. G. peice | Acid in Bowels. Remarks. 
} ¢.cS. eetilos) grms. 
4.11.84] 450] 1011 | 5°535 | 0°042 | notmoved | Weight of dog=12-23 kilos. 
5 550 | 1011 | 4:840 | 0-100 re Diet, Oatmeal=85 grms. 
6 550 } 1011 5650 07110 moved Milk, 300 c.c. | 
7 575 | 1013} 7977 | 07132 7 Euonymin 0°5 grm.=0-041 grm. per | 
kilo, ; 
8 | 500} 1015 | 7°358 0°083 3 Euonymin 0°75 grm.=0'06 grm. per 
| kilo. | 
9 | 465} 1014} 5°580 | 0109 | not moved 
10 470 | 1013 | 5°000 | 0-047 moved } 
11 | 450} 1013 { 5°265 0°060 a | 
12 | 455 | 1015 | 6°998 | 0°042 | copious soft | Eunoymin 1°5 grm.=0°1 grm. per | 
evacuation kilo. | 
13 500 | 1010} 4800] ... copious soft | Euonymin 2°0 grm.=0°16 grm, per | 
evacuation kilo. | 
Fig. 13 
germs. 
0°15 
0°10 
0°05 
Uric 
Acid. 


Exps. XVI. and XVII.—Excretion of Urea 
and Uric Acid under Ruonymin. 0°*5 grm. 
given ata; 0°75 grm. at; 1°5 grm. at a’ 
and 2°0 grms. at 0’. 


RELATIONSHIP OF UREA FORMATION TO BILE SECRETION. 303 


Average Daily Excretion of the various Constituents under Euonymin. 


aA 


Before | With | After Before | wi) |} Percentage 
Drug. | Drug. | Drug. |jand After. * || Change in 


——— = | 


Water, in c.cs. 56 537 461 488 537 || +10°0% 
Urea, in grms. || 5°341 7°667 5°432 5'386 7°667 +42°3% 
Urie Acid, ,, || 0°084 | 0-107 | 0-072 0078 | 0-107 || +37-14% 


Exp. XVII. 

This is simply a continuation of Experiment XVL., larger doses of the 
drug being given, 1°5 grms. on the 12th, and 2°0 grms. on the 13th. 
On the second day the purgative action of the drug was very marked, 
while even on the 12th the motions were soft and unusually copious. 
It will be seen that, in this experiment, the urea was not so markedly 
increased as in the previous one, in fact that on the 13th, when the 
purgative action of euonymin was so manifest, a fall below the mean 
took place. 

At the end of this experiment the dog was thin and looked ill. The 
diet was consequently increased and the uniform excretion of urea per 
diem was disturbed. 


Exp, XVIII. 

The dog had been upon the usual diet since November 14, and in 
spite of considerable daily variations, the average excretion of urea, from 
November 25 to December 1, was 7:02 grms. per diem. 

The experiment was commenced on December 1. Unfortunately 
the urines of the 2nd and 3rd were lost. On December 6, 1 grm. of 
green euonymin, made into a pill with absolute alcohol, was adminis- 
tered. Next day asimilar dose was given ; and on the 8th 1°5 grm. in 
the same form. 

It is of interest to observe that upon the 7th, when a loose copious 
motion was produced by the drug, the urea was not nearly so mani- 
festly increased as on the two other days. 

The uric acid participated in the irregularity of the urea in its rate 
of elimination, and it appeared in this case to be uninfluenced by the 
drug, unless indeed the rise on the 8th, 9th, and 10th are to be con- 
sidered as due to the euonymin. 


Exp. XVIII. 

; «| Uric 

Date. ied Sp. G. Urea in Acid in Rowels. Remarks. 
n¢.cs. grms. 
grms 
ee SSS OS eee 
1.12.84 | 650] 1008 | 6°240 0°154 moved Weight of dog=13'16 kilos. 
4 720 |} 1012 | 7°421 = not moved | Diet, Oatmeal, 113 grms. 
5 680 | 1010 | 7°820 | 0190 moved Milk, 320 c.cs, 
6 750 | 1018 | 8800 | 0150 | not moved | 1:0grm. Euonymin=0°08 grm. per kilo. 
7 750 | 1010 | 7°725 0°165 | soft motion | 1°0 + =0°08 ar 
8 750 | 1013 | 8-795 | 0-202 moved | 1°5 a =010 Ss 
9 650 | 1009 | 5°499 0°204 39 | 
| 


” 


505 | 1014] 7-777 | 07156 , 


id 
Ko 
oO 
a 
on 
_ 
o 
ms 
o 
on 
nr 
ive) 
we 
o 
is] 
So 
= 


304 DR D. NOEL-PATON, 


Fig. 14 
grms. 
9 


8 grms 
0°20 
Uric 
= Acid. 
‘ 0°15 
Urea 
6 0°10 


Exp. X VIII.—Excretion of Ureaand Uric Acid 
under Euonymin. 1°0grm. given at a, and 
at e, and 1°5 grm. atc. 


Average Daily Excretion of the various Constituents under Euonymin. 


, : as | Before | w; Percentage 
Before. | With. After. and After. | With. | Change Ba 


| 
| 


| 

Water, in'c:es. aie 683 | 750 573 628 750 | +19% 

Urea, in grms. . , 77160 | 8°440  6°286 6°723 8°440 | +20°37 | 
| 


| 


The uric acid is not given in this table, as a marked increase took place in the 
three days immediately following the administration of the drug. 


Results——1st. The water is slightly though distinctly increased 
under the action of euonymin. This is just what one would 
expect from the digitalis-like action of the drug. 

2nd. The urea is very markedly increased, especially when 
purgation is not induced. Experiments 18 and 19 show 
well the influence of purgation in preventing the full action of 
the drug upon the urea and uric acid. . 

3rd. An increase in the uric acid is also indicated in Experi- 
ment XVII. concomitantly with, andin Experiment XIX. follow- 
ing, the administration of the drug. 


Summary of Results. 
1. Salicylate of Soda—(A) In man. In dose of 0106 grms. 
per kilo, salicylate of soda causes no change in the amount of 
water passed, a slight increase in the urea and a very marked 


RELATIONSHIP OF UREA FORMATION TO BILE SECRETION. 305 


diminution in the uric acid excreted. This last change is by 
far the most manifest; in one experiment the diminution was 
as great as 64 per cent. 

(B) Jn dogs. In doses of from 0°45 grm. to 0°6 grm. per 
kilo, salicylate of soda caused a marked diminution in the water 
passed, a rise in the urea, and a very great diminution in the 
uric acid excreted. 

2. Benzoate of Soda, in doses of from 0°51 to 0°58 grm. per 
kilo, causes little or no change in the amount of water. The 
urea is greatly increased and the uric acid is diminished, 
though not so markedly as with salicylate of soda. 

3. Colchicum, in doses of 0:02 to 0:037 grm. per kilo of the 
acetic extract (B.P.) causes a very marked increase in the urea 
and uric acid. When the doses are small the water is also 
increased, but with large doses the water secreted may actually 
fall, while neither the urea nor the uric acid are so markedly 
increased as with smaller doses. 

4. Perchloride of Mercury, in doses of from 0:0015 to 00075 
grm. per kilo, causes an increase in the excretion of water, urea, 
and uric acid. 

5. Huonymin, in doses of from 0:04 to 0°10 grm. per kilo, 
causes a slight increase in the water excreted, and a very marked 
increase in the urea and uric acid. In larger doses, 0:16 grms. 
per kilo, it causes purging, with no diminution in the water 
passed by the kidneys, but without the marked rise in the urea 
excreted. 


We thus see that in dogs, ina condition of nitrogenous balance, 
stimulation of the flow of bile by means of these drugs is 
accompanied by an increased urea production. 

That an increased production and not merely an increased 
excretion of urea occurred is clearly shown by the fact that, after 
the administration of the dtug was stopped, the amount of urea 
merely returned.to the normal and did not manifest a fall 
corresponding to the initial rise. 

I would therefore conclude that the formation of urea in the 
liver bears a very direct relationship to the secretion of bile 
by that organ. On the nature of this relationship I have not 


touched in the present paper, but at an early date I hope 
VOL. XX. U 


306 RELATIONSHIP OF UREA FORMATION TO BILE SECRETION. 


to give the results of a series of experiments upon this sub- 
ject. 


considerable interest, especially of those which, dealing’ with the 
influence of salicylates and benzoates upon the uric acid 
excretion, afford a key to their mode of action in gout. No less 
interesting is the demonstration of the fact that colehicum 
increases and does not diminish the production of uric acid. 

In conclusion, I have to tender my thanks to Professor 
Rutherford, who, by suggesting to me as a subject for research 
the influence of the action of the hepatic stimulants on the 
composition of the urine, induced me to undertake the present 
series of experiments. Ihave also to thank him for encour- 
agement and advice during the prosecution of the work in the 
Physiological Laboratory of the University of Edinburgh, 


To the physician the results of these observations must have 


EEO OOOO 


RECENT HISTOLOGICAL METHODS. By Wu114m 
Hunter, M.B. (Edin.), M.R.C.S., Late Assistant to the 
Professor of Physiology, Edinburgh University ; former 
President of the Royal Medical Society, Edinburgh. 


WHILE engaged recently in working in the Pathological Institute 
in Leipsic, several methods employed in histology for hardening, 
embedding, cutting, and staining tissues, there in use, com- 
mended themselves so strongly for adoption, on account of their 
simplicity and effectiveness, that a short consideration of them 
at the present time may not be altogether without interest, 
perhaps value, to the many workers engaged in the same field 
of labour at home. 

Owing to the cheapness of alcohol in Germany, most of the 
hardening of tissues is done by means of this agent; a method, 
however, even if it were always the better one, which can only 
have a very limited application with us; but under this heading, 
I would desire, in passing, to refer to a rapid method of hardening 
in Miiller’s fluid, first recommended by Weigert, which, so far as 
my experience goes, is worthy of a more extended trial. 
Weigert. showed that, if the Miiller’s fluid be kept at a tempera- 
ture of about 30° to 40° C., the hardening process is considerably 
accelerated, the tissues being ready for use in the course of ten 
days or a fortnight, instead of six weeks. This method is 
specially applicable to nervous tissues, ey., brain or spinal cord, 
and, from what I have seen of it, I can strongly recommend it 
as both speedy and effective. The jar containing the tissues is 
placed in a brood-oven (hatching-oven), maintained at the above 
temperature for the above length of time, the fluid being changed 
on the second, fourth, and seventh days, if necessary also on the 
tenthday. Another method of hardening, I would merely note in 
passing, is one which is made great use of by Gaule, of the 
Physiological Institute, Leipsic, viz., placing the fresh tissue for 
twenty minutes or half an hour in a saturated solution of 
corrosive sublimate, and then hardening as usual in alcohol. 
The living tissues are in this way, as it were, fixed, and the 
method is specially suitable in carrying out minute histological 
observations. 


308 MR W. HUNTER. 


The chief purpose of this paper, however, is to direct attention 
to the method of embedding in celloidin. Since its introduction, | 
some two years ago now, into use in histology as an embed- 
ding agent, this substance has come to be very extensively 
employed, recommending itself, as it does, not only on account 
of its firmness, pliability, and its transparency when in thin 
sections, but also its cleanliness and the comparative ease with 
which it can be manipulated. It is obtainable either in the form 
of cakes, or as a fluid dissolved in ether. It is readily soluble 
in ether, slightly also in absolute alcohol. Although in mass it 
presents an opalescent appearance, in thin sections, clarified in 
the ordinary way by origanum oil or xylol, it is quite trans- 
parent, sometimes scarcely visible; and, at the same time that 
it firmly supports the tissue which is embedded in it, its 
presence in no way interferes with any of the necessary 
manipulations of the tissue, e.g., staining, &c. It thus combines 
in itself several very striking advantages. 

The method of embedding tissues in celloidin is the following:— 
The tissue, after having been hardened in alcohol, or, at least, 
immersed a sufficient length of time in alcohol to free it from 
water, is placed for twenty-four hours in a mixture of equal 
parts of alcohol and ether. If the tissue be of any size, it must 
be left in this mixture for twenty-four hours longer. It is then 
transferred into a thin solution of celloidin, made by dissolving 
a small piece of celloidin in an excess of ether, or better still in 
a mixture of equal parts of alcohol and ether, in which it is 
allowed to lie for twenty-four hours, after which it is thrown 
into a thicker solution of celloidin. In the latter it may be left 
for twenty-four hours, or even longer if-necessary, the time vary- 
ing with the size and thickness of the tissue. Asa general rule, 
three days suffice to complete the process ; twenty-four hours in 
alcohol and ether, twenty-four hours in thin celloidin, and 
twenty-four hours in the thicker celloidin. At the end of this 
time, the tissue has become thoroughly permeated with the 
celloidin, and our subsequent proceedings are directed towards 
enclosing it in a small solid block of this substance. This 
naturally can be effected by simply withdrawing the stopper 
from the bottle, and allowing the ether to evaporate till the 
whole of the celloidin solution has become solid, and we can 


o 
RECENT HISTOLOGICAL METHODS. 309 


then cut our tissue out of this mass in a small square block. 
But a more convenient way, and one which involves Jess waste 
of ether, is to pour some of the thicker celloidin solution into a 
small cardboard box,—the lid of an ordinary cover-glass box 
serves the purpose admirably, place the tissue in this, and cover 
over with more celloidin. 

It is now allowed to stand exposed to the air for a few minutes 
till the celloidin becomes somewhat firm, when it can be thrown 
into a mixture of alcohol and water, equal parts, or of the strength 
of 60 per cent. alcohol. When the celloidin has become quite 
firm round the tissue, the block can be cut out or turned out of 
the box, and for further use can be kept in alcohol of a similar 
strength. The rationale of the above proceedings will be at 
once apparent. It is necessary to have the tissue free from 
water, and, at the same time soaked in ether, before it is placed 
in the celloidin solution; hence the preliminary proceeding of 
soaking the tissue previously in alcohol and ether. Then, by 
placing it first in a thinner solution and afterwards in a thicker, 
we ensure that the celloidin reaches all parts of the tissue. a 
matter of importance when the tissue is delicate and all its 
parts require support. It is not only, however, for delicate 
tissues that this method of embedding recommends itself. 
Sometimes the tissue to be cut is so small or so thin that it 
becomes a matter of the greatest possible advantage to embed 
it in celloidin, by means of which each section is surrounded 
by a zone of celloidin which helps greatly in the after mani- 
pulation required. 

In cutting the tissues so prepared, the so-called “ dry ” method 
of cutting sections must be employed, if we wish to obtain the 
full advantages of this method of embedding. Acquaintance with 
this method—comparatively speaking so seldom employed in 
this ccuntry—so impresses one with the advantages which in 
many cases it offers over the ordinary method by freezing 
so much in use at home, that one cannot but feel a certain 
sense of astonishment at the tenacity with which we in this 
country have retained our old methods long after they have 
been rejected by our friends abroad. The instrument which has 
been recently turned out by the Cambridge Scientific Company 
will, therefore, supply a want long felt by histologists in this 


810 MR W. HUNTER. 


country. In Germany, the microtomes most in use are those 


made by Schanze, mechaniker, of the Pathological Institute in _ 


Leipzic, and by Jung, mechaniker, in Heidelberg. The latter 
often goes by the name of the “ Heidelberg” instrument or 
the “Thoma” microtome. The principle of these instruments 
is sufficiently well known as to preclude any necessity on my 


part for a detailed description of their working. In boto the | 


_ tissue is made fast within a clamp, which moves, in the ease of 
the Heidelberg instrument, along a finely inclined plane, in the 
Schanze microtome up a vertical plane, the movement in these 
planes being effected by a fine screw adjustment with millimetre 
scale, so that the thickness of each section can be accurately 
judged of and regulated. The knife, fixed on a heavy piece of 
steel, moves smocthly along a horizontal plane. In the 
Heidelberg instrument the screw adjustment is particularly 
fine, enabling one to cut sections of extreme thinness. In other 
respects, it is the instrument most suitable for cutting embryos 
an apparatus existing in it by means of which almost auto- 
matically all the sections can be made exactly of the same 
thickness. On the other hand, for general use the Schanze 
microtome will, in my opinion, be found the preferable one, 
especially if large sections be required, as of pons, medulla, &c., 
and it has the advantage of having a freezing apparatus in 
connection with it, worked by means of ether.! 

The tissue to be cut must as usual be well hardened. If it be 
of sufficiently firm consistence, ¢.g., liver or kidney, sections may 
readily be obtained from it by simply fixing it in the clamp, 
wedged in between two pieces of hard waxy liver to give support. 
This is, however, at best but a rough method, and is not 
applicable to tissues of any delicacy. Much the better plan is, 
first of all, to fix the tissue to the top of a cork by means of a 
drop of gum; on throwing the cork into spirit, the gum soon 
becomes perfectly hard, and thus the tissue is securely held. In 
doing this, it is convenient to stick a small leaden pin into the 
under surface of the cork, the weight of which sinks the cork, 
and at the same time keeps the tissue uppermost. For cutting, 
then, the cork is made fast within the clamp, with the tissue 


1 The price of these instruments varies from £4 to £10, according to size and 
completeness, 


4 


Te MS ee ee eee 


°° 


RECENT HISTOLOGICAL METHODS. 311 


securely attached to its uppersurface. The surface of the tissue, 
as it is cut, is kept moist with spirit, this being applied by 
means of a camel’s-hair brush, and the blade of the knife must 
also be kept wet; both objects being attainable by simply 
wetting the upper surface of the knife from time to time, the 
fluid passing from this on to the tissue each time the knife 
crosses it. The sections are transferred from the knife at once 
into methylated spirit, and after the desired number have been 
made, the cork with the tissue attached is thrown back into 
spirit till again required. No waste of tissue is involved by 
this method, a matter of no little importance, when,as occasionally 
happens, only a small piece is at our disposal ; and its convenience 
in private working, where only a few sections at a time are 
required, is extremely great. The result stands in marked 
contrast with that obtained by the freezing method, where, each 
time that a few sections are required, the tissue must be steeped 
in water for a period of twelve to twenty-four hours to free it 
from spirit, and where, as a rule, the tissue must be cut 
completely at the time, as its subsequent use after being frozen 
is generally out of the question. 

The tissue embedded in celloidin is also made fast to cork, 
but with celloidin instead of gum. This is done by placing it 
in some fluid celloidin on the top of the cork, and then pouring 
more celloidin over it. It is then allowed to stand exposed to 
the air till the celloidin solidifies, after which it is thrown back 
into a mixture of alchohol and water of the strength before 
mentioned. As the central portion of the celloidin remains 
longest fluid, air bubbles from the cork sometimes rise into it, 
and may interfere with the firmness of attachment. To prevent 
this occurrence it is advisable to have the corks previously 
steeped in alcohol and water. Sections can now be made of the 
tissue in the same way as before described. 

It now remains to say something of the method of handling 
the sections so obtained. The celloidin cuts very easily and 
smoothly. For staining, the sections can be treated in the 
ordinary way. Each section of tissue is surrounded by a 
somewhat stiffish zone of celloidin, which renders the after 
manipulation of delicate tissues very easy. After being stained, 
the sections are washed out first in spirit, or water if required, 


312 MR W. HUNTER, 


then transferred to alcohol. As previously mentioned, celloidin 
dissolves slowly in absolute alcohol, but for purposes of dehydra- 


tion this action is too slow to be taken into consideration. They ~ 


are then clarified—not in oi of cloves, which rapidly dissolves 
the celloidin but in origanum oil or xylol. The former is to be 
preferred ; its odour is more pleasant than that of xylol, but 
more important is the fact that xylol is liable to cause the 
tissues to shrink up into folds which are difficult of removal. 
Origanum oil has no such effect. The celloidin now becomes 
perfectly transparent, almost invisible, and the section is mounted 
in Canada balsam. We must be very careful to have the 
section free from water before placing it in origanum oil, as the 
slightest trace of water makes the celloidin opaque, and some 
time is required in the origanum oil for this opacity to clear 
up. 

I have sometimes, however, made use of clove oil for clarifying 
the tissues; under circumstances, viz., where the tissue has been 
embedded in celloidin to keep different portions of it together, 
rather than to give support to the individual tissues. In this 
way one has been enabled to keep the section intact during all 
the manipulations, and, then, at the last stage, after the section 
has been placed on the slide, by clarifying with oil of cloves, 
to remove at once all the celloidin and leave the tissue quite 
natural. In certain cases this method is convenient. 

Such is the method of embedding in celloidin. After 
considerable experience now in its use, I can strongly 
recommend it as a most valuable adjunct to our resources in 
Histology. At the same time, it is far from my intention to 
advocate any one method of embedding or cutting to the 
exclusion of all others. No one can be blind to the advantages 
which the method by freezing offers in most cases. Where 
large numbers of sections have to be made, and where the 
amount of material at our disposal is great, the method of 
freezing will always recommend itself for its simplicity, its 
cheapness, and the rapidity with which sections can be made. 
Hence, for class-teaching purposes this method is by far the best. 
But where an exact study is to be made of some delicate organ 
or tissue, or where it is desirable to obtain sections of the whole 
tissue, the method of embedding in celloidin and the use of the 


a a ee 


RECENT HISTOLOGICAL METHODS. 313 


“dry method” of cutting sections, are greatly to be preferred, 
the advantages which they offer being readily appreciable on 
even a brief acquaintance. 

Great as these advantages are, in suitable cases, they are even 
surpassed by those offered by another method of embedding, 
with which we have been long acquainted, viz. embedding in 
parafin. Owing to the general use of the method of freezing 
for cutting sections, and its inapplicability to the cutting of 
tissues embedded in paraffin, this latter method has hitherte 
not been so extensively adopted as it really deserves to be, for 
it is certain that by no method yet known to us can such delicate 
sections of tissues be obtained as after embedding in parafiin, a 
fact evidently of the greatest importance to histologists, but 
whose importance has hitherto not been fully recognised. To 
be successful, however, in embedding in paraffin, a definite 
procedure must be followed, the object of which, briefly stated, 
is to free the piece of tissue from all traces of water before 
placing it in the paraffin itself. The following method will be 
found to give very good results :—To effect this dehydration, the 
tissue must lie for a certain time in absolute alcohol, from which 
it is transferred to clove oil, where it is allowed to remain till it 
becomes saturated with the oil, a stage generally indicated by 
the sinking of the tissue in the oil. To remove this oil, and at 
the same time to have the tissue soaked in some fluid in which 
paraffin is soluble, the tissue is next placed for 4-5 hrs. (a 
much shorter time will suffice if small pieces of tissue are to be 
embedded, viz., 4-1 hr. in xylol or turpentine) in xylol, at 
the end of which time it is ready to be placed in paraffin. But 
here, as with celloidin, we first use a weak solution, viz., equal 
parts of xylol and paraffin, maintained at the melting point over 
a water bath; in this the tissue is allowed to lie for 3 hour, the 
dish being covered over during this time to prevent the evapora- 
tion of the xylol. It is then transferred to pure paraffin, 
maintained ina similar way at the melting point, where it is kept 
for 1-14 hrs., according to the size of the tissue, the dish now 
being left uncovered to allow all the xylol to evaporate. The 
tissue is then placed in a small paper boat, some of the fluid 
paraffin is poured over it, and the whole immediately floated on 
water to allow of rapid cooling. On cutting into the solidified 


314 MR W. HUNTER. 


paraffin, the tissue will be found completely incorporated with it, 
and of a dark translucent colour, and in this encasing it can be 
preserved an indefinite length of time. 

For purposes of cutting, the paraffin is made fast to a cork 
with the aid of a gentle heat, the surface of the cork being 
previously dipped in fluid paraffin, and sections are obtainable 
in the way already described. Sections of extreme delicacy can 
in this manner be obtained ; if desired, not thicker than a single 
layer of celis, a result, when obtainable, which fully compensates 
for any labour expended in suitably preparing the tissue for 
section. Each section as it is made tends to roll up, an 
occurrence to be avoided, since otherwise the section is as good 
as lost. To prevent this, it is only necessary to touch the edge 
of the section with a fine brush immediately the knife enters 
the paraffin ; the section then retains the “bent” thus given to 
it. If sections in series have to be obtained, they can be kept 
attached to each other in lines, like the segments of a tapeworm ; 
hence this method of embedding and cutting is invaluable in 
embryology. In such cases, the rows of sections are transferred 
directly to a large slide capable of holding 20, 50, or even more 
sections, and are permanently fixed on the slide by gently 
pressing with the finger the sections against the slide, or merely 
touching them with alcohol. The sections are thoroughly 
clarified, at the same time that the paraffin is removed from 
them, by throwing the slide over night into xylol, after which 
they are covered with Canada balsam, and a large cover-glass is 
placed over the whole series. In this way, in the course of a 
very short time, we can obtain the 2000 or 3000 sections of an 
entire embryo in complete linear series, without trouble and 
without any risk to the sections, a result obtainable in no other 
way yet known to us. The great advantage of the method is, 
that all the necessary manipulation is carried ont with the tissue 


as a whoie, so that the process is in reality an extremely simple 


one, and any trouble which it involves is amply compensated for 
by the ease of manipulation of the sections afterwards. Oil of 
turpentine can be used instead of xylol in embedding the tissue, 
as also in removing the paraffin afterwards, but the latter, when 
obtainable, is in my opinion to be preferred, as its solvent action 
on paraffin is almost instantaneous. 


en 


(oe . ol ee Eh) es +e ae * are 


- i tn: et 


* 
RECENT HISTOLOGICAL METHODS. 315 


As to the staining of tissues embedded in paraffin, two plans 
may be followed. In embryology, our procedure is much simpli- 
fied by staining the embryo entire before it is embedded. An 
extremely good staining agent for this purpose is alum carmine. 
This is made by adding 1 gramme pure carmine to 100 cc. of a 
warm 5 per cent. solution of alum, then boiling for twenty minutes, 
and filtering the solution after cooling. This isa very pretty stain- 
ing agent. It stains quickly, but has the great advantage of not 
readily overstaining. Sections can be left in it from ten minutes 
up to twenty-four hours without becoming overstained. It is 
generally convenient to leave the sections in the fluid over night, 
but if necessary ten to twenty minutes will usually suffice. An 
embryo must be left in for twelve to twenty-four hours. In the 
second place, it is very clean to work with; the tissue seems not 
to take up any excess of the stain, and hence the washing out 
in water can be completed in a few minutes, in which respect it 
stands in marked contrast with gentian violet, which stains very 
deeply. If an embryo has been stained in alum carmine, it is 
necessary to allow it to lie in water for about twelve hoars, 
changing the water two or three times daring this period. The 
resulting staining is limited for the most part to the nuclei, the 
colour partaking more of the purplish tint of hematoxylin than 
the reddish of carmine. It is necessary to have the embryo free 
from alcohol before placing it in the alum carmine, but, after it 
has been stained and washed in water, it can be placed in 
aleohol and embedded in the usual way in paraffin. The sections, 
therefore, on being cut are ready at once for mounting without 
further manipulation, and this is dene in the way already 
described. 

If the tissue cannot be stained en masse, the second of the two 
plans must be adopted. The sections, as made, are either fixed 
directly on the slide in the way already mentioned, or they 
are placed at once in xylol or turpentine. In the former case, 
the sections being permanently fixed to the slide are freed from 
paraffin by pouring some xylol over them. Before staining 
them, we wash out the xylol by passing a few drops of alcohol 
over them, and then, after dipping the slide in water, we 
stain the sections by placing a few drops of the staining fluid on 
the slide and allowing it to remain for a sufficient length of time. 


316 RECENT HISTOLOGICAL METHODS. 


The slide is then dipped in water, the sections dehydrated in 
alcohol, clarified in oil of cloves or xylol, and finally mounted in 
Canada balsam. The ease with which all this, apparently com- 
plicated, procedure can be carried out is very great. It is to be 
remembered that our sections are probably extremely thin, con- 
sisting perhaps of a single layer of cells; but they are perman- 
ently fixed to the slide, a number of them being on one slide, 
and hence they run no risk of being torn during all the subse- 
quent procedure, and we are dealing, not with one, but with a 
number of sections at the same time. 

If the sections have been placed in xylol or turpentine, and 
completely freed from all remains of paraffin, they ean be kept 
in alcohol, and used in the ordinary way for purposes of staining, 
&e. 

In conclusion, if one were to institute a comparison between 
the relative merits of these two embedding agents, celloidin and 
paraffin, one could only remark, that for general parposes, the 
celloidin will be found more generally useful than the paraffin, 
and especially in the study of the spinal cord, medulla and pons, 
and nervous tissues generally; but that for fine histological 
observation, as well as for embryological purposes, the method 
of embedding in paraflin is by far our best in histology, and 
gives results which can in no way be equalled by any other 
method yet known to us. 


THE SACRAL INDEX IN VARIOUS RACES OF MAN- 
KIND. By Professor WILLIAM TurNER, M.B., F.RS. 


IN my paper on “The Index of the Pelvic Brim as a basis of 
Classification,” published in the October number of this Jowrnal 
(1885), I briefly referred to variations in the relative length and 
breadth of the sacrum in different races of men, and pointed out 
that in some races the length exceeded the breadth, and that in 
others an opposite relation prevailed. These differences may be 
expressed numerically by computing a sacral index by multiplying 
the breadth of the sacrum by 100 and dividing by the length. 
When the sacral index is above 100 the breadth of the bone is 
greater than its length; when the index is below 100 the sacrum 
is longer than broad. The following descriptive terms may 
conveniently express these differences in the relative length and 
breadth of the sacrum. As the Greek word ‘vepor is the equiva- 
lent of the Latin sacrum ; the term dolichohieric would signify a 
sacrum in which the length exceeded the breadth, whilst platy- 
hieric would signify a sacrum in which the breadth exceeded the 
length. 

In considering the modifications in the sacral index, as in the 
index of the pelvic brim, it is important to bear in mind that sex 
modifies the relative proportions, and that in women the sacrum 
as a rule is broader in proportion to its length than in men. 

In working out the results at which I have arrived, I have 
measured a number of original skeletons, a few of which were 
brought home by H.M.S. Challenger, but the greater number of 
which are in the Anatomical Museum of the University of 
Edinburgh. The detailed measurements of these skeletons are 
given in the Tables in Part II. of my Report on the Human 
Skeletons now in type for the Challenger Reports. I have also 
examined the literature of the subject, so far as I have had 
access to it, and have analysed the observations on the length and 
breadth of the sacrum recorded by previous observers. 

Amongst Europeans M. Verneau has given, in sixty-three men, the 


mean length of the sacrum as 105 mm. and the mean breadth at the 
base as 118 mm., and in thirty-five women the mean length as 101 


318 PROFESSOR TURNER. 


mm. and the mean breadth as 116 mm. If an index be computed 
from these figures the males will be found to possess a sacral index of 
112-4 and the females one of 114°8. From Gértz’s measurements of the 
length and breadth of the sacrum in European women I have caleu- 
lated an index of 118°9. Dr Garson gives 101 mm. as the mean 
length of the sacrum in fourteen European women, and the mean 
breadth as 118-3, which yield an index of 116'8, slightly higher than 
that furnished by M. Verneau’s measurements, but not so high as those 
of Gértz. But from Carl Martin’s measurements of sixteen pelves, 
presumably German, the sacral length was 100 mm., the breadth 105 
mm., and the index therefore only 105. In Europeans, therefore, both 
males and females, the sacrum had a decidedly greater diameter at its 
base than in its long axis, or, in other words, it was platyhieric. 

In the Australians, again, an opposite relation prevails. In only 
one of the six adult males measured in Table I. was the breadth of 
the sacrum at the base greater than its length, in two these diameters 
were equal, and in the remaining three the length exceeded the 
breadth. The mean sacral index, therefore, of the males was only 98, 
and in the single adult female this index was only 101. Ina male 
measured by Keferstein the index was 88, in one by Barnard Davis 
90, and in five males measured by Spengel it was 111, 106, 97, 89, 
and 111 respectively. In the single male Australian measured by 
Verneau the sacral breadth at the base is stated to be 105 mm. and the 
length only 87 mm. ; but the latter diameter is so small for an adult 
male of this race, that one is disposed to think there must be either 
some error in the table or that the sacrum could not have been 
normal: the index furnished by these measurements, 120, is therefore 
exceptionally high. The sacral index of a female measured by 
Barnard Davis was 89, and the mean index computed from the two 
females measured by Verneau was 105°9, and the mean sacral index 
of the five women measured by Garson was 114. Excluding, there- 
fore, M. Verneau’s male pelvis for the reason given above, it is clear 
that in the Australian men the breadth of the sacrum was small in 
relation to its length, so that in a considerable proportion the index 
did not exceed 100, and the mean of the thirteen males was 98°5, 7.e., 
they were dolichohieric. In the women, again, the sacrum was 
relatively broader than long, though it did not attain the proportions 
reached in the European, for if we take the mean of the nine female 
pelves measured by B. Davis, Verneau, Garson, and myself, the sacral 
index was only 102°5. 

The proportions of the sacrum have been recorded in two Bushmen 
by G. Fritsch ; in the one the length was 95 mm. and the breadth 91 
mm., the sacral index being 95:8, in the other! the length was 94 mm. 
and the breadth 83 mm.,, the sacral index being 88. These two 
specimens, conjoined with the male measured in Table VL in my 
Challenger Report, give the mean sacral index of three males as 94. 


1This Bushman pelvis has had some of its characters described by Johannes 
Miiller. 


SACRAL INDEX IN RACES OF MANKIND. 319 


Verneau’s two Bushwomen had a mean sacral index 100; Girtz’s 
Bushwoman, Afandy, had a sacrum 87 mm. long and 90 mm. broad, 
the index being 103 ; whilst in an adult female recorded by G. Fritsch 
the sacral length was 97 mm. and the breadth 79 mm., the index being 
only 81. The mean of these four specimens was 94:7. There can, I 
think, be little doubt that it is the rule in the Bush race for the male 
sacrum to be longer than broad, z.¢., dolichohieric. It is not, however, 
quite so clear as to the relative proportion in the female, for although 
the mean of the four specimens is only 94°7, yet it will be observed 
that this low index is due to one of the specimens being only 81. 

G. Fritsch has also recorded the sacral length and breadth in the 
pelves of some Hottentots and Kaffirs from which indices may be 
computed. In a Hottentot woman the length of the sacrum was 95 
mm., its breadth at the base 81 mm., and its index was 85. In one 
male Korana Hottentot the sacral length was 95 mm., the breadth 90 
mm., and the index 94°7 ; in another the length was 105 mm., the 
breadth 79 mm., and the index only 75. In Wyman’s male Hottentot 
the sacral index was only 82. In all these specimens, therefore, the 
length of the sacrum exceeded the breadth, and the mean index of 
three males was 83:9. In the six male Kaffirs measured by Fritsch 
the highest sacral index was computed to be 106 and the lowest 82, 
the mean of the series being 92°8. In the single female the sacral 
length was 86 mm., the breadth 92 mm., and the index 107. In the 
male Kaffirs, therefore, the sacrum is as a rule longer than broad, and 
both in them and in the Hottentots it is dolichohieric. 

In all the Negro pelves measured in my Table III. the breadth of 
the sacrum exceeded the length, and the mean sacral index of the four 
males was 114; but though in one of the Negresses the sacrum was 
longer than broad, in the other the relation was reversed, and the 
mean index in the two specimens was only 99. In one of Spengel’s 
male Negros the sacral index was 114, in the other 97; whilst in a 
Negro from the Gaboon measured by Barnard Davis it was only 87. 
If we take, however, the series of twenty-two males described by M. 
Verneau from Guadeloupe, Mozambique, Nubia, or of unknown 
locality, the proportions are such as to give a mean sacral index of 97, 
whilst the seven females either from Guadeloupe or an unknown 
locality had a mean sacral index 105-5. In the Negress measured by 
G. Fritsch the sacral length was 93 mm., the breadth 86 mm., and 
the index 92. The mean of the twenty-nine males measured by 
Verneau, Spengel, B. Davis, and myself gave a sacral index 106, z.e., 
they were platyhieric. The mean of the ten females measured by 
Verneau, Fritsch, and myself was only 98°8 ; so far, therefore, as these 
specimens show, in the Negro race the sacrum presents the exceptional 
arrangement of being in the female not so broad in proportion to its 
length as in the male. 

In the only male adult Andaman Islander, as well as in the three 
adult females in my Table IV., the breadth of the sacrum exceeded the 
length ; the sacral index of the adult male was 114, that of a young 
male was 106; the mean index of the three adult females was 111, 


320 PROFESSOR TURNER. 


and the index of a young female was 96°5. In the series of eight 


male Andamanese measured by Professor Flower the mean sacral. 


length was 9771 mm. and the mean width was 91°3 mm., the index 
being 94, and in the series of nine females the mean length of this 
bone was 89°7 mm. and the mean breadth was 95:2 mm., the index 
being 106. In the single male specimen described by Barnard Davis 
the sacral breadth was so much less than the length that the index 
was only 77. It is clear, therefore, that the high sacral index in my 
male pelvis was an individual peculiarity, and that it is the rule in the 
male Andamanese for the length to exceed the breadth, so that the 
sacrum is dolichohieric. 

In Fritsch’s Nikobar Islander the sacral length was 101 mm. and 
the breadth was 90 mm., the index therefore being 89, or dolichohierie. 

The three male Tasmanians measured by Barnard Davis had a sacral 
index respectively 92, 86, 114, giving a mean of 97, so that the 
sacrum was dolichohieric ; but in the single female the index was 104. 

In each of my five female pelves from Oahu, Sandwich Islands, the 
sacral breadth exceeded the length, and the mean index was 113, and 
in the Tongan Islander the index was 115. In M. Verneau’s two 
Sandwich Island men the mean sacral index was also 113, and in his 
male Tongan it was 100. In his male Mangarevan the sacral index 
was 100°9, and in his Noukahivan it was 99. In Barnard Davis’s 
male Tannese this index was 100. In each of my two male New 
Zealanders the sacral index was 96. In Verneau’s Loyalty Islander 
the sacral index was 107°7, and in Barnard Davis’s male it was 103. 


In Verneau’s series of New Caledonians the mean sacral index in the — 


males was 102 and in the females 120, and in his single male New 
Guinea specimen it was 111°6. It is clear, therefore, that in the 
pelvis of these Pacific Islanders, of both sexes, whether we regard 
them as pure Melanesians or pure Polynesians, or as a inixture of the 
two races, the sacral breadth is as a rule greater than the length and 
the proportions of that bone are platyhieric. 

In my two male Guanche pelves the mean sacral index was 1085, 
and in M. Verneau’s male this index was 102. In my male Esquimaux 
the sacrum had the abnormal index 139, whilst this index in the 
female was 106. In my male Laplander the sacral index was 106, in 
the female 112; but the mean index in M. Verneau’s two male Lapps 
was only 92°6. The number of pelves in the Guanche, Esquimaux, 
and Laplanders is too small to form an average, but | think it not 
unlikely it will be found, both in the Guanche and the Esquimaux, 
that it is the rule for the sacrum to be broader than long. 

Of the natives of India the sacral index of the male Hindoo in my 
Table V. was 109. The dimensions of the sacrum in a male Hindoo 
presented to the museum by Dr. Anderson were- length 110 mm, 
breadth 122 mm., the index being 111; in the female Hindoo the 
index was 127. In all, therefore, the sacrum was platyhieric. In my 
Sikh the sacral index was 124:5, but in a male Sikh measured by 
Barnard Davis this index was only 91. In a male Bhutea, also 
measured by B. Davis, the sacral index was 93:6. 


SACRAL INDEX IN RACES OF MANKIND. 321 


The Chinese measured in my Table V. had a sacral index 98. In 
von Franque’s female specimen the length and breadth of the sacrum 
were equal at 111 mm., and the index was therefore 100. In 
Verneau’s female pelvis the sacral index was 120, but in his male, 
owing to the great length of the sacrum, 134 mm.,! this index was 
only 78. In Spengel’s male the sacrum had a still greater length, 139 
mm., and its breadth was 108 mm., the index being 77:7. Owing to 
the paucity of the specimens, and the wide diversity in the relative 
proportions of this bone in the pelves measured, it would be difficult 
to state, even approximately, the mean sacral index; but from the 
information at present before one it would seem as if in the male 
Chinese the length of the sacrum may exceed the breadth, though the 
possibility of Verneau having included the sixth vertebra in his 
measurements is not to be lost sight of. Im Verneau’s male Annamite 
also the sacral index was only 87°5. 

The female Aino in the Barnard Davis collection had a sacral index 
91; whilst in the male Aino, measured by Scheube, the length and 
breadth of this bone were equal at 115 mm. and the sacral index was 
100. 

In my male Malay the sacral index was 95. In von Franque’s 
female the sacral length was 102 mm. and the breadth 100 mm., the 
index being therefore 98. In Barnard Davis’s two male Javanese the 
sacral index was respectively 96 and 88, but in Verneau’s female, 
owing to the remarkable shortness of the sacrum, 72 mm., this index 
rose to 145. Of the twenty-six Javanese female pelves measured in 
Zaaijer’s Table I., the length of the sacrum exceeded the breadth in 
eleven specimens, it was less than the breadth in fourteen specimens, 
and in one of these the two dimensions were equal. As in the female 
pelvis it is the rule for the sacrum to be relatively broader than in the 
male, there can, I think, be little doubt that, in the Malay race, the male 
sacrum is longer than it is broad, and that this bone is dolichohieric. 

Of the North American Indians von Franque gives the length of 
the sacrum in the male Flathead at 112 mm. and the breadth at 114 
mm., the index being 101°8; whilst in the female the length was 107 
mm. and the breadth 108 mm., the index being 100°9. In Barnard 
Davis’s male Illinois Indian the sacral index was 115°8, and in 
Verneau’s female Mexican it was 108. Probably, therefore, in the 
North American aborigines the average breadth of the sacrum is more 
than the average length. 

The dimensions of the sacrum have been recorded in a few more 
specimens of South than of North American Indians. In a male 
Puelche measured by Barnard Davis the sacral index was 95°8, and in 
a male ancient Peruvian it was 92. From Verneau’s table of sacral 


1 Verneau states that in the Annamite and male and female Chinese pelves the 
sacrum consists of six pieces. The Chinese woman, notwithstanding this, has a 
relatively broad sacrum ; but if he has included the sixth piece in his measure- 
ments of the Chinese man and of the Annamite the low sacral index is accounted 
for It is not unlikely that Spengel’s specimen may have had a similar arrange- 
ment. 

VOL. XX. x 


322 PROFESSOR TURNER. 


measurements I have computed the sacral index in a male Charruan 
at 108, in a male Botocudon 117, in a male Bolivian 93, the mean of 
two male Peruvians 128, in a male Goytacazen 119; whilst the mean- 
of three female Peruvians was 121-7, and in a female Goytacazen it 
was 110°5. Of the eight males three had a sacral index below 100, 
but the others were so much above 100 that the mean sacral index of 
the series was 107°5, and the mean of the four females was 116. The 
measurements recorded by J. G. Garson, of four male Yahgans from 
Tierra del Fuego, gave a mean sacral length 103°2 mm., and a mean 
sacral breadth 112°2 mm., the mean index being 109, and in each 
specimen the index was above 100. In the South American Indians, 
therefore, I have little doubt that the mean breadth of the sacrum 
exceeds the mean length, so that the aborigines in both the north and 
south of that continent are platyhieric. 

From this analysis of my own measurements and of those of other 
anatomists it is clear that the proportion of the length to the breadth 
of the sacrum varies in different races of mankind. In some it is the 
rule to find it longer than broad, in others broader than long. 
Although additional observations are still greatly needed, especially on 
some of the races, yet it seems possible to make a provisional arrange- 
ment of the races of men into two groups, according as the sacral 
index is below or above 100, and in this Table, as in the one given in 
my previous paper, “On the Index of the Pelvic Brim,” the propor- 
tions are estimated on the measurements in the male sex, 


DoLIcCHOHIERIC, PLATYHIERIC. 

Sacral Index below 100. Sacral Index above 100, 

Australians Europeans 

Bushmen Negros 

Hottentots Melanesians 

Kaflirs Polynesians 

Andamanese Hindoos 

Tasmanians Guanclhie ? 

Chinese ? Esquimaux ? 

Aino ? North American Indians 

Malays South American Indians 


A comparison of this Table with that framed on the index of the 
pelvic brim will show that in many instances a dolichopellic brim is 
conjoined with a dolichohieric sacrum. For example, this is the case 
in the Australians, Bushmen, Kaffirs, Andamanese, Ainos, and Malays. 
Again a platypellic brim is conjoined with a platyhieric sacrum in 
the Europeans, American Indians, and probably the Guanches and the 
Esquimaux. The pelves which I have arranged in the mesatipellic, or 
intermediate division, partly belong, as regards the proportions of the 
sacrum, as in the Negros and Melanesians, to the platyhieric group, 
and partly, as in the Tasmanians, to the dolichohierie group. As the 
width of the pelvic brim is materially influenced by the breadth of 
the sacrum, it was to be expected that a platypellic pelvis would have 


a 


SACRAL INDEX IN RACES OF MANKIND. 323 


a relatively wide sacrum, and that a dolichopellic pelvis would have a 
relatively long sacrum. There are two races which, however, have an 
anomalous position in these two tables, viz., the Chinese and the Poly- 
nesians. The Chinese with a platypellic brim, it will be observed, 
are placed with pelves in which the sacrum is longer than broad. The 
position, however, which the Chinese occupy in this table can only 
be regarded as provisional, and when a larger series is examined, on 
the basis of only five sacral vertebrae being measured, it is not unlikely 
that their position in the table of sacral proportions will have to be 
altered. The position of the Polynesians in the table in the preceding 
paper is also provisional, as a wider series of observations may require 
them to be transferred to another column in that table. 

In connection with the relative dimensious of the conjugate and 
transverse diameters of the pelvic brim, and those of the length and 
breadth of the sacrum in the different races of men, a few words may 
appropriately be written on the corresponding relations in the pelvis 
in other mammals, at least in those which possess five vertebree in the 
sacrum. In the Anthropoid Apes the length of the sacrum is consider- 
ably greater than its breadth. In two orangs, which I have measured, 
the mean sacral index was 87 ; in two chimpanzees the mean was 77, 
and in a gorilla the index was 72; in an ox the sacral index was 87, and 
the sacrum in all these was markedly dolichohieric. The mean index 
of the pelvic brim in two chimpanzees was 133; and in an ox the 
pelvic index was 110. In these animals the conjugate diameter of the 
pelvic brim was materially greater than the transverse, z.e., the index 
was dolichopellic. When a human pelvis therefore is dolichopellic, 
and has also a dolichohieric sacrum, it corresponds in both characters 
with the more usual type of mammalian pelvis; and, as compared with 
the relations of parts met with in the Europeans, it possesses a degraded 
or animalised arrangement. 


THE BLOOD-FORMING ORGANS AND BLOOD-FORM A- 
TION: AN EXPERIMENTAL RESEARCH. By JOHN LOCKHART 
Gipson, M.D., formerly Senior Demonstrator of Physiology, 
University of Edinburgh. 


(Continued from p. 113, vol. xx.) 
Part II1.—ONn THE BLOOD-FORMING ORGANS. 


HAVING reviewed the different elements found in the blood, 
I now pass on to ascertain, if possible, the function of the so- 
called blood-forming organs. 

In considering blood-formation, I restrict myself entirely to 
blood-formation in extra-uterine life. 

The organs called “ blood-forming” are the spleen, lymph- 
glands, bone-marrow, and thyroid gland. I have endeavoured 
experimentally to discover if all of these organs produce red 
blood-corpuscles in extra-uterine life, and which of them are 
most active. That they all, with the exception of the thyroid, 
produce white blood-corpuscles, I think nobody will doubt. 

With regard to the formation of red corpuscles in extra- 
uterine life, the greater number of recent authors (Neumann, 
Bizzozero, &c.) have given the bone-marrow the first place as a 
producer of red corpuscles in extra-uterine life; though some 
authors deny that it has any blood-forming action. A smaller 
number (Rindfleisch, &c.) have given the spleen the first place ; 
a much larger number giving it a second place, and a few 
(Neumann and Ehrlich) giving it no place at all. A still 
smaller number (Zesas and Credé) look on the spleen and 
thyroid together as the chief producers of red corpuscles ; 
others denying to the thyroid any blood-forming action. The 
lymphatic glands receive, here and there, a little irregular and 
indetinite support. 

The points which seemed to me most doubtful, because differed 
about by men generally recognised as the most competent author- 
ities on the blood and on blood-formation, were :—1s¢, how the 
red corpuscles are produced in extra-uterine life ; 2nd, whether 
the spleen has any activity as a red-corpuscle-forming gland in 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 325 


extra-uterine life; 3rd, whether the lymphatic glands have any 
activity as producers of red blood-corpuscles ; and 4¢h, whether 
the thyroid gland has any blood-forming function. 

I shall consider—/irstly, the results of excision of the human 
spleen, and what can be learnt from the recorded cases, with 
regard to its blood-forming function; secondly, the experiments 
and results of the principal observers who have excised the 
spleen from animals and watched the effect; and thirdly, my 
own experiments on the function of the spleen, as also on the 
function of the lymphatic glands. The function of the bone- 
marrow will be considered together with the function of the 
spleen. 

The consideration of the thyroid gland will be more conven- 
iently left until the consideration of the spleen is completed, as I 
have been unable to find any grounds for supporting a blood- 
forming action of the thyroid. 

It has very recently been said that we do not know much 
more about the spleen than Aristotle did when he said: “The 
spleen is not an organ which is indispensably necessary to the 
body ;” and that we may still call it, as Galen did, the “organ 
full of mystery.” This, however, is hardly true. For although 
there is still but little uniformity of opinion as to the function of 
the spleen, yet much light has of late been thrown on the subject, 
more especially by the very able researches of Bizzozero. And 
I think that if it can here be shown that the spleen is an organ 
with an important action in the regeneration of the blood after 
heemorrhage, a function will be given it not unworthy even of 
an organ of its considerable size. 

As early as 1549 the spleen was successfully removed from a 
human subject;+ and previous to that date it had been ascer- 
tained that spleenless dogs not only lived after the operation, 
bat even seemed to thrive and grow fatter. 


Chronological lists of excisions of the spleen in man are given 
by Credé? and Zesas.2  Credé’s paper gives a list of thirty 


1 Fioravanti, Del tesoro della vita humana, libr. ii. cap. 8. (Fioravanti was the 
physician of the case, Zaccarelli the surgeon. ) 

2 B. Credé, ‘‘ Ueber die Exstirpation der kranken Milz beim Menschen,” Arch. 
J. klin. Chir. (v. Langenbeck’s), Bd. xxviii. (1882), pp. 401-410. 

3 Zesas, ‘‘ Ueber Exstirpation der Milz am Menschen und Thiere,” Arch. f. 
klin. Chir., Bd. xxviii. (1882), pp. 157-178. 


326 DR J. LOCKHART GIBSON. 


excisions of the diseased spleen, and also considers what has been 
learnt from the cases with regard to the blood-forming function of the 
spleen. Zesas, besides considering those cases where excision was 
performed for enlargement of the spleen, gives the results of twenty 
cases collected by him of excision of the spleen after penetrating 
wounds of the abdomen with injury to the spleen. Among these 
twenty cases Zesas could not find a single fatal result. 

Of the thirty cases quoted by Credé, sixteen were in leukemic 
patients, all of whom died during or shortly after the operation. 
From these sixteen nothing can be learnt regarding the function of 
the spleen, as the patients did not live long enough. Nor from them 
can we even conclude that excision of the spleen in leukemia is in itself 
necessarily fatal. For the patients were presumably not operated on 
till after all other remedies had been found powerless, and therefore not 
till the disease was far advanced, and their vital resistance reduced to 
a minimun. 

Of the fourteen other cases, five resulted fatally and nine were 
successful. Of the five unsuccessful cases two may be excluded. In 
one, where death occurred six hours after the operation, the ligature 
was found compressing the tail of the pancreas. The patient, who 
before the operation was in a very feeble condition, died of shock. 
In another, where death occurred two hours after the operation, 14 lbs. 
of blood was found in the abdomen. Neither of these cases can, in 
my opinion, have their fatal termination ascribed to the absence, or, in- 
deed, even to the removal, of the spleen. 

We have left, then, three fatal cases out of twelve ; which cannot be 
called a large mortality for such an operation, especially as so many 
of the operations were performed before attention was drawn to the 
immense importance of antiseptic precautions during operations on the 
abdomen. 

The information which can be gathered from these recorded cases 
with regard to the blood-forming action of the spleen is not very 
definite, but is still of value. The blood seems to have been examined 
after the operation in only five of the cases :— 

1. and 2. Two cases of Péan,! where some weeks after the operation 
there was a slight increase in the relative number of white corpuscles 
as compared with the red. 

3. Czerny records a case in which up to five weeks after the opera- 
tion there was apparently no change in the blood. The blood was not 
examined later. In this case the cervical and inguinal glands were 
painful for several weeks after the operation. 

4, Martin records a successful case where examination of the 
blood yielded negative results. 

5. By far the most carefully recorded case, however, is by Credé, in 
the above-mentioned paper. In it the changes in the blood seem to 
have been of a marked character. Two months after the operation 
(which was for a cyst of the spleen) the alteration of the blood was 
at its height, and the proportion of white to red corpuscles was 


1Péan, Tuimeurs de Vabdomen. 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 327 


1: 3o0r4. “There were also a great number of small red corpuscles 
(Mikrozyten), some of them nucleated.” The blood gradually 
changed back to normal again, and was quite normal 44} months after 
the operation. Four weeks after the operation a painful swelling of 
the whole thyroid gland appeared, which lasted for four months, dis- 
appearing soon after the blood resumed its normal condition. There 
was “neither swelling of the lymph-glands nor pain in the bone- 
marrow.” The patient was a man of 44 years, and he recovered his 
health entirely, so that he was able to continue his work, that of a 
mason. 

I was fortunate enough to hear Dr Credé’s son speak of the case at 
the Berlin Surgical Congress this year. The man was quite well, 
three years after the operation; and the white corpuscles had sunk 
below the normal number. Credé mentions particularly that the 
marked changes in the blood of his patient were not seen for some 
weeks after the operation, and that he did not find a return to a 
normal condition until four months after it; and says that the four 
other cases were not examined after they left their respective hosp1- 
tals, four or five weeks after the operation, whereas he watched 
the blood of his patiert for ten months. He thinks it possible that 
in the blood of the other patients, more especially of those of Péan, 
there may have been changes which were not marked until after the 
patients left the hospital, and therefore until observations on the 
blood had ceased to be taken. From these five cases, Credé gathers 
that the spleen has an important action in turning white into red 
corpuscles; that when the spleen is removed the retrogressive 
metamorphosis of the red corpuscles diminishes; and that the 
white corpuscles increase in number until another organ takes on 
itself the function of turning them into red ones. And this other 
organ may, he says, “possibly be the thyroid gland.” He believes 
that the lymph-glands carry on their function as before, and sees no 
reason why they should increase in size. ‘‘ The bone-marrow,” he says, 
“which in normal conditions produces only a very moderate number 
of small red corpuscles, appears to increase its function after the spleen 
is removed.” He gives his conclusions in the following tabulated form, 
and supports them by his own case and the two cases of Péan, and 
also by Zesas’ observations on animals, to be afterwards mentioned :— 

1. “The adult human subject bears the removal of the spleen 
without injurious effect. 

2. “The removal of the spleen causes changes in the condition of 
the blood, which afterwards pass off. 

3. “These changes are overcome by increased activity of the thyroid 
gland and the bone-marrow. 

4, “The spleen changes white corpuscles into red.” 


I have given these conclusions of Credé’s before going further, 
because they are the results of observations on the human 
subject, which of course are of much more value than similar 
observations on animals. Credé’s own case is, however, the 


328 DR J. LOCKHART GIBSON. 


only one which has been at all satisfactorily observed; and if 
we wish to know anything more definite with regard to the 
function of the spleen, we must pass to experiments on animals. 
From the time when Kolliker? found nucleated red cells in 
the spleen during intra-uterine life and after a year of 
extra-uterine life, and therefore stated that the spleen is a 
blood-forming organ, blood-formation was pretty generally 
believed to be at least one of its functions, until Neumann? 
in 1869 cast diseredit on this belief by saying that the blood- 
forming function of the spleen could not be demonstrated. 


In his paper Neumann supports and extends the observations he 
made on the bone-marrow in 1868°, when he found that the red 
marrow contained nucleated red corpuscles at all periods of life. He 
considers that the red bone-marrow is throughout the whole of extra- 
uterine life the only producer of nucleated red corpuscles, and there- 
fore of non-nucleated red corpuscles, and that after birth the spleen 
has no blood-forming function whatever ; and he even expresses doubt 
about the spleen having a blood-forming function in the foetus. He 
says that the nucleated red corpuscles found in the spleen were just 
caught there as they were flowing through it in the circulating blood.* 

Freyer,* a pupil of Neumann, found that the blood of the aorta 
was not poorer in nucleated red corpuscles than the blood in the 
spleen. He next attempted, by producing artificial anzemia in rabbits, 
to see if more such cells could be found in the spleen ; and obtained a 
negative result. 

In 1877 some valuable observations on the hone mnie were made 
by Litten and Orth.° These observations were rather in favour of 
Neumann’s views, and at any rate gave further proof cf the important 

part the bone-marrow plays in the formation of red blood-corpuscles. 
They found that the marrow in the shafts of the long bones was fatty, 
and that only the marrow in the spongy parts of the bones was fune- 
tional in the direction of producing nucleated red corpuscles. They 
tried, by producing artificial anemia in animals, to ascertain whether 
the fatty marrow also then became functional, to meet the increased 


* Kolliker, Handbuch der Gewebelehre. [In the statement about Kdlliker on 
page 101 of this volume (page 4 of reprint), the reader will please to put 
‘*spleen” instead of “liver,” and in the footnote put “Handb. d. Gewebelehre,” 
instead of “‘ Mikrosk. Anat., Bd. ii. p. 590.” The need for this correction is the 
fault, not of Dr Gibson, but of a reviser in his absence. ] 

* Neumann, ‘‘ Ueber die Bedeutung des Knochenmarkes fur die Blutbildung,” 
Arch, d. Heilk., x. (1869), p. 68. 

8 Neumann, Centralblatt f. d. med. Wiss., 1868, No. 44. 

4 Freyer, Ueber die Betheiligung der Milz bei der Entwickelung der rothen 
Blutkirperchen, Konigsberg, 1872. 

5 Litten and Orth, Berliner klinische Wochenschrift, 1877, No. 51. 


4 ° 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 329 


requirements ; and they found that the fatty marrow in the shafts of 
the long bones became red marrow, and was crowded with nucleated 
red corpuscles. 

Professor Bizzozero and Dr G. Salvioli of Turin then set themselves 
the question :—Does the spleen, which is at any rate said to produce 
nucleated red corpuscles at the beginning of extra-uterine life, renew its 
former function, and, like the fatty marrow, again produce nucleated 
red corpuscles, when artificial anemia is produced, and when 
therefore the reserve blood-forming capabilities of an animal are called 
into play? After elaborate researches, they found themselves able 
to give an affirmative answer to this question.! 

Their first step was to examine the spleens of healthy animals (they 
used dogs, guinea-pigs, and rabbits), in order to see whether they 
already contained nucleated red corpuscles. 

They found, like Kélliker, nucleated red corpuscles in the spleens 
of new-born dogs ; and even found them in one dog which was cer- 
tainly more than a year old. In the spleens of adult dogs, such 
corpuscles were, as a rule, not to be found; although single examples 
could be found by diligent search. 

In young guinea-pigs, numerous nucleated red corpuscles were 
found in the spleen. In adults, few were to be found; but they 
were very seldom entirely wanting. 

In rabbits, they found these cells only very seldom after birth ; 
while in the later period of youth and in adults they never found 
them. They therefore considered that the spleens of rabbits were 
peculiar after birth, in not containing any of these nucleated red cells. 

Their second step was to produce artificial anemia in animals and 
notice the effect on the spleen. 

For the purpose of estimating the amount of anemia, they employed 
the chromocytometer described by Bizzozero.2. They obtained positive 
results only in dogs and guinea-pigs. Finding their results from 
rabbits negative, they decided to discontinue their observations on 
these animals. 

Their first series of experiments was on guinea-pigs. They 
produced artificial anemia in five guinea-pigs, by removing 1 to 2 
per cent. (once 3 per cent.) of the body-weight of blood at a time. 
They never drew blood more than twice from the same animal ; and the 
second blood-letting was always after an interval of three days from 
the first. Three days after the second blood-letting, 7.¢., six days after 
the first, they killed the animal, and examined the spleen. In all 
cases the hemoglobin of the blood on the day of death was between 
50 and 60 per cent. of its original amount. 

In all cases the spleen was found swollen and rich in blood, some- 
times very much so, and always soft. It always contained “ very 
numerous ” nucleated red corpuscles. 


1 Bizzozero and Salvioli, ‘‘ Beitrage zur Himatologie,” Moleschott’s Untersuch- 
ungen, Bd. xii. (1881). 

* Bizzozero, ‘‘11 cromocitometro, nuovo instrumento per dosare l’emoglobina 
del sangue,” Accademia delle scienze de Torino, 1879. 


330 DR J. LOCKHART GIBSON. 


The marrow of the ribs and long bones also contained very numerous 
nucleated red corpuscles, sometimes more than the spleen, 


Having obtained such distinct results with guinea-pigs, they next 


treated a series of ten dogs in exactly the same way. They sometimes 
drew blood as often as four times from a dog, the amount of blood 
drawn at each time varying from 1} to 3} per cent. of the body-weight. 
About three days after the last blood- letting the animal was killed. 
The percentage of hemoglobin contained in the blood of the different 
animals on the day of death varied between 45 and 55 per cent. of 
its original amount. 

They constantly found the marrow of the ribs very rich in nucleated 
red corpuscles, the marrow of the long bones being less rich in them. 

In only one case (where there was only one blood-letting, and 
where the heemoglobin was reduced only to 62 per cent.) did they “fail to 
find nucleated red corpuscles in the spleen. In two cases they found 
a few, in three cases moderate numbers, and in four cases a great 
number, The spleen as a rule was slightly swollen; and in those 
cases where it contained numerous nucleated red corpuscles it was 
very much swollen, and very soft. Asa rule the marrow of the ribs 
contained more nucleated red corpuscles than the spleen: only in 
one case is it remarked that it contained a similar number, viz., in a 
dog which had been rendered more anemic than the others, the 
hemoglobin having been reduced to 25 per cent. They describe the 
naked- -eye appearances of the spleen as very characteristic: in fact, as 
so much so, that by simply looking at it they could say, “ there will 
be nucleated red cells here.” They describe it as “rose coloured and 
very much swollen,” and as contrasting very distinctly with the spleens 
of normal dogs, which are generally very poor in pulp. 

Their next step was to examine the blood of the splenic artery and 
the blood of the splenic vein :— 

lst, To ascertain whether the vein contained more red corpuscles 
than the artery. 2nd, To ascertain the number of white corpuscles in 
the splenic vein relatively to the number of red corpuscles, and to 
contrast this relationship with the relationship in the splenic artery. 

For this purpose they chose animals which had been rendered 
anemic by blood-letting, and whose spleens were therefore in a state 
of activity. {Im two cases they used very young dogs without 
producing anzmia, as they knew from previous observations that the 
spleens of such dogs are still active. 

The results they obtained were: that the blood in the splenic vein 
is richer in hemoglobin than the blood in the splenic artery, and that 
the relative number of white corpuscles, as compared with the red, 
is greater in the splenic vein than in the splenic artery. 

From these data, Bizzozero and Salvioli come to the conclusion 
that, as the ouly way in which the blood of the splenic vein can be 
richer in haemoglobin than the blood in the splenic artery is by its 
containing a larger number of red corpuscles, therefore the blood of 
the splenic vein contains more red corpuscles than the blood of the 
splenic artery ; and as they found the number of white corpuscles 
greater in proportion to the red corpuscles in the splenic vein than in 


THE} BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 331 


the splenic artery, they concluded that the addition of white corpuscles 
to the blood in its passage through the spleen is relatively still greater 
than the addition of red corpuscles. 

They did not count the absolute number either of red or of white 
corpuscles in a given quantity of blood, but used the old, and, accor- 
ding to them, most accurate method of ascertaining the relative 
number of red and white corpuscles, viz., that of mixing a drop of 
blood with a drop of a ‘75 per cent. solution of common salt, so 
putting on a cover-glass as to obtain only one layer of corpuscles, 
and then through an eye-piece micrometer divided into squares 
counting all the red and all the white corpuscles in a square, and so 
obtaining their relative numbers. 

They base their assumption that the hemoglobin of the blood can 
be increased only by the presence of an increased number of red 
corpuscles on the observations of Welcker, Vierordt, Heidenhain, 
Panum, and others, who demonstrated that after blood-letting the 
hemoglobin of the blood falls in percentage amount in consequence 
of the absorption of fluid plasma from the tissues; and on confirma- 
tory observations conducted by themselves, and published in Mole- 
schot?s Untersuchungen. 

By the observations last referred to, Bizzozero and Salvioli show that 
the fluid is not absorbed from the tissues at once, but that a time vary- 
ing from six to forty-eight hours is required, before the amount of fluid 
necessary to replace the®lost volume of blood has been absorbed, and 
accordingly before the hemoglobin in the blood has reached its mini- 
mum. They further show that the diminution of hemoglobin has a 
direct and constant relation to the amount of blood lost. For every 
per cent. of body-weight lost, there was a diminution of 11:14 per 
cent. of the hemoglobin of the blood. This proportion they estab- 
lished after experiments on a great number of animals. 

If these results of Bizzozero and Salvioli are compared with the 
results of J. F. Lyon,? who produced artificial anzemia in animals and 
counted the number of red corpuscles in the blood on the subsequent 
days until regeneration had occurred, it will be seen that the time when 
the number of red corpuscles suffered its greatest diminution coincides 
pretty closely with the time when Bizzozero found the blood poorest 
in hemoglobin. 

From their observations on anemic guinea-pigs and dogs, Bizzozero 
and Salvioli conclude that for these animals they have, in opposition 
to Neumann and Freyer, not only proved “the real production of colour- 
less blood-corpuscles in the spleen, but also placed beyond doubt its true 
hematopoiétic function, and the part it takes in the formation of coloured 
blood-corpuscles.” And they further, in support of their observations, 
and in opposition to those of Neumann and Freyer, bring forward the 


1 Bizzozero and Salvioli, ‘‘ Ueber die Aenderungen, welche der Hiimoglobin- 
gehalt des Blutes in Folge von Blutentziehungen erfihrt,” Moleschott’s Unter- 
suchungen, Bd. xii. 

* Lyon, ‘ Blutkérperzihlungen bei traumatischer Animie,” Virchow’s Archiv, 
Bd. lxxxiv. pp. 207-247. 


Son DR J. LOCKHART GIBSON. 


observations of Foa and Salvioli! on the embryo, which they think 


have placed beyond all doubt the fact that the spleen produces red _ 


blood-corpuscles in intra-uterine life. 

Bizzozero was, however, not satisfied with observations on dogs and 
guinea-pigs, but also made researches to ascertain the function of the 
spleen in other animals. Having found, as already stated, that the 
spleens of rabbits appeared to be inactive after birth, so far as the 
formation of red corpuscles was concerned, he turned his attention to 
birds; and a paper by Bizzozero and Torre? giving the results of 
observations on birds is published in Moleschott’s Untersuchungen. 
Their observations were chiefly on the pigeon, fowl, and finch; and 
they could find no indication of the formation of red blood-corpuscles 
in the spleens of these animals. 

These conclusions are directly contrary to those which Rindfleisch ® 
came to, after observations on birds. Rindfleisch particularly men- 
tioned birds as animals in which the spleen was “by far the most 
important producer of red blood-corpuscles.” 

Bizzozero and Torre found in birds, that after the production of 
artificial anzemia the fatty marrow became red, and actively produced 
red blood-corpuscles. They were able to trace all stages between the 
white corpuscles, which they believe are likewise formed in the bone- 
marrow, and the fully developed red corpuscles. 

These observations are in accordance with those of Theo. Korn, 
who found that the removal of the spleen was very well borne by 
birds (pigeons), and did not in any way affect regeneration of the blood 
after the production of artificial anzemia. 

Neumann considers that the experiments of Theo. Korn support 
the view he takes of the function of the spleen. But the error into 
which Neumann seems to have fallen is that, finding in rabbits that 
the spleen did not, at least in extra-uterine life, form red blood- 
corpuscles, he thence argued that the spleens of other animals, includ- 
ing man, were equally inactive. Bizzozero remarks, “if Neumann 
and Freyer had used dogs instead of rabbits, they would have come 
to the same conclusion as we did.” This is, however, not strictly 
correct, as will further on be seen. For in a later paper Neumann 
also denies the heematopoiétic function of the dog’s spleen. 

Bizzozero and Torre® also made observations on reptiles, amphibians, 
and fishes, the results of which were published in 1884, and are as 
follows :— 

1. In reptiles and tailless amphibians, the bone-marrow forms red 
blood-corpuscles, which divide and multiply in it ; but the spleen has 


 Foa and Salvioli, ‘‘Sull’ origine del globule rossi del sangue,” Archivio per le 
scienze mediche, vol. iv. p. 1. 

* Bizzozero and Torre, ‘‘ Ueber die Entstehung und Entwickelung der rothen 
Blutkorperchen bei Vigeln,” AMoleschott’s Untersuchungen, Bad. xii. 

3 Rindfleisch, Arch. f. mikrosk. Anat., Bd. xvii. pp. 21-42. 

4 Theo. Korn, Centralblatt f. d. med. Wiss., 1880, No. 41. 

° Bizzozero and Torre, Virchow’s Arch. f. Path. Anat. und Physiol., Bd. xev 
Heft 1. 


_— 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 333 


no function in the production of red blood-corpuscles, and is rather to 
be considered as a lymphatic gland. 

2. In tailed amphibians, the bone-marrow appears to lose altogether 
its blood-forming function ; and the spleen is the chief blood-forming 
organ. 

3. In fishes, the formation and regeneration of the blood-corpuscles 
takes place very slowly, and occurs chiefly in the spleen and in blood- 
forming parts of the kidney. 

Picard and Malassez,! from observations on dogs, came to the 
covelusion, that after loss of blood in spleenless animals the blood 
returns to its normal condition (as far as the number of corpuscles is 
concerned) as quickly as in animals still possessing a spleen. And 
they further found that extirpation of the spleen lessened the amount 
of hemoglobin in the blood, but that there was an increase rather than 
a decrease in the number of red corpuscles. 

Here again is seen the importance of distinguishing between the 
microcytes, which we have already considered, and the red blood- 
corpuscles. It seems to me that Picard and Malassez have counted 
the microcytes as coloured corpuscles, believing them to be young 
coloured corpuscles ; and that it is in this way they have found an 
increase rather than a decrease in the number of red corpuscles. 

Bizzozero attempted to ascertain the effect on the blood produced by 
removal of the spleen, using dogs for this purpose. He found that 
after spleen extirpation the hemoglobin of the blood diminished until 
it reached a certain degree, and that it then began gradually to rise 
again. He found, however, that it never reached its former percentage 
while the animal remained under observation. 

The experiments of Bizzozero and Salvioli to see whether after 
blood-letting in spleenless animals the blood returns to its original 
condition as quickly as in animals possessing a spleen, gave them 
results quite different from those of Picard and Malassez, inasmuch 
as they found that while the animals remained under observation the 
blood did not return to its former condition. They experimented on 
three spleenless guinea-pigs and four spleenless dogs ; but say that their 
experiments were not extensive enough to let them come to a perfectly 
definite conclusion, because in the case of one dog the blood seemed to 
return to its previous condition as quickly as in animals possessing a 
spleen. 

In 1869 Neumann? published a paper in which he showed that 
chronic diseases leading to general marasmus not only cause the 
red blood-forming marrow to extend itself very much, but also cause 
it to become very rich in nucleated red blood-corpuscles. Moreover, 
his observations have since received great support from those of Litten 
and Orth, already referred to. But, while making experiments to prove 
the active blood-forming function of the bone-marrow,? he also vame to 
the conclusion, already mentioned, that the spleen is “not a blood- 


1 Picard and Malassez, Gazette méd. de Paris, 1878, No. 15. 
? Neumann, Centralblatt f. d. med. Wiss., 1869, No. 19. 
3 Neumann, Arch. d, Heilk., xii. (1871), p. 187; xv. (1874), p. 470. 


334 DR J. LOCKHART GIBSON. 


forming organ,” at any rate “in extra-uterine life,” and that even in 
the foetus its action in this respect “is a very subordinate one.” In 


fact, he grants a blood-forming action to the spleen only in a very 


grudging manner. He says, ‘‘ in the embryo and for a short time after 
birth, nucleated red cells are found in the spleen ; and it is possible 
that some of them may arise in the spleen, as there are more of them 
in the spleen than in the heart blood.” 

These conclusions of Neumann are based on his own and Freyer’s 
researches on rabbits, already referred to; but, in the Zeitschrift fiir 
klinische Medicin' for 1881, he has, along with some observations on 
a case of anemia due to prolonged and frequent loss of blood, 
published the results of experiments on two dogs, which he considers 
further to support them. He drew large quantities of blood at a time, 


and repeated the blood-letting six and seven times in the course of 


six and five-and-a-half weeks respectively. The blood drawn from 
the first dog in the course of six weeks amounted to one-sixth of its 
body-weight, and the blood drawn from the second dog in five-and-a- 
half weeks amounted to one-eighth of its body-weight. The second 
dog was reduced to a state of such profound anemia that it died. The 
first dog was killed with cyanide of potassium; and the spleen was 
found to be very dry, and to contain “isolated nucleated coloured 
blood-corpuscles, which could only with difficulty be found amongst 
the colourless cells of the pulp.” In the second dog, being the one 
that died of anzmia, no nucleated red corpuscles were found in the 
spleen. In both dogs, the red marrow had extended itself into the 
yellow marrow, and contained a very large number of nucleated red 
corpuscles. 

The chief reason for objecting to the results of these experiments, 
as against the more numerous experiments of Bizzozero, seems to me 
to be the very high degree of anzmia produced. Under anzmia so 
great and so suddenly produced, an organ whose function had for a 
considerable time been laid aside or very little employed would very 
likely not be in sufficiently favourable circumstances for the resump- 
tion of that function. This reason, however, while applying to the 
spleen, would not in the same way apply to the bone-marrow. For at 
the very first loss of blood the bone-marrow, already in full activity, 
would have its activity enormously increased; although it is quite 
possible that its further action would be more or less paralysed by the 
profound state of aneemia ultimately produced. 

Neumann in this paper gives the history and post-mortem examina- 
tion of a very interesting case of anzemia from loss of blood in the 
human subject. The case was that of a woman, aged 38 years, who 
had suffered, at intervals, for three years, from great loss of blood from 
the uterine mucosa. She came frequently under treatment in hospital ; 
and left improved, only to return, after some months, with similar 
symptoms. She died during one of the attacks; and the examination 
of the blood-forming organs, conducted by Neumann, yielded the 
following :— 

1 Neumann, ‘‘ Ueber Blutregeneration und Biutbildung,” Zeitschrift fiir klin. 
Med., Ba. iii. Heft 3. 


OEE OEE 


— 


————— 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 335 


The spleen was neither enlarged nor succulent, but rather hard and 
dry. Liver was of normal size. Retro-peritoneal, mesenteric, and 
pelvic glands were distinguished by their flesh-red colour; parts being 
lighter, parts darker. They were not markedly swollen, and were of 
a hard consistence ; and their cut surface was dry. The bone-marrow 
of the ribs, vertebrae, and humerus was of a dark raspberry-red colour. 

Microscopic Examination.—F¥atty degeneration was very wide- 
spread. 

In the bone-marrow, from which the fat had almost disappeared, 
were an “astonishing number” of nucleated red blood-corpuscles, 
The number of these was so great that they were at least as numerous 
as the white corpuscles and colourless marrow-cells. There were more 
nucleated red corpuscles in the marrow than non-nucleated ones. 
“In contrast,” says Neumann, ‘‘ with the very distinct appearances in 
the bone-marrow, the results of the examination of the spleen and 
lymphatic glands were essentially negative.” 

In the spleen, he found nucleated red corpuscles; but so few of 
them that it was impossible to say that they were more numerous than 
in the aortic blood. 

In the lymphatic glands, he found chiefly leucocytes, “ but also, 
together with non-nucleated red cells, a few scattered nucleated red 
cells.” Some of the sinuses of the glands were filled with red blood- 
corpuscles, having among them large blood-corpuscle-holding cells 
(of 0°015 to 0°02 millimetres in diameter). 

This case Neumann considers very much against the blood-forming 
function of the spleen. But it will be noticed that some nucleated red 
corpuscles were found in the spleen, and that Neumann’s explanation 
that they were only caught there is as yet only a conjecture. Also 
that under this conjecture their absence from the spleen of one of his 
dogs (the one that died of anemia) cannot be explained. 

Regarding the interesting appearances found in the lymphatic 
glands, Neumann can hardly believe that either the nucleated or the 
non-nucleated red corpuscles were formed there; and is much more 
inclined to think that they found their way into the lymph-sinuses of 
the glands in consequence of a diseased condition of the arterial walls. 
I shall refer more particularly to the lymphatic glands of this case 
when I speak of the blood-forming function of the lymphatic glands. 

Neumann says that the post-mortem appearances in this case were 
so entirely like those of progressive pernicious anemia that the uterus 
alone prevented the case from being concluded to be one of that 
disease. And he considers that it supports the view of idiopathic 
anzmia which he published in 1877,! viz, that that disease is due, 
not to a diminished production, but to an increased waste or consump- 
tion of red cells, arising from unknown causes. 

In 1882-83 Dr D. G. Zesas? published two papers on the function 


1 Neumann, Berlin. klin. Wochenschrift, 1877, No. 47. 

* Zesas, ‘‘ Ueber Exstirpation der Milz am Menschen und Thiere.” Zesas, 
“‘Beitrage zur Kentniss der Blutveranderungen bei entmilzten Menschen und 
Thieren.”—Arch. f. klin. Chirurgie (v. Langenbeck’s), Bd. xxviii. 


336 DR J. LOCKHART GIBSON. 


of the spleen. He considers that the function of the spleen is to 
convert white into red blood-corpuscles. He experimented on rabbits, 
which, it will be remembered, were the animals used by Freyer when 
he obtained negative results as to a blood-forming action of the spleen, 
and which were also the animals laid aside by Bizzozero and Salvioli, 
because the results obtained from them were negative. He found 
that four to five weeks after excision of the spleen there was a decided 
increase in the number of the white corpuscles in the blood, and a 
decrease in the number of red corpuscles. These changes increased 
until the tenth week, ‘‘ when the blood was richest in white corpuscles, 
and very poor in red corpuscles.” He does not give any figures ; so 
it is impossible to say what he considers a great increase of white 
corpuscles, or to what extent the red corpuscles would have to be 
diminished before the blood could be considered ‘‘ very poor in them.” 

In the second of those papers, as a result of an experiment on a 
dog from which he removed, with a lethal result, both thyroid gland 
and spleen at the same time, Zesas argues that after removal of the 
spleen alone the thyroid takes on an increased activity, and in fact 
replaces it ; and that accordingly the removal of the spleen is borne 
only by an animal which still possesses a thyroid. 

When I consider more particularly the supposed blood-forming 
function of the thyroid, I shall attempt to show how precipitate and 
inaccurate Zesas was in coming to any such conclusion, without first 
ascertaining whether an animal survived the removal of the thyroid 
alone. 

He does not seem to have examined the blood-forming organs 
microscopically ; and in this paper he gives the bone-marrow credit 
for no blood-forming function whatever. His results are in favour of 
the blood-forming action of the spleen, though the experiments appear 
to me to have been too inexact to be of very much value. In his 
further experiments, which I shall mention under the observations on 
the thyroid gland, he advances further proof of the blood-forming 
function of the spleen (in cats and dogs). 


Having thus considered the more important views of others 
as to the blood-forming function of the spleen, and the experi- 
ments which seem to bear most directly on the subject, I shall 
now proceed to detail experiments made by myself, before 
attempting to draw and give my own conclusions. 

My experiments, though fewer and less complete than those 
of some of the authors mentioned, have yet, especially when 
taken in connection with those of Bizzozero, enabled me to come 
to definite conclusions regarding the action of the spleen in the 
production of red blood-corpuscles during extra-uterine life, 
And they have further enabled me to come to definite conclu- 
sions regarding the action of the bone-marrow and lymphatic 


glands. 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 337 


I removed the spleen from three dogs ; and the method I employed 
for observing the effect on the blood was the enumeration of the red 
and white blood-corpuscles by means of Gower’s “ hemocytometer.” 
I should have liked also to estimate the percentage of hemoglobin in 
the blood, but time would not permit me to do so. According, how- 
ever, to the observations and opinions of Bizzozero, already quoted, 
the percentage of hemoglobin in the blood has a direct relation to the 
number of red corpuscles; and all my observations have gone to sup- 
port this view, viz., that all the red corpuscles in the circulating blood 
have a similar quantity of hemoglobin, and that diminution or increase 
of hemoglobin ia the blood is due to diminution or increase of the 
number of red blood-corpuscles, and not to a smaller or greater amount 
of hemoglobin in any individual corpuscles. This I know is contrary 
to the opinion of most physicians, and of most French authors, who 
believe that the hemoglobin in the blood may be diminished even 
while there are a greater number of red blood-corpuscles than normal. 
I think, however, that this opinion has been due to including among 
the red corpuscles the bodies called by different writers “ globules of 
Donné,” “Koérnchenbildungen,” “ haemotoblasts,” or ‘‘microcytes.” If 
the view I have taken of these bodies in the first part of this 
paper be the correct one, the enumeration of them among the red 
corpuscles would quite account for a normal or increased number of red 
corpuscles being found along with a diminution in the percentage of 
hemoglobin. 

In freshly-drawn blood which is quickly examined, the red corpuscles 
have, to my mind, all the same colour. The hemoglobin, however, 
begins to diffuse out of the corpuscles very soon after the blood is 
drawn, and diffuses out of some more rapidly than out of others; so 
that if the examination of the preparation of blood be delayed, some 
corpuscles will appear to be less deeply coloured by hzmoglobin than 
others. The effect which slight pressure or slight capillary attraction 
has in causing the hemoglobin to diffuse out of the corpuscles is very 
marked, 

In speaking, at the end of this second part of my paper, of the 
development of the red corpuscles, I shall give further reasons for the 
view of the direct relation between percentage of hemoglobin and 
number of red corpuscles; and shall endeavour to show that the 
assumption of hemoglobin by the corpuscles takes place under the 
influence of their nuclei, at the same time giving reasons for thinking 
that after the nucleus disappears the corpuscle has no longer the 
power of adding to the quantity of hemoglobin it contains. 

I kept my dogs in as good hygienic conditions as possible, in a 
good-sized well-lighted room. They had a full and mixed diet, and 
their movements in the room were little or not at all restricted. 

In the excision of the spleen (and the same applies to excision of the 
thyroid) the animal was put under the influence of ether, which in 
such experiments should always be used. The animal takes the ether 
well, and it apparently has no bad after-effects. Antiseptic precau- 
tions were used during the operation ; but as these could not have been 
carried out thoroughly after the operation, I did not continue them, 

VOL. XX, Vi 


338 DR J. LOCKHART GIBSON. 


and used no dressing at all. I sewed up the wound with chromic-acid 


catgut, and latterly with horse-hair; and in every case of excision of. 


the spleen union by first intention was obtained. 

For excision of the spleen, after tying the animal out on its back 
and putting it under the influence of ether, I made an incision in 
the linea alba, immediately above the umbilicus, and about two inches 
long, into the abdominal cavity ; and I then put my fingers (previously 
thoroughly washed in 1-to-20 carbolic lotion) into the left hypochon- 
drium, where I quickly found the spleen, loosely attached to the great 
curvature of the stomach by the gastro-splenic omentum. I drew the 
spleen gently out through the wound, and secured the vessels passing 
to it in from four to six catgut ligatures, the tissues in which the vessels 
lay being included with them in the ligatures. The ligatures were 
then cut short, and the vessels entering the hilum were divided on 
the splenic side of the ligatures, The lips of the wound in the linea 
alba were then brought together with stitches of chromic catgut, and 
I was specially careful to bring the peritoneal surfaces into contact, 
in order that union of the peritoneal wound might at once take place, 
and the abdominal cavity be shut off, in case of the superficial part of 
the wound not healing by first intention. 

The dogs all recovered very quickly from the effects of the opera- 
tion. After coming out of the ether, they shivered for about a quarter 
of an hour; and two of them vomited once within the first half hour. 
But in all cases a few hours after the operation, 7.e., on the afternoon 
of the same day, the dogs were so well that no one going into the 
room would have supposed that they had been operated on that 
morning. At no time after the operation did the wound appear to 
give any pain. For the first day after the operation they got only 
milk as food ; on the second day they were given also a little meat ; and 
after that, their ordinary diet. 

Before operating I always kept a dog for a week, sometimes longer, to 
watch whether it would lose or gain weight under its new conditions 
of life, and also to obtain the average number of red and white 
corpuscles. In the tables here given, the averages of the different 
estimations of the number of corpuscles before the operation are 
placed at the head of the corresponding columns. 


The results of the examination of the blood after simple 
excision of the spleen were the following :’— 


ELaperiment I. 


A female dog, with short shaggy hair, somewhat of the Scotch 
terrier type. About three months old, and weighing 2 kilogrammes. 


1 Compare with enumerations of the blood in normal animals given by Lyon in 
Virchow’s Archiv, Bd. Ixxxiv. p. 207. The slight irregular variations in the 
number of the corpuscles in my estimations were due, I believe, to the attempts 
the remaining blood-forming organs were making to assert themselves. 


pers 


. 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 339 


The operation was performed on 26 September 1884, and the 
results of the estimations of the blood may be tabulated as follows :— 


Proportion of 
Leucocytes. | Leucocytes to 
Hemocytes. 


Hemocytes. 


Before Operation, . ‘ : 6,410,000 16,000 1: 400°6 


Operation on 26 September. 


26 September, 6 hours after, ; 5,940,000 28,000 5 a 1 
27 “ 1st day after, P 5,535,000 34,000 1:133°3 
28 i Qnd ,, .  .| 5,400,000 25,000 1: 216 
29 ES Bede ck Ye ive W- B.480,000 22,000 1 : 248-8 
2October, 6th ,, . .| 5,530,000 30,000 1:184°3 
4 br Rules Ty . | 5,540,000 31,000 1:1787 
9 :, 1S) ee me . | 6,000,000 24,000 1: 250 
a." :, 29nd ,, . «| 4,900,000 23,000 1:213 
re 30th ,, . «| 5,520,000 17,000 1: 308°8 
8 November, 43rd ,, . : 5,860,000 13,000 1 :450°7 
17 si 52nd ,, . «| 5,280,000 22,000 1 :236°3 
*25 i 606R 45) «3 . | 4,470,000 13,000 1 : 343°8 
3 December, 68th ,, . .| 5,290,000 14,000 1 :377°8 
ee > *,, S4th ,, .«  ~ |) 5,580,000 18,000 1 :310 
2January, 98th ,, . .| 6,080,000 17,000 1:357°6 
RE ‘ichhs.  .. - . | 5,610,000 11,000 1: 510 
29 7 UPA eerie . | 6,100,000 9,000 ioc rat 
26 February, 153rd,,_ . : 5,730,000 11,000 1 : 520°9 
13 March, WG6Sthies. 1): . | 6,500,000 10,000 1 : 650 


Experiment II. 


A female dog of the English terrier type, short black-and-tan hair, 
ears and tail clipped short. Weight about 5 kilogrammes. At least 
one year, probably two years old. 


Proportion of 
Hemocytes. Leucocytes. | Leucocytes to 


Hemocytes. 
Before Operation, . ‘ . | 7,520,000 16,000 1: 470°6 
Operation on 3 November. 

5 November, 2nd day after, . 7,280,000 21,000 1 : 346°6 
eee Hove eax hat. DO \O00 15,000 1:473°3 
1 ee eT, 6-4, fe 8}480, 000 16,000 | 1:528°1 
ne 1th, A. 4 vib 8,810,000 8,000 1 : 1076-2 
[ — 15th ,, . «|. 8,450,000 20,000 1: 422°5 

gen. 4, OR yet. = 8,870,000 9,000 | 1:930 
5 December, 32nd ,, . . | 8,980,000 10,090 1: 890 
meee deur a ©" 8 880000 9,000 | 1:98171 
1January, 59th ,, . «| 7,760,000 8,000 | 1:970 
*5 A 63rd. 55. : ; 6,590,000 17,000 1: 387°6 


340 DR J. LOCKHART GIBSON. 


Experiment ILI, 


A female dog, probably about eight months old, but seemingly full 


grown ; with short thick black-and-white hair. Weight slightly over 
5 kilogrammes. , 


Proportion of 


Hemocytes. Leucocytes. | Leucocytes to 
Hemocytes. 
Before Operation, : : : 7,000,000 6,000 1 :1168°2 
Operation on 15 November. 
19 November, 4th day after, . | 6,700,000 16,000 1: 418°7 
23 i 8th ney be : 7,220,000 12,000 1:601°6 
27 . Oth Se ee : 7,500,000 8,000 1 : 987°5 
2 December, 17th ,, -: : 6,850,000 6,000 1466 
9 7 4th ,, - Gee Je tle 6130, 000 22,000 1 :276°6 
22 fe Stun ek : 5,800,000 8,000 1: 728 
3 January, 49th ,, . ; 5,270,000 18,000 be 29257, 
Aes ie G0thy fe. 2 3 5,720,000 12,000 1: 478°5 
PAE | jord 2.8 D : 5,770,000 10,000 1 Dae 
“18 Hebruary, 90th 5.) © ‘ 4,810,000 10,000 1 : 481 
2 March, Ocoee = t 6,050,000 7,000 1: 864 
Be ae HOSths,. 97% : 6,590,000 6,000 1:1098 


In the dog of Huperiment J, the changes seem to have occurred 
quickly, chiefly within the first two months. They consisted in 
a decrease of the number of red corpuscles and a relative and 
absolute increase in the number of white corpuscles in the blood. 
The greatest decrease in the number of red corpuscles, as will 
be seen from the tables, was found two months after the opera- 
tion. After reaching a minimum, the red corpuscles then 
increased again in numbers, though not very regularly ; and the 
relation of the white corpuscles to the red gradually approached 
that which had existed before the operation. 

The increase in the number of white corpuscles, though it 
corresponded to a certain extent with the decrease in the number 
of red, did not correspond very regularly. The great increase 
for the first day or two was due, in part at least, to the operation. 
For the white corpuscles always rise in number, and the red 
corpuscles always fall in number, for a day or two after an 
operation; and that without any regular proportion to the amount 
of blood lost. Such changes due to an operation are always 
recovered from in about two days. 


1 Marked by *. 


ee Cn 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 341 


On the 15th of March, the day of the last estimation, and 
five-and-a-half months after the operation, the red corpuscles 
were slightly more than their original number. The fact, too, 
that the white corpuscles had then sunk below their original 
number seems to me to be likewise of significance, as indicating 
that the gland removed had for one of its functions the produc- 
tion of white blood-corpuscles. The important result of the 
blood estimations, however, is the distinct change observed in 
the number of ved blood-corpuscles. From this change it would 
appear that ‘the gland, if not taking a great part, must at any 
rate have taken some part in the formation of the red blood- 
corpuscles; and that although the other blood-forming organs 
had increased their activity, they had yet failed to bring the 
red corpuscles up to their normal number again until nearly six 
months after the operation. 

That the removal of the spleen had little or no effect on the 
growth of the animal, will be pretty evident when I state that 
the weight of the animal was 2 kilogrammes before the opera- 
tion; that it began to gain in weight almost immediately after 
the operation, and to increase in size; and that its weight on the 
26th of February (five months after the operation) was 4 kilo- 
grammes and 34 decagrammes. The dog was then in as good 
condition as on the day of operation, and, as will be seen, had 
doubled its weight (owing to increase in size), From the day of 
the operation to the day of its death, it was the picture of a 
healthy animal; and certainly there was not the slightest sign 
interference with digestion, which would be expected if Schifi’s } 
view of the relation of the spleen to pancreatic digestion were 
admitted. It took its food even more greedily after the operation 
than previously. In fact the behaviour of my dogs would rather 
support the statement that after the removal of the spleen the 
appetite is increased. 

In Experiment IJ., the results obtained were at first sight a 
little confusing: still they may, I think, be taken as likewise in 
favour of a blood-forming action of the spleen. It will be 
noticed that there wasa distinct fall in the number of corpuscles 
in the blood two months after the operation: the confusing point 
is that just after an initial slight fall, and apparently as the 


1 Schiff, Schweizer. Zeitschrift f. Heilkunde, Bd. i. (1862). 


342 DR J. LOCKHART GIBSON. 


immediate result of the extirpation, there was an increase rather 


than a decrease in the number of red corpuscles. Taking - 


all the facts together, I think the results obtained from 
the experiment (in which the animal, it will be remembered, was 
about two years old) still support the blood-forming function of 
the spleen, but point to that function being in adult life almost 
in abeyance. I think they show that even in adult life the 
spleen has some activity; and it seems to me that even the 
increase in the number of red blood-corpuscles after the 
operation, instead of being against this view, is rather in favour 
of it. For the increase might be explained by considering the 
spleen as a blood-forming organ, and supposing that on its 
removal the other blood-forming organs, which had previously 
been doing most of the work, were thereby stimulated to a greater 
activity that more than compensated for itsabsence. The explana- 
tion of the subsequent fall might be either that the bone-marrow 
and other blood-forming organs had been unable to maintain this 
high state of activity, or that the fall was merely part of an 
oscillation about the normal point. 

It will be noticed that the white corpuscles were diminished 
in number both relatively and absolutely, and that afterwards 
there was a slight increase in their number, concomitant with 
the decrease in the number of red corpuscles. 

An interesting point with regard to the animal is that it had 
an unusually well developed thyroid gland, which, however, did 
not enlarge after removal of the spleen. I shall subsequently 
consider the question of this dog’s thyroid under the effects of 
excision of the thyroid, in the third part of this paper. 

In Haperiment III, where a positive result was again 
obtained, we were dealing with a dog within the first year of 
extra-uterine life, though having already the appearance of 
being full-grown. 

In the case of this animal, the number of red corpuscles 
gradually decreased for the first six or seven weeks; next there 
was a hardly perceptible rise; and then they sank to their 
lowest point, reaching it three months after the excision. The 
white corpuscles increased both absolutely and relatively in 
number, though not in a very regular manner. Towards the 
middle of the fourth month they had begun to sink in number, 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 343 


and if the dog had been allowed to live longer they would 
probably have sunk even below their original number. The 
diminution of the red corpuscles in this dog was really a decided 
one, amounting to quite 2,000,000 out of every 7,000,000. 

The time of the greatest diminution of the red corpuscles 
corresponds pretty well in all three dogs, being between eight 
and twelve weeks; and it also corresponds with the time given 
by Zesas (ten weeks), as well as, interestingly enough, with the 
time given by Credé (two months), 

The last dog, like both the others, recover2d very quickly after 
the operation ; took its food very well; and after losing weight 
for the first day or two, increased again until it reached its 
former weight, at which it remained, weighing on the 3rd of 
March exactly the same as on the day of operation. Here also 
there were no signs of interference with digestion, and the dog 
would have been taken by any one for a perfectly healthy animal. 

During the examination of the blood of the dogs, I was never 
able to convince myself of the presence of any nucleated red 
corpuscles in it. 

Post-mortem Appearances.—Of the three dogs from which I excised 
the spleen, only one was allowed to live without further operation, 
viz., the dog of Experiment I. In the dog of Experiment III. I 
afterwards produced artificial anzemia, in order to see the condition of 
the blood-forming organs in a spleenless animal during the process 
of blood-regeneration. The dog of Experiment IT. was used also for 
excision of the thyroid, and its further history will be given under the 
head of that gland. 

The dog of Huperiment I. (on which no further operation was per- 
formed), on being killed after the expiry of five-and-a-half months, 
showed the following appearances :— 

Body well nourished ; neither more nor less fat than usual. No 
naked-eye abnormal appearances, with the exception of the absence of 
the spleen. There was not the least sign of there having been any 
irritation in the abdomen, either in the region from which the spleen 
had been removed or elsewhere. The liver, which has been noticed 
by Zesas to be larger in animals after removal of the spleen, was not 
distinctly enlarged, nor more than usually rich in blood. 


The tissues to which I paid particular attention were the bone- 
marrow, the lymphatic glands, and the thyroid gland. In the 
bone-marrow and lymphatic glands I found evidence of blood- 
formation, but not in the thyroid gland. 

As to the bone-marrow, I found evidence of blood-formation 


344 DR J. LOCKHART GIBSON. 


chiefly in the ved marrow of the ribs, the vertebral bodies, and 
the heads of the long bones; in short, in marrow of spongy bone. _ 
But I also found that the usually fatty marrow of the shafts of 
the humerus and femur had to some extent been transformed into 
red marrow, @.¢., was much redder than fatty marrow, though not 
so red as red marrow, and, further, presented microscopic appear- 
ances intermediate between red and yellow marrow. This 
change of the fatty marrow supports in an interesting way the 
blood-forming function of the spleen, especially if compared with 
the observations of Litten and Orth already referred to.—The 
marrow in the shafts of the other long bones had remained 
fatty. 

The marrow was examined fresh, and either undiluted or 
diluted with a neutral fluid. I preferred the latter method; 
because the only semi-fluid marrow could not be satisfactorily 
examined without dilution. For diluting fluid, a solution of sul- 
phate of soda, of sp. gr. 1022, was used; and as it was difficult to 
get an idea of the number of nucleated red cells in the marrow 
when the nuclei were uncoloured, the faintest trace of methyl- 
violet was added to the solution. It is impossible to say how 
much methyl-violet should be used: the amount needed is so 
small. It is best to use enough, but no more than enough, to 
colour the nuclei of the red and white cells. The addition 
of such a trace of methyl-violet to the “artificial serum” appears 
also to have the advantage of rendering it even less liable to 
alter the shape of the red corpuscles. This has been remarked 
by various observers. 

For the examination of the marrow, I first placed a drop of 
the diluting solution on a slide; then mixed the marrow with 
the drop, at the same time to some extent teazing it out; and, 
lastly, put on a cover-glass, pressing it gently down on the 
preparation so as to leave only a single layer of cells. In 
the case of spongy bone, I squeezed out the marrow by means 
of a pair of bone-forceps, and so obtained a semi-fluid marrow 
free from bony spicules. : 

The red marrow of a normal dog, when thus treated, with or 
without staining, shows numerous nucleated colourless cells of 
different. sizes, varying from about 8 micros. to as much as 16 
micros. in diameter. The nuclei are almost always relatively 


+ 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 345 


large; and they are more or less coarsely granular, this being 
due partly at least to an intranuclear net-work. The perinuclear 
protoplasm is faintly granular. Among the cells there are 
always a very considerable number of ordinary red blood- 
corpuscles. In addition, however, there are cells which can be 
distinctly recognised as nucleated red blood-corpuscles. These 
cells, as has already been said, are best seen when a trace of 
methyl-violet is added to the artificial serum in which they are 
examined. The methyl-violet has the effect of staining their 
nuclei, as well as the nuclei of the colourless cells of the marrow, 
and also of staining to a slight extent the perinuclear protoplasm 
of the colourless cells. Various stages may be seen in the 
development of the nucleated red cells in the bone-marrow, but 
the stage which is most characteristic, and which it would be 
difficult to mistake for anything but a nucleated red blood- 
corpuscle, is that where the cell is rather larger than a non- 
nucleated red blood-corpuscle, and has a nucleus which is not 
large as compared with the cell, and perinuclear substance 
which is as darkly hemoglobin-tinted as a non-nucleated red 
corpuscle (fig. 1, d).! Of this variety of cell, only a few can be 
seen in the red marrow of a normal dog. But there are numbers 
of what I consider to be earlier stages in the development of 
these cells from the ordinary cells of the marrow (fig. 1, d and c). 
Many of them, especially those of from 10 to 12 or even 14 
micros. in diameter, have a relatively very large nucleus with a 
very well defined outline (fig. 1, a). Filling up the interval 
between the nucleus and the capsule of some of these cells can 
be seen a thin yellow band, which I take to be the first appear- 
ance of hemoglobin in them (fig. 1, 6). So thin is the earliest 
yellow band, that it could be taken as significant of nothing, 
were it not for the fact that what must be later conditions of 
such cells can be seen, where it has become broader (fig. 1, c). 
And, indeed, all stages of transition can be seen, between the 
large cells, with their large nuclei and very thin band of 
hemoglobin-coloured perinuclear substance, and the typical 
nucleated red corpuscle, which is much smaller, and has a much 
smaller nucleus and a broad coloured band. As a rule, the band 
of hzmoglobin-coloured substance occupies the whole space 


1 The figure will be given in the next number of the Jowrnal. 


346 DR J. LOCKHART GIBSON. 


between the cell-envelope and the nucleus; but occasionally 


there can be seen a small amount of colourless substance 


immediately surrounding the nucleus, and still taking on the 
methyl-violet colouring. I have seen this remnant of the 
perinuclear finely granular substance of the cell only in the 
younger stages of development, never in a typical nucleated 
red cell. 

To return to the post-mortem appearances of the dog 
(Expt. I.) :-— 

The bone-marrow of the ribs, squeezed out, as stated, and 
mixed with the methyl-violet-tinted sodium-sulphate solution, 
showed great numbers of very young nucleated red cells; and 
here and there could be found nucleated red cells of the more 
typical variety. Nucleated red cells crenate very easily, thus 
giving an additional proof of their relation to the non-nucleated 
red cells. . 

The marrow from the heads of the femur and humerus showed 
enormous numbers of nucleated cells in the early stage of de- 
velopment: a greater number even than was found in the ribs. 
Also a few of the more typical variety. 

The partially reddened marrow of the shafts of the humerus 
and femur contained a few examples of developing red 
cells, 

In the lymphatic glands, very interesting and unexpected 
appearances were found. The glands of the mesentery were 
distinctly enlarged, when compared with those of normal dogs ; 
and they were distinctly succulent, though not decidedly redder 
than usual, A scraping from the cut surface of these glands, 
mixed with the methyl-violet-tinted sodium-sulphate solution, 
showed numbers of nucleated red cells, in earlier and later 
stages of development. There could be no doubt about them: 
for there were a few in every field; and quite as many of the 
later stages could be found as in the marrow of the ribs. This 
observation will be more interesting when it is compared with 
what I shall afterwards describe as having been found by me in 
the lymphatiz glands of an animal whose thoracic duct I had 
tied some time previously. 

The thyroid gland, when similarly examined, showed no signs 
of a blood-forming function. 


- 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 347 


I now pass to the dog on which Kuperiment III. was per- 
formed, and which I subsequently bled for the purpose of 
producing artificial anemia. On referring back, we see that 
on the 3rd of March, nearly four months after the operation of 
excision of the spleen, this animal had a number of red corpuscles 
per cubic millimeter of blood almost a great as the original 
number, and that the white corpuscles had almost their original 
relation to the red. 


Experiment IV. 


Dog previously used for Experiment III. 

On the 3rd of March I drew from the left carotid 20 e.c. of blood, 
being scarcely 4 per cent. of the body-weight ; and on the following days 
I estimated the blood, to notice the effect. In cases of drawing blood, 
as well as in my other experiments, I always put the animal under 
the influence of ether. 


Proportion of 
-Hemocytes. Leucocytes. | Leucocytes to 


Hemocytes. 
Before Operation, : : . | 6,599,000 6,000 1:1098°3 
Operation on 3 March. 
4 March, Ist day after, . . | 5,690,000 13,000 1 : 437°6 
a Indi, 5; : , | 6,060,000 10,000 1: 606 
Ot4 5; SED |! 43, P . | 6,250,000 8,000 1: 781°2 
wee? Ga . . | 6,270,000 5,000 1 :1254 


Six days after the loss of blood, which must be taken as an ex- 
tremely small one, the corpuscles had not quite returned to their 
previous numbers ; showing that blood-formation was not so rapid as 
usual.? 

As a proof, however, that in spleenless animals the blood is not so 
rapidly regenerated, the experiment cannot alone be considered of 
much value. 

On the 9th of March, I drew from the same carotid artery (the 
previous wound having in the meantime healed) a considerably larger 
quantity of blood, viz., 75 c.c., being about 1°8 per cent. of the body- 
weight. 

The dog recovered quickly from the operation, and was well next 
day, though not so lively as usual, being more inclined to sleep, and 


1 Compare with results of Lyon’s experiments, Virchow’s Archiv, Bd. lxxxiv. 


348 DR J. LOCKHART. GIBSON. 


less active in its movements. The amount of anemia produced will 
be evident from the following table :— 


Proportion of 


Heemocytes. Leucocytes. | Leucocytes to 
Heemocytes. 
Before Operation, : , : 6,270,000 | 5,000 1:1254 
Operation on 9 March. 
11 March, 2nd day after, . . | 4,400,000 7,000 1 :628°5 
Tai eee eno. tes . . | 4,490,000 9,000 1: 498°8 


The loss of blood had produced a very considerable degree of 
anzmia; and in this instance sufficient time did not elapse before I 
killed the aniutal to enable me to see to what extent the remaining 
blood-forming organs would be able to regenerate the blood, and how 
long it would take them to do so. 

Post-mortem appearances, after killing with ether, on the 14th 
of March :— 

Bone-marrow.—The marrow of the ribs contained a great 
many nucleated red cells, some of which had very small nuclei 
(fig. 1, e). 

The marrow in the whole length of the shafts of the humerus 
and femur was red, and contained great numbers of the early 
stages of nucleated red cells, as well as a fair number of the more 
typical nucleated red cells. 

The marrow in the heads of the long bones and the marrow of 
the bodies of the vertebrae contained a greater number of 
developing red cells than the marrow in the shafts of the long 
bones, but not so many as the marrow of the ribs. Most of the 
cells were in the earlier stages. 

The comparative rarity of the later stages of the developing 
red corpuscles, except in the marrow of the ribs, and the great 
numbers of the earlier stages, correspond entirely with the blood 
enumerations. For according to these the regeneration of the 
blood had even on the day of death hardly yet begun. 

Lymphatic Glands.-—Those of the abdomen were large and 
rather succulent, and if anything somewhat redder in the centre 
than usual. Some nucleated red cells were found, but not very 
many. There were, however, a considerable number of white 
cells with peculiarly clear perinuclear substance, which looked 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 349 


as if they needed nothing but hemoglobin to convert them into 
nucleated red cells. 

Thyroid Gland—Of usual size, most unusually pale, and 
presenting no appearance of a blood-forming function. 

In the abdomen there were not the faintest appearances of any 
irritation having followed the removal of the spleen. 

Up to this time I have been dealing with animals from which 
I had excised the spleen; and I think that my experiments 
furnish pretty conclusive proof that the spleen has in ordinary 
conditions a blood-forming action, if perhaps a subordinate one, 
throughout extra-uterine life, its activity being, however, pro- 
bably greater during the first year of life. The facts in favour 
of this action are :— 

1. The changes in the blood, consisting in a primary decrease 
in the number of red corpuscles and increase in the number of 
white, and in one case (Expt. I.) a secondary decrease of the 
white corpuscles below their original number after the red 
corpuscles had already returned to their original number. The 
secondary decrease in the number of white corpuscles, which 
corresponds with the observations on the blood of Credé’s 
patient three years after the extirpation of the spleen, points to 
the probable reason for the primary increase, viz., that an organ 
had been removed having, on the one hand, the function of 
transforming white corpuscles into red, and on the other hand 
the function of itself producing white corpussles. 

2. The partial transformation of the yellow marrow of the 
shafts of the humerus and femur into red marrow. 

3. The appearance of a considerable number of nucleated red 
cells in the lymphatic glands, only very few nucleated red cells 
being found in the lymphatic glands under ordinary conditions. 

I next tried to ascertain the truth of Bizzozero’s assertion that 
the spleen is brought into fresh activity by the production of 
artificial anemia. Experiment IV., already described, had given 
some evidence of a negative character in favour of Bizzozero’s 
view: for in it, after blood-letting in the spleenless animal, the 
regeneration of the blood did not take place as rapidly as, 
according to Lyon, it would in animals possessing spleens have 
done. The post-mortem appearances were similar to those found 
in the dog of Experiment I., being merely somewhat exaggerated, 


350 DR J. LOCKHART GIBSON. 


owing to the efforts of the remaining organs to overcome the 
artificial anzemia which had been produced. 

I was only able to perform one experiment for the purpose of 
ascertaining whether I could get any positive evidence in favour 
of Bizzozero’s conclusion, but from it fortunately got very 
decided evidence of the kind. 


- 


Experiment V. 


A very healthy female dog, about two years old, a mixture of pug 
and terrier. Weight 6 kilogrammes 20 decagrammes. . 

On the 10th of March I drew from the right carotid 140 cubic 
centimetres of blood, being about 2°6 per cent. of the body-weight. 
Then I left the animal for three days, feeding it as usual, and finding 
that during this time it did not lose weight. And then I killed it 
with ether, and examined the blood-forming organs. 

Spleen.—Unusually large, soft, and succulent, and of the rose-red 
colour which Bizzozero describes as so characteristic of a spleen 
containing many nucleated red cells. 

A scraping, examined in the usual way, showed very numerous 
nucleated red cells, many of which were in process of division. 
Many of the cells were in the earlier stages of nucleated red 
cells, but there were also a very large number in the later and 
more typical stages. There could be no doubt about these cells 
having been formed here. For their number, although smaller 
than the number found in the marrow of the ribs, was yet far 
above what could be accounted for by the supposition of nucle- 
ated red cells having been entangled in the spleen from the 
circulating blood. Moreover, the occurrence of the red cells in 
their different stages of development and the occurrence of the 
dividing red cells are very much against any such supposition. 

Lymphatic Glands.—Were perhaps a little larger than usual, 
but were no redder. A scraping from the the cut surface of a 
mesenteric gland showed many nucleated red cells, with very 
faintly coloured perinuclear substance, as well as a fair number 
of non-nucleated red cells. The bronchial glands showed the 
same appearances. 

Bone-Marrow.—The marrow of the ribs was of a very rich 
brown colour. In it were found enormous numbers of nucleated 
red cells, in all stages of development. These cells were more 
deeply heemoglobin-tinted than those found in the lymph glands, 


; 
‘ 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 351 


but were otherwise the same cells. The shafts of the femur 
and humerus were filled with a very rich red marrow, which 
contained numerous nucleated red cells, though not so many as 
the marrow in the heads of the bones. The marrow in the heads 
of the bones contained fewer than that of the ribs. 

There was a complication in this experiment, inasmuch as I 
had previously (three weeks before the blood-letting) removed 
from the animal one lobe of the thyroid gland. The case might 
therefore be interpreted as supporting the views of those who 
believe that the spleen and thyroid have a mutual compensatory 
function; and unfortunately I cannot altogether deny that such 
an interpretation is possible. But all my observations on the 
thyroid are greatly against its having a function similar to that 
of the spleen. I have, in fact, found no evidence of its having 
any blood-forming function. The dog in this experiment had 
suffered in no way from the removal of one lobe; and I shall 
further on, under the head of the thyroid gland, give other cases, 
both in animals and in the human subject, where removal of one 
lobe had no effect whatever. The examination of the remaining 
lobe of the thyroid gland, which was not distinctly enlarged, 
revealed no signs of a blood-forming function. 

I next, in order to be able to look at the question from as 
many sides as possible, estimated in two animals the numbers 
of red and white corpuscles in the splenic artery and vein, to 
see whether Bizzozero’s observations on this point could be 
confirmed. It will be remembered that he found, or concluded 
that he found, an increase in the number of red corpuscles, and 
a relatively still greater increase in the number of white 
corpuscles, in the splenic vein, over the numbers in the splenic 
artery. 

The results of my two observations were not very marked, 
but they pointed in the same direction as the results of 
Bizzozero’s experiments. 


Experiment VI. 


The first enumeration was taken from the blood of the splenic 
artery and splenic vein of a dog whose thoracic duct I had tied some 
time previously (with a result to be afterwards described). 

When about to kill the animal, I put it under ether, and cut 
directly down upon the spleen, on the outer border of the left rectus 


352 DR J. LOCKHART GIBSON. 


abdominis, just below the ribs. A good free incision was made, so 


that I could draw the spleen out of the abdomen without in any way ~ 


squeezing it, and without drawing on its blood-vessels. After clear- 
ing its vessels as carefully as possible, I made with a needle a puncture 
in the splenic vein sufficiently large to let the blood flow very freely. 
I made the puncture in the vein first, because a permanent wound in 
the vein would not affect the number of corpuscles in the blood of the 
artery, while if I had first taken blood from the artery and had 
happened to stop the flow in it, I should probably have found little 
or no blood in the vein when it was opened. After the required 
quantity of blood had been got from the vein, the gentlest pressure of 
a sponge for a minute sufficed to stop the flow from the wound in its 
walls ; and this seemed to have become closed, as the circulation went 
on as before. I then made a puncture in the artery, and took the 
required quantity of blood while it came freely spurting out. The 
enumeration of the red and white corpuscles gave the following :— 
Splenic Vein. 

Hemocytes 9,600,000, Leucocytes 11,000. Relative number of 

Leucocytes to Hemocytes 1 : 872°7. 
Splenic Artery. 

Hemocytes 9,410,000. Leucocytes 9,000. Relative number of 
Leucocytes to Hemocytes 1 : 1045-5. 

These figures support the results of Bizzozero, though not in 
a very pronounced manner, especially as the spleen in this 
experiment contained (as will later more particularly be stated) 
a moderate number of developing red blood-corpuscles, and was 
therefore more active than the spleen of a normal animal. And 
here, indeed, it must be remembered that Bizzozero’s own 
observations, made as they were on animals which had been 
rendered anzemic, were likewise made under conditions in which 
the spleen was actively producing red blood-corpuscles. 

We cannot expect that a very decided number of fresh red 
corpuscles should be added to every cubic millimeter of blood 
which passes through the spleen; and we must, I think, be 
content to allow that if the blood of the splenic vein contains at 
all a greater number of red corpuscles than the blood of the 
artery, then the spleen has really a blood-forming function. 

Moreover, the function of the spleen as a producer of white 
corpuscles is likewise supported, if indeed it needed any other 
than a histological support. 

I found no nucleated red corpuscles in the blood of either 
vein or artery. 


1 ee ie~ ewe +) 


st 


a oy Ae. 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 353 


ELeperiment VII, 


The second dog from which I got such an enumeration was the one 
I made artificially anemic (Experiment V.) tosee if the activity of 
the spleen would be increased. The steps taken for the enumeration 
were the same as in Experiment VI. The results, although in favour 
of the same conclusion as those of that experiment, were still less 
decided. This is remarkable, as the spleen was very rich in develop- 
ing red cells. 

Splenic Vein. 


Hemocytes 6,310,000. Leucocytes 9,000. Relative number of 
Leucocytes to Hemocytes 1 : 701-1. 


Splenic Artery. 
Hemocytes 6,250,000. Leucocytes 8,000. Relative number of 
Leucocytes to Hemocytes 1 : 781°2. 


These slight differences between the numbers of corpuscles in 
the vein and artery are quite in accordance with the observations 
of those writers who could find no difference between the con- 
tents of the vein and the contents of the artery in normal 
conditions. For if the difference is so slight when the spleen is 
actively forming red cells, it must be still slighter when the 
spleen is comparatively inactive. In other words, the addition 
of red corpuscles to a cubic millimetre of blood in passing 
through the spleen must in normal conditions be so small as 
to make it practically impossible to observe any numerical 
difference between the corpuscles in the artery and the corpuscles 
in the vein. 

But here another point must not be lost sight of, viz., what is 
stated by many authors to be one of the functions of the spleen, 
the breaking down of red blood-corpuscles. Later in my paper 
I shall speak of reasons for supporting this as one of the 
functions of the spleen. [f the spleen possesses any such 
function, the increase in the number of red corpuscles in the 
splenic vein over the number in the splenic artery would be 
no index of the number of fresh corpuscles added to the blood 
in its passage through the spleen. 


(Zo be continued in the next number.) 


VOL. XX. Z. 


Anatomical Notices. 


MALFORMATIONS OF PELVIS AND PELVIC ORGANS IN 
A FETUS. By Joun Patmer, L.R.C.P. Lond., M.R.C.S. Eng. 


Mrs J. was delivered of twins on the Ist of June 1884 (a month 
earlier than she expected). The first child was a girl, living and looking 
like a foetus of between 7 and 8 months, the second child was small 
and looked like a foetus at about the 6th month, this child lived five 
hours ; the first child is living now and is a very fine girl, the mother 
says as large as any of her children have been at the same age (14 
months). 


On superficial examination of the second child, the pelvis was found 
to be extremely small, and no external opening could be found ; in the 
middle line, a little below the lower end of sacrum, was a very small 
projection of skin with a small depression extending round it ; farther 
forward, a little below the symphysis pubis in the middle line, was a 
small nipple-like projection of skin, but no opening in or as it ; 
the appearance is represented in fig. 1. 


On post-mortem examination of chest, heart and lungs were found 
to be normal; and on opening abdomen, liver, spleen, suprarenal 
bodies, small and large intestines as far as sigmoid flexure were normal, 
cecum in right iliac fossa; in left iliac fossa the descending colon 
ended in a large closed dilatation measuring in length 2 inches, in 
width 1} inches; no trace of rectum either as a fibrous cord or other- 
wise could be found. The cavity of pelvis, which was very shallow 
and small, measured transversely ? of an inch, antero-posteriorly =, of 
an inch; in it was the lower part of a closed sac which measured in 
length from above downward 1} inches and transversely 14 inches. 
This sac on being opened was found to contain about two teaspoonfuls 
of milky-looking fluid, which under the microscope showed epithelial 
and fat cells and debris, the pouch at end of descending colon was 
attached to the upper and left side of this by connective tissue (their 
cavities did not communicate). Into the sides of this closed sac at its 
upper part were seen to be inserted two thickened cords which 
suddenly became fine tubes (Fallopian tubes), and were contained in 
the broad ligaments, the ovaries lying immediately below the fine 
tubes, the thickened portions apparently represented the two halves 


ANATOMICAL NOTICES. 355 


of an undeveloped uterus; the round ligaments were large, normal in 
position and destination. Reaching rather higher up than this sac, 
and somewhat to the left side, and extending downward behind to its 
base, being closely adherent to its posterior wall, was another sac, 
oblong in shape, the upper extremity pointed and continued into a 
tube which passed upward and was attached to the left kidney, higher 
than the hilus; this sac, on being opened, was found to contain clear 
urine, the walls of the sac were thin, into its base the right and left 
ureters entered. These two sacs, which represented the vagina and 
bladder, were almost entirely situated in the false pelvis and lower 
part of abdomen, the bladder being quite posterior. The left kidney 
measured 12 inches in length and 55, of an inch across, was wholly 
composed of cysts, and from it two tubes passed to the bladder, one 
the ureter, which was attached at the hilus, the other attached above 
the hilus and ending in the apex of bladder as before mentioned. 
The right kidney was very small, also cystic; it measured ;‘5 of an 
inch im length and } of an inch in width; the kidneys were in the 
normal positions, except that the right ureter being short, the corre- 
sponding kidney was somewhat lower down than usual. Fig. 2 
shows the relative positions of the internal parts. 

On examining the pelvis the symphysis pubis was found continued 
downward and backward to the posterior part of the tubera ischia ; 
the rami of ischia and pubes being close together and held in apposi- 
tion by fibrous tissue, so that there was slight mobility; the only 
outlet to pelvis was behind this and in front of sacrum, a space only 
large enough to admit the tip of the little finger; the two internal 
obturator muscles covered the cartilages and formed the floor of pelvis. 
At the umbilicus only one hypogastric artery was found and no 
urachus. 


Fic. 1.—The external appearance of perinzeum of fetus, legs acutely 
flexed on the abdomen. 


356 ANATOMICAL NOTICES. 


Fic. 2.—The relative position of genito-urinary organs; R.K., L.K., 


right and left kidney; B., bladder ending above in a fine tube attached - 


to upper part of left kidney; V., closed vaginal sac; B.L., B.L., broad 


Fig. 2. 


ligaments with halves of uterus internally and Fallopian tubes and 
ovaries in outer part; D.C., descending colon ending in large closed 
pouch ; R.L., round ligaments of uterus. 


ON AN ANOMALOUS MUSCLE IN THE FRONT OF THE 
NECK IN A HUMAN SUBJECT—A STERNO-PETROSO- 
PHARYNGEUS. By G. E. Rennin, B.A. (Sydney), Student 
of Medicine, University College, London. 


Tus peculiar muscle was met with in the cours? of a dissection of the 
anterior triangle of the neck, in a well-developed muscular male subject, 
which was brought into the dissecting room of University College, 
London, during the month of October 1884; and acting upon the 
suggestion of Professor Thane, I have prepared the following account 
of it for publication :— 


_— 


ee ee ee ee ee ee a 


ANATOMICAL NOTICES. 357 


The muscle took its origin from the anterior surface of the manu- 
brium, immediately internal to the sternal head of the right sterno- 
mastoid, by a thin flat tendon, nearly half an inch wide, and about 
half an inch long, extending higher on the posterior than on the 
anterior aspect of the muscle. The tendon gave rise to muscular 
fibres which formed a somewhat rounded bundle, some two or three 
lines in diameter, The muscle then passed up the right side of the 
neck, superficial to the sterno-hyoid, sterno-thyroid, and omo-hyoid 
muscles, then over the hypoglossal nerve, and then passing between 
the internal and external carotid arteries, disappeared beneath the 
digastric and stylo-hyoid muscles. On being traced subsequently to 
its termination, the muscle was seen to split up, at about two inches 
from its insertion, into three sets of fibres, the disposition of which 
was as follows:—(1) One set of fibres passed directly upwards, and 
was inserted into the vaginal process of the temporal bone, the inner 
fibres of insertion being in immediate contact with those of origin of 
the levator palati. (2) Another smaller set, turned slightly inwards, and 
passing beneath the lower border of the superior constructor, blended 
with the pharyngeal aponeurosis, under cover of that muscle. (3) A 
third set, much shorter than the preceding, turned a little outward 
and forward, and blended with the lower fibres of the superior 
constrictor muscle of the pharynx. 

The muscle was completely surrounded by the deep cervical fascia, 
as that structure passed forwards from the anterior border of the 
sterno-mastoid, and had the following relations to surrounding 
structures :— 

(a) To Muscles.—It rested upon the sterno-hyoid, sterno-thyroid, 
and anterior belly of omo-hyoid ; then upon the middle constrictor of 
the pharynx. It was crossed by the posterior belly of the digastric 
and the stylo-hyoid, and higher up by the stylo-pharyngeus; and in 
this case by an accessory slip from that muscle to the great cornur of 
the hyoid bone. To its outer border, for about half its length, and at 
a distance of half an inch from it, was the sterno-mastoid. 

(b) To Vessels.—It was crossed superficially at its lower part by the 
anterior jugular vein, and higher up by a communicating branch 
between the external and anterior jugular veins. It passed over the 
facial vein, and then sinking inwards between the two carotid arteries, 
lay to the inner side of the external carotid. A small vein from the 
substance of the muscle entered the internal jugular vein. 

(ec) To Nerves.—It was supplied by a small twig from the glosso- 
pharyngeal nerve which entered it on its outer surface. The muscle 
in its upward course crossed over the branch of the descendens noni to 
the anterior belly of the omo-hyoid, the hypoglossal, and the glosso- 
pharyngeal nerves. 

I have not been able to find any account of a similar muscle 
on record. Macalister, in Z'ransactions of Royal Irish Academy, 
1871, refers to one case in which a slip from the anterior margin 
of the sterno-mastoid was inserted into the tympanic ring; and 
this is the only record I can find of an insertion of any part of the 


358 ANATOMICAL NOTICES. 


sterno-mastoid into the petrous part of the temporal. I have also 


consulted Testut’s Anomalie Musculaire, Gruber’s Anatomische Notizen - 


in Virchow’s Archiv., and also Meckel’s Vergleichende Anatomie, but 
have failed to gain from them any information bearing upon this 
peculiar abnormality. 

In conclusion, I have to acknowledge my indebtedness to Professor 
Thane for kindly advice, and for lending me works of reference. 


NOTE ON A CASE OF CONGENITAL HYPERTROPHY OF 
THE LEG. (Diffuse Venous Nevus.) By GILBERT 
Baruine, M.B. 


THE specimen which I had the opportunity of dissecting was the left 
leg of a man, aged fifty years, who stated that at his birth the left leg 
was larger round than the right, and that the left foot was deformed, 
but, except for the weight of the limb, he had not suffered much in- 
convenience, and had until recently been able to get about quite 
actively. 

I had no opportunity of making comparative measurements of the 
two legs, but, roughly speaking, the left leg, from a little above the 
knee downwards, was about three times the size of the right. The 
enlargement was not quite uniform,. affecting chiefly the posterior 
surface, and not extending further on the foot than the base of the 
metatarsus. There was varicosity of the smaller skin veins, but there 


was no ulceration of the skin, nor had there been any. To the feel, 
the soft tissues of the leg gave the sensation of a lipoma with many 
dense fibrous septa. The foot was in a condition of calcareo- valgus. 
When the skin was removed, the subcutaneous tissue was found 
greatly thickened, partly from an increase of fat, but mainly from an 
abundance of fibrous tissue, in which were imbedded a large number of 
small, thin-walled and tortuous veins; from this thick layer, septa passed 
to the deep parts of the leg, ensheathing the muscles, &c. The posterior 
muscles appeared to have retained their shape and size, but the 
muscular fibres were almost replaced by fat, without, however, any 
nzvus condition. The tendo Achillis and its sheath were so blended 
with the surrounding fatty and fibrous tissues as to be indistinguish- 
able from them, but the other posterior tendons and their sheaths 
were healthy, except that the sheaths were thickened by the nevus 
tissue. 

The anterior muscles, though small, appeared to have healthy fibres, 
but all of them, except the tibialis anticus, were shortened, and their 
sheaths were generally thickened with nevus tissue. 


ANATOMICAL NOTICES. 359 


The peroneus, longus, and brevis, though of good size, had under- 
gone fatty change to a considerable extent, and they were shortened. 

On division of the tendons of the extenscr longus digitorum, the 
extensor proprius pollicis, and all three peronei, the foot was easily 
brought down to a right angle with the leg, but there still remained a 
considerable amount of outward rotation of the anterior part of the 
foot at the transverse tarsal joint. 

The posterior tibial vein was single and much enlarged, so much so 
as to admit of ones little finger being passed into it. 

The principal nerves were all thickened, but the posterior tibial was 
especially so, its circumference being quite as great as that of an 
ordinary sciatic. Microscopically, this enlargement was seen to be 
due to an increase of the epineurium and perineurium. 

The bones of the leg were fairly developed as regards size, but they 
were deficient in weight, and their surfaces presented numerous irre- 
gular outgrowths, such as are sometimes seen at the insertion of 
tendons in the lower extremity, though here quite independent of 
those attachments. 

The bones of the foot were in marked contrast to those of the leg, 
being small, feeble, and much distorted, the deformity falling chiefly 
upon the os calcis, which was hardly recognisable, as it was mostly 
represented by a thin vertical plate supporting a small horizontal plate 
projecting from its side. 


NOTE ON THE MANDIBULAR DENTITION OF THE 
SHREWS. By G. E. Donsoy, M.A., F.R.S. 


Tae Shrews (Sorictde) form a very compact family of Insectivora, 
the species of which, among other points of close resemblance one to 
another, are believed to possess the remarkable peculiarity of having, 
whatever may be the number of the upper teeth, invariably six pairs 
of mandibular teeth, of which the first on each side is considered an 
incisor, the second a canine, the third a premolar, and the remaining 
three molars. It was, therefore, with much interest that I lately dis- 
covered a rudimentary seventh pair of mandibular teeth in the other- 
wise also remarkable species Myosorex varius, Smuts., of South Africa. 
The very small additional tooth exists (in every specimen examined) 
on each side between the second and third teeth; it is quite invisible 
to the naked eye, but may be readily made out, by the aid of an 


ordinary lens lying between the upper surface of the hinder part of the 
base of the second tooth and the under surface of the overlapping fore 
part of the base of the third tooth, which leave its small extremity 
alone uncovered. The direction of the cusp of this tooth is such that, 


360 ANATOMICAL NOTICES. 


if produced, it would come in front of the first maxillary tooth, and 
therefore should, according to the system of dental notation now in 
use, be considered the lower canine, whence it would naturally appear 
to follow that no lower canine exists in any other species of the 
family. This and other interesting questions connected with the 
dentition of the Shrews I hope to deal with fully in Part ILI. of my 
Monograph of the Insectivora, which will be ready for publication in 
a few months. 


Journ. of Anat.& Phys, Jan® 1886. Vol. XX PL. 


fig. 74 


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Fournal of Anatomy and Phystology. 


ACTION OF INFUSED BEVERAGES ON PEPTIC 
DIGESTION. By James W. Fraser, M.D., C.M. Ed, 
M.R.C.S. Eng. 


THIS paper is a continuation of that which appeared in the 
Journal of Anatomy and Physiology (vol. xviii. p. 13 et seq), 
and is based on the results of the same experiments, the 
difference being that the amount of peptones dialysed, instead of 
being estimated as the total organic matter, as was done in that 
paper, is here estimated by the amount of organic nitrogen. 
It will be found by reference to the earlier paper that, after the 
various meats had been digested by a peptic fluid in presence 
of the various beverages, a measured quantity of the filtered 
solution of peptones was placed in a Graham’s parchment paper 
dialyser, and this suspended in a measured quantity of distilled 
water fora fixed time. The dialysed matters in solution were 
then evaporated to dryness in a water bath, a weighed quantity 
of common salt having first been added to them to make the 
residue a more manageable substance. Many of these residues 
had been preserved, the entire quantity not having been used in 
the former experiments, and it was on these that the experiments 
to be described were performed. Some of the residues had been 
entirely used or destroyed, and hence, in comparing the tables 
appended to this paper with those in the former one, gaps will 
be found. 

A process of estimating the nitrogen, sufficiently accurate to 
deal with the small quantities available, was much desired from 
the first, but it was only after many experiments that the 
following method was arranged and improved until it was 
satisfactory :— 

VOL, XX. 2A 


a 


362 DR JAMES W. FRASER. 


Small Bohemian glass combustion tubing, about j-inch diameter, 
was taken, and tubes about 4 inches long, drawn out and sealed at 
one end, were made of it. Into such a tube about 1°5 gram. of the 
residue was weighed, the tube itself being weighed first and then 
again after the substance had been placed init. To this was added 
four or five times its bulk of powdered recently ignited soda lime, and 
the tube was plugged with a small piece of recently ignited asbestos. 
It was then possible by shaking or lightly knocking the tube against 
a table to thoroughly mix its contents, without the smallest possibility 
of losing any part of them. The mixed matters should only about 
two-thirds fill the tube, and when mixed the asbestos should be 
pressed down the tube until it touches the surface of its contents, 
and, if necessary, the plug should be pierced with a coarse needle to 
prevent undue pressure behind it during the combustion, which would 
burst the tube. 

The ammonia given off in the combustion was so small in quantity 
that no alkalimetric process was sufficiently accurate to estimate the 
differences between that derived from various samples. A modified 
Nessler process of estimating the ammonia was therefore employed, 
and the apparatus used and the method followed were the following :— 

The combustion tube was united by a short piece of india-rubber 
tube to a right-angled glass tube. This was passed through one hole 
in an india-rubber stopper of size suitable to fit in one of the ordinary 
Nessler test glasses, The other hole in the stopper was occupied by a 
large glass tube drawn out below to fit the hole in the stopper, and 
having its upper part filled with broken glass. The Nessler test glass 
was filled up to the 50 c.c. mark with distilled water, and the stopper 
having been inserted, 1 c.c. of the normal volumetric solution of 
sulphuric acid was poured over the broken glass in the tube above 
mentioned. As the apparatus behind this tube was air-tight, the acid 
was retained in the tube. Before heating the combustion tube the 
india-rubber connection between it and the right-angled tube was pro- 
tected from the effects of the heat by wrapping a narrow piece of wet 
linen round it. The upper end of the large tube containing the 
broken glass and sulphuric acid was connected to a Bunsen filtering 
pump, which caused a negative pressure of about 4 inches of mercury 
to be kept up in the whole apparatus, and at once showed if any 
leakage had occurred, and prevented any loss of ammonia through the © 
leakage. The apparatus having been thus arranged, heat was applied 
to the combustion tube, and continued until the whole tube was red- 
hot and the water began to rise in the right-angled leading tube 


ACTION OF INFUSED BEVERAGES ON PEPTIC DIGESTION. 363 


through absorption of the ammonia. Then a clamp was placed on the 
tube leading to the filter pump, and the end of the combustion tube 
was broken off with wet crucible tongs, when the negative pressure 
existing in the Nessler tube caused a rush of air through the com- 
bustion tube, sweeping the last traces of ammonia with it into the 
water. ‘The combustion tube was then broken off with the wet tongs, 
between the asbestos plug and the india-rubber connection, and a 
current of air aspirated through the apparatus by the filter pump to 
carry any traces of ammonia, not caught by the water, into the broken- 
glass tube, there to be fixed by the sulphuric acid. The substances 
operated on not being quite dry, a few drops of condensed moisture 
were usually found in the horizontal arm of the leading tube, and 
were washed down into the Nessler glass with a little distilled water. 
The broken-glass tube, the under surface of the india-rubber stopper, 
the leading tube, and the walls of the Nessler glass, were then washed 
with distilled water, and the washings, mixed with the original 50 c.c. 
of water, were made up to 100c.c. In the greater number of these 
experiments the solution thus obtained, though too weak for alkali- 
metric estimation, was too strong to estimate by Nessler’s process, the 
colour given being too dark for small variations to be easily appreciated. 
Therefore 25 ¢c.c. were measured out, with a pipette washed with the 
solution, into another Nessler glass, and this having been made up to 
100 c.c. with distilled water, was Nesslerised, and the amount of 
ammonia estimated in the usual manner. 

The process of digestion was described in the earlier paper, and 
the only reference to it required here is to notice that, when the 75 cc. 
of mixed digestive fluid and beverage containing peptones in solution 
had been filtered, 25 c.c., or one-third of the solution, was used for 
dialysis. ‘The mixture of salt and dialysed matters obtained by the 
evaporation mentioned above was weighed, and its weight was called 
T. The weight of the quantity of the mixed substances taken for 
combustion was called a, and the amount of ammonia found £. 

aL Xo 


Xa a “peed 
Then aa ammonia in JT, and 7 aor ammonia in the whole 


3T 14, : 
of the 75 c.c. of liquid, and ae x pz =uitrogen in the same amount 


of liquid, and by this formula the calculations of the experiments 
were worked out. When, as described above, one-fourth of the 
solution of ammonia alone was Nesslerised, 8 was of course obtained 
by multiplying the amount of ammonia found by 4. 


The object of thus re-testing the results of former experiments 


364 DR JAMES W. FRASER. 


was twofold—firstly, to put the conclusions drawn from the 
total organic solid estimations to the proof, and secondly, to 
form a link between those experiments and a new series now in 
course of performance, and for which the old process was found 
unsuitable. 

The conclusions drawn from the experiments on which the 
first paper is founded may be summarised here, it being premised 
that they must be taken as applying to experiments performed 
under the conditions stated in that paper, viz., invariable amounts 
of meat, digestive fluid, and beverage, and invariable temperature 
and time of digestion; and that all actions on vital processes of 
secretion, movement, &c., are eliminated. The conclusions, as 
stated in the former paper, were the following :— 


(1) All infused beverages retard the peptic digestion of albu- 
menoid food-stuffs, with the four exceptions mentioned above, viz., 
ham and white of egg with coffee, and fish with cocoatina and with 
cocoa. 

(2) The digestion of the meats ordinarily used at breakfast, viz., 
ham, egg, and salt beef, is less retarded by the action of tea or coffee 
than that of other meats, and the same is true of roast beef; with 
cocoatina a somewhat similar grouping occurs, but with regard to the 
other beverages of the cocoa class no such division into a group of 
breakfast meats, and a group of those less suitable for breakfast use, 
has been observed. 

(3) That this retarding action is less as a rule with coffee than 
with tea, and less with either than with the beverages of the cocoa 
order. 

“(4) That the retardation is caused (a) in the case of ‘teas,’ by the 
tannic acid assisted by the volatile oil, the former precipitating the 
uncoagulated albumenoids of the food, and the syntonin and peptones 
as formed, tanning the gelatinous constituents of the meats, and 
removing some of the pepsin by entangling it with these precipitates, 
and the latter retarding the action of the pepsin. The alkaloid of tea 
appears to assist digestion, but its action is masked by that of the 
tannic acid and volatile oil. () In the case of ‘ coffee,’ the caffeo- 
tannic acid and volatile oil retard digestion, and the alkaloid assists it; 
and, therefore, in the cases where this beverage assists digestion, the 
alkaloid must be the active agent in producing the result, and in the 
cases where digestion is retarded, the caffeo-tannic acid and the volatile 
oil. (c) In the case of the cocoas, the tannic acid, volatile oil, and 


ACTION OF INFUSED BEVERAGES ON PEPTIC DIGESTION. 365 


alkaloid all assist in retarding digestion, but under the conditions of 
the Standard Process, the clogging action of the suspended matters is 
the most potent factor. 

“(5) In retarding the consumption of acid during digestion tea has 
the greatest effect, coffee has no more effect than water, and cocoa 
increases the cousumption. 

“(6) Coffee and cocoa cause the peptic digestion of albumenoids to 
pass on through the stage of peptones to the formation of leucine and 
tyrosine. 

“(7) Tea acts on the digestion of fresh meat so as to increase the 
production of flatus, but has no such effect with salt meat, and coffee 
has no more effect than water. 

(8) The addition of cream and sugar to the beverages reduces the 
retarding action of tea on digestion, but increases that of cocoa; and 
coffee appears to have its action reversed by these additions, but this 
result is doubtful.” 

Experiments had been performed (1) on the digestion of beverages 
alone, these being known as “ Factor” experiments, because the result 
obtained had to be subtracted from the result of the digestion of each 
meat in presence of that beverage; (2) on the digestion of meats in 
presence of the beverages called, for the sake of a name, “ Peptone” 
experiments; (3) “Time” experiments, in which the time taken for 
complete solution of a given quantity of white of egg was the variable 
on which the conclusions were based ; and (4) experiments to ascertain 
the causes of the effects, noted under the second and third heads. Of 
the first class, fifteen experiments were performed, but only in nine of 
these had the mixture of salt and peptones been preserved ; tea and 
cocoatina being among the absentees, new experiments, but made by 
exactly the old process, were performed, and therefore eleven results 
are presented in Table B. 

Of the second class, fifty-seven experiments had been performed, 
but of these the residues had only been preserved in thirty-five cases, 
and no new ones were prepared. Of the third class naturally there 
was no residue to preserve, and of the fourth class, forty experiments 
had been performed, but only sixteen residues were preserved, and no 
new ones were prepared. 

Of the above quoted conclusions the first, second, and third were 
re-tested in these new experiments. Some of the conclusions under 
the fourth head were also examined, while those under the fifth, 
sixth, and seventh heads were not meddled with. ‘The eighth set of 
conclusions were also examined. 


366 DR JAMES W. FRASER. 


Turning now to these new experiments, Table B (Appendix, 


page 381) is the first to be noted, and consists of three columns, - 


A, B,and C. Column A contains the “ Factors” obtained from 
the digestion of 25 cc. of each beverage, with 50 c.c. of diges- 
tive fluid, the results being estimated by the old process ; column 
B contains the factors expressed as the weight of nitrogen 
derived from the dialysable organic matter; and column C 
shows what percentage of this dialysable organic matter consists 
of nitrogen. 

The figures in A and B will be found, as a rule, confirmatory 
of one another. It is true that three teas, Chinese, Indian, and 
green, appear to give too small results as compared with the 
mixed tea, but in most of the others the mutual relations be- 
tween various sets of figures is maintained in the two columns. 
In all cases more nitrogen is found where the beverages have 
been digested than where the digestive fluid has only had water 
added to it, a result naturally depending on the highly nitro- 
venised nature of their alkaloids, which more than compensates 
for any loss of nitrogenous matter from the digestive fluid, by 
precipitation of albumenoids by the tannic acid or its homologue 
of the beverages, or by delay produced in digestion by their action. 

Turning to the percentage of nitrogen found in the dialysed 
matter, this is found in the cases of six beverages to fall below 
that when water is used as the beverage, viz., Chinese, Indian, 
and green tea, coffee with chicory, cocoatina, and cocoa. In the 
case of the three last the explanation is simple, the sugar, 
dialysed with the peptones, being free from nitrogen, naturally 
reduces the percentage of the latter in the total organic solids. 
And this is an argument in favour of the nitrogen process in the 
estimation of peptones, which are thus dissociated from all non- 
nitrogenised admixture. - In the case of the three teas, however, 
this want of uniformity in their percentages of nitrogen is un- 
explained by any reason which does not apply to other teas in 
which the ratio is much higher, and this adds to the suspicion 
with which these three factors must be regarded. 

Infusing tea with alkaline water is seen to extract a rather 
larger quantity of nitrogenous matter than when plain water is 
used for infusing the beverage, but the percentage of nitrogen is 
lower, showing that more hydro-carbonaceous matter (probably 


sa si 


_ ACTION OF INFUSED BEVERAGES ON PEPTIC DIGESTION, 367 


tannic acid uniting with the soda) is extracted in proportion. 
The nitrogen percentages of mixed tea and of cocoa nibs are 
both high, showing that these infusions, when properly made, 
contain much alkaloid and little non-nitrogenised matter. The 
percentage of nitrogen in the case of coffee is much higher than 
when chicory is added to it, this, of course, depending on the 
sugar contained in the latter. 

The largeness of these “ factors” and their differences among 
themselves show the necessity of subtracting them from the 
amounts obtained, when meats are digested in presence of the 
beverages, in order that serious errors may be avoided. Leaving 
now the factors, and turning our attention to the second series 
of experiments, in which 5 gram. of meat were digested by 
50 ec. of digestive fluid, in the presence of 25 c.c. of each of 
the beverages; the results of these experiments are embodied in 
Tables D to G inclusive. Table D consists of the results 
obtained by the estimation of the peptones, obtained by the 
dialysis of the products of digestion, as organic matter; while 
Table D, is comparable with it, and contains the results of their 
Da ailicn by the amount of nitrogen they contain, and Table D, 
shows the ratio between the two sets of figures thus obtained, 
i.e, the percentage of nitrogen contained in the dialysable 
matters. Table E, extracted like Table D from the former 
paper, shows the percentage digestive power of each mixture of 
beverage and peptic fluid, that of water and digestive fluid being 
taken as 100, and the figures therein contained were obtained 
by multiplying the result for each beverage by 100, and dividing 
by the result for that meat digested in presence of water. It is 
evident that if at the end of the experiment some meat was left 
undigested, these figures will give the percentage digestive 
powers of the various mixtures. Table E, is one similarly ob- 
tained from the nitrogen estimations. 

Some irregular results in these experiments were noted in the 
former paper, and are to be found in Table D. 

These consisted of the two minus results—egg digested in 
presence of chocolate, and bread in presence of the same bever- 
age—but, as these two residues were not among the number 
preserved, they do not appear in Table D,, and call for no notice 
here. The other irregular results were four in number, and 


368 DR JAMES W. FRASER. 


were more noticeable in Table E, where four figures, egg in 
presence of coffee, ham in presence of coffee, and fish in presence. 
of cocoatina and of cocoa, were found above 100 per cent. The 
residues of the ham experiments are not among those preserved, 
nor is that of fish in presence of cocoatina. Therefore, the only 
two to notice are (1) that of egg in presence of coffee, which 
receives remarkable confirmation from these new experiments.? 
This result will be seen to have been further confirmed by the 
“Time” experiments above referred to; and the new series of 
experiments, of which mention has been made as now progress- 
ing, also have the same result. Also, it was noted by Herzen 
(Revue Médicale de la Suisse romande, Janvier 1884) that, in a 
case of gastric fistula in a man named Band, on whom he made 
various experiments, café noir assisted the digestion of albumen, 
though he says tea is without action; a result not in consonance 
with the experiments here detailed. (2) The other irregular 
result, fish in presence of cocoa, is not confirmed by these new 
experiments, which make it appear that cocoa delays, not assists, 
the digestion of fish. It was said that the result making it 
appear to assist the digestion of fish was probably fallacious, and 
these new estimations confirm that view. 

But in Table E, five more irregular results appear, which were 
regular in Table E, viz., roast beef in presence of tea, cocoatina, 
and cocoa, fowl in presence of coffee, and bread in presence of 
cocoa. As to the first four, nothing can be said except that, 
possibly the residues not being thoroughly dry, and having been 
preserved nearly three years in tubes corked securely but only 
with ordinary corks, some loss of water may have taken place, 
and thus have caused the rise in the amount of nitrogen. The 
results in the case of beef were large in the former experiments, 
but kept below the 100 per cent. In the case of bread digested 
in presence of cocoa the error is enormous—so great as to bring 
the percentage digestive power of that mixture up to 464-0 per 
ceut., aresult perfectly impossible. The explanation is found in 
the note appended in Table D to this experiment, which note, 
though only attached to the cases of chocolate and coffee, really 
applies more or less to all the bread experiments, and which states 
that, in the evaporation of the residues some charring occurred. 

1 See Table E,. 


ACTION OF INFUSED BEVERAGES ON PEPTIC DIGESTION. 369 


This charring would naturally drive off much nitrogen, and leave 
a proportionately large percentage of carbon in the residue. Now 
the bread-cocoa experiment was the least affected by the char- 
ring, and hence contained most nitrogen; while the nitrogen in 
the other cases being reduced to a very small amount, the large 
result in the case of the least harmed specimen naturally 
followed. This result, from its obvious falsity, is left out in 
ealculating the average digestive power of a mixture of cocoa 
and digestive fluid. Again, though no more experiments show 
such obvious irregularities, yet, on comparing Table E and K,, 
the percentages in several cases are found to differ widely in the 
two tables, though the conclusions to be drawn from the results 
would be similar in both; that is to say, the results agree in 
direction, but differ in degree. Some of these differences are 
probably improvements in the nitrogen estimation, on the results 
obtained by estimating the total organic solids. Thus the result tea 
with soda in the Table E gave a high percentage digestive power 
to the mixture of this beverage and digestive fluid—a result for 
which no reason could be given, and which is in all probability 
properly altered in Table E;. Egg digested in presence of 
ordinary mixed tea is also considerably altered in position, in 
the direction of reducing the percentage digestive power of the 
mixture of beverage and peptic fluid. Again, the smaller per- 
centage in Table E, than in Table E in the case of bread-coffee 
has already been explained. ‘The other differences between 
Tables E and E, are not very enormous, and in all respects, 
except in the above noted cases, the two tables may be taken as 
confirming one another. Table D, shows the relation between 
the total organic solid experiments and the nitrogen experi- 
ments expressed as the percentage of nitrogen present in the 
organic matter. This table shows a set of figures acting as a 
sort of check on the others, for it cannot be conceived that any 
action of the beverage can split up the nitrogenous matters of 
the meats so as to cause more or less of the nitrogen to pass 
through the dialysis paper. Therefore, when any great varia- 
tion in the percentage of nitrogen is found from that found 
when water is the beverage, error may be assumed to have 
occurred either in estimating the nitrogen or the organic matter 
in the beverage residue or in the water residue. Such very 


370 DR JAMES W. FRASER. 


irregular percentages are found chiefly in the experiments which 


have been discussed above, such as the case of bread-cocoa,~ 


where the percentage of nitrogen is stated to be 28:8, while in 
the case of bread-water it is only 4°64. Again, egg-water, the 
percentage is 16°57, while egg-mixed tea gives a percentage of 
only 9°65, showing that in all probability the amount of organic 
matter in this case was stated too high, owing to accidents the 
nature of which was discussed in the former paper. However, 
in all the more reliable results the nitrogen percentage keeps 
within limits of variation quite compatible with a fair degree of 
accuracy in both determinations. 

Examining next the average results for digestion in presence 
of the five principal beverages, which results are found in Tables 
E and E,, it is found that by the total organic solids process the 
beverages in order of action on digestion ran thus:—Water, 
100; cocoatina, 89°96; tea, 89:06; coffee, 88°79; Epps’ cocoa, 
76°05, In the organie nitrogen experiments the order is 
rather different, being—Water, 100; coffee, 87°32; cocoatina, 
87:03; tea, 85°35; cocoa, 80°71. Coffee heads the list of 
infused beverages, and cocoatina and tea each go down one 
place. But if the same experiments are selected in both pro- 
cesses for drawing the average, several experiments with each 
beverage will have to be left out in the total organic solids 
experiments, and the averages under this head will now stand— 
Water, 100; tea, 92°06; coffee, 87°56; cocoatina, 81°68; and 
cocoa, 75°8. Cocoa stands lowest in all three; but while cocoa- 
tina heads the list of averages in the first set, it sinks to the 
second place in the nitrogen experiments, and to the third in 
the last set of averages. Coffee heads the list in the nitrogen 
averages, is second in the last list, and third in the first list. 
Now, as was remarked in the former paper, cocoatina headed 
the list of averages owing to the persistently moderate results 
it gave; while tea and coffee, though they had various very high 
results, were brought down by others which were very low. 
Now it happens in the case of tea that those residues which 
were preserved were chiefly those giving high results; and hence 
in the nitrogen averages and the total organic solid average of 
the same experiments, tea takes a higher place. Coffee in this 
latter set of averages is reduced to the third place, because some 


oe. fe . 


ACTION OF INFUSED BEVERAGES ON PEPTIC DIGESTION. 371 


of its highest results, ham, salt beef, and fish have to be omitted, 
and it is brought to the head of the nitrogen averages by the 
irregular result with roast fowl. The less important beverages 
take the following order in the total organic solids experiments— 
Tea with soda, 97; cocoa nibs, 78°8; Chinese tea, 72°8; coffee 
with chicory, 62°9; green tea, 51:6; and Indian tea, 45:4. In 
the nitrogen experiments the order is—Chinese tea, 74:27; 
coffee with chicory, 69°35; cocoa nibs, 66°69; green tea, 66:14; 
Indian tea and tea with soda, 62°98. Tea with soda has in the 
nitrogen experiments to leave the head of the list and join 
Indian tea at the foot, while cocoa nibs infusion has to yield 
two places. The rest of the beverages follow the same order in 
both sets. 

Examining individual results in the cases of the principal 
beverages, it was found in the former paper that the meats 
grouped themselves, as regards the effect of the beverages on 
their digestion, into a set less injuriously affected, called break- 
fast meats, and a set more injuriously affected. In the case of 
tea, the less affected meats were—roast beef, 96:4 per cent.; salt 
beef, 94:8 per cent.; ham, 95:98 per cent.; and white of egg, 
91°7 per cent.; and the more affected—fowl, 66°73 per cent. ; 
lamb, 88°4 per cent. ; fish, 88:26 per cent.; and bread, 89°23 per 
cent. In the nitrogen experiments the higher set contains— 
roast beef, 125°71 per cent.; salt beef, 84:24 per cent.; and 
bread, 89°68 per cent.; and the lower—fish, 72°35 per cent.; 
and egg, 54°8 per cent.; while lamb, ham, and fowl are wanting. 
Here roast beef and salt beef on the one hand, and fish on the 
other, remain in the same sets in both estimations, while ege 
and bread have changed places. But bread is in both cases a 
doubtful experiment, especially in the nitrogen estimations, for 
reasons above alluded to. 

For coffee the total organic solid results gave a higher division, 
consisting of—ege, 106°45 per cent. ; ham, 100-44 per cent. ; roast 
beef, 98°8 per cent.; fish, 96°33 per cent.; and salt beef, 93-4 
per cent.; and a lower—bread, 58:27 per cent.; lamb, 69:95 
per cent.; and fowl, 86°74 per cent. The nitrogen results 
are, in the higher set—egeg, 109-1 per cent.; roast beef, 97°18 
per cent.; and fowl, 10474 per cent.; and the lower set only 
contains the suspicious bread, 38°28 per cent., here agreeing with 


372 DR JAMES W. FRASER. 


the organic solid result; salt beef, lamb, ham, and fish are 


wanting, and fowl] has passed from the lower into the higher set. 


With cocoatina the division into two classes is less evident, the 
higher class containing, in the former experiments—fish, 139°7 
per cent.; egg, 85°25 per cent.; roast beef, 88°09 per cent.; salt 
beef, 86°88 per cent.; ham, 88:96 per cent.; and bread, 87:22 
per cent.; and the lower—lamb, 77:04 per cent.; and fowl, 
66°52 per cent. In the nitrogen experiments the higher set 
contains—egg, 96°10 per cent.; roast beef, 110-42 per cent. ; salt 
beef, 83°81 per cent; and the lower—fowl, 57°82 per cent. ; 
lamb, ham, and fish being wanting, but all those which remain 
occupying the same places in both estimations. In the case of 
cocoa, no order or arrangement could be detected in the former 
experiments, and the same is the case here. The higher set-— 
roast beef, 110°9 per cent.; fish, 82°65 per cent.; bread, 464:0 
per cent.; and the lower-—egg, 59°79 per cent.; salt beef, 69°52 
per cent.; lamb, ham, and fowl do not appear. Examining the 
figures in the case of tea, coffee, and cocoatina, the notable 
differences produced by the two methods of estimation are—(1) 
white of egg having passed in the lower set in the case of tea, 
and fowl into the higher in the case of coffee. The results with 
roast beef in the cases of tea and cocoatina are much larger in 
the nitrogen than in the other estimations, but remain in the 
same division, Summing up these results, it may be stated 
that the meats least acted on in digestion by tea, coffee, and 
cocoa are roast beef and salt beef; that if ham were not want- 
ing in the nitrogen results it would probably come into the list; 
and that egg is little acted on by cocoatina, and its digestion 
positively assisted by coffee, while tea retards its digestion, 
The positions of bread and fowl in the higher list in the nitro- 
gen estimations, and of fish in the same list in the organic solid 
estimations, are doubtfully correct. Thus these new estimations 
confirm the remarks made about the older set, that the common 
use of salt beef, ham, and egg, as well as of roast beef, as break- 
fast dishes—that is at the only meal at which custom sanctions 
the mixture of infused beverages and meats—is founded, pro- 
bably unwittingly, on scientific principles. 

Experiments called “Time” experiments have been men- 
tioned above, and were described in the former paper. In these 


ACTION OF INFUSED BEVERAGES ON PEPTIC DIGESTION. 373 


the percentage digestive powers of the mixtures of beverages 
and digestive fluid were calculated from the time required for 
complete solution of a fixed quantity of white of egg. In Table 
G, column A, the percentage digestive powers thus calculated 
are shown; in column B the percentage digestive powers are 
caleulated from the “ Peptone” experiments with white of egg, 
see Table E; in column C from the averages of the peptone 
experiments ; in column D from the same experiments as in B, 
but calculated from the nitrogen estimations; and in column E 
as in O, but also calculated from the nitrogen estimations. Here 
in the case of tea the difference between the total organic solid 
and the nitrogen estimation of its action with white of egg is 
seen, while the average results with this beverage agree pretty 
well with each other and with the “Time” result. In the case 
of coffee this is reversed, the averages agreeing with each other 
but disagreeing with the Time experiment, which comes nearer 
to the estimations with white of egg only. This, of course, 
depends on the fact that coffee assists the digestion of white of 
egg, but retards more or less that of otlier meats. 

In the cases of cocoatina and cocoa the figures in each of the 
columns agree pretty well one with another, and as no nitrogen 
estimations were made with chocolate no notice need be taken 
of it. 

Summing up the results, very similar words may be used to 
those used in the former paper: (1) that all infused beverages 
retard the digestion of all meats, with five exceptions, these being 
white of egg with coffee, and perhaps fowl with coffee, and roast 
beef with tea, cocoa, and cocoatina; (2) under this general rule 
some subdivisions must be noticed ; thus tea has less action on 
roast beef and salt beef and doubtfully on bread; and coffee has 
less action on roast beef and assists the digestion of white of 
egg, and perhaps of fowl; and again, cocoatina has less action on 
the digestion of egg and of salt beef, and appears to assist the 
digestion of roast beef; while with cocoa no subdivision into 
two classes can be traced. * 

The results obtained in the former estimations, and which 
have been, as a rule, confirmed by the later detailed here, 
having been examined, some attempts were made to arrive at 
their causes by the performance of various special experiments. 


374 DR JAMES W. FRASER. 


The results of the “Cause” experiments were re-tested, in the 
suitable cases which remained, by the nitrogen process, 

One of the theoretical causes which might assist in producing 
the retardation of digestion by various of the beverages was 
the fact that precipitates formed in the digestive fluids by tea 
and cocoa (coffee gave no such precipitate) would carry down 
pepsin entangled with them (according to a well-known property 
of this ferment), and this pepsin would not be entirely re- 
dissolved during digestion. This theoretical consideration was 
put to the practical test by the experiments, the results of 
which appear in Tables H, I and I,. In Table H factors 
are shown obtained by digesting mixtures of beverages and 
digestive fluid (mixed in the proper proportions), but which 
mixtures had been filtered before digestion, thus preventing 
any resolution of the pepsin. The removal of the pepsin in 
the case of tea is equivalent to a loss ‘001 grm. of organic 
matter, or ‘0022 erm. of organic nitrogen, the loss of nitrogen 
being slightly greater in proportion than that of organic matter, 
as will be seen by comparing the percentage of nitrogen in 
this table with that in Table B. In the case of coffee, where no 
precipitate formed, the deviations are trifling and irregular, and 
depend on experimental errors. In the case of cocoa, the re- 
moval of the suspended matters of the beverage from the action 
of the digestive juice reduced the factor in column A from ‘53 
erm, to ‘412, while the nitrogen factor in column B is practically 
the same in both cases, the difference being only ‘0004 grm. of 
nitrogen, and this in favour of the filtered solution, showing 
that little if any pepsin is removed by the cocoa precipitate, 
and also that the substances extracted by digestion from the 
suspended solids must chiefly consist of non-nitrogenous matter, 
such as tannic acid, sugar, &c., this being shown by the nitrogen 
percentage being higher in the case of the filtered than of the 
unfiltered beverage. 

When white of egg is digested in presence of these beverages, 
filtered after mixing with the digestive fluid, it is found,.as 
might be expected, that but little different effect is produced 
in the case of coffee, whether filtered or not, the percentage 
digestive power in the former case being 108°33, in the latter 
109'10. In the case of cocoa a wide difference is observed 


ACTION OF INFUSED BEVERAGES ON PEPTIC DIGESTION. 375 


between the results with the filtered and unfiltered beverage, 
the digestive power in presence of the former being 209-45 per 
cent., of the latter 59°79.1_ The former large result in the 
estimation by the nitrogen process is similar in direction, but 
widely different in degree, from that in the total organic solid 
estimation, viz., 12603 per cent., but both of these depend, 
not on any action of the infusion itself, but on the removal of 
the suspended matters, which, falling to the bottom of the 
beaker used in the digestive operations, coated over the white 
of egg, and prevented the digestive fluid from acting on it. 

No nitrogen estimation in the case of white of egg, digested 
in presence of a mixture of tea and digestive fluid, filtered after 
mixing, was made, but from the agreement between the organic 
solid and the nitrogen “ Factors” for this mixture it may be 
assumed that the organic solid experiment, in the case of exg, 
would have been confirmed by the nitrogen experiment. 

The second theoretical cause, viz., the contraction by the 
gelatinous matters of the meat by the tanning action of tea and 
cocoa; and the third, viz., coagulation of any albumenoids of the 
meat by the action of any of the beverage, rest sulely on the total 
organic solid estimations, no new estimations having been made. 

The same is true of the fourth possible cause, viz., the retard- 
ing action of the volatile oils on digestion; and of the fifth 
cause, viz., the action of the alkaloids, which was shown in 
the case of theine or caffeine to be favourable to digestion. 
The sixth possible cause, viz., the precipitation of syntonin and 
peptones as formed,' by the tannic acid or its homologue 
present in the beverage, was re-tested as far as the syntonin 
is concerned, the results of both estimations being shown in 
Tables N and N,; the process, detailed in the former paper, 
consisted in mixing 25 cc. of a solution of syntonin with 
25 cc. of a digestive fluid of double strength, thus giving 
an unknown but invariable quantity of syntonin, dissolved in 
a digestive fluid of the normal strength; to this 25 ce. of 
the beverage was added, and digestion and the remainder of 
the process carried out. 

The total organic solid estimations show a great reduction of 
the amount of syntonin converted into dialysable peptones in the 

1 See Table Ej. 


376 Dk JAMES W. FRASER. 


case of tea and coffee, the amounts dialysed being only 146 
and 17:4 per cent. respectively, of what passed through when 


water was the beverage, but with cocoa the reduction is only — 


to 50°9 per cent. The nitrogen estimations, though all agreeing 
in kind with these, vary much in degree; thus the result for 
tea is a minus quantity, that is to say, not enough nitrogen was 
dialysed to supply the whole nitrogen factor in tea. With 
coffee, again, the percentage of nitrogen dialysed was 76°54 per 
cent. of that dialysed in presence of water as the beverage. This 
agrees better with what has already been shown of the action 
of coffee on digestion, and may probably be more correct than 
the organic solid estimation. With cocoa the percentage is 
only 19°34, which may or may not be more correct than the 
organic solid percentage. 

The seventh possible cause applies only to beverages contain- 
ing much albumenoid matter, and is that the accumulation of 
peptones from this albumenoid matter may arrest the digestive 
power of the pepsin. No special experiment on this was 
performed, but as confirming the principle, attention may be 
called to the figures for digestion of white of egg and beef 
in undiluted digestive fluid, Tables D to E,. The superiority in 
accuracy of the nitrogen process over the organic solid process 
of estimation is well shown in the case of these two results in 
Table E,, where, instead of disagreeing by more than 12 per cent., 
as they do in Table E, they agree within -2 per cent., a result 
which is evidently correct, as they depend on a property of the 
digestive fluid. They show that reducing the amount of water 
in the digestive fluid by one-third reduces its peptonising 
property by almost exactly the same amount. The eighth and 
last possible cause has already been referred to, and is that 
the suspended matter of beverages like cocoa clogs the action 
of the digestive fluid) As an example of this the action of 
cocoa on digestion before and after filtering may be referred to. 
Summing up the causes of the action of the beverages on 
digestion, so far as they have been re-tested by the nitrogen 
process of estimation, it is found— 

(1) That tea, as a type of all the “teas,” retards digestion as 
a rule, and acts by causing precipitates which entangle and 
carry down the pepsin, and by precipitating the syntonin as 


= 


5 


ACTION OF INFUSED BEVERAGES ON PEPTIC DIGESTION. 377 


formed in digestion. It was shown further in the former 
experiments that it was chiefly the tannic acid which had 
these properties, and that it also acted by precipitating the 
peptones as formed and by tanning the gelatine and albumenoids 
of the meats; that the volatile oil also had some action in 
reducing digestion ; and that the action of the alkaloid, if any, 
was favourable to digestion. 

(2) The action of coffee, taken as a type of the “coffees,” 
depends on the action of its caffeo-tannic acid in retarding 
digestion by various of the above actions. It has been seen 
from the nitrogen results that coffee does not act by precipitating 
pepsin, and that it has less action than the other beverages in pre- 
cipitating syntonin. When coffee assists digestion it appears from 
the former estimations that the alkaloid is the active ingredient. 

(3) Cocoa has been shown by the nitrogen experiments to 
precipitate syntonin as formed, and also, by the clogging action 
of its suspended matter, to reduce the activity of digestion. 
This latter action would in all probability not occur in the 
stomach, the active movements of which would prevent this 
stagnation. In respect of the other actions, the results of 
which were not re-tested by the nitrogen process,it is intermediate 
in activity between tea and coffee; and further, its alkaloid 
appears slightly to retard digestion. 

The first, second, third, and fourth of the conclusions quoted 
above have been reviewed, and as the fifth, sixth, and seventh 
left no results to re-test, there only remains the eighth and last, 
viz., that relating to the effects, on the actions of beverages on 
digestion, of the addition to these beverages of milk and sugar, 
in proper proportions. To the 25 c.c. of beverage, 5 e.c. of milk 
and 2°15 grain. of sugar were added. Factor experiments were 
performed without meat, their results being found in Table R, 
where column A contains the total organic solid estimations, 
B the nitrogen estimations, and C the percentage of nitrogen in 
the former results. 

It was found by the organic solids process that tea slightly 
reduced the amount of dialysable matter produced in the 
digestion of the milk as compared with the results where water 
was the beverage, coffee somewhat increased the amount, and 
cocoa increased it considerably. In the nitrogen results it is 

VOL, XX. 2B 


378 DR JAMES W. FRASER. 


found that the result with tea is slightly greater than that with 
water, and that the percentage of nitrogen is also greater, point- 
ing to the fact that the nitrogen of the alkaloid is more than 
sufficient in this to make up for any loss of albumenoids from 
the milk or the digestive fluid. In the case of coffee, the 
nitrogen result, though considerably greater than that with 
water as the beverage, still has the nitrogen percentage some- 

what smaller than with tea, though very slightly. In the case 
of cocoa the percentage of nitrogen, more than one per cent. 
higher than in any other case, is suspicious for the accuracy 
of this result. The smallness of these percentages of course 
depends on the large quantity of sugar in the dialysed matters. 

The results of digestion of white of egg with peptic fluid, in 
presence of water alone, and of the three typical beverages, with 
milk and sugar, are shown in Tables S and §,. 

The nitrogen estimations all agree in direction with those 
obtained by the organic solids process, but differ greatly in 
decree. The result with coffee confirms what was said in the 
former paper, that the addition of milk and sugar to this 
beverage reverses its action on the digestion of white of egg. 
In the case of cocoa the addition of milk increases its action 
on digestion by increasing the amount of suspended matter, 
which covers over the meat, and by increasing the amount 
of albumenoid matter in the beverage, which, by becoming 
peptonised, reduces the action of the peptic fluid on the meat. 
Therefore the minus result in this case is not without proba- 
bility. But tea was believed from the former experiments to 
have its retarding action on digestion lessened by the mixture 
with it of milk and sugar. This is not confirmed by the nitrogen 
results, and therefore the real effect of milk and sugar on the 
action of tea on digestion remains for the present doubtful. 
teverting to the conclusions of the former paper, it is found 
that the first conclusion, viz., that all beverages retard the peptic 
digestion of all albumenoid food-stuffs, is confirmed, but the 
exceptions are rather different, being in the new estimations 
white of egg with coffee; roast beef with tea, cocoatina, and 
cocoa; and fowl with coffee. Fish with cocoa ceases to be 
an exception to the rule; and ham with coffee, and fish with 
cocoatina, were not examined. 


ACTION OF INFUSED BEVERAGES ON PEPTIC DIGESTION. 379 


(2) The grouping into breakfast meats is more or less retained 
in the cases of tea, coffee, and cocoatina, but white of egg is 
removed from this group in the case of tea. Cocoa, as before, 
shows no such grouping. 

(8) This retarding action is less with coffee than with tea as 
a rale, and less with either of these than with the cocoas. 

(4) As to the causes, the only ones examined in these new 
estimations were the precipitation of syntonin by the tea and 
the precipitation of pepsin by the precipitates caused by the 
tea; the occurrence of these causes of retardation being con- 
firmed. The action of caffeo-tannin in these directions was 
shown to be slight, and the same was shown to be true with 
regard to the tannic acid of cocoa. The principal cause of the 
action of cocoa on digestion was shown to be the clogging action 
of its suspended matters. Therefore the causes of the actions 
of the beverages, as shown by the organic solids process, are 
confirmed by the nitrogen process in the cases in which they 
were re-tested. 

(8) The addition of milk and sugar to cocoa was shown to 
increase its retarding action on digestion, and the reversal of the 
action of the coffee by these additions was confirmed. But no 
confirmation was given of the result with regard to tea obtained 
in the former estimations. 

In the former paper certain practical deductions from the 
experimental results were drawn, but as these remain the same 
under the new estimations it is unnecessary to recapitulate 
them here. One conclusion, however, deserves reference, viz., 
the large loss which results from eating meat and drinking 
infused beverages at the same time. 

Sir L. Playfair quotes as a mere “subsistence” diet that of 
a London sempstress, and states that it consists of— 


Nitrogenous matter, F . 2:33 oz. or 65:95 grm. 
Fat, : : ; : . 0°84 oz. or 23°83 grm. 
Carbohydrates, ; ; . 11°69 oz, or 331°4 grm. 


the food being reckoned as dry solids. This diet, excluding 
fats, would be represented by— 
Bread, . i : ; . 19°83 oz. or 550 grm. 
Meat, ; : ; . 10°5 oz. or 300 grm. 
reckoned as they appeal as food and containing water. It 


380 DR JAMES W. FRASER. 


was calculated that in the United Kingdom, in the year end- 
ing March 31, 1882, the infused beverages consumed amounted 
to 35°3 gallons per head, or ‘75 pint per diem. This London 
sempstress would drink at any rate her share of tea, even if she 
did not greatly exceed it. 

But assuming that she only consumed this “75 pint per diem. 
It was seen by the former experiments that this quantity of tea, if 
drunk along with meat or bread, would have the effect of reduc- 
ing the peptones derived from 85 grms. of bread to what they 
would have been if only 75°8 grms. had been eaten; and those 
derived from 85 grms. of beef to what they would have been 
had only 81:9 grms. been eaten, or if fish were the meat, from 
85 to 75 grms. These calculations are based on the percentage 
results in Table E. If the results in Table E,, derived from the 
nitrogen experiments, be used in the same way, it is found that 
if bread be digested in presence of °75 pint of tea 85 grms. only 
yield the same amount of peptones as if 76°22 grms. were 
digested in presence of water. If fish be the meat, 85 grms. 
only yield the peptones of 61:40 grms. If beef be used, accord- 
ing to these latter experiments the same amount of digestive 
fluid which will in the presence of water only digest 85 grms. of 
beef, will in the presence of °75 pints of tea digest 106°85 grms. 
In the other two cases it is evident that loss occurs in eating 
the meat and drinking the tea at the same time, and even if by 
the use of tea tissue waste is prevented and food saved, yet it 
would be much more economical to take the tea on an empty 
stomach, and then when the meal time came less food would be 
needed and no waste occur. Coffees have in some cases an 
action like that of tea on roast beef, but usually they retard 
digestion, though less than tea. Cocoas retard digestion, but 
chiefly by the clogging action of their suspended solids, and this 
clogging action will probably not occur in the stomach, owing 
to its active mixing movements. 

These actions are harmful to the very poor by wasting their 
food, and can only be beneficial in the case of those who 
habitually eat too much nitrogenous matter, which, if absorbed, 
might overstrain the kidneys and perhaps cause gout or kidney 
disease. For such, tea would probably be the best beverage. 
The beverages, as shown by their factors, and especially when 


> 


ACTION OF INFUSED BEVERAGES CN PEPTIC DIGESTION. 381 


taken, as they usually are, with milk and sugar, are the means 
of conveying some considerable amount of nourishment into the 
body. The breakfast meats above mentioned should be those 
eaten when infused beverages are drunk, and this rule appears 
generally carried out, whether by some instinct or by experience 
cannot be said. The best time of the day for drinking these 
infused beverages is undoubtedly about 5 p.m. when the lunch 
has been digested, and the cup of tea or coffee carries the energies 
on till dinner time. 


APPENDIX. 


The Tables are similarly lettered to those which appeared in the 
former paper. 


TABLE B. 


Results of digestion of 50 c.c. of digestive fluid alone, with 25 c.c. 
of water, and with 25 c.c. of each beverage. 

‘A. Total organic solid estimations. 

B. Organic nitrogen estimations. 

C. Percentaze of | nitrogen in organic solids. 


jai) eal 


Per cent. | Per cent. | Per cent. 


Digestive fluid undiluted, 5 : : "192 0146 76 
is »» with water, . : , "192 0146 76 
a » With tea(mixed) .  . 26 0243 10°8 
FS » With tea (Chinese), . : 272 "0199 731 
; » with tea (Indian), . . 318 0187 5°88 
PS », With tea (green), . ; "413 0217 5°25 
5 » with tea with soda . ; "259 0248 9°57 
4 », With coffee,  . 269 0258 9°59 
os » with coffee with chicory, . ; 336 0226 6°72 
3 » With cocoatina, . ; i 237 0174 7°33 
35 » With cocoa, E . : 530 0297 5°60 
», With cocoa nibs, ' : 297 0320 10°77 


[Tape D. 


382 DR JAMES W. FRASER. 


TABLE D. 


Results of the peptone experiments. 


Ey 3 cele = I = a 
sa/8 [32/8 ]/e}/e 2] 
me atele| Ms 3 
So a S43 a = a oO =] 
Rx s As ES rst 3 = a 
g = 
cf ce mM >) [S) S S 
= [=] (=a) a Q 
Grms.| Grms.| Grms.| Grms.] Grms.| Grms.| Grms.| Grms. 
Digestive fluid undiluted, . 9 . | °376a] *771b) .. ae e ae as ee 
50 with water, . - . | *4£34a} 1°0426] 1°121 | -649 | °897 | *920 | °792 | °446 
ee with mixed tea, . . | 7398 | 170056) 1°063 | -574 | *861 | *614 | *699 | °398 
aa with Chinese tea, . “B17 = ao oe aa = 
By with Indian tea, . anima 
4 with green tea, . . | °224 
with tea with soda, Sl eral 
es with maté, . . | *359 ah oe a8 Ae ste 
x with coffee, : -462a| 1-03c | 1-047 | -454 | -901 | -798 | -763 | -26d 
“3 with coffee and chicory. 273 ae ae oe Bo a 
= with Arab coffee, . 2 | °295 55 3- ee ze % 
“5 with cocoatina, . . | -390 | -918c] -974] -500 | -798 | -612 |4-107 | -389 
SS with cocoa, . . | 164] *815c} -973) -542 | -628] .. 812 | -330 
‘ with chocolate, . . |-"005] .. *588 | -407 | °648 | -795 | *700 |-"05: 
na with cocoa nibs, . Sil) $248 te ore ae és oe = oe 30 
ny with guarana, “ . | °224 | 


Results marked (a) belong to a different set from the rest of the experiments with white of 
egg. Results marked (0) belong to a different set from those marked (c) in the experiments with 
roast beef. 


(@) Charring took place in the evaporation of the dialysiate in these cases. 


TABLE D,. 


Results of peptone experiments estimated as organic nitrogen. 


bo : a j a ; 
S13 |oa) | 2 | Siege 
~- |@& |22| 8 \)8 |e) ee 
; | ¢ |ee| 2 | 2 
2 | 8 |93) 2 2 |G 
|e Ce) eo ee 
Siry p F Grms.| Grms.| Grms.| Grms.| Grms.| Grms.| Grms.| Grms. 
Digestive fluid undiluted,. . . .| -0519|-1084 = 
af with water, . «| 0719 | +1497 | -2101 ». | +1305 | -1306 | -0206 
9 with mixed tea, . | ‘0394 | -1882 | -177 of aa .. | 70945 | -0185 
7 with Chinese tea, . .|°0534| .. - ca Bi ne ale 
.; with Indian tea, - | 0453 F 
> with green tea, - «| "0476 ov 
is with tea with soda, . | 0453 ee Es ee ‘3 oi a 
s with coffee, . 0785 | "1455 | .. |.0999| -. | *E405)" cS 0079 
5 with coffee with chicory, 70499 | .. $5 x ae 
49 with cocoatina, , “0691 | 1653 | °176 | *1146] .. | :0780 
+5 with cocoa, + + «| 70430 | 1660 |°1461 | 0965] .. 2 ‘1079 0958 


as with cocoa ‘nibs, - «| 0480 


4 


. 


ACTION OF INFUSED BEVERAGES ON PEPTIC DIGESTION. 383 


TaBiE D,. 


Percentage of nitrogen in the total organic solids results. 


White of Egg. 
Roast Beef 
Boiled 
Salt Beef. 
Roast Lamb. 
Boiled Ham. 
Roast Fowl. 
Boiled Fish. 
Bread 


Per | Per | Per | Per | Per | Per | Per | Per 
cent. | cent. | cent.| cent. | cent.| cent.| cent. | cent. 


Digestive fluid undiluted, . . ./13°81/14:05| .. Be ae es - 
9 with water, . .  . | 16°57|14:36|18°75| .. .. | 14°67] 16-48] 4-64 
a with mixedtea, . . | 9°65 | 17°17 | 16°65 ‘S a re 13°5 4°65 
ee with Chinese tea, . . | 16°84) .. at ne = oe a Pe 
% with Indian tea, . . | 23°00 
s with green tea, . . | 21°20 
be with tea with soda, a ROs@Gs| ae 7" Be Se “8 me ts 
& with coffee, . 16:99) TAT) Pe 2099. cy APB 3°03 
a with coffee with chicory, 18:31} 2. ne a2 oe ee op .. 
a with cocoatina, . 17°71 | 18°00 | 18-05 | 22:92} .. |12°75] .. ae 
— with cocoa, . e - | 26°24 | 20°07 | 15:00 | 17°80 ae 3c 13°29 | 28°8 
% with cocoa hibs, i ee L957E || 2. a is Ss ag es ¥ 

TaBLe EF. 


Percentage results of the peptone experiments. 


i ee a 

aad ce 0 ee i ee a ee 

So | a Se tas & a 3 BR 5) 

a=) 2 = nl 3 a 3 S = is) 4 

= & f=} fea} & ==) 
Per | Per | Per | Per | Per | Per | Per Per Per 
cent. | cent. | cent.| cent.| cent.| cent. | cent. cent. cent. 
Digestive fluid undiluted, . Fie TOTO et iis as “3 + - of 80°3 

< with water, . : .|100 {100 |100 |100 |100 {100 100 100 100 
a with mixed tea, . - | 91°7 | 96°4 | 94:8 | 884 | 95°98} 66°73] 88:26} 89:23 | 89-06 
ey with Chinese tea, . . | 72°8 Bs ar Ae i 5 ae ne 72°8 
re with Indian tea, . - | 45°4 a os “ic a = a ae 45°4 
5 with green tea, . . | 51°6 es a ve an os fa aE 51°6 
a with tea with soda, - | 97 oe ae Ap 32 “ ae ce 97 
= with maté, . F - | 82°7 =f Se oF fs a a 82°7 
a with coffee, . : » 106-45 | 98°8 | 934 | 69°95 |100-44| 86-74 | 96-33 58°27 | 88°79 
a with coffee and chicory, | 62:9 Pi ¥ ee Sq Be - r 62°9 
= with Arab coffee, . = SOG | es “E ae aie 4 ae be 67:97 
+e with cocoatina, . . | 85°25 | 88:09 | 86°88 | 77°04 | 88°96 | 66°52 13977] 87-22 | 89:96 
“ with cocoa, . -  .| 87°78] 78°2 | 86:8 | 83:35) 70 -- {102°5 73°76 | 76-05 
= with chocolate, minuelt .. | 52-45| 62-71) 72-25] 86-41| 88:38 mae 72-44 
x with cocoa nibs,  . - | 78°8 et a ea Je Yc ae oe 78'S 
As with guarana, : Pa oeoy G3 * 3 ae fs Ee a BF os Be 57:14 


$n 


384 DR JAMES W. FRASER. 


TABLE E,. 


Percentage results of peptone experiments estimated by nitrogen 
process. 


o 
) ~ o eSsina ~ = fais ol > 
Hw] Ba (Pad! Se | Sh | ee | se] = | s 
Go of = 6S = ae 2 nd 
E Blea |g@o|e5| en | ee | see eee 
Per |-Per | Per | Per | Per | Per |-Per | Per | Ber 
cent. | cent. | cent. | cent, | cent.} cent. | cent, | cent. | cent. 
Digestive fluid undiluted 12°24 | 72:42). an eS < a oe 72°31 
‘i with water, > =» 200) 1400 ROOF .. {100 {100 {100 {100 
“ with mixed tea, : 54°80 |125°71 | 84:24 | .. Br ed 72°35 | §9'°68 | 85°35 
; with Chinese tea, . TAT ee eS oF ae Le ce oe, | S42e 
ay with Indian tea, . 62°98} .. Ae xe sc ac “5 a 62°98 
59 with green tea, : 6614} .. #0 Woe Be ae a8 ae 66:14 
ss with tea with soda, 62°93) .. AG a He aye ts eo 62°98 
“1 with coffee, Ste 2 I LOOTO} ESTES 55) .. |10474) .. | 38:28 | 87-32 
Be with coffee and chicory,| 69°35) .. 3 é sf #s Re 69°35 
7 with cocoatina, a 96°10 |110:42] 83:S1 | ..a BTs82i\ eee .. | 87:03 
= with cocoa, . . 59°79 1109 |6952 | ..a 82°65 |464:0b) 80°71 
of With cocoanibs, . | 6669] .. Se : ae .. | 66°69 


a, Absent because there is no water experiment for comparison. 
6. Omitted in calculating average. 


TABLE G. 


Percentage digestive powers of mixtures of digestive fluid and 
beverages deduced—A, from time experiments; Bb, from peptone 
experiments with white of egg; C, from average of peptone experi- 
ments; D, from nitrogen experiments with white of egg; and HE, 
from average of nitrogen experiments. 


A. B. C. D. E. 
Per cent. | Per cent. | Per cent. | Per cent. | Per cent. 
Digestive fluid with water, . . 100 100 100 100 100 

7 with mixed tea, . 89°26 91:7 89°06 54°80 85°35 
< with coffee, ,. . 115-02 106°45 88°79 109-10 87°32 
ap with cocoatina, . 72°04 85°26 89°96 96°10 87°03 
35 with cocoa, . . 89°26 37°78 76°05 59°79 80°71 
3 with chocolate, . 76°33 ate 72°44 Av aie 


[Tanie H. 


- 


ACTION OF INFUSED BEVERAGES ON PEPTIC DIGESTION. 385 


TABLE H. 


Results of digestion of 50 c.c. of digestive fluid with 25 c.c. of the 
three principal beverages, filtered after mixing. A, Total organic 
solid estimations; B, Organic nitrogen estimations; C, Percentage of 
nitrogen in organic solids. 


Tne B. C. 


Grms. Grms. Per cent. 
Mixed tea, . : : “225 ‘0221 10 
Coffee, P : 5 "263 0263 10 
Cocoa, , : , "412 ‘0301 73 
TABLE I, 


Results of digestion of 5 grms. white of egg with 50 c.c. of digestive 
fluid, and 25 c.c. of beverage, filtered after mixing. 


Actual Weights. | Percentages. 
Grms. Per cent. 
Water, . : ‘ : : 434 100 
Mixed tea, . : : : ‘34 78°34 
Coffee, . : ‘ : : 462 10645 
Cocoa, . : : : - 547 126°03 
TaBLe I. 


Results of digestion of 5 grms. of white of egg with 50 c.c. diges- 
tive fluid and 25 c.c. of beverage, filtered after mixing. Results 
estimated by nitrogen process. 


Actual Weights. Percentages. 
Grms. Per cent. 
‘07195 100 
vane T: 07795 108°33 
z F z 5 f "1506 209°45 
TABLE N. 


Digestion of syntonin in presence of water, and of the three principal 
beverages. 


Actual Weights. Percentages. 
Grms. Per cent. 
Water, . E F : d "1206 100 
Mixed tea, . ; 5 3 ‘0176 14°6 
Coffee, . : F “ ’ 021 17°4 


Cocoa, . : : ; ‘ 0615 50°9 


386 DR JAMES W. FRASER. 


Tas.e N,. 


Digestion of syntonin in presence of water and of three principal 
beverages. Estimated by nitrogen process, 


Actual Weights. Percentages, 
Grms. Per cent. 
Water, . 2 ; : 4 0162 100 
Mixed tea, . . : : — 00649 “ae 
Coffee, . . ; , 5 0124 76°54 
Cocoa, . ; E : : 0031 19°34 
TABLE R. 


Results of digestion of 25 ¢.c. of beverage, with 5 c.c. of milk and 
1:25 erm. of sugar, in presence of 5 c.c. of digestive fluid. A, total 
organic solid estimations; B, organic nitrogen estimations; C, per- 
centage of nitrogen in organic solids. 


A. C. 
Gris. Per cent. 
Water, : : : 1345 2°29 
Mixed tea, Bee 1-275 2:54 
Coffee, : : . 1:413 2°52 
Cocoa, : : 1677 3°69 
TABLE S. 


Results of digestion of 5 grms. of white of egg in presence of water, 
and of the beverages with milk and sugar. 


Actual Weights. Percentages. 


Grms. Per cent. 
Water without milk, . 4 "434 100 


Tea with milk and sugar, . "426 98°15 
Coffee with milk and sugar, . ‘237 54°6 
Cocoa with milk and sugar, . 04 9:2 


ey 


= Pee def 


al 


- 
ACTION OF INFUSED BEVERAGES ON PEPTIC DIGESTION. 387 
TABLE §). 


Results of digestion of white of egg in presence of water, and of the 
beverages with milk and sugar. Estimated by nitrogen process. 


Actual Weights. Percentages. 
Grms. Per cent. 
Water without milk, &., 07195 100 
Tea with milk, &., : 0189 26°31 
Coffee with milk, &., .  . 0154 21:37 


Cocoa with milk, &e., . : — 00546 


SOME VARIATIONS IN THE HUMAN SKELETON. By 
W. Arputunot Lane, M.S., F.R.C.S., Senior Demonstrator 
of Anatomy, Guy’s Hospital, Assistant Surgeon to the Hospital 
for Sick Children. 


Asymmetry of the Spinal Column, Chest, and Skull; Bifid Ribs 
and Cartilages ; Extent of Lower Limit of Pleura, &e. 


I WILL give a detailed account of these cases, and note the varia- 
tions from the normal as they occur, discussing any points of 
interest in connection with them. 


The first case was a man about 5 feet and 8 inches in height, with 
a chest of average size. His sternum was abnormally thick and broad. 
The greatest breadth of the manubrium was 3} inches, and the least 
2} inches. The greatest breadth of the gladiolus was 1? inches. The 
length of the manubrium was 22 inches, that of the gladiolus 4 inches. 
They were united to one another by a thin layer of dense fibrous tissue, 
the joint so formed allowing of very slight movement. 

The first ribs were almost exactly symmetrical in size and in posi- 
tion. The span of each was 34 inches. The first cartilage was placed 
a little more horizontally than usual ; its antero-posterior diameter was 
unusually great, and it was more than normally flattened on its upper 
surface. It was ossified in almost the whole of its thickness. In each 
an amphiarthrodial articulation was present. On the right side it was 
situated in its usual position, namely, half an inch from the extremity 
of the rib; and on the left side it was placed in the centre of the 
ossified cartilage. The other costal cartilages were quite free from any 
ossific change. Owing to a slight obliquity of the articulation between 
the two pieces of the sternum, the left second costal cartilage was 
attached to the sternum at a slightly higher level than the right. 

The third right costal cartilage was nearly normal in depth at its 
attachment to the sternum. As it passed outwards it rapidly increased 
in vertical measurement, and just before its junction with the corre- 
spondingly enlarged extremity of the third rib it was perforated by a 
pemen 2 inch in diameter. At this point the cartilage was 14 inches 
deep. 

The third left costal cartilage was about # inch deep, and was 
attached to the sternum a little above its fellow. 

The right fourth cartilage was normal throughout, 

The left fourth cartilage was 14 inches deep at its inner end, where 
it was perforated by a foramen half an inch in diameter. Its depth 
diminished as it passed outwards, though it had not reached its normal 
measurement where it joined the enlarged extremity of the fourth rib. 


= A aa 


- aitiiealaedaai 


SOME VARIATIONS IN THE HUMAN SKELETON. 389 


The right fifth cartilage was nearly an inch deep in the greater part 
of its length, though at its inner end it was somewhat narrower. 

The left fifth cartilage was 3 inch deep, and attached to the sternum 
above its fellow. 

The right sixth and seventh cartilages were normal, and articulated 
with the sternum. 

The left sixth cartilage was 14 inches deep at its inner end. It 
increased considerably in breadth as it passed outwards, and at a point 
1} inches from the sternum it split into two cartilages, which enclosed 
a foramen measuring 34 inches in its transverse diameter. These two 
rods of cartilage united with the bifurcated broad extremity of the 
sixth rib, which itself bounded the foramen. 

The seventh left cartilage was broader than usual and articulated 
with the sternum. A point of some clinical interest was the very 
narrow interval in the front side of the chest between the left sixth 
rib and cartilage and the adjoining ribs and cartilages, especially the 
seventh. This circumstance would have caused some inconvenience 
in the event of the ordinary operation for an empyema being per- 
formed here, as it would have been impossible to drain the cavity 
through such a narrow interspace without resecting a portion of the 
rib. 

The left costal cartilages articulating with the sternum were, with 
the exception of the first, at least a quarter of an inch longer than those 
of the right side. 

The bones forming the spinal column were dense and strong. They 
presented no lateral curve or extensive pressure change. The upper 
dorsal vertebree were thrown slightly forwards. There were no old 
pleuritic adhesion or chronic lung affection. There was the same con- 
dition of the vertebra which was present in a case I described in the 
last volume of the Jour. of Anat. and Phys., “ Vertebral Asymmetry,” 
&c. Up to the present time that case was the only instance of verte- 
bral asymmetry I have seen in the dissecting room which was not due 
to pressure cr costalleverage, In this case the left transverse processes 
were directed in a more upward and backward direction than usual, so 
that on this side there was a narrower groove between the transverse 
processes and spines than on the opposite side. The difference on the 
left side, on an average, was about half an inch in the middle eight 
dorsal vertebra, and about an eighth of an inch in the upper two dorsal 
vertebre. The left were also on a higher level than the right. This 
difference in level in the upper and central part of the dorsal spine 
yaried from about half an inch to about a quarter or an eighth of an 
inch in the lower dorsal vertebre. 

Chiefly on account of the alteration in the direction of the left 
transverse processes, but slightly also owing to a diminution in the 
length of the ribs between the angles and heads, their angles of the 
ribs on the right side were more distant from the middle line than 
those on the opposite side. This difference amounted in some to 
half an inch, and in others to three-quarters of an inch. In this 
chest we have, besides a certain amount of asymmetry in the attach- 


390 MR W. ARBUTHNOT LANE. 


ment of the costal cartilages to the sternum, asymmetry in the form 
and length of the cartilages and ribs, and of the laminz and trans- 
verse processes of the dorsal vertebre. 

One would have expected the cartilages on the left side to be 
the shorter, as they are attached to the sternum at points higher than 
their fellows. This asymmetry of the two sides of the chest is very 
interesting, both from a clinical as well as from an anatomical point of 
view ; and, as I have already shown in previous papers, it is by no 
means an uncommon condition. 


I would here protest strongly against the clinical habit of 
gauging too accurately the size of the heart by the relation 
borne by the apex-beat to the nipple and to the interspace. 
The chest wall is frequently so variable in its bony framework 
that, in a doubtful case, its constituent parts should be carefully 
defined and measured, and a broad margin allowed for the posi- 
tion of the healthy apex, should the framework of the chest 
deviate to any great extent from what is regarded as the normal, 
or to speak more accurately, the more common type. The more 
I examine the bodies of subjects in the dissecting room, the 
more am I convinced of the importance of this fact, and | 
do not think it is sufficiently recognised, if it is so at all, by 
clinical observers. 

It is also by no means uncommon to find considerable 
asymmetry of the skull. In the case that I have just described the 
skull was nearly symmetrical, the left half being slightly the 
larger of the two. It is curious that the cavity of the skull 
should be so frequently asymmetrical, and yet it is not so difficult 
to attempt an explanation in the case of the skull as it is in the 
case of the framework of the chest. I have examined and 
measured a number of skulls, and in the large majority I have 
found the left half to be larger than the right. I measured 
them by stretching a string from the skull immediately above 
the crista galli to a point immediately above the centre of the 
longitudinal sinus at the internal occipital protuberance, the 
skull cap having been removed in the usual way. At right 
angles to this, other cords were stretched, and the distances from 
the middle line measured As a rule, the increase was tolerably 
general on one side, but in many cases it was limited to the 
front or back of the skull. Given that one half of the cavity is 
larger than the other, I think we may assume that the half 


Se. 


é eatin Tore Om od lens 


z 


SOME VARIATIONS IN THE HUMAN SKELETON. 391 


of the cerebrum contained in it is also larger than the other 
half. 

As the right limbs have for generations been used much more 
extensively than the left, not only in obedience to very com- 
plicated efferent impulses, but also, in the hand especially, for 
the purpose of gathering afferent impulses of a very complex 
nature, it is I think fair to suppose that, as the result of this, the 
left half of the brain has become larger than the right. I hope 
soon to have more accurate data as to the relative frequency of 
the side and portion of the side enlarged. 

In torticollis occurring at an early period of life, we see what 
appears to be an atrophic ora less developed condition of the 
head and face on the affected side. This does not seem to be 
explicable on the same physical basis as are the curves and 
changes produced in the spinal column, and the consequent 
alterations in the form of the chest. I have not yet been able 
to examine the skeleton of a subject affected by torticollis 
accompanied by asymmetry of the skull and face, though I was 
fortunate in being able to examine the bodies of two adults in 
whom the cranial changes were not present. One is described 
in the Zvansactions of the Pathological Society, 1885. 

During four years I have seen three other bodies in which 
bifid ribs or cartilages were present. In two of these the left 
fourth cartilage and rib were affected, but of the third I have 
lost the notes, and only remember that one rib, or one rib and 
cartilage were affected. Professor Struthers, in his paper! in the 
Jour. of Anat. and Phys., describes five cases of this condition. 
In three of these the abnormality was ascertained to be of the 
fourth rib, another probably of the fourth, the other two probably 
either of the fourth or fifth. In three it occurred on the left 
side only, in one on the right, in one on both sides. 


I have also seen two subjects in which the costal cartilages were 
bifid, but the upper limb of the divided cartilage ended in a free 
rounded extremity, and did not, as in the previously described cases, in- 
clude a foramen. The fourth costal cartilage was the one affected in 
both cases. In the first, which was a female subject, the manubrium 
was 21 inches long, and the gladiolus 3} inches. They articulated 
with one another by an amphiarthrodial joint. The right fourth costal 
cartilage was 3 inch broad at its attachment to the sternum. At abcut 


1 “Variations of the Vertebre and Ribs in Man,” vol. ix. 


392 MR W. ARBUTHNOT LANE. 


half an inch from its inner extremity it split into two, the lower 
division taking on the functions of the normal fourth cartilage; the 
upper division, which was 1 inch long, ending in a free rounded 
extremity. The breadth of the fourth rib was increased abruptly at a 
point about 1 inch from its anterior extremity, showing a tendency 
to bifurcation of the rib also. 

In the second case, which was a male subject, the manubrium was 
12 inches long and the gladiolus about 4 inches. These bones articu- 
lated by an amphiarthrodial joint. The left fourth costal cartilage was 
more than # inch deep at its inner extremity. It bifurcated in a simi- 
lar manner to the last, the rib being, however, normal in form and 
outline. The free end of the upper division terminated abruptly in 
the intercostal muscles as in the preceding case. 


Another point I would call attention to is the relative 
lengths of the clavicles on both sides. I find it stated in 
Gray's Anatomy that the right clavicle is frequently shorter 
than the left. I have measured a great many, and have not 
been able to verify this statement, either in people who had 
or who had not led laborious lives, nor do I see for what 
reason the right clavicle should be shorter than the left. 
Long before the clavicle would yield and alter its form under 
the influence of pressure, the first rib would have yielded to 
a relatively greater extent, and in it even a slight depression 
is rare. I have described fully the manner in which force is 
transmitted from the arm and shoulder to the chest wall in 
papers contained in the Zvransactions of the Pathological Society, 
1884-1885, “ Case of Costo-Chondral Dislocation,” “ One Mode 
of Fracture of the Sternum,” and “Mode of Fracture of First 
Rib alone,” so will not do more than refer to them here. 


Whenever I have found the first costal arch equal on both sides, as 
is usually the case, the clavicles have also been symmetrical, but when 
the span of one arch was larger than its fellow, the clavicle on the 
same side was correspondingly so also. Asa very marked instance of 
this, I would refer again to the cases already mentioned as having been 
described in the paper on ‘“‘Supernumerary Cervico-Dorsal Bearing 
Ribs,” in which the span of the costal arch on which the right 
clavicle rested was 32 inches, while that crossed by the left clavicle 
was only 3 inches. The right clavicle was 6 inches long, and the left 
51 inches. ‘This is a considerable and an unusual difference, but the 
difference between the costal spans was also great. 

Returning to the ease I was describing, the clavicles were equal in 
length. ‘Their inner extremities were peculiar in that they were much 
flattened on their under surfaces, the articular surfaces on their inner 
aspect being continuous, with large flat facets on the under surface. 


Se 


SOME VARIATIONS IN THE HUMAN SKELETON, 393 


This last articulated with a correspondingly well marked depression on 
the upper surface of the first cartilage. The ligaments connecting the 
clavicles to the sternum and first costal cartilage were very well 
developed, and this was particularly the case with the interclavicular 
ligament, which was very thick and strong, and had hardly any con- 
nection with the sternal notch. There was no rheumatoid change in 
this or in any other articulation in the body. In the resting position, 
the clavicle lay on the first cartilage at a point half an inch internal to 
the attachment of the first rib, being much within the normal point of 
crossing. Also, on pressing back the shoulder, the clavicle could not 
be made to press on the rib where it received the insertion of the 
scalerius medius. This was due in part to the remarkable change in 
the form of the sterno-clavicular articulation, and in part to the short- 
ness and great strength of the rhomboid ligament, and to the slight 
difference in the direction of the ossified cartilage. 

On the left side the subclavius muscle was entirely absent, and 
the costo-coracoid membrane was ill developed. On the right side 
both were fairly well marked. In this case I did not see this part of 
the body till this region had been partly dissected, so cannot testify 
with absolute certainty to the complete absence of the left subclavius 
muscle, I was then unable to find any relic of it, and was quite satis- 
fied that, if it had been present, it must have been very small indeed. 

The coraco-humeral ligament on the right side was replaced by the 
tendon of the pectoralis minor. The left was normally attached. 
The right coraco-brachialis arose by a large fleshy belly from the 
whole length of the coracoid process as well as by the tendon common 
to it and the biceps. 

I mention these abnormalities as I have frequently noticed that, 
when you get an abnormal condition of the thoracic wall, you not 
uncommonly get other parts varying also, especially structures about 
the clavicles and shoulders. I would refer again to a case I described 
in which there were thirteen cervico-dorsal vertebra bearing ribs. In 
that there were on both sides chondro-epitrochlearis and three headed 
biceps muscles, besides several other muscular abnormalities. In the 
ease which I have already referred to twice, in which there were appa- 
rently eight cervical vertebra, the omohyoids had each only one fleshy 
belly, the anterior, and this arose by an aponeurosis from the clavicle. 
The subclavius muscle was also absent on both sides, 


Simple costal and sternal asymmetry may be regarded as a 
condition of retrocession to the arrangement found normally in 
the orang-utans, and Professor Turner has demonstrated the 
fact that in Cetacea cervical ribs also exist not unfrequently, 
that they may fuse with the first thoracic rib, and that the 
upper two thoracic ribs may also unite, though not with the 
same frequency. Similar variations in the number of the ribs 
and in their degree of development have been described by Pro- 

VOL. XX. 2¢ 


394 MR W. ARBUTHNOT LANE. 


fessor Struthers among other animals. I have not succeeded 
in finding instances of bifid ribs or cartilages in any animal 
except man, but my experience is limited to museum specimens 
alone. . 

The next important point I will allude to is a very remark- 
able change in the outline of the shoulder-joint, and consists in 
an alteration in the relation of the head of the humerus to the 
acromion process. I need not do more than refer to the normal 
form of the shoulder as seen in people who do not lead very 
laborious lives. Its rounded shape is unbroken by any pro- 
minence of the acromion on its posterior aspect, or of the 
humeral head on its anterior aspect. The head of the humerus 
is sheltered by the acromion above, the small interval being 
filled in by the deltoid. 

In some subjects, who have-led hard laborious existences, 
I have found that the head of the humerus did not occupy the 
normal relative position to the acromion, and that the direction 
of the glenoid cavity was more anterior than usual. This 
modification in the form of the shoulder-joint does not appear to 
be limited to subjects whose spinal columns presented any par- 
ticular labour curve. 


The alteration consists in a forward displacement of the head 


of the humerus, so that it is no longer covered by the acromion 
to the same extent as before, and in the variation of the direction 
of the glenoid cavity from the normal, which I have just described. 
I was inclined to regard these changes as the product of strain 
thrown upon the articulation in certain employments. 

The shoulder-joint of this subject displayed the above con- 
ditions in an extreme degree, so much so, that before the 
removal of the skin the appearance presented was that of 
double subcoracoid dislocation, in which a new socket had 
formed for the head of the humerus, and this socket had 
trenched considerably on the glenoid cavity. The anterior 
extremity of the acromion projected outwards behind the centre 
of the head of the bone, and the posterior part of the acromion 
projected back some way beyond the humeral head, so as to 
produce a considerable interval between these points. This 
was even more marked on the left side. The direction of the 
glenoid cavity was altered very distinctly. 


o 


SOME VARIATIONS IN THE HUMAN SKELETON. 395 


The man was powerfully built, and his muscles were well 
developed, especially the deltoid and trapezius. 

The question now arises as to the mode of origin of this 
condition, Is it the result of the occupation of the individual 
or of some fault in deportment, and produced in the same way 
as pressure changes elsewhere? In a subsequent paper * I hope 
to show that the shape of the bead of the femur and the form 
and extent of the acetabular cavity is modified very considerably 
by the mode of labour of the individual, and that there can be 
no reasonable doubt in that case as to the manner of its pro- 
duction. In it the result produced by pressure is similar to 
that present in this shoulder-joint. I might also refer to changes 
in the direction of other joints as the result of pressure, which 
I have described in a paper in the last number of the G's 
Hospital Reports, “ Senile Changes,” &c. 

Or is this condition of the shoulder-joint one that is peculiar 
to the individual and congenital, as are his bifid ribs, &., and 
due to some cause of which we have no knowledge ? 

Though the vertebrze were strong and dense, this man’s spinal 
column presented no pressure curve, except a slight forward 
curve of the upper dorsal vertebree and an increase in the size 
of the lumbar and sacral spinous processes. His first ribs, as 
well as their ossified costal cartilages, were particularly strong, 
The sternal articulation was broad and firm. As I said before, 
the ligaments in connection with the clavicle were very strong, 
the interclavicular having but slight connection to the sternum. 
The clavicles also rested on the first cartilages and not on the 
ribs, and the line of direction of the cartilage and rib was altered 
from the normal. 

Each of these circumstances taken separately does not prove 
much, but all taken together suggest strongly that the man had 
been engaged in some occupation in which he was in the habit of 
carrying heavy weights simultaneously in both hands, and that 
the conditions present resulted from the long-continued pursuit 
of this form of labour. Observing the different classes of 
labourers, one sees that there are but few whose function it is to 


1 «« Pressure changes in the joints of the extremities, including senile changes,” 
Transactions of the Pathological Society, vol. xxxvii., and ‘‘Dupuytren’s con- 
traction,” &c., Guy's Hospital Reports, vol. xliii. 


396 MR W. ARBUTHNOT LANE. 


carry weights for any long time in this way, though of course 


many combine this largely with others. Perhaps milkmen more - 


than others would seem to supply the necessary requisites for 
the conditions present in this skeleton. 


Fusion of the First and Second Costal Cartilages on the 
Left Side. 


I am describing this specimen as it is a good instance of an 
unusual condition, and also because its characters will be seen 
to deviate slightly from those usually present in these cases, 
in regard to the span of the upper ribs, and the relative 
measurements of the manubrium and gladiolus. In a paper in 
the Guy’s Hospital Reports, vol. xlii., “ Cervical and Bicipital 
Ribs in Man,” I gave descriptions of three instances of this 
form of abnormality, and the chests of these subjects were 
characterised by the relative diminution in the capacity of their 
upper part, including the manubrium, and by the relative 
increase in that of the lower part, including the gladiolus, 
and in the number of ribs articulating with that bone. 


In the first case (fig. 6) the manubrium was 1? inches long, the 
gladiolus 43 inches, and eight cartilages articulated with the sternum 
on the side opposite to that on which fusion was present, namely, 
the left. 

The cartilages articulated symmetrically with the sternum, except 
below, where the sternum itself deviated to the right. The first and 
second cartilages were fused through at least an inch of their 
extent. 

In the second case (fig. 8) the right first and second costal cartil- 

ages were fused along 2-inch of their length. 

The span of the first rib was 2,3, inch, the manubrium was under 

2 inches, and the gladiolus over 4 inches. 

On either side eight ribs articulated symmetrically with the sternum. 

In the third case both first and second costal cartilages were fused 
in half an inch of their extent. The manubrium was ik inches, and 
the gladiolus 4} inches long. There were on either side eight costal 

cartilages articulating asymmetrically with the sternum. As one would 
expect, the first and second alone of the upper costal cartilages ever unite. 

This new specimen existed in the body of a well-built male subject 
with a good large chest. 

The greatest span of the right first rib was 32 inches. That of the 
left was 3} inches, 

The span of the right second rib was a third of an inch greater 
than that of the left. 

The manubrium was 2 inches long, the gladiolus 41 inches. 


cy YP ny 


Pade 


eine tc 


SOME VARIATIONS IN THE HUMAN SKELETON, 397 


There were seven ribs articulating symmetrically with the sternum. 

The twelfth rib was 6 inches long. 

The first and second right costal cartilages were normal in their 
form and arrangement. The first was ossified nearly throughout, and 
presented an arthrodial articulation in the normal situation. 

The two upper left cartilages were fused in the inner inch of their 
extent, five-eighths of the first being free and half an inch of the 
second. The fused cartilage, as in fig. 6 referred to above, only 
articulated by a small surface with the gladiolus. 

I would call attention to the extent to which this fused cartilage 
has undergone ossification. That part of it formed by the first costal 
cartilage was completely ossified, and presented an arthrodial joint at 
its outer extremity, that portion which was not fused articulating 
with the fused part. The line of ossified cartilage crossed obliquely 
downwards and inwards that part of the fused cartilage corresponding 
to the second costal cartilage, the remainder of it being free from any 
osseous change. There was no ossification of the other costal cartilages. 
Though the whole of the fused cartilage was subject to the same 
influence in respiration as was the first costal cartilage on the right 
side, yet, as we have seen, only that portion of the fused cartilage cor- 
responding to the jirst costal cartilage underwent any osseous change, 
and the process of ossification was so far advanced as to necessitate 
the presence of a loose arthrodrial articulation. 


In previous papers! in the Zrans. Path. Soc. I have de- 
scribed the mode of formation of this joint in the ossified 
cartilage, first in the sternum, and in the articulations con- 
nected with it, also the joint formation in the less frequently 
ossified lower cartilages. I ascribed the joint formation in the 
first cartilage as being due to two factors, the movements of re- 
spiration and the leverage action of the clavicle on the manubrium 
and first costal arch. I considered that the ossification of the 
first cartilages, at a period long antecedent to that of the other 
cartilages, was due almost entirely to the strain exerted on 
it by force transmitted through the clavicle. I then supposed 
that the leverage action of the clavicle and the movements of 
the sternum in respiration were the causes which determined 
the joint-formation in the ossified cartilage, and that of these 
two the former was the most important factor in its production. 

I think the conditions presented by this fused first cartilage 
go far to prove the truth of that hypothesis. Against it, how- 
ever, there is the fact that where other cartilages besides the 


1 “* Gase of Costo-chondral Dislocation,” vol. xxxiv.; ‘‘ Mode of Fracture of 
the Sternum,” vol. xxxv. ; ‘‘ Mode of Fracture of First Rib alone,” vol. xxxvi. 


398 MR W. ARBUTHNOT LANE. 


first ossify, they also develop an articulation. For further 
details I would refer to the papers already mentioned. 

This man had been a labourer, and his spinal column showed 
extensive pressure changes. 


Two Subjects with Six Lumbar Vertebre, the First Lumbar 
Vertebra in one possessing Lumbar Ribs. Bifid Spinous 
Processes in Lower Dorsal Region. Separation of the First 
Piece of the Sacrum. Fusion of the Sacrum and First 
Lumbar, &e, 


Before commencing the description of these cases, I will 
refer for the third time to the case in which there were thirteen 
cervico-dorsal vertebra bearing ribs,’ besides seven cervical and 
five lumbar vertebra. In that paper I entered into what now 
appears to me to have been rather a barren discussion, namely, 
as to whether the extra vertebra was a dorsal vertebra with ribs 
deficiently developed, or an eighth cervical with cervical ribs, 
and I then concluded in favour of the last supposition. The 
question would now appear to be a less obscure and a more 
general one. 

There was one point I did not mention in that paper and 
which will bear on the question. It is one that must be of 
great obstetric interest. That is, that the oval facet on the 
upper surface of the sacrum occupied a lower position than 
usual, and that the line prolonging its direction forwards cut the 
symphysis just below its upper margin. The reason I did not 
describe this at that time was because I had not then succeeded 
in separating very distinctly the various modifications in form 
which the lower part of the spinal column undergoes under the 
influence of continued and exaggerated superjacent pressure, or 
of ordinary pressure acting on an enfeebled subject, and I then 
thought that the condition of the sacrum present in this subject 
was produced in a like manner, especially as there were marked 
pressure changes in the whole column. Thinking it had no 
bearing on the subject under discussion, I left it out of the 
paper. 

Since that time I have been able to separate and classify the 


1 «* Supernumerary Cervico-Dorsal Bearing Ribs,” Journal of Anat. and Phys., 
1885. 


SOME VARIATIONS IN THE HUMAN SKELETON. 399 


pressure changes in the lower part of the spinal column, and I 
have described them fully in a paper! in the Z’ransactions of the 
Pathological Society. 1 then found that the conditions pre- 
sented by this sacrum were quite distinct from those produced by 
pressure, though resembling them superficially. I was also able 
to show in that paper that the larger number of the numerous 
modifications in form of the lumbar and sacral spinous processes, 
and in the form and direction of the articular processes, were due 
to pressure, and were not instances of congenital deviations from 
the normal, as supposed up to the present time by those who 
have observed them. 

On examining other specimens, in which there were six 
lumbar vertebree, I found the same condition of the sacrum 
present. 

In the normal sacrum, if the direction of the plane of the oval 
articular surface be continued forwards, it will be seen to pass 
about an inch and a half above the symphysis, and the upper 
limit of the sacral body is found to be considerably above the 
level of the brim of the true pelvis. 

With these facts before me, I examined the two following 
cases very thoroughly with the view of clearing up the difficulty. 


The first subject was a well-formed male, above the average height. 
His bones were dense and strong, and he still retained most of his 
teeth. 

The spinal column showed no marked pressure change. The lumbar 
convexity was very prominent, and much longer than usual. 

The direction of the oval facet on the upper surface of the sacrum 
cut the symphysis immediately below its upper limit, and the facet was 
seen to be on the same level as the ileo-pectineal line, therefore con- 
siderably below its normal position. 

As a result of this, the upper aperture of the true pelvis was heart- 
shaped, the apex of the heart corresponding to the back and of the sym- 
physis, and being therefore flattened. The true pelvis was small, and 
its antero-posterior diameter at the inlet particularly so. I have unfor- 
tunately mislaid the measurements. 

There were six lumbar vertebra, the uppermost having on either 
side ribs one inch long, and articulating with either side of the body 
by a small facet with a synovial membrane, and posteriorly with the 
small transverse process. The transverse process of the third lumbar 
vertebra was longer than any of the others. The first lumbar nerve 
gave off the ilio-hypogastric and the ilio-inguinal, the second gave off 


1 <«* Pressure Changes in the Lower Part of the Spinal Column,” vol xxxvi. 


400 MR W. ARBUTHNOF LANE. 


the genito-crural, and the obturator and anterior crural nerves came off 
from the second, third, and fourth lumbar nerves. The fifth lumbar 


passed down over the upper surface of the sacrum, and gave. off the’ 


superior gluteal. The sixth lumbar nerve passed down to form part 
of the sacral plexus. The first sacral nerve joined with the two last. 
The second sacral nerve was small compared to the first, and was no 
larger than the normal third sacral. There was a very small third 
sacral, and it took the normal distribution of the fourth. The fourth 
resembled in distribution the normal fifth ; it did not enter into the 
formation of the sacral plexus. 

On making a vertical median section of the lumbar spine and 
sacrum, the latter bone was seen to consist of five parts, the limits 
of the bodies of the first three being still indicated by layers of fibrous 
tissue. 

On comparing with this section another obtained from a normal 
column, it was then seen that the first piece of the sacrum had become 
separated so as to form the sixth lumbar vertebra, which the first 
coccygeal had united with the sacrum, so that it still consisted of five 
pieces. As we have already seen, the mode of distribution of the 
sixth lumbar was that of the normal first sacral. 

On removing the fibro-cartilage between the sixth lumbar vertebra 
and sacrum, and then making these two bones articulate directly, the 
appearance given by the normal sacrum is obtained. 

This shows that at least in this case and in similar ones an extra 
vertebra is not interposed between the occiput and attachment of the 
iliac bones to the spine, but that the first sacral vertebra, which does 
not seem to form a necessary part of the sacro-iliac articulation, is 
separated from the rest of the bones, the sacrum having joined to it 
the first coceygeal vertebra. Modifications in the form of the thorax 
are also present in these cases of dissociated first sacral vertebrae, and 
they consist in an increase in the size of the lower part of the chest, 
and a corresponding decrease in the capacity of the upper part. In 
this subject the manubrium was one and a half incheslong. The span 
of the first rib was two and three-quarter inches. Light cartilages 
articulated symmetrically with the sternum, and the twelfth ribs were 
more than 7 inches long. The costal cartilages were shorter than 
usual, but they were equal on both sides. The pleura extended for 
an inch and a half below the lower margin of the last rib. 


The LOWER LIMIT OF THE PLEURA is a point that deserves much 
attention, especially now that the kidney is so frequently re- 
moved or explored from the loin, since opening of the pleural 
cavity at this point has caused both immediate and subsequent 
danger. This I have alluded to in describing the operation.! 
The lower limit of the cavity behind can be determined clinically 
by measuring the lower ribs, and especially the twelfth. 


1 “Nephrectomy :” Manual of Operative Surgery. 


cain geal winsome on 


Sane eee Cee 


. 


SOME VARIATIONS IN THE HUMAN SKELETON. 401 


It will be found that if the twelfth ribs are under two inches 
in length, the pleura may not even reach its upper margin, or, at 
the most, it may extend but a little way over the inner half of 
its anterior surface. 

As the length of this rib increases, the pleura extends still 
lower down, and in some instances, as in that just described, it 
may reach a point an inch and a half below the lower border of 
the rib. 

If the last rib be over two inches in length, the lower limit 
of pleura crosses it obliquely at its centre, so that it bears a direct 
relation to the length of the rib. This relationship of the pleura 
to the length of the last rib shows that there need be no risk 
of opening this cavity in the ordinary operation of lumbar 
nephrectomy. In the description of the extent of the pleura in 
Quain, vol. ii., the following somewhat vague reference 1s 
made :—* Behind, the lower extent of the pleura is as far down 
as the vertebral end of the twelfth rib, or even in some cases as 
far as the transverse process of the first lumbar vertebra.” The 
great and frequent variations in the length of the lower ribs, not 
always on both sides, and the consequent modifications in the 
form of the thorax, are not considered by anatomists in their 
description of the pleural limits. For instance, Luschka says 
that in the axillary line the right pleura extends down to the 
lower edge of the ninth rib, while the left pleura reaches to the 
upper edge of the tenth. In the many bodies I have examined 
for this purpose, I have been unable to verify this greater down- 
wards extent of the pleura on the left side ; in fact I have found 
the reverse more frequently true. Taking the average of a 
number of cases, the lower limit of the pleura crossed the 
seventh costal cartilage obliquely about three-quarters of an 
inch below its articulation with the rib, then the end of the 
eighth rib, or the cartilage immediately below it, the ninth rib a 
quarter of an inch above its extremity, the tenth rib three- 
quarters of an inch, and the eleventh one and a quarter inches 
from its outer end. In the axillary line, which is at least some- 
times defined as a line passing vertically down from the head of 
the humerus, I found that in the large majority of cases the 
lower limit crossed the tenth space or the eleventh rib, not 
unfrequently lower on one side than on the other, but not much 


402 MR W. ARBUTHNOT LANE. 


more frequently on one side. Luschka probably used the term 


in a different acceptation, yet that fact would not modify the~ 


relative extent of the limit on both sides, 

The second case, in which there were six lumbar vertebre, 
resembled the last pretty closely. It was also a male subject. 
The lower part of the chest was relatively large and the upper 
part small. The sternum had been removed before the subject 
came into the room. The subclavius muscle was abnormal on 
both sides, the tendon splitting into two divisions, of which each 
had a muscular belly, one being attached to the upper margin 
of the scapula, and the other to the normal attachment of the 
muscle to the clavicle. ‘The sacrum consisted of five pieces, but 
on making a vertical median section of the pelvis it was seen that 
the last piece was the first coccygeal, which had become fused to 
the sacrum. The pelvis presented the same changes as those 
described in the last case. It was small, considering that the 
man was much above the average in size and build. The upper 
aperture of the true pelvis was typically heart-shaped. Its 
antero-posterior diameter was only three and a half inches, while 
its greatest transverse diameter measured five and a quarter inches. 
In the average male pelvis these diameters are four inches and four 
and a half inches respectively, so in this case the conjugate was 
considerably decreased, while the transverse diameter was increased. 
The interval between the anterior superior spines was ten inches, 
the normal being seven and three-quarter to eight and three- 
quarter inches. The sacro-vertebral angle measured 130°, this 
angle averaging usually from 110° to 125°. The breadth of the 
sacrum was about normal, namely, four and a half inches. 

The separated first sacral vertebra resembled the last lumbar 
in appearance, except that the right transverse process was very 
large and articulated with the surface of the ilium by an ex- 
panded flat facet. The left transverse process was longer than 
that of the normal fifth lumbar. The formation of the lumbar 
and sacral plexuses was identical with that described in the 
last case. 

The lumbar curve was very prominent indeed, producing 
well marked lordosis, and consequent backward protrusion of 
the buttock and an increased obliquity of the position of the 
pelvis. This condition of sacral dissociation is probably the 


ee we 


SOME VARIATIONS IN THE HUMAN SKELETON. 403 


reason of the lordosis and considerable gluteal prominence which 
one sees occasionally, and which dates back in these cases to 
early life. It may be very easily overlooked, as the superficial 
deviations from the normal are very slight indeed. 

The spinous processes of the lower two dorsal vertebre and 
the upper three lumbar vertebrae were very peculiar. The 
spine of the tenth dorsal vertebrae was very broad and expanded 
at its posterior extremity, where it presented a vertical groove 
in its centre. The spine of the eleventh dorsal was also very 
thick and strong. It was completely bifid at its extremity, the 
divisions also being thick and strong. That of the twelfth was 
similar in form, except that both the spine and its bifurcation 
was even stronger. The spine of the first lumbar vertebra was 
very thick, and terminated in a great flat end, measuring trans- 
versely three-quarters of an inch, and five-sixths of an inch in 
the vertical diameter. The spine of the second lumbar was 
similar in form and structure, and ended in a smaller plate than 
the last. 

The spine of the third, fourth, fifth, and sixth were like those 
of the ordinary lumbar vertebra in form. They were very 
strong, but their depth was not exaggerated, so that they did 
not articulate directly with one another. On the lower margin 
of the second, third, and fourth lumbar spines there were on 
either side, at a point half an inch from the apex, large 
prominent nodules receiving the insertion of tendons of the 
multifidus spine. 

Between the bifid extremities of the tenth, eleventh, and 
twelfth dorsal spines and the spines of the adjacent vertebre 
there were pairs of interspinous muscles covered by a strong 
layer of supraspinous ligament. Between the spines of the first 
and second lumbar vertebre there was a single layer of inter- 
spinous muscle, beneath a very dense supraspinous ligament, and 
between the spines of the lower lumbar vertebree and the sacral 
spinous process there was an extremely thick and dense liga- 
mentous tissue. 

The erector spine and multifidus spine muscles in the 
lumbar region were remarkably large, and the aponeurosis 
covering them particularly strong and dense. 

The reason of all this increase in the size of the spinous pro- 


404 SOME VARIATIONS IN THE HUMAN SKELETON. 


cesses, and in the bulk and strength of the ligaments and 


muscle, was to compensate for the great concavity and length of - 


the lumbar region, and the consequent greater force required to 
raise the flexed upper part of the trunk upon the sacrum. 

I have never before seen this splitting and enlargement of the 
lumbar and dorsal spines which were present in this specimen, 
nor have I read of it anywhere. 


The necks of the thigh-bones joined the shaft at an extremely 
oblique angle. I mention this fact as I see that the same remarkable 
obliquity of the neck was present in a specimen of what appears to 
have been dissociated first sacral vertebra, which is described by 
Professor Struthers in page 83, “ Variations of the Vertebra and Ribs 
in Man.” 

This man also had a remarkably large head, whose increase in size 
was most marked in its antero-posterior diameter. The left side was 
the larger throughout. The increase in the transverse diameter and 
the decrease in the conjugate diameter of the brain are in a direction 
the reverse of retrocession to a lower type, as is also the increased 
capacity of the cranial cavity. 

With this case we might compare an example of fusion of the sacrum 
with the fifth lumbar vertebra. This is distinctly a condition of retro- 
cession, as the last lumbar vertebra not unfrequently joins with the 
sacrum in the gorilla, orang, and chimpanzee. It was a male subject, 
whose chest was of an average size; the manubrium and gladiolus 
having their average relative size. The last ribs were about 8 
inches long, consequently there was but a narrow interval between it 
and the iliac crest. The margin of the fifth lumbar was united to 
the first sacral by bone, the intervening fibro-cartilage being mueh 
diminished in depth. The lumbar vertebre formed with the sacrum 
an angle of 145°. The conjugate of the brain measured 44 inches, 
and the transverse diameter 8} inches. The neck of the femur joined 
the shaft at an angle slightly less than normal. This was not a very 
typical case of fusion of the sacral and last lumbar vertebrae, as the 
union of these two bones was not complete. After the bone connect- 
ing their margins had been cut through with the contained fibro-carti- 
lage, the laminz and spinous processes were found to be quite free, 
the articular surfaces of the last lumbar vertebra looking directly for- 
wards. Probably on this account the pelvis does not deviate very 
markedly from the normal. 


NOTE ON A CASE OF BICIPITAL RIB. By R. L. Mac- 
DonNnELL, M.D., Demonstrator of Anatomy in M Gill 
University, Montreal. 


THE recent article of Professor Turner on “Cervical Ribs and 
the so-called Bicipital Ribs in Man,” in the Journal of Anatomy 
and Physiology, vol. xvii., as well as his previous contribution 
on the subject of the so-called two-headed ribs in whales and in 
man, vol. v., leaves very little to be said upon the subject of 
fused ribs; but inasmuch as the occurrence of this abnormality 
is rare, it may be well to place upon record a description of a 
case of bicipital rib which came under my notice in the dis- 
secting-room of M‘Gill University in the winter session of 
1884-85. 

In a male subject the first and second ribs on the right side 
were joined up to a distance of an inch from their cartilages. 
The first rib has a normal head articulating in the usual manner 
with the first dorsal vertebra. The neck of the rib is rather 
more bent upon the shaft than is usually the case, and its width 
from before backwards, at the tubercle is beyond the normal. 
The junction with the second rib takes place at a distance 
of five-eighths of an inch from the tip of its tubercle, the fusion 
being apparently from above downwards, as if the ribs were 
laid one on the top of another. For a little more than one-half 
the distance between the tubercle and the sternal extremity, 
the first rib in outline is distinct from the second, there being 
a shallow groove between the two bones, but after this point 
the junction of the two ribs takes place more by their bor- 
ders than by their surfaces, so that they form a large flat 
plate of bone gradually spreading out to a maximum width 
of two inches. On holding the specimen up to the light the 
two ribs are seen to be separated from one another by a very 
thin transparent plate of bone for a distance of half an inch 
from their point of fusion. On the outer surface of the fused 
body there is the usual rough surface on the second rib for 
the attachment of the serratus magnus muscle. The rough 
line commonly seen on the upper surface of the first rib, 


406 NOTE ON A CASE OF BICIPITAL RIB. 


which gives attachment te the scalenus medius is not dis- 


tinctly marked upon the first rib, but is plainly seen extend- - 


ing from its usual site over that portion of the fused body 
which belongs to the second rib. The scalene tubercle is not 
distinct. The bifurcation takes place anteriorly, at a distance 
of 3 inches from the tubercle of the first rib, The upper 
branch of it ends abruptly in the cartilage of the first mb, 
while the lower branch, the continuation of the second rib, is 
one and a half inches long, three-quarters of an inch wide at 
its outer, and one-third of an inch wide at its sternal end. 
The costal cartilage to which this lower branch is attached 
ends in the usual position of the second costal cartilage. 

On the under surface the fused body presents a smooth 
surface for the protection of the upper part of the pleural 
cavity. 

Owing to an unfortunate occurrence,—the removal of the 
ribs soon after their peculiarity had been observed,—I am 
unable to give any account of the dissection of the soft parts. 

In this same subject it was found on cleaning the skeleton 
that the first dorsal vertebra presented a remarkable abnor- 
mality. The laminz of the neural arches of each side do 
not run on the same plane, the one with the other; but one, 
that of the left side, is higher than that of the right, so that 
the spine of the vertebra is formed by the placing cf one 
lamina over the other. 


i 
f 
a 
y 


OSTEOLOGY OF CONURUS CAROLINENSIS. By R. W. 
SHUFELD?, M.D., Medical Corps U.S. Army; Membr. A.0.U. ; 
Membr. Soc. Nat. E.US.; Membr. Philosoph. Biol. Socs. of 
Washington, &c, (PLATES X., XI.) 


WE learn from the best authorities on the subject, and those 
who have given special attention to the Pszttact, that we may 
reckon something over three hundred and fifty well determined 
species of Parrots in the world’s fauna, so far as it is at present 
known. The strongly marked characters and the unmistakable 
external appearance of any of the forms that go to make up this 
wonderfully interesting group of birds, have long served to 
clearly distinguish them from other, and perhaps not so well 
defined, orders of the class. Fortunate as ornithologists have 
been in this respect, the matter of drawing the lines for the 
minor divisions used in classification has, since the earliest 
days of the science, owing to the perplexing interrelation of the 
majority of the forms, proved a far more difficult task. The 
several schemes proposing as many arrangements into families 
were all more or less unsatisfactory, until the subject came 
under the master-hand of Garrod, who, through his able and 
well-directed investigations into the structure of the leading 
types of the order, showed how the entire group could first be 
divided into two primary series, viz., the PALHORNITHID, and 
the Psirracipé.! The former of these (excepting Cacatua) 
having two carotids, the left one being normal, and no ambiens 
muscle; while, on the other hand, the remaining series have the 
two carotids present, the left one being superficial, and the am- 
biens, present in some of its genera, is found to be missing in 
others. 

Fourth among the divisions of the second series, or in the 
Psitiacide, we find the Avine, characterised by the presence of 
the ambiens muscle, tufted oil-gland, and union of the clavicles 
at their lowest and median point, into a complete os furculum. 


1 Garrod, A. H., ‘On some Points in the Anatomy of the Parrots, which bear 
on the Classification of the Sub-order,” (P. B. S., 1874, pp. 586-98). 


408 DR R. W. SHUFELDT. 


To this sub-family Avine belongs the sole representative of 
the Order in the avifauna of the United States, the Carolina ~ 
parrot, and it is to the description of the skeleton of this form 
that the present memoir is devoted. 

Formerly the geographical distribution of this bird spread 
over a much larger area, but of late years, between the interests 
of feather-dealers, and its general wanton destruction, the com- 
plete extermination of the species seems to be as certain as it is 
no less a much to be regretted fact. 

My first attempts to secure a skeleton of this bird, for the 
purpose of description, failed, and it was not until Mr James 
Bell of Gainesville, Florida, cheerfully came to my assistance 
that success at last was met. This gentleman, always ready 
with his aid in the interests of science, forwarded me two very 
complete skeletons, a male and a female of this species, to meet 
the end I had in view. 

This very acceptable gift came to my hands in March 1885, 
at a time when every moment of my spare hours was being 
devoted to a large mass of material, chiefly osteological, illus- 
trating the American sub-polar avifauna, and belonging to the 
Smithsonian Institution, so that it was not possible until the 
present month, August 1885, when my labours in that direction 
ceased, that I could at last give my attention to placing 
on record an account of the skeleton of our little Carolina 
parroquet. 

Of the two skeletons in question, I chose the male to make 
my drawings from in the plates, and I found it to be somewhat 
larger than the female specimen. Notwithstanding, both were 
evidently adults. 

Of the Skull (Pl. X. figs. 1-4).—In Conuwrus the culmen of 
the superior mandible or its middle line lies in the are of a 
circle extending from the cranio-facial axis to the tip of this 
part of the beak. Posterior to the subcircular osseous nares 
this culmenar surface is broad and nearly flat, but beyond these 
apertures it is convex, both transversely, and as has been said, 
along its middle line. The dentary margins of this mandible 
are cultrate and deeply notched at their middle points, as shown 
in figure 1, where the skull is represented upon lateral aspect. 

Complete absorption of the nasal bones has taken place, so 


OSTEOLOGY OF CONURUS CAROLINENSIS. 409 


that in the adult their existence would never be suspected, and 
their exact limits can only be guessed at, as no sutural traces 
remain. 

The under side of this mandible is bounded behind by a 
straight, transverse line, beneath which the palatines are in- 
serted. Its general surface, otherwise, is unbroken, and evenly 
concave throughout (fig. 4). 

The interior of this part of the skull is more or less filled up 
with an osseous, spongy tissue, that presents a more compact 
nature where it forms the anterior wall to the rhinal chamber. 
One cannot very well judge as to how much, if any, of this 
structure can be claimed as representing the maxillo-palatines 
in this parrot. 

Speaking in general terms, Professor Huxley says of the 
Psitiaci that “the maxillo-palatines are very large and spongy 
in texture, and unite with one another and with the ossified 
nasal septum so as to fill up almost the whole base of the beak. 
Above, however, a nasal passage is left on each side; and, 
below the maxillo-palatines stop short, so that, in the dry skull, 
a passage, leading into the cavity of the rostrum, is left on each 
side of the septum.”! 

These remarks are illustrated by the under view of the skull of 
Cacatua galerita, which bird, I think, from the drawing, must 
have its nasal septum as well as this spongy mass which sur- 
rounds, produced much further backwards than it is in the 
subject we have in hand. However, the parts no doubt are 
homologous in both of these forms, though one would hardly 
have suspected the mass in question to have represented a part 
of the skull deserving of a special name, had the Carolina parrot 
been the only bird examined. 

Among the points that have always attracted the most 
attention in the skeleton of the Psittaci, the cranio-facial hinge 
is here in Conwrus as perfect in its mechanism as we perhaps 
will find it in any of the order. 

Its structure is too well known to enter upon its details here ; 
I find, however, that neither in this parrot nor any other of the 
group that I have ever examined is this feature one whit better 
developed than it is in Sula bassana. 


1 Huxley, T. H., ‘‘ On the Classification of Birds,” P, Z, S., 1867, p. 442. 
VOL. XX. 20 


410 DR R. W. SHUFELDT. 


Passing now to other parts, we find the union between 


the sphenotic process and the descending portion of the la-. 


chrymal bone to be complete, forming an external orbital 
periphery or ring, which is very nearly circular (figs. 1, 4, o7.). 

According to Parker, this is through the intervention of 
the os wneinatwm, which in some parrots, by union with the 
zeyomatic process of the squamosal, bridges over the temporal 
fossa.t 

The lachrymal itself has indistinguishably, so far as a suture is 
concerned, merged above with the frontal bone, while its union 
with the sphenotic process, just alluded to, is equally well 
obliterated. 

Internally it unites in a similar manner with the small 
pars plana, a circular foramen for the olfactory nerve passing 
between it and the ethmoid, while externally the antero-inferior 
arc of the orbital ring is marked by a longitudinal concave 
notch. 

As for the orbital cavity itself, its walls are but fairly entire, 
the pars plana being small, and the exit from the brain-case for 
the first nerve being far larger than this branch demands. 
Moreover, the palatines being vertical plates in this situation, 
and the pterygoids slender, the floor of the cavity is necessarily 
badly provided for in this regard. 

In both these specimens the foramina for the exit of the optic 
and the third, fourth, and sixth nerves are distinct, and scarcely 
any greater in size than the structures they are designed to 
transmit are in calibre. 

The interorbital system is without vacuities, and directly 
throughout merges with the rostrum of the sphenoid beneath it, 
the lower margin of the whole plate being sharp, both inferiorly 
and in front, 

Anteriorly the ethmoid bone proper is very broad, being spread 
out as an abutment against, and bordering for all its width, the 
posterior line of the cranio-facial hinge, The body of the bone 
is thickened and filled in with diploic tissue. 

That portion of the skull which lies behind and below the 
orbital ring presents for examination, above the lateral aspect of 
the evenly-convex vault of the cranium, and below, the long 


1 Parker, W. K., Morphology of the Skull, p. 264. 


se 


OSTEOLOGY OF CONURUS CAROLINENSIS. 411 


squamosal process separated from the sphenotic by a well- 
defined valley. 

The bony ear conch is circumscribed by sharp margins, while 
to its under side a vertical plate is thrown down the temporal 
wing of the exoccipital. 

As is well known, the characteristic feature of a parrot’s 
quadrate bone is, that its mandibular facet is single, and placed 
in such a way that its long axis is in the same straight line with 
the longitudinal axis of the pterygoid; to the possession of this 
narrow, convex, and long facet Conwrus forms no exception, 
Above it, the body of the quadrate is flattened from side to side, 
with a conical projection on its outer aspect, posterior to its middle 
point, that has a pit as its apex to accommodate the apophysis 
on the inner side of the hinder extremity of the quadrato-jugal. 

The orbital process of the quadrate is spiculaform and well 
developed, while the mastoidal limb of the bone rises as a stout 
subeylindrical rod, with two convex articular facets at its 
summit. These are divided by a notch, and the inner one of 
the two is very small, not presenting more than one-tenth of 
the amount of articular surface the outer one does. The pneu- 
matic foramen is found near its most usual place, on the inner 
and back part of the mastoidal shaft. 

Viewing the skull now from above as shown in fig. 3, we 
observe that the narial apertures can also be seen upon this 
aspect surrounded above by a few minute vascular foramina. 

The cranio-facial hinge bordering the superior mandible 
behind, is found to be a transverse and depressed line extending 
all the way across. 

Between the superior orbital peripheries the frontal region of 
the skull is smooth and nearly flat; as we proceed backwards 
it gradually becomes convex, to form the beautifully rounded 
vault of the cranium. 

A few perforating foramina are found just within the two 
edges of the orbits in the frontal bones. 

Turning to the under side of the skull (fig. 4), the most 
remarkable feature that confronts us is the extraordinarily 
fashioned palatines. These bones, as they occur in the Psittaci, 
have been described by a number of anatomists, so their pecu- 
liar conformation is well known. Conurus, like most true 


412 DR R. W. SHUFELDT. 


parrots, has either of these bones horizontally flattened in front, 


where it is inserted above the hinder portion of the superior- 


mandible to meet the lower part of the nasal septum, but not 
the palatine of the opposite side. 

Proceeding backwards from this horizontal extremity, the 
palatine is seen to contract, then immediately afterwards to 
form a broad, oblong, and vertical plate. This plate has a 
certain limited portion of its antero-superior part curled towards 
the median line, where it meets a corresponding edge of the 
fellow of the opposite side; and the two here form, by the 
assistance of the palato-pterygoidal articulation, the usual longi- 
tudinal groove for the under edge of the rostrum. 

Behind this the superior margin of the palatine plate is 
sharp, and has the anterior two-thirds of the corresponding 
pterygoid resting upon it; the posterior margin shows a deep 
notch, while the inferior margin of this part of the bone is 
rounded, becoming in front continuous with the dilated anterior 
end. 

Both the internal and external surfaces of these palatine plates 
may develop processes and ridges for the better insertion of 
muscles, which in life are thereto attached. A broad, spindle- 
shaped vacuity exists between these palatines in front, while 
posteriorly the angle separating their plates is somewhat less 
than the angle of divergence of the pterygoid bones (fig. 4). 

These latter elements are long, nearly straight and cylindrical 
rods articulating as shown in the figures in the Plate (pt.). 
They are at some distance below the basis cranii; and in no 
parrot, so far as is at present known, do they develop basiptery- 
goid processes. 

The maxillary portion of either of the infraorbital bars (ms.) 
is inserted by a somewhat horizontally-flattened end, just within 
the posterior edge of the beak, on a higher plane than the inser- 
tion of the palatines, and at a point where I take the foot of the 
nasal to be. The remainder of the bar almost immediately 
becomes of a uniform calibre, and at first being concave out- 
wards, passes just beneath the orbital wing, directly downwards 
and backwards to its articulation with the quadrate (fig. 4, 
ML., JU). 

At the cranial base we find a basitemporal area of small 


or 


——_—-~S—i‘ ie 


OSTEOLOGY OF CONURUS CAROLINENSIS. 413 


extent, triangular in outline, bounded on all its sides by raised 
lines, and having its apex directed anteriorly, terminating at 
the point where occurs the naked and external double-tubed 
entrance of the Eustachian canals. 

On either side of these apertures are seen from three to four, 
or sometimes only two, minute foramina. Well to the outer 
sides of these are the conspicuous foramina ovalia. 

As has already been said in a former paragraph, the temporal 
wings of the exoccipitals are very prominently produced ; and, 
as usual, to their inner sides at the basal angles of the basi- 
temporal triangle are found the ordinary group of foramina for 
the entrance and exit of vessels and nerves. 

The foramen magnum is of a subcircular outline, and the 
plane of its periphery makes an angle of some 20° with the 
backwardly-produced plane of the basis cranii. The condyle is 
comparatively large, hemispherical in form, and sessile. 

Rising almost perpendicular to the basitemporal triangular, 
the occipital area is well defined by an elliptical bounding are, 
which sweeps round on either hand to the apices of the temporal 
wings. In the middle of this space a moderately prominent, 
unpierced, supraoccipital elevation is to be seen. In removing 
the cranial vault I find that the tables are very closely juxta- 
opposed, and, of a consequence, but a little diploic tissue present. 
The several cerebral fossee are sharply defined by out-jutting 
lamelliform ledges of bone. 

At the base of the sella turcica there seems to be but one 
carotid opening, and the posterior clinoid wall to this fossa is 
very thin, and usually exhibits one or two perforations. 

The mandible (figs. 1 and 2) of Conwrus is somewhat horse- 
shoe shaped, with very deep and smooth ramal sides, which are 
deficient anteriorly, leaving a semicircular opening with cutting 
edge all round. When the horny mandibular sheath is care- 
fully removed in the fresh specimen, this edge has filiform 
prolongations of soft tissue standing out from its middle third 
below, which, after they have dried and become more or less 
shrunken, look something like a row of delicate teeth. 

The ramal sides of this bone slope away as we proceed back- 
wards, and the mandibular ends are truncated at about the 
same angle. 


414 DR R. W. SHUFELDT. 


To the inner sides of these articular ends a ledge is thrown 
out to support the facet for either quadrate. Behind these- 
longitudinal articular grooves single pits are found, at the 
bases of which the pneumatic foramina occur. 

The under borders of the mandible are smooth and rounded 
(fig. 2). | 

Of the Hyoid Arches (Pl. X. fig. 5)—Notwithstanding the 
fact that the glossohyal which supports the thick, short, and 
fleshy tongue of this parrot remains in cartilage throughout life, 
the ceratohyals (ch.) are very completely developed, meet in 
the median line, and ossify up to the very hinder body of this 
element anterior to them. Where they unite at the mesial 
point behind, an articular surface is formed for the first basi- 
branchial (bh.). This last-named element is unusually long, 
and anchyloses with the second basibranchial (0.b2.), the point 
of mergence being enlarged to accommodate the heads of 4 
eeratobranchials (c.br.), and anteriorly to support a peculiar | 
osseous outgrowth (0/.), that, so far as I am at present informed, 
is restricted to the Psittaci; indeed, this is the only form in 4 
which I have observed this latter feature. ; 

The ceratobranchials (¢.b7.) are very long, subeylindrical, and 
rather stout rods of bone; while, on the other hand, the 
epibranchials (c.br.) are notably short and but feebly developed, 
As thus constituted, the thyrohyal elements show but little 


curvature along their continuities, and still less disposition to 
curl up behind the cranium. 

Of the Remainder of the Aaial Skeleton (Pl. XI. figs. 8, 9, 
10, 18, 15, 17, and 18).—Conurus carolinensis has thirty-five ; 
vertebree in its spinal column, and a large pygostyle at its 
terminal extremity, 

The atlas is characterised by a broad neural arch above, : 
an unperforated cup for the occipital condyle, and a prominent | 
process extending backwards from the pseudo-centrum behind. J 

Axis vertebra has a very inconspicuous odontoid process, j 
strongly developed neural and hypapophysial spines, and : 
tuberous postzygapophyses. This segment, like the rest of 
the column and the pelvis, is pneumatic; to this statement, 
however, the last five caudal vertebree and pygostyle prove 
an exception. 


OSTEOLOGY OF CONURUS CAROLINENSIS. 415 

Both third and fourth vertebree have strong hypapophysial 
spines, and neural ones scarcely less marked. In these, too, 
the lateral canal are seen, but the processes at their hinder 
margins are, as yet, but feebly produced. The zygapophysial 
arms are short, and their being joined from before backwards 
in each case by bone extension lend to these two segments a 
width upon their dorsal aspects and a solid appearance not 
possessed by any of the other vertebrae below. 

In the fifth vertebra the dorsal and ventral spines have lost 
not a little of their prominence, while the parapophyses are 
much longer. This segment has the postzygapophyses mani- 
festly lengthened, whereas but little change has taken place in 
the anterior pair. 

The sixth vertebra loses the neural and hypapophysial 
spines altogether ; the parapophyses gradually diminish in size 
from this segment down the chain, until they, with the 
pleurapophyses, again become prominent as free ribs.  Like- 
wise the neural and lateral canals, which are here quite small, 
also increase in calibre as we proceed in the same direction. 

This vertebra has also a short carotid canal present in place 
of the hypapophysis. 

And this last feature is still better marked in the seventh 
vertebra, though it remains open below. These are the only 
two which have it in this parrot, in the eighth its site being 
again occupied by a low, median hypapophysial spine. 

In all these segments, as well as in the few succeeding ones 
that we find before coming to the true dorsals, the pre- and 
postzygapophyses are diverging limbs of the most usual form in 
aves. The articulations among the centra are heteroccelous. 

Ninth vertebra has the neural spine commencing to make 
its appearance again, and is here a low tubercle, more promi- 
nent in the tenth, and thus on till it assumes the dorsal form 
of this spine. The hypapophysial plates in both the ninth and 
tenth vertebre are deep, long, and of a quadrate form, and 
from the lateral masses being low on the sides of the centra, 
they appear sunken between these protuberances. 

We find that the twelfth vertebra has much the general 
aspect of one of the dorsals, and moreover, its pleurapophyses 
have become freed as a tiny pair of ribs. These attain quite 


416 DR R. W. SHUFELDT. 


a respectable length in the thirteenth vertebre, while in the 
fourteenth, where they are still unconnected with the sternum, 
they possess small unciform processes. 

We may term the fifteenth to the eighteenth vertebre in- 
clusive true dorsals, for they all have ribs connecting them 
in the usual way with the sternum. They also have inter- 
locking neural spines, and their transverse processes are 
strengthened by each one developing a single spiculaform 
interlacing metapophysis at its outer extremity. Prominent 
hypapophyses are found upon the thirteenth, fourteenth, 
fifteenth, and sixteenth, and a small one sometimes on the 
seventeenth vertebra. 

The ribs have broad unciform processes anchylosed to them, 
but there are still two other pairs that come from beneath the 
pelvis, belonging as they do to the sacrum, that also meet 
costal ribs below, which do not have these appendages. 

Sometimes abortive ribs are found anchylosed to the 
twenty-first and twenty-second vertebra; these being the third 
and fourth segments appropriated by the sacrum. 

Now, in my male specimen of this parroquet, I find the 
nineteenth to the twenty-ninth vertebra, inclusive, form the 
pelvic sacrum, while in the female an additional segment, 
which in the male remains a free caudal, has become firmly 
attached behind. 

This circumstance gives the male six free tail vertebra, 
whereas the other specimen has but five. Such a freak as 
this, however, not unfrequently happens among birds, where 
the count for the entire number of vertebre in the column 
remains wonderfully constant for the species. 

The caudal vertebre (fig. 13) have spreading transverse pro- 
cesses, and stumpy neural spines; the ultimate two having 
strong bifid hypapophyses, 

Of an irregular quadrilateral outline,the pygostylehasthickened 
hinder and lower margins, while the remaining two are cultrate. 

Giving our attention now to the pelvis, we find this com- 
pound bone in Conwrus (figs. 9,10) devoid of any very striking 
features, it having all the general characteristics of this part 
of the avian skeleton, without any to particularly distinguish 
it beyond the general facies of its kind. 


OSTEOLOGY OF CONURUS CAROLINENSIS. 417 


Viewed from above, it will be seen that the pre- and 
postacetabular areas are about equal in extent, the ilium being 
concave where it forms the first, and the reverse where it 
constitutes the latter. For the entire length of the sacrum 
these bones are firmly sealed to its outer margins, forming the 
most complete “ilio-neural canals” anteriorly, which do not 
even open posteriorly as in some birds; while behind it lends 
to the postacetabular area a very unbroken aspect, that is 
rendered even more so from the absence of all but a few 
small foramina among the sacral diapophyses. 

Upon the lateral aspect of the pelvis we note that the propubis 
is not developed, and that the inner periphery of the cotyloid ring 
is nearly as large as the outer one. The small obturator foramen 
is rendered complete by a pretty thorough meeting between the 
ischium and the somewhat slender postpubis immediately be- 
hind it. 

The obturator space is long and spindle-shaped, but the lower 
angle of the ischium does not need the postpubic shaft beneath 
it, as it does in so many birds. 

The antitrochanter and the elliptical ischial foramen are both 
of comparatively moderate size, and these several features, con- 
stituting as they do a group of notable occurrence on the lateral 
aspect in nearly all avian pelves, are here in harmony, both as 
to size and position, with the same as they are found in the 
class’s majority. 

On the under side we find that the lateral processes of the 
leading four sacral vertebree are thrown out as abutments against 
the nether sides of the ilia; beyond, or rather behind these, the 
usual cavity of the pelvic basin occurs, and the succeeding di- 
apophyses of these consolidated vertebree are less manifest than 
common, being all elevated and having their extremities in the 
roof above. 

The foramina for the exit of the sacral nerves are double, in 
each case one being placed above another, and the swell to 
accommodate the myelonic enlargement in this part of the cord’s 
track is here well pronounced. 

Conurus, in common with many other parrots, has for its 
general size comparatively a large sternwm (figs. 8 and 18). 
Seen from above we observe that the costal processes are but 


418 DR R. W. SHUFELDT. 


scarcely produced above the lateral borders, which latter rise 
cradually to their summits. These costal borders each support 
six facets for the hemapophyses, the concavities among them 
being pierced by small groups of pneumatic foramina. 

The space occupied by one of these costal borders is equal to 
about half the whole length of the lateral sternal margin. 

Posterior to them, on either side, the margins are sharp all the 
way round the xiphoidal extremity, this part of the bone having 
a shield-shaped outline, being concave above, though not nearly 
so much so as that part of the sternal body lying between the 
costal borders in front. 

In this latter section we sometimes find a few scattered pneu- 
matic foramina down the median line; the most constant, and a 
large one of these, however, is close up to the anterior border of 
the bone, which curls backwards over it, and the fossa thus 
fo-med is always spanned over by a median longitudinal bridge 
of bone. 

The anterior sternal body is thickened, and directly over its 
sharpened edge in front we find a continuous coracoidal groove ; 
beyond this there rears directly up a broad quadrate manubrium, 
which is continuous with, and has its lateral surfaces in the same 
plane with, the carina below. 

Extending the entire length of the body of the bone the keel 
of this sternum is comparatively a very deep one. Both its 
lower and anterior borders are convex, the latter being quite 
sharp. ‘The carinal angle formed by the meeting of these edges 
is rounded off, so that the lines form really one common curved 
line (fig. 8). 

That anterior vertical and thickened column of bone which 
is present in the keel of nearly all avian sterna is here well 
developed, but situated at some little distance back from the 
anterior margin. Moreover, it does not descend so far as is 
usually seen, being apparently interrupted by the muscular line 
which longitudinally marks the bone. 

The muscular lines of the pectoral aspect are roughly parallel 
to the costal borders, and remain quite distinct as we proceed 
towards the xiphoidal extremity, nearly as far as the elliptical 
fenestra that there occur, one on either side. 

In the shoulder-girdle (figs. 15 and 17) we find a scapula with 


OSTEOLOGY OF CONURUS CAROLINENSIS. 419 


rather a short blade though a stout one, having the usual sabre- 
hike form with obliquely truncated extremity posteriorly. 

It contributes the usual amount of articular surface to the 
glenoid cavity (g.), but when im situ does not occupy the entire 
length of the superior line of the scapular process of the coracoid, 
nor have any connection with the furculum except through a 
slight ligamentous attachment. 

I have represented in the Plate the right coracoid (c., fig. 17) 
as seen from a posterior view. 

It will be at once observed that in Conwrus this element of the 
girdle has a form that partakes much of the pattern it assumes 
among birds generally. ' 

Its tuberous summit is inclined slightly forwards and towards 
the median line, when articulated im situ, and has resting against 
it the frail clavicular head of that side. The scapular process 
already alluded to is well developed, but here chiefly given over 
to quite extensive ligamentous attachment. 

The coracoidal shaft is strong, comparatively of good length, 
and subcompressed in the antero-posterior direction, being 
faintly marked at the usual sites by muscular lines. 

At the sternal extremity of the bone we find the expanded 
portion, the form of which can best be seen in fig. 17, where 
we note that the lateral process at its externo-inferior angle is 
well marked. 

Many parrots are notorious for having incomplete furecula, in 
others the union at their medio-inferior points is very feeble. 

In this particular they resemble the Strigidw, a group they 
undoubtedly have some remote affinity with, in structure. Our 
Carolina parroquet has a complete os furculum, as shown in 
fig. 17 (cl.); it is, however, a very weak bone, and functionally 
accomplishes little more than an ossified ligament in the same 
position. Indeed, it reminds one very much of such a structure, 
for when duly articulated it is but little in advance of the 
imaginary plane that is tangent to the anterior surfaces of the 
coracoidal shafts, and consequently but little dissociated from 
the ligaments that descend from the coracoidal summits to meet 
for attachment on the top of the sternal manubrium. It is in 
form of the U-shaped pattern, and without a hypocleidium at 
the clavicular junction below. 

As already intimated above, and so far as the light I have on 


420 DR R. W. SHUFELDT. * 


the subject will at present permit me to judge, I believe that the 
shoulder-girdle of Conwrus more nearly resembles these parts 
in some of the owls than it does the corresponding lines in any 
other class of birds with which I am acquainted. 

Of the Appendicular Skeleton (Plates X., XI., figs. 6, 7, 11, 
12,13, and 16).—A glance at the drawing in the Plate of the 
pectoral limb of this parrot will be sufficient to convince us that 
it presents no very striking deviations from the average skeleton 
of the wing as found in existing birds. 

The bones are all harmoniously balanced both as regards their 
relative lengths and calibres. 

Pneumaticity is enjoyed by the hwmerus alone (h.), and this 
bone is here characterised by a short, though not inconspicuous, 
radial crest, an ulnar crest devoted, as usual, to forming a 
canopy over the pneumatic fossa, in which are found the air- 
holes leading to the interior of the humeral shaft: This latter 
is but little curved in any direction, being subcylindrical and 
smooth. At the distal extremity of the bone we find the 
trochleze for articulation with the antibrachial elements prom- 
inently produced, while on the obverse aspect a broad and a 
narrow gutter are seen, which guide the passing tendons in life. 

The radius is nearly straight for its entire length, differing 
from the w/na in this particular, it having a considerable curve 
along its shaft, the concavity of which is on the radial side, 
and gives rise to quite a wide interosseous space. 

This curvature of the ulna is not well seen in the drawing, 
because the bone is rotated there to a position that brings it on 
the side towards the observer, and consequently makes the shaft 
appear nearly straight. 

The carpal elements (re., we.) are two in number as usual, and 
they have the form most commonly presented by these bones 
throughout the majority of the class. 

In the manus we find a carpo-metacarpus of the ordinary form 
for birds generally. Its rather large pollex phalanx is without 
a claw, this feature being likewise absent from the tip of the 
distal digit. 

My drawing presents all these bones of the size they attain 
in an adult male parrot, with every important detail brought 
out, and held in such a position that their greatest lengths are 


OSTEOLOGY OF CONURUS CAROLINENSIS. 431 


represented, and can be easily measured and appreciated from 
the Plate itself. 

None of the bones of the pelvic extremity in Conwrus have air 
admitted to their interiors, and they ail become dark and greasy 
in the ordinarily prepared skeleton. 

The femur has a large, semiglobular head, with a shallow, 
though quite extensive excavation upon it, for the round liga- 
ment. A broad articular surface is found at the summit of the 
bone for the antitrochanter of the pelvis, and the suppressed 
trochanterian ridge does not rise above this. 

The shaft of the bone is but little bent in any direction, and 
it has the usual cylindrical form. At the distal extremity the 
condyles are fairly well developed, not strikingly large, the 
outer one being the lower when the bone is held in the vertical 
position, In front the rotular channel does not extend upon 
the shaft above them, while behind the popliteal depression is 
shallow also. 

The cleft for the fibular head marks the posterior aspect of 
the external condyle, dividing it, as usual, into two parts. 

Our subject possesses a small patella of a subcordate form, 
maintaining its usual relations with the bones of the leg and 
thigh (fig. 12, P.). 

Tibio-tarsus (fig. 12, 7%.) has its cnemial crest but very 
slightly produced above the articulating surface at the summit 
of the bone, while below it the pro- and ectocnemial ridges are 
but feebly manifested, and very soon merge into the shaft. 

This latter is quite straight and smooth, being slightly com- 
pressed in the antero-posterior direction. 

At the distal extremity the inner condylar protuberance is 
decidedly the more prominent, both upon front and rear aspects. 
The valley between these two eminences is quite wide and well 
defined, even to the posterior side of the bone. 

The osseous bridge for the extensor tendons is present. 

Marked feebleness in development is displayed on the part of 
the fibula of the Carolina parrot, for this bone is found not to 
extend below the ridge it articulates with on the side of the 
shaft of the greater leg-bone. Below this point the inferior 
apex of the fibular shaft is produced and replaced by a ligament 
of hair-like dimensions. 


4293 DR. Rk. W. SHUFELDT. 


What there is of the fibula in this bird, however, is fully as 
well developed as we usually find it in the class; simply its 
apparently useless prolongation, as seen in many birds, has never 
ossified. 

In the skeleton of the foot we find a short, thick-set tarso- 
metatarsus, with spreading trochlear extremity. Three views 
are presented in the Plate of this bone, besides the one where it 
is shown on a lateral view with the foot as a whole (figs. 13 and 
16). 

Its shaft is short and straight, being much compressed from 
before backwards, On the anterior aspect it is convex from 
side to side, while behind it is longitudinally excavated. The 
hypotarsus is a narrow, projecting ledge with one vertical, 
cylindrical perforation (fig. 13, st.) near its centre, and scarcely 
erooved for the other tendons behind. At the summit of this 
bone we note the two condylar depressions for the trochles of 
the tibio-tarsus. 

The usual arterial foramen pierces the shaft at its ordinary 
site at the distal end. 

As is well known, Psittaci are permanently zygodactyle birds 
by reversion of the fourth toe, while they not only possess a 
well developed and free accessory metatarsal, but the usual 
number of joints to the digits. Conwrus carolinensis agrees in 
all these particulars. 

Whenever I can I make it a rule to fully illustrate in the 
figures the tibio-tarsus and skeleton of the pes, as the points pre- 
sented by these parts stand among the most important in this 
all-important system of the bird’s anatomy, for when sufficient 
data of this kind become available they will be not only valuable 
as an aid in classification, but help to determine the affinities of 
existing birds with such fossil forms as may from time to time 
be discovered. It will be seen that I have not overlooked this 
fact in the present instance. 


Synopsis of the Skeletal Characters of Conwrus earolinensis. 


1. Superior mandible arched as in Raptores; osseous nares 
small, subcircular, separated by nasal septum. Dentary margins 
of mandible cultrate, and notched. 


OSTEOLOGY OF CONURUS CAROLINENSIS. 493 


2. Orbital ring complete. 

3. Cranio-facial hinge as in other Psiétacz. 

4, Lower margin of rostrum cultrate. 

5. Quadrate has a large and small facet on mastoidal head, a 
well-developed orbital process, and a single, longitudinal mandi- 
bular facet, which is laterally compressed and convex in both 
directions. 

6. Pterygoids long and slender rods, anterioriy articulating 
with each other and with the palatines. 

7. Major portion of either palatine—a large vertical plate, 
directed downwards and backwards. These bones curl towards 
each other and form a limited articulation in the median line; 
anteriorly they are horizontally flattened, and are hinged to 
the mandible beneath the spongy mass, which constitutes the 
maxillo-palatine and nasal septum. 

8. Mandible truncated in front; rami and symphysis deep and 
gradually round into each other.* 

9. Hyoid apparatus with large, united ceratohyals, and a 
peculiar bony outgrowth on either side of the first basibranchial, 
extending forwards. 

10. Manubrium of sternum erect, large, and continuous with 
the deep carina. Xiphoidal extremity of this bone has an ~ 
elliptical fenestra on either side. Costal processes low, and 
usually six hamapophysial facets on each costal border. 
Coracoidal grooves unite in front. 

11. Furculum of shoulder-girdle firmly united below. 

12. The humerus only is pneumatic in the pectoral limb. 

13. A well-developed patella present. Fibula short, extend- 
ing only so far as the lower end of fibular ridge of tibio- 
tarsus. 

14, The tendinal bridge at antero-distal end of tibio-tarsus 
at right angles to long axis of shaft. The inner condyle the 
larger and more elevated. 

15, General skeletal characters of pes agree with other 
Psitiaci. 


Negatwe Characters, 


1. Vomer absent. 
2. Basipterygoid processes not developed. 


424 DR R. W. SHUFELDT. 


3. No hypocleidium on os furculum, and this bone does not 
meet the scapular process of coracoid. 

4, Propubis of pelvis absent. 

5. Pelvic limb non-pneumatic. 


DESCRIPTION OF PLATES X., XI. 
Reference Letters. 


b. Basal view of left tarso- ju. Jugal. 
metatarsus. m’, m'', Middle metacarpal and its 
bf. Bony outgrowth from basi- phalanx. 
branchial. me. Maxillary. 
b.bh. Second basibranchial of or. Orbital ring. 
hyoid apparatus. P. Patella. 
bh. First basibranchial. p. Pollex phalanx. 
. Coracoid, pl. Palatine. 
c.br. Ceratobranchial of hyoid pt. Pterygoid. 
apparatus. q. Quadrate. 
ch. Ceratohyals. rd. Radius. 
cl. Os furculum. re. Radiale. 
e.br. Epibranchial of hyoid ap- st. Summitof tarso-metatarsus. 
paratus. Tt. Tibio-tarsus. 
Fb. Fibula. ue, Ulnare, 
. Glenoid cavity. ul, Ulna. 
. Humerus. 
i,t’, v. Second metacarpal and 


index digit. 


Fig. 1. Left lateral view of skull, and detached mandible of Co- 


nurus carolinensis, . Life size from the specimen. 


Fig. 2. Mandible of Conwrus carolinensis; seen from above. 


size, same subject. 
Fig. 3. Skull of Conurus carolinensis. 
Life size from the same specimen. 
Fig. 4. Under view of the skull of Conurus carolinensis ; mandible 


removed. 


from below, same specimen, x 2. 
Fig. 6. Anconeal aspect of left pectoral limb of Conurus carolinensis. 


Life size, same subject. 


show them to best advantage. 
Fig. 7. Palmar aspect of left humerus of Conurus carolinensis. 
Life size, same specimen. 
Fig. 8. Right lateral view of sternum of Conurus carolinensis. 
Life size, from the same specimen as before. 


Life 


Seen upon superior aspect. 


The mandible has-been removed. 


Life size, same specimen as before. 
Fig. 5. The hyoidean apparatus of Conuwrus carolinensis. 


Seen 


Bones somewhat dislodged, and moved to 


OSTEOLOGY OF CONURUS CAROLINENSIS. 425 


Fig. 9. Right lateral view of the pelvis of Conurus carolinensis. 
Life size, same specimen as in the former figures. 

Fig. 10. Superior aspect of pelvis of Conurus carolinensis. Life 
size, same bone as shown in fig. 9. 

Fig. 11. Anterior aspect of left femur of Conurus carolinensis. 
Life size. 

Fig. 12. Anterior aspect of left tibio-tarsus, fibula, and patella of 
Conurus carolinensis. Life size. 

Fig. 13. Anterior aspect of left tarso-metatarsus of Conuris caro- 
linensis, showing also basal and summit views of the same bone. All 
these bones, including the foot (fig. 16), are taken from the same 
specimen used for the former figures. Life size. 

Fig. 14. Right lateral view of the five vertebra and pygostyle con- 
stituting the skeleton of the tail of Conuwrus carolinensis, Life size, 
same specimen. 

Fig. 15. Superior aspect of left scapula of Conurus carolinensis. 
Life size, same specimen as in former figures. 

Fig. 16. Inner aspect of left pes of Conurus carolinensis. Life 
size, same specimen. 

Fig. 17. Right coracoid and os furculum of Conurus carolinensis, 
seen from behind. Life size, same specimen. 

Fig. 18. Sternum of Conurus carolinensis. Pectoral aspect. Life 
size, same bone shown in fig. 8. . 


VOL. XX, 2k 


A NAVAJO SKULL. By R. W. Suuretpr, M.D., Captain 
Med, Corps, U.S. Army, Membr. A.O.U., Membr. Soe. Nat. 
E.U.S., Memb. Scientif. Socs. of Washington ; Cor. Memb. Soc. 
Ital. Antropologia, Etnologia, e Psicologia Comp., Florence. 
(PLATE XII.) 


Ir is a well-known fact that for many years past the 
majority of descriptive anthropotomists, in describing the 
skull, have divided the bones composing it into those of the 
face and those of the skull. So that if we adopt the nomen- 
clature of Dr J. Barnard Davis, the valuable and interesting 
specimen which forms the subject of this paper would be 
considered a calvaria, as it lacks the lower maxilla. Accord- 
ing to this authority, too, a craniwm was regarded as being 
composed of the entire number of bones of the head and 
face, while the calvaria was made up of the bones of the 
skull alone. In these days, when the knowledge of the 
general structure and physiology of vertebrates has become 
absolutely indispensable to the anatomist, be his particular 
line of research what it may, such artificial landmarks are 
eradually becoming obsolete. 

At the best of times Navajo Indian skulls are difficult 
objects to obtain, so I considered myself particularly fortunate 
when some time during the early autumn of 1885 the present 
specimen came into my possession. It was collected by a 
young man on one of their burial grounds upon the hills in 
the vicinity of Fort Wingate, New Mexico, and handed to me 
immediately afterwards to make such use of it as I saw fit. 

This skull is from a male subject of about forty years of age, 
who came to his death by a gun-shot wound of the head. The 
results of this fatal injury are not far to seek in the specimen, 
and they may be seen in part in my illustrations of it in the 
Plate. 

We find the large wound of entrance has pierced the left 
outer angle of the supraoccipital bone, and destroyed the adja- 
cent mastoidal process of the pars mastoidea of the temporal, 
resulting in a magnificent example of that rare condition, a 


= 
-~ 


ON A NAVAJO SKULL. 427 


fracture by contre coup, the external appearance of which may 
be seen upon the right frontal bone in three of the figures. 

Five years ago, when Otis published the list of the human 
crania contained in the Anatomical Section of the United 
States Army Medical Museum, this extensive and unrivalled 
collection of several thousand specimens had in it but twenty- 
two, perfect or imperfect, Navajo Indian skulls, 

At present this collection is not available to me, but from 
the excellent catalogue in question, I am enabled to select cer- 
tain data of the highest value for comparison with similar 
observations made by myself upon the specimen in hand. 

Of the twenty-two skulls alluded to I have chosen eight as 
nearly perfect ones as possible, and of the same sex as ‘our 
subject, with a slight variation in age. From the data afforded 
by these in the catalogue, the averages exhibited in the sub- 
joined table, for comparison with corresponding ones in our 
specimen, have been computed. Very wisely, Dr Otis adopted 
the metric system in all of his measurements, and the same is 
employed here. At the headings of the several columns of the 
table I have used certain abbreviations; among these, where 
an explanation seems necessary, ‘“ Cran: cap. c.c.,” stands for 
cranial capacity in cubic centimetres ; C is the circumference 
taken (in my case) with a steel metric tape measure, upon the 
periphery of the figure formed by the plane which passes 
through the glabella, the occiput, and outstanding lateral points ; 
F is facial angle; L is longitudinal diameter, from glabella to 
highest point of occipital prominence ; H refers to the height, 
measured from the middle of the anterior margin of the foramen 
magnum to the highest point on the cranial vault; ZD, is the 
zygomatic diameter; and finally, B, the greatest transverse 
, parietal diameter, or the breadth. 

During the time Dr Otis lived (and it was my good fortune 
to see many of the human crania there measured while he had 
charge of that part of the Museum) No. 8 shot was the medium 
by which the internal capacity of the specimens was ascer- 
tained; but since then methods involving greater accuracy 
have been adopted. 

As I have already stated, circumferential measurements were 
obtained from the specimen in my hands by a flexible steel 


* 
%, : 


428 DR R. W. SHUFELDT 


tape measure, while diameters were taken with an accurate 


pair of calipers. In computing the cranial capacity, I filled 


all the foramina leading into the brain case so carefully 
that the fillings were made flush in every instance with 
the internal cranial wall. After this was done the skull 
would contain water without leaking, but instead of using 
that vehicle, I employed the shot known to collectors as “ dust 


Table of Averages. 


Museum No. |Age and| Cran. C. L. 156 Zia: B. 
of Specimen. Sex. |Cap. cc.} mm. f mm |mm.| mm, mm. 
2 788 35, 6. | 1480 OT Tie 180 137 142 148 
97 30,6. | 1190 479) | 9° 164 124 127 140 

130 45, 3 1335 AOD | Wits 168 133 129 144 

18 30, 6 1480 511 ssl 183 142 137 

633 30, 3 1325 499 | 76° 175 138 125 130 
784 40, 3 1550 5138 | 74 182 144 139 136 

785 55, 3 1435 490 | 77° 163 140 138 148 
1086 45, 3 1560 521 | 82° 187 138 143 142 


Averages. 39+ 1419+) 503+] 77°+}) 175+) 187 134+} 140+ 


Data from the 
present 40+ x 1520 527 80° 169 151 150 160 


Specimen. 


shot” (smaller than No. 12). It will at once be seen that this 
gives a higher figure, yet at the same time a more accurate re- 
sult than No, 8 shot can do, as used by Dr Otis. Quicksilver 
would come still nearer the mark, as it would perfectly accom- 
modate itself to all the surfaces of the walls bounding the 
cranial casket within. In consulting the above “ Table of 
Averages,” the fact that I used smaller-sized shot in taking the 
cranial capacity must be borne in mind. 

Next, with an excellent camera, which I possess and employ 
to do work very similar to the present, I photographed this cal- 
varia in four different positions, and from these photographs, 
by a process in which inaccuracy is reduced to a minimum, I 
made the four drawings presented in the Plate. 

So far as any single specimen of a skull can be, this one, no 
doubt, is typical of the skull as found in the Navajo Indians 
at least for the adult male portion of them. 

In past times the field of study in human craniology was 


- 


ON A NAVAJO SKULL. 429 


rather an unsatisfactory and discouraging one, owing to the fact 
that the greatest diversity was found to exist among skulls re- 
presenting individuals of the same race; but now, thanks to 
composite photography, the vista which opens before us is far 
more hopeful. Especially is this true of Galton’s methods of 
comparing the components with the composite. After the ideal 
composite has been formed by the proper employment of the 
eight types of components, we then have the starting point from 
which generalised data may be deduced, and subjects such as 
the present can be advantageously compared. By comparing 
the figures with the data in the table, and the latter for the 
several specimens introduced, much may be brought out of a 
very suggestive nature, even in such a brief sketch as I here 
offer—indeed, a great deal which falls without its proper scope, 
as originally intended, which was simply to illustrate by mea- 
surements and drawings the leading characteristics of an adult 
male skull of a representative of one of the most prominent 
tribes of existing North American Indians. 


DESCRIPTION OF PLATE XII. 


Fig. 1. Calvaria of Navajo Indian, ad. ¢, nearly direct anterior 
aspect, considerably reduced from the original. 

Fig. 2. The same specimen seen from above. 

Fig. 3. A rear view of the same upon a somewhat larger scale than 
employed in figs. 1 and 2. 

Fig. 4. The same specimen, lateral view, the scale being the same 
as in fig. 3. 


ADDITIONAL NOTE ON THE NAVAJO INDIAN SKULL. 
By Professor Sir WILLIAM TurRNER, M.B., F.R.S. 


As Dr Shufeldt has very kindly sent me the Navajo skull, 
described in his paper, for presentation to the Anatomical 
Museum of the University of Edinburgh, I have had the oppor- 
tunity of examining this interesting specimen, and wish to 
supplement his description with a few additional particulars. 

The skull presented a well-marked parieto-occipital flattening, 
obviously due to artificial pressure, which had been applied so 
as to cause the suprasquamous part of the occipital bone and the 
posterior ?ths of the parietal to slope upwards and forwards. 
The frontal region did not exhibit any flattening, so that in this 
individual, and it may be in his tribe of Indians, the pressure 
applied in infancy was apparently limited to the back of the head. 
Owing to this artificial distortion, the longitudinal diameter of 
the head was diminished, and the cephalic index, 94°6, computed 
from Dr Shufeldt’s measurements of the length and breadth, was 
therefore higher than it would have been in an undeformed skull. 
The cranium was hyperbrachycephalic. 

The height of the skull was also very considerable, and reached, 


as may be seen from the table, 151 mm.; the vertical index was © 


89, so that the skull was hyperakrocephalic. In all probability 
the pressure during infancy, which shortened the skull in its 
antero-posterior direction, forced the vertex upwards and added 
to the height of the cranium, so that the high vertical index was 
occasioned both by diminished length and increased height. 

The skull was cryptozygous, for not only was the breadth in 
the parietal region great, but the stephanic diameter was 137 mm. 
The glabella was not very prominent, but the supraciliary ridges 
were thick and strong. The bridge of the nose was concave 
forward, so that the tip projected to the front. The basi-nasal 
diameter was 105 mm.; the basi-alveolar 98 mm.; the gnathic 
index was 93, and the skull was orthognathic. The nasal spine 
of the superior maxilla was moderate. Where the side walls of 
the anterior nares joined the floor, the margin of the opening 


- 


ADDITIONAL NOTE ON THE NAVAJO INDIAN SKULL. 431 


was rounded. The transverse diameter of the orbit was 40 mm.; 
the vertical diameter 36 mm.; the orbital index was 90, and the 
orbit was megaseme. The nasal height was 48 mm.; the nasal 
width 25 mm.; the nasal index was 52, and the nose was meso- 
rhine, The palato-maxillary length was 56, the palato-maxillary 
width was 72 mm.; the palato-maxillary index was 128, and 
the roof of the mouth was brachyuranic.!. The teeth were all 
erupted and not much worn, The cranial sutures were all un- 
ossified. The parieto-sphenoid suture in the pterion was 19 mm. 
in antero-posterior diameter. There were no Wormian bones. 
The anterior end of the inferior turbinated bone was almost in 
the same plane as the anterior nares. 


1 See my Report on Human Crania in Reports of ‘‘ Challenger” Expedition, 
part xxix., 1884, for the introduction of this term. 


ON THE ORIGIN OF CERTAIN CYSTS—OVARIAN, 
VAGINAL, SACRAL, LINGUAL, AND TRACHEAL. 
By J. Buanp SurTon, F.R.CS., Hrasmus Wilson Lecturer 
on Pathology, Royal College of Surgeons, England. (PLATE 
XTi 


In the present communication the mode of origin of certain 
forms of ovarian cysts will be considered, also the relation 
of Gartner’s ducts to vaginal cysts, and the part played by the 
post-anal gut in producing congenital sacral cystic tumours. 
Some remarks will also be made on lingual and tracheal cysts. 

The material which has supplied me with the facts to be 
recorded in this communication concerning the ovaries and 
Gartner’s ducts consisted of the uterus and its appendages 
taken from seventy cows, varying in age from nine months to 
ten years. Many of these were virgins or heifers, others 
had borne several calves, and many of the specimens were 
pregnant at the time of dissection, These uteri were obtained 
from animals slaughtered for the purposes of food; all the 
specimens were taken consecutively, so as to represent as far 
as possible average conditions. 

At the same time, and under precisely similar conditions, 
the uterus and appendages were examined from fifty sows. 
Also the generative organs of twenty aged mares, in addition to 
fifty reported upon last year. | 

These specimens are supplementary to the great number 
of animals which come under my observation at the gardens 
of the Zoological Society. 

The dissection of these specimens was undertaken for the 
purpose to trace, if possible, the mode of origin of ovarian 
cysts, and the relation of Gartner’s ducts to cysts of the broad 
ligament, and the upper part of the vagina. 


Ovarian Cysts. 


[ do not propose to spend time in discussing the various 
theories that have been raised to explain the mode of origin 
of ovarian cysts. Nearly everything that enters into the com- 


ae 


a 


ON THE ORIGIN OF CERTAIN CYSTS. 433 


position of this wonderful organ has been pressed into the 
service. Briefly, my aim is to show in no uncertain way the 
mode of origin of some of the cysts. 

In this Journal (vol. xix. p. 139) I drew attention to the 
fact that two-thirds of all mares that attain the age of eight 
years present cystic ovaries, the cysts varying in size 
from a grape to an orange: it was also stated that some of 
these cysts probably arise in Graafian follicles, and that others 
are of parovarian origin. After the publication of the paper 
referred to, I still pursued the investigation, and was fortunate 
enough to encounter in the ovary of a mare, a cyst of the size 
of a nut occupying the centre of a large corpus luteum. This 
specimen is figured and described in the Zrans. Path. Soc., 
vol. xxxvi. This case seemed to support the view that there 
is reason to believe in the origin of some ovarian cysts in 
degenerate corpora lutea, that I resolved to pursue the ques- 
tion in the ovaries of cows, for, of all domestic animals, they 
seem to be the most liable to ovarian cysts. Accordingly, 
seventy sets of generative organs were obtained from these 
animals. 

The result of the investigation was eminently satisfactory, 
for it goes to show in a most conclusive manner that ovarian 
cysts do arise in corpora. lutea. 


In fig. 1 (p. 434) the various stages are represented as follows :— 


A.—A section through a normal] ovary from a heifer; the small 
cysts are immature Graafian follicles. 


B.—A corpus luteum of pregnancy. The uterus contained a calf 
about two months old in the corresponding uterine cornu. 


C.—The centre of the corpus luteum in this case is occupied by 
two distinct cavities, lined by a delicate membrane, and con- 
taining fluid. Two distinct cavities are only occasionally 
present in one corpus luteum. 


D.—In this stage the cavity has attained the size of a nut; two 
old cysts of similar organ are also seen in the ovary. 


E.—The cyst here represented was of the size of an orange, and 
filled with fluid. Its relation to the preceding cysts is de- 
monstrated indubitably by the circumstance that the same 
yellow-coloured material forms a lining to the cyst immedi- 
ately beneath the peritoneal investment. 


434 MR J. BLAND SUTTON. 


In addition to the principal cyst, a number of smaller ones 
may be seen on its sides. It is important to draw attention to 
this, for they may be mistaken to represent secondary cysts, 
whereas in reality they are cysts similarly developed in old 


Fic. 1.—A series of drawings to show the origin of ovarian cysts in corpora lutea. 
Figs. A to E are from the ovaries of cows. F represents a cyst which ot- 


curred in the ovary of a tiger. 


corpora lutea, but quite independent of the larger one; they 
form part of the cyst wall because they, with the ovary, became 
flattened out and incorporated with the cyst walls, due to the 
pressure of the abnormal mass. I have been able thus to 
account for similar multiple cysts in the human female. 


ON THE ORIGIN OF CERTAIN CYSTS. 435 


This series supplies evidence beyond all dispute of the 
gradual formation of a cyst from a corpus luteum. Evidence 
is not wanting to show that if two or more corpora lutea in 
the same ovary become cystic, and this is far from being 
uncommon, a multilocular cystoma may arise. To me, the 
most satisfactory feature of the investigation is this,—anyone 
with a little patience may obtain twenty-five cows’ ovaries and 
demonstrate all the stages represented in fig. 1 with the greatest 
of ease, and without any other assistance than that afforded 
by a sharp knife. 

Besides in cows, the process may be traced in mares and in 
pigs, but not so readily as in cows. Cysts also occur after the 
same fashion in tigers, and one of fair proportions from this 
animal is shown in woodcut 1, F. Other observers have suggested 
that the same process holds good for human females, and I have 
seen specimens which support these observations in women and 
in monkeys. 

One other question suggests itself. In the specimens figured 
the cysts occurred for the most part in animals which had borne 
young. Do the same statements hold good for virgins ? It may be 
mentioned that old animals or those which had borne young were 
selected, because the corpora lutea were larger, and illustrated 
the condition more clearly, but virgins are also liable to precisely 
the same changes in all the species dissected. 

It is now essential that we should endeavour to explain why 
these bodies should soften in the centre and form cysts. For- 
tunately a very satisfactory explanation is forthcoming. In the 
ripe ovarian follicle we can distinguish a distinct wall derived 
from, and continuous with, the stroma of the ovary. The inner 
aspect of the boundary wall is lined by layers of epithelial cells 
forming the membrana granulosa. Similar cells surround the 
ovum, but are polyhedral or columnar in shape; a collection of 
cells usually forms at one part known as the discus proligerus. 
When the ovum escapes from the follicle the cavity becomes 
filled with effused blood. After a time (about fourteen days) 
the blood clot is in a great measure replaced by a material of a 
distinct yellow colour, forming the well-known corpus luteum, 
This tissue is present in corpora lutea of menstruation and of 
pregnancy, the only difference being one of quantity. 


436 MR J. BLAND SUTTON. 


It is by no means a decided question as to the source of the 
yellow material. Under the microscope it presents cells of 
various shapes and sizes intersected by capillary blood-vessels 
derived from the wall of the follicle, the tissue according to one 
observer resembling the cortical substance of the supra-renal 
capsules. ‘There are three possible sources of this tissue :— 

(1) It may arise from proliferation of the cells of the mem- 
brana granulosa, or (2) be derived from the follicular wall; (3) 
it may result from the organisation of the blood-clot. 

For my own part I think it is due to the two last causes 
combined. Certainly the clot is in an exceedingly favourable 
condition for organisation. Whichever view is the true one the 
following fact’ remains inviolate, viz.; the centre of a corpus 
luteum is usually occupied by the débris of the original clot, 
which is in a gelatinous condition, the material becomes sur- 
rounded by a delicate membrane formed by the coalescence of 
some of the cells composing the yellow tissue. J¢ is by sub- 
sequent distension of the space thus formed that cysts arise so 
Srequently wn these bodies. 

With regard to the ovaries of the sows it is unnecessary to 
enter into details. The origin of cysts from degenerate corpora 
lutea could be traced as easily in them as in the cows; with the 
exception that the cysts do not attain so large a size relatively. 
Fig. 2 represents the ovary of a sow, showing numerous cysts 


Fic, 2.—Ovary and Fallopian tube Fie, 3.—An ovary and Fallopian tube of 
of a sow. a sow, showing hydrosalpinx. 


which have arisen in corpora lutea, for in the case of these 
animals six or seven ova escape from each ovary at the time 
of heat, producing as many corpora lutea. In fig. 3 is repre- 
sented also a sow’s ovary studded with cysts, and at one end of 


ON THE ORIGIN OF CERTAIN CYSTS. 437 


the organ the Fallopian tube has become firmly attached by 
strong adhesions, and in such » way that its ostium abdominale 
is completely occluded. As a result of this, the tube has be- 
come dilated so that the little finger could easily be accommo- 
dated in its interior. The natural dimensions of the tube are 
represented in fig. 2. 

On carefully examining the Fallopian tubes of all the sows 
(fifty in number) three well-marked cases of distention of the 
tube were found, constituting veritable cases of hydrosalpinx. 
In all cases the abdominal end was firmly adherent to the 
ovary, and completely occluded. In my previous paper a case 
of pyosalpinx was described which occurred in a kangaroo; a 
second specimen has since come to hand. Now that the affec- 
tion has been found also in pigs and in the cow, there can be 
little doubt that, if carefully looked for in other mammals, by 
those who have opportunities, other cases will doubtless come to 
light. 

Before leaving the ovary it will be of interest to consider 
further certain details connected with the “tissue of the hilum.” 
For several months I have systematically examined the ovaries 
of every female human fetus, from the eighth to ninth month 
of intra-uterine life, which has come into my room for dissection. 
The ovaries were preserved entire in Miiller’s fluid in the usual 
way, and when properly hardened were cut into sections for the 
microscope. In this work I was fortunate enough to have the 
assistance of three of my pupils—Messrs Sibley, Nash, and 
Marris—in cutting and mounting many of the specimens. 

The results of this work will be briefly stated. In all cases 
the “ tissue of the hilum” was very conspicuous, but varied con- 
siderably in amount. It consists chiefly of connective tissue, 
containing a very large number of veins. These veins, later in 
life, may undergo abnormal dilatation, constituting varicocele in 
the female, which is in every respect analogous to the same con- 
dition occurring in relation with the testicle. 

In the majority of the ovaries exainined, the tubules consti- 
tuting the parovarium could be traced with ease into the sub- 
stance of the ovary, but they were in the majority of cases 
entirely confined to the hilum of the organ. Associated with 
the tissue of the hilum are relics of the mesonephros (Wolffian 


438 MR J. BLAND SUTTON. 


body), named by Waldeyer “the paroophoron.” The paroophoron 
agrees histologically as well as in the matter of development with 
the paradidymis (organ of Giraldés.) Critical examination of 
the “tissue of the hilum” has served to convince me, not 
merely that it contains remains of the mesonephros, but that 
the connective tissue composing it is derived directly from the 
degeneration of that organ. 

It occasionally happens that at birth an ovary will be found 
divided by a deep longitudinal furrow. In some of these speci- 
mens the furrow actually divides the ovary into two histologi- 
cally distinct parts—parenchyma, and mesonephritic remains. 
This is very well seen on Plate XIII. fig. 1, where the part P 
represents true ovarian stroma, and O a very large paroophoron : 
the section being carried across the two edges of the longitudinal 
furrow before referred to, 

In one case in which the ovary was found to be cystic, some 
of the cysts being as large as a pea, the remains of the meso- 
nephros exceeded the bulk of the true ovarian stroma three 
times. 

The transverse section (that is, in the direction of the minor 
axis) of the foetal ovary on Plate XIII. fig. 2, represents another 
example of an excessive amount of mesonephritic tissue. In this 
instance the foetus was anencephalic. In this ovary, as well as 
in the one represented in fig. 1, the distinction between the 
two parts was so obvious that it was easy to determine the 
parenchyma from the “tissue of the hilum,” when the sections 
were made transparent and fixed upon a slide. 

The relation of the various parts of the ovary to each other, 
and to cystic formations occurring in that organ, are represented 
in Plate XIII. fig. 3. This figure has been compounded from a 
series of sections of a human ovary at the ninth month. It is 
intended to show the parenchyma, with a ripe ovum lying in a 
follicle, also a corpus luteum. Running into the hilum are 
several parovarian tubules, and lying among them is an incipient 
papillary cyst, which is prone to occur in this part of the ovary. 

Seeing that the “ tissue of the hilum” really has its origin in, 
and often contains the remains of part of the mesonephros, 
and is really equivalent to the structure named by Waldeyer 
“ paroophoron,” it seems unnecessary to employ the term “ tissue 


ON THE ORIGIN OF CERTAIN CYSTS. 439 


of the hilum,” but to regard the ovary as being compounded of 
two parts, oophoron and paroophoron. 

The investigation has not only been interesting concerning 
the relation of the tissue of the hilum to the paroophoron, but it 
surprised me in the very large proportion of foetal ovaries which 
present a cystic condition at birth. I have already seen seven 
examples, out of a total of more than forty foetuses, which have 
been specially examined in regard to this point. After birth, 
doubtless, many of these cysts disappear, or cause no trouble ; 
nevertheless, there can be little doubt that many ovarian cysto- 
mata which later in life give trouble, had already commenced 
their growth in the ovary at the time the unfortunate individuals 
who possess them commenced their extra-uterine existence. 

Broap LicamMent Cysts.—In a criticism on my previous 
communication regarding cysts in connection with the repro- 
ductive organs of animals, in this Journal, vol. xix. p. 121, 
Mr Doran suggested that in all probability some of the cysts 
described as parovarian in origin might really be simple broad 
ligament cysts, unconnected with the parovarian tubules. An 
examination of some fresh ovaries of mares has convinced me 
that some of the cysts found in the mesosalpinx are uncon- 
nected with the parovarium. On Plate XIIL fig. 4,is shown the 
Fallopian tube of a mare with three of these cysts in situ: 
they were situated between the layers of the mesosalpinx, 
possessed thin walls, and were filled with fluid. Running 
among the cysts and in relation with one of them are large 
dilated lymphatic vessels. These lymphatic vessels in cases of 
obstruction caused by cysts are easy of detection, for imme- 
diately after death the lymph coagulates, and they become dis- 
tended as by a natural injection. Probably many, but not all, 
simple broad ligament cysts arise from dilatation of lymphatic 
channels. 


Gartner’s or Gaeriner’s Ducts. 


Whilst engaged in examining the generative organs of cows, 
I seized the opportunity of further testing the question as to the 
probability of cysts of the broad ligaments and of the upper 
part of the vagina arising in these ducts or their rudiments, 

The material was exceedingly favourable for the investigation, 


440 MR J. BLAND SUTTON. 


because in cows these ducts attain, so far as female mammals 
are concerned, a very high development. Among the seventy 
specimens the following variations in the condition of Giirtner’s 
duct were met with. 

In all young animals both ducts were present as pale streaks 
on the ventral wall of the uterus, immediately beneath the peri- 
toneum. In the majority of the specimens they became gradu- 
ally incorporated with the tissue of the cervix uteri, their lumen 
gradually suffering obliteration at this point; in some cases, but 
quite exceptionally, they opened on the mucous surface of the 
vagina, midway between the os uteri externum and the meatus 
urinarius. Anteriorly these tubes were continuous with the 
longitudinal tube of the parovarium. In older cows the ducts 
in some cases disappear in various parts of their course, in a few 
instances completely. The persistent portions of the tube, how- 
ever, dilate, especially in the neighbourhood of the cervix. and 
vagina, forming tubulo-cysts, which in some cases are as large as an 
orange, and bulge intothe vagina. In cows which have borne 
several calves, the ducts in relation with the cervix exhibit 
extreme thickening of the walls, as though from chronic inflam- 
mation. If the ducts remain pervious throughout their whole 
length they increase in dimensions with the uterus, often attain 
the size of a quill pen, and are filled with fluid resembling serum. 

There is yet one other point of some interest to be considered 
in relation to the mode of termination of these ducts in the 
vagina. Anatomists had long been of opinion that two minute 
openings in the human vagina, near the opening of the urethra, 
represented the terminal orifices of Gartner’s ducts, which have 
become familiar as Skene’s tubes. Further attention was centred 
upon them when it was found that they are liable to inflame and 
lead to trouble in the human subject. Recently, however, the 
relation of the orifices to Girtner’s ducts has been questioned by 
Dr Schiiller? and others, and the openings have been described 
as belonging to two glands lying in the wall of the vagina, near 
the termination of Girtner’s ducts. A knowledge of these 
opinions induced me to write cautiously on the mode of termi- 
nation of the ducts in my previous papers on the subject, until 


1 See Doran, Tumours of the Ovary, page 43. 


ON THE ORIGIN OF CERTAIN CYSTS. 441 


leisure and opportunity should enable me to inquire into the 
matter afresh. 

It is a fact thoroughly established that the vasa deferentia of 
the testicles are the male representatives of the ducts of Gart- 
ner in the female. Immediately before the vasa terminate on 
the floor of the urethra two diverticula are found, the vesiculz 
seminales. In virgin cows a similar but very much smaller 
diverticulum arises from the end of Gartner’s duct, and is situated 
in the wall of the vagina. In very many cases the duct, im- 
mediately above the glandular diverticulum, often becomes 
obsolete as mentioned before. Under such conditions a thin 
probe passed into the orifice of the duct when existing would 
lead into the gland, but not into the duct above, and thus give 
rise to the fallacy of the independence of the gland in question 
from Giartner’s duct. See woodcut +. 

In this diagram (1) represents the vas deferens in its relation 
to the vesicula seminalis. (2) 
Shows the duct of Gartner and 
the glandular diverticulum I have 
described, the diverticulum bear- 
ing the same relation to the duct 
as the vesicula seminalis does to 
the vas in the male. (38) This is 
intended to represent a not un- Fic. 4. 
common condition of the glands 1. The Vas deferens and vesicula semi- 
in question; the tube becoming lis. 2. Gartner's duct and its 
: : : xe diverticulum. 3, The terminal 
ee yun immediately above the orifice and diverticulum of Girt- 
spot where the gland-duct joins ner’s duct in its most common 
thetube. Thisarrangementclearly condition. 
explains the apparent independence of the gland and Girtner’s 
duct. Judging from the number of specimens I have dissected, 
it is easy to understand, from the frequent amount of mucoid 
fluid they contain, that they could easily become involved in 
local catarrhal inflammation, and lead to considerable annoyance. 
But that they are independent of the ducts of Gartner is 
erroneous. 

From the foregoing statements the following are the chief in- 
ferences which may be drawn :— 


1. In cows, mares, pigs, tigers, goats, and human beings, some 
VOL. XX, 2 F 


442 MR J. BLAND SUTTON. 


cysts of the ovaries arise in corpora lutea either of pregnancy 


or of menstruation. Waldeyer, Beigel, and De Sinéty have 


pointed out that even before birth the ova ripen, atrophy, and 
form a kind of corpus luteum. I have had ample opportunities 
of confirming these statements, and have convinced myself that 
the process goes on throughout childhood, and gives rise to cysts. 

2. Pigs and cows, as well as kangaroos, are occasionally liable 
to dilatation of the Fallopian tubes due to adhesion of the 
fimbriated end to the ovary, Hydrosalpinz. 

3. Giirtner’s ducts are potential sources of vaginal cysts. 

4, Skene’s tubes (so-called) are diverticula of Gartner’s ducts, 
and represent in the female the vesiculze seminales of the male. 


Sacral Cystic Tumours. 

A vast number of tumours, varying in character from a simple 
cyst to a more or less fully formed individual, have been placed 
on record as occurring in connection with the sacral and coccy- 
geal regions of human beings. The term sacral tumour has been 
used by pathologists in a generic sense to include all the 
varieties of these singular formations. 

Within the last few years some observations have been 
recorded which tend to throw some definite light on these cases 
and recent advances in the science of embryology, especially in 
connection with the caudal extremity of the embryo, offer clear 
explanations of the cystic varieties of sacral tumours. 

In the first place it must be pointed out that it is necessary 
to separate the various tumours into classes. 

Braune, who has collected a vast number of these cases 
(95 in all) in his admirable monograph, Die Doppelbildungen 
und angeborenen Geschwiilste der Kreuzbeingegend, 1862, divides 
them thus :—(1) Parasitic, or attached foetuses; (2) Hygromata, 
or cysts; (3) Lipomata. The first of these three groups I do not 
propose to consider, but the cystic formations and lipomata will 
claim especial attention. 

Braune clearly recognised that among the sacral hygromata 
we have to deal with two distinct varieties, one form arising as 
a dilatation of the caudal extremity of the spinal meninges, the 
other he endeavoured to account for rather fancifully, as arising 
from the degeneration of Luschka’s coccygeal gland. 


ON THE ORIGIN OF CERTAIN CYSTS. 445 


The first type case is from a preparation preserved in the 
Meckelian Museum at Halle. The tumour was attached to the 
lower sacral region, and measured 10 c.m. in length, 6 cm. in 
breadth, and hung pendulant by a pedicle 1 c.m. in thickness. 
The details of its relation to the vertebral column are shown in 
fig. 5. An accurate dissection of the relation of the coverings 


Fic. 5.—A congenital sacral cystic Fic. 6. — Congenital cystic sacral 
tumour due to dilatation of the tumour lying anterior to the sa- 
terminal portion of the spinal crum. (After Braune.) 7.2., inno- 
meninges. (After Braune.) s.c., minate bone; s.c., spinal canal ; 
spinal canal ; p.m., dura mater. L.A., levator ani; G, gut. 


of the tumour to the fascia, &c., of the trunk was not possible 
on account of the long soaking of the specimen in spirit. The 
following points, however, could be made out with certainty. 
The central cavity of the tumour was in relation with the spinal 
canal, as in the case of a spina bifida, but with this difference, 
the communication occurred through the normal Aiatus sacralis. 
The walls of the cyst were made up of skin, fascia, connective 


444 MR J. BLAND SUTTON. 


tissue, muscle fibre, and a fibrous lining to the cyst resembling 
the dura mater. : 

The second type case was also preserved in the Meckelian 
collection at Halle. It was a seven months’ female fcetus, 
which had long been preserved in alcohol. The anatomical 
details are shown in woodcut 6. In this instance the tumour 
is unconnected with the spinal canal, which is completely closed, 
and lies on the anterior (ventral) aspect of the coccyx. Between 
it and the rectum is the levator ani muscle, and a fibrous 
envelope which surrounds it is directly continuous with the 
coccygeal periosteum. The centre of the mass is traversed by 
trabecule giving rise to a cystic condition. It is this form of 
sacral congenital tumour that is often described as sarcomatous. 
In this case Braune considers the tumour to have arisen as a 
result of the degeneration of Luschka’s coccygeal gland; in this 
opinion he is followed by others. This view, however, is 
rendered somewhat untenable by the investigations of embry- 
ologists in relation to the post-anal gut. 

In adult animal forms it is the normal condition that the 
rectum should terminate by an external opening called the anus. 
In 1871 Kowalevsky! drew attention to the remarkable circum- 
stance that the alimentary canal is brought into direct communi- 
cation with the central canal of the spinal cord by means of a 
passage known as the neurenteric canal, which turns round the 
posterior end of the notochord. He? also pointed out the 
interesting fact that primarily the gut is continued posteriorly 
beyond the anus, this prolonged portion being named by Balfour 
the “post-anal gut.” The general relation of the anus, post- 
anal gut, neurenteric, and neural canals to one another are repre- 
sented in fig. 7. 

Subsequent researches show that this arrangement of the 
primary intestinal canal probably holds good for all Chordata 
from amphioxus up to man. 

Since the announcement of its discovery by the great Russian 
researcher it has been found in Acipenser, Axolotl, Bombinator, 
Plagiostomi, Teleostei, the hen, rabbit, and in the human embryo. 
These later observations are of great interest, for on the first dis- 


1 Archiv fiir Mikr. Anat., Bd. vii. S. 114. 
2 Tbid., Bd. xiii. S. 194, 195. 


ON THE ORIGIN OF CERTAIN CYSTS. 445 


covery of the neurenteric canal it was thought to be a peculiar 
Ichthyopsidian character. Careful observations on various forms 
of the higher vertebrata go to show that a neurenteric passage 


Fic. 7.—Longitudinal section of an embryo Bombinator igneus. A, anus; B, 
brain ; N, notochord ; M, mouth; 1, liver; N.c., neurenteric canal ; M.c., 
medullary canal (after Goette). 


and a post-anal gut are characters perhaps as constant as a 
notochord. 

For a general review of the literature of this interesting 
subject, and a detailed account of the embryology and structural 
characters of the part, the reader should consult Balfour’s 
Embryology, vol. 11. pp. 267 and 634.1 The points most import- 
ant for our present purpose are the following :— 

The post-anal gut is always better developed in the lower 
than in the higher forms of vertebrate life. In Elasmobranchs 
it is very well developed, and persists during a considerable 
portion of embryonic life, and at one period presents itself as 
a terminal vesicle at the end of the tail, but communicates by 
a narrow neck with the neurenteric canal. These curious 
relations of parts have led embryologists into some very 
interesting speculations regarding the original position of the 
anus, but interesting as they are, we cannot further follow them 
in connection with this subject. The first attempt to asso- 
ciate the origin of sacral cysts with post-anal gut was that by 
Middeldorpf,? who, in reporting a case of congenital sacral cyst, 
consulted Wiedersheim in relation to the development of the 
parts in animals. This induced him, in the paper mentioned, 
to connect the cyst with developmental conditions, and he further 

1 Balfour's Development of Elasmobranch Fishes, p. 91 and 218, may‘be con- 


sulted with advantage. 
? Virchow Arch., Bd. 101, p. 37. 


446 MR J. BLAND SUTTON. 


points out that abnormal dilatations of this section of the gut 
belong to the same category as cases of abnormal persistence, 
and in many cases, undue dilatation of a vitelline duct. 

The lining tissue of the cyst was also confirmatory of the 
opinion of Middledorpf as to the origin of the cyst, for it not 
only presented a similar structure to gut under the microscope, 
but solitary follicles were also present in great quantity. 

This case is an eminently satisfactory one, but it only 
disposes of a part of the question; nevertheless it proves that 
certain cystic sacral tumours may arise in connection with the 
post-anal section of the alimentary canal. We must now con- 
sider some of the arguments which associate a few of these 
cystic tumours with the medullary canal. 

Virchow’s Archiv, Bd. c. s. 571, contains an article written 
by the great pathologist who edits the Archiv, entitled 
“ Ueber einen Fall von Hygroma cysticum glutaeale congenitum.” 
In this instance Virchow received from Dr Ludwig Wolff a 
tumour, apparently a lipoma, which had been extirpated from 
the gluteal region of a new-born negro child. Its appearance 
and situation is represented in the accompanying drawing 
(fig. 8), modified from the original figure. It is in shape 


Fic, 8 . — Congeni- 
tal sacral cystic 
tumour, in a new- 
born negro-child. 
(Modified from 
Virchow. ) Fie. 9. 


like a large fig, and measures 7:3 c.m.in length and in width 
3°7 cm. On section the interior was found to be occupied by 
two cysts. The walls of the tumour were thick, and composed 
of skin externally, fibrous tissue, transversely striated muscle 
fibre, nerve fasciculi, and arteries with thick walls. The inner 
wall of the cyst was lined with flat granular epithelium, From 


ON THE ORIGIN OF CERTAIN CYSTS. 447 


a careful consideration of the case Virchow comes to the con- 
clusion that we have to deal, notwithstanding its lateral situation, 
with a derivative from the spinal sheath. As in the variety of 
these cysts previously considered, we have positive evidence that 
hygromata may be clerived from the hernial protrusions of spinal 
membranes, as in spinee bifida; the sac may become peduncu- 
lated, the pedicle finally close, and the sac become separated 
from the spinal sheath, as is shown in the specimen represented 
in the diagram (fig. 9). 

The case from which this diagram has been constructed was 
the remarkable one described by Mr Solly (Med. and Chir. Soc. 
Trans., vol. xl. p. 19). The tumour was noticed at birth, but 
on account of its connection with the spinal canal its removal 
was deferred until the patient was twenty-nine years old. It 
was satisfactorily removed by Mr Solly. It was united by a 
ligamentous pedicle to the spine, but no communication existed 
with the spinal canal. The section of the tumour exactly re- 
sembled the pedunculated lipomata occurring in connection with 
the sacrum. Three figures illustrate the case in the original 
paper, but the details are represented diagramatically in fig. 9. 

This second variety of sacral cyst arises in connection with 
the structures on the dorsal aspect of the neurenteric canal. 
These two cases, Middledorpf and Virchow’s, confirm Braune’s 
views, that one form of these cysts lie anterior to the coccyx, and 
the other posterior, exactly as represented in figs. 5 and 6, 
and testify to the correctness of the view that the first variety 
arises as a distension of the spinal meninges» Whether, in as- 
cribing the second variety to a degeneration of Luschka’s gland, 
Braune is in keeping with facts must now be discussed. The 
arguments against this view of their origin are the following :— 

(1) Luschka’s gland is composed chiefly of a coil of blood- 
vessels and connective tissue. Congenital sacral tumours 
contain few blood-vessels. 

(2) In carefully-examined specimens of the congenital sacral 
tumours of the cystic variety, the cysts and duct-like passages 
are lined for the most part with cubical epithelium, and are held 
together by young connective tissue. The epithelium is usually 
columnar. These two circumstances alone are sufficient to 
exclude the coccygeal gland as the germ which gives rise to 


448 MR J. BLAND SUTTON. 


these tumours. On the other hand, the structure of the post- 


anal gut eminently accords with the histological details of the 


cystic masses in question. 

(3) When we remember the tendency functionless ducts and 
tubules have to become cystic, and consider the position and often 
pedunculated but usually impervious connection these tumours 
have with the rectum, and the close agreement of the epithelial 
lining in the two cases, it seems to me that the evidence as to 
the origin of these cysts in the post-anal gut is beyond doubt, 
and it is a more satisfactory explanation than calling in the aid 
of Luschka’s gland. 

We must now consider the third group—lipomata. Several 
instances have been collected by Braune and others of the occur- 
rence of congenital lipomata in the sacral region, and, as we saw in 
Virchow’s case, the growth was thought to be a lipoma by the 
operator who removed it. From a careful consideration of many 
recorded cases of fatty tumours occurring in unusual situations, 
and of several of which I have had opportunity of examining, I 
ventured to publish a paper in the 7rans. Med. and Chir. Soc., vol. 
Ixviii., dealing with the subject, and endeavoured to show that 
many of these uncommon lipomata are really to be regarded as 
being originally composed of higher tissues, such as muscle, nerve, 
&c., which, in consequence of having no function, have retrograded 
into fat. So in these examples of sacral lipomata; doubtless they 
originated as diverticula from the spinal meninges or post-anal 
gut, as the case may be, and the tissues covering them, muscles, 
nerves, &c., have degenerated into fat. 

Careful consideration of the anatomical and _ histological 
characters of these tumours seems clearly to indicate that one 
variety arises from a dilatation of the meninges of the spinal 
cord; the second variety is associated with the post-anal gut, 
and the third group or lipomata may arise in connection with 
either, but seems to be more frequently associated with the 
spinal variety. 

REFERENCES. 


In addition to the eighty-six cases of sacral tumours (hygromata, 
lipomata, spinal diverticula and coccygeal gland ? tumours) recotded 
by Braune in his classical work already referred to, the following cases 
may be mentioned :— 


ON THE ORIGIN OF CERTAIN CYSTS. 449 


Hurcurnson, Illustrations of Clinical Surgery, fas. xiii. This paper 
also contains several references. 


Saattock, Trans. Path. Soc., vol. xxxii. p. 197. 
Maonamara, Trans. Path. Soc., vol. xxxii. p. 199. 
Treves, Trans. Path. Soc., vol. xxxiii. p. 285. 
Wacstarre, St Zhomas’ Hosp. Rep., vol. iv. p. 213. 
Vircnow, Archiv, Bd. 100, p. 571. 

Mippteporrr, Virchow’s Archives, Bd. 101, p. 37. 
Hotmrs, System of Surgery, vol. iii. p. 780. 3rd edition. 


The standard works of Gurlt, Forster, Ahlfeld, and of Geoffrey St 
Hilaire contain numerous cases of this nature. 


Dermoid Cysts of the Tongue. 


In the preceding account of sacral congenital tumours it 
seems fairly evident that we have to deal with cystic formations 
arising in connection with “obsolete” canals—obsolete in the 
sense of being disused. There are many examples of these 
disused canals and passages in the body, eg., the pouch of 
Rathke, at the top of the pharynx; the infundibulum; the 
branchial arches; the central canal of the spinal cord; the 
neurenteric passage; the post-anal gut; the tubules of the 
Wolffian duct, &. 

In nearly all these passages we have a striking arrangement 
of parts, for these canals served during some period of embry- 
onic life to bring the three blastodermic layers, epi-, hypo-, and 
mesoblast, into intimate relation. 

In this section I shall confine my remarks particularly to 
dermoids of the tongue, for in this situation the relation of 
teratomata to obsolete canals is illustrated in a very striking 
and remarkable manner. 

The best account of lingual dermoid cysts, and one easy of 
access, is by Mr Barker, Trans. Clin. Soc., vol. xvi. p. 215. Mr 
Barker not only gives an interesting description of a case occur- 
ring in his practice, but has collected and analysed sixteen other 
recorded examples, and furnishes the following rules concerning 
them :— 


1. They may be unilateral, lying between the genio-hyo-glossi 
and the mylo-hyoid muscles, on one side or the other. 


450 MR J. BLAND SUTTON. 


2. They may be central, lying between the genio-hyo-glossi 

muscles. ; 

3. They may be bilateral, lying between the mylo-hyoid and 

genio-hyo-glossi muscles. 

Histologically they consist of tough fibrous walls, with a thin, 
smooth lining membrane. They contain sebaceous matter and 
embedded hairs, and perhaps teeth. 

Mr Butlin’ says of these cysts :—“ Cases in which their actual 
existence has been noticed soon after birth have been very few, 
and the very large majority of dermoid cysts of the mouth have 
been observed in adults, or at upwards of thirty years of age. 
One case has been described in which the patient was more than 
sixty years old.” I shall now proceed to show that dermoids in 
the tongue receive an explanation on anatomical grounds. 

Professor Wilhelm His, in a recent work, Anatomie mensch- 
lichen Embryonen, Heft. iii. p. 100, 1885, gives an account of a 
narrow duct lined with epithelium, which runs from the foramen 
cecum on the dorsum of the tongue downwards, between the 
genio-hyo-glossi muscles, to end blindly in the hollow of the 
basi-byoid. This is termed by His the lingual canal. Asso- 
ciated and often in connection with the duct, is a third or 
middle lobe to the thyroid gland. This, too, is in connection 
with a duct known as the thyroid canal. In embryonic life the 
two ducts are continuous, and constitute a canal known as the 
ductus thyreo-glossus. | 

The existence of these ducts admits of no doubt whatever. In 
the tongue of the human fcetus at birth a thin bristle may often 
be passed from the foramen cecum to the hollow of the basi- 
hyoid. In the adult it is best demonstrated by carefully remov- 
ing the body of the hyoid bone, as shown in Plate XIIL fig. 6, 
then carefully dissecting the cellular tissue lying between the 
genio-hyo-glossi, the duct, if present, is easily found. Up to 
the present I have searched for the lingual duct fourteen times, 
and found it present in a complete form thrice; in other speci- 
mens it could only be distinguished in parts of its course; but in 
the majority of cases no trace of it could be detected except the 
foramen cecum, and in some specimens even this recess was 
wanting. 

1 Diseases of the Tongue. 


ON THE ORIGIN OF CERTAIN CYSTS. 451 


The morphology of this duct I do not propose to consider at, 
present, for it will be essential to work out its leading features 
as far as possible in a variety of mammals. The interest it has 
for us in this paper is that, as in the case of sacral cungenital 
tumours and pituitary teratomata, dermoids of the tongue occur 
in the immediate neighbourhood of, if not in some way con- 
nected with, an obsolete canal, which traverses a large mass of 
mesoblastic tissue and brings the buccal epiblast and the hypo- 
blast of the branchial clefts into intimate relation. 

It is of course possible that these cysts arise in the same 
way as cutaneous proliferous cysts, occurring at the angle of 
the orbit, between the dura mater and the skull, and on the 
bodies of animals (see Plate XIII. fig. 5), viz., that they are in- 
voluted islands of superficial epiblast, which play the part of 
tumour germs. In the case of the skull we know that the skin 
and dura mater are in embryonic life in contact, an association 
which becomes disturbed when osseous material is deposited to 
form the skull. It is easy to see, as in a case reported by Pro- 
fessor Turner and more especially one by Dr Ogle,” that the 
eyst may have had its origin in small islands of skin left on the 
dura mater abnormally. In Professor Turner’s specimen, although 
there was no visible defect in the occipital bone, as in Dr Ogle’s 
specimen, nevertheless the cyst occurred at a spot where for a 
long period of intra-uterine life a defect or cleft exists in the 
squamo-occipital bone, and where skin and dura mater were in 
close relation. 


Tracheal Cysts. 


An interesting cyst came under my observation in an Emu, 
Dromeus nove-hollandie. Before entering into the details of 
the case it will be necessary to give a brief account of the normal 
anatomy of the trachea of this singular bird. 

If the integument be carefully dissected from the neck of an 
emu, a curious deficiency in the anterior wall of the trachea 
will be exposed, usuaily corresponding to a variable space be- 
tween the fiftieth and sixtieth rings. The general aspect of this 
singular opening may be fairly inferred from Plate XIII. fig. 7. 
Although the number of tracheal rings which are defective vary, 


l Barth. Hosp. Rept., vol. ii. 2 Path. Soc. Trans., vol. vi. p. 12. 


452 MR J. BLAND SUTTON. 


yet for the most part the gap rarely exceeds seven rings in longi- 


tudinal extent. The opening is about two and a half inches in ~ 


length. 

In connection with this aperture is a sac, varying in dimen- 
sions with different birds, according to age and sex. In an emu 
five weeks old I failed to find any evidence of a sac, the defect 
in the trachea being a mere slit; in the young emu the sac is of 
insignificant size, but exact observations are yet required to 
decide whether it differs in capacity in the male and female; 
but there is some probability that this is so. In adult birds of 
this species the sac attains considerable dimensions ; in one case, 
according to Knox, the cyst was as large as a man’s head. In 
Dr Murie’s case the sac measured fourteen inches in its greatest 
length. The walls of the sac are composed of white fibrous 
tissue, its exterior being overlaid in parts with striped muscle 
fibre in very thin layers; the inner layer is continuous with the 
mucous membrane of the trachea, and they become continuous 
with each other through the tracheal opening, indeed the sac 
could well be described as a “hernia of the tracheal mucous 
membrane through a deficiency in the anterior walls of the wind- 
pipe,’ for such it seems to be. As far as could be made out in 
my specimen, the epithelium was flattened on the surface, but 
spheroidal in the deeper layers. There were small glands 
(mucous) dotted over the membrane. During the pairing season 
the bird distends this sac with air, and it is due to the chamber thus 
formed that these birds are able to make the curious drumming 
noise so characteristic of them. For all that concerns the normal 
anatomy of this remarkable structure the reader must refer to 
Dr Murie’s admirable account in Proc. Zool. Soc., 1867, p. 405. 

Mr Bartlett invited me to examine an emu which had a large 
swelling in the neck, and as the bird refused food and seemed to 
be dying, it was deemed expedient to attempt its removal. 
After a severe tussle the bird was thrown; manipulation 
quickly informed one that it was a cyst with fluid contents, and 
an incision was promptly made, and the contents, which con- 
sisted entirely of mucus, evacuated. On attempting to arouse the 
bird, to our astonishment it was dead. Dissection made clear 
the cause of the fatality. The tracheal cyst, which has been 
described above, had inflamed and led to such an abundant 


ee 


ON THE ORIGIN OF CERTAIN CYSTS. 453 


secretion of mucus that the sac had become so distended as 
to measure in length 18 inches and in width 14 inches, 
When the bird was thrown for the purpose of laying open the 
cavity the dependent position had caused the mucus to flow into 
the trachea and obstruct the bird’s respiration; in fact it was 
drowned in its own mucus. 

The cyst in the emu’s neck is to be regarded as a hernia of the 
tracheal mucous membrane through a deficiency in the rings of 
that tube on its anterior aspect; in this sense it is very remarkable. 
Tracheal cysts in the human subject occasionally occur, but are 
usually found protruding through the muscular and fibrous tissue 
which fill up the deficient space on the posterior aspect of the 
trachea, and often form cavities of considerable size; in rare 
instances they become so large as to require surgical interfer- 
ence. Virchow quotes Textor as having operated in one of 
these cases with success.! 

That the mucous membrane of the trachea should become 
herniated in this way is not at all remarkable; every other 
similarly constructed tube is liable to the same abnormality. 
In the intestines false diverticula may be found. As many as 
two hundred have been seen in the same subject. 

In the bladder the corresponding condition is known as 
sacculation. The cesophagus is the seat of similar lesions often. 
In this Journal, vol. ix. p. 134, Dr Morrison Watson described a 
large diverticulum which was connected with the pharynx, due 
in all probability to the same cause. Indeed, any tubular 
structure may become so affected, be it intestine, artery, vein, 
ventricle of the brain, or even the spinal cord. 

The cyst of the, emu has other points of interest, for it is 
doubtless an example of a pathological condition which has 
been perpetuated, so as at length to become a specific character. 
That this cyst is the direct result of the imperfection of some 
portion of the tracheal rings cannot be doubted. In the young 
bird the cyst does not exist, but as life advances the hernial pro- 
trusion occurs. The defect in the trachea is inherited; the 
escape of the mucous membrane through the opening is the 
necessary result. 

Among all species of birds no example is known of a similar 


1 Tumours, French edit., vol. i, p. 263. 


454 MR J. BLAND SUTTON. 


arrangement of the trachea such as pertains in the Emu, but the 


Bustard (Otis tarda) affords valuable testimony in support of the - 


view here advocated regarding its origin. 

At certain times of the year the Australian Bustard (Hupo- 
dotis australis), and the Bustard (Otis tarda) exhibit the peculiar 
phenomenon termed “showing off” In both birds during the 
“show off” there is a distension of the neck with air. In the 
case of the Australian Bustard this tube is simply the dilated 
cesophagus, but in the case of Otis tarda, according to some 
observers, it is due to a special sac opening under the tongue. 

According to the evidence it appears that this gular pouch of 
the bustard is present in the adult male only, but is not a 
constant character. Garrod points out, however, that in the 
young bird there is a singular arrangement of the frenum 
lingue. In the young bustard this structure, instead of being 
median and single, consists of two slight lateral vertical folds, 
with a median interval a quarter of an inch across. This 
arrangement is exceedingly well adapted for permitting the 
development of a pouch during the sexual season; when the air 
passages are inflated during the showing-off, the fold of mucous 
membrane is very weak, and the continued pressure causes it 
to stretch. The statement concerning the pressure is not a 
gratuitous assumption, as the following case will show. When 
dissecting a young specimen of Otis tarda Garrod was surprised 
to find what appeared to him to be a crop projecting from 
the posterior aspect of the cesophagus. Further investigation 
revealed to him the fact that this was really an abnormal 
diverticulum of the cesophagus, probably produced by the same 
mechanism that was responsible for the gular pouch. 

In 1882 Mr W. A. Forbes rendered Garrod’s opinion more 
probable by finding in an Australian Duck (Siziwra lobata) a 
similar arrangement of the frenum lingue and a gular pouch 
in two specimens of the male, such as is found in the adult male 
of Otis tarda. 

Taking into consideration the anatomical arrangement of the 
freenum, the non-existence of the gular pouch in the young bird, 
its absence in the female, and its inconstancy in the male bird, 
as well as the evidence afforded by the two Australian birds 
(Eupodotis and Bizivra), it will not be inconsistent to regard the 


ON THE ORIGIN OF CERTAIN CYSTS. 455 


gular pouch as a pathological phenomenon brought about by the 
habit of “showing off,’ the tendency to cystic dilatation, or 
rather hernia, of this part of the buccal mucous membrane 
being inherited. 


Further details concerning the gular pouch of the bustard will be 
found on reference to the following papers :— 

Morte, Proceedings of the Zoological Society, 1868. 

Garrop, Proc. Zool. Soc., 1874, p. 471, and Collected Papers, p. 242. 

Forses, Proc. Zool. Soc., 1882, p. 455, and Collected Papers, p. 354. 

Dr Murie’s paper contains several other references of value. 


EXPLANATION OF PLATE XIII 


Fig 1. Longitudinal section of the ovary of an anencephalic human 
foetus at the eighth month. P., paroophoron; O., oophoron. The 
two parts were separated by a distinct furrow. [ x 10. 

Fig. 2. Transverse section of the ovary of a human feetus at birth. 
P., paroophoron, or remains of the mesonephros, unusually large; 
Q., oophoron, or true ovarian parenchyma. [ x 10.] 

Fig. 3. Diagrammatic section of an ovary. /., parovarian tubules, 
the seat of simple (parovarian) cysts ; Pa., paroophoron, the seat of 
papillary cysts, P.C.; O., oophoron, the seat of cysts arising in 
corpora lutea, C.LZ. A ripe follicle is shown at RF. 

Fig. 4. Simple cysts of the broad ligament ina mare. J, a dilated 
lymphatic. 

Fig. 5. Piliferous cyst from the back of a cow. 

Fig. 6. The root of the tongue and hyoid bone, to show the lingual 
duct. G., genio-hyo-glossus ; H.G.1., hyo-glossus muscle ; H., hyoid 
bone; 7.H.M., the thyro-hyoid membrane; 7.C., the thyroid carti- 
lage. Duct, the lingual duct. 

Fig. 7. Tracheal cyst of the Emu. The figure shows the tracheal 
window. 


THE BLOOD-FORMING ORGANS AND _ BLOOD-- 


FORMATION: AN EXPERIMENTAL RESEARCH. 
By Joun Locknart Gipson, M.D., Formerly Semor 
Demonstrator of Physiology, University of Edinburgh. 
(PLATE XIV.) 


(Continued from p. 353, vol. xx.) 


Function of Lymphatic Glands,—The function of the lym- 
phatic glands as regards the formation of red blood-corpuscles 
has in this paper already been alluded to; but it will now be 
necessary to consider an experiment which has a more direct 
bearing on this function. 

After thinking over what I wished to do on the blood and 
drawing up a plan of work, I consulted Professor Rutherford as 
to whether this plan would be likely to yield results. He was 
good enough to draw my attention more particularly to the 
possible action the lymphatic glands might take in the forma- 
tion of red blood-corpuscles, an action I had been inclined to 
doubt. He suggested that in this relation a lymphatic gland 
should be excised from one animal and planted under the skin 
of another animal, so that it might be nourished by the lymph 
of its host, while its own lymph would probably be confined in 
its interior; and that by this means one might be able to dis- 
cover whether the lymph corpuscles when confined in a lymph 
gland for a longer time than usual would show stages of transi- 
tion towards red corpuscles. 

On considering this suggestion, I thought its advantages 
might be obtained by another method of procedure, which, 
although much more difficult, would probably be more certain, 
and would allow of other observations being made at the same 
time. 

This other method was to tie the thoracic duct at its opening 
into the veins at the root of the neck, and first watch the effect 
on the blood before the death of the animal, and then observe 
the condition of the lymphatic glands after the death. I say, 
“after the death,” because I expected the animal to die within 


a short time of the operation, from interference with the 
absorption of fat. 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 457 


Experiment VILLI. 


A male dog of the English terrier type, weighing 5 kilogrammes 
70 decagrammes. Apart from being rather thin, the animal was per- 
fectly healthy. Must have been at least three years old. 

On the 2nd of February 1885, four hours before the time proposed for 
the operation, the dog was given as much milk as it would drink, in order 
that the thoracic duct might be full of chyle at the time of operation, 
and so, by its white colour, help one to find it. The animal having been 
put under ether, a V-shaped incision was made at the lower part of the 
neck, one of the limbs of the incision being parallel to and just above 
the clavicle, and the other parallel to the posterior border of the 
sterno-mastoid muscle. Then I reflected back the triangular flap of 
skin so marked out, divided the platysma, and dissected downwards 
with the finger, following the course of the external jugular vein, as 
far as its junction with the internal jugular. To give a little more 
room, some of the clavicular fibres of the sterno-mastoid muscle were 
divided. In this way one got deep down behind the clavicle; and 
after separating the structures as gently as possible, one noticed what 
appeared to be a white vessel passing towards the angle of junction 
between the jugular vein and the subclavian vein. By very gentle 
tracing this vessel was found to join the jugular at the angle of junc- 
tion with the subclavian vein. A chromic acid catgut ligature was 
then tied round it, about a quarter of an inch from the vein ; and it 
at once seemed to become somewhat distended behind the lgature. 
After a second thread of catgut had been laid round the vessel at a 
greater distance from the vein, in order that the vessel might, when 
necessary, be ligatured again, a pretty considerable opening was made 
in the vessel, and immediately a white fluid poured out so freely as to 
fill the wound. Then the other ligature was tied to stop the flow of 
the chyle ; but instead of being tied behind the wound in the vessel, 
it was accidentally tied over it, so that there was still an escape of the 
white fluid. A third catgut ligature was therefore put round the 
vessel, distinctly behind the wound in its walls, and about three- 
quarters of an inch from the junction with the vein; and this was 
firmly tied. That the vessel tied was actually the thoracic duct, the 
chyle which flowed out of it proved. 

The dog did not seem to suffer in the least from the operation. It 
was ordered lean meat that it might not be troubled with unabsorbed 
fat. It ate its food well, and had regular stools which did not show 
an unusual quantity of fat. Its weight varied slightly, but at no 
time showed any decided diminution. On the 4th of March, we, a 
month after the operation, its weight was 6 kilogrammes 10 deca- 
grammes, or 40 decagrammes more than on the day of operation. It 
was killed on the 11th of March, and then weighed 5 kilogrammes 75 
decagrammes. 

As in the previous tables, the corpuscles of the blood are enumerated 
in columns. In another column is the weight of the animal at different 
dates after the operation. 

VOL, XX. 2G 


458 DR J. LOCKHART GIBSON. 
Relation of 
Hemocytes. | Leucocytes. | Leucocytes to Weight. 
Hemocytes. 
Before Operation, . § | 8,726,900 14,000 1 :622°8 | 5 kilos. 70 decas. 


Operation on. 2 February. 
3 Feb., Ist day after,| 8,590,000 24,090 | 1:316°2 | 5 kilos. 61 decas. 


Be oi and 5 8,690,000 19,000 1:452'1 | 5 kilos. 45 decas. 
Dr ees 3rd » | 8,420,000 17,000 1:495°2 5kilos. 67 decas. 
Si 5 6th > | 7,830,000 14,000 1+ 523°5 | 5 kilos. 60 decas. 
10 ts, 8th 9 7,470,000 12,000 1:€22%5 | 5 kilos. 65 decas. 
LCS: 16th » | 7,890,000 10,000 1: 782 | 5 kilos. 35 decas. 
3 Fl ee 23rd =s,,_~—=s |_—s« 7,620,000 10,000 1: 762 | 5 kilos. 95 decas. 
4 March, 30th ein Ni 5,000,000 9,000 1: 841-1 | 6 kilos. 10 decas. 
ib eae 37th 5 8,630,000 12,000 1: 7194 | 3 kilos. 75 decas. 


The examination of the blood-forming organs yieldel the 
following :— 

The spleen was not remarkable either for size or for softness, 
If anything, it was rather larger and softer than usual. A 
scraping, treated as usual, showed very large numbers of 
“)lood-corpuscle-holding cells.” These cells, which are pretty 
constantly found in the spleen and bone-marrow, will be con- 
sidered later. Only a few nucleated red cells were found in 
each preparation. Still there were many more than in the spleen 
of a normal dog, where they either are not found at all or are 
found only after very diligent search (Bizzozero and others, and 
confirmed by myself). 

Bone-marrow.—The whole length of the shafts of the humerus 
and femur contained nearly fully reddened marrow, in which a 
few nucleated red cells were found, most of which were in the 
earlier stages, some, however, being in the more typical later 
stages. Numbers of “ blood-corpuscle-holding cells” were found 
in the marrow of all the bones examined. I found more of 
these cells in the spleen and bone-marrow of this dog than in 
any other dog. The marrow of the heads of the bones contained 
less fat than that in the shafts of the bones, and showed numbers 
of young nucleated red cells and of nucleated red cells intermedi- 
ate between the very young ones and the more typical, as well as 
a fair number of the really typical. The marrow of the ribs con- 
tained a fair number of nucleated red cells. There was quite as 
much fat as usual in the red marrow of all the bones, and the 
marrow in the shafts of the long bones other than the humerus 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 459 


and femur had its usual fatty appearance. To summarise the 
naked-eye and microscopic appearances of the marrow: those 
parts of the bones which always contain red marrow contained 
marrow richer than usual in nucleated red corpuscles, while in 
the humerus and femur the usually fatty marrow of the shafts 
had been replaced by red marrow. 

Lymphatic Glands.—In the left axilla there was an unusually 
large and succulent lymphatic gland, which was redder than 
usual; and in the right axilla a gland larger than usual, though 
not so large as that in the left axilla. A scraping from the cut 
surface of the gland in the left axilla, treated in the usual way 
with artificial serum and methyl-violet, showed some distinctly 
nucleated red cells of the later and more typical variety. There 
were also an unusual number of non-nucleated red corpuscles. 
In the gland of the right axilla, careful searching gave one or 
two nucleated red cells in each preparation. The most distinct 
appearances, however, were found in the glands of the mesentery. 
These glands were much larger than usual, very soft and succu- 
lent, and somewhat red on section. In scrapings from them 
were large numbers of nucleated red cells, of whose nature there 
could not be the slightest doubt. Most of these were of the 
later and more typical variety, and there were comparatively 
few belonging to the earlier stages. There were also large 
numbers of non-nucleated red corpuscles. No “ blood-corpuscle- 
holding cells” were found in any of the lymph glands examined. 
The inguinal glands were very decidedly enlarged. 

Thyroid gland of usual size. No evidence of blood-formation 
could be found in it. 

From this case, then, it would appear that the lymph glands 
even in normal conditions change a certain number of colourless 
into coloured corpuscles. This is supported not only by the 
microscopic appearances in the glands, where the nucleated red 
cells were almost as numerous as the colourless cells, and were 
more numerous than the nucleated red cells in the marrow of 
the ribs, but also by the enumeration of the blood-corpuscles 
during the life of the animal. For the table given shows that a 
very few days after the operation the red corpuscles showed a 
decided decrease, and that they remained very distinctly below 
their original number for fully a month after the operation. 


460 DR J. LOCKHART GIBSON. 


Soon after the fall in the number of red corpuscles, the colour- 
less corpuscles likewise suffered a diminution in their number, ~ 
the diminution being both absolute and relative, and remaining 
until the end of the observations on the animal. Between five 
and six weeks after the operation, the red corpuscles had prac- 
tically returned to their former number, the red-corpuscle-form- 
ing function of the lymphatic glands having been taken up by 
the marrow of the shafts of the humerus and femur. This, at 
least, is the only explanation I can find for the transformation 
of the originally fatty marrow of the shafts of these bones into 
red marrow. 


Very interesting was the condition of the thoracic duct. Its dissec- 
tion was very easy, as it was rather dilated. The contents, however, 
which must have been the cause of the dilatation, had disappeared, 
the duct being empty. The duct had a clear semitransparent pink 
appearance, which seemed to be due to the colour of its coats. I 
followed it upwards to its opening into the vein, dissecting very care- 
fully, to avoid destroying any branch. At three-quarters of an inch 
from the opening into the vein, the distinct tube became suddenly 
replaced by a white opaque cord. The junction of this apparently 
obliterated part with the rest of the duct exactly corresponded with 
the position of the most external of the ligatures ; and in this part of 
the duct one could with perfect distinctness feel three hard nodules, 
each about the size of a pin’s head, in the positions where the three 
ligatures had been placed. To make certain as to the condition of the 
duct, I opened it a short distance before the apparent obliteration ; 
and easily passed a fine thread of catgut along it as far as the begin- 
ning of the constriction, where it was stopped. And I then tried a 
horse-hair, but it also failed to pass into the obliterated part of the 
duct. There can, therefore, be no doubt that the thoracic duct had 
been tied, and that the result of the tying had been complete oblitera- 
tion. As to the possible existence of an important branch of the duct, 
with a corresponding persistence of the circulation of the chyle, it may 
be stated that I searched very carefully along the whole length of the 
duct, but could find no branch passing off from it, nor any second 
duct passing upwards from the receptaculum chyli. I have been 
unable to find a good description of the lymphatic system in dogs ; 
and from one case, and without careful injection of the thoracic duct, 
it would, of course, be absurd positively to conclude that the chyle 
had been altogether prevented from reaching the blood by a collateral 
channel: still, the facts are in favour of such a conclusion. 


Apart from what was found in the lymph glands, the case is 
extremely interesting from the number of possibilities it suggests. 
If the passage of the chyle into the blood by the thoracie duct 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 461 


be prevented, is there any other way by which it can enter the 
circulation? And if there is no other way, how was it that this 
dog did not suffer from fat starvation? In answer to this last 
question, it may be said that dogs can obtain fats from proteids 
more easily than a human being can, and that in them the con- 
version of proteids into fats can take place to such an extent as 
to render the absorption of fat from the intestine superfluous. 
Amyloids cannot in the case of this dog be supposed to have 
been a sufficient source of fat, as the animal was fed almost en- 
tirely on lean meat. The meat was, however, of course, not 
carefully deprived of its fat. 

In this connection, it is important to mention an interesting 
paper of William Turner’s! on obstruction of the thoracic duct 
in the human subject. He in two cases found the thoracic duct 
entirely obliterated by a thoracic aneurism without emaciation 
having resulted. He thinks that in all cases where obstruc- 
tion of the duct comes on gradually collateral channels are 
established, and that there is no sufficient proof of the state- 
ment that obstruction of the duct by an intrathoracic growth 
leads to emaciation. He also mentions cases where collateral 
branches have been found, as well as communications between 
the lymphatics and veins elsewhere than at the root of the 
neck. Turner’s idea of the gradual establishment of collateral 
circulation will not apply in my case, because there the oblitera- 
tion of the duct was sudden. There may, however, previous to 
the operation, have been unusual communications between the 
abdominal lymphatics and the venous system. Anyhow, the 
obliteration of the duct must have caused some obstruction to 
the flow of lymph, or I should not have found so many develop- 
ing red cells in the lymph glands. 

To return to the light thrown by this case on the function of 
the lymphatic glands: it seems to me that there is no link 
wanting in the chain of evidence in favour of the lymphatic 
glands having and exerting in normal conditions the power 
of producing red blood-corpuscles. (1) The enumeration of the 
blood-corpuscles during life showed, for at least a month after 
the operation, a distinct diminution in the number of red 
corpuscles, while a diminution in the number of white cor- 


1 Turner, Ed. Med. Journal, 1859. 


462 DR J. LOCKHART GIBSON. 


puscles occurred only after the fall in the number of red. 


(2) Nucleated red corpuscles were found in great numbers in _ 


the abdominal lymphatic glands. (3) It was found that the 
red blood-forming marrow had extended into the shafts of the 
humerus and femur. 

The nucleated red cells found in the lymphatic glands of the 
spleenless animals point to the same conclusion. And some 
other recorded observations, when taken together with this case, 
put, in my opinion, this function of the lymphatic glands entirely 
beyond the region of doubt. 


The first of these observations is the case recorded by Neumann, 
already alluded to. It will be remembered that he had recourse to 
the hypothesis that owing to some disease of the vascular walls the 
red corpuscles had found their way into the terminal lymphatics and 
lymph sinuses. It seems to me much more likely that he found 
nucleated and non-nucleated red cells in the lymphatic glands because 
they were formed there. 

Secondly, Weigert! found at the post-mortem of a case of pernicious 
anemia that the lymph glands were large, soft, and red; that their 
lymph sinuses were filled with lymph uncommonly rich in red blood- 
corpuscles; and that the large lymph vessels had similar contents. He 
gives as the explanation of this that first struck him the possibility of 
there having been some abnormal condition of the vascular walls, 
which had allowed the red corpuscles to pass out more easily than in 
normal conditions. But he remarks that, on reading Rindfleisch’s 
paper,” he began to think that it might have been a case of blood-for- 
mation in the lymphatic glands, and regrets that he had not examined 
the glands in the fresh condition with special reference to the presence 
of nucleated red cells, adding that on hardened preparations he had 
failed to satisfy himself. 

The paper of Rindfleisch quoted by Weigert contains the account 
of a case where there was general sclerosis of the bones, as the result 
of rachitis, and where the marrow cavities were practically obliterated. 
Rindfleisch found the spleen enlarged and soft, and the lymph glands 
enlarged and resembling the spleen in consistence and appearance. 
In the lymph glands he found numerous ordinary red cells and 
numerous nucleated ones. He looked on this as a case of vicarious 
function of the lymphatic glands. 


Having thus, on the subject of blood-formation, given all my 
own experiments except those performed in connection with the 


' Weigert, ‘‘ Perniciése Animie mit ausgedehnter Lymphangiectasie. Erfiil- 
lung der Lymphbahnen mit blutéhnlicher Lymphe,” Virchow’s Archiv, Bd. 1xxix. 
p- 390. 

* Rindfleisch, Archiv f. mikrosk. Anat., Bd. xvii. (1879). 


. 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 463 


thyroid gland, and also given what seem to me the more im- 
portant experiments and opinions of other writers, I shall now 
proceed to review the facts on which my own conclusions are 
_based. The conclusions themselves are more or less embodied 
in what has already been said, but a more categorical statement 
will not be amiss. 


In this paper it has already been assumed that the nucleated red 
cells found in the bone-marrow and elsewhere are the forerunners of 
the non-nucleated red blood-corpuscles, and it will perhaps seem 
curious I should only now begin to give my reasons for holding them 
to be so. In explanation, I would point out, on the one hand, that 
the function of the blood-forming organs could not have been treated 
without the assumption, and, on the other hand, that the life-history 
of the red cells could not have been thoroughly treated without a pre- 
vious consideration of the blood-forming organs. Under these circum- 
stances, I have chosen simply to indicate at the beginning of the 
paper what the nucleated red cells are, and then, for the time being, 
assume that they develop into non-nucleated red cells. 

It will here be convenient first to dismiss the hematoblasts of Hayem. 
In the first part of this paper I have pretty clearly indicated my 
own opinions as to these elements; as also the opinions of Neumann 
and Bizzozero. It seems to me that Bizzozero finishes the discussion 
very well, when he expresses the hope that Hayem may “one day 
see the nucleated red cells in the bone-marrow ;” for he thinks that 
if he does so, and compares them with the nucleated red cells of the 
embryo, his allegiance to his hematoblasts will vanish. And 
Neumann remarks on Pouchet’s explanation of the nucleated red 
cells found in the bone-marrow (for Pouchet does not deny that they 
exist), that if Pouchet had compared them with the nucleated red 
cells of the blood of the embryo “ he would not have fallen into the 
unfortunate fabrication, ‘ déyénérescence hémoglobinique,’ which leads to 
the absurd conclusion that the nucleated red cells in the blood of the 
embryo are nothing else than hemoglobin-degenerated cells.” 

Neumann, in his paper in the Zeitschrift fiir klin. Medicin, already 
mentioned, gives a very good criticism of all the views as to the 
origin of the non-nucleated red blood-corpuscles, and refuses to accept 
Schiafer’s and Ranvier’s view that they may arise endogenously as 
non-nucleated bodies in the. protoplasm of cells, for instance in the 
subcutaneous tissue of the embryo. And he also refuses to accept 
the intermediate position taken up by Rollet, viz., that they arise in 
two ways, either as nucleated daughter-cells or as non-nucleated 
offspring of pre-existing cells. Rollet thinks with Ranvier that the 
former mode of development occurs only in the embryo. 

Merkel, in Virchow’s Jahresbericht, says of this paper of Neumann’s 
that its “‘ very well deserved destructive criticism of the views of 
Hayem, Pouchet, Schafer, and Ranvier,” is worthy of very special 
attention. 


464 DR J. LOCKHART GIBSON. 


Neumann considers that there is at all periods of life only one way 


in which non-nucleated red corpuscles originate, viz., as nucleated red _ 


cells, like those of the embryo. Rindfleisch, too, in his paper already 
quoted, says that ‘“‘we must entirely banish the idea of their arising 
in any other way than as nucleated red cells, throughout life as well 
as in the embryo.” And Bizzozero is equally certain on the point. 
In fact, in opposition to Hayem and Pouchet, the opinion that the 
nucleated red cells in the bone-marrow and other blood-forming organs 
are the only forerunners of the non-nucleated red corpuscles is almost 
universal, And it will be evident that my own observations, so far 
as they go, support it very strongly. 

While agreeing that the nucleated red cells are the forerunners of 
the non-nucleated ones, the different observers on the blood disagree 
both as to the details of the transition and as to the origin of the 
nucleated red cells themselves. 

To consider first what becomes of the nucleus :— 

Kélliker and Neumann say that it disappears from the cell by 
breaking down, Neumann evidently thinking that in breaking down 
it becomes absorbed in the general cell substance. K@lliker’s observa- 
tions were made on the nucleated red cells of the embryo, and 
Neumann’s on the nucleated red cells found in the embryonal liver, 
and in the marrow throughout the whole of life. Neumann gives 
drawings showing the last remains of the nucleus as a small body of 
the size of a colourless microcyte, and Kélliker says that it breaks up 
into two, three, or four fragments. 

Rindfleisch, in opposition to Neumann and Kolliker, says that the 
nucleus, surrounded by a small amount of undifferentiated protoplasm, 
passes bodily out of the cell. The further history of the nucleus 
‘“may,” says Rindfleisch, ‘‘be a matter for discussion.” 

Bizzozero, without giving any distinct opinion, is rather in favour of 
Rindfleisch’s view. 

From what I have myself seen, I should prefer to support Neumann’s 
view. For I have seen in the bone-marrow nucleated red cells in which 
there was apparently only a small remnant of the nucleus, and have, 
like Neumann, entirely failed to find a ring of undifferentiated proto- 
plasm surrounding the nucleus of a typical nucleated red cell. 

To the view of Obrastzow! and Arndt,? viz., that the nucleus in 
the nucleated red cells has not the character of a real cell nucleus, 
but is a post-mortem appearance, analogous to the precipitation of 
fibrin, Neumann returns the best answer, when he calls attention to 
the researches of Fleming on the indirect division of cells. Of the 
different stages of such indirect division (‘“ Kariokinesis”) in the 
nucleus of a nucleated red cell, elaborate drawings are given by 
Lavdowsky * and Bizzozero.* 

The question of the origin of the nucleated red cells is still more 


Obrastzow, Centralbl. f.d. Med. Wiss., 1880, No. 24. 
Arndt, Virchow’s Archiv, Bd. lxxxiii. p. 18. 
Lavdowsky, Virchow’s Archiv, Bd. xevi. (1884). 

4 Bizzozero, Virchow’s Archiv, Bd. xcv. (1884). 


1 
2 
3 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 465 


undecided. Neumann, from observations on the bone-marrow, thought 
that they arose from the leucocytes; but his faith in that idea has 
evidently been shaken by his observations on the embryonal liver,’ 
where he finds that the red cells arise from nuclei which appear in 
the protoplasm of large cells like the proper cells of the liver; that 
these nuclei first come to possess a very thin ring of hemoglobin- 
coloured perinuclear substance ; that this thin ring gradually increases 
in breadth until typical nucleated red cells are produced ; and that 
these nucleated red cells then pass out of the mother-cell, to multiply 
by division in the liver and in the circulating blood. The “ mother- 
cells” were first discovered by Gerlach,? and were called by him 
* blood-corpuscle-holding cells,” because he thought they contained 
young red blood-corpuscles. Gerlach, however, changed his opinion of 
them when Kélliker* and Ecker® pointed out that they did not con- 
tain young red blood-corpuscles, but were probably cells which had 
taken up extravasated blood-corpuscles. 

In spite of the very strong testimony of such an authority on the 
blood as Neumann, I think that the view taken by Kolliker and Ecker, 
or a modification of it, will turn out to be the right one ; and that even 
the nucleated red cells found by Neumann in the “ blood-corpuscle- 
holding cells ” had been taken up from the tissues. I shall return to 
these “blood-corpuscle-holding cells” when I consider similar cells 
found in the spleen and bone-marrow. In his last work on the blood, 
Neumann says : “ As to the origin of the early stages of the nucleated 
red blood-corpuscles, which, as is known, is still very little under- 
stood, I need here say no more.” Not only Neumann, but also 
Rindfleisch, Bizzozero, and many others, affirm that the nucleated red 
cells increase in number by indirect division ; and Bizzozero, in his 
last paper,® states that they increase only by indirect division, and 
that all the other views which have been advanced as to their origin 
(for example from the leucocytes) are mere hypotheses. In making 
this statement, however, which he applies to all classes of vertebrates, 
Bizzozero seems to forget that he very strongly incriminates himself. 
For in his paper on the development of blood-corpuscles in birds,’ he 
not only speaks of the nucleated red cells as arising from the colour- 
less cells of the marrow, but gives diagrams of the different steps of 
development, and says that the colourless cells pass out of the marrow 
pulp into the veins of the marrow before assuming hemoglobin. In 
his last paper, he urges that the nucleated red cells cannot arise from 
the white cells, because they are never found in the lymph glands or 
lymph follicles, where white corpuscles are most numerous. This objec- 
tion, however, as my experiments have already shown, may be got over. 


1 Nenmann, Archiv der Heilkunde, Bd. xv. 

2 Gerlach, Zeitschr. fiir rat. Med., vii. (1849), p. 79. 

3 Gerlach, Handbuch der Gewebelehre, 2. Aufl. (1860), p. 56, 58. 
4 K6lliker, Zeitschr. fiir wiss. Zool., ii. (1850), p. 117. 
5 Keker, Zeitschr. fiir wiss. Zool., ii. (1850), p. 276. 

6 Bizzozero, Virchow’s Archiv, Bd. xev. (1884). 

7 Bizzozero, Moleschott’s Untersuchungen, Bd. xii. 


466 DR J. LOCKHART GIBSON. 


As will have been seen from my descriptions of the appearances in 
the bone-marrow and other blood-forming organs of the animals on 
which I experimented, I have come to definite views as to the origin 
and development of the nucleated red cells. And it will also have 
been noticed that my conclusions agree with those of Kolliker and 
Fahrner?! and others, who hold that the nucleated red cells are derived 
from the colourless corpuscles. 

My description of the origin of the nucleated coloured cells in the 
marrow corresponds almost exactly with that given hy Malassez ;? but 
as to their further history differs very much from that of Malassez, and 
agrees with that given by Neumann and Bizzozero. 

Malassez says :—‘‘ They arise in the bone-marrow from cells which 
have a large nucleus which nearly fills the whole cell. The next 
stage is shown by the presence of a trace of hemoglobin in the cell, 
the cell still having a large nucleus, which is granular, and is not so 
strongly coloured by staining agents as the nucleus of the earlier stages 
of the cells. The protoplasm becomes more and more hyaline, the 
heemoglobin-stained contents at the same time increasing in quantity, 
and the nucleus becoming still less easily affected by staining agents.” 
The last stage in this process he calls that of “cellules hémoglo- 
biques.” 

Up to this point, z.e., to the production of the ‘“ cellules hémo- 
globiques,” I can quite agree with Malassez’s description; but here 
my description diverges very much from it, and agrees with those of 
Neumann, Bizzozero, and others, 

According to Malassez, buds appear on the side of his hzmoglobin- 
coloured cell, in some animals (young goat) many buds appearing, in 
other animals (rabbit, calf, cat, and ox) only one bud being evident ; 
and these buds, according to him, enlarge and become non-nucleated 
red blood-corpuscles, the cell remaining and enlarging to produce more 
buds. 

Bizzozero’s explanation of the presence of these buds is, that as he 
could never see them in preparations treated with neutral fluids, there- 
fore they must have been produced by the action of the osmic acid 
with which Malassez prepared his specimens. And this explanation 
receives great support when we remember that by means of reagents, 
such as tannic acid, buds can be produced on the ordinary red blood- 
corpuscles. 

I do not at all deny that one of the methods by which the nucleated 
red cells increase in number is indirect division. That they can and 
do increase in that manner, has been put beyond all doubt by the 
researches of Bizzozero, Neumann, and others. I do, however, say 
that there is also a fresh production of the nucleated red cells 
throughout the whole of life ; and that in the blood-forming organs of 
the animals observed by me there were not a sufficient number of 
dividing nucleated red cells present to account for the numerous 


nucleated red cells found, while there were always numerous examples — 


1 Kolliker, Zeitschr. fiir rat. Med., Bd. iv. (1845), p. 112. 
2 Malassez, Gaz. méd. de Paris, 1881, No. 49. 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 467 


of the intermediate stages between the colourless marrow cells and 
nucleated red cells. 


To describe the transition stages more particularly :— 

The condition of one of the colourless marrow cells before 
beginning to assume hemoglobin is that of a cell of from 10 to 
12, or sometimes even 14, micros. in diameter, with a relatively 
very large nucleus, almost filling the cell, and finely-granular 
perinuclear protoplasm (fig. 1, a). The first appearance of 
hemoglobin in this cell is in the form of a thin ring at the 
periphery of the cell; and between this ring and the nucleus 
a small amount of the finely-granular perinuclear protoplasm 
can often still be observed. More frequently, however, the 
band of hemoglobin-tinted substance seems to fill the whole 
space between the cell-envelope and the nucleus, and the 
perinuclear substance has the same clear appearance as the 
substance of the non-nucleated red corpuscles (fig. 1, 0). As 
this hzmoglobin-tinted perinuclear substance increases in 
amount, the nucleus appears to become smaller and to retreat 
towards the centre of the cell, the cell as a whole at the same 
time becoming smaller (fig. 1, ¢). 

I have, in my preparations, been able to see all stages of 
transition, between the comparatively large young nucleated red 
cells and the much smaller typical nucleated red cell, measuring 
about 9 micros. in diameter, where the nucleus, too, is much 
smaller, and the perinuclear coloured substance is much broader 
(fig. 1, d). My conception has been, that previous to the assump- 
tion ef hemoglobin the nucleus of the colourless marrow cells en- 
larges and comes to occupy almost the entire cell, and that under 
the influence of the nucleus the peripheral part of the cell 
assumes hemoglobin, the nucleus at the same time becoming 
smaller. That the nucleus enlarges previous to the assumption 
of hemoglobin, I infer from the fact that there are in the bone- 
marrow and other blood-forming organs many large colourless 
cells which contain nuclei not so very large as compared with 
the cells and yet in other respects resemble the cells of fig. 1, a, 
supposed by me to be marrow cells in the condition immediately 
preceding the assumption of hemoglobin. These large colour- 
less cells, with nuclei not so disproportionately large, are the 


468 DR J. LOCKHART GIBSON. 


cells in the blood-forming organs that seem to me to divide most 
actively. In particular, they seem to divide much more actively 
than the nucleated red cells, of whose division, however, I have 
seen indubitable examples. 

It would accordingly appear that the nucleus takes an active 
part in enabling the cell to assume hemoglobin, and then, as 
being of no further use, disappéars from the cell; and my belief 
in this function of the nucleus is one of my reasons for thinking 
it unlikely that the non-nucleated red corpuscles, while circulat- 
ing in the blood, possess different amounts of hemoglobin. For 
I cannot think that without the presence of a nucleus the red 
corpuscles have it in their power to add to the hemoglobin they 
contain. Perhaps, however, it would be premature to deny that 
the amount of hemoglobin assumed under the influence of the 
nuclei may in some diseases be abnormally small. 

I could not obtain evidence of small nucleated red cells 
increasing in size: the small nucleated red cells which were seen 
in my preparations could, I thought, always be accounted for by 
the effect of the diluting fluid. The young nucleated red cells 
show a still greater tendency to creuate and contract into 
small globular bodies than the non-nucleated red céllsdo. Those 
nucleated red cells which were dividing were always large 
enough to produce two daughter-cells of nearly the same size 
as the ordinary red cells. The developing red cells in the 
lymph glands were in the younger stages not so large as those 
found in the marrow, but in other respects tle development 
seemed to be the same. In the spleen, the development of red 
cells is the same as it is in the bone-marrow. 

Those white cells which are transformed into nucleated red 
cells in the bone-marrow are not necessarily all produced there: 
it may be that many of them are produced in the lymphatic 
glands, and carried to the bone-marrow, there to enlarge and 
pass through the variotis stages of the transformation. 

The white blood-corpuscles, whether produced in the lymph- 
glands, spleen, or bone-marrow, seem to me to be the fore- 
runners of the nucleated red blood-corpuscles :— 

(1) Because those cells in the bone-marrow which belong to 
the stage in the development of the nucleated red cells previous 
to the appearance of hemoglobin are exactly similar to enlarged 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 469 


white corpuscles, while the bone-marrow contains besides such 
cells also many white corpuscles of ordinary size. (2) Because in 
the lymphatic glands nucleated red cells were found which had 
evidently been developed there, and also white cells with pecu- 
liarly clear perinuclear substance, which looked as if they needed 
nothing but hemoglobin to convert them into nucleated red 
cells. (3) Because when the lymph is prevented from entering 
the blood the number of red corpuscles in the blood falls. (4) 
Because after excision of the spleen, which is an organ for 
forming white blood corpuscles, the white corpuscles in the 
blood on the one hand increase in number while the red cor- 
puscles decrease, and on the other hand may sink even below 
their original number as the red corpuscles rise again to theirs. 
(5) Because a leucocyte is typically an embryonic cell capable of 
further differentiation. 

I entirely agree with Neumann, Rindfleisch, and Bizzozero, in 
thinking that the only forerunners of the non-nucleated red 
corpuscles, during the whole of extra-uterine life as well as 
during intra-uterine life, are the nucleated red cells found in 
the blood-forming organs. Now that it can be shown that these 
cells exist in the blood-forming organs throughout the whole of 
life, there is really no necessity for having recourse to micro- 
cytes, and endogenous cell-formation, in order to explain the 
blood-forming process. For why should the origin of the blood- 
corpuscles in extra-uterine life be different from their origin in 
intra-uterine life 2 And, on the other hand, it has been abund- 
antly shown that the nucleated red cells increase in number 
in the blood-forming organs in direct proportion to the neces- 
sity for the formation of blood, and accordingly increase most 
when the blood is being most actively regenerated. So that we 
cannot have any stronger proof that they are young red blood- 
corpuscles. 

These cells, then, being accepted as young red blood-corpuscles, 
the question arises, “ Which is the organ most active in their 
formation in extra-uterine life?” In answering it, I must, as 
the result of my own experiments, agree with Neumann and 
Bizzozero that the bone-marrow is the most active producer of 
red blood-corpuscles during extra-uterine life: that is, in the 
generality of animals; Bizzozero having found that in tailed 


470 DR J. LOCKHART GIBSON. 


amphibians and in fishes the chief blood-forming organ is the 
spleen. i 

From personal experience, I can, of course, speak only of the 
blood-forming organs of dogs. In these animals, the red 
marrow appeared to be the chief organ; though the spleen, too, 
at any rate within the first year of extra-uterine life, produces 
red blood-corpuscles, and does so even under normal conditions. 
I should say that the spleen probably throughout life forms a 
certain number of red blood-corpuscles. 

Experiment V., where the dog was at least two years old, 
gave the strongest possible evidence in favour of Bizzozero’s 
view that the spleen actively produces red corpuscles when the 
blood-forming capabilities of an animal are called into play by 
loss of blood. 

As to the function of the lymphatic glands in the formation 
of red blood-corpuscles, Experiment VIII. proved—as it seems 
to me, conclusively—that the lymphatic glands do, during the 
whole of extra-uterine life, take part in the formation of the red 
blood-corpuscles. But, from the cases of excision of the spleen, 
it appears that their activity in this respect is at any rate less 
than that of the bone-marrow; and I think it very probable 
that it is also less than that of the spleen. 


As to how the young red blood-corpuscles formed in the bone-marrow 
pass into the circulation, there are two opinions—that of Rindfleisch, 
which.is adopted by Neumann, and that of Bizzozero, Rindfleisch 
supports the view which Hoyer! and Riidinger? took in 1869, viz., 
that the greater part of the capillary system in the bone-marrow pos- 
sesses no special wall, but that the thin-walled arteries open into 
tunnels in the marrow-substance (i.e, into the interstices of the 
marrow-pulp), and that the veins take origin from these tunnels. 
He argues that it would be quite superfluous for the capillaries of 
the marrow to possess walls, because the circulation in the marrow 
must be a restricted one, from the hard bony surroundings making 
the inflow and outflow of blood always correspond. Bizzozero,® 
on the other hand, considers that the capillaries of the marrow 
are real capillaries, and have distinct thin walls, and that the 
veins arise as very wide channels, also with very thin walls. He 
further considers, or rather considered,* that the colourless marrow- 


1 Hoyer, Centralbl. f. d. med. Wiss., 1869. 
2 Riidinger, Centralbl. f. d. med. Wiss., 1869. 
3 Bizzozero, Morgagni, 1869. 
+ Bizzozero, Moleschott’s Untersuchungen, Bd. xii. 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION, 471 


cells, before assuming hemoglobin, pass through the thin walls of the 
veins into their lumen, there to assume hemoglobin ; and that they 
never contain hemoglobin until they enter the veins, no hemoglobin- 
containing cells being visible outside the vessels. As, however, in 
the same paper, he talks of its being a doubtful question whether 
the blood-channels in the spleen have or have not distinct walls, we 
may take his view of the condition of the circulation in the bone- 
marrow with some reserve. The question is, of course, a very difficult 
one to settle ; for it is difficult to get the marrow so hardened as to 
allow of satisfactory sections being made. I think it, however, more 
likely that Rindfleisch’s opinion will turn out the correct one. 


A word must be said about the so-called “ blood-corpuscle- 
holding cells ” :— 


As has already been mentioned, these cells were discovered, in the 
liver of the embryo, by Gerlach, and were at first thought by him to 
contain developing red blood-corpuscles. Gerlach, however, afterwards 
gave up this view, and accepted that of Kélliker and Ecker, who con- 
sidered them to be cells which had taken up extravasated blood-cor- 
puscles. Neumann rediscovered these cells, and recognised them to 
be the same as those described by Gerlach, but considered Gerlach’s 
original opinion about them to be correct. Gerlach’s original opinion 
and description were as follows:1— He said he found “coloured 
blood-corpuscles in large numbers lying as cell-contents inside large 
colourless cells ;” and ‘said, pds that ‘these enclosed bubble- like, 
reddish-yellow nuclei (viz., the ‘coloured blood-corpuscles’), become 
free, and, perhaps by the formation of an envelope (Hiille) and the shar- 
ing of the colouring matter with this envelope, become transformed into 
coloured blood-corpuscles.” This description of Gerlach’s exactly 
applies to ‘‘ bluod-corpuscle-holding cells” which I have found in the 
spleen and bone-marrow, and supports the view taken by KGlliker and 
Ecker. Neumann objects to this view chiefly because he found 
nucleated red corpuscles in these large mother-cells and found, as he 
thought, stages of transition between “free nuclei” contained in the 
cells (but differing from the proper nucleus of the cell) and the typical 
nucleated red corpuscles. To turn Neumann’s explanation of Hayem’s 
hematoblasts against himself, one may ask, Is it not possible that in 
embryonal life some of the nucleated red corpuscles are, as easily as the 
non-nucleated ones, extravasated and taken up by the large cells, and 
that the transition-stages in the “‘ development ” of the nucleated red 
corpuscles observed by Neumann in “mother-cells” are simply stages 
of breaking-down of such corpuscles in cells which had taken them up? 


I have no personal knowledge of the “ blood-corpuscle-holding 
cells” in the embryonal liver, but can speak of similar cells in the 
spleen and bone-marrow—cells whose function seems to be to 
take up breaking-down red blood-corpuscles, complete their break- 

1 Gerlach, Zeitschrift fiir rationelle Medicin, vii. (1849), p. 79. 


472 DR J. LOCKHART GIBSON. 


ing-down, and perhaps in some way prepare their hemoglobin 
for being again made use of. In the accompanying drawings 
my reasons for supposing this function will be seen (fig. 2). 
The cells vary from about 8 or 10 micros. to perhaps 40 micros. 
in diameter. The nucleus is, as a rule, pushed to one side, and 
often flattened, like the nucleus of a fat cell. In some cells, 
however, I was, like Neumann, unable to distinguish a nucleus. 

The peculiar contents of these cells are well described in 
Gerlach’s words, as “ bubble-like reddish-yellow nuclei” ; except 
that they cannot be called “ nuclei,” having, in fact, none of the 
characters of nuclei. They are perfectly clear, show no granula- 
tion, and are darkly coloured with hemoglobin. Of course, 
Gerlach’s after-admission that they are breaking-down coloured 
corpuscles shows that he gave up the idea of their being nuclei. 

These “ blood-corpuscle-holding cells” contain, generally ac- 
cording to their size, one or several “ bubble-like reddish-yellow ” 
bodies, which vary very markedly in size, and vary in colour 
directly with their size. In some of the cells there can be seen, in 
the centre, a round or irregular coloured body, often many times 
larger than a normal red corpuscle; which at first may appear 
single, but when carefully focussed is seen to be composed of a 
number of smaller bodies closely packed together (fig. 2, 6). 
As a rule, around this darkly-coloured mass there are many 
small straw-coloured fragments, which have exactly the same 
appearance as the fragments of a broken-down red corpuscle. 
The general finely granular protoplasm of the cell has, as a rule, 
a faint hemoglobin tint. Different examples of these cells are 
shown in fig. 2. 

It would appear that under the influence of these cells the 
hemoglobin-substance of the broken-down red corpuscles is 
fused into a common mass; but whether their function is to 
prepare hemoglobin to be again made use of as hemoglobin 
or to prepare it for being carried to the liver to be there made 
use of, it would be difficult to say. 

Before giving a summary of my conclusions, I should like 
again to draw attention to the excellent advice of Neumann and 
Bizzozero, who urge that the substance of the blood-forming 
organs should, like the blood itself, always be examined fresh, 
and either undiluted or diluted with newtral fluids. 


THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 473 


The conclusions are :— 

1. Nucleated red cells, derived from white corpuscles and 
colourless marrow cells, are the only forerunners of the non- 
nucleated red blood-corpuscle, throughout the whole of life. 

2. The transformation of the colourless cells into nucleated red 
cells takes place in the bone-marrow,spleen,and lymphatic glands, 

3. The colourless cells and the nucleated red cells multiply 
in the blood-forming organs by division. 

4. The red bone-marrow plays the most important part in the 
production of red blood-corpuscles during extra-uterine life. 

5. After the production of anzemia, some of the fatty marrow 
becomes red marrow, and joins in the formation of red blood- 
corpuscles. 

6. The blood-forming action of the spleen is in extra-uterine life 
a subordinate one, but when the reserve blood-forming capabili- 
ties of an animal are called on its activity is greatly increased. 

7. After excision of the spleen, a portion of the formerly fatty 
marrow becomes red marrow, and the lymphatic glands increase 
their activity as regards the production of red blood-corpuscles. 

8. After excision of the spleen, the red corpuscles in the blood 
decrease in number, and there is a consequent increase in the 
number of white corpuscles. The red corpuscles return to their 
normal number within six months, after which there may be 
a decrease in the number of white corpuscles. 

9. The chief function of the lymphatic glands is the pro- 
duction of white corpuscles, but they also, even in normal 
conditions, produce a certain number of red corpuscles. Their 
activity in the latter respect increases with the necessity for the 
production of red corpuscles. 

10. The spleen and bone-marrow, and possibly also the lym- 
phatic glands, contain cells whose function appears to be to 
break down red blood-corpuscles. 


1 With reference to Dr Gibson’s thesis, Dr Creighton has called the Editors’ 
attention to a paper by himself (Dr C.), ‘‘ Illustrations of the Pathology of Sar- 
coma,” published in 1880, in the April number of this Jowrnal, claiming that it 
contains an important contribution to the study of blood-formation, and that he 
has developed his hematoblastic doctrine further in the article ‘‘ Pathology” in 
the Encyclopedia Britannica. As Dr Gibson sailed for Queensland some months 
ago, it has not been possible to bring Dr Creighton’s papers under his notice, but 
the Editors take this opportunity of referring to them. 

VOL, XX. 2H 


474 THE BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 


EXPLANATION OF PLATE XIV. 


Fig. 1. a, Colourless marrow-cells, that is, marrow-cells before the 
appearance of hemoglobin in them. 0, First appearance of hemo- 
globin in marrow-cells. c, A later stage of the marrow-cells. d, Typi- 
cal nucleated red cell. e, Probably shows the remains of a nucleus in 
a red cell. #, Ordinary non-nucleated red corpuscles, to serve as a 
scale. 

Fig. 2. a, Small “blood-corpuscle-holding cell,” containing one 
pretty large “ bubble-like reddish-yellow” body. 0, Large blood-cor- 
puscle-holding cell, containing many broken-down and partially fused 
red corpuscles (bubble-like reddish-yellow bodies). c, The same as 3, 
but with fragments more completely fused together in centre. Many 
small isolated fragments of red corpuscles in the periphery of the cell 
d, Shows small nucleus (unstained), pushed into a corner. Various 
bubble-like bodies: some very faintly hemoglobin-tinted, one very 
deeply tinted. e, Large “ blood-corpuscle-holding cell,” with nucleus 
(unstained) pushed to one side and flattened. Many bubble-like 
bodies. The larger the bodies, the more darkly are they tinted with 
ne eee ont. f and g, Other examples of “ blood-corpuscle-holding 
cells,” 


(Zo be completed in the next number.) 


INVESTIGATIONS IN THE RELATION BETWEEN CON- 
VERGENCE AND ACCOMMODATION OF THE 
EYES. By Ernest E. Mappox, M.B. Edin, Syme 
Surgical Fellow in the University of Edinburgh? 


I. Introductory Sketch. 


Wuy, if we see separately with each eye, do we not see double 
when both are used? This problem has taxed the ingenuity 
of many busy minds in past ages, and its history is by no 
means one of uniform progress. 

Euclid, two or three centuries B.c., had advanced so far beyond 
some at a far later date as to recognise that both eyes were employed 
in unison, and that their dissimilar pictures were in some way united. 
Galen surmised that the union of the optic nerves at the commissure 
supplied a clue. Both he and Herophilus assumed that the two 
nerves were there united by mysterious pores; doubtless to permit 
the free passage and intercourse of the little spirits of both sides, 
whose remarkable unanimity in fitting the pictures together was 
evidenced by single vision. Later on Gassendus, Tacquet, and Joan 
Baptista Porta, the inventor of the camera obscura, escaped the 
difficulty altogether by assuming that one eye only at a time was 
engaged in vision. 

In 1613, Francis Aguillon (Aguilonius), a learned Jesuit, called in 
the aid of what he termed a “ common sense,” which “imparts its aid 
equally to each eye, exerting its own power equally in the same 
manner as the eyes are converged by means of their optical 
axes.” This was an advance, for the two pictures, we may truly say, 
are mentally united by a “common sense,”? of the real nature of 
which we probably know little more than Aguilonius, though we 
may notice more of its effects. 

Dr Briggs appears to have been the first to have suggested “ cor- 
responding ” or “identical” points in the two retine, that is, that each 
point on the inner side of one retina has a corresponding point on the 
outer side of the other, so that when images are thrown by an object 
upon these identical points, they are mentally united. This was a 
great advance, though the theory of “identical points in the field of 
vision” is now considered more correct. But he explained it in a 


1 The original of this memoir was the successful essay submitted in com- 
petition for the Syme Surgical Fellowship in April 1884. Before publication it 
has been revised and enlarged. 

? It is now located in a theoretical ‘‘ fusion centre.” 


476 MR ERNEST E. MADDOX. 


curious way, by ascribing to each jibre of the optic nerve a different 
degree of tension, like the strings of a violin or piano, each vibrating 
in unison with its own retinal area,—‘“‘a tension,” argued Porterfield, 
‘‘impossible in the soft and pulpy structure of the nerve fibres.” 

From the fact that ‘in animals which look the same way with 
both eyes, the optic nerves meet before they enter the brain, while 
this union does not occur in those which do not, such as fishes and 
the chameleon,” Sir Isaac Newton suggested au arrangement of the 
optic fibres at the commissure, which exactly tallies with that now 
generally received—“ the fibres on the right side of both (optick) 
nerves uniting there at the commissure, and, after union, going 
thence into the brain in the nerve which is on the right side of the 
head, and the fibres on the left side of both nerves uniting in the 
same place, and, after union, going into the brain in the nerve which 
is on the left side of the head.” I quote from the 13th Query 
at the end of his “ Treatise on Opticks” (1718), the more remarkable 
because it was the belief of anatomists, like Vesalius, that no 
decussation occurred at the commissure, and that it consisted of 
fibrous tissue. 

Dr William Porterfield of Edinburgh is believed to have first 
enunciated the correct, though still very partial theory of binocular 
vision. In his “Treatise on the Eye” (1759)! he showed that when 
the eyes are accommodated for any object their two visual axes are 
also exactly converged upon the same point, and “since each eye 
possesses the power, either intuitively or by acquisition, of localising 
points in space, the object must appear single, it being impossible 
for us to conceive two objects existing in the same place at the same 
time. 


Single binocular vision therefore requires a perfect concert 
between the efforts of accommodation and convergence. The 
former secures distinct vision; the latter single vision. 

Accommodation affects the nature of the images thrown on 
the retin ; convergence affects their position on the retin, so 
that they still fall on the same portions whether the object 
looked at is near or distant. If distant, both accommodation 
and convergence are 7i/. With every approach or recession of 
the object, they increase or decrease simultaneously. The two 
efforts are not only associated in their daily exercise, but the 
nervous centres which govern them are linked in the brain by 
strong nervous ties, so that the slightest action of one affects 
the other. This is shown by Donders’ experiments, for, though 
they demonstrate that the desire for single vision has power to 
overcome the nervous ties within limits, when lenses or prisms 
are used, yet they show also that the slightest alteration in 


1 To which I am indebted for most of what precedes. } 


| 
| 


CONVERGENCE AND ACCOMMODATION OF THE EYES. 477 


convergence shifts both limits of the possible play of accommo- 
dation in the same direction. 

Further evidence was given by Dr Loring, who, while look- 
ing at an object through concave lenses, reduced the desire for 
fusion by placing coloured glass before one eye, and thus pro- 
duced diplopia. The distance between the two images varied 
with the strength of the lenses worn, showing that “for every 
degree of tension of the ciliary muscle there is a corresponding 
degree of tension of the interni.” 

Convergence, like accommodation, is brought about by a single 
effort. Hering’s theory may well be mentioned here, since it 
receives striking and repeated confirmation in the following 
pages. It is that “each eye is supplied by two innervations— 
one directed to the turning of both eyes to the right or left, the 
other to turning both eyes inward or outward.” “Both eyes are 
used in the service of the sense of sight as a single organ con- 
sisting of two separate limbs.” 

The movements of both eyes to the right or left may for con- 
venience be called “ranging” movements. They depend on two 
distinct mechanisms, which have no known connection with each 
other. Of these, one supplies the external rectus of the right 
eye and the internal rectus of the left, and turns both eyes to 
the right; the other supplies the remaining lateral recti, and 
turns both eyes to the left. When both ranging centres evolve 
an equal quantity of nervous energy the result is simply in- 
creased tension of all four lateral recti, since each internus 
antagonises its fellow externus. If one centre predominates, 
both eyes are deviated to the right or left as the case may be.! 
Stimulation of Ferrier’s area 12 in the frontal lobe causes 
among other movements turning of both eyes to the opposite 
side. It is clear, therefore, that “convergence” or intersection 
of the visual axes is not provided for by this innervation. It 
is brought about by a separate and superadded effort, and is 
provided for by a mechanism which affects both eyes equally. 


1JIn the nates Adamuk finds a common centre for both eyes, stimulation of the 
right side producing movements of both eyes to the left, of the left side move- 
ments to the right, while stimulation in the middle line behind causes a down- 
ward movement of both eyes with convergence of the axis, and in the front an 
upward movement with return to parallelism, both accompanied by the naturally 
associated movements of the pupil.—Michael Foster. 


478 MR ERNEST E. MADDOX. 


When an object is viewed in the mesial plane the effort of 
convergence causes the two visual axes to intersect at the point 
of fixation, and no effort is needed on the part of either ranging 
centre. But if the point of fixation is carried ever so little to 
the right or left of the mesial plane, convergence must be sup- 
plemented by an effort. of one of the ranging centres to carry 
the point of intersection into the required plane. 

Is the central connection between the efforts of convergence 
and accommodation complete? Though the nervous association 
can be partly overcome when necessary by prisms or lenses, it 
does not follow that it should be naturally incomplete, and it 
has generally been supposed that a normal eye when excluded 
from vision would remain im statu quo. Consistently with 
this, since the demand for accommodation is relatively greater 
in a hypermetrope and less in a myope than in normal eyes, it 
has been supposed that under the same conditions the eye of 
every myope would deviate outwards, and that of every hyper- 
metrope inwards. We shall find this is far from being the case. 


Il. The Blind-spot Method of employing the “Viswal Camera.” 


The object of this method is to ascertain the behaviour of an 
eye placed subjectively in the dark when the other eye is 
employed in vision. The blind spot, or “punctum cecum,” 
is a nearly circular gap in the field of vision of each eye dis- 
covered by Mariotte, and shown by Donders to be due to the 
fact that the entire surface of the “ optic disc” (the extremity of 
the optic nerve at its entrance into the eye) is wholly insensible 
to light. When one eye is closed, therefore, there is an area in 
the outer part of the field of vision of the other entirely devoid 
of visual impressions, and large enough, according to Helmholtz, 
for eleven full moons to stand in a row in it (Handbuch der 
Physiologik Optik, 1867). The method of its employment for 
our purpose is illustrated in fig. 1, which represents a dark box 
or camera of a flattened pyramidal shape, measuring about a foot 
from side to side and nine inches from before backwards.!_ The 
narrow end contains two visual apertures, pierced through slides 
(a, a), which permit their mutual distance to be regulated as the 
eyes of different observers require. 

To be obtained from Messrs Pickard & Curry,£Gt. Portland St., London. 


CONVERGENCE AND ACCOMMODATION OF THE EYES. 479 


The curved border of the box is built up of two arcs (d, d) 
united by a straight line nearly 2} inches long, and therefore 
equal to the average distance between the centres of the two 
eyes, while each arc is part of a circle drawn from the centre of 
motion’ of the eye of the same side. This end of the box is 
provided with three luminous points, one fixed (e) and two 


22: line 
ye lines. 


Fic. 1.—View of the visual camera with the roof removed. 


(Erratum.—The dotted lines should cross iz the crystalline lens instead of 
behind it; 224 lines should be 284 lines. ) 


movable (7,7). They are tiny apertures, which become luminous 
when the box is held up to the light. The central one (e) is 
stationary, and since it is used as the point of fixation, should be 
provided with a piece of ground glass, a letter, or cross wires, to 
fix attention.? The lateral points (7, /) are preferably coloured, 


1 This point is about 13 mm. (Donders) behind the anterior surface of the cornea. 
Nearly half an inch is allowed for the distance of the corner from the visual 
apertures, so that since the box is 9°2 inches from before backwards, points on 
its further border are 10 inches from the dioptric centres, and therefore when 
looked at require 4 dioptres of accommodation to be in exercise. A dioptre is the 
chosen unit of refractive power ; it is that possessed by a spherical lens of the 
focal length of a metre (nearly 40 inches). Fowr such lenses would represent the 
inerease in the refractive power of the crystalline lens required to focus on the 
retina distinct images of points 10 inches distant. 

? In default of these it suffices to moisten a piece of printed paper and apply it 
to the outside of the aperture. 


480 MR ERNEST E. MADDOX, 


and are pierced through brass slides (s, s) which travel in 
grooves, so that each aperture can be moved at pleasure along 
its own half of the curved end independently of the other and 
of the central one, and without the admission of any additional 
light. This is brought about by a system of long slits so cut in 
the brasswork that the two slides and the side of the box against 
which they are apposed mutually overlap each other's slits, and 
yet permit the points of light to be seen through. A graduated 
scale of degrees (made by taking as a radius the centre of the 
eye of the same side) is attached to the outer surface of the arcs, 
and indicates the angular interval between each of the movable 
points and the central one. 

The camera is nearly divided into two lateral compartments 
by a median vertical partition (>), which runs forward to within 
an inch or two of the central luminous point. It is interrupted 
by a small cross-piece of wood called the “ stop” or “ obstructive ” 
(c), which is let in through a slit in the roof, and can be made to 
travel shortly from side to side so as to intercept at pleasure the 
view of the central point (e) by either the right or left eye. 
This is shown to the right in dotted outline (g), but the central 
point (e) is perfectly visible by both eyes, so lony as the “ stop” 
is in the middle of its slit, as represented by the shaded portion 
of the figure (c). 

Since the optic nerve enters the eye to the inner side of the 
visual axis, and since all projections are reversed in position, 
there is an area on each side of the curved end of the box (repre- 
sented by a shaded circle) which corresponds to the projection 
of the blind spot of the eye of the same side, and which may be 
called the “blind area.” Tach is about an inch in diameter at 
this distance from the eye. It may be observed that vision of 
the movable points is always monocular, since the medium 
partition (b) cuts off the view of each from the opposite eye; 
whereas vision of the central point is either monocular or bin- 
ocular at pleasure according to the position of the “stop,” the 
motion of which is too short to interfere with the view of either 
of the movable apertures, though wide enough to interfere (when 
desired) with the view of the central one by either eye. 


Exp. 1.—As a preliminary, push the deft brass slide inwards until 
the point it bears is overlapped by the brass work ‘and thus 


CONVERGENCE AND ACCOMMODATION OF THE EYES, 481 


disposed of. It is not needed in the observation. Put the stop 
in the middle of its slit, and leave the ight movable point within 
the usual limits of the right blind area. Now let the subject of 
the experiment hold the camera up to the light and look steadily 
with both eyes at the central fixation point. The right luminous 
point, being in the blind area, is then out of sight so long as 
the stop is in the middle. Now push the stop to the right, and 
it will be found that though the observer does not know what 
has happened, and still thinks he sees as before with both eyes, yet 
in most cases, after the lapse of a moment or two, the hitherto 
hidden point springs into view, showing that the eye has deviated 
from its former position, and has allowed the image of the luminous 
point to fall on a sensitive portion of the retina, as in fig. 2. 
The only effect of which the observer 
is conscious when the stop is pushed 
to the right is that the fixation aper- 
ture appears less bright,! yet by so 
doing the right eye is excluded from 
vision entirely, and placed subjectively 
in the dark, since of the two apertures 
the fixation one is cut off by the stop 
and the other throws its image on the 
blind spot where it produces no im- 
pression. He is aware neither of the 
exclusion of the eye nor of its devia- 
tion. 

If now, after the eye has deviated, 
the right brass slide is drawn out- 


Fic. 2.—The vision of the central 
: ” aperture (¢) being cut off by the 
wards, the movable point it bears stop from the right eye, its axis 
again becomes lost to view in the has deviated frome to 7, and its 
blind area, showing that the devia- blind area (b') has moved to ex- 


ti herent Wikadech extent actly the same extent, so that it 
10n was outwards. S exact exten no longer conceals the point of 


may be measured in degrees by read- _licht (f). The Jeft blind area (5) 
ing off from the graduated scale, the does not move, showing that only 
position of the inner border of the  ® eye deviates. 

blind area before and after the eye has deviated, that is, first with the 
stop in the middle and then to the right. The difference between the 
two records gives the angular deviation of the visual axis. In my own 
eyes it is about 5° as a rule, though it varies from 3° to 7° or even 8°, 
according to the time of day, the temporary comparative anemia or 
congestion of the brain, the previous occupation of the eyes, and 
doubtless many other conditions. It appears to be greater in the 
morning than in the evening, and less after much reading, or with con- 
gestion of the eyes from close work or hot rooms. That there should 
be any outward deviation at all in my case was an unexpected result, 
owing to the presence of at least 2 D of hypermetropia, for it has 
hitherto been supposed that when excluded from vision a hyperme- 


1 The central aperture sometimes also appears to move slowly to the right, but 
this is not generally noticed unless attention is called to the fact. 


482 MR ERNEST E, MADDOX. 


tropic one deviated inwards.! I believe, however, that a great many 
eyes with minor degrees of hypermetropia would be found to devi- 


ate outwards, and that if this were duly estimated some of those 


difficult cases might be more readily relieved which are so sensitive 
to any disturbance of the requisite relation between convergence and 
accommodation. 

The psychical factor furnishes an occasional difficulty in the observa- 
tions when there is a constant expectation of seeing the hidden point 
appear. It may be guarded against by registering the position of the 
outer as well as the inner border 
of the blind area in both records, 
which thus mutually correct each 
other, since the same mental effort 
which might prematurely bring the 
hidden point into view when one 
border is being tested would do the 
very reverse when the other is 
under trial. Moreover, if the re- 
corded breadth of the blind area be 
found equal in the two observations, 
before the deviation and after it, 
the coincidence is reassuring as 
to the exactness of the records. 
Variations in the shape and size 
of the “disc” in no wise affect the 
experiments, since the same definite 
point in each border is taken as 
the index of deviation. The shape 
Fic. 3.—AcB was the optic angle be- Of the curved end of the box is 

fore the right eye deviated. AdB,is such that each movable aperture in 

the optic angle after deviation; it any part of its range still throws a 

is less than before, by the angle pet tiny and distinct image upon the 

deviation cBd. Wait : : Pe 
retina instead of a diffused one ; for, 
as Donders has said, “in the emmetropic eye the whole curvature of 
the retina lies in the focal surface of the dioptric system.” The 
image is about ;',th the size of the aperture, so that the latter being 


half a line wide its image is about ;4,th of an inch in width 
400 


* T am indebted to Mr Brudenell Carter’s ‘‘ Defects of Vision” for the fact 
that Hansen has recorded a few instances of ‘‘ central defect,” though Mr Carter 
had not identified them (1877, p. 141), and says: ‘‘ In every case of myopia the 
tendency of the visual axes would be towards divergence, and in every case of 
hypermetropia the tendency would be towards convergence as soon as the con- 
trol exercised by the demand for fusion was withdrawn” (p. 138). To Hansen 
then belongs the first notification of the fact that in ‘‘a few persons” an excluded 
eye diverges with the ordinary tests at reading distance. I think, however, the 
camera will show that instead of being a rare exception, this is the normal con- 
dition, though not the invariable one. Doubtless Hansen’s cases were, in one 
sense, really exceptions to the normal, in that the degree of deviation was large 
enough to be detected by the ordinary methods. 


CONVERGENCE AND ACCOMMODATION OF THE EYES. 483 


It may be stated as a simple geometrical necessity! that the angular 
deviation of either eye alters the “optic angle” (or “angle of con- 
vergence ” contained between the two visual axes), by the same number 
of degrees (fig. 3). When both eyes fix the central aperture the optic 
angle is 14°. A deviation therefore of the excluded eye to the extent 
of 5°, reduces the optic angle from 14° to 9°. From this it is easy to 
calculate that, while accommodation still remains in both eyes for a 
distance of 10 inches, the visual axes intersect at a distance more than 
half as much again (15°7 in.), and which, if it in turn became the point 
of fixation, would need 14 dioptres less of accommodation to be in 
exercise (24 D instead of 4 D).? I have tried a sufficient number of 
cases to assure myself that owtward deviation of the excluded eye 
is the rule where refraction is apparently normal or only slightly 
hypermetropic, though here and there an exception is found. Of ten 
recorded cases the average deviation was 44°, as shown in the follow- 
ing table, which also gives the angular interval between each border 
of the blind area and the visual axis before deviation—the difference 
between them gives angular dimensions of the blind spot. 


Table I. 
| | 
N Inner border of | Outer border of | Breadth of blind | ae “ 
nes | blind area. blind area. area. DEVIATION. 
a!) ae ee) ee oe eevee S Peer oe es he | 
| | 
Tey 123° 18}° 53° 0° 
2. | 123° 183° 6° 1° or 3° 
3. 124° 19° 64° 91° 
5. ll’ 17° 6° “2 
5. 123° 183° 6° 44° | 
6. 125° 183° 6° 5° 
ae 13° 19° 6° 63° 
8. 13° 185° bse HS 
9. 113° Wis 5s° (si 
10. 123° 183° 6° 77 
| 
Average, ? 44° 


If this table is at all representative (and I expect it is fairly so), it 
shows that, while deviation occurs in nearly all, its amount varies greatly 
in different individuals; in No. 10 only 63° of convergence is left, as 
attached centrally to the accommodative effort—less than one half, A 
more extensive set of observations is much to be desired to arrive at a 
more reliable average, and to seek, if possible, to note some of the 
causes of these variations, but for taking records the “ direct method,” 
to be described presently, is far to be preferred. 


1 Eue., bk. i. prop. 32. 
” See the footnote on page 479. 


484 MR ERNEST E. MADDOX. 


It has been considered by Donders, a fact at present un- 


accountable, that only a small proportion of hypermetropes - 


should develop strabismus, and that the same refractive anomaly 
should lead to squint in some cases and not in others. No 
doubt an explanation is afforded by these great variations which 
exist in the amount of convergence naturally attached to the 
effort of accommodation. So long as every hypermetropic eye 
was supposed to deviate inwards when excluded there was no 
reason why all hypermetropes should not squint. The minor 
degrees of deviation which the camera detects come thus to 
have importance. The advantages of angular measurements 
over linear ones are obvious. The latter would vary with 
camerze of different sizes, and would not permit of direct com- 
parison, whereas the former are invariable. 

It is evident from the results obtained that the central con- 
nection between the efforts of convergence and accommodation 
is still considerable, though not complete. If there were xo 
central connection the excluded eye would deviate outwards 
nearly 14° instead of only 43°. If the connection were com- 
plete it would not deviate at all. In ordinary vision there is 
perfect concert between the two efforts, since the two visual 
axes meet exactly at whatever point is accommodated for. To 
bring this about a “supplementary” effort must be in exercise 
whenever central connection is insufficient. This effort is con- 
nected with the instinctive desire for single vision, of which the 
seat is yet unknown, so that we may say the relatively complete 
convergence of ordinary vision is maintained partly by central 
connection with accommodation and partly by this additional 
effort, which is fist roused into activity by the sensible presence 
of double images, and then maintained in exercise by the fact, 
of which the nervous centre is every moment kept sensible, that 
were the effort abated the mental image would immediately 
resolve itself visually into two. To keep it from doing so the 
joint sensations from the retinee must all the while be bearing 
between them the message of continually impending (yet as 
quickly averted) double vision, by threats of double images so 
slight and frequent that they produce the required effect with- 
out our being conscious of their existence. It is difficult to 
conceive the exquisite mechanism at work so assiduously when 


CONVERGENCE AND ACCOMMODATION OF THE EYES. 485 


we remember that, if double images are produced artificially or 
by disease, it is impossible for the mind to tell to which eye 
each image belongs—whether, therefore, the visual axes are 
crossed or not, and whether convergence needs to be increased 
or relaxed to bring the images together. 

By Hering’s theory, convergence is a single effort, exerted in 
equal amount in each eye. 

It is also clear that impressions from both eyes are neces- 
sary to maintain the supplementary factor in convergence con- 
nected with the abhorrence of double images. When, therefore, 
the obstructive in the experiment 
is placed before the right eye, 
and vision is confined to the left 
only, this common effort ceases, 
and both internal recti receive 
correspondingly diminished im- 
pulses from the converging 
centre. Were this all that hap- 
pened, ¢.g., in my own case, each 
eye would deviate outwards 24° 
as represented by the dotted 
lines in fig. 2. As a matter of 
fact, however, the active one 
remains stationary, fixing the rye, 3.4.—Convergence of the visual 
central aperture, while the uncon- — rst ae Sse sera nese 

2 ellected by the coliverging innerva 
trolled one moves outwards 5°. tion; but they are jointly deflected 

This can be proved by com- {0H "ght hand cross by the rang: 
mencing the experiment with both Hering’s theory. 
lateral apertures in their respective blind areas, when it will be 
found that if the stop is pushed to the right, although the right 
lateral aperture comes into view, the left one remains hidden 
the whole time; if the stop be pushed to the left the left aper- 
ture appears while the right one continues hidden, showing clearly 
that in each case it is the seeing eye which continues stationary, 
and the excluded one which deviates, Another innervation, 
therefore, distinct from that of convergence, must come into 
play to keep both the eyes from deviating equally. This is 
found in that centre whose ordinary function it is to turn both 
eyes to the right, and which, therefore, presides over the internal 


486 MR ERNEST E. MADDOX. 


rectus of the left eye, and the external of the right eye. Jt 


compensates by a slight effort for those impulses which the left: 


internal rectus has lost from the converging centre ; but since it 
governs both eyes equally, while it maintains the convergence 
of the left eye, which would otherwise fall back 24°, it moves 
the right eye through an additional 24° (sce fig. 3 A). 

The effort put forth by this fresh innervation is determined 
entirely by the requirements of the seeing eye; it only affects 
the deviating eye because it cannot help influencing one as 
much as the other. Its intervention is proved by the next two 
experiments. The result is that exactly half the deviation of 
the right eye is due to relaxation of the internal rectus, and the 
other half is due to contraction of the external rectus; but since 
in the left eye the diminishing converging effort and the in- 
creasing ranging effort have each to do with the internal rectus, 
it remains stationary. 


Exp. 2.—With the stop in the middle, fix the central aperture with 
both eyes, and try to place the right forefinger exactly upon the central 
aperture from outside. The attempt will succeed in proportion to 
the perfectness of the observer's muscular sense. Now push the stop 
to the right, and repeat the attempt. The finger will be found to 
have missed its mark, and to be actually on the right side of it; and 
similarly to the left side of it if the stop is pushed to the left. The 
miscalculation will be slight if the attempt is made directly after the 
exclusion of the eye, and greater with every increase in the interval 
which elapses till the maximum miscalculation is reached, which in 
my case is about a distance which corresponds to 23° on the gradu- 
ated scale. The right eye, we have seen, has meanwhile moved 5°, 
It may therefore be accredited as a rule that the angle of miscalcula- 
tion is half that of the deviation of the excluded eye; it is slight at 
first, because the deviation is slight, and they increase together in the 
proportion of 1 to 2. 

It has long been known that when one eye is closed, and a finger is 
pushed forward from under a book, it misses its mark to the side of 
the closed eye; but I believe this phenomenon will be absent in 
those with whom deviation of an excluded eye does not occur at the 
distance of the test; and that the extent of miscalculation will be 
found to depend entirely on the amount of the deviation, and to be 
half as great. 


Expr. 3.—If the central aperture is very closely watched its apparent 
position may be observed to move slowly to the right as soon as the 
stop is pushed to the right. Now, it is remarkable that the point of 
view should seem to be moving when not only is the point really 
stationary but also the image it throws on the retina, and the retina 
itself. Since only one eye is in this case engaged in vision, and that 


CONVERGENCE AND ACCOMMODATION OF THE EYES, 487 


(as may be shown by the immobility of its blind area) keeps quite still 
the whole time, there cannot be the slightest change in the comparative 
tension of its recti, to account for the apparent movement of the 
image. Moreover, though the excluded eye deviates, we shall see 
later that the oculomotor muscular sense is purely central and not 
peripheral, since the same degree of tension in a muscle is mentally 
estimated or mentally ignored, according to the central source of the 
impulses which cause the tension. The stillness of the seeing eye 
therefore proves that the illusion is due to some alteration in central 
nerve effort of which the mind takes (what is now) unnecessary cog- 
nizance, and thus forms a false estimate. 

The new effort is also shown by the nature of the apparent move- 
ment to be the one which the mind has been accustomed to associate 
with lateral displacement of the point of fixation, and with the joint 
movement of both eyes to the right, which such displacement makes 
necessary in the ordinary vision of nature. The illusion cannot be 
due to the diminution of converging effort, because that, as we shall 
see, is mentally associated only with the idea of distance, not at all 
with the angular departure of the object from the median plane, or its 
position in the field of vision. The slowness of the apparent move- 
ment is a striking feature ; it shows how gradually the ranging effort 
is put forth, consistently with the gradual diminution of the converg- 
ing effort for which it exactly compensates. 

It is a fact which affords some food for thought, that although the 
stumulus which causes the “supplementary ” converging effort ceases 
suddenly when the stop is pushed to the right, yet the effort itself 
continues for some time decreasing only gradually. This is in strik- 
ing contrast to the speed with which full convergence is again effected 
when the stimulus is restored. The gradual relaxation of the con- 
verging effort when the stimulus is withdrawn, causes loth internal 
recti to receive growingly feebler impulses from the converging centre, 
so that each eye has a constant and momentary tendency to deviate 
outwards, which is only prevented in the left one by the wonderful 
vigilance of the nervous mechanism which every instant appreciates 
this tendency, and as quickly compensates for it, not by again stimu- 
lating the flagging convergence, but by causing a strictly propor- 
tionate and gradual increase of that effort whose output causes in the 
mind the impression that the point of view (really stationary) is 
moving to the right. It need hardly be said that all this naturally 
accords with and establishes Hering’s theories mentioned on p. 477. 
The apparent movement of the central aperture is through half the 
angle and at half the rate of the real movement of the deviating eye. 
A little reflection on the preceding experiment will show the truth of 
this, as nearly as it can be determined, and also that when an object 
is fixed not far from the middle line its position is mentally referred 
to the vertical plane which bisects the angle of convergence, and 
which, as we shall see, runs through a point midway between and 
a3) behind the centres of the two eyes. (See the line yp in 

g. 3. 
After a few attempts to touch the point thus miscalculated, the 


488 MR ERNEST E. MADDOX. 


mind allows for the error, and the attempts begin to succeed. It has 
already been suggested that thousands of such attempts in childhood 
contribute to the wonderful correlation between the muscular sense of 
the eye and the hand. How perfectly they may by practice be made 
to co-operate is seen in a good cricketer or marksman. 

The senses are there to begin with, but the mental apprehension of 
their import, both singly and jointly, seems to be largely left to be 
perfected by education. Indeed, it is known how any sense itself 
may be quickened by receiving a larger share of psychical attention, 
or dulled by its prolonged abstraction. 

The human body is thus made capable of adapting itself within 
limits to adventitious circumstances ; it is not made, like an ordinary 
loom, capable only when once set cf turning out material of one tex- 
ture,—but it is like a loom, if one can be conceived, made with such 
wonderful skill and forethought that it can automatically adapt itself 
to the requirement of any new material and other altered circum- 
stances. 

I find, on trying to touch the central aperture with my left hand, 
that when the stop is to the right, instead of missing its mark to the 
right side of the central aperture aimed at, it misses it to the left side, 
and when the stop is in the middle it misses it still more to the left 
side, though its miscalculation is not very precise. Its muscular sense 
is therefore less perfect. 

Exp, 4. On first opening the eyes in the morning the divergence 
is greater than during the day; it falls just after the mid-day meal and 
perhaps after the others. 

Exp. 5.—When vision is directed through either the central aper- 
ture or the left lateral one at an object placed at different distances, 
accommodation is, of course, diminished in proportion. It will be 
found that the excluded eye moves outwards with each removal, and 
inwards with each approach of the point of fixation. This shows 
how delicate is the connection between the two efforts, since the 
slightest difference in accommodation causes an alteration in the 
degree of convergence. 

Exp. 6.—If convex glasses of increasing strength be placed in turn 
before the active eye, the blind area of the obstructed eye moves out- 
wards with each increase in the refractive power of the lens employed. 
With concave glasses, on the other hand, it moves inwards with every 
increase. This experiment, of course, differs only from the last in 
the method employed ; which, indeed, is far less satisfactory, owing to 
the- fallacy introduced by prismatic action of the lenses, if their 
optical centres are not placed exactly in the line of vision—a pre- 
caution of great difficulty. 

Exp. 7.—When the box is sloped downwards from the eyes, I 
have records which show that the deviation of the obstructed eye is 
reduced by 2° or 3°. I am not quite satisfied, however, with the 
observations—the bridge of the nose almost obliges the box to be 
held at a greater distance. The way to get over the difficulty would 
be to use prisms with their bases upwards, which would permit the 


CONVERGENCE AND ACCOMMODATION. OF THE EYES. 489 


box to be held horizontally, and yet record the effect of a downward 
direction of the visual axes. The ordinary circular prisms used in 
practice are not available for this purpose, owing to the difficulty of 
placing the centre of the base exactly in the vertical line which bisects 
the prism. A slight shift to either side not only reduces the vertical 
deflection of the line of vision, but introduces a still greater /ateral 
deflection, which vitiates the result. Small prisms fixed 7m the visual 
apertures would be most satisfactory. 

Exe. 8.—If the central aperture be fixed by the left eye, with the 
obstructive to the right, it is possible to place the right lateral 
aperture so precisely upon the inner border of the right blind area 
that the point of light alternately appears and disappears, showing an 
evident tendency in the nerve centre to rhythmic, or at least irregular 
action. ‘This irregularity furnishes a striking contrast to the fixed- 
ness of gaze and precision of movement in ordinary binocular vision. 
It devolves upon the supplementary effort in single binocular vision 
to fill in these irregularities in the fluctuating basis, besides meeting 
the new and changeful requirements constantly introduced in glancing 
from point to point. It is interesting to notice that this fluctuating 
effect in the converging centre is connected with the evolution of a 
stewly stream of nervous energy from the accommodating centres. It 
may perhaps bear some comparison with the rhythmic automatism 
which manifests itself in the vasomotor centre under the uniform 
stimulation of venous blood, as evidenced by Traube’s curves. 


Exe. 9.—With both eyes fixing the central aperture, and with the 
obstructive in the middle, place the right lateral aperture in the outer 
part of the blind area at a definite number of degrees from its inner 
border. Push the obstructive to the right, and note how long a time 
elapses before the hidden point comes into view, by listening to a 
clock pendulum beating half-seconds. As might be expected from 
Exp. 8, the interval is a variable one. Thus, at one sitting, my right 
eye was engaged from 124 to 22 seconds in rotating outwards 33°. 

Exp. 10,—After wearing convex spectacles for some hours, I find 
that for a’time the relative divergence is diminished (by the training 
the converging centre has undergone in the increased relative demand 
made upon its energies). How long this effect lasts I have not been 
able to observe. 

Exe. 11.—Measurement of the Blind Spot.—I have found the 
angular dimensions of the blind spot in its horizontal meridian, as 
far as the box measures it, very uniform. In nearly all cases it was 
approximately 6°. So far as the observations are worth, they go 
therefore to confirm Landolt’s estimate of 6°, rather than Helmholtz’s 
of nearly 7° (6° 56’).1 The method they both employed was that of 
moving a pencil on a piece of paper till the point became lost to view. 
With one who has thoroughly practised indirect vision this suffices, 
but for others it is very uncertain. Thus Helmholtz says: ‘I have 
even seen men of education and information—doctors, e.g.—not able 


1It must be remembered, however, that any error of the box from not 
measuring the exact horizontal meridian tends to give too small a result. 
VOL, XX. 21 


490 MR ERNEST E, MADDOX. 


to prove the disappearance of small objects on the blind spot.” 
Hanover and Thomson, in 22 eyes (quoted by Helmholtz), found 
the breadth to vary from 3° 39’ to 9° 47’. I believe cases of less 
than 5° or more than 7° will be found exceedingly rare. In taking 
measurements, the stop should be either in the middle or to the 
opposite side of the eye under examination. I believe it is better to 
start with the point hidden, and let the observer exclaim at its first 
appearance at either border, rather than to note its disappearance, 
though the two may check each other. 

A point of light is peculiarly fitted for the purpose, owing to the 
comparatively great susceptibility of the peripheral parts of the retina 
to light. Brewster? stated that astronomers, when they cannot see a 
minute star by looking directly at it, may often bring it into view by 
looking somewhat away from it. Landolt,® however, finds “ the per- 
ception of light remains almost exactly the same throughout the whole 
extent of the retina.” He instances that in his right eye the percep- 
tion of light at a part 30° from the centre remains the same, while the 
visual acuteness is reduced to 4; but certainly, in my own eyes, the 
point of light appears to be more easily discerned on its emergence 
from the znne (macular) border of the blind area than from the outer 
border—it may not be so with others. Clinically, the measurement 
of the blind spot may be useful, both to determine the increase of the 
posterior staphyloma of progressive myopia and to trace the progress 
and decline of such affections as optic neuritis, in which the adjacent 
retina loses its perception awhile by infiltration. 

A disadvantage is, that in the original instrument the two lateral 
apertures are not upon the same level, and therefore one of them (the 
highest) measures the blind spot above its horizontal diameter, and 
gives a uniformly smaller and fallacious record. ‘This may be rectified 
by using, instead of slides, two flexible ribbons arranged circularly, 
so as to have the lateral apertures on the same level. 

It is well to have the point coloured due, since the peripheral parts 
of the retina perceive this colour most readily. If we assume that 
an angle of 4°, with its apex at the optical centre of a normal eye, 
subtends 1 mm. of the retina, then 6° would subtend 1} mm.; 
showing the close coincidence between the anatomical and physio- 
logical dimensions of the disc. The angular distance between the visual 
axis and the border of the blind area I have not found so uniform as 
the breadth of the blind spot. Landolt and Dobrowolsky found the 
interval greater in hypermetropes and smaller in myopes.? It would 
be well to confirm this by the camera. 


! Optique Physiologique, p. 735. 

* Brewster on Stercoscope, 1856, p. 44. 

3 Landolt, on Examination of the Eyes (translated by Dr Burnett, 1879, 
Philadelphia), p. 214. 

4 Examination of the Eyes, Landolt, 1879, p. 216. 


—— 


' 


CONVERGENCE AND ACCOMMODATION OF THE EYES. 491 


Ill. Zhe Direct Method. 


This method is far more useful clinically, and not less inter- 
esting physiologically. The eye is not placed in the dark, nor is 
the blind spot made use of. It depends upon the fact, that when 
each eye receives a single image upon its median vertical 
meridian, from whatever points they are thrown, the two are 
mentally referred to the same vertical line. 


Exp. 12.—Place the /efé aperture out of sight and the obstructive to 
the right ; the observer then sees 
the central and the right lateral 
apertures. As he looks, they ap- 
pear to approach. The right slide 
is then pushed inwards till they 
seem to lie in the same vertical 
line. The process is now com- 
plete; it will be found that a 
real interval separates the ap- 
parently superimposed apertures. 
This interval expresses in de- 
grees the relative divergence of yye.3p,—Illustratesthe “direct method.” 
the eyes, for one visual axis The apertures appear superimposed 
passes through one aperture, while — though really separated by the deviat- 
the second lies either above or 8 angle of the eye. 
below the other. I have found this method quite easy in a child of 
six.! In comparing its results with those obtained by the blind spot 
method, I found that they coincided, showing that the mere addi- 
tional presence of an image upon the retina does not affect the con- 
vergence and accommodation, so long as the desire to unite double 
images is eliminated. In the blind spot method there is an image in 
one eye, in the macular method in both. Its explanation is simple. 
Since the view of the right point by the left eye is intercepted by the 
median partition, and that of the central aperture by the right eye is 
cut off by the obstructive, each eye sees only one point, and that a 
different one, as shown in fig. 3B. From the nature of the curve at 
the base of the camera, accommodation is required from each eye in 
equal amount (or practically so). If now the brain relationship were 
complete, when attention is directed to one aperture, say the central 
one, both visual axes would converge toward it, while the image of the 
right point would fall to the znner side of the macula of the right eye, 
and would be correctly referred outwards to its real position in space. 
This, in fact, does continue momentarily, when first the points are looked 
at. As soon, however, as relative divergence commences, and the 
right eye deviates outwards, the image of the right point approaches 


' It is convenient for children to remove altogether the little wooden slides 
bearing the visual apertures. 


492 MR ERNEST E. MADDOX. 


the macula, or, more correctly, the macula approaches the image, for 
it is the eye which moves and not the point. While this is going on, | 
————— the two stationary apertures 
appear to be getting nearer to 
each other, for the cerebral 
centres are unconscious of the 
divergence, and make no 
allowance for it. The images 
do not appear to meet com- 
pletely until each falls upon 
the median vertical meridian 
of its eye. It is well to begin 
the experiment with the aper- 
tures at some distance from 
each other, and after allow- 
ing a short time for them to 
approach naturally as far as 
they will, to push the right 
slide inwards, and let the 
observer say when they come 
into the same vertical line. In 
this part of the process the 


Fie. 4.—The direct method. Each lum:n- : : . 
ous point throws an image on the fovea of @/@ remains stationary while 
the eye on the same side, so that both the 7mage ts moved, on to its 
images are mentally referred to. the plane median vertical meridian. 


which bisects the angle of convergence. The dialocue would be 
C 
o 


something like this :-— 

Q. What do you see!—A. Two bits of light. 

Q. How far apart?—A. An inch or two. 

(. What happens? (pushing on the right slide slowly).—d. The 
right one is moving to the left. 

@. Say when they are quite together, that is, when the right point 
comes to be exactly below the left.—A. Now! 

This concludes the observation. The real interval between the two 
points, automatically recorded by the graduated scale at the base of 
the box, has only to be read off to give in degrees the relative diverg- 
ence of the eyes. This method dispenses with the use of prisms and 
the fallacies which attend them; it saves the trouble of special 
measurement, and gives an angular instead of a linear record, which is 
therefore always ready for comparison. It is equally available by 
daylight or artificial light. 


But the best practical evidence of its efficiency is afforded by 
the ease with which it reveals the physiological prevalence of 
relative divergence in near vision, while the ordinary methods 
have only hitherto detected the grosser pathological exceptions. 
I may not be acquainted with all of them, and therefore cannot 
indicate the reasons of their failure, but I think I can suggest 


CONVERGENCE AND ACCOMMODATION OF THE EYES, 493 


what they are in Von Graefe’s well-known test, which when 
carried out as usually directed, does not reveal the slightest 
relative divergence in my own eyes, though, as we have seen, 
5° really exists on exclusion. I have not had access to Von 
Graefe’s own directions. I may quote those in Mr Carter's 
valuable treatise on Defects of Vision, az [ followed them :— 

“Tn this more delicate test the object of vision is a small black dot, 
bisected by a vertical line. A card thus marked is fixed in the 
median line at a distance of 8 or 10 inches from the eyes, and the 
patient is directed to look at it steadily. A prism of ten or twelve 
degrees, with its base either upwards or downwards, is then placed 
before the eye ; and as the power of the superior or inferior rectus to 
overcome double vision is very limited, this prism necessarily produces 
a vertical diplopia. The patient will therefore see two dots, one above 
the other. If the original convergence for the object is accurately 
maintained, the duplication of the vertical line will only cause it to 
appear elongated, and the two dots will be seen one above the other 
on the same line. If, on the contrary, the convergence be not main- 
tained, the patient will see two lines with a dot upon each ; and when 
the diplopia is a consequence of relative divergence of the optic axes, 
the double images will be crossed, and the extent of the divergence 
will determine the distance between them.” 

On carrying out these instructions the dot truly duplicates and 
the line elongates, but thatisall. The line still continues single. 
The reason of this becomes evident when the further step is 
taken of covering one eye for a short time ; on again uncovering 
it, two lines appear, separated by a considerable interval, but 
they quickly run together again. This shows that the desire for 
fusion, though doubtless weakened, is not removed altogether, for 
the overlapping portions of the two linear images are sufficient 
to excite it. We shall see that images need not be similar in 
shape to excite an effort to unite them. Indeed, in ordinary 
vision the two pictures, as illustrated by the stereoscope, are 
slightly dissimilar except when the objects viewed are at a 
practically infinite distance. But I find if the upper part of the 
line be drawn very wavy, and the lower part straight, so that in 
the experiment the wavy portion overlaps the straight portion, 
there appears to be no attempt to unite them, though even then 
would not be quite sure that there is not a faint effort to keep 
them nearer to each other than they would otherwise be. 

The fallacy may also be demonstrated in another way without 
temporary exclusion of either eye, by simply holding the line at 


494 MR ERNEST E, MADDOX, 


first horizontally (with the prism as before) and then quickly 


returning it to the vertical position; the two images for a - 


moment or longer are quite separate, and hesitate a little before 
they run together. 

“Why then,” it may be asked, “ if the test does not eliminate 
the fusion effort, does it ever reveal relative divergence?” It does 
so because, though it does not, like the camera, remove the desire 
for single vision, yet it lessens it to such an extent that it 
becomes inadequate to the demands made upon it in certain 
pathological conditions. The test weakens the desire for single 
vision, not only by the effect on one of the images of the slight 
light-absorbing (especially when the prism is not perfectly clean 
and free from moisture) and chromatic properties of the prism, 
but also by shortening the linear extent of the overlapping por- 
tions of the two images of the line. It would therefore detect 
relative divergence in such conditions as (1) those probably very 
rare cases in which the normal desire for fusion is defective. 
By lessening the desire still further it might be rendered incap- 
able of rousing a sufficient “supplementary ” converging effort. 
(2) Where the mechanical difficulties which attend convergence 
are so great that no effort can overcome them unless prompted 
by a strong fusion stimulus, as in some extreme cases of myopia, 
or where there is weakness of the internal recti or functional 
disability in their innervation. (3) Where almost the whole of 
the required convergence devolves on the fusion effort. 

In all cases of myopia a larger share falls to the fusion effort 
than in the normal eye, because there is less demand for the 
effort of accommodation in looking at any point, and therefore 
the degree of convergence due to central association is correspond- 
ingly small. The smaller it is, the more work it leaves for the 
fusion effort, so that, “ ceeteris paribus,” the greater the refractive 
anomaly the larger is the required proportion of supplementary 
or fusion effort. 

A great effort needs a great stimulus. The latter is so weak- 
ened by the prism that, while still adequate for the requirements 
of normal refraction, it may be inadequate for those of high 
myopia, in which, moreover, mechanical difficulties almost 
always exist as well from the altered shape of the globe. 

To make the test of any relative value even in these cases, 


CONVERGENCE AND ACCOMMODATION OF THE EYES, 495 


care must be taken to make the line of always the same length, 
or if not, to adjust its distance from the eyes in proportion; so 
that the reduplicated portion of the line may always be of the 
same length, and thus ensure wniform diminution of the desire 
for fusion, otherwise the test might at one time detect an insuffi- 
ciency and at another time not. Moreover, the line which joins 
the apex and base of the prism must be exactly at right angles to 
the line uniting the centres of the two eyes (intercentral line) ; 
otherwise, though the lines continue parallel, their very opposi- 
tion would only prove that convergence is not complete—if it 
were so, the lines would be separated by an interval determined by 
the strength and degree of rotation of the prism. Even when 
the prism is held correctly, if the dine looked at is not also held 
exactly at right angles to the intercentral line another fallacy 
ensues, for the linear images, though still parallel are oblique, 
so that coincidence of their overlapping portions, instead of show- 
ing convergence to be complete, can only take place when it is 
incomplete, for were it complete an interval would separate 
them, varying as before with the degree of rotation of the card. 

These difficulties, 1 would suggest, may be overcome by the 
use of a double prism composed of two prisms, each of 2°, fused 
together by their bases! (see fig. 5). The patient, shutting the 
left eye, holds this prism before the right one, and looks through 
it at a card marked with a single dot or short line. Two false 
images appear, one 2° above and the other 2° below the real 
position of the dot, and both are seen by the right eye. It is 
easy for the patient to hold the prism so that the two images 
appear in the same vertical line, and then when the left eye is 
opened as well to say whether the veal image of the dot lies to 
the right or left of this line. Even if the first two are not held 
vertically, if all three images are in one straight line it shows 
that convergence is complete. If the central one lies to the 
right of th2 line, uniting the other two, there is relative 
divergence ; if to the left, there is relative convergence. 

Simple as this expedient is, and though it yields the same 
result as the camera, it is inferior to the use of the latter by the 


1 In reality, of course, it is a single prism of 176° though double in its use, 
since three faces are used instead of two. The large face (or base) should be 
towards the eye, the two smaller faces towards the object. 


496 MR ERNEST E, MADDOX. 


direct method. The camera ensures uniformity in the distance 
of the object from the eyes without the trouble of measurement ; 
it needs less intelligence in the patient, and gives an automatic 
angular record. The double prism, however, would I think be 
found useful for rough analysis at greater distances. The 


Fic. 5.—Side view of the right eye and the double prism. The false images seen 
by the right eye are dotted. The central one is seen by the left eye. 
radical difference between Von Graefe’s test and the camera is 
that in the latter a separate object is used for each eye, while in 
the former the same object is reduplicated by a prism. The 
camera also not only reduces the desire for single vision, but 
abolishes it altogether when the lower of the two lateral apertures 


7° 6° 5° ye 3° 2° 19» 90% Be: 1:20 568-2 7° 
B, 


A 5e 40. go-go. f0 jo." 20° 30 he “Ge 6? 7° 
Ul 


79 © §°. 49 39 2° yo 2° 839 4e 5? 6°. 7° CG 


C, 


Fic. 6.—To illustrate how relative divergence is measured by the double prism. 
A is the only device on the card, and is seen by the left eye; B and C 
are false images of it, and are seen by the right eye. In this instance 5° of 
deviation are seen recorded. If the two lowest arrows are made continuous by 
rotating the prism, the middle one points to twice the divergence, for as C 
moved to the right, B moves equally to the left, A of course remaining 
stationary. The arrows would all but touch the lines above them when 
the card is held at the appropriate distance of 10 inches. 


is used in conjunction with the central one, so that the eye 
takes a position determined solely by the converging effort which 
is associated with the accommodation. 


CONVERGENCE AND ACCOMMODATION OF THE EYES. 497 

If when, in the “direct method,” the two images are in the 
same vertical line, as in fig. 3 B, an effort be made from outside to 
place the finger on them, it will miss both, for it will be just 
half-way between the two actual apertures, which, though they 
appear superimposed, are, as we have seen, really separated by 
an interval of nearly an inch, so that the vertical plane in which 
the two images appear to lie is that which bisects the angle of 
convergence, as represented in fig. 4. At present we have only 
to do with movements of the eye in the horizontal plane, and 
with the head stationary. The converging apparatus appears to 
be solely connected with the union of double images and the 
estimation of distance. With the relative position of points 
along the horizontal meridian of the field of vision it has 
nothing todo. This must be determined entirely by— 

(1) The part of the retina on which images fall. 

(2) The innervation which turns both eyes to the right or 
left. 

As regards the first indication, since eack image falls on the 
median vertical meridian of its eye, the effect is the same as 
though they were both thrown from one vertical line, for which 
convergence were complete; and, since the relaxation of the 
converging effort is not taken into account, there is no reason 
why the images should not be referred to the median plane, for 
there is nothing so far to give any preponderance in favour of 
either side. 

As regards the second indication, however, as seen in fig. 3, 
while convergence occurs to the left-hand cross, both axes 
are directed to the right-hand cross by the ranging innervation 
of which the mind does take cognisance, Now, the inclination 
of the plane which bisects the angle of convergence, to the median 
plane, exactly represents the angular effect of the ranging 
energy which is in exercise—hence the images are referred to 
this line. It is now easy to observe the fluctuations in the 
stream of nervous energy noticed with the blind spot method 
on p. 487, for one point continues to make tiny excursions to 
the right and left of the other, though without any regular 
rhythm. This makes it useless to take very exact records in 
minutes and seconds. It is also clear that a more accurate 
method is scarcely to be desired, since it would only magnify 


498 MR ERNEST E. MADDOX. 


these irregularities. Care must be taken that the difference in 


level of the two apertures is enough to avoid continued effort to - 


unite them. It is remarkable that, when their vertical separa- 
tion is only slightly more than enough to prevent optical union, 
a tendency may be noticed for them to keep near the same vertical 
line. Even when one is pushed a little way to the right or 
left the other is apt to follow it, and this in spite of their 
dissimilar shape. It is even noticeable when the apertures are 
coloured differently ; but disappears very rapidly with increasing 
vertical separation of the two points, and is not in my own 
eyes detected in the slightest degree when the Jowest of the 
two lateral apertures is the one employed, in conjunction with 
the central one, which is, of course, the highest of the three, 
being separated from the Jowest movable point by an angle 
of 2° (from the eyes), and from the highest point by slightly 
less than 1°,in the camera with which I experimented. This 
latter interval is one which can be overcome at times by the 
superior or inferior rectus in order to satisfy the desire for 
fusion, especially when the eye has succeeded a few times, and 
acquired the facility. After allowing it to do this for a time, 
the following experiment may be made :— 


Exp. 13.—Place the stop to the deft. Let the left aperture be as 
before entirely occluded, and the right one be placed 3° or 4° away 
from the central aperture. Look through the camera thus for about 
a minute, during which interval the right eye sees both the images, 
and the left neither, so that the latter is deviating outwards. 

Now, push the stop from the left to the right. This proceeding 
transfers the view of the central aperture from the right eye to the 
left one, which, being deviated 5°, miscalculates its position, and 
refers it to the right of the other point still seen by the right eye. 
The two points thus separated now by a small interval run together, 
though to do so it is clear the relative divergence is diminished by a 
slight converging effort. If to start with the right lateral point is 
placed 6° away from the central one instead of 4° when the experi- 
ment is repeated, though the points appear separated by the same 
interval as before yet their position is of course reversed ; yet they still 
run together. In this case the relative divergence is increased to 
meet the desire for fusion, instead of being diminished. Whether 
this is brought about by inhibition of the centres for the internal 
recti, or by antagonism of the external, remains yet to be found out. 

Exp. 14.—Were the right lateral aperture, to start with, placed 
5° away from the centre, and the last experiment repeated with the 
stop to the left, the two points would appear separated by nearly an 


CONVERGENCE AND ACCOMMODATION OF THE EYES. 499 


inch, and with the stop to the right they would appear in the same 
vertical line. This enables the observation of a patient by the “ direct 
method” to be easily confirmed, for all that is needed after taking the 
observation to ensure that fusion effort is eliminated, is to push the 
stop back to the right, and again to the left, to see whether the point 
at its first reappearance occupies the same position as before. 


Exp. 15.—Use the highest lateral aperture on the right side, and 
with the stop to the right, move it till it meets and appears to 
fuse with the image of the central aperture. After fusion, push the 
right brass slide inwards slowly and steadily, and it will be found that 
the two blended images move to the left together, for the one which 
really moves carries the other with it to preserve fusion, and this goes 
on until the moving aperture travels right up to the central one, 
or at least as near to it as the construction of the box will permit, 
so that even this fw/se fusion has sufficient power to undo the whole of 
the relative divergence, 

If on the other hand the brass slide with the movable point it 
bears is drawn outwards, resolution of the two images does not occur 
till the points themselves are really separated by 10°. The desire to 
continue the false fusion is thus strong enough to double the previous 
divergence. Albeit the experiment makes the eyes water and feel un- 
comfortable. Whether this discomfort is due to per/pheral antagonism 
of two sets of muscles, the external recti and the internal, I cannot 
tell; or whether it is due to a central struggle to overcome by inhibi- 
tion a nervous connection probably never before invaded to that 
extent. Perhaps if the two points were on the same level the attain- 
able divergence might be still greater. As it is, the deviation of 10° 
brings the eyes to within 4° of parallelism. This false fusion is of the 
same nature as that described by Sir D. Brewster, when, in looking at 
a patterned wall it is possible to converge the eyes for a point so far 
behind, or in front of the wall, that fusion of the laterally adjacent 
patterns takes place. The strength of the effort put forth to maintain 
accomplished fusion is much greater than that instigated by the desire 
to unite the two images when they are separated to begin with. The 
relative divergence attainable by the present experiment is much 
ereater than, and must not be confused with, that attainable by the 
effort to overcome a prism, for in the latter case the two images are 
separated to begin with by the act of placing the prism before the eye. 
It may be deduced from what has preceded that, when the brass slide 
bearing the right luminous point is drawn outwards or pushed inwards, 
the fused images appear to follow at half the rate. But if they are 
not fused only one of them appears to move, and that at a rate equal 
to that at which the slide travels; the difference being that in the 
latter case both eyes are stationary, whereas in the former, while the 
left eye remains fixed the right one moves at the same rate as does the 
brass slide which bears the point of light it perceives. If the slide is 
moved jerkily slight momentary separations of the images result. If 
the effort to maintain false fusion be so strong, probably that to 
maintain ¢rue fusion is greater still. ‘To measure it a camera should 
be used with two apertures at the same level, or else with a prism let 


500 MR ERNEST FE. MADDOX. 


into one of the small wooden slides before the eyes to just rectify the 


difference in level. It is the effort to maintain existing fusion which 


is tested by the common practice of approaching a finger to the eyes 
till one of them rolls out, though in this test accommodation increases 
at the same time, while in the camera accommodation is unchanged. 
I may note in passing that in my own eyes the nearest point of single 
vision by the finger test is closer to them than that of distinct vision, 
which illustrates a fact almost self-evident, that the relative divergence 
which occurs on the exclusion of one eye does not indicate deficiency 
in the converging function itself, but only in the dink which connects 
it with accommodation. Accommodation assists convergence, and 
convergence accommodation, but they do so only through the central 
link which connects the two efforts, and enables one to influence 
the other. The slighter the link the less the effect one has on the 
other—but that has nothing tu do with the individual strength of 
either, 

Donders has shown that hypermetropes fix more easily when they 
look through prisms which make them converge more strongly. Is 
this because the converging effort assists that of accommodation by 
means of the central link between them? Apart from any pathological 
affection of either centre it is reasonable to suppose, since the sympathy 
is mutual, that if accommodation exerts only a weak influence on con- 
vergence, convergence will have a correspondingly weak influence on 
accommodation ; a unit of either will contribute less than usual to the 
other. 

If this be so, relative divergence, as revealed by the camera, since it 
indicates imperfection in the channel of mutual assistance, would 
lessen the advantage of the prisms above mentioned—though it would 
remove in their use all fear of their causing squint. But this is 
theory and needs practical confirmation. 

A little confusion has arisen from the incorrect supposition that the 
s'rength of the internal recti is tested by prisms base outwards, and 
the ability to overcome them ; whereas it is the conditions of the con- 
verging reflex as a whole which are thus estimated, including the 
existing intensity and activity of the desire for single vision. This 
is clear from the fact that when both eyes are directed to the 
right or left the contraction of the internal rectus may be greater than 
can possibly be attained by converging effort, the innervation called 
into play being a different one. Inability to overcome such prisms of 
high power might of course be due only to weakness of muscles, but 
in that case the ranging and converging movements would be equally 
impaired. Moreover, since accommodation remains unchanged, such 
prisms only indicate the limits of attainable relative divergence 
or convergence, which depend largely on the strength of the 
central nervous connection between convergence and accommodation. 
Approaching the finger to the eyes till one rolls outward is another 
method of testing the strength of converging effort, though sf it is 
not the efficiency of the internal recti only that is indicated, but of the 
whole converging sensory-motor apparatus—afferent, central efferent, 
muscular, and mechanical. Strength of fusion effort is also influenced 


CONVERGENCE AND ACCOMMODATION OF THE EYES. 501 


by the nature and doubtless by the size and number of the images to 
be fused, as well as by the amount of attention directed to them. 


Expr. 16.—There are some cases of strabismus, especially of the 
divergent kind, in which, as Donders has pointed out, the mind be- 
comes conscious of the direction of each eye. , Such a person can cor- 
rectly calculate the position of any object seen by either eye, and, 
indeed, employs one or the other just as convenience requires, being 
able to distinguish very readily which he is using for observation. 

In testing a case of slight external strabismus with the camera, one 
would rightly expect to find that when the two images by the direct 
method are in the same vertical line there would be a very great 
interval between the actual apertures. This would be so in recent 
cases, or any in which one eye is disused in ordinary vision, for 
however much deviation might really exist, the images on both 
maculze would still be mentally referred to the same vertical line. 

But in cases like those mentioned by Donders the fact is that the 
eyes correctly estimate the distance between the two apertures, so that 
the images do not appear superimposed at all, unless the apertures 
themselves are made so, which the construction of the camera does not 
quite admit. The axis of the deviated eye does not follow the moving 
point of light, but correctly estimates its position as its image travels 
along the retina. An instance of this rather puzzling anomaly was 
tested with a camera by Dr Joseph Bolton. The “direct method” is 
useless for such cases, and will not detect even any deviation, being 
like the usual prismatic ones too subjective ; but the “blind spot 
method ” enables the exact position of either eye to be noted. <A 
careful examination of a few of these cases might yield interesting 
results. All that is necessary to discover them is to try the two 
methods and see whether their records differ. 


Exe. 17.—When the two images in the “ direct method” are looked 
at for some time with a dim illumination they may be observed to 
alternately disappear altogether. This favours the current view that 
the part played in vision by the visual apparatus of the two eyes alter- 
nates in intensity ; as each in turn becomes tired, the other gives it a 
rest. 


Exp. 18.—While looking at the two images by the ‘‘ direct method” 
shut the right eye; the image seen by it moves upwards and to the 
left, showing the eye itself has moved downwards and to the right. 
Why? 

Exp. 19.—To ascertain the vate of the deviation of an excluded eye. 
With the stop to the right, and the right luminous point a consider- 
able distance from the centre, look through the camera till convinced 
that no more deviation will occur. Move the right point inwards till 
its image seems just below the other, and eave tt in position. After a 
timed interval of rest, take up the camera again and look at the cen- 
tral aperture with the stop in the middle, listening to the beats of a 
clock pendulum. At one end of a beat push the stop to the right, 
and count the number of seconds which elapse until the two lumi- 
nous points meet, for the right one has been left in position all the 


502 MR ERNEST E, MADDOX. 


while. The interval varies considerably in different cases and at 
different times, from half a minute to a minute and a half. In 
one case the total divergence of 64° appeared to take from 68 to 73 
seconds, 

In the same case, when the two points were separated to begin with 
by 5°, the images took 40 seconds to meet. When separated by 2° 
they took 5 to 8 seconds. In this way the rate can be estimated for 
each degree. It is clear that the eye deviates outwards with diminish- 
ing rapidity, at any rate in the latter parts of the deviation. 


IV. Central Method. 


Exp. 20.—Place both lateral apertures out of sight, and use the 
central one only, looking at it with the stop to the right, for some 
seconds. Then push the stop quickly to the left. The first image 
disappears, and a second one takes its place to the left, at a distance 
from the former determined by the degree of divergence which has 
occurred, Hach eye rotates through the same angle, sometimes as 
quickly as the stop is pushed, sometimes not until after an appreciable 
interval. 

This is proved by repeating the experiment with the right luminous 
point out of view to start with in the right blind area, while the 
stop is to the right. This hidden test-point sometimes springs 
into view when the stop is pushed to the left almost simultaneously 
with the appearance of the left image of the central aperture, 
but sometimes not till a moment or two after. With the stop 
to the right, the left eye fixed the central aperture while the 
right deviated. Pushing the stop to the left excludes the left eye 
and lets the right eye see, so that either at that moment or a little 
later both eyes swing through an angle equal to the deviation, to 
let the right eye fix and the left eye deviate. Why, then, does the 
new image of the central aperture not seem to move? Because the 
converging innervation is unaffected by the change, and to it alone 
are all false estimates in the camera primarily due. The only inner- 
vation called into play for the movement in question is the “ rang- 
ing” one, of whose efforts the centres are so well-informed that 
though the eyes move the image does not seem to. So wonderfully 
is the correction made that even in nystagmus, though the eyes con- 
tinually oscillate unknown to the patient, he never, according to 
Helmholtz, sees fixed objects moving. So truly, also, through the 
higher centres does the mind estimate the amount of this effort 
which is in exercise, that artists are said to be able to judge more cor- 
rectly the lateral distance between the two objects by glancing rapidly 
from one to the other, than by any other visual method. 


Exp. 21.—To ascertain how soon the deviation begins. Let a thin 
strip of india-rubber connect one end of the stop with the left side of 
the box, while a piece of string passes from the other end of the stop 
round the forefinger of the right hand. Give the string, to begin with, 
just such a degree of tension as to keep the stop in the middle while 
both eyes look through the camera at the central aperture. Listen 


} 
} 


CONVERGENCE AND ACCOMMODATION OF THE EYES. 503 


to a clock pendulum beating half-seconds. At one end of a beat pull 
the stop instantaneously to the right, and at the other end of the beat 
let it fly back to the left; in so doing it exposes a transitory double 
image of the central aperture, or at least a perceptible widening. This 
shows that divergence has commenced in less than half a second. In 
how much less still remains to be noted, which may be done by short- 
ening the pendulum; but great quickness of observation would be 
needed to attain any degree of accuracy. 


Exp. 22.—Let the central aperture be alone used, and push the stop, 
alternately from left to right and right to left, at definite intervals. If 
the intervals are not too short relative divergence sets in, as shown by 
the apparent displacement of the image to right or left at every move- 
ment of the stop. The vision of the central aperture is alternately 
monocular and binocular, the proportion between the two being deter- 
mined by the rapidity of each movement and the length of the interval 
between them. At certain rates the relative divergence gradually in- 
creases, as shown by the greater and greater apparent displacement 
with each excursion, which proves that a certain proportion of bin- 
ocular vision to monocular is required to overcome the natural tendency 
to divergence. The desire for single vision is, as it were, diluted by 
interruptions. There appears to bea certain rate at which the relative 
divergence neither increases nor diminishes, and a quicker one still 
at which, if divergence is present to begin with, it slowly diminishes, 
and yet a further rate at which it does not appear at all; but a 
mechanical apparatus would have to be used to obtain really 
reliable results. The width of the stop, of course, greatly affects the 
proportion of binocular vision, as also does the length of its sht. If 
the stop were just wide enough to cut off in the mzddle of its course 
the view of the central aperture by both eyes, vision would be wholly 
monocular ; with each diminution of size there would be a larger pro- 
portion of binocular vision. 

It may be that this method would furnish a comparative means of 
estimating the efficiency of the fusion centre, by noting the amount 
and nature of the dilution required to make the desire for fusion in- 
capable of preventing either the occurrence of relative divergence or 
its continuous increase. Two points would have to be considered— 
the frequency of the interruptions, and the /ength of each; the former 
would depend on the nwmber of side to side movements per minute, 
the latter on the rate and length of each movement, the pause at the 
end of each, and the width of the stop. It is possible that a certain 
duration of the two pictures in the brain is necessary to elicit the 
desire to fuse them at all. 

If a wide stop were used, and an up and down movement given to 
it in the slit, binocular vision would be diluted, not by monocular, but 
by intervals of no vision at all. Whether the result would be the 
same I do not know. 


Exp. 23.—TIf, after looking at the central aperture for a little time 
with the stop to the right, the latter then be pushed quickly to the 
middle, the central aperture appears duplicated for a moment by the 


504 MR ERNEST E. MADDOX. 


addition of another image to the left, and the two run quickly and 
with equal velocity into one. The appearance of this second image 
shows, of course, that the eye has deviated. The apparent angular 
separation of the two images is equal to the angle of deviation. If 
they could be kept in their first position (and this we shall see may be 
done), an effort to touch the right one would place the finger 23° to 
the right of the middle line, and an effort to touch the left would show 
it to be likewise 24° to the left of the middle line. We have seen how 
when the stop was at first pushed to the right, and the right eye 
deviated 5°, that the only then existing image of the central aperture 
seen by the left eye appeared to move till it was referred 23° to the 
right of the middle line. The right eye was then excluwled, but now, 
when the stop is put back in the middle, it receives an image on the | 
retina 5° to the right of the macula, and which is therefore referred 5° 
to the left of the image seen by the left eye.t Since the latter image 
appears 24° to the right of the middle line, the former must be 25° to 
the left of it. The left eye remains stationary throughout, while the 
right one (no longer excluded) now returns through the same angle 
through which it deviated. Why, then, if the right eye only is moving, 
and that through an angle of 5° (and this is it easy to verify by testing 
the positions of the b/ind areas), does each aperture appear to traverse 
an equal angle of 23°? It is just the wndoing of what happened when 
the obstructive was pushed to the right. Then the image seen by the 
left eye seemed to move slowly to the right for 25°; now it quichly 
appears to return, because the desire for single vision has aroused the 
supplementary converging effort, which so affects the innervation of 
the left internal rectus that there is no longer any need for that effort 
which usually turns both eyes to the right; it therefore ceases, and 
just as the mind took cognisance of its introduction, and imagined the 
point seen to move to the right, so it takes cognisance of its cessation, 
and refers the point again to its original position. In like manner the 
second image (seen to the left by the right eye) though it traverses 5° 
of the retina, only seems to move 23°, because that is the only moiety 
of the movement which is due to cessation of the mentally-recognised 
ranging effort ; the other half being due to positive converging effort, 
of which no mental account is taken as regards horizontal position. Half 
the movement of the deviated and returning eye is due to relaxation 
of the external rectus, and the other half to increased contraction of 
the internal rectus. The former is taken into account mentally, the 
latter is not. It is quite clear from this that the oculo-motor muscular 
sense is purely central, for the same contraction of a muscle is appre- 
ciated or not according to the central source of the effort. 

Exp. 24.—(a) Place the right lateral aperture, to begin with, 4° or 
5° away from the central one, and the stop to the left. Two images 
are now seen by the right eye, the right lateral and the central one, 
and their relative distance is correctly estimated, though the position 
of both is miscalculated 234° to the left. Now push the stop in the 
middle ; this uncovers the deviated left eye, and reveals another image 
of the central aperture miscalculated 24° to the right, so that it appears 

1 These angles have their apex at the principal dioptric centre. 


CONVERGENCE AND ACCOMMODATION OF THE EYES. 505 


just above the right lateral aperture, which is miscalculated 24° to the 
left, as we saw. But the two images of the central aperture are only 
thus separated for a moment, for they quickly run together, and 
normal fusion takes place. 

(6) If, however, instead of starting with the stop to the /eft, the 
stop be first placed to the right, and then replaced in the middle, the 
images do not run together, though the relative position of the three 
points is exactly the same as before. In the latter trial, the desire for 
false fusion at the near distance is greater than the desire for true 
fusion at the greater one; but why it should not be so in the first 
trial I cannot certainly explain. It may be that the desire for false 
fusion of two objects not in the same vertical line takes longer to 
develop its strength than that for true fusion, and has not time in the 
first trial to do so before the movement for the fusion has begun. 
The effect of attention, as seen in Exp. 26, has probably more to do with 
it. From the construction of the camera, the vision of the right 
lateral aperture cannot be other than monocular, so there must be only 
one image of zt, but the vision of the central aperture, though mon- 
ocular when the stop is either to the left or the right, is binocular 
when it is replaced in the middle, so that, deviation having in the 
meanwhile occurred, there are two images of it. The only difference 
between the two trials lies in the order in which these two images 
appear. In a third modification they may be made to appear simul- 
taneously. 

(c) Push inwards the left brass slide till it just occludes the 
central aperture. Let the right luminous point (now the only one 
visible) be placed as before, and the stop in the middle. On quickly 
drawing out the left brass slide both images of the central aperture 
appear at once, and generally run together, though the result depends 
somewhat on the position of the right lateral point and the amount of 
deviation that has been permitted to occur. This experiment may be 
repeated with many variations by anyone desirous of ascertaining the 
laws of fusion; the left lateral aperture, e.g., may be used instead of 
the right, and each in different positions, or both may be used. 


Exp. 25. To ascertain the effect of attention on the desire for single 
vision. Place the right lateral aperture, coloured blue, 24° to the right 
of the central one, which is covered with a piece of paper or ground 
glass, and therefore white. On looking into the camera with the stop 
in the middle, the two points are seen in their true positions, the 
white one appearing nearly half an inch to the left of the blue one. 
Now push the stop to the right; the two images begin to move slowly 
together till they come to be in the same vertical line, and again sep- 
arate by each pursuing its movement till they have just changed places. 
The white image is now nearly half an inch to the right of the blue 
one. When the stop is replaced in the middle, another white image 
appears nearly half an inch to the left of the blue one, which now has 
a white image on each side of it, and at equal distances from it. I, 
while moving the stop, attention is directed to either of the whzte images, 
they quickly run together, while the blue returns to its original posi- . 
tion. But if the attention is concentrated on the blue point the whole 

VOL. XX. 2K 


506 MR ERNEST E. MADDOX. 


time, the white ones do not run together, but remain as at first, one 
on each side of the blue one. This shows that the mere presence of 
double images, when they are perfectly well defined, does not excite 
the desire for single vision, unless one of them becomes the special 
object of attention. 

If the blue point is not exactly halfway between the two white 
ones, move the brass slide which bears it until it becomes so, and then 
read off its angular position, which, when doubled, will give the 
angular deviation, or relative divergence of the eyes. 

This, then, is the third method of measuring it by the camera, not 
of any clinical value, but useful as showing that the deviation is 
practically the same in extent under such varying tests, for— 


(1) In the “‘blind-spot” method there is one image upon the fovea of 
one eye. 

(2) In the “direct” method there are two images, one upon the 
fovea of each eye. 

(3) In this “central” method one eye receives an image on its 
fovea, and each eye receives an image away from its fovea. 

It would be wearisome to recount more experiments, as the use of 
the camera permits of so many variations. Before passing to the next 
section on “ Distant Vision” it may be well to give a convenient 
summary of the results obtained in near vision. 


1. When one eye is excluded from vision and placed sub- 
Jectively in the dark, it nearly always deviates outwards 
(Exp. 1). 

2. The same deviation occurs if each eye is made to receive an 
image or any number of images, provided that the desire for 
fusion is in abeyance (Exp. 12). 

3. The average angle of deviation appears at present to be 
about 44° with vision for 10 inches. 

4. There are four methods of measuring this angle—three by 
the camera, and one by a double prism modification of Von 
Graefe’s test. 

5. When the record by the “blind-spot ” method differs from 
that of the other three, it is because the mind has learnt to 
estimate the position of each eye separately (Exp. 16). 

6. There are reasons why Von Graefe’s clinical method has 
not revealed physiological divergence. 

7. The divergence begins in less than half a second (Exp. 21), 
and continues gradually at decreasing speed (which may be 
measured at any point) for from half a minute to a minute and 
a half (Exp. 19). 

8. Half the deviation of the excluded eye is due to contrac- 


CONVERGENCE AND ACCOMMODATION OF THE EYES. 507 


tion of its external rectus, the other half to relaxation of the 
internus. 

9. The oculo-motor muscular sense is purely central; the same 
contraction of a muscle is mentally appreciated in one way or 
another according entirely to the central source of the effort (Exp. 
23). 

10. The truth of Hering’s theory that the horizontal move- 
ments of the eyes are governed by two innervations, each acting 
on both eyes as a single organ, is repeatedly demonstrated. 

11. The object fixed by the seeing eye appears during the 
deviation to move in the same direction; the apparent move- 
roent is at half the rate and through half the angle of the real 
movement of the excluded eye (Exp. 3). 

12. An image on the fovea, whatever the real position of the 
eye, is referred to the plane which bisects the angle of converg- 
ence, and which therefore passes through a point midway betwen 
and slightly behind the centres of the two eyes (Exp. 2 and 3). 

13. A fixed object seen by a stationary eye may appear to 
move, and the same fixed object seen by a moving eye may 
appear stationary according to the innervations in play. 

14. The degree of deviation which occurs on exclusion is 
greater in the early morning, often falls after meals, and is sub- 
ject to oscillations, according to conditions which affect the 
nervous system. 

15. A large degree of convergence is still centrally connected 
with the accommodating effort, though its amount differs greatly 
in different persons, being in some more than twice as much as 
in others. 

16. It is probable that these differences account for the fact 
that squint develops in many cases of hypermetropia where 
there is less refractive abnormality than in other cases where 
squint shows no tendency to occur. 

17. The connection between the converging and accommodat- 
ing efforts is still very delicate; the slightest alteration in the 
latter is accompanied by an alteration in the former (Exp. 
5). 

18. The degree of convergence centrally attached to accom- 
modation is subject to slight waverings (Exp. 8). It may toa 
certain extent be either increased or diminished by the desire to 


03 CONVERGENCE AND ACCOMMODATION OF THE EYES. 


unite two images, and to a still greater extent by the desire to 
maintain their fusion (Exp. 13 and 15). 

19. Images at slightly different levels in the two eyes, even 
when the difference in level is great enough to prevent their 
fusion, are often kept near each other by the desire for it. 

20. This tendency decreases rapidly with increasing difference 
in level, and is not perceptible with the images separated by a 
vertical angle of 2° or 3°. 

21. The desire for this false fusion at a near distance may be 
greater than the desire for true fusion at a greater distance, 
though this is affected by the order in which the desires are 
roused (Exp. 24). 

22. When the images are colowred differently the desire for 
fusion is weakened but not altogether removed. 

23. The desire for single vision can be interrupted to any 
required extent by causing alternations of binocular and mon- 
ocular vision, so regulated that, with different rates, deviation may 
be either prevented, retarded, arrested at any part of its course, 
or made slowly to retrogress (Exp. 22). 

24, The effect of attentwn exerts a well-marked influence om 
the desire for fusion (Exp. 25). 

25. The ordinary test of placing a prism (base in or out) 
before one eye estimates simply the degree of relative divergence 
or convergence which is attainable by the desire to rectify the 
diplopia created by the prism, and which is compatible with the 
existing effort of accommodation. 

26. Approaching a finger to the eyes tests the power of mazn- 
taining existing fusion with proportionately increasing accom- 
modation. This is true up to the nearest point of distinct 
vision, within ¢hat it tests the relative convergence attainable 
by the desire to maintain fusion complicated with inereasing 
indistinctness of the images. 


(To be continued.) ! 


' The writer will be greatly obliged for the pointing out of any omissions and 
errors detected in this paper.—Address, E. E. Maddox, M.B., Shipton, Chipping 
Norton, Oxon. 


A CONTRIBUTION TO SPLENIC HISTOLOGY. By 
Ropert Rosertson, M.D. (Edin.), Assistant Physician to 
the Ventnor Consumption Hospital. (PLATE XV.) 


(THE following observations were made in 1879 while working 
under Dr D. J. Hamilton in the Pathological Laboratory of 
Edinburgh University. Circumstances have prevented a re- 
sumption of the work hitherto, hence the incomplete form of 
the present paper.) 

Controversy in splenic histology of late has mainly centred 
about the minute structure of the splenic pulp, and the manner 
in which the circulation is carried on therein. It cannot yet be 
said that the channels by which blood reaches the veins from 
the arteries have been determined beyond dispute. W. Miiller? 
is the representative of those who maintain that the capillary 
plexus, which in other organs intervenes between arteries and 
veins, is in the spleen replaced by an adenoid reticulum through 
which the blood filters in its passage to the veins, the arteries 
ending by their endothelial lining becoming continuous with the 
cells of the reticulum, while in the veins the fibres of the 
reticulum gradually approximate until the vessel wall and its 
endothelial lining are again complete. Others again, while 
agreeing with this view as to the absence of a continuous system 
of vessels, differ in details,—Billroth,? for instance, holding that 
the arteries terminate by open mouths in the reticulum of the 
tissue which intervenes between the “capillary veins;” Klein,’ 
more recently, denying the presence of a reticulum of fibres, 
which he says is an appearance due to section of the septa of a 
honeycomb of membranes. 

On the other hand, Kolliker* and Grohe® maintain that the 


1 Stricker’s ‘‘ Human and Comparative Histology,” New Syd. Soc., 1870, vol. i. 
p. 357. 

2 «Zur normalen und pathologischen Anatomie der Menschlichen Milz,” Virch. 
Arch., vol. xx. p. 413. 

3 “ Observations on the Structure of the Spleen,” Quart. Jour. Micr. Sci., vol. 
xy. p. 368. 

4 Manual of Histology, p. 369, London, 1860; New Syd. Soc., p. 451. 

> “* Zur Geschichte der Melanémie nebst Bemerkungen iiber den normalen Bau 
der Milz und Lymphdriisen,” Virch. Arch., 1861, vol. xx. p. 325 et seq. 


510 DR ROBERT ROBERTSON. 


circulation in the spleen is carried on in a precisely similar 


manner to that of all other organs of the body; that is to say, 


there is a continuous system of vessels out of which, except by 
rupture, the blood never passes; that the arteries break up into 
capillaries, which are again continuous with the minute veins. 
But while Kélliker thinks it probable that an endothelial lining, 
too delicate to be demonstrable in the finest vessels, is con- 
tinuous throughout, Grohe thinks that a layer of fusiform cells 
is the only lining membrane. 

The uncertainty which these opposing views excites is in- 
creased by the difficulty of explaining completely by either 
certain appearances found in the pathological condition known 
as “diffuse waxy spleen.” Stained with methylaniline iodine, 
a section of such a spleen shows, even under a low power-of the 
microscope, that the organ consists in great part of an infinite 
number of canals and their intervening septa. Sections of these 
canals, in all directions, transverse, longitudinal, and oblique, are 
recognisable, bounded by limiting lines of rose-stained “ waxy” 
material, and if the disease is in an early stage between two 
adjacent rosy lines, a blue portion of the septum forming a 
narrow strip of apparently retiform tissue can easily be found. 
Are these canals with their intervening septa adventitious ap- 
pearances due to the morbid condition, and if not, to what do 
they correspond in the normal organ ? 

To determine this question, the spleen of a healthy young 
adult was examined. The organ was markedly “congested,” 
but in no other respect peculiar. Hardened in dilute chromic 
acid and stained with logwood, sections were half cleared up in 
clove oil, and then mounted in dammar. By this method the 
canals or vessels were demonstrated which had already been 
seen mapped out by morbid material in “waxy disease,” but 
without “half-clearing ” in clove oil the canals could not be recog- 
nised. In the septa and walls of these canals cells were seen, 
which, when isolated, had large rounded or more or less ovoid 
nuclei, surrounded by a small quantity of matrix substance, and 
projecting on the inner border of the narrow band or fibre into 
which the matrix substance was prolonged on either side of the 
nucleus. The general outline of the inner border, except where 
the nucleus bulged, was concave; the opposite border was con- 


CONTRIBUTION TO SPLENIC HISTOLOGY. 511 


vex and wavy, refracting the light; the ends of the fibre were 
occasionally found curled towards the inner border, not unlike 
what is seen in fibres of yellow elastic tissue. In the section 
these cells were seen arranged side by side in the long axis of 
one of the “canals” referred to, so as to form an apparently con- 
tinuous membrane, and transverse section may thus show a row 
of “leucocytes” projecting towards the lumen of the canal. 

In the septa the coloured blood-corpuscles, often in linear 
arrangement, and nucleated cells and fibres gave the appearance 
of an areolar tissue (such, indeed, Billroth regarded it, though 
he ultimately abandoned the view that the capillaries end in it 
by open mouths, finding that they pierced the canal walls. 

There exists then in the splenic pulp an abundant canal 
system, the canals being of considerable size, and separated by 
septa usually relatively small and of loose fibro-cellular structure. 
The walls of these canals consisting largely, so far as we have 
yet seen, of peculiar fusiform cells with nuclzar body projecting 
towards the inner aspect of the cell. 

To ascertain the relation of this canal system to the arteries, 
various methods of injection from the artery were tried with 
spleens of pigs and sheep. It was found very easy with fluid 
injection to drive the liquid freely out through the vein, but on 
preparation of sections of these spleens, the tissue was seen to 
be very unequally permeated by the injection fluid; and in those 
parts where the injection had entered most freely the appearance 
was that of a general infiltration of the tissue with the injection ; 
no arterial branches except those of larger size could be made 
out. 

On injecting, however, by the vein with an open cannula in 
the artery it was discovered that after sixteen ounces of the 
liquid had been thrown into the spleen of a sheep not a drop 
came out through the artery. The cannula in the vein was 
then closed, and further injection attempted from the artery, 
but only a very small quantity could be forced in, though in the 
attempt the cannula was driven off the syringe. Additional 
injection through the vein was then tried, and with ease another 
8 oz. of fluid was thrown into the organ before the capsule 
ruptured. 

Double injection was also tried, blue gelatine mass being 


512 DR ROBERT ROBERTSON. 


thrown into the veins, and when this was cold a carmine in- 
jection mass into the arteries, The special object in view was to 
inject the capillary plexus of the Malpighian bodies from the 
artery ; and on naked eye examination of a section of the 
injected organ it seemed probable that this had been accom- 
plished successfully, for the Malpighian bodies were conspicuous 
by their carmine colour in ap area of dark blue. On microscopic 
examination, however, no injection of the Malpighian capillaries 
could anywhere be found, but everywhere a zone of carmine 
surrounding the uncoloured Malpighian body, and separating it 
from the dark blue of the injected pulp-substance. 

Injection with carmine mass of a human spleen in an early 
stage of waxy disease was tried also. The injection was thrown 
into the artery only, and after hardening, sections were stained 
with methylaniline, and mounted. It was hoped that, the canals 
being mapped out by the morbid material, the relation of the 
injected arterioles to them might be more easily discovered. This 
hope was to a certain extent realised on examination of sections 
microscopically. It was found that minute injected vessels ran 
in the septa between the canals, and such a vessel was seen 
after running for some considerable distance in the middle of a 
septum bifurcating, and the two branches diverging approached 
the walls of canals, and one partly encircling a canal in trans- 
verse section was lost sight of just when apparently about to 
pierce the canal wall. 

Injection of sheep’s spleens with } per cent. nitrate of silver 
solution was also tried, and in this connection it may be 
mentioned that in injecting from the artery a zone of pigment- 
ation was obtained around the Malpighian bodies, correspond- 
ing to the carmine zone in the double injections described 
above. 

Thus we have seen that the splenic arteries do break up 
into minute or capillary vessels which run for a considerable 
distance in the septa that intervene between the canals already 
described, and sooner or later they approach closely to the 
walls of these canals, and probably pierce them. In the 
Malpighian bodies it would seem that the surrounding pres- 
sure empties the capillaries of their contents, the injection 
appearing in the portions of the canal system with which the 


CONTRIBUTION TO SPLENIC HISTOLOGY. 513 


Malpighian body is related, and with which its vessels apparently 
communicate. 

We have also seen that some valvular arrangement exists, 
which, while it permits the blood or injection fluid to flow 
freely from artery to vein, prevents a return current from vein 
to artery. What this arrangement probably is we shall presently 
see ; in the meantime it is necessary to consider the endothelial 
lining of the arterial capillaries and of the canal system. 

A fresh sheep’s spleen was injected from the artery with 
4 per cent. solution of nitrate of silver, and portions of the 
silvered tissue afterwards carefully teased out. In such speci- 
mens the endothelial lining of the arterial capillary was well 
demonstrated, and in a few specimens it could be traced until 
an infundibular expansion of the vessel took place to twice or 
three times its previous size. Nuclear bodies lying in the long 
axis of the capillary vessel outside the endothelium were found 
to belong to fusiform cells like those already described in the 
walls of the canals, and where a capillary vessel had been 
broken across, the curled end of one of these cells might be 
found projecting beyond the endothelium. The difficulty of 
demonstrating the continuity of the endothelium of the arterial 
capillaries with that of the canal system was due to the exces- 
sive delicacy of the walls of the canals, which, during the 
preparation by teasing, generally became entirely broken up, 
and to the elasticity of the spleen in lower animals, which 
prevented the ready demonstration of the canal system in 
sections of silvered spleens. Fortunately the spleen of a 
healthy human adult was obtained and injected five hours after 
death, and silvering of the endothelium by injection of the artery 
with nitrate of silver solution was successfully accomplished. 

The tissue was hardened in dilute chromic acid and in 
spirit, and preparations half cleared up in clove oil, and 
mounted in dammar, were examined. The tissue was also 
examined by teasing in the fresh state after exposure to light, 
It was found that the arterial capillaries were lined by an 
endothelium, as already described in the sheep’s spleen, but 
that unlike their infundibular expansion in the sheep’s spleen, 
the vessel ended in an abrupt expansion (or flange), producing 
an appearance like the end of a candlestick. 


514 DR ROBERT ROBERTSON. 


On examination of hardened sections, stained with logwood, 
and mounted as described, the canals were found to be lined 
with an exceedingly delicate endothelial layer of cells, the out- 
lines of the cells being faintly silvered, their shape long and 
narrow, and arranged generally across the direction of the canal, 
while outside this endothelium the fusiform cells, with their 
nuclei stained blue, were arranged in the long axis of the 
vessel. 

Under a power of 1000 diameters, circular openings of small 
vessels could be seen with a clearly maintained continuity of 
endothelium. In some places the opening of the small vessels 
into the canals was conical to some extent, but this did not 
appear to be commonly the case. 

Having thus traced the continuity of artery and venous 
canal, and shown that a continuous and unbroken endothelial 
lining does exist, we can now understand by what arrangement 
regurgitation from veins to arteries is prevented. It has been 
mentioned that the septa in which the arterial capillaries run 
are relatively small as compared with the size of the canals, 
and that they present a loose fibro-cellular texture. By dis- 
tending the venous canals with blood or other fiuid, these septa 
are pressed upon, the lumen of the capillary vessels becomes 
obliterated, and the liquid can neither flow back through them, 
nor can any pass from the arterial side, except by the use 
of very high pressure. The relative size of the two sets of 
vessels sufficiently explains how when no more fluid could be 
driven into the arterial system in the experiment described, a 
third more than the organ then contained was easily injected by 
the vein, and the organ ruptured. 

Malpighian Corpuscles—These bodies have been incidentally 
referred to, as mapped out by an injected zone, in silvered 
human and sheep’s spleens, and in double injection of the 
spleen. This injected zone was in all cases sharply defined 
towards the Malpighian body in the condensed stroma of the 
pulp, but passed gradually away into the surrounding pulp- 
substance. 

In the injected spleen of a cat the plexus of capillaries in the 
Malpighian body was well seen, and it was observed that they 
formed loops towards the periphery, from which branches passed 


CONTRIBUTION TO SPLENIC HISTOLOGY. 515 


into the pulp-substance beyond. This zone of injection, there- 
fore, is no doubt the fluid which has passed into the Malpighian 
capillaries, and, by surrounding compression, has been forced 
into the canal system of the pulp, with which the capillaries 
communicate. 

In “half-cleared” sections, stained in logwood, the con- 
tinuity of the stroma of the Malpighian body with the tissue 
of the septa of the pulp-substance was recognised. 

So that the Malpighian bodies are, as it were, localised over- 
growths of the tissues of the septa which separate the venous 
canals, cells and fibres multiplying, and the arterial capillaries, 
instead of bifurcating, breaking up into a plexus; but the 
stroma and capillaries maintain their continuity with the 
adjacent septa and canals, only, from compression by the over- 
growth, these latter are condensed and form what was thought 
formerly to be the capsule of the Malpighian bodies. 


EXPLANATION OF PLATE XV. 


Fig. 1.—Endothelium of funnel-shaped junction of arterial capil- 
lary (a) and venous canal (b), from sheep’s spleen (silvered). (x 450.) 

Fig. 2.—Spleen of woman injected with nitrate of silver five hours 
after death, showing expansion of arterial capillary (a) into a venous 
canal (b); the outlining of the endothelial plates is well seen. 
{x 450.) 

Fig. 3.—Small portion of tissue from same spleen, x 800; a, 
venous canal, with silvered marking ef endothelium; 2%, arterial 
capillary ; c, section of vessel in communication with both a and D. 

Fig. 4.—Arterial termination (waxy spleen x 450); a, mouth of 
minute arteriole; (b’, b”, b'"), venous canals; c, nuclei of fibre cells ; 
e, waxy infiltration of wall of eanal. 


SUPERNUMERARY LEG IN A MALE FROG (Rana 
palustris). By FREDERICK TUCKERMAN, M.D., Amherst, 
Massachusetts. (PLATE XVI.) 


I rirst wish to thank Professor Warner of the Massachusetts 
Agricultural Collegé, to whom the frog belongs, for his kindness 
in placing it at my disposal. 

The remarkable abnormality about to be described occurred 
in a frog which was blown, while blasting with gunpowder, out 
of a crevice in a ledge of mica schist 12 feet below the surface. 
[t was immediately picked up, and, apart from being stunned, 
had apparently sustained no injury. 

The same explosion also brought to the surface five other 
frogs, of different ages and sizes, none of which were killed or 
injured. 

At the time of its unexpected appearance the frog was 
supposed to be about a month old. From this time until it was 
killed, two months later, it was not observed to eat anything. 

The external opening of the crevice in the ledge measured 
only a few lines at its widest point, and flowing into it was a 
small stream of water, which undoubtedly conveyed either the 
egos or the frogs in the larval stage to the interior of the rock, 
as the breach in the ledge was much too small even to admit 
the passage of a very young frog. 

The pigmentation of the skin covering the extra limb corre- 
sponded in form and colour with that of the rest of the body, 
with the exception that the dark Lrown bands or spots towards the 
distal end of the leg completely encircled it. It was attached to 
the trunk slightly to the left of the mesial line, and to the left of 
the posterior end of the tip of the urostyle, just above the open- 
ing of the cloaca. ‘he first part, about a line in length, seemed 
to be chiefly tegumentary in character, and was quite narrow. 
This was followed by an enlargement 24 lines in length, and a 
little less in width, chiefly in a ventral direction. Then came a 
second constriction greater than the first, which immediately 
widened into a large oval-shaped dilatation or sac, ? of an inch 
in Jength and 3 of an inch in width, the walls of which were 


SUPERNUMERARY LEG IN A MALE FROG, 517 


not at all tense, and offered less resistance to pressure than was 
the case in the first enlargement. The space within was par- 
tially filled with fluid. It was quite evident, however, that it 
contained something else beside fluid, for, when rested upon 
one end, it not only kept its oval form perfectly, but could not 
be possibly depressed in the direction of its long axis. The 
remainder of the limb to the distal tarsals was straight, and 
measured 4 an inch in length and } of an inch in width. It 
was jointed above and below. The superior articulation ad- 
mitted of but slight movement, the inferior was quite movable. 
There were two digits present, the inner one the longer, and a 
rudiment of a third. The usual position of the toes was that of 
adduction. 

The limb measured, from the point of its attachment to the 
trunk to the tip of the last phalanx of the inner digit, 24 inches. 

The circulation of the blood in the web between the two 
digits, as seen with the aid of the microscope, presented nothing 
unusual in appearance. 

When stimulation was applied to the distal end of the limb, 
or to any part of it, the frog would usually draw itself, or jump 
quickly away, dragging the extra limb after it. Sometimes, 
however, often-repeated stimulation would call forth a slight 
muscular contraction in the limb itself, and this was occasionally 
seen to take place when the animal was left entirely to itself. 
The distal end of the limb would then be abducted a little to 
the right, the toes would partially separate, and at the same 
time a slight drawing of the whole limb toward the trunk would 
be perceptible. 

The following appearances were observed on opening the 
limb :— 

The narrow portion close to the point of attachment to the 
trunk contained a few tendons, blood-vessels, and a nerve. The 
tendons took their origin from the symphysis pubis. The 
blood-vessels could not be made out with any degree of certainty, 
with the single exception of a vein which appeared to join the 
left pelvic vein. The nerve was apparently a division or branch 
of the sciatic. In tracing it down the limb no division of it 
above the knee could be discovered, nor were any lateral branches 
made out. The space in the first enlargement was almost 


518 DR. FREDERICK TUCKERMAN. 


completely filled by a mass of muscle to which the tendons 
belonged, and in which the vessels ramified. Neither bone 
(femur) nor cartilage were present at this point. The second 
constricted portion, smaller and narrower than the first, con- 
tained the tendons, vessels, and nerve, which it conveyed to the 
parts beyond. Within the large oval-shaped sac was a bundle 
of muscles running the entire length of it. There were no 
adhesions uniting the skin to the subjacent parts except at the 
ends. Two-thirds of the space not occupied by muscles was 
filled with a yellowish-red fluid, having a slightly acid reaction, 
and quite rich in red and colourless blood-corpuscles. The 
muscles were somewhat smaller than usual, and a few were 
wanting; but otherwise they presented no special variations 
from those found in this region. The gastrocnemius, tibialis 
posticus, peroneus, and tibialis anticus were easily distinguished. 
On cutting away the muscles a bone (crus) was brought into 
view. The shaft was long, cylindrical, and very slender. No 
groove was detected indicating the line of union between the 
fibula and tibia. The articular expansion at the inferior extre- 
mity was very large and out of proportion to the shaft above it. 
This formed a movable joint with the articular process of the 
bone below it. At the superior end of the shaft was an articular 
process, quite small, which had only a tendinous connection with 
the region above it. In the proximal tarsal region the skin 
adhered quite firmly to the parts below it. The muscles were 
much atrophied, and the vessels and nerves could not be clearly 
followed. In place of the usual two elongated bones (astragalus 
and caleanewm), connected above and below by epiphyses, 
there was a single long bone, unmarked by any longitudinal 
groove, articulating slightly with the ankylosed (?) tibia and 
fibula. 

At the distal end of the bone no extra ossicles were found. 
The right digit was composed of a metatarsal bone and two 
phalanges. The inner digit (would be the fourth counting from 
the allux), the longer, consisted of a metatarsal bone and three 
phalanges. The rudiment corresponded to the middle digit. 

From this description it will be seen that the foot belonged 
to the right side of the body, although the limb sprang from the 
left side. 


SUPERNUMERARY LEG IN A MALE FROG. 519 


With the exception of the accessory limb the frog appeared 
to be in a perfectly normal condition. 


EXPLANATION OF PLATE XVI. 


It was found necessary, in order to show the rudimentary digit 
well, to turn the pes over, thus bringing the dorsal surface into view. 

1, Enlargement filled with muscles corresponding to the position of 
the femur. 2. Sac partly filled with fluid containing muscles and 
crus. 3. Proximal tarsal region composed of a single long bone. 
4. Rudiment of the middle digit. 


[Editorial Note——The malformation in this very remarkable frog 
will, without doubt, recall to some of our readers the case of Jean 
Battista dos Santos, a native of Portugal, who visited this country in 
1865, being then in his nineteenth year, and whose case was described 
in the Lancet of July 29, and subsequently by the late Dr P. D. Han- 
dyside, in the Edinburgh Medical Journal, March 1866. In Dos 
Santos’s case of imperfect double monstrosity there were two complete 
sets of external organs of generation in addition to the intermediate 
lower limb. A case of a somewhat similar kind, in a female foetus, 
but with two lower limbs, was described by Von Baer, in Mem. de 
VAcad. de St. Petersb. VI. Series, and his figures are reproduced in 
Forster's Missbildungen des Menschen, Jena, 1865. The writer of this 
note had also the opportunity of seeing, in February 1885, through 
the courtesy of Mr R. Urquhart, Lecturer on Pharmacy, Edinburgh, 
a new-born child with a similar deformity. The child lived a few 
days, but no dissection of it was permitted by the parents.—W. 7’.] 


THE NATURE OF THE RELATIONSHIP BETWEEN 
UREA FORMATION AND BILE SECRETION. By 
D. Noki-Paton, M.D., B.Sc., F.RS.E., Biological Fellow of 
the University of Edinburgh. 


(From the Physiological Laboratory of the University of Edinburgh.) 


IN a previous paper, published in the last numbers of this 
Journal, vol. xx pp. 114 and 267, I endeavoured to demonstrate 
that a direct relationship existed between urea formation and 
bile secretion. It is now necessary to inquire into the manner 
in which these two processes are thus connected. In investi- 
gating this subject we must remember that the bile constituents 
and urea are produced in the liver, and that our attention 
therefore must be directed to changes occurring in this gland. 

The chief functions of the liver are the formation of glycogen 
and the secretion of bile, and the latter of these functions may 
be divided into two elements, the production of bile acids and 
the elimination of the effete blood-pigment as bilirubin. I shall 
endeavour to show that urea formation is connected with the 
latter of these, the excretion of effete haemoglobin. 

That blood-pigment is converted into bilirubin and excreted 
by the liver as such is clearly demonstrated by the work of 
Frerichs, of Stideler, of Kiihne, of M. Herman, and others. 

Tarchanoff was perhaps the first (Pfliiger’s Arch., Bd. ix. pp. 53 
and 187) to connect this production of bilirubin with an increased 
flow of bile. After injecting 100 cc. of a dilute solution of 
hemoglobin into the veins of a dog with a biliary fistula, he 
observed an immediate and enormous rise in the amount of 
bile and in the quantity of bile pigments excreted. A similar 
rise was observed after the injection of distilled water, which set 
free heemoglobin by dissolving the blood-corpuscles. He observes 
that, while the bile pigments were so greatly increased, the bile 
acids were diminished in amount. 

Stadelmann (Arch. f. Exp. Pathol., Bd. xv. p. 337) records a 


1 Towards the expenses of this research, a grant was made by the British 
Medical Association on the recommendation of the Scientific Grant Committee of 
the Association. 


UREA FORMATION AND BILE SECRETION. 521 


similar series of experiments in which he injected hemoglobin 
in much larger quantities, and in mugh more concentrated solu- 
tion—20 to 40 grammes in a 20 percent. solution. He observed 
that the bile first became thick and viscid, and that its rate of 
excretion diminished, the percentage amount of bile pigments 
beiug enormously increased. This initial fall was followed in a 
few hours by a great increase. Obviously the results of these 
experiments are to be explained by the large quantity of heemo- 
globin used, and the concentration of the solution tn which 
it was injected. By the increase tn the solid constituents of the 
bile its passage along the bile ducts was markedly impeded, and 
it was only upon a more fluid bile being produced that it eseaped 
freely and allowed the increased secretion to manifest itself. 

In Tarchanoff’s experiments, on the other hand, the hemo- 
globin reached the liver in a dilute solution and in moderate 
amount, so that no blocking of the bile passages by the viscid 
bile occurred. 

Such methods of injecting hemoglobin in solution are by no 
means satisfactory, since the altered condition of the specitic 
gravity of the blood and the changes in the circulation possibly 
induced, tends to vitiate the results obtained. 

Employing the method of injecting into the blood some power- 
ful hemolytic, such as toluylendiamin, Afanassiew (Zésch. f. Clin. 
Med., Bd. vi. H. 4) has shown that the bile secretion is greatly 
increased. And ina paper, published in Pfliiges’s Archiv., Bd. 
xxx., he says:—“ Da nach Ergebnissen der Versuche mit Toluylen- 
diamin die Gallenbildung durch massenhaften Zerfall der rothen 
Blutkorperchen gesteigert wird—tritt den Gedanken nahe, dass, 
unter normalen Verhaltnissen, der Zerfall rothen. Blutkérper- 
chen, selbstverstindlich innerhalb bestimmten physiologischen 
Grenzen, ein Anregung zur Gallen-secretion gebe.” 

Paschkis (Wiener Med. Jahrb., 1884, 8. 295) has further 
shown that these well-known blood-corpuscle-destroying agents, 
the salts of the bile acids, are exceeding powerful cholagogues. 

All evidence on the subject, therefore, goes to show that the 
destruction of blood-corpuscles, and the consequent setting free 
of hemoglobin in the blood, does lead to a great increase in the 
secretion of bile. 

That urea formation also is connected with destruction of 

VOL, XX. . 21 


522 DR D, NOEL-PATON. 


blood-corpuscles was first suggested by Fiihrer and H. Ludwig 
(Vierordts, Arch., Bd. xiv. 8. 307) while Addison in 1864 (Brit. 
Med. Jour., vol. i. p. 202), reasoning from different data, came 
to the same conclusion. The arguments of these authors are 
full of fallacies, and the facts upon which their theory is founded 
are capable of various different interpretation. 

More recently Meissner, in a paper entitled “ Ein Beitrage zur 
kenntniss des Stoffwechsels im thierischen organismus (Zésch. 
J. rat. Med., Bd. xxxi. 8. 234), adduces very strong evidence 
for the production of a great part of the urea in the liver from 
blood-pigment. It is impossible to discuss fully this long and 
ably reasoned paper, but one or two of his arguments must be 
mentioned here. | 

No tissue of the body functionates so long as the blood,—to the 
last moment of life. The urea excretion during starvation also 
keeps up in like proportion. Now, fresh corpuscles must be con- 
stantiy manufactured up to the last moment of life, and when 
food no longer supplies the material the muscles and fat are then 
used. That during the period of inanition blood-corpuscles are 
being destroyed is shown by the fact (Bidder and Schmidt) that a 
not inconsiderable amount of bile is formed up to the time of 
death, and it is no longer doubtful that the biliary colouring matter 
are derived from the hemoglobin. On the other hand, the blood- 
corpuscles do not diminish much during starvation. Since Voit 
found that in a cat the blood lost only 4:8 grms. of fixed sub- 
stances, a much smaller loss than is sustained by any other 
tissue in the body. Panum also found a relatively small 
reduction of the quantity of blood, and no diminution in the 
corpuscles during starvation. The lymph stream also persist 
till death, and in the lymph system blood-corpuscles are formed. 
The chief argument for his theory of the formation of urea he 
considers to be its appearance in the liver, and the relation of 
the formation and destruction of blood-corpuscles to the liver— 
the former very doubtful—the latter certain, the liver being 
the only organ of the body where a copious destruction of the 
blood-corpuscles can be detected. The fact which before all 
other supports this destruction in the liver, is the separation of 
bile pigments which are derived from the hemoglobin (see. 
Funke’s Lehrbuch der Physiol, 4 Autl. Bd. I. 8S. 262). David has 
shown that it is undoubted that destruction of blood-corpuscles 


UREA FORMATION AND BILE SECRETION, 523 


occurs in the liver (Hin Beitriige zur Frage iiber die Gerinnung 
des Lebervenenblutes, Dorpat, 1866). He entirely denies their 
formation in the liver. 

Meissner’s arguments are of no small value, because they are 
founded on well-established facts, such as the connection of the 
bile secretion with blood destruction on the one hand, and with 
urea excretion on the other; and on the formation in the liver 
of both bile and urea. 

That heemocytes really are broken down in the liver has 
been, since the appearance of David’s paper, definitely shown 
by Nicolaides (Arch. de Physiologie, T. x. p. 581, 1882), who 
found a very marked diminution in the number of corpuscles in 
the portal vein, a diminution which, during active digestion, was 
frequently as great as from 1,000,000 to 2,000,000. In four 
observations made on fasting animals this difference was found 
to be much smaller. 

In investigating this connection between urea formation and 
excretion of hemoglobin as bilirubin by the liver two methods 
are open tous. Wemay either directly inject heemoglobin and 
study its effect on the urea excretion, or we may destroy blood- 
corpuscles within the body of the animal by the administration 
of drugs which have outside the body been proved to have 
hemolytic action. 

In order to avoid the infliction of pain, I have adopted the 
latter method. 

For the purpose of breaking down corpuscles I have employed 
pyrogallic acid, and the salts of the bile acids, the direct influence 
of which on heemocytes has been for long well known. 

In these experiments healthy dogs in a state of nitrogenous 
balance (stickstoffgleichgewicht) were used, and a detailed ac- 
count of the method of observation will be found in the Journal 
of Anat. and Physiol., vol. xx. p. 268. 

Exp. I. 

A healthy setter bitch, weighing 14°96 kilos., had been kept upon a 
diet of porridge and milk till the urea excretion became constant. 


Pyrogallic acid was then administered by the mouth. Its influence on 
the urea excretion is shown in the accompanying chart and in fig. 1. 


524 DR D. NOEL-PATON, 


PAM AD Ure! So i, i Oe eee 
: uantity of | age 
| ed . Tanne in Sp. G. tg Remarks. 
ah c. CS. ae 
| 1. |(765-6-734} 1011 | 7:404 | Weight of dog—14°96 kilos. 
2 ( 765-—6°734} 1010 6064 | Diet—Oatmeal, 113 grms. 
3 885 1009 7°368 Milk, . 320 c.ces. 
4 855 1009 7°463 
5 810 1009 | 6°966 il grm. pyrogalic acid. 
6 975 1012 13°182 1=5 grm._,, § 
7 780 1009 7°956 
8 | 835 1009 7°480 
| 9 1 835 1009 7480 


Fig. 


Urea in 
gms. 


11 


10 


Exp. I.—Influence of pyrogallie acid on excretion of urea. 
1 grm. given at a, and 1°5 grm. at b. 


Exp. Il. 

In the second experiment I employed the mixed salts of the bile 
acids—the ordinary “ crystalline bile” derived from ox bile. This 
was given by the mouth, and hence only acted when exhibited im 
large doses, since, as we know, much of these acids is decomposed in 
the intestines, and therefore a small part is absorbed unchanged, and 
ina form capable of destroying corpuscles. 

The same dog was used, and the conditions of the experiment are 
precisely similar to those in Exp. I. 


UREA FORMATION AND BILE SECRETION, 525 


| Day. |U rine in ees, Sp. G. ioe, Remarks. 
1 650 1019 | 67500 Weight of dog=14°96 kilos. 
2 750 1009 | 67600 Diet as in previous Exp. 
3 660 1009 | 6°468 2 germs. crystalline bile. 
4 580 1010 | 5°742 2°5 ,, ‘ " 
5 600 1014 | 8640 gece * . 
6 380 G15”) joke. 
considerable 
| quantity lost 
ae 860 1008 6°656 
8 690 1011 7°404 
9 840 1010 | 6°064 | | 
Fig. 2. 
Urea in 
grms. 


8 


Exp. I1.—Influence of salts of bile acid on urea excretion. 
2 grm. given at a, 2°5 grm. at 6, and 6 grm. atc. 

These experiments show that the action of these hemocytic 
agents is accompanied by an increased production of urea. 

But more than this is required to establish the fact that this 
increase is solely and purely due to the destruction of blooid- 
corpuscles. We must be able to show a definite relationship 
between the number of corpuscles destroyed, zc. the amount of 
hemoglobin to be excreted, and the urea produced. I have 
accordingly undertaken the following experiments, in which the 
urea was daily estimated and the hemocytes daily enumerated 
before and after the administration of some hzmocytic agent, in 
order that a calculation might be made of the increase in the 
production of urea on the one hand, and the amount of hiemo- 
elobin liberated on the other. 

From such a method of research only approximate results can 
be expected, since, in the first place, the only available method of 
enumerating the hemocyte, by Hayem’s or Gower’s hemocyto- 
meter, is not absolutely correct. In addition to this, for such a 


26 DR D. NOEL-PATON, 


or 


research as the present, where the object is to ascertain the total 


number of corpuscles destroyed, the disturbing element increased - 


production, which in all probability follows increased destruction 
of corpuscles, just as it follows any loss of blood must tend to 
vitiate results. During the first day or two of any increased 
destruction, this increased production will make itself less felt 
than after the economy has had time to attempt to re-establish 
the normal condition. 

Another source of fallacy, which must be borne in mind, is 
the difficulty of calculating the amount of hemoglobin in rela- 
tionship to the weight of the dog. In estimating the amount of 
blood, I have followed Bischoff and Welcker, and in regard to 
percentage of hemoglobin present I have accepted Preyer’s re- 
sults (Die Blutkrystalle). 

Two hypothetical formule have been suggested to repre- 
sent the disintegration of the hemoglobin molecule. That of 
Zuelzer (Untersuch wi. die Seimdelogie des Harns, Berlin, 1884) 
need not here be considered, since it implies a great production 
of bile acids, which, according to the observations of Tarchanoff 
and of Stadelman, does not occur. Charles (Brit. Med. Jour., 
1885, vol. i. p. 820) suggests the following decomposition of two 
molecules of hemoglobin :— 


Cyooo Hyoo9 Ngog Fes Sg Ogs¢-+ 501 0+ 182H,0 + 6H,SO, - Fe, O5. 


= (eh H. Ni O. 
2 Bilirubin, . ; j 64 72 8 19 
150 Urea, ‘ : j 150 600 300 150 
32 Glycogen, . . ; 960 1600 — 800 
26 Carbonic Acid, : : 26 — — 52 
1200 2272 308 1014 


Tt is interesting to observe that in both these hypothetical 
decompositions a process of oxidation is supposed to occur, a fact 
which fully agrees with Ehrlich’s recent experiments in regard 
to the reduction of methyl blue in the liver as well as in the 
lunes (Cot. f. Med. Wissensch., Feb. 21, 1885). 


Ge oF . 54°00 Fe, 0:42 
ne ba: Give . 0°68 
Ni . 16-25 Qian os . 21°45 


Considering the percentage composition of hemoglobin from 
these data we may calculate the initial amount of hemoglobin in 
the blood of an animal, and the amount set free by the disinte- 


UREA FORMATION AND BILE SECRETION. 527 


gration of corpuscles, we see that 200 grms. of this substance 
corresponds to 32°5 germs. of nitrogen, and from the above 
formule it appears that about 4 erms. must go to the formation 
of bilirubin, leaving 28 grms. to supply the nitrogen of urea. 
So 200 grms. of hemoglobin may thus yield 60 grms. of urea,— 
the percentage of nitrogen in urea being 46°6,—that is, 1 grm. 
of urea will represent 3°3 grms. of hemoglobin. 


Exp. I1.—Pyrroeariic Aci. 
In this experiment a large black retriever, weighing 15°43 kilos., was 
used, The diet and details of the experiment are given in the accom- 
panying table and figure :— 


. 
Day of | Urinein | Urea in Hemocytes Weight iat ste 
| Exp. c.cs. erms, Be Ge hin kilos. Remarks. 
5 of blood. } 
eS ee eS SS #* — 
1 { 500 5800 | Diet.—Oatmeal, 169°8 
2 (500 | 5800 erms. as porridge. 
3 ( 565 5998 | Milk, 320 c.es. 
4 | 565 5998 
5 400 5°450 
6 \ 455 5643 15°42 
7 ( 455 5°643 | 7,460,000 
f 2 erms. of pyrogallic acid 
8 600 9°483 6,615,000 | urine dark in colour— 
| no albumen. 
9 | 485 | 6617 | 7,275,000 | Urine still dark. 
10° | (475 | 6-220 
11. | (475 | 6-220 | 7,270,000 | 15-42 
Urea in Hemocytes, 


erms. per ¢.min. 
9 
S 8,000,000 
7 7,000,000 
b 6,000,000 


Exp. III.—Influence of pyrogallic.acid on urea excretion and the 
number of hemocytes. 2 grms, administered at a. 


Before the administration of the drug the corpuscles were 7,460,000 
per c.mm., but at the end of the second day, after the exhibition of 


~ 


528 DR D. NOEL-PATON. 


pyrogallic acid, they fell to 6,615,000. Now, the blood of a dog is 


about 51, of its weight, therefore in this case the dog had 1186°3 grms. . 


of blood. The hemoglobin, according to Preyer, is in the dog 13:8 
per cent. of the weight of the blood—in this case probably a little less, 
as the corpuscles did not reach the usual number, about 7,800,000 
to 8,000,000, which would make the percentage about 13°4 instead of 
13:8, and the weight of the hemoglobin 129 grms. 

Now, under the administration of the drug the corpuscles fell from 
7,460,000 to 6,615,000, so 15 grms. of hemoglobin must be set free ; 
and since 3°3 grms. of hemoglobin equal 1 grm. of urea, this will equal 
4°54 germs. of urea, an amount corresponding very closely to the actual 
increase of 4°556 grms. for the two days. 


Exp. 1V.—Pyroeatnic Acip. 


For seven days before the exhibition of the drug the dog, a healthy- 


setter bitch, had passed on an average 7°679 germs. of urea per diem. 
The corpuscles at the end of the 6th and 7th day were 7,866,000, 
and 7,850,000, being an average of 7,860,000 per c.mm. The weight 
of the dog was 13°607 kilos. Calculating from Bischoff’s results we 
see that the blood weighed 1046 grms., which will, according to Preyer, 
contain 144°348—say 144 germs. of hemoglobin. 

On the 8th day 2 grms. of pyrogallic acid were given, and another 
1-5 grm. were administered on the 9th day. 

At the end of the 10th day the corpuscles had fallen to 5,190,000 
per ¢.mm., indicating a destruction of 2,670,000 pere.mm. Calculating 
from the above data we find that this indicates a liberating of 48-9 
grms. of hemoglobin. And as we have already seen that 3°3 erms. of 
hemoglobin is equivalent to 1 grm. of urea, we should expect to find, if 
all the heemoglobin was excreted as bilirubin urea, &ec., an increase of 
14°8 grms. of urea above the normal. Buta considerable part of the 
hzemoglobin escaped the action of the liver, and was excreted in the 
urine as such. Accordingly we find a rise of only 11°855 germs. of 
urea above the normal. 

During the next twenty-four hours the corpuscles fell to 2,604,000 
per c.mm., indicating a further liberation of 40 grms., of hemoglobin, 
corresponding to 12°1 grms. of urea. But the urine still contained 
blood-pigment and we find an increase of only 10°855 grms. of urea. 

Next day the corpuscles fell from 2,604,000 to 1,876,000, which 
represents the liberation of 11°2 grms. of hemoglobin, yielding 
34 grms. of urea. In reality the urea in the urine was 13°4 germs. 
above the normal. Such an excess does not, however, tell against the 
formation of urea from liemoglobin, since at this period of the experi- 
ment the increased production of blood-corpuscles tended to mask 
the increased destruction. That this is the explanation is rendered 
probable by the appearance at this time of numerous microcytes in the 
blood, 


UREA FORMATION AND BILE SECRETION, 529 


| 
Day of | Urine Sp. G. 


Exper. | in ¢.cs. 


1 | (660 1011 
2» | 1660 10 
3 700 1009 
4 750 1009 
5 680 | 1010 
6 720 ~—-| 1010 
7 620. | 1011 
8 720 | 1013 
9 560 | 1015 
10 675 1018 
el 380 | 1034 
12 680 | 1033 
1B 360 1032 
14 300 ‘| 1032 
15 600 ‘| 1014 
16 570 1016 
17 530 1010 
18 570 —'| 1010 
19 530 | 1011 
20 510 1014 
21 600 | 1011 
22 500-1013 
23 580 | 1012 
24 610 1010 | 
25 300 1030 | 
26 None, 
27 450 | 1040 


Urea in 
grms. 


“4 


6-460 
6°100 
7728 


Heemocytes 
per ¢.mm. 


5,195,000 


2,604,000 


1,876,000 
1,800,000 
2,330,000 
2,546,000 


2,893,000 
3,290,000 
3,456,000 
3,630,000 
4,616,000 
4,935,000 
4,400,000 
5,606,000 


Weight 
in 
kilos. 


13°607 


13°100 


11°70 


Remarks. 


2 grm. pyrogallic acid. 

1°) grm. pyrogallic acid. Urine 
dark, some albumin. No bile 
pigments. 

Urine contains abundant albu- 
min and hemoglobin. No 
corpuscles, no bile pigments. 

Urine contains much hemoglo- 
bin, but no corpuscles. Dog 
takes food but is markedly | 
icteric. | 

Urine bloody. Dog is dull. 
Vomited matter richly bile 
stained ; jaundice less marked, 

Urine contains much less alb., 
and is free from hemoglobin. 
Dog took 650 es. of milk. 

Urine gives a faint reaction of 
bile acids. Dog takes food 
well. 

Urine contains no blood, and a 
mere trace of albumin, with 
light tawny port wine colour. 


Urine free from albumin, 


6 grms. salicylate of soda. 

6 grms. salicylate of soda. 

Urine dark colour, contains a 
trace of albumin, but no 
heemoglobin, 

Dog appears ill, so experiment 
stopped. 


I here introduce an experiment on the action of salicylate of 
soda on the blood-corpuscles and urea production. At a later 
part of this paper I shall enter more fully into the hemocytic 
This experiment is not so satisfactory as the 
two former, because at the time the dog was recovering from the 
anemia produced by pyrogallic acid, and blood-production was 


action of this drug. 


greatly in excess af destruction. 


Nevertheless, the effect of the 


destruction of blood on the first two days of the experiment was 
well marked, though on the third day the urea produced was not 
connected with any apparent fall in the corpuscles—in all 
probability because here again increased production completely 
masked the process of disintegration. 


Ko 


53 DR D. NOEL-PATON. 


) 


000,000 


000,000 
000,090 
2,000,000 


4,000,000 
ia | 
1,000,000 


6,000,000 


vo, 
» 

9,U 
) 


At a’ 6 germs. of sali- 


2 grms, given at @, and 1°5 grms. given at 0. 


In luence of pyrozallic acid and salicylate of soda on urea production and a 
oO 
cylate of soda were given, and a similar dose was administered at 0’, 


val 
o 
a) 
ie) 
o 
° 
a 
Ss 
+) 
— & 
a 
a“ 
ae 
a -~ 
o 
=> 2 
Hoa 
S 
s 
I 
mS 
re) 
Ss 


18 
16 
14 
J 

10 


Exp. V.—SAvICYLATE OF SODA, 

This experiment is merely a continuation of Exp. 1V. From the 17th 
to the 25th day the average excretion of urea was 6°303 grms. per 
diem. From the 13th to the 24th the corpuscles had increased at the 
average rate of 354,400 per c.mm. per diem. Thus if no increased 
destruction had been induced they would have reached 5,774,000 on 
the 25th day, 6,028,800 on the 26th, and 6,883,000 on the 27thé 


» 


UREA FORMATION AND BILE SECRETION. 5351 


We may therefore consider that upon the 25th a fall of corpuscles 
from 5,674,000 to 5,200,000 occurred. Now, calculating from our 
previous data, and allowing for the loss of weight in the dog, we see 
that such a fall will set free 8°3 grms. of hemoglobin, which will 
correspond to 2°5 arms. of urea. In reality the excess over the normal 
on this day was 1:424 germs. of urea. 

Next day the fall in the number of corpuscles must be calculated at 
from 5,200,000 + 354,400 = 5,754,400 to 4,555,000, which will yield 
157 grms. of hemoglobin, equivalent to 4°757 germs. of urea; but this 
even with the surplus 1-076 grms. of the previous day falls short of the 
actual increased production by 0°911 grms. As is probably indicated 
by the marked increase on the next day, when the corpuscles, in spite 
of the presence of the drug in the system,—as demonstrated by its appear- 
ance in the urine,—rose from 4,555,000 to 5,105,000, the increased 
production was already tending to mask the amount of destruction. 
On the next day it is impossible to say whether or not any destruction 
occurred, since a rise instead of a fall in the number of hzemocytes was 
noted. 


We thus see that both the secretion of bile and the production 
of urea depend in large measure upon the destruction of blood- 
corpuscles, and that through this they necessarily bear a direct 
relationship to one another. 

In confirmation of these conclusions, I shall,in the next place, 
give the results of a series of observations on the action on the 
red blood-corpuscles of some of the more important cholagogues 
which I have already shown increase the urea production. 


(To be continued.) 


ON THE HIND-LIMB OF ICHTHYOSAURUS, AND ON 
THE MORPHOLOGY OF VERTEBRATE LIMBS. By 
D’Arcy W. THompson, B.A., Professor of Biology in Univer- 
sity College, Dundee. 


I aM indebted to Professor Turner for the opportunity of study- 
ing a remarkable skeleton ascribed to Jchthyosaurus platyodon 
in the Anatomical Museum of Edinburgh University. The 
interest of this specimen centres in the hind-limb, which presents 
several exceptional features. In the first place, the femur has 
articulated with it three bones, identifiable as tibia, intermedium, 
and fibula, as in Marsh’s Sauwranodon (Baptanodon), and as in 
the limb figured as Pliosaurus portlandicus by Owen (Fossil 
Fig. 1. 


° 
° 
° 
° 
° 
ce) 
° 
| 

| 

| 

\ 

\ 


ol 


1. Hind-limb of Sawranodon (after Marsh). 2. Left hind-limb of Jehthyosaurus 
platyodon. 3. ‘Typical hind-limb of Jehthyosawrus (after Hulke). Jes 
femur; 7'./., tibia and fibula; 4. 7, tibiale, fibulare; 7, intermedium ; 
ce, c', ce’, centralia. 

Reptiles), but aseribed to Plesiosaurus Manseli by Hulke 

(V.S.G.8., 1883, Suppl. p. 52).2 This, therefore, is an additional 


1 The substance of this paper was communicated to the British Association at 
Aberdeen, in August 1885. 

* Long ago Buckland, in his Bridgewater Treatise, figured /. p/latyodon with 
three elements in the proximal tarsal row; but the circumstance passed un- 
noticed, 


MORPHOLOGY OF VERTEBRATE LIMBS. 533 


proof that the primary location of the intermedium is in the 
propodial segment of the limb. So far the Edinburgh specimen 
is only equally archaic with the American Sauranodon, but in 
the following feature it surpasses it in primitiveness. 

The limb of Sauranodon contains in its next segment four 
bones, and so probably, to judge from its articular surfaces, did 
that of Pliosaurus. That is to say, granted that the bones already 
mentioned are rightly identified, we have in the proximal segment 
of the tarsus a ¢ibiale,a jibulare, and two centralia. In the Edin- 
burgh Ichthyosaurus we have one centrale only ; and, moreover, 
we have again in the next succeeding segment ¢iree bones only 
(tarsalia), whereas we have five in the corresponding region of 
Marsh’s Sauranodon. 

So far then we have three longitudinal series of bones in 
perfect symmetry, formed of three bones in the propodium, three 
in the proximal region of the tarsus, and three in the distal. 
And these three longitudinal rows continue distinct to the distal 
extremity of the limb. Two other longitudinal series of bones 
exist; one, a somewhat irregular series of rounded discoid 
ossicles, separate from one another and without articular facets, 
is applied to the tibial side of the limb, commencing immediately 
distal to the tarsus but not directly articulated with it; the 
other, commencing at the same level, is inserted between the 
median and the external or fibular row. 

The appearance of the limb suggests at once that both of these 
rows are accessory and not primitive. If we may consider the 
sround-plan of the limb apart from them, we have simply three 
segmented rays or longitudinal series of bones symmetrically 
articulated with a basal segment. 

While the limb of Sauranodon seemed to be a weighty argu- 
ment in support of Gegenbaur’s theory of the primarily double 
nature of the centrale, the present example seems to mea still 
more potent argument against it; for the common type of 
Ichthyosaurian limb must be considered intermediate between 
that of the present example and the typical cheiropterygium of 
the higher vertebrates; and we pass from our present case to 
the typical Ichthyosaurus, by transverse cleavage of the centrale 
and apportionment of its outer moiety to the interstitial digit. 

It is perhaps equally easy to pass downwards from this limb to 


534 PROFESSOR D’ARCY THOMPSON, 


the fin of fishes. Assuming the femur to represent the basi- 
pterygium (Balfour), we have here three basalia, which by elon~ 
cation and segmentation may be supposed to have given rise to 
the distal portion of the fin. 


Pelvis and hind-limb of Fishes. 1. Secyllium; 2. Callorhynchus; 3. Polyp- 
terus ; 4. Protopterus. p., pubis ; pp., pre-pubic process ; @., ilium ; bp., 
basipterygium ; J., basalia. 


And the limb has a close parallel resemblance to that of 
Polypterus, which is derivable by similar processes (with con- 
temporaneous degeneration of the pelvis) from the Elasmobranch 
limb. In Polypterus (where I concur with Davidoff that the 
bone commonly called pelvis is actually the basipterygium) the 
basalia are reduced to four, or asin one specimen which I have 
dissected, actually to three. 

It need hardly be said that this new conception of the 
Enaliosaurian limb is wholly incompatible with Gegenbaur’s 
view. For Gegenbaur looks upon the whole radial (tibial) side of 
the Enaliosaurian or higher vertebrate limb, beginning with and 
including the humerus, as homologous with the basal series of the 
Selachian metapterygium (—basipterygium), the other elements 
of the limb being attached laterally to the base, on the ulnar 
(fibular) side. While on the present view, the humerus (femur) 
is the only basal representative, and to it are attached radial, 
ulnar, and intermedial rays. 

It remains to be seen whether an explanation can be found 
for the Dipnoan limb, which bas remained unexplained since 
the view became untenable that it was superlatively archaic. 
In the fin of Protopterus, the segment articulating with the 
pelvis is, I take it, a true basipterygium. To it I have found 


MORPHOLOGY OF VERTEBRATE LIMBS. 535 


attached two small nodules of ‘cartilage, between which is the 
main axis of the fin. Here we seem to have three basalia, two 
aborted, and one only continued into a segmented ray, as are all 
in our Ichthyosaurus. While in Ceratodus, we are reduced to 
supposing that this one segmented ray has branched laterally, 
in order to give breadth and strength to the fin. 

In the fore-limb of the Edinburgh Ichthyosaurus, the inter- 
medium no longer articulates with the humerus, and the centrale 
is already double; so that here we have, as usual, a marked pre- 
ponderance of archaic features in the hind as compared with the 
fore-limb. 


Editorial Note.—TYhis specimen of Jchthyosaurus from Lyme Regis, 
Dorsetshire, was at one time the property of Sir William Jardine, 
Bart., of Applegarth. It was purchased at the sale of his scientific 
collection by Professor Archibald Geikie for the University of Edin- 
burgh, and is now displayed in the vestibule of the Anatomical 
Museum in the New Buildings of the Medical School.—W. T. 


THE LUMBAR CURVE OF THE SPINAL COLUMN IN 
SEVERAL RACES OF MEN. By Professor Sir WILLIAM 
TuRNER, M.B., LL.D., F.B.S. 


Ix the course of the investigations into the modifications of 
the skeleton in different races of men, which I have been con- 
ducting in connection with my Report on the Human Skeleton 
for the Reports of H.M.S. “Challenger,” I have measured the 
bodies of the lumbar vertebrie, with the view of ascertaining if 
modifications existed in their vertical diameter, anteriorly and 
posteriorly, which might affect the lumbar curve of the spine. 
If the vertical diameter of the series of lumbar bodies be less 
anteriorly than posteriorly, then the lumbar region possesses 
(if this be not compensated for by modifications in the thick- 
ness of the intervertebral discs) an anterior concavity, continu- 
ous with the anterior concavity in the dorsal region; and an 
approximation to the curvature of the spine is produced, such as 
is characteristic of the spinal column in all mammals except man. 

Anatomists are in the habit of teaching that the human spine 
is convex forward in the lumbar region, so that a lumbar con- 
vexity is interposed between the thoracic and sacral concavities, 
and contributes to the alternating series of concavo-convex 
curves of the spinal column, which are associated with the erect 
attitude of man. My belief in the universality of the view 
that the human spine is invariably convex forward in the 
lumbar region was disturbed some years ago, when Charles 
Robertson, Esq., of the Oxford Museum, showed me the skeleton 
of an aboriginal Australian in that museum, which he had 
articulated in 1873. Mr Robertson told me that the skeleton was 
that of an adult male of the Tomki tribe of the Richmond River, 
N.S.W. In it there was a continuous curve, concave forwards 
through both thoracic and lumbar regions. As the skeleton was, 
however, artificially articulated, the question naturally arose in 
one’s mind if this modification in the lumbar curve might not 
have been produced by some peculiarity in the method of 
articulation, and was not therefore natural to the spine. Since 
I saw this skeleton, however, Mr Robertson has written to tell 
me of another adult male from Port Augusta, South Australia, 


THE LUMBAR CURVE OF THE SPINAL COLUMN, 537 


articulated in 1878, which exhibited a similar concavity in the 
lumbar region, and that the articulated skeletons of a Gilbert 
Islander and a male Andaman Islander have a similar lumbar 
concavity, though not so well marked. Before I had heard, 
however, of these later specimens in the Oxford Museum, I had 
examined the lumbar vertebra in the series of spines at my 
disposal, and had obtained some interesting results. 

Two important factors contribute to the curve in the lumbar 
region, viz., the vertebral bodies and the intervertebral discs. The 
exact share contributed by each of these parts can only be 
ascertained with precision by the method of observation which 
Professor D. J. Cunningham, of Trinity College, Dublin, is con- 
ducting, of making longitudinal mesial sections through the long 
axis of the spine in frozen subjects, and then carefully measur- 
ing the relative thickness both of the vertebral bodies and the 
discs. In the absence, however, of the fresh bodies of Austral- 
iaus and other aborigines, I have been precluded from obtain- 
ing any information on the thickness of the discs, and have been 
restricted to the examination of the vertebrze themselves, so far 
as they have been preserved in the skeletons which have reached 
me. I have measured, therefore, the vertical diameter of the 
body of each lumbar vertebra, both in front and behind, and 
have noted the difference in each vertebra, and in the series of 
lumbar vertebree in each spine. 

In order to obtain some data for comparison I measured the lumbar 
vertebree in twelve adult European spines, the majority of which were 
males, and found that the vertical diameter of the anterior surface of 
the bodies of the five vertebre in each spine was collectively greater 
than the vertical diameter of the posterior surfaces in the same spine. 
The maximum difference between the collective depth of these sur- 
faces in the series of five vertebrae was 11 mm. in one skeleton, and 
the minimum difference was 1 mm. in another skeleton. The mean 
collective depth of the five vertebre in the twelve European skeletons 
was 137 mm. for the anterior, and 131-4 for the posterior surface, the 
mean difference therefore was 5°6 mm. in favour of the anterior sur- 
face. If we were to assume that in these spines each intervertebral disc 
was of equai thickness throughout, then the greater thickness of the 
vertical diameter of the bodies in front than behind in each spine 
would give a slight convexity forwards to the spinal column in the 
lumbar region. But there is reason to believe that this difference in 
vertical diameter is not limited to the vertebral bodies, and that some 
of the discs also are thicker anteriorly than posteriorly so as to increase 


the anterior convexity. 
VOL, XX. . 2M 


538 PROFESSOR TURNER. 


If we now examine the individual lumbar vertebre in each of these 
European spines we shall find that with only two exceptions the body 
of the lst lumbar vertebra was deeper behind than in front, in one 
instance 6 mm., in another 4mm. but usually not more than 1 or 2 mm.; 
in the exceptional cases the anterior and posterior vertical diameters 
were equal. The body of the 2nd lumbar was deeper behind than in 
front in six spines; they were equal in depth in four spines, and the 
anterior surface was deeper than the posterior in two spines. The 
body of the 3rd lumbar was deeper in front than behind in ten spines, 
and in two they were equal. The body of the 4th lumbar was deeper 
in front than behind in eleven spines, and deeper behind than in front 
in one specimen. The body of the 5th lumbar was deeper in front 
than behind in all the specimens. From these spines it is evident 
that, whilst in the 1st and 2nd lumbar vertebre the body was deeper 
behind than in front in a considerable proportion of the specimens, 
in the 3rd and 4th lumbar the reverse occurred, until in the 5th 
lumbar the bodies of all the specimens had a greater vertical diameter 
anteriorly than posteriorly, and this indeed is a character of the 5th 
lumbar that has long been recognised by the descriptive human 
anatomist. In this series of twelve European spines, if we take the 
vertical diameter of the body of the 4th lumbar in the series we find 
that it amounts to 336 mm. for the anterior surfaces collectively, and 
to 313 mm. for the posterior surfaces collectively. In the 5th lumbar 
the vertical diameter of the anterior surfaces collectively amounted to 
337 mm. and the posterior surfaces to 281 mm.; the mean anterior 
depth was 28 mm., the mean posterior 23:4, and the mean difference 
in favour of the anterior surface was 4°6 mm. Hence it follows that 
of all the lumbar vertebrz the 5th has much the greatest proportional 
depth at the front than at the back of its body, and that it contributes 
more than any of the others to the anterior convexity of the lumbar 
portion of the spinal column. 


For the purposes of comparison of the lumbar region in 
Europeans with that in the spines of other races of men, it may 
be well to frame a lumbar index both for the entire region and 
for the body of the 5th lumbar. If we assume the vertical 
diameter of the bodies of the five vertebre anteriorly to = 100, 
posterior diameter x 100 

anterior diameter 
required. If this be applied to the lumbar region in Europeans 
the mean index in them of the series of five vertebre is 95, and 
the mean index of the 5th lumbar vertebra itself is 83. 


then the formula would give the index 


During the past few years I have collected the skeletons of seven 
adult aboriginal Australians,—six men and one woman. In four of the 
men the lumbar spine is complete, in one the last lumbar vertebra has 
been lost, in another the 3rd, 4th, and 5th lumbars are absent ; in the 
woman all the lumbars are present. In each of the five skeletons in 


THE LUMBAR CURVE OF THE SPINAL COLUMN. 539 


which the lumbar spine was complete, the vertical diameter of the 
bodies of the five vertebra collectively was deeper behind than in front ; 
the maximum difference observed in three male skeletons was 9 mm., 
the minimum in the woman was 2mm. The mean collective depth of 
the five vertebra in the five perfect Australian skeletons was 112°2 mm. 
for the anterior surface of the bodies, and 118°8 mm. for the posterior 
surface ; the mean difference, therefore, was 6°6 mm. in favour of the 
posterior surface. In the relation of the vertical diameter of the 
posterior surface to the anterior surface the opposite condition prevailed 
to that which was found in the Europeans. In the skeleton in which 
the 5th lumbar was absent the collective diameter of the four lumbars 
was 3 mm. greater behind than in front. Before, indeed, I had 
measured the vertebre in these Australians, I found that, when the 
lumbars in each spine were articulated together, the bodies gave a 
concave curve forward, and not a convex curve as in the European spine, 
so that I was not surprised to see, when the bodies were measured, that 
collectively they were deeper posteriorly than anteriorly. 

When the measurements of the individual lumbar vertebre in the 
series of Australian spines were examined, it was seen that the body 
of the 1st lumbar vertebra in every instance was deeper behind than 
in front, in four skeletons as much as 4mm. The body of the Znd 
lumbar was with one exception deeper behind than in front, in two 
specimens as much as 4 mm.; in the exceptional vertebra the depth 
in front was 1 mm. greater than behind. The body of the 3rd lumbar 
in four skeletons was deeper behind than in front; in one skeleton 
they were equal, and in another—the adult female—the anterior 
diameter was 1 mm. deeper than the posterior. The body of the 4th 
lumbar was deeper behind than in front in three skeletons ; these 
diameters were equal in one, and in two the anterior diameter was 
greater than the posterior. The body of the 5th lumbar was deeper 
in front than behind in all the five complete skeletons, the maximum 
difference between the two surfaces being 3 mm. 

When these dimensions are compared with those obtained from the 
European spines, it will be seen that in the Ist, 2nd, and 3rd lumbars the 
body was more constantly deeper behind than in front in the Australians 
than in the Europeans. In the 4th lumbar, whilst it was the exception 
in the Europeans for the body to be deeper behind than in front, in 
the Australians one-half the skeletons exhibited this relation. In all 
the Australians, as in the Europeans, the body of the 5th lumbar was 
deeper in front than behind ; the mean vertical diameter of the anterior 
surfaces was 23'2, and of the posterior 21-2, a difference of 2 mm. only 
in favour of the anterior surface; whilst in the Europeans the anterior 
surface was on the average 4°6 mm. thicker than the posterior. 

The mean lumbar index in the Australians was 105°8, and the mean 
index of the 5th lumbar vertebra was 91. 

In my single male Bush skeleton the collective vertical diameter of 
the bodies of the five lumbar vertebrae: was 108 mm. anteriorly, and 
115 mm. posteriorly. In the Ist, 2nd, and 3rd lumbars the posterior 
diameter exceeded the anterior; in the 4th these two diameters were 
equal, and in the 5th the anterior diameter was 1 mm. greater than 


540 PROFESSOR TURNER. 


the posterior. The proportions in this skeleton closely corresponded 


to what was seen in the Australians. ‘The lumbar index was 106, and 


the index of the 5th lumbar vertebra was 95. 

In my series of Andamanese skeletons only two had the lumbar 
vertebree complete. In one the vertical diameter of the five vertebrae 
collectively was 113 mm. anteriorly, and 112 mm. posteriorly ; in the 
other 125 mm. anteriorly, 124 mm. posteriorly. The lst and 2nd 
lumbars in both skeletons were thicker behind than in front, The 
3rd lumbar in one skeleton was of equal diameter on both aspects, and 
in the other was 1 mm. thicker behind than in front. In both skele- 
tons both the 4th and 5th lumbars were thicker in front than behind, 
in the one skeleton the anterior surface of the 5th lumbar being 3 mm., 
in the other 5 mm., thicker than the posterior. The mean lumbar 
index of the two skeletons was 99, and the mean index of the 5th 
lumbar vertebra was 84, 

In three Negro skeletons I was able to measure the vertical diameter 
of the bodies of the lumbar vertebrae both in front and behind. In 
each of the three skeletons the collective vertical diameter of the five 
lumbar bodies was slightly greater in front than behind ; the maximum 
difference, however, was only2 mm. ‘The mean collective depth of the 
five vertebre in the three Negro skeletons was 121 mm. for the anterior 
surfaces, and 119°6 mm. for the posterior surfaces ; the mean difference, 
therefore, was 1:4 mm. in favour of the anterior surface. In all three 
skeletons, both the 1st and 2nd lumbars were slightly deeper behind 
than in front ; the 3rd lumbar was equal in depth both anteriorly and 
posteriorly, whilst both the 4th and 5th lumbars were somewhat 
deeper in front than behind. The mean lumbar index was 98:9, and 
the mean index of the 5th lumbar vertebra was 89. 

In a Maori skeleton, from Otago, the vertical diameter of the series 
of five vertebra, was the same both in front and behind, viz., 101 mm. 
The Ist and 2nd lumbars were slightly deeper behind than in front, 
the 3rd and 4th were equal in depth on both surfaces, and the 5th 
was 3 mm. deeper in front than behind. The lumbar index was 100, 
and the index of the 5th lumbar vertebra was 85. 

In each of two female skeletons from Oahu, in the Sandwich 
Islands, the collective vertical diameter of the five lumbar bodies was 
ereater behind than in front; in the one skeleton the difference was 
7 mm., in the other 4 mm., in favour of the posterior surface. The 
mean collective depth of the five vertebra in the two skeletons was 
117°5 mm. for the anterior and 123 mm. for the posterior surfaces ; 
the mean difference, therefore, was 5‘5 mm. in favour of the posterior 
surface. In both skeletons the bodies of the Ist, 2nd, 3rd, and 4th 
lumbars were all deeper behind than in front, whilst the 5th lumbar 
was deeper in front than behind. The mean lumbar index was 104°7, 
and the mean index of the 5th lumbar vertebra was 87. 

In one of three Hindoo skeletons, a tall male,! the vertical diameter 
of the series of five lumbar bodies was 137 mm. anteriorly, and 146 
mm, posteriorly. The 1st, 2nd, and 5th lumbars were deeper behind 

1 This skeleton, presented by Dr John Anderson, F.R.S., was estimated as 
belonging to a man 6 feet high, 


. 


THE LUMBAR CURVE OF THE SPINAL COLUMN. 541 


than in front, the 3rd was 1 mm. deeper in front than behind, and in 
the 4th these two diameters were equal. The lumbar index was 106, 
and the index of the 5th lumbar vertebra 107. In the two other 
Hindoo skeletons, a male and a female, the vertical diameter of the 
bodies of the five lumbars was somewhat deeper in front than behind, 
and the mean lumbar index was 97°8. In each of these skeletons the 
vertical diameter of the 5th lumbar vertebra was deeper in front than 
behind, and the mean index was 89. In the skeleton of a male Sikh, 
the vertical diameter of the five lumbar bodies was 130 mm. anteriorly 
and 133 mm. posteriorly, being a difference of 3 mm. in favour of the 
posterior diameter. In this skeleton the Ist and 5th lumbar bodies 
were deeper behind than in front, but the 2nd, 3rd, and 4th were 
each of equal diameter on both aspects. The lumbar index was 102, 
and the index of the 5th lumbar vertebra was 108-7. 

In a Chinese skeleton the vertical diameter of the five lumbar 
bodies was 145 mm. anteriorly, and 123 mm. posteriorly. In each 
vertebra, except the lst, the vertical diameter was deeper in front 
than behind, and in the Ist the two diameters were equal. The 
lumbar index was 84°8, and the index of the 5th lumbar vertebra was 
70. In a male Malay skeleton, the vertical diameter of the five 
lumbar bodies was 127 mm. anteriorly, and 125 mm. posteriorly. In 
the Ist, 2nd, and 3rd, the posterior diameter was deeper than the 
anterior; in the 4th and 5th the anterior diameter was deeper than 
the posterior. The lumbar index was 98, and the index of the 5th 
lumbar vertebra was 77:7. 

In a female Esquimaux, the vertical diameter of the series of five 
lumbar bodies was the same in front and behind (127 mm.), so that the 
lumbar index was 100. In a male skeleton the vertical diameter of 
the bodies anteriorly was 120 mm., and posteriorly 116 mm., and the 
lumbar index was 96-6. Both in the female and male the lst and 
2nd lumbars were deeper behind than in front, but the 4th and 5th 
lumbars were deeper in front than behind. The index of the 5th 
lumbar vertebra in the female was 81, and in the male 71. Ina 
male Laplander the vertical diameter of the five lumbar vertebre was 
111 anteriorly and 110 posteriorly, and the lumbar index was 99. In 
a female Laplander the vertical diameter was anteriorly 121 mm., 
and posteriorly 118 mm. and the lumbar index was 97°5. In both 
skeletons, whilst the lst lumbar vertebra was deeper behind than in 
front, both the 4th and 5th lumbar were deeper in front than behind. 
The index of the 5th lumbar in the male was 86, and in the female 88. 


From the data which are recorded in the preceding pages of the 
measurements of the lumbar region in thirty-six spines of various 
races of men, it will be seen that differences occur, often to a con- 
siderable degree, in the vertical diameter anteriorly and poste- 
riorly of the bodies of the series of five vertebree. These differ- 
ences are expressed numerically by the lumbar index, computed in 
the manner already explained. The lowest index (848) wasina 


542 PROFESSOR TURNER. 


Chinese skeleton, and the highest (106) in a Bushman and ina 
male Hindoo, with a mean of 105°8 in a series of five Australians, - 
whilst the mean index of twelve Europeans was 95. The number 
of Europeans measured may, I think, be regarded as sufficient on 
which to frame an average. But it would not be safe to speak 
so definitely of the mean index in the other races, on account of 
the few skeletons which have as yet been measured. Still, from 
the fact that each of the five Australian skeletons presented the 
character of having the vertical diameter of the series of lumbar 
bodies deeper posteriorly than anteriorly, and from the peculi- 
arities of the two Australian skeletons in the Oxford Museum 
articulated by Mr Charles Robertson, there can, I think, be little 
doubt that in that race it is the rule for the lumbar vertebree to 
have an opposite relation, as regards the depth of the body, to 
what is found in Europeans. 

So far then as one can judge of the configuration of the lumbar 
spine by the measurements of the bodies of the vertebra with- 
out the intervertebral discs, this region may present one or other 
of three forms in different races of men. It may be convex 
forwards ; or straight; or concave forwards, and to each group a 
numerical limit may be assigned, based on the lumbar index. 
We may assume that a spine with the lumbar index from 98 to 
102, both inclusive, is a straight spine, Ortho-rachic; one with 
an index above 102 is a spine concave forwards, Koilo-rachic ; 
and one with an index below 98 is a spine convex forwards 
Kurto-rachic. From the data before me the skeletons which 
I have measured would be arranged as follows :—The Chinese and 
Europeans would have convex lumbar regions, the Andamanese, 
Negros, Maoris, Sikhs, and perhaps Hindoos, Esquimaux, and 
Lapps would have straight lumbar regions, whilst the Australians, 
Bush, and Sandwich Islanders would have concave lumbar 
regions. But this arrangement is of course entirely provisional 
aud will doubtless require to be modified as observations on 
the lumbar vertebre are multiplied. Variations in the anterior 
curvature of the spine in the lumbar region would in all proba- 
bility affect the outline of the back of the body in the lumbar 
region, as one would not expect the back to have so well marked 
a hollow in that region, when the spine is concave forwards as 
when it possesses the anterior convexity. How far these depart- 


THE LUMBAR CURVE OF THE SPINAL COLUMN 543 


ures in the lower races of men from the well-recognised lumbar 
convexity of the higher races may serve to modify the spine in 
the erect attitude can only be definitely settled when the interver- 
tebral discs, as well as the vertebral bodies, have been measured. 

As regards the 5th lumbar vertebra, in all the races the 
vertical diameter of the anterior surface of the body is deeper 
than that of the posterior. There are without doubt ditferences 
in the relative depth. The anterior diameter is proportionally 
greater than the posterior in the Chinese, Malay, Esquimaux, 
Lapps, Europeans, and Andamanese, than in the Australians, 
Bush, Negros, and Hindoos. In one Hindoo skeleton, and in 
the Sikh, the posterior diameter of this vertebra was deeper than 
the anterior, but these were probably individual exceptions, and 
this greater depth would assist in giving the high lumbar index 
exhibited by these two skeletons. 


Anatomical Hotices, 


FLOATING KIDNEY. By W. Arsursnor Lanz, M.S. 


Ix a female subject in the dissecting room I found the right kidney 
freely movable. It was completely enclosed in peritoneum, and was 
attached to the abdominal wall by a mesentery, whose attachment to 
the kidney extended from the upper extremity of its inner margin to 
the lower limit of the hilum. This mesentery was connected to the 
abdominal wall in a direction extending obliquely downwards and to 
the right from the origin of the right renal artery from the aorta, The 
mesentery at its attachment to the abdominal wall was 4} inches in 
breadth. The renal artery and vein ran along its free inner margin, 
which was 34 inches long, its outer margin measuring 24 inches. The 
ureter, which was lax and tortuous, ran up behind the kidney, where it 
passed between the two layers of the mesentery, and expanded to form 
the pelvis. The kidney was placed obliquely, its direction being 
downwards and to the left. It was considerably smaller than the left 
kidney, and had a cyst in its anterior wall ¢ of an inch in diameter. 
The right suprarenal capsule occupied its normal position. The kidney 
could be easily made to lie on the left side of the spinal column, and 
to reach the front and left side of the fifth lumbar vertebra and the 
lumbo-sacral articulation. The left kidney was more extensively 
covered by peritoneum than usual. The liver and spleen were not 
enlarged. The woman was not very stout, but her tissues were soft 
and flabby. Her vessels were all remarkably tortuous. She had 
worked hard, as her spinal column showed well-marked pressure 
changes. In this specimen the kidney possessed a more complete 
peritoneal investment than in the case described by David Hepburn, 
M.B., in the Journal of Anatomy and Physiology, Jan. 1885. 


AN INTERCLAVICULAR MUSCLE IN THE HUMAN 
SUBJECT. By W. Arsutunor Lanz, M.S. 


I round the following remarkable muscle in the body of a powerfully- 
built male subject. It consisted of two symmetrical fleshy bellies 
of considerable size. These were joined by a round tendon nearly as 


ANATOMICAL NOTICES. 545 


thick as that of the long head of the biceps muscle. Each fleshy 
belly arose from the clavicle immediately in front of the attachment of 
the rhomboid ligament, and from the front of the rhomboid ligament 
just below its clavicular insertion. The muscle measured 1} inch in 
transverse measurement at its origin. Its tendon crossed the front of 
the manubrium nearer its lower than its upper limit, but it was in no way 
connected to it. On the front of the manubrium, on each side of the 
middle line, it was joined by a rounded tendon. This tendon, which 
was not an aponeurotic expansion, was derived from the upper part 
of the sternal portion of the pectoralis major; namely, that portion 
which usually arises from the manubrium sterni and the inner portion 
of the first costal cartilage. In this case this part of the muscle had 
no insertion into bone or cartilage, but ended in a tendon, which 
passed freely over the sternum, and fused with the tendon of the 
digastric interclavicular muscle. The portion of the pectoralis major 
below this, and the clavicular division of the muscle, had their usual 
attachments. The latter covered a great part of each belly of the 
interclavicular muscle, and between it and the upper portion of the 
sternal division of the muscle there was an angular interval of con- 
siderable size, in which the belly of the interclavicular muscle ap- 
peared. The interclavicular muscle had no connection with the sub- 
clavius, from which it was separated by the costo-coracoid membrane. 
Its function was evidently to approximate the clavicles. That portion 
of the pectoralis major joining it formed practically an interhumeral 
digastric muscle. It could be separated as a distinct muscle from the 
remainder of the pectoralis major for a very considerable distance. 
The subclavius, costo-coracoid membrane, rhomboid, and coraco-clavi- 
cular ligaments were normal. 

This variety in the supernumerary muscles attached to the clavicle 
is rare, and the nearest approach to the form above described is one 
named by Professor Gruber, musc. interclavicularis anticus digastricus, 
and described by him in Reichert u. du Bois-Reymond’s Archiv.,1865, 
vol. vii. p. 710, pl. xviii. fig. 3. 


VOL. XX. 


| mS] 
A 


546 NOTICES OF NEW BOOKS. 


Hotices of Hew Pooks. 


Revue d’ Anthropologie, Dirigée par Paul Topinard, Troisieme Série, 
Tome i., Premier Fascicule. Paris, 1886. 


In 1872 the late M. Paul Broca founded the Revue d’ Anthropologie, and 
up to the time of his much lamented death was actively engaged both in 
writing for it and in editingit. At his death the direction of the Revue 
was intrusted to M. Topinard, Secretary of the Anthropological Society 
of Paris. From its foundation it has been the recognised medium 
for the publication of a number of valuable memoirs in the French 
language on various branches of anthropology. With the commence- 
ment of the present year a new series, the third, of the Revue is begun, 
under the direction of M. Topinard, whose recent most elaborate work, 
entitled Eléments d’ Anthropologie Générale, has placed him in the first 
rank of living anthropologists. Co-operating with him in the direction 
are such well-known names as MM. de Quatrefages, Hamy, Mathias 
Duval, General Faidherbe, Dr Gavarret, the Marquis de Nadailfiac, 
Baron Larrey, MM. Jules Rochard, L. Rousselet, and d’Arbois de 
Jubainville. 

The number for January contains original articles by Dr Topinard, 
Dr Verneau, M. Seeland, M. de Nadaillac, and M. Ledouble. Im addi- 
tion, there are critical and prehistoric reviews, and abstracts of various 
memoirs from the German, English, and French. 


Report of a Committee of the Clinical Society of London, nominated 
November 10, 1882, to investigate Spina Bifida and its Treatment 
by the Injection of Dr Hunter’s Iodo-Glycerine Solution. 


Tus report, which has been lately issued, bears the signatures of 
Howard Marsh, A. Pearce Gould, H. H. Clutton, and hk. W. Parker, 
hon. sec. Before attempting to discuss the results of the treatment, 
the committee thought it of essential importance to determine, more 
clearly than had hitherto been done, the pathological conditions in- 
cluded under this term, and with that object undertook examination 
of all the specimens contained in London museums, as well as those in 
Cambridge and Glasgow, amounting to 125. They thus went over 
much the same ground as I had done, and their conclusions are, in the 
main, confirmatory of the conclusions given by me in the Lancet, 
March 25, 1885, and in vol. xix. of this Jowrnal,—viz., (1) the spinal 
cord and nerves are, in most cases, prolonged into the sac, and ex- 
panded in its walls. The nerves proceed from the sac in regular order 
through the intervertebral foramina, some running forwards with the 
cord into the spinal canal; and the ganglia upon the posterior roots 


NOTICES OF NEW BOOKS. 547 


are normal, though, in a few cases, as shown by specimens in the Cam- 
bridge Museum, two or more are blended together. (2) When the 
spina bifida is in the upper lumbar or dorsal region the cord may be 
traced again from the sac into the lower part of the spinal cord. (3) 
The arachnoid is traceable into the sac, as well as the dura mater; and 
the fluid is in the subarachnoid space. I conclude it is implied that 
the fluid is, as I have stated it to be, in the anterior subarachnoid 
tissue. (4) As a rule, the base only of the tumour is covered with 
skin, which ends abruptly in a thin membrane in which the various 
tissues, including the nervous, and the membranes of the cord, are 
blended. (5) The instances in which the cavity of the sac is a dila- 
tation of the central canal of the cord are very rare. With reference 
to this there is the following important addition to our knowledge :— 
“The histology of the sac-wall in a typical case of meningo-myelocele, 
by demonstrating the integrity of the central canal of the included 
portion of the cord, settles beyond doubt what must otherwise be 
matter of conjecture only, that neither does the neural furrow remain 
unclosed in spina bifida, nor, after having been closed, is it subsequently 
distended by dropsy and ruptured ;” and “this examination seems to 
complete the refutation of the view held by Forster and many subse- 
quent German writers, viz., that spina bifida, in the great majority of 
cases, is due to a dropsy of the central canal of the cord.” (6) The 
malformation is attributable to imperfect development of the meso- 
blastic tissues on the posterior aspect of the cord ; but the difficult ques- 
tion of the relation of the collection of fluid to the imperfection of 
development is not discussed. The fixation of the cord in the sac from 
an early period of embryonic life of course prevents its ascent in the 
spinal canal during development, and in some cases reverses the ordi- 
nary direction of the nerves, causing them to pass upwards to their 
intervertebral foramina instead of downwards. 

The pathological anatomy of the three varieties—Spinal meningocele 
(protrusion of the membranes only), meningo-myelocele (protrusion of 
the membranes with the cord and nerves), and Syringo-myelocele (dila- 
tion of the central canal), is said to be strictly parallel with those occur- 
ring in the head, viz, Meningocele and Encephalocele ; but by what 
contrivance the three spinal conditions are to be made to correspond 
with the ¢wo cerebral we are not told. 

Though not denying the occurrence of spinal meningocele, I was not 
able to find a single unequivocal example of it; and I must confess 
to feel great doubt about some, at any rate, of the ten examples of this 
condition mentioned in the report. 

Analyses of the fluid by Dr Halliburton show slight alkaline 
reaction, slight opalescence on boiling, small quantity of solid matter, 
consisting of sodium chloride, phosphates, and carbonates, with small 
_ fraction of proteids, which was composed of globulin, 

Some unusual varieties are mentioned, such as subdivision of the 
sac and the presence of bony outgrowths across the spinal canal in the 
_ neighbourhood of the tumour, five examples of the latter being given. 


Camprincr, January 1, 1886. G. M. Humpnury. 


548 NOTICES OF NEW BOOKS. 


Die Chirurgische Anatomie in ihrer Beziehung zur Chirurgischen Diag- 
nostic, Pathologie und Therapie, ein Handbuch fiir Studirende 
und Aerzte, von Prof. Dr Max ScHt ier, in Berlin, Heft. 1., 
Die obere Extremitat, 1885. 


Tus promises to be the most complete and exhaustive treatise on 
surgical anatomy that has hitherto appeared in any country. It is 
illustrated by numerous woodcuts, 1s written in a simple style, and 
will be valued both by students and practitioners. It gives surface 
markings, measurements, disposition of fasciz, vessels, muscles, &c., 
descriptions of joints, directions of dislocations and fractures, the 
causes of displacements, the courses which matter is likely to take, the 
position of aneurisms, wounds of arteries, the methods of compression 
and ligaturing the several arteries, re-setting joints, and a variety of 
information bearing upon the practice of surgery. The action of the 
several muscles is given, with the effects of paralysis of them, and the 
appropriate points for Faradization. 

When fluid is injected into the shoulder-joint in the dead body the 
arm is said to be thereby slightly raised, abducted, and rotated 
inwards, the head of the humerus to be pressed a centimeter away 
from the glenoid cavity. At the same time the lower angle of the 
scapula is thrown a little backwards—this and the abduction of the 
arm being due to the comparative looseness of the capsule at its under 
part. The swelling and fluctuation are most perceptible in front and 
behind; and the synovial processes upon the biceps-tendon and the 
subscapular muscle become distended. When the cavity of the elbow- 
joint is in like manner filled the forearm becomes bent upon the arm 
nearly to a right angle, and semi-prone. The capsule, as shown in a 
woodcut (p. 198), is distended all round, but especially in front and 
behind on the sides of the brachialis anticus and the triceps; and the 
radius and ulna are found to be distanced from the humerus one or 
two millimetres. In the case of the radio-carpal joint the hand 
remains in a straight line with the fore-arm, or slight dorsal-flexion 
may be induced, and the articular surfaces are separated three or four 
millimetres from one another. 

At page 309 is represented the dissection of a specimen of contrac- 
tion of the palmar fascia (Dupuytren’s contraction) in which the pro- 
cesses of the fascia extending into the fingers are seen to be thickened 
and contracted, while the flexor tendons, as the author remarks, are 
unaffected. He notes the fact that the fascial processes of the ring 
finger are commonly first affected, which he rightly attributes to the 
circumstance that pressure from a stick or other body held in the hand 
is most felt at this part, and acts as a source of irritation to the skin 
and fascia here. 

Enough has been said to show that the work, in this its first instal- — 
meut, is a valuable practical addition to anatomical literature. ; 


Vol XX PLA. 


RW.Schufeldt del. aN F. Huth, Lith? Edint 


CONURUS CAROLINENSIS. 


—— 


Journ. of Anat. dé Phys, Aprit 1886. Vol. XX, PU. XI. 


W.R.Schufeldt del. F. Huth, Lith® Edin® 
GONURUS CAROLINENSI!IS. 


> 


Journof Anat. dé Phys, tort 1886 


W.R.Schufeldt del 


NAVAJO 


INDIAN. 


Vol. XX, Pt. XT. 


F. Huth, Lith® Edin? 


Vol.XX, Pl. XU. 


| Journ. ot Anat.& Phys, April 1886. 


t of a Mare. 


Samen 


Cysts in the broad Li 


Trachea of Emu. 


From a Cow. 


ith Edint 


> 
Li 


F Huth, 


OIRAEGAN OIF = Cie Sores: 


Journ.ot Anat é Phys, April [886 VoLAX, PLATV. 


BLOOD. FORMING CELLS. 


F. Huth, Lith’ Edin* 


Journ. of Anat. é Phys, tprul [886 Vol. XX, PLAXV. 


F Huth, Lith? Edin™ 


HS MOL iOIG:Y JOlFS (SIP E SEIN. 


wh ery @| 


Sourn.ot Anat.é Phys,.tpril 1886 Vol. XX, PLXV. 


MAEFORMED FROG. 
From Photograph F. Huth, Lith™ Edin® 


Fournal of Anatomy and Phpstology. 


ON THE PHYSIOLOGY OF THE HEART OF THE 
ALLIGATOR. By T. Westey Mis, M.A, M_D., 
Lecturer on Physiology, MGill University, Montreal, Canada. 


THE only paper bearing on the physiology of the heart of the 
Crocodilia is one by Gaskell, in vol. v. No. 1 of the Journal of 
Physiology. The anatomy of the sympathetic is considered by this 
writer in a paper in Nos. 4, 5, and 6 of vol. v. of the same. 

With some of the conclusions in the former short communica- 
tion my observations accord; with others they are entirely at 
variance. I shall therefore refer in some detail to the experi- 
ments leading me to opposite conclusions from the writer 
referred to above. 

My experiments were made upon specimens of Alligator 
Mississippiensis, part of them at the Marine Laboratory of the 
Johns Hopkins University at Beaufort, N.C.; and part of them 
in the Biological Laboratory of the same institution in Baltimore, 
to the directors of both of which I desire to acknowledge my 
indebtedness for their kindness in facilitating this work. 
Gaskell has stated, in the paper referred to, that in the Crocodile 
“the vagus seems to contain purely inhibitory fibres.” That 
such is far from being the case, the following extract from notes 
of my experiments on the Alligator will amply show. It is 
scarcely to be supposed that the case would be different for 
creatures so alike as the Crocodile and the Alligator. 

For stimulation the interrupted current of a Du Bois’ in- 
duction coil fed by one Bunsen’s cell was used. 


I. The Results of Stimulation of the Vagus Nerve. 


The following extracts from my notes bear on this subject :— 
VOL, XX. 20 


550 DR T. WESLEY MILLS. 


Experiment.— 
Cardiac rhythm = 42. 


1. Stimulation of right vagus with interrupted current led to arrest, 
and the following after-rhythm :— 


During the Ist minute 43 


4 2nd. ,, 44 
es pid Jc, . ako 
4th —,, 43 


Witer 10 minutes R. = 42. 


2. Stimulation of left vagus led to arrest, and the following after- 
thythm :— 
During the 1st minute 40 


us Dad. 40.5. oo 
ri Std: — 43 ae 
Ef Ath oe 


8th ,, 44 
The original rhythm was 40 to 41. 


In another case, in which the heart was much depressed in action 
and the rhythm slow, a single stimulation raised the rhythm from 10 
to 17, with marked increase in the force of the beat. 


I have given the exact rhythm in the above cases to show the 
amount of acceleration. This is quite as much as would be 
expected for a rate of beat so high as 40, and quite as much as 
oceurs usually in the Chelonians. 


IT, How the Vagus arrests the Heart. 


As in the Chelonian, a current that suffices to stop the auricle 
proper (“bulged” part) will not arrest the sinus or sinus 
extension (“basal wall”) in all cases; so long as the sinus 
venosus continues to beat the ventricle will follow, at least 
while the beat is not very weak indeed. 

Often a momentary stop of the ventricle follows the arrest of 
the auricles, but it is very short in such cases. 

Practically, as in the Chelonians, the sinus is the controlling 
part of the heart so far as rhythm is concerned; when it is 
arrested the auricles or ventricle ceases to pulsate. 

As to whether the arrest is due solely to the influence of the 
vagus in the sinus is considered in my paper on the Sea-Turtle. 

Sometimes, in consequence of this greater power of the vagus 
over the auricles, a very weak current will produce, when first 


PHYSIOLOGY OF THE HEART OF THE ALLIGATOR. BAT 


applied, an arrest of both auricles and ventricle of brief duration, 
when the continuance of the stimulation will give rise not to 
arrest but to acceleration ; that is, the auricle breaks away and 
participates in the good effects of vagus-stimulation exactly as if 
the current had been withdrawn. As in the Chelonians, the 
heart is arrested promptly by the vagus, and not by reducing the 
force of the beat to zero. 

A fter-Effects of Stinulation of the Vagus.—These vary with a 
variety of circumstances, and follow much the same laws as in 
the Chelonians. 

2. When the rhythm at the time of stimulation is rapid, the 
after-rhythm is not much in advance of the original; it may for 
some time even fall slightly below it, but if the rhythm be very 


slow the proportionate increase may be very great. 


2. But in all cases there is an increase in the force of the 
beat, seen in ventricles as well as auricles. This is best noted 
when the original beat is feeble, or when one beat is of force 
disproportionate to another. In the Alligator this increase of 
the strength of the beat has been the most striking phenomenon. 

3. Stimulation of the vagus removes irregularities both in the 
force and frequency of the beat; this follows, not only when the 
heart is slightly disturbed, but when its whole action is very 
much depressed. 

4, The arrest of the beat usually follows after a very brief 
period of latency; and after the cessation of stimulation the 
beat recommences after a latency which varies with the con- 
dition of the heart. When this organ is strong, and at an early 
stage of experimentation, the latency last referred to is usually 
from three to five seconds; but later this may be doubled or 
trebled. 

5. The acceleration that follows vagus-stimulation reaches its 
maximum sooner, and declines more speedily when the heart’s 
nutrition has begun to suffer than when the heart is fresh. The 
difference in this respect is very great; often the maximum 
acceleration is reached under bad conditions of nutrition in one 
minute, while in a fresh condition of the heart the maximum of 
acceleration, as my notes of one experiment given above show, 
may not be reached for several minutes. This holds also for the 
Chelonians, 


552 DR T. WESLEY MILLS. 


6. After stand-still, when the rhythm is resumed, the sinus is 


the first to beat, and may be followed by the auricles and ven- 


tricles, or by the ventricles only at first, ie, the auricles are 
often the last to recover, as in the Chelonians. 

The results of stimulation of the vagus in the Alligator may 
be thus generalised :— 


Stimulation of the vagus weakens the heart’s action, 
and may cause an arrest of the auricles alone, or of the 
auricles followed by a brief stop of the ventricles; with 
a sufficiently strong current the entire heart is arrested, 
and remains so during stimulation, with marked increase 


in the usual diastolic relaxation ; after a variable period ~ 


of latency, the length depending on the condition of 
nutrition of the heart at the time, the beat is resumed ; 
frequently the auricles are the last to begin; the beat of 
the heart is marked by acceleration, reaching a maxi- 
mum in a variable time, and declining slowly or the 
reverse, according to the condition of the nutrition of 
the heart ; the beat is also characterised by greater force, 
evident in both auricles and ventricles. The force and 
frequency are increased in inverse proportion to those 
prevailing at the time of stimulation. 


It will be seen that the conduct of the heart in the Alligator 
under vagus-stimulation is very similar to that of the Chelonians, 
though in the hearts of the Crocodilia, with their two auricles 
and paired ventricles well-defined and separated,—in the shape 
of the latter especially, in the general character of the beat, 
&c.,—there is a considerable departure from this class. One 
looking at the heart of a small Alligator beating rapidly might 
believe, on superficial observation, that he had before him the 
heart of a mammal or bird; the colour of the blood issuing 
from wounds, especially about the head and upper parts of the 
Alligator, is much brighter than in the Chelonians, denoting a 
circulation admitting of better oxidation of the blood, and 
pointing to an advance in the animal scale. 


* 
. 


*” 


PHYSIOLOGY OF THE HEART OF THE ALLIGATOR. 553 


III. Accessory Vayi. 


From some previous experience with similar small nerves, in 
connection with the Sea-Turtle, to those I am about to describe, 
I was not wholly unprepared for a very remarkable result. 
These nerves may be traced in the Alligator from the glosso- 
pharyngeal, soon after its exit from the skull, down the neck, 
underneath the trachea, over the vessels, and to the heart. 
They were not more than a fifth or sixth of the thickness of the 
vagus in the specimens examined. They did not run close to 
the vagus but diverged from it, as they passed downwards more 
and more. 

An extract from my notes of one experiment will give some 
idea of their function. 

Experiment.—Nerve divided and peripheral end stimulated. 

When the rhythm was 28, stimulation led to a fall to 20; when 12 
to 6, 

Force of beat is diminished; short stops, leading to an irregular 
rhythm. 

On cessation of stimulation the rhythm rose from 28 to 35. 

During the pause the normal diastolic relaxation was increased. 

It is thus seen that in this case these nerves behaved exactly 
like the vagi; and, considering their origin and how closely 
related the vagus and glosso-pharyngeal nerves are anatomically, 
it seemed not unreasonable to suppose that some of the vagus 
fibres may have wandered off and taken a separate course to the 
heart. 

This subject is further considered in my paper on the Sea- 
Turtle. 

Whether they are always present, or have always the same 
action, I am unable to say without further examination. 


IV. Accelerator Cardiac Nerves, 

There passes from a ganglionic enlargement of the eleventh 
metamere towards the heart, and quite alone, a large well-defined 
nerve, as Gaskell and Gadow! have described. 

I subjoin the results of two stimulations of this nerve in one 
case as illustrative of its influence. 

1 Vol. v. Nos. 4, 5, and 6 of the Journal of Physiology. 


554 DR T. WESLEY MILLS. 


Experiment.—Heart beating with a regular rhythm of 29, a short 
stimulation of 11th metamere raises the rhythm to 32 from 29. 
2. A second stimulation had the following effect :— . 


Rhythm during 1st minute of stimulation, 34 
- 4 one 9 32 
3 7, wore & 32 
but it is to be noticed, that while during the second and third minutes 
the rhythm fell the force of the beats increased. 

Thus it appears that the true test of an augmentor nerve is 
the amount of work the heart is enabled to do under its 
influence. I desire to call especial attention to this result, for 
it does not stand alone, and to a certain extent applies to the 
action of the vagus also. 

Not only are cardiac accelerators augmentors and should be 
so called, but they increase the work done by the heart; for 
when there is a fall in rhythm it is so slight as to be much over- 
balanced by the greater force of the beat. Indeed, so far as my 
observation goes, this increase of the force of the beat is by far 
the most important matter. It is easy to see how one, on super- 
ficial examination of either an accelerator or the vagus, might be 
led into erroneous conclusions, especially if the rate of beat 
were alone considered. 


V. Relative Power over the Heart of the Vagi, and the Results of 
their prolonged alternate Stumulation. 

So far as I can judge from a very limited number of experi- 
ments, the right vagus, as in the Chelonians, has somewhat 
greater influence over the heart than the left; but both are 
usually very efficient. 

I have also found that the heart is capable of prolonged 
inhibition by alternate stimulation of the vagi; but in the 
Alligator the nerves seem to die more rapidly than in the 
Chelonians, and it is doubtful if as prolonged inhibition could be 
maintained as in the latter; so far as my limited experience 
in this matter goes this is not the case. 


VI. Peculiar Cardiac Inhibition followed by Acceleration. 
In this connection I wish to discuss some peculiarities of 
reflex (?) inhibition, and describe an experiment which appears to 
be unique in physiology. 


‘ 
4 
3 
= 
J 
= 


PHYSIOLOGY OF THE HEART OF THE ALLIGATOR. 555 


Lxperiment.—Small Alligator, about 1} feet long. 

Medulla oblongata destroyed completely for more than one hour ; 
both vagi divided; the latter dead throughout the greater part of their 
course. 

Sharp tapping over stomach and liver, with an ordinary dissecting 
forceps, causes slowing, weakening, and brief stops of the heart. This 
lasts for about one minute after cessation of the stimulation, and is 
followed soon by a greatly accelerated rhythm (from 40 to 50). This 
rhythm is at first quite regular, but gradually diminishes and falls into 
irregularity. 

This experiment is repeated three or four times with precisely 
sunilar results. 


The only experiment known to me at all comparable with 
this is that mentioned by Marshall Hall) and discussed by 
M*William on p. 239, Nos. 4 and 5, vol. vi. of the Journal of 
Physiology. 

The last-mentioned writer would explain Hall’s result as due 
to mechanical stimulation of the vagi nerves, by the jar caused 
by a blow to the stomach with a hammer in Hall’s experiment, 
the brain and cord being destroyed. But in my experiment the 
vagi nerves were already dead throughout most of their length, 
and the jar of a blow from a pair of forceps caunot be considered 
as very great. On the other hand, the fact that the first effect 
was followed by regularity and acceleration of rhythm, certainly 
points to the vagus nerve. 

I wish now to call attention to several phenomena which 
seem to me to demand careful consideration. 

1. I have found in the fish that stimulation of certain parts, 
notably of the anus and tail, led frequently, and with moderate 
currents, generally to acceleration, either followed by slowing or 
not. 

2. A similar result has been observed on stimulating the liver 
in the Sea-Turtle. In both the cases referred to the medulla 
and cord and vagi were intact. 

3. On p. 271 of my paper on the Terrapin’s heart,? reference 
is made to irregularity of rhythm as the result of stimulation of 
the main sympathetic stem, and I have observed on several 
occasions, what is not therein much insisted upon, that on first 
applying the electrodes to the part of the sympathetic therein 


1 Todd’s Cyclopedia of Anat. and Phys., art. ‘* Heart.” 
2 Journal of Physiology, vol. vi. Nos. 4 and 5. 


556 DR T. WESLEY MILLS. 


defined, a brief slowing or stop of the heart has preceded the 
usual acceleration and augmentation of the beat. ; 

M¢William states that stimulation of the abdominal sympa- 
thetic does not lead in the Eel to cardiac arrest, and maintains 
that the afferent impulses are transmitted through the spinal 
cord to the medullary centre, when cardiac inhibition takes place 
after stimulation of the tail of the Eel. 

But how upon M¢William’s hypothesis is my result of ae- 
celeration under such circumstances to be explained ? 

There is one fact brought out in my experiments on the fish’s 
heart with atropin which favours M°William’s view. 

After the free application of this poison to the heart of the fish 
I have not found it possible to get reflex cardiac inhibition as 
usual; that is to say, we may get from this an argument that 
the vagi nerves, and these only, are concerned ; but again, if we 
assume that all the inhibitory fibres for the heart do not run in 
the vagus, necessarily this experiment does not carry with it so 
much force. 

It is true that hitherto it has been believed that inhibitory 
fibres were confined to the vagus; but in this paper I have shown 
that in the Alligator this does not seem to be so, and the case for 
the Sea-Turtle also favours such a view, as also perhaps what is 
referred to under (3) above. 

As mentioned on p. 254 of my paper on the Terrapin, stimula- 
tion of the main sympathetic stem does in that animal produce 
the most decided cardiac inhibition, and that when stimulation of 
the brachial plexus is ineffective. Why is this, if the course of the 
impulses is along the cord ? 

M‘William explains the after-acceleration of retiex cardiac 
inhibition (stimulation of the vagus itself not being followed by 
such), by assuming in the Eel a “constant controlling vagal in- 
fluence,’ which is weakened in the former case during stimu- 
lation. Asaresult of my eighteen experiments on this subject on 
the Slider Terrapin (see pp. 252 and 253 of the paper on that 
animal) no very strong case is made out for this view as regards 
this one cold-blooded animal. 

It seems to me, when considering all these facts, that we 
should begin to seek for explanations more satisfactory than 
some of those now prevalent; at present I have no view to 


PHYSIOLOGY OF THE HEART OF THE ALLIGATOR. 557 


present that is free from difficulties; but many of the 
phenomena I have cited in this paper, and treated elsewhere, 
seem to point in the direction of possible cardiac inhibition 
other than through the main vagus stem, or possibly even its 
ultimate branches, though the latter have, apart from the main 
stem, been but little considered, in short, that some such view 
as that suggested by me on p. 271 of my paper on the Terrapin 
may be rendered tenable by accumulating facts; but my object 
in this discussion of a very obscure subject has been rather to 
emphasize facts, and point out difficulties in the way of the 
complete acceptance of prevailing theories, than to offer new 
explanations. 

If we assume that in the case of the fish and Sea-Turtle, as 
referred to in 1 and 2 above, the impulses pass along the 
sympathetic chain and then get into some cardiac accelerating 
branch, or even first pass along the spinal cord but not to the 
medulla, it is possible to understand the acceleration that first 
* ensues under the circumstances referred to, without introducing 
the vagus at all. 

In the case of Marshall Hall’s experiment on the Eel, and my 
own on the Alligator, we may suppose the impulse to travel 
primarily along some inhibitory fibres not in the vagus stem, or 
not in the usual cardiac branches of that stem, and that these 
are finally overpowered by the influence of accelerating fibres. 
I found no more difficulty in this connection with M°William’s 
objection that stimulation of the abdominal sympathetic in the 
Eel does not produce eardiac arrest than that stimulation of the 
same part in the Chelonians does not produce cardiac acceleration 
for stimulation of another part of the main sympathetic does. 
Impulses may have and do have a great variety of choice in the 
tracts they follow. 


VII. Stimulation of the Heart with the rapidly interrupted 
Current. 


The results depend in part on the strength of the current and in 
part on the condition of the heart at the moment of stimulation. 
1. When the heart is fresh and in good condition the sinus 
can be arrested with as great ease as in the Chelonians, but when 
the heart is much exhausted, and the animal has been long 


558 PHYSIOLOGY OF THE HEART OF THE ALLIGATOR. 


under experiment, arrest by any strength of current is quite 
impossible. ; 

2. Stimulation of the auricles with a weak current causes 
dilation around the point of contact of the electrodes, and a 
weakening of the rhythm beat of the auricle. A strouger current 
wholly arrests the auricle, and the local dilation spreads over a 
wider area, finally embracing almost the whole auricle. 

3. A moderate or strong current applied to the ventricles, 
when the heart is in good condition, causes a rapid intervermi- 
form movement, followed by frequent stops. On the cessation 
of stimulation a long pause in the rhythm follows. 

In all cases, no matter what part of the heart is stimulated, 
at the exact points at which the electrodes are applied, very 
small light-coloured areas may be seen, as in the case of fishes 
and Chelonians; also the usual blue appearance in the parts 
locally dilated, but 1 have observed in the Alligator an effect 
which is not to be seen in the fish or in the Chelonian heart. 
On the application of a very strong current to the edge of the 
auricles they are seen to exchange their natural bluish colour for 
a pallor—almost a whiteness ; by carrying the electrodes along, 
more and more of the auricle takes on this appearance. The 
portion affected in this manner is thrown out of action. This, 
it seems to me, is the reverse of dilation, is, in fact, due, as also 
the finer smaller light-coloured points seen at the exact points 
of application of the electrodes when a weaker current is used, 
to a very marked and probably tetanic contraction of the heart- 
muscle. The dilation (local paralysis) may possibly be due, at 
least in part, to nerve influence, while the other is the result of 
the direct effects of the current on the heart-muscle itself. 


THE FUNCTIONS OF THE TONSILS. By R. Hriyaston 
Fox, M.D., M.R.C.P. 


Ir is remarkable that, easy of inspection as the tonsils are, 
and frequently as they are affected in a large number of diseases, 
so little is certainly known as to their function. 

The tonsils are the largest and most conspicuous portions of 
a ring of lymphatic structures, which extends around the pharynx 
and fauces. Thus there are very abundant nodules of adenoid 
tissue on the back of the tongue at its root, and numerous similar 
nodules on the sides of the pharynx about the orifices of the 
Eustachian tubes, whilst others are disposed on the hinder wall 
of the pharynx so as to form a band across it at this level. 

The tonsil itself has been described as the largest mass of 
adenoid tissue in the body, being an aggregation of round nodules, 
cumposed of this tissue, and often, though not very appropriately, 
spoken of as “follicles.” It is unnecessary to describe adenoid 
or lymphatic tissue in detail, further than to say that it consists 
almost entirely of small amceboid cells similar to those of the 
blood, but many of them presenting double or partially divided 
nuclei, these cells being enclosed in a retiform matrix. There 
is an abundant supply of blood-vessels and of lymphatic plexuses, 
and the free surface of the gland is covered by a thick layer of 
stratified squamous epithelium. This surface presents many 
involutions, forming blind depressions, the crypts of the tonsil, 
into which the ducts of small mucous glands open, and around 
which the adenoid nodules are disposed. Such crypts are found 
also on the back of the tongue. 

There is one further fact which may be here alluded to. It 
is stated by embryologists” that the fauces is the seat of what 
may be termed a developmental junction; that at this spot the 
inflected layer of epiblast which forms the lining membrane of the 
mouth meets the layer of hypoblast, which forms the major part 
of the alimentary canal. I lay no special stress upon this state- 
ment, but if it is well grounded it is of some interest. It has 
long been pointed out that new growths are apt to arise at such 


1 Kolliker, quoted in Quain’s Anatomy, section Pharynz. 
2 Quain’s Anatomy, 9th ed., vol. ii. pp. 878, 883, 884. 


560 DR R. HINGSTON FOX. 


junctions, where tissues of differing affinities met. Now, it is 
curious that the fauces, if it be such a junction, should be the 
spot selected by so many diseases for the production of inflam- 
matory lesions. 

Passing on to inquire into the function of the tonsil, we may 
ask first, Does this gland belong to the respiratory or to the 
digestive tract ? Unquestionably, to the digestive tract. When 
the position of the gland is considered, and its relation to the 
surrounding structures during the acts of respiration and degluti- 
tion, no other conclusion can I think be arrived at. 

Let us take the condition when the mouth is at rest and 
closed. The cavity of the mouth is then nearly obliterated ; 
some space is left between the tongue and hard palate, but 
further back the velum palati (or, at least, its lower edge) lies 
in apposition to the tongue surface. If the fauces be attentively 
examined whilst the mouth is closing, it will be- seen that the 
relaxed soft palate tends to assume a very sloping direction down- 
wards and backwards, perhaps nearer to the horizontal than to 
the vertical, and that the pillars on either side arch outwards, 
also in a direction tending towards the horizontal. 

Now the posterior arch of the soft palate is at a lower level 
than the anterior, and the hinder pillars are nearer together than 
those in front. This gives to the faucial arch a hollowed appear- 
ance in front, and this hollow is, in my belief, filled, when the 
mouth is closed, by the root of the tongue, which is convex from 
side to side as well as from above down. If this be so, the 
hinder pillars of the fauces lie against the tongue surface, as do 
those in front, and the tonsils, being situated on each side in the 
interval between the pillars, lie also against the tongue, and are 
shut off from the pharynx. 

In support of these statements, I may allude to the 
moulded shape which the tonsil often assumes when it is 
enlarged and soft: thus, I have more than once observed the 
are distinctly mapped out upon its surface—that facet which 
lay in contact with the tongue, separated by a ridge-like line 
from the facet which was in contact with the swollen uvula. It 
is also common, when a patient opens his mouth for inspection 
of the throat, to see vanishing strings or sheets of mucus, stretch- 


1 Cf. Dr H. Ashby in Practitioner, vol. xxxi. pp. 407 sqq. 


——— EE 


FUNCTIONS OF THE TONSILS. 561 


ing between the pillars of the fauces or the tonsils, and the 
tongue surface with which they have just been in contact. 

It will not be contested that normal respiration, after the 
period of infancy, takes place through the nose, the mouth being 
kept shut. From the posterior nares the respiratory tract will 
then pass down the hinder surface of the velum, and over the 
root of the tongue against which it lies, without entering the 
mouth, or coming in contact with the tonsils. 

In the act of deglutition, on the other hand, each morsel of 
food, being grasped first by the anterior and then by the 
posterior lamina of the faucial arch, must necessarily be brought 
into contact with the tonsils,—indeed, it must actually be rubbed 
against their surface during its passage through the fauces 
Surely the function of these glands, whatever it may be, must 
have some relation to that which is swallowed. 

If further argument be needed to sustain the position that the 
tonsils belong to the digestive tract, it may be pointed out that, 
whilst there is no other considerable collection of adenoid tissue 
in any part of the respiratory tract, there exist in the lower part 
of the digestive canal bodies almost identical in structure with 
the tonsils, and connected with them by a close pathological 
relationship—the solitary and agminated glands of the intestine. 
It would be out of place to enlarge here upon this relationship, 
but it is a matter of great interest. 

Besides, then, the presence of adenoid tissue in fine layers in 
the coats of the digestive tube, in the substance of the villi, &c., 
as indeed it is found in all the important tracts and organs of 
the body, there are these large and obvious collections of adenoid 
tissue in the wall of the canal—the tonsils and neighbouring 
nodules at the fauces, and the solitary and agminated glands in 
the intestines. As a class, these organs have been termed the 
“follicular lymphatic glands,” to distinguish them from the 
“conglobate” or ordinary lymphatic glands of the body. 

The follicular lymph glands are separated only by the ordinary 
epithelial lining from the contents of the digestive canal. It is 


1 It need hardly be said that these latter differ only in their more complicated 
structure, being traversed by lymph channels, so adapted that the fluid may be 
brought into intimate contact with the adenoid nodules in its passage through 
the glands, 


562 DR R. HINGSTON FOX. 


hardly possible to escape the conclusion that an interchange of 
some kind must take place between the glands and the food 
materials. What is the nature of this interchange? Their 
structure forbids the idea that these organs are secreting glands 
in the ordinary sense. They consist of closed nodules, without 
ducts, and without any gland cells other than the small white 
amceboid corpuscles alluded to. 

Yet the writer of the article “Tonsil,” in Quain’s Dictionary 
of Medicine, states, without any question, that the office of that 
organ is to secrete a lubricating fluid to moisten the fauces and 
aid in deglutition! This view appears to me wholly untenable. 

As regards the solitary and agminated glands, we are not, 
however, left in any doubt. The process of absorption of fat 
particles in large quantity by the amceboid cells of these glands 
has been repeatedly observed by Schifer? and others. That the 
glands are absorbent in function is what we should infer from 
their structure, 

I am not aware that any observations of this kind have been 
as yet made upon the tonsil, except that Stohr has observed 
leucocytes passing apparently between the epithelial cells, and 
reaching the mouth? But in view of its structure, and of the 
ascertained function of the closely allied glands in the intestinal 
canal, it appears to me that we may safely conclude that the 
office of the tonsil also is one of absorption. 

It will be objected to this view that the epithelial layer 
covering the gland is so thick as to render absorption through it 
improbable. The observation of Stohr just quoted would, if 
quite reliable, meet this objection, the more so as it has recently 
been shown‘ that the absorbent processes which take place in 
the digestive canal are largely carried on through the agency of 
leucocytes. There is no primd facie reason why white cells or 
their pseudopodia should not pass between the cells of a fairly 


1 Page 1647, 

2 International Journal of Anatomy and Histology, vol. ii. No. 1, 1885, pp. 
6-29. Iam indebted to Mr J. McCarthy, Lecturer on Physiology at the London 
Hospital College, for a reference to this paper. See also review of Zawarykin’s 
researches, in Lancet, 1883, ii. p. 64. 

3 Stohr, in Virchow’s Archiv. quoted by Schiifer (Joc. cit.), and by Landois. 

4 Schiifer, Joc. cit. Philipson states that absorption on epithelial surfaces takes 
place through the intercellular cement substance (Lancet, 1884, ii. p. 309). 


FUNCTIONS OF THE TONSILS. 563 


thick layer of stratified epithelium in the moist living condition. 
Pathological considerations lend a further support to this view, 
for there is strong reason to believe that the tonsils absorb 
morbid poisons directly from the saliva. 

It is not indeed likely that any considerable absorption can 
take place from the bolus of food as it passes rapidly through 
the isthmus of the fauces, encased in its glutinous covering of 
salivary fluid. But the fauces give passage to something besides 
food. I allude to the Saliva. During the intervals between 
meals} a constant stream of this secretion is very slowly 
trickling through the fauces, converging from the floor of the 
mouth on each side, and from the dorsum of the tongue, to pass 
down over the root of that organ into the pharynx. In its 
passage the saliva not only fills the crypts and bathes the 
lymphatic nodules, with which this part of the tongue surface 


_is richly furnished, but it must also, and especially, bathe the 


a 


tonsils, since these organs lie in the groove on each side of the 
tongue and probably against its surface. 

I believe the function of the tonsil to be connected with this 
stream of saliva, which is poured over it without cessation day 
and night. 

It has been stated that the bulk of all the secretions which 
are furnished to the digestive canal, are reabsorbed by the blood- 
vessels of the segment below.2 That this is so, for example, 
with some of the chief constituents of the bile, is generally 
believed. I ask then, Is it not reasonable to think that the 
tonsils reabsorb from the saliva, in the intervals of meals, 
certain of its constituents which would otherwise be wasted ? 

In conformity with modern views, we may regard the adenoid 
tissue of which the tonsils are composed as the birthplace of 
leucocytes. The materials which would be perhaps wasted in 
the stomach are thus, if my theory is correct, intercepted by 


1 That the saliva is secreted in considerable quantity when food is not being 
taken is evidenced in cases of salivary fistula, and by the dropping of saliva 
from the mouths of persons during sleep, which is often observed. The saliva 
is also, as we are all aware, not usually subjected to definite acts of deglutition, 
but passes imperceptibly through the fauces. 

2 See review of Brinton on ‘‘ Food,” in Med. Chir. Review, vol. xxx. p. 243. 
Dr Allchin in Quain’s Dict., p. 496. 

3 See especially Tappeiner, reviewed in Lond, Med. Record, Oct. 1885, p. 423. 


564 FUNCTIONS OF THE TONSILS. 


these glands, and made to minister to the growth of white 


cells. 
The tonsils are apt to atrophy in middle and later life. 


Adenoid tissue everywhere is more largely developed in child- 
hood, when not only nutrition but growth has to be provided 
for, than it is afterwards. And there is nothing surprising in 
the fact that these nurseries of young leucocytes (permit the 
fancy), planted here as it were by the river side, and drawing 
their sustenance from the nutrient stream, should dwindle in 
later life when the demand for white cells has become much 


less. 


INVESTIGATIONS IN THE RELATION BETWEEN 
CONVERGENCE AND ACCOMMODATION OF THE 
EYES. By Ernest E. Mappox, M.B., O.M. Edin., Syme 
Surgical Fellow in the University of Edinburgh. 


(Continued from p. 508, vol. xx.) 
V. Distant Vision. 


We have seen that with near vision the visual axis of an 
excluded eye generally deviates outwards from its fellow; it 
appears to be just as usual with distant vision for an excluded 
eye to deviate inwards. This is easily shown by pricking two 
pin-holes through a piece of paper at a distance of rather less 
than 23 inches from each other. On holding them horizontally 
before the eyes, and looking through the left aperture with the 
left eye at a distant object, the two circular images vary in their 
apparent relative position according to the distance of the paper 
from the eyes in such a way as to demonstrate the presence of 
relative convergence. More need not be said about this, since 
the camera acts upon the same principle.? 


Exp. 26.—The central and right lateral apertures? are used as in 
fig. 6, the stop being to the right. 

The observer, instead of looking 
at the central aperture (£), as in 
testing for near vision, now looks 
through it at any very distant ob- 
ject, and while doing so moves the 
right slide till its aperture appears 
to lie just below the image of the 
central one. 

If then the distance between the 
actual apertures be measured, they 
will be found separated by an in- 
terval rather Jess than the distance 
between the centres of the two 
eyes. Now, since the left eye is aaa 
looking directly at the object, the Bien ibn Hs pe ay 
image of the object, as wellas that 4 should be pone pe eee 
of the aperture which encircles it with ¥’.) ae 


1 A notice of this experiment was kindly communicated for me by Prof. Crum 
Brown to the Proc. Roy. Soc. Edin. in 1883-1884. 

* At the time when the figure was made I used the right or higher lateral aperture. 
I now use the left or lower one to eliminate more completely the desire for fusion. 

VOL. XX. 2P 


566 MR ERNEST E. MADDOX. 


as in a frame, must fall on the left fovea centralis or point of acutest 
vision. The encircled image therefore is referred—where all foveal 
images are referred—to the line which bisects the angle of convergence. 
But the other aperture has been placed so that its image appears to be in 
the same line, or rather slightly below it; it therefore must fall exactly 
above the right fovea, on the median vertical meridian of the right 
retina. Since each image falls on a median vertical meridian, it 
follows that if the apertures themselves were separated by an interval 
equal to the intercentral distance, the visual axes would be parallel ; 
if the interval were greater there would be relative divergence, but 
as it is, the interval is less, showing relative convergence. 

Moreover, if, while the apertures are still kept in position, both eyes 
be made to observe distant objects through them, 
the images none the less appear superimposed ; the 
amount of convergence attached to distant vision re- 
mains unaltered, whether one eye or both is used.! 
Since the natural outflow of energy from the con- 
verging centre when the desire for fusion is absent is 
a delicate comparative index of the accommodating 
energy, this fact shows that the activity of the accom- 
modating centre is no greater when both eyes are 
used than when vision is confined to one, and corro- 
borates the statement that accommodation is the work 
of a single innervation affecting both eyes equally at 
the same time. The object thus seen by the right 
eye, through the lower aperture, is one which really 
lies in a space to the deft of the object seen by 
the left eye through the higher aperture. Thus, 
if in figs. 7 and 8, a and 0 are two 
distant objects, a is seen by the 
right eye throngh the lower 
aperture, and b by the left eye 
through the higher one. Fig. 
9 shows that for this to occur 
the visual axes must cross some- 
where between the camera and 
the distant objects. This cross- 
ing point is at anaverage distance 
of about 112 inches from my 
own eyes. 

Another glance at figs. 7 and 
8 will make it evident that for 
the same object (b) to be visible 
by both eyes, the lower of the 


Figs. 7 and 8.—Objects 
(a, b) seen through two apertures must be drawn 


the apertures of the away from its apparent position just under the 
camera. other to the right. Let this be done till “0” is 
visible in both apertures, the actual distance between them will 


Fig. 9. 


This presumes the possession of eyes of equal refraction, 


CONVERGENCE AND ACCOMMODATION OF THE EYES. 567 


then be the same as the distance between the centres of the two 
eyes, while the apparent distance between them will indicate the 
degree of relative convergence present. To measure it, we need only 
take the difference between the number of degrees now recorded and 
that previously recorded when the lower aperture appeared just under 
the other. In taking these observations on others it is essential, to 
obtain accurate results, that the arms should be supported ; or better 
still, that the camera itself should be hinged (as mine is) on a stand, 
so that by telescopic action it can be raised or lowered to any required 
height, or inclined at any angle. It is difficult otherwise to maintain 
the requisite steadiness, and the arms get tired before the observation 
is complete. 


The amount of Relative Convergence with negative accommoda- 
tion.—This varies in different individuals, being apparently much 
greater in some than in others. In my own case fifty observa- 
tions of the amount of relative convergence, associated with very 
distant vision, gave the average of 1° 18’ 43”. By more than 
neutralising my half dioptre of manifest! hypermetropia I have 
never succeeded in reducing the convergence lower than 28’ 30”. 
For a hypermetrope such convergence excites no surprise; it 
would indeed be looked for, since the ciliary muscle, unlike that 
of an emmetrope, is never quite relaxed in distant vision, and a 
certain amount of attached converging effort might therefore 
well be expected to cling to it. But in fact the convergence 
noted is less in degree than with most of the emmetropic eyes I 
have tested. The unexpected feature is the comparative small- 
ness of it, in the presence of hypermetropia. Were the connec- 
tion between the two efforts as complete as it was once thought 
to be, as much as three and a half degrees of convergence would 
accompany each dioptre of hypermetropia. 

Dr Bolton in six cases, without known hypermetropia, ob- 
tained an average inward deviation of 1° 38’, and I found that of 
a similar number of records to be 2° 16’ 14”. The variations in 
these twelve persons were from 0° to 4°. Large relative conver- 
gence in distant vision appears to be generally associated with 
small relative convergence in near vision, and vice versd, though 
not without frequent and sometimes striking exceptions. In 
one instance slight divergence was found by Dr Bolton with 
negative accommodation, and in another, relative convergence 


1 “Manifest” lhypermetropia is that which is discoverable without the use of 
atropine, while the remainder is latent. 


568 MR ERNEST E. MADDOX. 


noted for some time gave way, after a great nervous expenditure, 
to parallelism.or even slight divergence of the visual axes, which 
continued for several days. I find that in the absence of excep- 
tional causes of disturbance the amount of relative convergence 
attached to negative accommodation goes through a fairly 
uniform variation through each day, becoming greater as the day 
advances, but suffering a fall after each meal, though more 
especially after the principal midday one. The average A.M. 
record was 1° 5’12”,and the P.M. record 1° 24’ 33”, while the after- 
dinner one was 1° 3’ 4”, The relative divergence in near vision 
(at 10 inches) diminishes through the day, though by far the 
greatest fall occurs during the first hour after rising. The con- 
vergence attached to negative accommodation and that attached 
to great positive accommodation do not, however, rise and fall 
together, for the relative divergence in the latter as noticed by 
the camera is lessened shortly after a meal, especially after the 
midday one, showing that the attached convergence is increased 
at the same time that that of negative accommodation is dimin- 
ished. The average deviation, with vision for 10 inches, before 
the eyes were otherwise opened in the morning, was 7° 36. In 
this I have not included an exceptional record of about 3° 35’ 
after disturbed sleep, consequent on the uncustomary taking of 
a supper on the previous night. It is known how irritation of 
the prime vie may cause temporary nervous strabismus in 
children ; this record is so interesting, as showing how the same 
condition which causes a pathological effect at one age may at a 
later one cause only an unnoticed effect on the physiological 
condition of the centres, that I give here the observations of 
that day, and the one before and after it. 


| | 
7.15 A.M. | 7.55 A.M. 


8.20 A.M.| 9 A.M. 10 A.M. 
Hirst daisy ¢-.\cu he 7° 45’ “y, 6° we (ae 
Second day,. . . 3° 35! mee 4° 15’ 5° 46’ 6° 26’ 28” 
hirdiday,*; '% SQ VOT SFT | My Mi ate 


It may be noted that while the records of the first and third 


' Owing to an imperfect marking of the camera these records are all slightly 
too large, though uniformly so; they serve for comparison only. The one marked 
* was earlier than the time given, 


er 


CONVERGENCE AND ACCOMMODATION OF THE EYES. 569 


days diminish as usual, those of the second day gradually increase 
to the usual amount. Before attempting to estimate the effect 
of drugs and different conditions on the brain centres, it would 
be well for the observer to become acquainted with his owa 
diurnal variations. 

Point of Coincidence—When an object is moved along that 
horizontal line of the sagittal plane which is level with tbe 
pupils, and looked at with one eye, there is a certain distance at 
which the meeting-place of the two visual axes coincides with 
the object. I name it the “point of coincidence,” since here 
the attached convergence coincides exactly with the accommo- 
dation to which it is attached. At this point there is neither 
relative convergence nor divergence ; when the object is beyond 
it, there is the former; when within it, the latter. In some it 
might be rather a region than a point. Were the object moved 
along different radiate lines in the horizontal plane with direct 
oblique vision, the points so discovered would constitute a 
horizontal “line of coincidence,” and similarly a vertical line of 
coincidence could be found. 

My own point of coincidence in the line first mentioned, 
which runs straight forward from the root of the nose, has varied 
from 56 inches to 14 inches distant. As might be expected from 
the fuctuations in the amount of deviation attached to near and 
distant vision, it shifts from minute to minute, according, doubt- 
less, to the occupation of the eyes as well as the conditions ot 
the nervous system. 


The camera furnishes the most exact method of finding the dis- 
tance of the point of coincidence, but the double prism is a much 
more ready weans. With the former an object is approached till it 
is visible in the central and one of the lateral apertures at the same time 
that the apertures themselves appear superimposed ; or, better still, a 
card coloured red to one side of a vertical line and green to the other, 
and provided with printed matter to insure exact accommodation, is 
carried on a traveller of a strip of wood hinged on to the stand of the 
camera. The left brass slide of the camera is withdrawn more than 
half its own length, so as to leave a short stationary s/i¢t instead of an 
aperture. With the stop to the left, place the card so that the median 
line on it is visible through the central aperture by the right eye; it 
is then made to approach or recede till the lines seen in the slit and 
the aperture appear continuous. This method permits of great nicety 
as well as speed. It is very important to insure that the eyes are 
accurately accommodated for the object. When this precaution is 


570 MR ERNEST E. MADDOX. 


taken, the variations are far less. Strangely enough, I have uniformly 


found the distance of the point of coincidence greater after very near _ 


vision of fine print, and less after distant vision. Another method I 
have employed is to use a long graduated strip of wood, at one end of 
which the double prism! can be fixed at pleasure; it rests on the 
mantelpiece or any convenient horizontal situation, and a night-light 
or other appropriate object is moved along it till the three images are 
exactly in a line. Objects should be avoided which have vertical 
edges if they are long enough to overlap. For self-trial a wooden 
traveller is used, which can be pulled backwards and forwards by a 
piece of string. This is the easiest expedient, but is not so exact or 
trustworthy as that with the camera. 

When diplopia is heteronymous, it shows the point of coincidence 
is beyond the object, which must therefore be made to recede, and 
conversely to approach when diplopia is homonymous. It is easy to 
see the bearing of this point on the well-known clinical test, in which 
a flame at some distance serves for the object of view and a prism 
(base vertical) is held before one eye; of the two images which appear, 
one passes to the right or left of the other, according to the existing 
relationship between accommodation and convergence. The distance 
of the flame trom the observer has been thought a point of little con- 
sequence, but the whole result depends upon it. If the distance is 
less than that of the point of coincidence, relative divergence will 
appear, while if greater, relative convergence will betray itself; pro- 
vided, of course, that the prism is held absolutely vertical. If neither 
deviation is observed, the flame is exactly at the point in question ; 
but if this happened with the flame at the usual distance of 7 or 8 
feet, there would almost certainly be observed a wide deviation out- 
wards if near vision were tested. Since the distance of the point of 
coincidence is generally less than that of the flame, slight relative 
convergence would be almost uniformly detected were the method 
sufficiently accurate, though the prismatic errors liable to occur in this 
test are far less than in those which require prisms of higher powers. 
The smaller the refracting angle of a prism, the smaller is the degree 
of accidental displacement of the image which attends any rotation of 
the prism from the vertical. 

Seven cases examined by Dr Bolton showed an average of 43 inches 
for the distance of the point of coincidence. In one case it was 97 
inches, about the usual distance of the flame in the clinical test spoken 
of, the result of which, therefore, in this case would have been nega- 
tive, though with vision for 10 inches there was a deviation outwards 
of 7° recorded by the camera. Since the average distance of the point 
is less than that of the flame, relative convergence would generally 
show itself if the test were carried out with sufficient accuracy. There 
might be great outward deviation in near vision, or inward deviation 
in distant vision, or both, and yet neither be detected by the test as 
usually employed. 


1 The prism used is of a square outline, which gives no trouble in adjusting to 
the vertical, since one side rests on the wood. 


CONVERGENCE AND ACCOMMODATION OF THE EYES. 571. 


The point of coincidence is of interest, because whenever vision 
of daily life wanders beyond it, the fusion effort, instead of acting 
through the converging apparatus, acts through some diverging 
mechanism, which is doubtless connected with the external recti, 
though it is just possible that it acts by inhibition of the converging 
centre. The nearer the point of coincidence is to the eyes the more 
fusion work falls through the day to the diverging mechanism and the 
less to the converging mechanism, and vice versd, so that its position 
is what determines the “division of labour” for these two innerva- 
tions in their connection with the desire for single vision. 

I may add that when an object is seen just within the ‘‘ near point” 
with the double prism before one eye, relative divergence may for a 
moment disappear, or even give way to relative convergence, so great 
is the fruitless effort to accommodate more. The effort shows itself 
by the sympathetic effect on the converging centre—a good illustra- 
tion of the fact that it is not accommodation and convergence that are 
united, but the nervous efforts in the ganglia of the 3rd nerve which 
cause them. For this experiment care must be taken to avoid 
obliquity of vision or to allow for its presence. 


If two horizontal slits are cut in a piece of paper or card- 
board, so that the inner end of each terminates in the vertical 
middle line of the card, and yet so that one is on a level about 
Linch higher than the other, the point of coincidence can be 
found in a very simple and inexpensive way, without either 
prism or camera, by looking through the slits at any vertical 
linear object till it appears continuous in each. It is not very 
trustworthy, however, in a patient’s hand, since obliquity may 
cause fallacy.t 

The Three Grades of Convergence—We may now distinguish 
the three elements of that complete convergence which occurs 
with binocular vision of a near object. 


12 Initial Convergence—which is that attached to negative 
accommodation, 

2. Accommodative Convergence—which is that, in addition, 
attached to positive accommodation. 

3, The Fusion Supplement—which is that excited by the 
desire for fusion to make up, when it can, for the 
deficit (or excess) of the other two. 

1 Two horizontal stenopaic slits at different levels, placed in the ordinary trial- 
frame, would act on the same principle. 
? It is convenient to use contractions for these—I.C., A.C., and F.S. respect- 


ively; also, R.C. for relative convergence, R.D. for relative divergence, and 
P.C. for the point of coincidence. 


572 MR ERNEST E. MADDOX. 


Fig. 10 illustrates these three factors in looking at a point O. 


The first brings the visual axes from parallelism pp to w, the 


second to aa, and the third to O. Were O at the point of 
coincidence it would coincide with 
aa, and the third factor would be 
engaged only in steadying the eyes 
by counteracting the tiny inequalities 
in the evolution of that nervous 
energy which maintains the other 
two. For objects beyond the point 
of coincidence there are only the 
first two factors in convergence in 
Fic, 10.—The three elements of eXercise, the fusion supplement being 
eens OF. done 0. binocular employed no longer in augmenting 
but in partly neutralising their effect. 
The distinction between the initial and accommodative converg- 
ence is important, because they are unequally affected by dif 
ferent conditions. Thus we have seen that the effect of a meal 
is to diminish the former and increase the latter so much, that it 
more than makes up for the initial diminution. Initial con- 
vergence is affected directly by the sympathetic system through 
the external recti, while the accommodative convergence is only 
affected by changes in the cerebral centres of the 3rd nerve when 
vision is near, and the 3rd and 6th when vision is more remote. 
We have already seen how to measure the initial convergence 
and the relative divergence in near vision—the difference be- 
tween the two gives the accommodative convergence. 

Squints may be analysed into their initial and accommodative 
elements by careful measurement, first with negative accom- 
modation, and then with positive, by the direct or blind spot 
method, as required. 

The accommodative convergence can itself be analysed by 
finding how much of it belongs to each dioptre of accommoda- 
tion. To do this objects are viewed at 1m., } m.,4 m., and $m. 
distance in turn; but an average of a great many experiments 
must be made to get results of any value. Dr Bolton’s experi- 
ments! show that the convergence attached to the /ir'st dioptre is 
the greatest in his case, being 3° 18’ 27”. To the 2nd D was 

' Kindly made at request, February 1886. 


CONVERGENCE AND ACCOMMODATION OF THE EYES. 575 


attached only 45’ 18”; to the 5rd D—2° 54’; and to the 4th D— 
2° 20’. The excess of the first over the others fully accords with 
the fact that the point of greatest R.C. is not at practival in- 
finity. Objects at the same distances give in my own case 
1°53’ 42”—1° 13’ 87’”—1° 41’ 40”—and 2° 44’ 24” of accommoda- 
tive convergence for these four dioptres respectively. The first 
is greater than the next two, the second is, as with Dr Bolton, 
the least, but the fourth is the greatest of all, owing doubtless 
to the greater effort needed for this dioptre than for the previous 
ones, just as a man must put forth more effort in raising a 
weight through the fourth foot from the ground than through 
the first foot owing to increasing mechanical disadvantages. <A 
dioptre is a unit of work, not of effort, and since in a hyperme- 
trope these four dioptres are not the /irst four, the effort required 
for each begins to increase more rapidly than in an emmetrope. 
No doubt each succeeding dioptre, and each succeeding degree 
of convergence, needs more effort than the preceding one, but the 
resistance experienced by one effort may increase more rapidly 
than that experienced by the other, and the amount of converg- 
_ence attached to each unit of accommodation varies accordingly 
—at least in near vision uncomplicated by any diverging effort. 
In testing the ¢rue initial convergence of a hypermetrope, the 
compensating lenses worn must have their optical centres separ- 
ated by the exact intercentral distance of the observer, to avoid 
their prismatic fallaey—even then an error exists, proportionate 
to the strength of the lenses and the degree of initial converg- 
ence, for which a subsequent formula gives the correction. I 
prefer, to avoid this, to place a single lens of 94 inches greater 
focal length before the central aperture, or else before the slit 
made by largely withdrawing the left brass slide, the end of 
which can then be placed apparently under the central aperture. 

The prismatic action of such a lens introduces no error at all. 
The farther a lens is from the eyes the more it relieves accom- 
modation, because its focus is brought by so much the nearer to 
the retina; hence a weak lens at a distance has the same effect 
as a more proximate and stronger one. For this reason old 
people sometimes prefer to wear their spectacles on the tip of 
the nose. 

Since spectacles are almost never worn nearer than half an 


574 MR ERNEST £. MADDOX. 


inch from the dioptric centre their power does not exactly ex- 


press the relief they give to the accommodation—half an inch- 


at least must be substracted from their focal length to do this. 
Hypermetropia therefore is always slightly greater than the trial 
lenses indicate, and, conversely, myopia is always less. But this 
is a digression—it is to illustrate why a lens at the base of the 
camera must be weaker than one before the eyes. I have as yet 
by such lenses only been able to reduce my initial convergence, 
and have never obtained complete parallelism of the axes. 

Point of greatest relative Convergence—In glancing with one 
eye at objects intermediate between practical infinity and the 
point of coincidence I had thought that the amount of relative 
convergence diminished uniformily with each approach of the 
point of view, until at the point of coincidence it gave way to 
commencing relative divergence. But Dr J. 8. Bolton dis- 
covered that in his own case the relative convergence at first 
increased as vision became less remote, reaching its maximum 
when an object is viewed at 5 metres; within which it gradually 
diminished up to the point of coincidence. His “ initial conver- 
gence” was 1° 10’; relative convergence increased, with accommo- 
dation for 8 metres, to 1° 44’; for 6 m, to 2°; and for 5 m. to 2° 12’ 
36”. It was here therefore nearly twice as great as the initial 
convergence. At 4 m. it fell again to 2°; at 2 m. to 1° 42’, soit 
was still greater than the initial convergence; at 1 m. to 1°; then 
came the point of coincidence at the distance of ‘85 m. (34 in.). 

At half a metre (20 in.) there was relative divergence of 1° 24’; 
and at the distance of the apertures of the camera (‘25 m.) the 
divergence reached 6°. On testing another subject he found 
“the point of greatest relative convergence” to be 6 m. from the 
eyes. In this case the increase was not so marked, the “initial” 
convergence being 1° 5’ 24", while at 6 m. the record gives only 
1° 45° 36"., At 2 m, the relative convergence was still greater 
than the initial, being 1° 18’. The point of coincidence was at 
1 m.; and the relative divergence, though being 2° 15’ 36” at 
half a metre, was at the distance of the camera 6°. In these 
two cases the conditions with vision for 10 inches were therefore 
the same, but the smaller initial convergence in the latter seemed 
to throw the two points of “ coincidence” and of “maximum re- 
lative convergence” to a slightly greater distance. 


4 
* 


CONVERGENCE AND ACCOMMODATION OF THE EYES. 575 


The results in my own case vary too much to be of any value 
in regard to this phenomenon, the relative convergence at 
8 m. being sometimes more and sometimes less than the initial 
convergence; but since the refraction is abnormal, this is no 
guide to the physiological conditions. 

To summarise: If we look at accommodation and the con- 
vergence attached to it as two racers beginning from infinity, 
convergence has the first start, and gains still more up to the 
point mentioned by Dr Bolton; it then slowly falls behind, till 
at the point of coincidence the two are abreast; it then con- 
tinues to fall behind, and thus creates the relative divergence of 
near vision; but just at the near point—the end of the race—it 
again gains ground, and comes in abreast or even in front of 
accoinmodation. 

The Point of Repose.—It has been known for some time that 
the visual axes are not parallel when the eyes are completely at 
rest, z.¢., without being engaged in the vision of any special 
object, or affected by any act of volition;! but it is difficult to 
determine at what point the axes meet. It is not the point of 
initial convergence, for that is attached to negative accommoda- 
tion, And it is well known that the ciliary muscle can by very 
few, if by any, be completely relaxed without the aid of a distant 
object to excite the desire for distinct vision ; so that in a state 
of rest there is some positive accommodation, which cannot exist 
without being accompanied by a due proportion of attached con- 
vergence of the visual axes in addition to the initial convergence. 

The natural position of eyes in the dark I have tried to de- 
termine by the following experiment. 

Exp. 27.—Use the central and left lateral apertures of the camera, 
with the stop to the left; but in looking into it cover the apertures 
outside with a black pad, so as to produce the impression that the 
eyes are looking into absolute darkness. A momentary displace- 
ment of the pad (a penwiper does well) reveals the two apertures and 
covers them again before there is any attempt to accommodate for 
them. The left slide can be moved by repeated trials till, on moving 
the pad, the images appear in the same vertical line. The real dis- 
tance between them then, of course, records the separation of the 


visual lines at the distance of the base of the camera. The records 
vary in my own case from 5° to 8° 30,! being generally only slightly 


1 Mr Brudenell Carter has expressed his belief that ‘‘in states of rest the 
internal recti possess the physiological preponderance over the externi which 


576 MR ERNEST E. MADDOX. 


different from the disposition of the axes when one eye is occupied in 


vision at 10 inches. This is somewhat unaccountable, since the radiate 


appearance of the apertures shows that accommodation does not occur 
for them.! The average of observations taken on nine different occa- 
sions was 6° 26’ 16”, while that of the relative divergence at 10 inches 
on the same occasions was 6° 49’ 36”, so that the visual axes would meet 
at the distances of 184 and 194 inches from the eyes respectively. 
Many of the observations were taken in the morning before the eyes 
were otherwise opened; but though the point was at that time rather 
more remote, the difference was not greater than that of the natural 
relative divergence in near vision at the same period. It is difficult 
to believe that so near a point of meeting of the visual axes should be 
the real position of rest. It may be that a psychical effect is pro- 
duced by looking into blank darkness, since near objects are the ones 
that would be instinctively looked for in such a state. Still, on 
closing the eyes and then opening them suddenly, the apparent posi- 
tion of the apertures only confirms the results. When the light was 
so reduced that the mind could only detect the dimmest indications 
of apertures, they appeared together when really separated by 10°, so 
that the visual axes did not meet until nearly 34 inches from the eyes. 
On looking at a dull surface without attempt to accommodate, the 
results were intermediate between those of the last two methods 
(8° 15’). The position taken by the eyes of a patient under chloroform 
is not a very trustworthy indication of the position of rest ; the effect 
on the pupils points to probable disturbance of other ocular centres. 
That the external rectus is partly supplied by the sympathetic is an 
interesting fact, as showing to that extent an antagonism between the 
sympathetic and the 3rd nerve in the ocular movements we are now 
considering, analogous to their antagonism in the movements of the 
pupil ; chloroform would he likely to affect them in a similar way. 
In one case I noticed marked external strabismus under chloroform, 
which continued no longer than the anesthesia.? The eyes made con- 
stant tiny oscillations from side to side, and the margin of the cornea 
was } inch above that of the lower lid. The pupil at first widely dilated, 
but shortly after the disappearance of the conjunctival reflex it began 


flexor muscles commonly possess over extensors, and the eyes during sleep or 
anesthesia are just a little convergent, in the same way as the limbs are slightly 
flexed.” ‘‘ This preponderance,” he says, ‘‘is probably increased by the fact that 
in the conditions of civilised life the externi are less used than the interni 

1 For these experiments, as indeed for all, a piece of black silk depends from 
the lower margin of each wooden slide at the narrow end of the camera, to ex- 
clude vision of other objects. 

* To give full attention to the respiration, and yet have one hand free for the eyes, 
it is useful in administering chloroform to women to hold the left end of the towel 
between the thumb and the forefinger of the left hand, so that the tips of the middle 
and ring fingers rest continuously against the clavicle, and appreciate, the former 
the rolling movement of its anterior surface against the side of the last phalanx, 
and the latter its npward movement against the tip. The right hand is then free, 
when not adjusting the towel, to open both eyelids by its first and little fingers. 


CONVERGENCE AND ACCOMMODATION OF THE EYES, 577 


gradually to contract, taking a long time to get to the size of a pin’s 
head. On the removal of the chloroform it remained thus for some 
time, then suddenly dilated nearly to its full, this preceding the return 
of the conjunctival reflex. On reapplying the chloroform it slowly 
began to contract again, taking as before a long time to reach its 
minimum size. This was its uniform behaviour during the two hours’ 
administration for an intra-peritoneal operation. The strabismus may 
have been reflex from the abdomen. 


The intercentral distance—There are many methods of finding 
the distance between the centres of motion of the two eyes. (a) 
The simplest and best is to look at a linear vertical object at a 
ereat distance through the camera, moving one of the slides till 
the object appears to bisect the central and one of the lateral 
apertures, as attention is directed to each in turn. The distance 
between the apertures gives the intercentral distance ;! while the 
number of degrees recorded between them expresses the exact 
magnitude of the “optic angle” (z.e., the angle included between 
the visual axes) of the person tested when he looks with both 
eyes at the central aperture; since it is through the “alternate ” 
angle that the one visual axis must rotate out from that point 
in order to become parallel with the other (Euclid, I. 29). This 
angular record is the one most convenient to preserve, since to 
obtain the “initial convergence” at any time it is only necessary 
to deduct from it the record given when the simple observation 
is taken of looking at a distant object through one aperture and 
placing the other apparently beneath it. To convert the angular 
dimensions into linear ones, it suffices to know the linear equiva- 
lent of each degree. It is ‘1789 inch, with vision for 10 inches. 
From this a table may be made. 


Table of the Linear Value of Degrees at the Base of the Camera. 


Inches. mm. Inches. mm. Inches. mm. 

For 21468 54:468 13° 30’ 2°4151 61°276 14° 50’ 2°6537 67°329 
12°10’ 2°1766 55:224 13° 40’ 2°4450 62:°033 15° 2°6835 68°085 
12° 20’ 2°2064 55°981 14° 2°4748 62°789 15° 10’ 2°7133 - 68-841 
12° 30’ 2:2362 56°737 14° 2°5046 63°546 15° 20’ 2°7431 69°598 
12° 40’ 2°2661 57°494 14°10’ 2°5344 64°303 15° 30’ 2°7729 70°353 
12° 50’ 2°2959 58:25 14° 20’ 2°5642 65:°059 15° 40’ 2°8027 71°11 

1 2°3257 59:007 14° 30’ 2°5940 65°816 15° 50’ 2°8326 71-867 
13° 10’ 2°3555 59°763 14° 40’ 2°6239 66°572 16° 2°8624 72°623 
13° 20’ 2°3853 60°52 


1 The measurement being partly on a curve, there is a slight error of ‘004 inch. 
It may be avoided where great accuracy is required by using the left aperture 7° 
away from the middle line instead of the central one, 


578 MR ERNEST E. MADDOX. 


From this table, if either the intercentral distance or the 
optic angle in binocular vision for 10 inches be known, the 
other may be found. The radius is 10:25 inches, since the 
centre of motion is behind the optical centre. 

(b) It is not always easy to find a suitable linear object at 
sufficient distance. If the camera is mounted on a stand it can, 
after adjusting the central aperture for any distant object, receive 
a slight tilt to bring the other aperture to the same level. The 
tilt would always be through the same angle, and can with my 
own stand be mechanically provided for when this method is 
employed. 

(c) In practice, however, the object is far more convenient 
within the room. To open a window is not advisable always, 
and to look through it may introduce some prismatic error from 
the panes. Any linear object may be viewed—if at 17 ft. 1 inch 
from the base of the camera the distance between the apertures 
must be increased by 315th, if at half that distance by 7th. I 
now use a card marked with a vertical strip of white, inch 
wide, down the middle, to the right of which is all coloured red, 
and to the left green. After adjusting the camera so that the 
white strip is visible in the central aperture with an equal band 
of colour on each side, it is easy to know whether to push the 
lateral aperture in or out by whether the red or green colour is 
that seen through it. This expedient greatly compensates for 
want of intelligence in the patient. The one source of fallacy 
in all methods with a near object springs from the varying dis- 
tance in different patients between the cornez and the apparatus; 
but the greater the distance of the object the less the error. The 
card can either be connected to the base of the camera by a 
string of the required length, or it can be placed on the strip of 
wood already mentioned at the distance of 41 inches from the eyes; 
the latter is now my usual practice, being much the handiest 
and quickest indoor method. The record must be increased by 
adding a third as much again. An object nearer than this is not 
advisable. In testing for the relative convergence or divergence, 
when accommodation is exerted for objects nearer than practical 
infinity, it is useful to know how many marked degrees at the 
base of the box would be intercepted between the visual axes, 
were they both directed accurately at the object viewed. To 


CONVERGENCE AND ACCOMMODATION OF THE EYES. 579 


find the amount of deviation it is then only necessary to deduct 
from this the degrees recorded when, in looking at the object 
through one aperture, the other is placed apparently beneath it. 
The separation of the visual lines at the distance of the base of 
the camera in binocular vision of any object is given by the 
ixd 
cepted degrees, 7 is the intercentral distance, and d the dis- 
tance in inches of the object from the eyes. 


formula z= where zis the required number of inter- 


VI. Direct Oblique Vision. 


So far we have considered the relation between convergence 
and accommodation apart from any influence which may be 
exerted upon either by one of those innervations which turns 
both eyes to the right or left. Whenever the point at which 
the visual axes meet lies to one or other side of the median 
plane, a new innervation comes into play. Does the interven- 
tion of this new effort in any way disturb or alter the relation- 
ship we have been considering? We can only judge of efforts 
by their results, and these results are in oblique vision compli- 
cated by the fact that accommodation and convergence are 
required in differing proportions when an object is viewed with 
different degrees of obliquity. This, indeed, is in itself ample 
reason why the two are not so inflexibly bound together as 
they were once thought to be, and as the two accommodating 
mechanisms actually are. Complete central connection might 
at first sight appear the simplest conception. But in daily life 
the eyes need constantly to wander from side to side, and as 
they do so, accommodation and ccnvergence are required in as 
constantly differing proportions, which make a certain “play” 
between the two innervations a necessity, and for which pro- 
vision is made by superadded functions which can reflexly 
increase or diminish either as required. But might not these 
have been done without, by such an adaptation of the central 
connections that each ranging effort would in a direct way pro- 
duce the required modification in the others? No, for up to a 
certain geometrical degree of obliquity accommodation needs to 
be relatively increased, while beyond it convergence needs to be 


580 MR ERNEST E. MADDOX. 


increased. Why, then, if to attain this play central connection 
must in part be overcome, is there any such connection at all ?. 
It is a great saving of the reflex fusion-effort. 

It is evident that geometrical considerations must precede the 
physiological study of the subject of oblique vision. 


(a) Accommodation.—Apart from any connection with con- 
vergence, disproportion between the accommodative requirements 
of the two eyes is brought about by the slightest deviation of 
the point of fixation 
from the median plane. 
Fig. 11 illustrates this 
when any flat object is 
looked at, as in reading 
abook. The prolonga- 
tions of the visual lines 
on the distal side of 
the line AB _ repre- 
sent the disproportion. 

Fig. 11. Thus, when both eyes 
are looking at “w,” the object is nearer to the right eye than to 
the left by the distance tw, and so on. Every departure of the 
point of fixation from the middle line lessens the required 
accommodation in the opposite eye, while at first it increases 
that of the eye of the same side till the fixation point has tra- 
versed a distance op equal to half the intercentral distance, 
after which it falls through a similar interval pq to the original 
amount, and then continuously diminishes. But the centres 
for accommodation are so intimately connected that one eye 
cannot accommodate more than the other. When variations, 
therefore, exist either in the refractive power or requirements of 
the two eyes, “that eye has the bright image which attains it 
most easily at the expense of the other” (Donders). They do 
not split the difference ; if they did so there would be diffusion 
circles in each eye. Since accommodation with normal refrac- 
tion implies positive effort, that eye which is farthest from the 
object, and can see it with least effort, determines the accommo- 
dation for both. Fig. 12 therefore represents “the line of equal 
accommodation” for near vision, in whatever point of which the 


CONVERGENCE AND ACCOMMODATION OF THE EYES, 581 


object is placed accommodation remains the same. It is com- 
posed of two arcs of equal radius, described from the centres of 
their opposite eyes. 
In hypermetropiathe 
line is similar; the 
reasons for it being 
so are intensified, 
since accommodation 
is a greater effort ; 
but in myopia accom- 
modation is negative outside the line which limits the far point, 
and which is made by drawing each are from the centre of the 
eye of the same side. Since in these cases relaxation is often 
attended with more effort than accommodation (owing to spasm 
of the ciliary muscle), the “line of equal convergence” would 
be of the same shape as the “far point line” for some distance 
within it.! 

(b) Convergence.—Fig. 11 shows that convergence, as well as 
accommodation, diminishes with oblique vision; but that they 
do not diminish equally is made clear by the fact that the line 
of equal convergence (fig. 12) is not of the same shape as that 
of equal accommodation (fig. 13).2. The former is part of a circle 
which passes through the centres of motion of the two eyes (not 
like the horopter through the dioptric centre), and -possesses 
these properties. (1) The angle of convergence is the same 
whatever point in it is made the point of binocular fixation. 
(2) In glancing from any one point in it to any other, both 
visual axes traverse equal angles; thus in fig. 14 the angles OBe 
and OAc are equal; while, in contrast to this, fig. 11 shows that 
in glancing from O to @q or O to »p, the left axis passes through 
a greater angle than the right; as it does, indeed, whatever point 
is looked at to the left in the straight line AB. (3) The line 


Fig. 12. Fig. 13. 


1 While what has been said holds good in normal vision, since attention is 
seldom directed to one side for long at a time, a fallacy must be guarded against 
in experiments which require sustained obliquity of the visual axes ; the visual 
apparatus of the farthest eye might weary, and for a time permit the accommo- 
dation of both eyes to be governed by that of the nearest one, and thus be 
increased. 

2 These were called in the original thesis the ‘‘Isostigmal” and ‘Isogonal ” 
lines respectively, but I do not see the necessity of naming them. 


VOL, XX. 2Q 


582 MR ERNEST E. MADDOX. 


which bisects the angle of convergence is the one to which 
hypothetically objects upon the maculz should be mentally 
referred. Whether, in fact, 
they are so physiologically 
remains to be proved. The 
line is obtained by uniting 
the point of binocular fixa- 
tion (c, in fig. 14) to the 
posterior point of the circle 
(b). The position of this 
posterior point shifts, of 


\ course, with every variation 
aS a in the size of the circle; 

b the line cd itself is inclined 
to the median plane by an 
angle which measures the 
obliquity of vision, since it is equal to the angle which each 
visual axis has traversed in looking from the anterior point of 
the circle (0) to any other point in it (c). This angle, therefore, 
represents the “ranging” work to be done in looking at any 
such point. I do not say the ranging effort, for, as before men- 
tioned, effort is often disproportionate to work, owing to greater 
resistance or other disadvantages. A certain evolution of nervous 
energy from the converging centre produces a definite angular 
deflection inwards of both visual axes, and similarly an effort of 
a ranging centre produces a definite deviation of both visual axes 
to the right or left. The nervous impulses perform angular 
work, if I may so say, as far as vision is concerned, and there- 
fore we may assume that it is by angles that the mind judges of 
the work done in estimating the projection of the field of vision ; 
but since the judgment is based solely upon the effort put forth, 
any discrepancy between effort and work would show itself in 
angular misjudgment, unless by habit the mind had come to 
associate a certain degree of effort with the work it wswally per- 
forms, instead of with the work it should perform compared with 
a smaller effort. Such allowance is no doubt made, in whole or 
in part, except for unusual obliquities. Were “effort” and 
“work” exactly proportionate in both the converging and 
ranging innervations, all objects seen by the fovea of one eye or 


Fig. 14. 


CONVERGENCE AND ACCOMMODATION OF THE EYES. 583 
both, however obliquely, would be referred to the line which 
bisects the angle of convergence, since its inclination to the 
median plane would exactly express the angular impression in 
the mind produced by the ranging effort. I find, however, that 
if a large piece of card be held obliquely a few inches outside 
and in front of either eye, with a mark upon its upper border, 
looked at with both eyes, a finger passed up behind it generally 
at first misses the mark to the outside by nearly an inch, showing 
that the ranging effort is relatively so far greater than the work 
it accomplishes that the mind estimates as if more angular work 
were done, and mentally displaces the object from the median 
plane by an angle greater than that of the line which bisects the 
angle of convergence, and which only measures the work actually 
accomplished, not the effort put forth to accomplish it. 

In fig. 15 the dotted arcs represent the line of equal accorn- 
modation, so applied to that of equal convergence as to illustrate 
the fact already mentioned, that within a certain degree of obli- 
quity of vision the pro- 
portion of convergence 
to accommodation is 
ereater than in the me- 
dian plane, while for 
ereater obliquity the 
proportion is less. At 
the points dd, where 
the two lines intersect, 
the proportion between 
convergence and accom- 
modation is the same as 
at O; within these points 
convergence must be re- 
latively increased ; out- 
side of them relatively 
lessened. The distance 
of each d from O is always exactly equal to the intercentral 
distance of the observer. 

The same figure shows how, in looking obliquely at any point, 
the convergence and accommodation required may be compared 
with that in the median plane, All that is necessary for any 


584 CONVERGENCE AND ACCOMMODATION OF THE EYES. 


given point (c) is to describe a circle through it and through the 
centre of each eye, as in the figure, and from the centre of the 
farthest eye (B) to draw the are ex from ¢ to the median line. 
The circle is the line of equal convergence for that point, and the 
arc is part of the line of equal accommodation for the same. In 
binocular vision, therefore, of the point ¢, convergence must occur 
as if for O, and accommodation as if for z, while both eyes 
are deviated to the right through the equal angles OBe, oAec. 
The point d gives the base of the line which bisects the angle of 
convergence, and which is made by joining dc, while the inclina- 
tion of this line to the median plane expresses the angular 
ranging movement of the two eyes. Were the relation between 
convergence and accommodation inflexibly complete for objects 
in the middle line, there would be diplopia for any object out of 
the middle line, except at one point on each side, within which 
diplopia would be heteronymous from relative divergence, and 
without which it would be homonymous from relative con- 
vergence. The nervous ties, therefore, though strong enough to 
relieve fusion-effort, are not so strong but what they can be 
overcome to meet such requirements. 

The lines of equal convergence and accommodation, if rotated 
round the intercentral line as an axis, would describe surfaces of 
equal convergence and accommodation respectively.” 


1 This may be done by drawing from A and B straight lines at right angles to 
cA, cB respectively. From the point where they meet draw a straight line to ¢, 
the bisection of which gives the centre of the required circle. Or Euclid, iv. 5. 

2 (1) To find the line of equal convergence for any required angle of con- 
vergence—the radius of the required circle is found by dividing half the inter- 
central distance by the sine of the angle. (It may be found geometrically by 
Euclid, iii, 33.) (2) To find the angle of convergence when an object is viewed 
in the median plane at a given distance from the eyes—half the intercentral dis- 
tance, divided by the distance of the object from the eyes, gives the sine of half 
the angle. 


(To be continued.) 


SIX SPECIMENS OF SPINA BIFIDA WITH BONY PRO- 
JECTIONS FROM THE BODIES OF THE VERTE- 
BRZ INTO THE VERTEBRAL CANAL. By Professor 
Houmpury, F.RS. (PLates XVII, XVIII.) 


1. Tue first specimen was taken froma child who died a few days 
after birth. The sac, which was very thin, and covered the 
sacral and lumbar regions, had burst, and a bony prominence was 
felt in its middle. 

The vertebral canal is widely open below the 1st lumbar 
arch, the laminze of the four lower lumbar and the sacral 
vertebre being all directed horizontally outwards. The arches 
of the 11th and 12th dorsal vertebrie are directed normally, but 
are incomplete, the vertebral canal being at this part widely 
open. The arch of the 1st lumbar vertebra is, however, com- 
pleted by cartilage; and a curved bony process (or bar) ascends 
from, and has grown out of, the back of the body of the 2nd 
lumbar vertebra (rather, as the section, fig. 35, shows, from the 
intervertebral substance between the 1st and 2nd lumbar bodies), 
bisects the vertebral canal, and impinges upon the under edge of 
this cartilaginous arch. From the back of the bodies of the 3rd 
and 4th lumbar vertebre rises a second much larger osseo-carti- 
laginous process, bridging over and uniting these two bodies, 
and rising considerably above the level of the dorsal and lumbar 
arches. It forms one mass at its base, but is partially bifid 
towards its summit. It slants forwards, and its base partially 
extends over the body of the second lumbar vertebra. The 1st 
and 2nd sacral vertebre appear to be normal, with the excep- 
tion of the lateral direction of their arches; but the vertebrae 
below these are irregular and confused and curved to the left; 
and the cocecyx is represented only by a cartilaginous knob 
(fig. 36). The dorsal bodies and the arches, above the 11th, are 
normal, except that the spine is bent a little to the left, and the 
arches and transverse processes of the 9th, 10th, and 11th ver- 
tebree, where the curve is most marked, are, on the left side, 
approximated and anchylosed, and the extremities of their trans- 
verse processes are covered and united by a thin band of carti- 
lage (fig. 2, ¢). 


386 PROFESSOR HUMPHRY. 


A vertical section (fig. 3) from before backwards, in the median 


line, or in parts slightly to the right of it, shows the bodies of - 


the vertebree in regular serial order and in normal form. But 
the body of the 5rd lumbar vertebra is small, not being more 
than half its proper size, and that of the 4th is somewhat 
smaller than natural. Placed bebind these, separated from them 
and from one another by cartilage resembling that between the 
bodies of the vertebree, are two spheroidal spongy bony masses 
resembling vertebral bodies. That over the body of the 3rd 
lumbar vertebra, which appears stunted by it, is the larger. 
These form the base of the large projection into the spina bifida; 
and, connected with their posterior aspects by cartilage, are bony 
processes, somewhat denser, with foramina! between them, sug- 
gestive of rudimentary and misshapen vertebral arches, which 
form the summit of that projection. The cartilages intervening 
between the spongy bone-masses, and separating them from the 
vertebral bones in front of them, resemble the intervertebral 
cartilages, inasmuch as each is, like them, composed of two 
cartilage-plates appertaining to the adjacent bones with an 
intervening softer plate foreshadowing, apparently, an interver- 
tebral disc. Indeed, the section of the projection is very sug- 
gestive of its being formed by two supplementary vertebrie 
developed behind the bodies of the 5rd and 4th lumbar bodies, 
and growing into the vertebral canal. The azygos bony process, 
situated further forward, is seen to extend from the cartilage 
between the 1st and 2nd lumbar vertebral bodies to the carti- 
laginous arch of the 1st lumbar vertebra, and to be continuous 
with both these cartilages. 

The spinal cord and membranes passed down to the 1st lum- 
bar vertebra, retaining the normal condition to this point, and 
presenting no bulging at the vacant space in the arches of the 11th 
and 12th dorsal vertebrae. Beneath the arch of the 1st lumbar 
vertebra it apparently divided, passing on the sides of the mesial 
bony bar, and then expanding upon the under surface of the 
back part of the sac, as is usual in cases of spina bifida.2 It and 
the wall of the sac were quite free from the bony eminence, 


‘ These foramina did not, so far as I could discover, contain or transmit any 
neural elements. 
* See paper by me on spina bifida, this Jowrna/, vol. xx. p. 500. 


SIX SPECIMENS OF SPINA BIFIDA. 587 


which was smooth and covered by the dura mater and the lining 
of the sac. The nerves leaving the cord traversed the sac; 
being separate and free in it, on their way to the lumbar and 
sacral intervertebral foramina. One or two of them, however, 
were applied upon the sides of the large bony process, and 
connected themselves with, or traversed, the membrane cover- 
ing it. 

The peculiarities in this specimen are— 

First. The deficiencies in the vertebral arches above the spina 
bifida, and separated from it by the complete arch of the Ist 
lumbar vertebra, below which the spina bifida commenced, this 
deficiency not being attended by any abnormality in the cord or 
its membranes at this part. There was merely a deficiency in 
the growth of the arches, without that deficiency in the segmen- 
tation of the spinal cord and its coverings, which is usual, and 
is indeed a prime feature, in spina bifida. 

Secondly. The bony processes projecting from the bodies of the 
vertebre into the vertebral canal. Processes, similar to the 
upper one of the two in this specimen, passing from before 
backwards in the middle line, and bisecting the spinal cord, 
have occurred, and are mentioned in the Report of the Spina 
Bifida Committee (Transactions of the Clinical Society, vol. xviii.); 
but in this instance there was a second large osseo-cartilaginous 
erowth projecting into the sac, covered by the anterior wall of 
the sac, and free from the posterior wall, and consisting 
apparently of the bodies and processes of two vertebree placed 
behind two of the lumbar vertebre, and connected with them 
and with each other in a manner similar to that which usually 
obtains between the vertebral bodies at the time of birth. 
Whether these two are to be regarded as displaced vertebra, 
vertebre shunted, as it were, behind the others, or as super- 
numerary vertebre, is difficult to decide. Indeed, the morpho- 
logical significance, if they have any, of these pseudo-vertebrae 
and the azygos bar in front of them, I must leave for the 
consideration of those who are better versed than I am in the 
vagaries of the developmental processes. 

Thirdly. The deficiency in development of the lower part of 
the sacrum and of the coccyx, the curves in the spine, and the 
anchylosis of the vertebral arches, are worthy of note. 


588 PROFESSOR HUMPHRY. 


2. Professor Stewart, at the College of Surgeons in London, 
has been so good as further to dissect for me and make drawings 
(Plate XVIII.) of the interesting specimen (No. 276c) in that 
museum presented by Mr Jackson of Wolverhampton, and 
represented and partially described in the Report on Spina 
Bifida given in the Zransactions of the Clinical Society, vol. xviii. 
This further dissection shows that the specimen really consists 
of 41 vertebree, viz., the 11th and 12th dorsal and the 1st, 2nd, 
and half of the 3rd lumbar, and that the bodies of the 12th 
dorsal and 1st lumbar are fused together, forming what appears 
as one body. The evidence of the individuality of these two 
is furnished by their arches and processes being separate, with a 
separate intervertebral foramen on each side, which was of quite its 
natural size, and had a nerve passing throughit. There is alsoa 
band of intervertebral substance surrounding the apparently 
single body, and dividing it into an upper and a lower portion; 
and a slight extension of this substance upwards and downwards 
indicates the line of division between the bodies and the pedicles 
of thearches. The horizontal bony process traversing the verte- 
bral canal passes backwards from between the bodies of the Ist 
and 2nd lumbar vertebre. It is broad at its base, where it 
appears to have encroached upon the bodies of the contiguous 
vertebree, especially of that above it; and its irregularly quadri- 
lateral extremity, which has a deep pit in its middle, occupies the 
ends between the lamin of these vertebre (the Ist and 2nd 
lumbar), being connected to them by fibrous tissue, except on 
the left side, where there is bony union between it and the 
lamina of the 2nd lumbar vertebra. 

The spinal cord passes in two divisions on the sides of this 
process unsymmetrically, as described in the report. The central 
canal, which is dilated in the upper part of the specimen, is con- 
tinued into the right or larger division, and a blind tubular 
pouch projects from it through an interval between the arches 
of the 12th dorsal and 1st lumbar vertebree—the arches, that is, 
of the vertebra, the bodies of which are united. This pouch is 
accompanied and surrounded by a process of dura mater, which 
forms a covering over it. The central canal is not dilated below 
the bony process. The mass of the swelling into which the 
pouch projects appears to be formed of soft connective or mucous 


an ea: 


ty A Rl TT 


SIX SPECIMENS OF SPINA BIFIDA. 589 


tissue, covered by skin or structure continuous with the skin 
and subcutaneous tissues. That, however, it is not easy to decide 
in the present state of the specimen. 

3. In the specimen in St Thomas’ Hospital (No. LL, 123), de- 
scribed and represented in the same report (the skeleton of a 
fcetus with spina bifida affecting the last dorsal and the lumbar and 
sacral vertebre), a cylindrical process bisects the vertebral canal 
just above the spina bifida, and closely resembles the upper, or 
azygos, process in my case. It passes backwards (from between 
the bodies of the 11th and 12th dorsal vertebrae apparently) to 
the arches of the 10th and 11th, which are pressed together in 
consequence of a slight curve forwards at the junction of the 
dorsal and lumbar parts of the column. There is a median 
depression, giving the appearance of two ossific centres in the 
fore part of the body of the 11th dorsal vertebra in front, and in 
that of the 12th behind. 

4, In the specimen (No. 3485) in St Bartholomew’s Hospital, 
mentioned in the same report, of lumbo-sacral spina bifida, 
where a median process extends from before backwards across 
the vertebra canal, perforating the spinal cord immediately 
above the spina bifida, there is evidently great irregularity in 
the bodies of the vertebre and in the formation of the lower 
part of the spine. 

5. In the specimen in University College (No. 5195), also 
mentioned and represented in the report as an instance of parti- 
tioned spina bifida and bifid spinal cord, with a small process . 
of bone in an antero-posterior direction, crossing the vertebral 
canal a little above the sac, and lying between the halves of the 
cord, Mr Topham has been good enough to bisect the spinal 
column. This shows that the spina bifida commences at the 
11th dorsal vertebra, that the bodies of the vertebre along the 
whole column are serially and numerically correct, but that the 
bodies, from 5th to 9th inclusive, of the dorsal vertebre are 
small—some smaller than others—and that placed behind these, 
in close contact with them and dwarfing them, is a second, 
or supplementary set of bodies or apparent bodies. These are 
separated from one another and from the bodies in front of 
them by lines (apparently) of intervertebral substance, as in my 
own case. Furthermore, at the back of these are three smaller 


590 PROFESSOR HUMPHRY. 


osseous pieces, also resembling vertebral bodies, which are pro- 
duced into processes projecting into the vertebral canal, and the’ 
halves of the bisected spinal cord pass on the sides of these 
processes. 

The further examination of these four specimens already de- 
scribed in the report above mentioned has disclosed some addi- 
tional facts; but I do not know that it has thrown any light 
upon the real nature and morphological relations of the pro- 
cesses Which are thus found to project into the vertebral canal 
in some cases of spina bifida. In all, except that in the College 
of Surgeons, and the lower one in the first case which I have 
described, the processes are above the spina bifida. In two, as 
in my specimen, they are immediately above it; in that in 
University College (No. 5) they are higher up. There may or 
may not be some irregularity in the bodies of the vertebre ; and 
when there is an irregularity it is difficult to see any connection 
between it and the abnormal processes. The supplementary 
bodies, or apparent bodies, behind the normal ones, and project- 
ing into the vertebral canal, found in the University College 
specimen, resemble those in mine; but they are situated above 
the spina bifida, where the vertebral bodies and arches were 
in other respects normal; whereas in my specimen they pro- 
ject into the spina bifida. 

6. Mr Batteham, house surgeon to the Wolverhampton In- 
firmary, has been good enough to send me a specimen of sacral 
. spina bifida, taken from a male child, et. 16 days. It was 
removed three days after death, when the soft parts were not in 
a fit condition for minute examination. When I received the 
specimen it was dry, and the soft structures had been in part 
removed. After it had been soaked in water I find that the 
arches of the lower dorsal vertebre are complete, that all 
those of the lumbar vertebre are incomplete, but, as in my case, 
have their proper direction, and the cord passes normally beneath 
them. The sacral arches are laterally deflected, giving great 
width to the vertebral canal at that part. The sacrum. appears 
to terminate abruptly at the second vertebra; but a plate of 
bone, which is probably a continuation of it, runs horizontally 
forwards from it between the two iliac bones. There is no trace 
of coceyx. The lowest part of the vertebral column is there- 


SIX SPECIMENS OF SPINA BIFIDA. 591 | 


fore very imperfect. A median hard bony process arises from 
the back of the space between the 3rd and 4th lumbar vertebre, 
aud expands like a T into a transversely flattened plate, which 
meets, on the two sides, the laminz of the 3rd lumbar vertebra 
and completes the arch. A similar process, consisting of carti- 
lage and bone, projects from the back of the bodies of the Ist 
and 2nd sacral vertebra. It expands, and, inclining to the right, 
is connected with the right side of the 5th lumbar vertebra, 
thus bridging over the right side of the vertebral canal. The 
spinal cord passes in two divisions, one on either side of these 
median processes. The right division, passing under the arch 
formed by the lower median process, is connected with the back 
of the sac of the spina bifida. That on the left side has been in 
great part cut away; and all traces of the sac on this side have 
been removed. It may be inferred that the left division of the 
cord passed beneath the arch formed by the upper process, and, 
like that on the right side, reached the sae of the spina bifida. 


EXPLANATION OF PLATES XVII., XVIII. 


Fig. 1. View of interior of sac of spina bifida from which the right 
side has been removed. a, upper cut edge of sac; a’, lower cut edge 
of sac; 0, spinal cord (right division of it) passing under the carti- 
laginous arch of the lst lumbar vertebra, and expanding upon the 
under surface of the wall of the sac, with the nerves passing from it, 
through the sac, to the intervertebral foramina. Someof the nerves pene- 
trate the wall of the sac where it covers the bony projection into the 
sac. The under wall of the sac is clear from this bony projection. d, 
the lateralised arches of the lumbar and sacral vertebre; e, carti- 
laginous lump representing the coccyx. : 

Fig. 2. View taken from the back and right side of the vertebrie 
at and adjacent to the spina bifida after the soft parts had been cleared 
away. a, the complete cartilaginous arch of the 1st lumbar vertebra. 
A needle has been passed beneath the curved azygos bar, which passes 
from the middle of this arch to the middle of the intervertebral sub- 
stance between the Ist and 2nd lumbar bodies. 6, the incomplete 
arches of the 11th and 12th dorsal vertebree ; c, the anchylosed arches 
of 10th and 11th dorsal vertebre ; d, the osseo-cartilaginous projection 
into the spina bifida ; ¢, lateralised arch of 1st sacral vertebra. 

Fig. 3. View of section of column. a, cartilaginous arch of 1st 
lumbar vertebra ; a bristle is placed beneath the azygos bar. 1, 2, 3, 


, 592 SIX SPECIMENS OF SPINA BIFIDA. 


4, 5, bodies of the five lumbar vertebre, with the supplementary bones’ 
(vertebral bodies and arches?) situated above (behind) the bodies of 
the 3rd and 4th lumbar vertebre, and forming the osseo-cartilaginous 
projection into the spina bifida ; 0, cartilaginous knob representing the 
coccyx. 

We. 4, External view of specimen in College of Surgeons. : 

Fig. 5. Sectional view. 11, 12, 1, 2, 3, the bodies of the lowest two _ 
dorsal and the upper three lumbar vertebre ; a, the band of cartila- 
ginous intervertebral substance encircling the fused bodies of the 
12th dorsal and 1st lumbar vertebrz. The bony process is seen pro- 
jecting backwards from between the lst and 2nd lumbar bodies, its — 
hollowed posterior part presenting in section a bifid appearance ; b, — 
central canal of spinal cord ; c, tubular prolongation of it between the 
arches of the 12th dorsal and 1st lumbar vertebrae, covered by an ex- — 
tension of the dura mater; d, upper and lower cut ends of the smaller — 
division of the cord, which passed on the left side of the osseous 


process. 


ON THE PENTADACTYLOUS PES IN THE DORKING 
FOWL, A VARIETY OF THE GALLUS DOMESTICUS, 
WITH ESPECIAL REFERENCE TO THE HALLUX. 
By JouHNn Cowper, Student of Anatomy, University of 
Edinburgh. 


In the common fowl, as is well known, the foot possesses only 
four toes, but in the breed known as the Dorking, there are 
normally present five digits on each foot. In Tegetmeir’s 
standard work on poultry (p. 92) the following statement 
relating to this bird is found :—“The feet must be five-toed, the 
extra toe well developed and distinctly separated from the 
others.” 

I have dissected the feet of four specimens of this fowl, with 
the view of determining the arrangement of the extra toe. The 
tarso-metatarsus is divided inferiorly into three articular 
surfaces for the three outer digits, and, as in other birds pos- 
sessing more than three digits, there is also present a rudimentary 
lst metatarsal bone, articulating with or attached to the lower 
end of the inner aspect of the tarso-metatarsus. It is with 
the distal part of this bone that the toe which it is customary to 
call the hallux articulates. In the case of the Dorking the 
rudimentary 1st metatarsal bone presents, however, two distinct 
articular surfaces on its distal aspect, the inferior and smaller of 
the two being for articulation with the so-called hallux, while 
the superior and larger is for the proximal phalanx of the extra 
digit normally present in the Dorking fowl. This digit is 
almost exactly on the same antero-posterior plane as the so- 
called hallux, but is placed superior to it. The so-called hallux 
possesses two phalanges, while the extra toe possesses three, 
which latter are usually larger than those of the so-called hallux, 
and the distal phalanx of the extra toe carries a claw, just as 
does the corresponding phalanges of the other toes. 

On the dorsal aspect of this extra toe a tendon is situated, 
which is inserted into the proximal end of the dorsal aspect of 
the middle phalanx, and it may be also into the terminal 


594 MR JOHN COWPER. 


phalanx, by means of a prolongation ; the tendon, when followed 


up, is found to take origin opposite the upper end of the tarso- 


metatarsus, from the common extensor tendon of the three 
outer toes—this latter tendon coming from a muscle situated on 
the anterior aspect of the crus. The so-called hallux possesses 
an extensor tendon, which is seen to be derived from a muscle 
arising from the anterior aspect of the upper end of the tarso- 
metatarsus. It has also a flexor tendon inserted into the 
proximal end of the inferior aspect of the proximal phalanx, 
which is derived from a muscle arising from the inner aspect of 
the upper end of the tarso-metatarsus. There is also a second 
tendon inserted into the terminal phalanx on its inferior aspect, 
and this tendon is common to the so-called hallux and the extra 
digit; it is derived from a muscle of the crus, and bifurcates, 
one half going to be inserted into the inferior aspect of the 
terminal phalanx of each of these toes; just before bifurcating 
it sends tendinous slips to the flexor tendons of the three outer 
digits. This extra digit is truly transmitted, so much so that it 
constitutes one of the essential characteristics of the Dorking 
fowl, and it is stated in Mr Darwin’s Animals and Plants under 
Domestication to be generally transmitted to the offspring, even 
when crossed with other breeds which only possess four toes. 
What is the morphological value of this extra toe ? 
Professor Huxley states, in his Anatomy cf Vertebrates :— 


In all birds, even in Archzeopteryx, the 5th digit of the pes remains 
undeveloped, and the 2nd, 3rd, and 4th metatarsals are anchylosed 
together. 


Professor Owen states, in his Comparative Anatomy and 
Physiology of Vertebrates, vol. ii. p. 83 :-- 


The toes of birds never exceed four in number, and of these three 
are usually elongated and directed forward, diverging, while one is 
short and directed backward. The hind toe articulates on the same 
plane as the others in grasping and perching birds, but on a higher 
level in terrestrial and aquatic kinds. By the analogy of the number 
of the phalanges of these toes with those in lizards, the back toe is the 
innermost, answering to “hallux,” the inner of the front toes is the 
2nd, the middle one is the 3rd, the outer one is the 4th; it will 
be seen that the number of phalanges progressively increases from two 
to five. The 5th toe of the four phalanges in the Monitor is not 
developed in any bird. 


o 


THE PENTADACTYLOUS PES IN THE DORKING FOWL. 595 


In the foot of the Dorking fowl I believe that the toe second 
in position and possessing two phalanges represents the so- 
called hallux of other four-toed birds, because it occupies the same 
morphological position, has the same number of phalanges, and 
articulates with the rudimentary metatarsal placed on the inner 
aspect of the tarso-metatarsus, If then the extra toe, possess- 
ing three phalanges, and placed internal te the so-called hallux, 
is to be regarded as a constituent part of the formation of the 
foot, and not as a monstrosity—aud its constant presence in the 
Dorking’s foot is, I think, sufficient evidence that it is not a 
monstrous formation—then it, and not the digit usually called 
“hallux,” occupies the position of the hallux, and must be regarded 
assuch. If this beso, it follows that the so-called hallux, possess- 
ing two phalanges, is really the 2nd toe, and that the toes usually 
called 2nd, 3rd, and 4th, are the 3rd, 4th, and 5th; the missing 
toe in a four-toed bird being not the 5th but the hallux, and the 
so-called hallux being the 2nd tve. In sucha case, the rudi- 
mentary metatarsal, with which the 1st and 2nd toes of the 
Dorking fowl articulate, probably represents the elements of the 
Ist and 2nd metatarsals, and the tarso-metatarsus represents 
the 3rd, 4th, and 5th metatarsal bones. 

Taking the Reptilia as a class, they possess five digits in the 
pes; in the Mammalia, however, the digits of the pes are very 
commonly reduced in number, and Professor Flower states—“ if 
one toe only is absent it is the first or hallux,” so that, if the 
above view of the morphology of the foot be correct, the bird 
follows the mammalian rule in this respect. 


THE VITALITY OF WILD ANIMALS UNDER FIRE. 
By Brigade-Surgeon WiLtiAM Curran, A.M.D. (PLATE 
XIX.) 


THE comparative immunity with which soldiers and others 
engaged in warfare receive wounds in vital parts is well known. 
Many instances of this immunity are on record, and specimens 
in point may be seen any day in our leading anatomical 
museums. They are also described in great detail by surgical 
writers, but the particulars of these would be unsuited to these 
pages, and the books containing them are open to all. It is not 
so well known that our fellow mortals, “the beasts that perish,” 
are equally favoured in this respect, and one would look in vain, 
even through the writings of professional veterinarians,! for any 
exhaustive account of this immunity. We therefore think 
that the matter stands in need of elucidation, if only from an 
artistic point of view, and what we have to say on the point is 
fortunately based on our own personal experience, or on the 
statements of friends whose competence and veracity are beyond 
the reach of doubt. 

It would not be easy to say what species of animal exhibits 
the largest amount of this kind of tolerance, nor has the question 
been, so far as we know, ever raised. Anyhow, it has not been 
determined, and, even admitting that a comparison is possible, it 
is not easy to see how one could be instituted, with any approach 
to accuracy, between the heart of a whale, which fills a large 
tub, and that of a mouse ora weasel that can be taken up on the 
end of a pin or swallowed ata bite by a small puppy. All we 
need say is that as “the structure of the lungs, the mechanism 
of respiration, the arrangement of the pulmonary vessels, the 
cavities of the heart, and the general disposition of the arteries 
and veins of the systemic circulation, differ in no material 
circumstance (in the Mammalia generally) from what is met 
with in our own persons,” so also there would be no more 


1 Williams’s Veterinary Medicine (p. 519) is the only work that contains, 
so far as we know, any information of this kind. 


VITALITY OF WILD ANIMALS UNDER FIRE. 597 


difference in this respect between man and beast than their 
respective sizes, endurance, or muscular development would 
naturally enforce or imply. 

And this, too, is found in practice to be really the case. For 
though the tiger, the bison, the deer, and other creatures of that 
kind may exhibit greater momentary activity after wounds in 
their hearts than man usually does, yet is this their activity not 
sustained. Their ungovernable rage, fear of man, or efforts at 
escape only tend by stimulating the circulation, or preventing 
the closure of their wounds to aggravate the iatter, and we find, 
accordingly, that whilst some wounded beasts are often able to 
execute fierce charges after they are fatally stricken, or that they 
can run considerable distances without betraying any sign of 
vital failure, yet do they as assuredly succumb in the end, and 
that, too, in a shorter time than their wiser or less impulsive 
fellow-sufferer—man. But then the latter is not obliged to fly for 
his life as the beasts are, and that, we fancy, makes all the differ- 
ence in the case. 

Owing to the bungling of a certain executioner in this 
country, or to the facial contortions of persons beheaded in 
France, we have lately had discussions in the Press as to the 
best or least painful method of taking away the lives of our 
criminals, Some writers advocate the use of prussic acid or other 
instantaneously fatal poison, others prefer electricity, and a few 
have suggested chloroform or some similar narcotic. In other 
words, the advocates of these measures imply that the best means 
of effecting their purposes are such as operate through the brain 
and the nervous system, or through the heart—the central organ 
of the circulation. They want to destroy the sensibility of the 
one, or paralyse the action of the other, but life is not thereby 
necessarily extinguished in either case, and it is now generally 
admitted that consciousness survives decapitation by the 
guillotine. 

On the other hand, very awkward accidents have followed the 
use of the rope through inexperience, timidity, or intemperance 
on the part of the executioner, so that measures directed to the 
stoppage of the heart would seem to be those that make the 
strongest appeal to our sympathies. But are there any such 


measures, or is it quite so certain that the cessation of the heart’s 
VOL. XX. 2R 


598 MR WILLIAM CURRAN. 


action necessarily entails death? We know asa matter of fact 
that such is not the case, and that many persons whose hearts 
had ceased to beat to all outward appearance during periods cf 
so-called suspended animation—catalepsy, &c.—have subse- 
quently recovered, and lived long afterwards. Per contra, life 
would seem to survive, if we are to believe hagiologists and 
others, the actual removal of this organ from its owner. Thus 
Burnet,! describing the execution of one Hackston a covenanter, 
says that “his heart when cut out continued to palpitate some 
time after it was in the hangman’s knife,” and Bishop Challoner? 
adds that Edward Jennings “uttered the words Sancte Gregori 
ora pro me, after his bowels had been thrown into the fire, and 
his heart had been torn out by the executioner.” Facts of like 
import are mentioned by toxicologists,’ and the very structure of 
this organ, consisting as it does of involuntary muscular fibre, 
would seem to imply that life may coexist with, nay, may even 
be restored, after it had apparently ceased to act. 

Let us now take the heart of a sheep—a very accessible 
object by the way—and compare with it, as with a standard, the 
lesions or injuries which the hearts of some other animals are able 
to bear without immediately failing. It would be readily seen 
that this is a perfectly healthy organ, very like that of man, 
though somewhat smaller, and that it possesses substantially the 
same structure and elasticity as his. One would suppose that 
any leak, however small, in this central fountain would prove 
necessarily, if not instantaneously, fatal. But such is not the 


1 History of his Own Times, p. 338. He notes, as ‘fa matter of no small 
astonishment,” in his History of the Reformation, vol. i. p. 533, that Cranmer’s 
heart remained ‘‘ entire and not consumed among the ashes” that were produced 
by the burning of his body. This organ often survives, we believe, the decomposi- 
tion of the other fleshly parts of its receptacle. 

* Martyrs to the Catholic Faith, p. 182. Bacon mentions, in his History of Life 
and Death, that ‘he saw the heart of one that was bowelled for high treason leap, 
on being cast into the fire, a foot and a half high, and after, by degrees, lower and 
lower, for the space, as he remembered, of seven or eight minutes.” We may put 
any construction we like on such narratives, and it was stated in a pamphlet of 
the day that when the body of Bellingham (who was hanged for the murder of Mr 
Percival in 1812) was dissected, ‘‘it was noticed that his heart continued its 
functions for four hours after he was laid open.” 

3 See, for a very interesting case in point, Dr Chevers’ Manual of Medical 
Jurisprudence for India, pp. 645-7 ; and Taylor adds that ‘‘ Duval saw the heart 
of a criminal, a quarter of an hour after decapitation, beating with great 
distinctness. ” 


— 


A 8 ee Ry Sh eee ieesinihcaitieagiahin os 


VITALITY OF WILD ANIMALS UNDER FIRE. 599 


case, and considerable portions of its body may be taken away 
without apparently diminishing the activity of its owner, or 
instantaneously extinguishing life. We will now endeavour to 
establish the truth of this statement by facts drawn from our 
own or other’s experience. 

One day, while shooting with a friend near Lucknow, we came 
across a small herd of the Black Buck or Antelope of tie plains. 
One of these, a female, fell to the rifle of our companion, and as 
this succumbed to a wound of the head, we subjoin a sketch of 
its heart, which, it will be at once noticed, was quite intact. The 
black spot or star on it is intended to show the place in which 
one of its fellows was struck, about the same time, by ourselves.’ 

This latter ran, after receiving our fire, for fully 40 yards or 
so, as if there was nothing the matter with it, and then rolled 
over, “all of a heap,” and fell dead. Till this took place we were 
under the impression that we had missed it. On examining its 
heart afterwards we easily put our forefinger quite through its 
centre (fig. 1). The ball which inflicted the wound was a sharp- 
pointed conical one,and was fired out of one of Sharp’s patent rifles. 

The animal (Felis tigris) whose heart is here reproduced (fig. 2), 
was shot by us at night, while watching from a muchan (a stage in 
a tree), a buffalo that was tied, as a bait, beneath us, and which 
it had sprung upon and killed. We fired from a distance of 
forty paces, but the tigress did not acknowledge our shot, as is 
usual in such cases when they are struck, but gave one bound 
and disappeared. We naturally thought that we had missed her, 
and knowing that our shot and its smell, &c., would effectually 
scare away any other beast that might be prowling about in the 
neighbourhood, we retired to our tent for the night. On return- 
ing to the spot whence we had fired on the previous night, early 
on the following morning, we found her lying quite dead, about 
120 yards from the same. 


1 This tolerance is not peculiar to the Antelope of India. Mr Myers found a 
very similar amount of vitality in one of its fellows in Africa. Describing his 
adventures in a portion of that Continent, he says (Life among the Hamran 
Arabs, p. 215) that ‘‘an Antelope, in whose side such a hole had been made that 
the lung protruded, not only survived for some time, but actually ran away”; and 
he adds further on, that ‘‘a Gazelle managed to gallop off more than one hundred 
yards after most of its interior had been knocked out by a low shot with an ex- 
panding bullet.”” It has not been considered necessary to include this sketch 
in our plate. 


600 MR WILLIAM CURRAN. 


She had, meanwhile, bitten her thighs through in two or three 
places, but this is not a very uncommon occurrence, and we have 
even heard of an instance in which a wounded tigress worried 
her own young ere she died. The fatal wound was just behind 
the shoulder, and,after skinning her we opened, as was our wont, 
her chest. The heart was completely shattered, even more so 
than in the drawing. The ball we used was a spherical gauge, 
and was fired from a smooth-bore barrel which contained 5 
drams of powder. 

The heart here figured is that of a panther (Felis leopardus), 
and its owner was killed in much the same way as the tigress 
just referred to (fig. 5). Our bait in this instance, however, was an 
old sheep, and not a buffalo, and we fired from a distance of only 
5 or 6 feet. The brute bountled away with a growl (a good 
sign), and we found it dead the next morning about 40 yards 
from where we stood. On examination the heart was found to 
be shattered. Our projectile on this occasion was a copper 
bottle No. 12 bore shell, filled with chlorate of potash and 
sulphuret of antimony, and it was propelled by 4 drams of 
powder. 

So far, omitting minor episodes, for our own personal experi- 
ence on the point; let us now turn to the experience of com- 
panions or others whose feats we either witnessed ourselves or 
had them rehearsed to us, viva voce, by their authors. A field 
officer of our acquaintance (a noted Shikari), answering a recent 
interrogatory of ours, says :—“ I once fired into a full-grown tigress 
that had been badly wounded in the belly the previous day, as 
she was going full split in pursuit of prey. My rifle was a 
16 bore, and this contained a conical (Jacob’s) shell of about an 
ounce and three quarters weight. It did not explode, which was 
due, I think, to the fact that I fired broadside on, as she was 
about 1 foot from my muzzle. I fired from her left side, and 
(being 6 feet high) directed my aim heartwise, and the ball 
tore away a good sized piece (of flesh) from the apex of her 
heart from above downwards.” 

She then passed on as if untouched, chased for 15 yards or 
so a gun-carrier,and pulled him down. She continued to worry 
him alive while I was reloading from the muzzle one barrel, and 
then sank back quietly all at once with the man in her mouth. 


oe eo ay)) ate ee ee 


VITALITY OF WILD ANIMALS UNDER FIRE. 601 


He further describes the charge of a full-grown tiger, whose 
heart had been “smashed to smithereens” by a shell explosion 
within his chest, and concludes by saying that “I have seen 
several black-buck, that were shot through the heart itself, go 
full tilt as if untouched, and then roll over dead.” 

And apropos of the leading passion strong in death, as well in 
beast as in man, I may be here permitted to interpolate a fact in 
this connection that was related tome several years ago by a bye- 
stander. It runs to the effect, that an officer who subsequently 
achieved a high position in the service was carried bodily from 
the houdah of an elephant, at a spring, by a tiger through whose 
heart a bullet was subsequently ascertained to have passed. 
The officer in question was deposited on the ground badly 
bruised and scratched, and was only released from a worse fate 
by the death of his assailant on his body.! 

And though these statements are sufficient of themselves to 
establish the truth of our thesis, yet have we considered it desir- 
able to add, as far as in us lay, to their cogency. We have 
applied with this view to two medical friends, both well-known 
shots, to favour us with their experience on this point, and they 
have answered us in confirmatory terms. One of them, an 
officer of long standing in the Indian Army, assures us that he 
was once charged by a panther, “the whole of whose thoracic 
viscera—including, of course, the heart—had been torn to 
pieces,” while on another occasion a nylghaie (a species of cow), 
whose right auricle and root of aorta were wounded by a No. 12 
spherical bullet passing through them, went away “ for more 
than 100 yards as if there was nothing the matter with him.” 

The other, an old friend and confrére, writes to us as follows 
on the same subject :—“I can give you two instances (of surviv- 
ance after gun-shot injuries of the heart, &c.), one shot through 
the heart with a large shell, and the other shot through the aorta, 


1 We may quote, in corroboration of this fact, the observation of Major 
Shakespear (Wild Sports of India) to the effect that ‘‘ those who have not actually 
seen it will scarcely credit that a tiger will often go in his charge several yards, 
with all the power and capability to strike down everyone in his path, after the 
bullet has gone through his heart or crashed through his brain.” Describing his 
adventures in South America, Mr Masters says (At Home with the Patagonians, 
p. 53), apropos of the Puma, that ‘‘to kill this animal with a gun is rather a 
difficult matter, as, unless the ball enters his skull, or strikes near the region of 
the heart, he has as many lives as a cat.” 


602 MR WILLIAM CURRAN. 


close to the heart, by a bullet, which clean divided this vessel in 
its entirety. The first was a tigress, which stood about 20 
yards from me; the shell entered just behind the right shoulder, 
passed through both ventricles of the heart, and emerged behind 


the other shoulder. The beast ..... ran a little over 100 
yards and then dropped dead.” 
“« B—— shot a tiger with one bullet through part of the 


shoulder and brisket, and with the other through the right 
shoulder and aorta as it ran away. This brute gave no sign 
whatever of being hit, and my friend thought he had missed him, 
but it was not so; on following up the track we found him dead 
about 40 or 50 yards ahead of us. I made a post-mortem ex- 
amination in both these, and can vouch for the truth of what I 
have written,” and it is painful to add that these two officers 
are now dead. One of them succumbed to injuries that were 
inflicted by a tiger in the Nepal Terai, the other died of a fever 
that was contracted, in the pursuit of game, in the same locality, 
and both were men of much promise in their respective spheres. 

We might easily adduce similar records in respect of the 
Himalayan bear, the bison of the American Prairies,? and the 
ovis ammon, oorial or wild sheep of Thibet, and the Sevaliks, did 
space so allow. But it does not. We have, we fear, already tres- 
passed too far on the indulgence of our readers, and moreover 
we have had no experience of our own in connection with these 
animals. We prefer, therefore, leaving our case to rest on its 
own merits, such as they are as above, rather than encumber it 
with details for which we cannot ourselves personally vouch. 
Books of sport and hunting adventure are common enough, and 
such of our readers as desire further information on these points 
may be referred to the writings of Dunlop, Kinloch, Markham, 
and others, who have specially dealt with this phase of the 
question. 


1 Sir Edward Melville wonders (Zhe Last of the Arctic Voyages, vol. i. p. 64) 
that a bear he met in Melville Bay showed fight ‘‘even after a ball had passed 
through his brain.” 

2 See Ruxton’s Adventures in Mexico and the Rocky Mountains, p. 268, and 
Captain Trench Townshend’s J'en Thousand Miles of Travel, &c., &c. The writer 
has been recently informed, by an officer who served at the Cape, that ‘‘a bustard 
once flew (in his presence) several hundred yards after he had received in his 
chest the contents (buckshot) of a heavily loaded gun, which caused the viscera 
of his thorax to protrude, 


TOR oy inteticeaah tation dictate * 


WET eenp Se a, ee 


* 


VITALITY OF WILD ANIMALS UNDER FIRE. 603 


EXPLANATION OF PLATE XIX. 


Fig. 1. Heart of the black buck of the plains of India, showing 
wound in the right ventricle. 

Fig. 2. Heart of a tiger, in which the greater part of the ventricles 
was shot away. 

Fig. 3. Heart of a panther, the ventricles of which were destroyed 
by a 12 bore shell. 


THE ALIMENTARY CANAL AND PANCREAS OF 
ACIPENSER, AMIA, AND LEPIDOSTEUS. By 
A. B. Macattum, B.A., Fellow of University College, 
Toronto, Canada. (PLATE XX.) 


THE investigation, some results of which appear in the present 
work, was begun two years ago, with the object of treating fully 
of the digestive organs of fishes in general. Since I have now 
had for some time but few opportunities for continuing the 
research further, and do not expect to be more fortunate in this 
respect for a year or so to come, I have been led to abandon 
that object wholly. The results of my studies, however, so far 
as they relate to Ganoid fishes, seem to me to have some value, 
and I have therefore decided to publish them. 7 

The description here given, so far as it pertains to Lepidosteus, 4 a 
is based on forms of 7-10 em. in length, which I was forced to 
accept for the purpose, since the adult forms are to be obtained 
from remote places only, and always in a poor state of preserva- 
tion for histological examination. As a full description of the 
histology of the intestinal tract in young Lepidostei would not 
correspond in every respect to the conditions obtaining in the 
adult forms, I have endeavoured to omit references to peculi- 
arities of structure such as may be affected by age. 

The terms fore-gut, mid-gut, and hind-gut are here employed 
for the purpose of description, as they answer to the conditions 
of the alimentary canal in fishes better than any others that can 
be suggested. They have already, to a certain extent, been 
sanctioned by usage, since they are employed in the English 
edition of Gegenbaur’s Elements of Comparative Anatomy. 


ns 


Ira, ForE-GUrt. 

Esophagus.—In Aecipenser, that part of the alimentary canal 
lying behind the branchial chamber, and terminating about 3 em. 
behind the membrane separating the pericardial and peritoneal 
cavities, is usually considered to be the cesophagus. It is com- 
paratively short, and is at once recognisable by the large number 
of papillee arranged in longitudinal series on its inner surface. 


THE ALIMENTARY CANAL AND PANCREAS. 605 


In its wall are found the two layers of striated muscular fibre, 
commonly present in the cesophagus of fishes. 

There are, however, elements in the structure of this part of 
the alimentary canal which throw some doubt on its being 
homologous with the cesophagus of other vertebrates. It is 
lined with an epithelium formed of many layers of fusiform, 
polyhedral, or flattened cells, which do not differ to any appreci- 
able extent from those of the membrane lining the posterior 
part of the pharyngeal chamber. This kind of epithelium is 
rare in the cesophagus of fishes! There are also to be found in 
this part taste-buds, occurring usually on the summits of the 
papille already referred to. I have never seen any reference to 
the oceurrence of such structures in the cesophagus of any verte- 
brate,2 but they are to be met with commonly in pharynx, 
especially in fishes. That part of the fore-gut in Acipenser 
immediately following the so-called cesophagus is devoid of 
striated muscle, and therefore might be regarded as part of the 
stomach. But this is not a sure method of distinguishing the 
two divisions of the fore-gut, for if the anterior part possesses 
elands which secrete a digesting principle, the absence of volun- 
tary muscular fibre is a necessary condition, in order to allow 
the secretion to perform its function. The anterior part of the 
fore-eut in Acipenser does possess glands of this description, 
hence the absence of striated muscular fibre from its wall. It 
will be seen farther on, that the division of the fore-gut into 
cesophagus and stomach is not very marked in Amia and Lepi- 
dosteus, the passage from one to the other part being at best but 
a gradual one. That part following the papillated section of the 
canal in Acipenser must be regarded, therefore, as the true 
cesophagus, and from its structure, as described below, it may be 
correctly said to terminate a short distance behind the opening 
of the duct of the air-bladder, near the commencement of the 
castric loop. 

The papillated portion is then but the posterior extension of 
the pharynx. Its inner surface is greyish-white, and the few 

1 According to Edinger (Arch. fiir Mikr. Anat., Bd. xiii. p. 651) a several- 
layered scale-like epithelium is present in the cesophagus of Polypterus and of 
some Selachians. 


2 In Amphioxus I have, however, determined the presence of taste-buds as far 
posteriorly in the alimentary canal as the opening of the ‘‘ hepatic”’ cecum, 


606 MR A. B. MACALLUM. 


folds that are present run longitudinally between the series of 
papille. It terminates about 3 cm. behind the pericardial 
chamber. The epithelium lining it is constituted of from five to 
ten layers of cells, which are usually fusiform, often polyhedral 
or squamous, the superficial layer being, however, formed of 
cylinder and goblet cells almost invariably. F. E. Schulze! 
found the cylinder cells provided with cilia. I am unable to 
confirm this, although I have studied the epithelium in fresh 
and in hardened condition. The theca of each goblet cell is of 
an elongated oval shape and is filled with mucus, containing a 
large number of feebly refracting granules. Many of the 
cylinder cells also exhibit a mucigenous transformation of their 
contents. The basal processes of both forms of cells are exceed- 
ingly fine and delicate. The replacement of the cast-off goblet 
cells, into which all the cylinder cells are metamorphosed, is 
auecomplished by those of the layer next below them. 

The taste-buds are not different from those found in the mouth, 
except perhaps in their slightly more elongated form (fig. 2). 
Several may be found on the summit of a papilla. 

The true cesophagus possesses to the naked eye all the charac- 
teristics of the stomach. The colour of its lining membrane is, 
in the fresh condition, chocolate-red, and its folds pass uninter- ; 

| 
; 


ruptedly, without increase or decrease in size into, those of the 
stomach. The calibre of both parts of the fore-gut is the same. 
The only distinctions to be found between them lie in the histo- 
logical structure of the mucosa. The point of transition between 
the cesophagus and stomach, as already stated, is near the gastric 
loop, up to which point the epithelium of the former retains its 
cilia, while the peptic tubules found in it are somewhat different 
from those of the loop. 

In Amia the cesophagus is short, and terminates immediately 
behind the opening of the duct of the air-bladder. As in 
Acipenser, the inner surfaces of both cesophagus and stomach 
are alike in every respect when viewed with the naked eye. 
The folds in both parts are continuous and arranged longitudin- 
ally, while the dull reddish tint of the mucosa of the stomach is 
possessed to some extent by that of the cesophagus. Where one 
part ends and the other begins can be determined approximately 


1 Arch. fiir Mikr, Anat., Bd, iii. p. 174. 


- 


THE ALIMENTARY CANAL AND PANCREAS. 607 


only, the lining epithelium and the glands of the anterior part 
passing without any abrupt change into those of the posterior. 
Taking the presence of striated muscular fibre as a guide in this 
determination, although a very uncertain one, the two parts may 
be divided as cesophagus and stomach at the point intimated. 

In Lepidosteus the cesophagus commences immediately anterior 
to the opening of the air-duct; where it terminates posteriorly 
it is difficult to say. The distinction between the two parts of 
the fore-gut is not less indefinite, perhaps more so, than in the 
other two genera. For a third of the extent of the straight tube 
forming the fore-gut, the epithelium in character and the glands 
in both character and number differ considerably from the 
corresponding structures in the posterior two-thirds, although, so 
far as the glands are concerned, the transition from one to the 
other part is a gradual, not an abrupt one. The anterior third, 
which may therefore be termed the cesophagus, does not seem to 
have any striated muscular fibre in its outer wall, but the speci- 
mens from which my preparations were taken were not of such 
an age as to allow me to speak with certainty on this point. 

The opening of the air-duct in Amia and Lepidosteus isa 
longitudinal slit on the dorsal side of the cesophagus. This 
opening is provided with two folds, one on the right, the other 
on the left, which can be approximated by the striated muscular 
fibre present in them to shut off, after the fashion of a glottis, 
all connection between the duct and the cavity of the cesophagus. 
The muscular fibre is very abundant in Ama, in which it 
radiates out over the duct and on the surface of the air-bladder. 
In Acipenser, in which the opening of the duct is funnel-shaped, 
there is apparently no mechanism by which the cavity of the 
air-bladder can be completely closed off from that of the ceso- 
phagus. 

In Acipenser, Amia, and Lepidosteus the epithelium of the 
cesophagus is ciliated. The cilia in the first named form a thick 
brush on each cell, and all usually matted together by mucus; 
in the other two genera they are much longer and finer. The 
cells in all are cylindrical, with fine basal processes which pass 
into and are interwoven with the connective tissue of the 
mucosa. In Acipensev,in the outer or peripheral halves of a 
large number of these cells, the contents are swollen like those 


608 MR A. B. MACALLUM. 


of the goblet cells, but the staining with various dyes is uniform 
in the inner and outer cellular portions, the absence of mucigen 
being thereby shown. In Amia many of the cells have lost 
their cilia and are changed into goblet cells; this is more com- 
monly the case in the shallow crypts which are to be found 
abundantly over the mucous surface. All the cells exhibit the 
mucigenous transformation. In Lepidostews the cells are of 
regular form, with their outer portions containing a quantity of 
mucigen. Cilia were observed on all the cells. In this genus, 
as well as in Acipenser, some of the cellular elements must, 
before they break up or are destroyed, open peripherally to dis- 
charge their mucus. 

In the opening of the air-duct in the three genera the cylinder 
cells are short,.and in the middle of the extent of the duct they | 
become cubical. In Lepidosteus the folds at the side of the 
opening are covered with short columnar cells. 

In Acipenser and Lepidosteus the derivatives of the epithelium 
in the esophagus may be referred to three forms :—(1) elongated - 
crypt-like insinkings ; (2) elongated crypts terminated in dilated 
sacs; (3) true gland tubules. In Amia in addition to these | 
three classes of structures there is a fourth, examples of which | 
are to be found in the neighbourhood of the mouth of the air- 

; 


duct, in the latter and even in the air-bladder. 

The elongated crypt-like insinkings of the epithelium are 
apparently most common in Acipenser in which they are 
specially noticeable at the commencement of the cesophagus. 
They are not always straight in their course through the 
mucosa. The ordinary epithelium of the general surface is 
continued into them for half their length, while the lower half of 
the cylinder cells, though still retained, are shortened, much 
so in Amia, and their cilia less prominent. In these the 
nuclei are placed next the membrane of connective tissue 
surrounding the crypt; the contents of the central portions of 
the cells appear to be mucigenous when viewed in sections of 
the hardened tissue, but when observed after isolation in serum 
the mucigen is obscured by the presence of fatty matter in the 
form of minute globules, which are more abundant near the 
lumen. In Amia the corresponding cells are shorter and the 
central half of each is packed full of granules, whether of 


THE ALIMENTARY CANAL AND PANCREAS. 609 


zymogen or not 1 am unable to say. In Lepidosteus these 
insinkings of the epithelium are found only at the posterior end 
of the first third of the fore-gut, and their cells are much short- 
ened, often reaching a cubical form. 

In <Acipenser these crypts may branch more than once, 
although as a rule they do not branch at all. In this genus 
they are not found farther than 2cm. behind the commencement 
of the cesophagus, where they become changed into the structures 
of the third class mentioned above. This change is a gradual 
one, the cells at the bases of the crypts, as the latter are 
followed backward, becoming shorter and more cubical, assuming 
at the same time a more specialized glandular character. In 
Amia also, these insinkings are least numerous where the true 
gland tubules are most so, and passage-forms between the two 
elasses of structures are present. 

Elongated crypts terminating in dilated sacs are not numerous 
in Acipenser, and Lepidosteus and in Ania, in which they appear 
to occur in the largest number, they bear a very small proportion 
to the structures of the first class. As to their character they 
are most highly developed in Acipensev and least so in Lepi- 
dosteus. The tubular part is lined with cylinder cells which 
decrease in length as the sae is approached, while the cilia 
become shorter and tess abundant. There is in Acipenser a 
slight constriction of the tubule where it terminates in the sac; 
at this point the cilia vanish and the cells attain their greatest 
decrease in height. The cells lining the sae and their nuclei 
are flattened, but the form never becomes squamous. What 
little protoplasm there is surrounding the nuclei appears clear 
and transparent. 

Sometimes in Ama one or more gland tubules are found to 
open into one of these sacs. This is the case in young forms 
particularly. I have not, however, observed a like arrangement 
in the two other genera, but whether it occurs in them or not 
these sacs are still remarkable on account of their structure, 
since they have not, as far as I am aware, been observed in any 
other fish. Their fewness of number and their limited distri- 
bution in Acipenser also render them worthy of note. They 
cannot serve in any case for the purpose of absorption, since 
they are placed too far in front of the seat of digestive changes, 


610 MR A. B. MACALLUM. 


and as they are undoubtedly devoid of a glandular function, one 
is compelled to consider them to be transudatory organs. A 
careful study of the histology of the fore-gut in a large number 
of fishes may show the presence of these structures to be more 
ceneral. 

True gland tubules are most abundant in the cesophagus of 
Acipenser and fewest in Lepidosteus ; in the latter they are only 
to be found at the posterior end of the cesophagus. In the first 
named genus they are usually much elongated, the elongation 
increasing as they are followed backward. Large cubical cells 
line them and enclose a lumen of considerable size. The 
contents of the central half of each cell stain with difficulty 
in carmine, while the contrary is the case with the outer half in 
which the nucleus is placed. The edge of the cell touching on 
the lumen is very irregular and ragged in appearance, owing to 
the abundance of granules present. 

The neck of each tubule and the crypt into which it opens 
constitute from one-third to one-half the length of the three 
parts, and in this respect there is a marked contrast to the 
elands of the anterior portion of the stomach in Aczpenser, or 
to those in any part of the cardia in the other genera. In the 
crypt the epithelium is the same as that of the general surface, 
being ciliated and cylindrical. In the neck of the gland the 
cells are transitional in their form, and contents between those 
of the crypt and those of the body of the tubule, being nearly 
cubical in form, and containing a non-granular protoplasm with 
a narrow zone of mucigenous matter in each, which borders on 
the lumen. 

There is in Ama, as already mentioned, a fourth variety of 
tubules occurring at the mouth of the air-duct, and appearing to 
be similar in every respect to that found in the air-duct itself, 
and on the inner surface of the inferior wall of the air-bladder. 
Tubules of this sort are not numerous. They and the erypts 
into which they open are very short. Their cells are narrower 
and shorter than those of the other kinds of tubules described. 
Sometimes the contents of the cells are strongly granular, at 
other times they are constituted of mucus almost wholly. The 
lumen of the tubule is more frequently indistinct. 

I had not, unfortunately, an opportunity of studying these 


THE ALIMENTARY CANAL AND PANCREAS. 611 


tubules in a fresh condition, when a careful examination of 
them might reveal more about their function. They are in all 
probability but degraded forms of the true gland tubules of the 
cesophagus. A wider knowledge of the histology of the air- 
bladder in fishes may show that the occurrence of tubules of 
this description is not confined to Ama, since the air-bladder, 
before its function had become hydrostatic, probably acted as an 
accessory secreting organ provided with the same histological 
structure as the fore-gut from which it originated as an out- 
crowth, and ought, therefore, still to have in many cases traces, 
though rudimentary, of this similarity. 

Stomach.—The stomach in the three genera shows a consider- 
able difference of form. The division of it into cardia and 
pylorus is easily distinguishable, much more so than is the 
division of the fore-gut into cesophagns and stomach. 

In Acipenser the stomach is thrown into a loop which ex- 
tends forward in front of the mouth of the air-duct, where the 
cardiac portion terminates in a retort-like pylorus. The wall 
of the cardia does not differ in thickness from that of the ceso- 
phagus, but in the pylorus the thickness is from one and a half 
centimetres, especially at the upper and under surfaces, at each 
of which the muscle fibre is collected into a thick plate-like 
mass; this in the relaxed condition of the pylorus lies flat on its 
fellow of the opposite side. At the right and left borders of 
these thickened plates the wall of the pylorus is as thin as in 
the cardia. In the latter the colour of the mucous membrane 
in its fresh condition is chocolate-red and the folds are arranged 
longitudinally, while in the pylorus the mucous membrane is 
dull white and its surface smooth. Viewed from without there 
is a contrast in the colour of the two parts of the stomach, that 
of the pylorus being a glistening white. In a sturgeon measur- 
ing 66 cm. the cardia was 16 cm., and the pylorus 3 cm. in 
length. 

In Amia the cardia is dilated and its cavity is continued 
behind into a conical cecum. The pyloric tube originates 
from the lower side of the cardia and passes forward and 
outward, undergoing a constriction where it unites with the 
mid-gut. The wall of the stomach is of equal thickness 
throughout, except for a slight increase of the muscular tissue 


612 MR A. B. MACALLUM. 


near the termination of the pylorus. The mucous membrane is 
folded, the direction of the folds being longitudinal; its 
colour is the same in the first half of the pylorus as it is in the 
cardia and cecum. The surface of the membrane in the 
posterior half of the pylorus is nearly smooth. The colour here 
is a dull white. 

In Lepidosteus the calibre of the stomach increases as it 
passes backward. Its caecum, if it can be said to have any, is 
very short. Its mucous membrane is raised into longitudinal 
folds. On the inferior surface the narrow pyloric tube passes 
forward a short distance, then turns to the right, and terminates 
in the mid-gut immediately in front of the lobulated pyloric 
appendage. 

The epithelium of the stomach in Acipenser is somewhat 
different in character from that in the cesophagus, and it differs 
even in the cardia and pylorus. The cells constituting it in the 
cardiac portion are longer and more slender than those in the 
cesophagus, and smaller than those in the pylorus They are 
unciliated, a peripheral membrane is present in each, and their 
contents do not apparently show any traces of that transforma- 
tion into mucigen usually observable in the gastric epithelium 
of other vertebrates. The staining in my preparations is quite 
uniform throughout each cell. Only when Miiller’s fluid or a 
solution of potassic bichromate had been used as a hardening 
reagent did any of the cells, and then only a few, exhibit a 
peripheral opening. When alcohol alone had been used in 
preparing the epithelium the peripheral wall of each cell appears 
thick, probably owing to a deposit of mucus on the outside. 

In the pylorus of Acipenser the majority of the cells of 
the epithelium are open peripherally; from this, and from 
their being sometimes swollen, they bear a resemblance to 
voblet cells. The contents of the distal half of each cell are 
almost completely mucigenous. A thick dense layer of mucus 
is often found to cover the epithelium, but it does not dip down 
into the crypts, although it accommodates itself to the general 
surface of the mucous membrane. Scattered throughout this 
layer of mucus a large quantity of granules together with nuclei 
of broken-down cells can be observed. 

In Amia the epithelium is ciliated throughout the cardia, 


THE ALIMENTARY CANAL AND PANCREAS, 613 


cecum, and pylorus. The cells are long and cylindrical, slightly 
swollen at their peripheral halves, and they are covered in 
chromic acid preparations by a thin coatof mucus. The contents 
in the peripheral half of each cell do not appear in fresh condition 
to differ from those in the central half, but when hardened and 
stained it is usual to find the outer half of the cell completely 
uncoloured. The cilia are obscured by the coat of mucus, but 
in the deeper crypts when this is absent they can be easily 
observed. 

In Lepidosteus the epithelium of the posterior two-thirds of 
the fore-gut is unciliated, except in the posterior half of the 
pyloric tube. The cylinder cells are always provided with a 
peripheral membrane. They contain mucigen, as they are 
stainable only in their central halves. 

The glands of the cardia in Acipenser are of the type to be 
usually found in fishes (fig. 7). The crypts of the epithelium, 
into which several gland tubules may open, have evidently been 
mistaken by Leydig! for glands, and they form, according to 
him, as such the only kind observable in Acipenser. Edinger, 
commenting on Leydig’s description, remarks that it is doubtful 
if Acipenser has true gastric glands, for the structures figured and 
described by Leydig bear no resemblance at all to the peptic 
glands of other fishes. Wiedersheim,® also misled by Leydig’s 
description, doubts the existence of peptic glands in the Sturgeon. 
I am unable to explain how Leydig came to overlook the true 
elands, unless it is that in the species studied by him the epi- 
thelial crypts are actually the only approach to the form of glands 
present. This is not probable, however, and further attention 
given to the structure of the gastric mucosa in A. nacari and 
nasus, the species employed by Leydig in his research, will in all 
likelihood show glands not differing to any appreciable extent 
from those observed by me in A. rubicundus. 

The cells of the tubules may be classed according as they 
oceupy the neck or the body of the gland, Those in the neck 
evidently do uot secrete pepsin, as they lack granules and gene- 


' Untersuchungen iiber Fische und Reptilien, Berlin, 1853, p. 16 ; also Lehrbuch 
der Histologie, 1857, p. 313. 

2 Arch. fiir Mikr. Anat., Bad. xiii. p. 651. 

3 Lehrbuch der Vergleichenden Anatomie der Wirbelthiere, p. 581. 

VOL, XX. 28 


614 MR A. B. MACALLUM. 


rally take a uniform stain with hematoxylin or carmine. They 
are transitional in their form and characters between the ordinary 
epithelial cells and those of the body of the tubule, being similar 
to the latter in size and shape. In this part of the gland the 
lumen is distinct in the body, usually it is not. The true gland 
cells are rhombohedral in shape ; the nucleus in each is placed in 
the half of the cell removed from the lumen, and the contents of 
the central half are very granular and difficult to stain. In pre- 
parations made from a Sturgeon which had undergone previously 
a prolonged period of hunger, the staining which these cells took 
was more uniform. Ordinarily they are seen, when isolated, to 
have each a fine process, which, when the cell occupies its posi- 
tion in the tubule, is directed downwards between the membrana 
propria and the cell next below. These processes are to be ob- 
served in the gastric glands of many fishes, for example in Perea, 
Amiurus, Gasterosteus, Stizostedion, Esox, &c., and they seem to 
point out that the development of epithelial cells into gland cells 
is not a complete one in fishes, so far as form is concerned. 

These glands are more numerous, but shorter near the pylorus. 
In the pylorus they are replaced by the less numerous mucous 
tubules, which are very much elongated, and provided each with 
a wide lumen (fig. 9). The cells constituting them are of two 
kinds : short goblet cells aud columnar cells, the latter being 
sometimes wholly replaced by the former. The columnar cells 
are closed and their contents slightly granular. In the goblet cells 
the greater part of each is formed of a moderately swollen theca, 
the contents of which are clear, unstainable, and apparently 
formed of mucigen. The nuclei in both forms of cells are 
placed next the membrana propria of the tubule, These mucous 
clands are evenly distributed throughout the pylorus; the re- 
placement of the cardiac glands by them is an abrupt, not a 
eradual one. 

In Amia one or more gland tubules may open into each short 
crypt of the epithelium. As in Acipenser the cells of the neck 
and those of the body of the tubule are quite different. Those of 
the body are provided with fine processes also, The lumen is 
distinguishable usually only as a line in the axis of the tubule, 
but it is seen more clearly in transverse section (fig. 8). The 
gland cells are small and but shghtly granular. Since all my 


THE ALIMENTARY CANAL AND PANCREAS. 615 


preparations were made from specimens of Amia which had 
gone without food for nearly two months, Iam unable to speak 
of the condition of these cells during activity. As it is, they and 
their nuclei stain deeply and nearly uniformly. The cells of 
the neck are fewer in number compared with those of the body 
of the gland than in Acipenser. In the pylorus they become 
more numerous at the expense of the gland cells. About the 
middle of the extent of the pylorus the neck cells form about 
three-fourths of the tubule, while a few millimetres farther on 
the gland cells vanish completely. The tubules then in the 
posterior half of the pylorus are formed of cells, not differing 
from those in the necks of the cardiac glands except in the 
ereater abundance of cilia. 

In Lepidosteus the gland cells in the cardiac tubules are small, 
but their nuclei are large. Granules were not observed in the 
interior of the cells during either their resting period or their 
functional activity. Naturally a different result would be 
obtained in the case of a fully adult form. The cells stain uni- 
formly in carmine or hematoxylin. Mucous glands, such as are 
figured by Edinger from the posterior part of the stomach, were 
not developed in the specimens from which my preparations 
were made. In the shallow cavity answering to a cecum the 
few tubules present had a larger lumen than those in the main 
portion of the stomach, but are shorter in length. 


THE Mip-Gvt. 


The mid-gut in Acipensev has a different arrangement as 
regards its coils and loops from what it has in either Ama or 
Leprdosteus. 

In Acipenser the anterior or duodenal portion is a straight 
tube of about 12 cm. length (in a specimen measuring 66 em.), 
and terminates in a median portion of 7 cm., which is directed 
forward to pass into the posterior, or valvate section of the 
intestine. The latter portion, which tapers into the rectum 
or hind-gut, contains the spiral valve. The median part is 
as a rule of smaller calibre than the others, from which it is 
marked off by a slight constriction at each end. The valvate 
part is of largest diameter and provided with the thickest wall. 
It is supplied with a vein and an artery, both of considerable 


616 MR A. B. MACALLUM. 


size, which accompany the coils of the spiral valve about the 
axis of the intestine. The course of the two vessels appears 
usually very distinct through the muscular layers and the 
serosa, but sometimes the amount of pigment present is such as 
to obscure them. It is noticeable in the other portions of the 
intestine, as well as here, that pigment tissue is distributed 
chiefly along the course of the vascular channels. The valvate 
portion of the mid-gut measures about 20 cm. while the hind- 
gut is not more than 1 cm.; the former is also greater in length 
than the anterior and median sections taken together. 

In Amia the anterior straight portion of the mid-gut measures 
18 cm. (in a full grown specimen of 55 em.), nearly occupying 
the length of the peritoneal cavity. It is connected on the left 
with a median portion of 9 cm., which is directed forward, and 
terminates opposite the czecum of the stomach in the posterior 
part answering to the valvate section of the mid-gut in 
Acipenser, but having a very small portion of itself occupied 
by the spiral valve. The rectum or hind-gut does not measure 
more than 2 cm. The spiral valve is 3 cm. long, the total length 
of the mid-gut in front of it being 32 em. 

In figures of the intestine in Amia which have been given 
hitherto, the mid-gut is represented wrongly with a calibre 
ereatly decreasing as it extends backward. The only decrease 
in size which occurs at all is to be found in the rectal portion. 

In Lepidosteus there seems to be a great difference in the 
number and arrangement of the loops, according to the size of 
the specimen examined. In young forms it is as represented in 
fig. 1. Consulting, however, the figure of the intestines, as given 
by Balfour and Parker! in their work on the structure and 
development of Lepidosteus, one sees there a more complicated 
looping of the mid-gut, which, according to these authors, 
possesses three compact coils. The specimen of Lepidosteus from 
which their figure was drawn measured 100 cm. I was 
consequently at first of the opinion that the figure given 
here represents a very early condition, but such a condition is 
indicated by van der Hoeven? in a case where the alimentary 


1 Phil, Trans., 1882, pt. i. p. 359. 
2 “Ueber die zellige Schwimmblase des Lepidosteus,” Miiller’s Archiv, 1841, 
and pl. x. 


THE ALIMENTARY CANAL AND PANCREAS. 617 


canal measured 30 cm., and, moreover, notes made three years 
ago, when I was dissecting a smaller specimen, seem to show 
that this arrangement persists for a large part of the life of the 
fish. 

It is, however, not surprising that great differences should be 
found in the mid-gut of Zepidosteus and of fishesgenerally. I have 
elsewhere! called attention to the unequal growth of this part 
of the intestine in Amiurus nigricans during its life history. In 
this fish it was shown that the mid-gut of one of 60 cm. body- 
length measured more than treble that of a specimen of 38 cm. 
This disproportion in the growth of the mid-gut, in relation to 
the growth in length of the body, is noticeable in other fishes 
also. In consequence, the difference in the arrangement of the 
loops in the mid-gut in Zepidosteus is explained on the ground 
that an increase in size of the body calls for a greater increase 
in the length of the principal part of the absorptive tract, which 
is thereby thrown into a larger number of coils. 

The arrangement of the loops in a young Lepidosteus is inter- 
esting on account of its similarity to that in the mid-gut of 
Amia. It appears probable then that this was the disposition 
of the parts in the mid-gut in both genera before the spiral 
valve had acquired its present rudimentary condition. 

The pyloric valve in Acipenser and Ama takes the form of a 
tube projecting into the cavity of the mid-gut for a centimetre 
ormore. A valve like this is present in Lepidosteus, according 
to Balfour and Parker; it is present in young forms of 7-10 cm. 
length, but itis not fully developed. In Acipenser and Amia 
it is provided with a thickened ingrowth of the fibre of the 
muscularis nucose. 

The inner surface of the anterior, or duodenal section of the 
mid-gut in Acipenser is covered with a thick mucous membrane 
honey-combed with crypts of various size, the openings of the 
majority of which are very plainly visible. There is otherwise 
but little unevenness of surface. If the membrane is in a bad 
state of preservation, however, or if its epithelium has been 
macerated away, it then presents the appearance of a network 
of minute folds, a condition erroneously ascribed to the intact 
membrane. In the median part of the mid-gut, that which is 


1 Proceedings Canad. Inst., Toronto, new series, vol. ii. No. 3. 


618 MR A. B. MACALLUM. 


directed forward, the crypts are as a rule smaller and less 
numerous, while the mucosa apparently is thinner. 

In Amia the mucous membrane of the anterior section of the 
mid-gut shows a prominent network of folds, at the junction 
of every two of which occurs a prolongation outwards in the 
form of a villus. The surface is uneven, and the crypts visible 
to the naked eye seem to be but shallow cavities between the 
folds. As in 4cipensev, the median portion possesses in its 
mucosa the characters of the anterior section, but in a less 
marked degree: its folds are smaller, its surface smoother, and 
its crypts less numerous. 

In my preparations of Lepidostcus the mucous née is 
folded alike over the whole of the mid-gut, and presents insink- 
ings of the epithelium to form structures analogous to the crypts 
in Acipenser and Amia. 

In Acipenser the mucous membrane of the valvate section of 
the intestine is as thick as that of the anterior or duodenal 
section, but in the size and the number of the crypts it is like 
that of the median part. The crypts are all but absent from 
the surface of the spiral valve. In Amia the corresponding 
section of the mid-gut is in its anterior two-thirds lined with a 
mucosa of a character like that of the median portion which is 
directed forward. In the posterior third, containing the spiral 
valve, the mucosa of its wall, between the threads of the spiral, 
is provided with narrow longitudinal folds, oblique, or transverse 
ones being wanting. The larger crypts visible to the naked 
eye, occurring on the spiral valve, which is almost as smooth as 


in Acipenser, are very few in number and may be completely — 


absent. 

The number of turns in the spiral valve in Acipenser is usually 
eight, and the distance between them averages about 2°5 em. 
The valve appears sometimes thin and membranous, at other 
times it is found to be very thick, the difference in size being 
no doubt due to the degree of distension to which the lymph- 
und blood-vessels are subjected. This portion of the intestine 
is always provided with a thick and unyielding muscular 
wall. 

In Ama the spiral valve makes nearly four turns. It is not 
as thick nor as high as in Acipenser, and the turns of the spiral 


THE ALIMENTARY CANAL AND PANCREAS. 619 


are more closely approximated, being about three-fourths of a 
centimetre apart. 

In Lepidosteus the valve makes three and a half turns, and in 
specimens of 7-10 cm. in length is so large as to fill completely 
the lumen of the intestinal tube. There is evidently a reduction 
in the number of turns of the spiral in advancing age, for, 
according to Balfour and Parker, in the fully adult specimen 
examined by them the valve does not complete the second turn. 

A transverse section of the anterior portion of the mid-gut in 
Acipenser shows under the microscope a mucosa crowded with 
elongated crypts or tubules, each possessing a large lumen of 
varying diameter. ‘They may open separately on the epi- 
thelial surface, or several may be united so as to open by a 
large common mouth. They are a continuation of thuse of the 
pylorus, but have acquired a differently constituted epithelium, 
which is, however, the same as that of the ordinary surface of 
the anterior part of the mid-gut. It is formed of one layer of 
cylinder cells, but there are at least two layers of nuclei observ- 
able (tig. 11). The nuclei of the lower layer are those of young 
cells destined to replace the cast-off or waste cells above them, 
The superficial cells have granular contents, often holding also 
fatty particles, and they possess a hyaline peripheral wall sur- 
mounted by a short fringe of cilia. The latter are to be 
observed only in carefully preserved material, and then they 
present very often the appearance of a coat of mucus on the 
ends of the cells.. Goblet cells are not common. 

In Amia a somewhat different appearance is presented by a 
transverse section of the anterior part of the mid-gut. Here the 
tubules densely crowding the mucosa are very much elongated, 
rarely branch, and radiate straight outward from the epithelial 
surface. These, as well as the epithelium, are constituted of 
short cylinder cells resting on four or five layers of nuclei. 
The cylinder cells are, apparently all of them, closed _peri- 
pherally, and are fringed with strong cilia. It is impossible 
to trace in hardened preparations the outlines of the cells for 
more than a third of the depth of the epithelial stratum, the 
cause of this difficulty being the granulation and the extreme 
abundance of nuclei which are no doubt those of young 


cells. 


620 MR A. B. MACALLUM. 


The adenoid tissue of the mucosa is very scanty between the 
tubules, this being specially the case in Aza. 

In Lepidosteus the tubules of the mid-gut are, in my prepar- 
ations, but wide-mouthed pouches clothed with cylinder cells, 
having only one series of nuclei. The cells are longer and 
slenderer than in Acipenser and Amia, with delicate, often not 
detectable, cilia. Goblet cells are common; they occur very 
often in an exhausted condition; the body of the cylinder is 
more slender than usual, a small portion of the peripheral end 
remaining swollen and containing mucigen. 

The histological relations of the median portion of the mid-gut 
in Amia and Acipenser are the same as those of the anterior 
portion. In the section containing the spiral valve in both 
genera, the epithelium is different in some respects, and the 
tubules fewer and shorter, while the adenoid tissue of the 
mucosa is more abundant. Goblet cells are more common, and 
the cylinder cells larger and more granular in their contents, 
the cilia also being longer and thicker. 

In Acipenser the epithelium of the spiral valve is constituted, 
one-third at least, of goblet cells. The cylinder cells are very 
much elongated, and if observed in fresh condition they usually 
appear charged with fatty granules. Their peripheral ends are 
closed by a hyaline wall, covering which is a long brush of 
cilia (fig. 13). 

Many of these cells, when isolated carefully after maceration, 
show very fine processes which are sometimes dendritic at their 
terminations. The fact that the processes, more frequently 
appear to be divided than is the case with other epithelium of 
the alimentary tract, may be due to the greater ease with which 
they are isolated from the fibrous tissue of the mucosa, without 
injury or breakage. The outer halves of the goblet cells are 
much inflated. Their transformed contents are very granular, 
and project beyond the epithelial border in the form of a 
stopper. 

In Amia and Lepidosteus the epithelium of the spiral valve 
does not differ from that of the neighbouring intestinal wall. 


1 According to F. E. Schulze these cells are provided with striated outer borders. 
It is possible that what was taken by this observer for a striation was in reality 
a fringe of cilia. I have been unable to find the original description by Schulze, 
and only saw a reference made by Edinger to it. 


Carine 404 


THE ALIMENTARY CANAL AND “PANCREAS. 621 


In Acipenser the tubules of the spiral valve are comparatively 
few in number and usually very short. The majority of them 
are but shallow insinkings of the epithelium, and in all of these 
goblet cells are very numerous. There are others, however, 
which differ somewhat from them, although a further study of 
fresh material may sbow both forms to completely resemble one 
another. Such tubules, of which but very few were seen, open 
into shallow pouches of the epithelium, and being much 
elongated they sometimes reach through half the thickness of 
the valve. They lack goblet cells. The fringe of cilia is 
much shorter and finer than that of the ordinary epithelium. 
Whether these structures are the representatives of the tubules 
in the anterior part of the mid-gut and in the neighbouring 
intestinal wall I do not know. They are probably derived from 
the latter, and if the present description of them is correct they 
form a class of structures which are unique in themselves, an: 
in all likelihood perform the functions of the Lieberkiihnian 
elands in higher vertebrates. In Amal have not found similar 
structures on the spiral valve, the tubules and crypts of which 
are alike in every respect to those found on the ordinary 
intestinal wall. 

In the axis of the spiral valve in Acipenser and Amica there is 
present a large quantity of unstriated muscular fibre, which in 
the last-named genus is aggregated into a single bundle. In 
Acipenser there are several bundles, arranged irregularly in 
direction and position. In vertical sections of the valve they 
sometimes appear separated by large lacunar spaces, which 
probably are the parts of lymph channels. At other times a 
muscular bundle appears almost surrounded by a closed follicle 
tilled with leucocytes. In every case the bundles are distinctly 
marked off from the adenoid tissue of the mucosa. Scattered 
so irregularly as they are through the valve, it is almost im- 
possible to conceive that they represent excessively developed 
portions of the muscularis mucose. 

Lymph follicles are very abundant in the spiral valve of 
Acipenser, as many as seventeen having been counted in a single 
vertical section. They are usually round or oval in shape when 
near the epithelium, but of irregular form when nearer the centre 
of the valve, and always of varying size. When very numerous 


622 MR A. B. MACALLUM. 


the valve appears to be made up in bulk almost wholly of them. 
Sometimes they are placed so near to the epithelium that the 
latter appears to rest immediately on them. Around each 
follicle ordinarily the connective tissue is collected into a dense 
sheath, in which one can observe a number of corpuscles 
(fig. 14). The structure of the interior of the follicle can be 
made out only after the greater number of its cellular elements 
are removed, and this is accomplished by well agitating a thin 
section of the valve in water. Then, when stained, the close 
fibrillar network comes out clearly in the interior of the follicle, 
which still contains a few lymph corpuscles in the meshes of 
this network. At the same time, too, there can be seen in 
transverse section a large number of arterioles scattered over 
the field of the follicle. 

Hyrtl + observed in the axis of the spiral valve in Acipenser 
ruthenus a large compact lymphoid organ, which he considered to 
be homologous with a similarly situated structure in Lepidosiren. 
Ayers” has found lymphoid capsules in the same species, and 
has given a figure of a vertical section of the valve to compare 
as to these capsules with one of the valve in Lepidosiren. They 
are undoubtedly homologous structures, although the anterior 
part of the organ in Lepidosiren, found outside the intestine, is 
lacking in Acipenser. A compact organ such as Hyrtl observed 
dues not exist in Acipenser rubicundus, in which lymphoid 
follicles are more highly developed than in the species used by 
these authors for investigation. 

Scattered collections of lymph corpuscles are very common in 
the various parts of the intestinal tract in fishes, and as such 
they simply represent an overloading of the adenoid tissue of the 
mucosa and submucosa with these structures, the limits of the 
deposit being rarely sharply definable. A compact lymph organ 
has been observed in the cesophagus of some Selachians, and, 
according to Edinger,’ who has given the most detailed account 
of it, it possesses a capsule of connective tissue, from which 
trabeculz of fibrils penetrating the interior divide and redivide, 

1 «*Tepidosiren paradoxa,” Abhand. der bihm Gesell. der Wiss., 1845. I have 
had no oppertunity for consulting this work, but found a quotation made by 
Ayers from it, which contains the view referred to. 

2 « Beitrige zur Anat. and Phys, der Dipnoér,” Jenaische Zeit., Bd. xviii. 

3 Op. cit. 


THE ALIMENTARY CANAL AND PANCREAS. 623 


cells similar to those of lymph being enclosed in the meshes 
thus formed. It would seem, from the descriptions of this and 
other authors, to be constant in presence and position, and it 
cannot therefore be one of those accidental infiltrations of the 
tissue with leucocytes which are so commonly found in every 
part of the intestine. It appears then that structures which 
may be correctly termed lymph follicles are to be found in the 
intestinal tracts of Acipenser, Dipnoi, and some Selachians only 
among fishes. Further research may, however, correct or extend 
this list. 

In Acipenser the lymph follicles of the spiral valve are the 
representatives of the closed follicles forming the Peyer’s patches 
in the higher vertebrates. They can usually be seen by the help 
of a magnifying hand-glass, as minute, whilst spots unequally 
distributed in the valve, which has, contrasted with them, a dull 
white colour. At some points they are so densely crowded 
together as to form an almost complete resemblance to a Peyer’s 
patch, 

The remaining part of the mucosa in the neighbourhood of 
the spiral valve in Acipenser and Amia.is loaded with lymph 
corpuscles, although not nearly to the same extent as the follicles 
of the former. 


Enb-GUt. 

The relative lengths of this part of the intestine in Acipenser 
and Amaia have been given above. 

In some specimens of Acipenser there was practically no end- 
gut, the last turn of the spiral valve extending to the anal aperture. 
Where, however, it acquires any length, the mucosa covering it 
is either smooth or thrown into slender longitudinal folds, and 
is sometimes provided with minute crypts. The epithelium 
consists of cylinder and goblet cells, which gradually shorten to 
cubical cells, and at the vent to flattened cells, forming several 
layers in thickness. Anteriorly the cylinder cells bear a short 
fringe of cilia. 

In Amia the inner surface of the end-gut is folded longi- 
tudinally ; sometimes it is smooth and provided with crypts. 
When the epithelium is removed by maceration, the surface has 
a reticulated appearance. The epithelium consists of ciliated 


624 MR A. B. MACALLUM. 


cylinder cells with a few goblet cells. 1t undergoes a gradual 
decrease in height towards the vent. 

In Lepidosteus the epithelium is of the character of that of the 
mid-gut, but the cilia are thicker and longer. At the vent the 
cells become cubical, which form gives no evidence of the 
possession of cilia. 

I have observed in Lepidosteus the mesentery which connects 
the end-gut with the ventral wall of the peritoneal cavity, and 
which was discovered and described by Balfour and Parker.’ 


THE PyLoric APPENDAGE. 


The pyloric appendage in Acipenser and Lepidosteus differs 
from similarly situated organs in other fishes, in that it is a 
compound structure with its pouches communicating with the 
intestinal cavity by a common duct, whereas in the great 
majority of fishes each pouch opens singly into the mid-gut. 
The latter arrangement is the primitive one; it can be seen in 
very young Lepidostei, in which the ceca arise as isolated out- 
crowths of the intestinal wall, and assume later a common duct, 
this being the manner of development in the Sturgeon also, 
according to Balfour. 

In Acipenser the organ is flattened on its inferior face, and 
appears uniform. The duct opens on the left side of the mid- 
eut, about a centimetre from the pyloric vaive. It is of such a 
dimension in very large Sturgeons as to permit readily the intro- 
duction of the index finger into it for some distance, but in 
Sturgeon of ordinary size it is of the diameter of a goose-quill. 
Ten to twenty ceca, arranged in a radiate fashion, open into it at 
a distance of a centimetre or so from its intestinal aperture, all 
the czeca being enclosed in a common sheath of muscular and 
connective tissue. The mucosa of the duct and ceca is arranged 
in folds, crypts, and tubules, in much the same way as it is in 
the mid-gut. When denuded of its epithelium it appears 
possessed of the same characteristic network of the subepithelial 
tissues. 

In a thin vertical section of the appendage, placed under the 
microscope, the structure of the mucosa is seen to differ un- 


t ODF tit. 
2 Comparative Embryology, vol. ii. p. 632. 


THE ALIMENTARY CANAL AND PANCREAS. 625 


essentially from that of the mid-gut. The epithelium possesses 
more nuclear layers, the cilia are shorter, the cylinder cells 
slightly larger, and the goblet cells more extended than in the 
intestine. The connective tissue between the tubules is very 
abundant, and at various points in it there are collections of 
leucocytes. 

The two muscular layers of the mid-gut are extended over the 
duct and cca of the organ, but on the latter are arranged 
obliquely with regard to each other. 

In Lepidosteus the organ appears more lobed, and presents the 
shape of a bean in young forms, in which it extends over the 
inferior face of the gall bladder and pancreas for some distance. 
Its lobes or ceca are very numerous, the common duct into 
which they lead opening into the mid-gut immediately behind 
the pyloric valve. ‘The epithelial lining of both duct and ceca 
is of exactly the same character as that of the mid-gut. 

In Ama there is no pyloric appendage. 

Various theories have been put forward as to the function of 
these organs. The earliest opinion, and one for a time generally 
adopted, was that they act collectively as a pancreas in the 
absence of the latter organ. Whatever support this view had 
was taken from it when it was shown that pyloric ceca and a 
pancreas may consist in some fishes, notably Lota. 

In 1860, Mordecai! advanced the view that these organs 
collect and store up fluid nutritious matter, which, during the 
passage of the fish to its spawning ground, is absorbed and 
utilised to repair the wasting processes of the body in the 
absence of ordinary food. His observations were made on 
Alosa prestabilis (Clupea sapidissima, Wilson), in which there 
are from sixty to one hundred separate ceca, and these, during 
the ascent of a river by the fish, were found distended with a 
brownish mucus, which was absent at other times of the year. 
Mordecai found also that the number of the ceca increase with 
age, and that in Zabrax and Dvioplites, some rudimentary in 
younger forms are functional in the fully adult. 

According to Edinger,? the ceca serve to suck up the liquid 


1 His pamphlet is republished in the Bull. U.S. Fish Commission, 1881, 
p. 227. 
2 Op. cit. 


626 MR A. B. MACALLUM. 


and digested food matter as it escapes from the stomach, and 
absorb it. Wiedersheim! also adopts this view, and tries to 
show that the development of the pyloric appendages stands in 
inverse relation to that of the spiral valve. He mentions two 
instances of this: Polypterus, in which the pyloric appendage is 
but a short ezcal pouch, while the spiral valve is highly 
developed, and ZLepidosteus, which has a well-developed pyloric 
appendage, but no spiral valve. This view of Wiedersheim, it 
must be said, cannot be held as generally correct, even when 
applied to the Ganoid fishes alone, for in Acipenser, in which 
the cxeca being provided with a common duct is an evidence of 
the great development of these organs, there is a well-marked 
spiral valve, and in Ama, in which the spiral valve is almost as 
rudimentary as in Lepidosteus, there are no pyloric ceca. 

Krukenberg ? found in the ceeea of Acipenser sturio evidence 
of the presence of diastase, pepsin, and trypsin. In other fishes 
these organs gave extracts containing one or more of the same 
enzymes. Krukenberg is, however, inclined to believe, after a 
long series of experiments, that the pyloric appendages perform 
specially the function of absorption. 

According to Stirling’s® view, these organs subserve different 
functions in different fishes, being absorptive in some, and in 
others glandular, secreting trypsin in some cases in important 
quantities. 

Blanchard + found that the extract of the pyloric cea in 
various fishes effects the transformation of boiled starch into 
orape sugar, and that when boiled white of egg or fibrin is added 
to the extract, whether this is alkaline, acid, or neutral in its 
reaction, peptones are formed. He believes that a tryptic 
ferment alone is present to accomplish the latter result. 

During the summer of last year, I made a number of experi- 
ments with extracts of the pyloric organ in Acipenser, in order 
to determine the presence of the enzymes, which Krukenberg 
affirms he found there. For this purpose large fish were 
employed. The organ after its removal was slit open, the 
mucous matter in the duct and cieca quickly washed away, to 


' Lehrbuch der Vergl. Anat., p. 565. 

* Kiihne’s Untersuchungen, Ba. ii., 1882. 

3 Journal of Anat. and Phys., vol. xviii. p. 426. 
4 Comptes rendus, xevi. 1241-1244, 


Spb Gato Sin 


THE ALIMENTARY CANAL AND PANCREAS. 627 


remove such enzymes as might have gained entrance from the 
intestine, and which would vitiate the result. It was then in 
some cases finely minced, and in this condition covered with 
distilled water in a flask surrounded by ice for twenty-four 
hours ; in others it was carefully dried, and in a finely divided 
state extracted at a temperature of 35° C., with a 0-2 per cent. 
solution of hydrochloric acid, or with a 0°5 per cent. solution of 
sodic carbonate. With extracts made with distilled water, no 
conversion of starch into grape sugar could be obtained, and in 
no case could the presence of the latter substance be detected. 
Almost always an emulsion occurred when a quantity of the 
extract was shaken with some olive oil and allowed to stand. 
From the acid or alkaline extracts of the finely divided organ 
I was unable to obtain the slightest traces of digestion when 
fibrin or boiled white of egg was treated with them; if, however, 
the organ was slightly sponged, instead of being quickly washed 
with water, traces of pepsin and trypsin were found, but no 
diastase, although grape sugar could be detected in very slight 
quantities. The presence of trypsin can be easily explained in 
this case, as the opening of the pancreatic duct is just opposite 
that of the pyloric organ, and in the first contraction of the 
intestine after the pancreatic fluid escapes some of the fluid 
must find its way into the duct and ceca, even if it does not 
reach them by any other means, ¢.., diffusion. The pepsin 
gains access in like manner from the fluid contents of the 
stomach, immediately after passing through the pyloric orifice. 

The difference between the results obtained by Krukenberg 
and those here detailed, as to the presence of enzymes in the 
pyloric appendage of <Acipenser, may be accounted for on the 
ground that in all the material used by me the stomach and 
intestine were completely empty, while it is barely possible that 
it may have been otherwise with that employed by Krukenberg, 
since enzymes are more likely to be present in detectable 
quantities when food matter is in the intestinal canal. 

Although it is an important matter to establish whether the 
enzymes referred to are present or not in the pyloric cca, and 
in what quantities, yet such an accomplishment must not be 
allowed to decide definitely what the function of the organs is 
in the great majority of fishes. Pepsin and trypsin may be 


§28 MR A. B. MACALLUM. 


normally present as a result of their easy diffusion, as is some- 
times observed in organs not immediately connected with the 
alimentary canal in higher vertebrates, or they may be present 
as the result of the active secretion of the ceca. That such a 
secretion can occur and does, is possible, apart from the fact 
that in certain fishes, eg.,in the cod, according to Stirling, the 
czeca are provided with gland structures of the usual form. It 
is true that the caca are lined with eylinder cells usually, to 
which there has not yet been attributed the power of secreting 
either pepsin or trypsin. But when it is seen that the long 
eylinder cells lining the intestinal tract and the so-called liver 
in Amphioxus must secrete whatever enzymes are used in the 
digestion of the food swallowed, one is compelled to ask, Why 
should not a somewhat similar epithelium in the pyloric ceca of 
fishes perform a like function? The secretory function of this 
epithelium, as Stirling points out, does not exclude that of 
absorption, this also being exemplified in the case of Amphioxus. 
We see, therefore, that it is not easy to determine definitely the 
function of these organs in all fishes ; in some, absorption may 
overshadow secretion, in others, the reverse may occur, while io 
others again, the two functions may occur in equal proportions. 
This would explain the contradictory results obtained by 
different observers in this field of research. 

In any case, it is fair to conclude that the pyloric céca are 
rudimentary structures once actively glandular. They are in 
their development remarkably similar to other outgrowths of 
the intestinal wall in their immediate neighbourhood, @e., the 
liver and pancreas. It seems extremely probable that at one 
time in the history of vertebrates there opened into the 
alimentary canal, which had a simpler form than it now 
possesses, a large number of pouch-like diverticula, and that 
by gradual specialisation of some of these arose posteriorly 
the liver and pancreas, anteriorly the air bladder, and, if they 
are not homologous with the latter, the lungs, while others 
retained their primitive structure and arrangement, persisting as 
pyloric cca. In consequence of this specialisation, all the 
functions, such as those of absorption and of the secretion of 
enzymes, possessed by the original ceca, would necessarily be 
retained in an impaired degree by those persisting in that form. 


ee 


ee ante~ 


THE ALIMENTARY CANAL AND PANCREAS. 629 


It is to be expected, also, that the digesting principles secreted 
by the cca partake more or less of the characters of those 
of the primitive organs. We may in this way explain the 
occurrence of such an enzyme as Blanchard found in the pyloric 
appendages, which digests in an alkaline, acid, or neutral 
solution ; this, probably, was the character of the primitive 
seeretion, not only of these organs, but of the whole intestinal 
tract, and by a course of selection operating in the organs and 
their functions the secretion came in the course of time to 
present in the stomach the properties of pepsin, in the pancreas 
those of trypsin. We have then a clue to an explanation of 
the supposed presence of both enzymes in the pyloric ceca ; 
one enzyme in reality is present, which digests fibrin in an 
alkaline or an acid solution, and which partakes of the 
characters of both pepsin and trypsin. 


THE PANCREAS. 

It is over half a century since the first observation on 
the pancreas of fishes was published, and it is only twelve 
years ago that the presence of such an organ was demonstrated 
to be general in this class. To Legouis belongs the honour of 
having accomplished the latter task. 

This observer! has given a full history of the researches of the 
various anatomists on the subject, and on this account further 
reference to the general literature bearing on it may be avoided. 
With regard, however, to that treating of the pancreas of Ganoid 
fishes a few words here are necessary. 

In 1833 Alessandrini? discovered a pancreas in the Sturgeon. 
After a number of anatomists had failed to confirm this dis- 
covery, Leydig’ recognised the accuracy of Alessandrini’s 
description, and, what the latter had not done, gave a short 
account of the histological structure of the organ. 

Alessandrini’s and Leydig’s statements have not been confirmed 
by the researches of a third observer up to the present date. 
Wiedersheim ‘ describes as the pancreas an organ extending from 


1 Ann. des Sciences Nat., 1873. 

2 Ann. des Sciences Nat., t. xxix., 1833; also Acad. Scien. Inst., Bonon, 1835, 
tom, il. 

3 Untersuch, tiber Fische und Reptilen, Berlin, 1853. 

4 Lehrbuch der Vergleich. Anat., p. 605. 

VOL. XX. 2T 


630 MR A. B. MACALLUM. 


the pyloric appendage down the left side of the anterior portion 
of the mid-gut, while Krukenberg! found no organ specially 
performing the function of a pancreas, and he maintains that 
the organ of Wiedersheim, which he regards as the spleen, is 
neutral as to digestion, its extracts having no action whatsoever. 
Ayers? has given a figure of a section of the supposed pancreas, 
and he recognises it as constituted of lymphoid tissue. 

No reference has hitherto been made in the literature to the 
occurrence of a pancreas in Ama. 

In Lepidosteus Balfour and Parker* found a pancreas in young 
forms, it being situated, according to their description, behind 
the liver in the loop of the pyloric section of the stomach. 
They found also that it arises in the embryo in the same 
manner as in other vertebrates, and that late in larval life it 
envelops the anterior section of the bile-duct. In the fully 
adult Lepidosteus they observed a small, apparently glandular, 
mass connected with the bile-duct, and occupying a position 
similar to that of the pancreas in the larva. They were unable 
to decide on its nature, as its poor preservation did not permit a 
histological examination. 

In Acipenser, the organ considered by Wiedersheim to be the 
pancreas is really the spleen, as a careful study of its histology 
will show. . 

The true pancreas in this fish is a disseminated one, 2.¢., its 
tissue is not localised, but variously extended throughout the 
the right half of the peritoneal cavity. It occupies on the whole 
the position described by Alessandrini and Leydig. Its tubules 
and ductlets entwine about the branches of a_blood-vessel 
(arteria ceeliaco-mesenterica), which it accompanies down the 
valvate portion of the intestine, on the median and anterior 
portions, and into the liver (fig. 16). It was in sections of the 
last-named organ that I found it first. It is more disseminated in 
some specimens of Acipenser than in others; for example, in one 
I found that it extended 8 cm. down the length of the valvate 
portion of the mid-gut, while usually the distance extended is 
only about 3 em. The greater bulk of the organ is found 


1 Kiihne’s Untersuchungen, Ba. i. and ii. 
* Jenaische Zeitschrift., Bd, xviii. p. 479, taf. xvii. fig. 52. 
3 Phil. Trans., 1882, pt. ib p. 359, 


THE ALIMENTARY CANAL AND PANCREAS. 651 


between the liver and the second bend of the mid-gut on the 
right side, and is easily detectable from the thickness of the 
walls of the blood-vessel referred to, consequent on the pancreatic 
tissue enveloping them. Asa branch of this artery enters the 
accessory spleen (fig. 16, asp), vertical sections of the latter 
organ sometimes betray the presence of the pancreatic tubules. 

The serosa covering the pancreatic tissue and the vessels has 
usually a dark colour, sometimes perfectly black, owing to the 
presence of a large number of pigment cells. 

The portions of the pancreas which are attached to the intes- 
tinal walls are always closely applied to the latter and included 
within the serosa. 

The pancreatic duct opens into the mid-gut on the dorsal side; 
its aperture and that of the bile duct being found on a prominent 
papilla on the inner intestinal surface, not more than halfa 
centimetre usually from the tip of the pyloric valve. I have 
never succeeded in injecting the pancreatic duct with coloured 
fluids so as to show its course. It is easily distinguishable from 
the bile duct by the greenish tinge which the latter always has. 

In Acipenser sturio Alessandrini found the bile duct and the 
pancreatic duct to open on separate but similar papillz on the 
_ inner wall of the mid-gut, that on which the opening of the 
pancreatic duct is found being placed more than 2 cm. from the 
pyloric valve. I cannot find this to be the case in Acipenser 
rubicundus, in which, by snipping off the point of the single 


papille above referred to and spraying the stump with water,< 


one can easily see the two apertures placed side by side on it. 

In Amia the pancreas is disseminated in a manner similar to 
what it is in Avipenser. A part of it is found adherent to the 
extreme anterior end of the mid-gut (fig. 10, pz), but a greater 
part envelops the larger branches of the portal vein in the 
interior of the liver. It forms a large portion of the bridge 
between the right and left hepatic lobes, and it covers, to a 
certain extent, the arch of the bile duct. 

The pancreatic duct is here also hard to trace. I did this, 
however, in a young Amia of about 6 cm. in length, by means 
of aseries of vertical sections made by Professor Ramsay Wright, 
who was the first to observe the presence of a pancreas in this 
fish. A careful dissection of the tissues about the bile duct in 


632 MR A. B. MACALLUM. 


the adult gave the same result as the study of the series of 
sections. The pancreatic duct is much narrower in diameter 
than the bile duct, to which it is nearly parallel, and near the 
intestinal wall they both have. a common investment of con- 
nective tissue, the one thereby appearing to open into the other. 
They remain completely separate, and their apertures are to be 
found on the apex of a slender papilla placed about 1:5 em. from 
the pyloric valve. The course of the two ducts is from before 
backward, almost in a line with the anterior part of the mid- 
gut. 

In Lepidosteus the pancreas is not so widely disseminated as 
in either Acipenser or Ania. It envelops the portal vein as it 
runs along partially sunken in the dorsal face of the liver, in the 
posterior third of this organ, while it reaches behind accompany- 
ing the same vein to a point opposite the posterior extremity 
of the pyloric appendage (fig. 1, P). Some of its tubules are 
adjacent to the superior face of the gall bladder, others twining 
about the bile duct for some distance. In the liver the tubules 
radiate frequently out from the vicinity of the portal vein 
between the hepatic lobules. Both the bile duct and that of the 
pancreas run side by side backward to the mid-gut, the pan- 


creatic duct being placed on the gastric side of the other. — 


Adjacent to the wall of the mid-gut the two ducts become fused. 
In a transverse section of this part of the body it is possible to 
distinguish this common duct from a pyloric czecum only by 
tracing it from behind forwards, since the histological structure 
of both near their termination in the intestine are alike in every 
particular. Opposite the pyloric appendage, and above the 
pylorus, is a small quantity of lymphoid tissue, which, though it 
contains a few tubules of the pancreas, is not the latter organ, 
us Balfour and Parker suppose. This, usually as large in 
vertical section as the mid-gut, and, traversed by the portal vein, 
seems to be an accessory spleen, the pancreatic tissue forming 
but a very small part of it. 

I have not had an opportunity of studying sections of the 
liver of an adult Lepidosteus, but I have no hesitation in saying 
that such sections, if taken through the posterior third of the 
organ, would show well-developed pancreatic tissue surrounding 
some of the larger vascular channels near the dorsal surface, and 


“a 


THE ALIMENTARY CANAL AND PANCREAS. 633 


that the pancreas in this fish is not inferior, structurally and 
functionally, to what it is in either Acipenser or Amia. 

With regard to the finer histology of the pancreas in the 
three genera a short description only can be given here, as the 
quantity of material at my disposal did not allow very varied 
methods of preparation. 

In Acipenser the structure is very much as in the higher 
vertebrates. In the necks of the tubules, which may also be 
termed the intermediary canals of the alveoli, the lining 
epithelium is constituted of cubical cells. In the alveolar 
portions the cells are large, the nuclei distinct and arranged in 
the peripheral halves of the cells, while the central halves are 
filled with granules. In my preparations the nucleus may be 
said to be the only part of the cell which stains in carmine or 
hematoxylin, as it is rare that the protoplasm in the immediate 
vicinity of the nucleus takes any tint at all. The alveolar 
Iumen is usually distinct and of a zigzag course. Transverse 
sections of the tubule in its various parts exhibit the usual 
characteristic appearances. 

Among the cells and in the lumen of each alveolus, there are 
other cellular elements, which are noticeable at first sight on 
account of the deep and nearly uniform staining which they take, 
and also on account of their varied shape, sometimes oval or 
polyhedral, and sometimes amceboid. Granules are lacking, the 
protoplasm appears clear, and no reticulation was observable in 
it. The nuclei always stain much more deeply than those of 
the ordinary cells of the alveoli. The cells are oftenest to be 
found in the lumen, and then they appear sometimes to send out 
processes between the gland cells (fig. 17, C.). They are, I think, 
remarkably similar to the centro-acinar cells of Langerhans. I 
have never met with these structures in any other fish, 

The arrangement of the tubules about a branch of the arteria 
celiaco-mesenterica is indicated in fig. 17, A. Trabeculee of 
connective tissue radiate outward between the tubules to the 
serosa, the latter possessing a large number of pigment cells, to 
which the dark colour commonly associated with the pancreas in 
this fish is due. 

In Amia the pancreatic tubules imbedded in the liver are 
twined about the larger branches of the interlobular veins in the 


634 MR A. B. MACALLUM. 


manner represented in fig. 18. The connective tissue of the 
interlobular septa run between and separate the neighbouring 
tubules, which in a vertical section of the liver appear cut in 
various directions, oftener longitudinally. The cells of the 
aveoli are, in their central portions at least, crowded with 
cranules, and stain very feebly apart from their nuclei. Near 
the lumen the limits of the cells are indistinct, owing to the 
quantity of granules present. The lumen sometimes is, and 
sometimes is not, visible. 

In Lepidosteus an alveolar and an intermediary portion can be 
distinguished in each pancreatic tubule. 


The pancreas would appear to be rudimentary or absent in 
but very few fishes. There are conditions when such an organ 
seems to be useless, and these are found in small fishes only, 
such as some of the Cyprinoids, in which digestion by the stomach 
alone prepares enough of nutritious material for absorption to 
satisfy the demands of the organism, and in which the mid-gut 
does not serve to submit the food matters escaping from the 
gastric cavity to a second digestive process but to a further 
absorption. No such reason can be advanced for the supposed 
absence of the pancreas in large fishes. Its presence is so 
diseuised in many forms that one finds it impossible to decide 
definitely whether it is absent in others. In Ganoids, as above 
described, it is partially imbedded in the liver, in some Siluroids ! 
completely so. Why may it not be connected in a like manner 
with the liver in other forms in which it is now supposed to be 
absent? That this is frequently the case is probable from the 
occurrence of trypsin in the extracts of the liver of various fishes, 
as shown by Krukenberg. The liver and pancreas thus fused 
into one organ can of course in no case be considered as a 
hepato-pancreas. 

! In Amiurus the pancreas is imbedded in the interior of the liver, its tubules 


being entwined about the interlobular veins, (see Proceedings Canad. Inst. , 
Toronto, new series, vol. ii. No. 3). 


THE ALIMENTARY CANAL AND PANCREAS. 635 


EXPLANATION OF PLATE XX. 


Fig 1. The alimentary canal and pancreas of Lepidosteus, from a 
specimen measuring 10 cm. a, cesophagus ; /, the commencement of 
the stomach anteriorly ; ¢, the pylorus; ¢, the anterior or duodenal 
section of the mid-gut; f, the posterior, or valvate, section of the 
same; at, the hind-gut; ap, the pyloric appendage ; », the pancreas. 
(Reduced. ) 

Fig. 2. A longitudinal section of one of the papille of the so-called 
cesophagus in Acipenser’. 

Fig. 3. An oblique section of a dilated sac from the wsophagus of 
Acipenser. 

Fig. 4. A transverse section of a dilated sac from the cesophagus of 
Amia. 

Fig. 5. A tubule from the mouth of the air-duct in Amia. 

Fig. 6. A vertical section of a fold of the mucosa of the cesophagus 
of Lepidosteus, showing ciliated epithelium and dilated sacs. 

Fig. 7. Two glands from near the posterior part of the cardia in 
Acipenser. 

Fig. 8. A gland from the cardia of Amia. a, one seen in trans- 
verse view. 

Fig. 9. Part of a mucous gland from the pylorus of Acipenser. 

Fig. 10. A vertical section of the wall of the anterior part of the 
mid-gut in Amia. pn, the pancreas; m, the mucosa; 7, the inner, 
o, the outer, muscular layer. 

Fig. 11. Epithelium of a crypt of the mid-gut in Ac/penser. 

Fig. 12. Part of a vertical section of the spiral valve in Acipenser, 
showing the short pouches of the epithelium, and the mucosa under- 
neath richly laden with lymph corpuscles. 

Fig 13. Epithelium of the same section more highly magnified. 4, 
a young epithelial cell. 

Fig. 14. A section of a lymph follicle from the spiral valve of 
Acipenser ; the greater number of lymph corpuscles have been re- 
moved from the section by agitating it in water. 

Fig. 15. Epithelium of a tubule of the spiral valve of Ama. 

Fig. 16. A view of the mid-gut and the organs attached to it in 
Acipenser. Ap, the pyloric appendage; sp, the spleen; «a.sp, the 
accessory spleen; p, the main portion of the pancreas surrounding a 
branch of the arteria celiaco-mesenterica ; p', p", pancreatic tissue 
adhering to the smaller branches of this vessel, and to the wall of the 
mid-gut ; /, the liver, the course of the pancreatic tissue in which is 
represented by dotted lines ; m, the anterior portion of the mid-gut ; 
v, the valvate portion of the same. 

Fig. 17. A, A section of a branch of the A. caliaco-mesenterica in 
Acipenser, with pancreatic tubules surrounding it, a portion only of 
these being represented ; B, a part of one of the tubules more highly 
magnified ; C, a transverse view of one of the tubules. In both B and 
C the centro-acinar cells have a darker shading. Bb and C highly 
magnified. 


636 THE ALIMENTARY CANAL AND PANCREAS. 


Fig. 18. From a vertical section of the liver in Ama, showing the 
manner in which the pancreatic tubules are arranged around a larger 
interlobular vein ; pt’, one of the tubules seen in transverse section ; ~ 
vt, an interlobular vein filled with blood-corpuscles. 

Fig. 19. From a vertical section of the dorsal face of the posterior ~ 
part of the liver in Lepidosteus, showing the arrangement of the 
pancreatic tubules pé about the portal vein (vp); h, liver tissue; J, 
the bile duct. 


THE NEURAL SPINES OF THE CERVICAL VERTEBRAi 
AS A RACE-CHARACTER. By D. J. Cunnincuay, 
M.D. (Edin. et Dubl.), Professor of Anatomy in the 
University of Dublin. . 


Sin Ricwarp OWEN,! in one of the numerousimportant articles 
which he has contributed upon the Anatomy of the Anthropoid 
Apes, has compared the vertebral column of the Gorilla, Orang, 
and Chimpanzee with that of several of the different races of 
man. In this he remarks that “in the skeleton of a Boschisman, 
as well as in that of the female Australian in the Museum of 
the Royal College of Surgeons, the spines of the five lower 
cervical vertebre are simple.” He further observes that the 
neural spines of the 3rd cervical vertebra in a male Australian 
is “not bifurcated as usually in Europeans.” 

Before my attention was drawn to these observations of Sir 
Richard Owen, I had collected statistics regarding the condition 
of the neural spines of the cervical vertebrze in a number of the 
lower races of man, and, as these extend considerably the data 
recorded on this subject, I have thought that their publication 
might not be without some interest. It will generally be noticed 
that in low races the cervical neural spines are relatively shorter 
and more stunted than in the European. More especially can 
this be observed in the case of the neural spine of the 3rd 
cervical vertebra. This condition is obviously for the purpose 
of allowing a greater freedom of movement in the cervical 
section of the spine, and is to be associated with other peculi- 
arities present in the lumbar region of the low races which give 
to the loin segment a greater flexibility and suppleness.2 But 
the feature which especially attracts attention is the condition of 
the extremities of the cervical neural spines. As Owen has 
pointed out in the case of the Australian and Bushman, the 
cervical spines in the low races are as a rule non-bifid or only 
very slightly so. 


1 “Comparison of the Lower Jaw and Vertebral Column of the TZ'roglodytes 
gorilla, Troglodytes niger, and Pithecus satyrus and Different Varieties of the 
Human Race,” Trans. Zool. Soc., 1851, vol. iv. 

2 Vide Nature, Feb. 16, 1886, ‘‘'The Lumbar Curve,” by D. J. Cunningham. 


638 PROFESSOR D. J. CUNNINGHAM. 


With the view of determining the comparative frequency of 
this character in the different races of man, I have examined 
the spines of 15 Europeans, 7 Australians, 6 Negroes, 4 
Andamans, 3 Tasmanians, 1 Bushman, and 2 Esquimaux, and in 
recording the results which I have obtained I must give expres- 
sion to my sense of the deep obligations under which I lie to Dr 
J. G. Garson of the Royal College of Surgeons in England, and 
to Professor Macalister of Cambridge. The number of spines of 
the low races in the Trinity College Museum were ‘too few in 
number to allow of me carrying out this investigation in Dublin, 
and I had necessarily to ask the permission of these gentlemen 
to examine the specimens under their charge. This was 
accorded to me in the freest manner possible. 

The spine of the 7th cervical vertebra is never bifid, and 
that of the 6th is rarely cleft even in the European. In the 
following tables, therefore, the condition of the spines of the 2nd 
Srd, 4th, and 5th cervical vertebre is alone recorded. It will be 
observed that the spines are arranged in three groups—(1) the 
first comprises those which are distinctly bifid; (2) the second 
includes those in which there is merely an indication of bifidity ; 
and (3) the third those which show no trace of it whatever. 


Condition of Neural Spines of Cervical Vertelre in Fifteen 


Huropeans. 
{ 
| Bifid. Feebly Bifid. Non-Bifid. 
2nd cervical vertebra, . : é 15 ok 
3rd cervical vertebra, . 5 ‘ 10 3 2 
4th cervical vertebra, . : + a 9 t 2 | 
5th cervical vertebra, . : E 9 5 1 


Condition of Neural Spines of Cervical Vertebree in Seven 


Australians. 
Bifid. Feebly Bifid. | Non-Bifid. 
2 
2nd cervical vertebra, . : «7 7 “he sri 
3rd cervical vertebra, . : ral 2 see | 5 
4th cervical vertebra, . ; Ay 1 2 4 
5th cervical vertebra, . : : 2 2 3 


NEURAL SPINES OF THE CERVICAL VERTEBR&. 639 


Condition of Neural Spines of Cervical Vertebree in Three Tasmanians. 


| 
| Bifid. Feebly Bifida. Non-Bifid. 
2nd cervical vertebra, . 4 : 3 | ae < 
| 3rd cervical vertebra, . , : eS 1 2 
| 4th cervical vertebra, . ; : ix 1 2 
_ 5th cervical vertebra, . : : aii 1 2 


Condition of Neural Spines of Cervical Vertebrae in Six Negroes. 


Bifid. | Feebly Bifid. | Non-Bifid. | 
= a an Ss = | 
2nd cervical vertebra, zy 3 i! 
3rd cervical vertebra, if 2 3 
4th cervical vertebra, = 1 1 4 | 
5th cervical vertebra, . f : | 1 2 3 


Condition of Neural Spines of Cervical Vertebru: in Four Andamans. 


Bifid. Feebly Bifid. | Non-Bifid. 
| 
| | 
2nd cervical vertebra, . : : 3 1 Re: 
3rd cervical vertebra, 2 | 2 
4th cervical vertebra, 2 2 
5th cervical vertebra, 2 2 


In one Bushman the neural spine of the 2nd cervical vertebra 
was bifid; that of the 5th feebly bifid, and those of the 5rd and 
4th non-bifid. In two Esquimaux the neural spines were feebly 
bifid throughout in one specimen, whilst in the other the only 
neural spine which was bifid was that of the 2nd vertebra ; 
those of the remainder were non-bifid. 

The following table gives the combined results of the 24 
low races that have been examined :— 


Condition of Neural Spines of Cervical Vertebiw in Twenty-four of 
the Low Races of Man. 


| 
Bifid. Feebly Bifid. Non-Bifid. | 
Qnd cervical vertebra, . 2 : 18 5 1 
3rd cervical vertebra, . ; . | 3 6 15 
4th cervical vertebra, . : a 2 7 15 
5th cervical vertebra, 3 10 11 


640 NEURAL SPINES OF THE CERVICAL VERTEBRA, 


By comparing the results given in this table with those which 
are recorded for the Europeans, a very marked difference in the - 
condition of the neural spines will be observed. The only 
cervical vertebra which presents an invariably bifid neural spine 
in all the races is the axis vertebra. In only one instance, viz., 
in a Negro, have I found it non-bifid. This character, however, 
is shared by the Chimpanzee, and also in some cases a trace of 
the same bifidity may be noticeable in the corresponding neural 
spine of the Gibbon, and likewise even in the Orang and Gorilla. . 

In the Europeans the neural spines of the 3rd, 4th, and 5th 
cervical vertebree are as a rule bifid. In only one case did I 
find all these spines non-bifid in the same individual. 

In the low races, however, the reverse condition is the rule, 
and in this respect they resemble the corresponding neural 
spines of the Chimpanzee. They differ from the latter, however, 
in being more stunted and not so sharp at the point. 

It will be observed from the tables that have been given that 
the two neural spines in the low races which are least frequently 
bifid are the 3rd and 4th. These also are the most stunted of 
the series. By this arrangement the longer spines above and 
below are allowed to close over them in extension of the neck. 


[Zditorial Note—As additional references, we may say that 
M. Hamy, in his excellent description of the skeleton of an 
Aéta Negrito woman (Nouv. Archives du Museum, 1879, vol. ii.), 
has also directed attention to the tendency in the coloured races 
for the cervical spines, the axis excepted, to be non-bifid, and a 
number of examples have also been recorded by the writer of 
this note in his “Challenger” Report on the Bones of the 


Human Skeleton, part ii, now in type, and shortly to be 
published. W. T.] 


VARIATIONS IN THE NERVE SUPPLY OF THE 
FLEXOR BREVIS POLLICIS MUSCLE. By H. Sr 
JOHN Brooks, B.A. and M.B. (Dnb.), Demonstrator of 
Anatomy in the University of Dublin. 


OnE of the earliest facts impressed upon the student, when he 
takes up the study of Practical Anatomy, is the relatively slight 
tendency shown by nerves to vary; in particular, the nervous 
supply of muscles is supposed to be immutable. This dogma 
has been extended to Comparative Anatomy by the Heidelberg 
School, and Professor Ruge “asserts, with Gegenbaur, that a 
muscle is to be regarded as the end-organ of a nerve, and 
therefore when a muscle alters in position and connections, its 
original and typical relations can always be identified by its 
nerve of supply.” ! 

In the foot, where in man the muscles are arranged upon a 
more primitive plan than in the hand, we find that the abductor 
and flexor brevis pollicis are supplied by the internal plantar 
nerve, while all the other intrinsic muscles receive their supply 
from the external plantar. It is interesting to note the deviation 
from this fundamental arrangement among the Mammalia as 
worked out by Professor Cunningham. He says :— 

“Tn the Elephant the internal plantar nerve supplies the 
flexor brevis indicis; in the Hyrax the internal plantar nerve 
supplies the flexor brevis indicis, the adductor indicis, and the 
second dorsal interosseous muscle; in the Beaver a still more 
remarkable deviation is found. From the internal plantar nerve 
proceed the twigs of supply for the abductor hallucis, flexor 
brevis indicis, flexor brevis medii, and the first and third dorsal 
interossei. Lastly, in the Fox-bat there is an example of the 
external plantar nerve encroaching on the domain of the 
internal plantar, by supplying a twig to the outer head of the 
flexor brevis hallucis.” ? 

This is a most interesting case of two nerves struggling, as it 

? Morphologisches Jahrbuch., 1878,—quoted by Professor Cunningham. 


Challenger Reports, part xvi. p. 49. 
2 Op. cit., p. 135. 


642 MR H. ST JOHN BROOKS. 


were, for a group of muscles, sometimes the internal plantar 


gaining ground, at other times the external plantar extending. 


its domain. 

In this paper I have to record a similar struggle (if we may 
call it so) between the two homologous nerves of the human 
hand, the median and the ulnar; and we shall find that the 
nerve-supply to muscles is not immutable, even when our 
observations are confined to one animal—Man. 


Table of Variations in the Nerve-Supply of Flexor Brevis 
Pollicis in Man. 


Outer head supplied by deep branch of ulnar alone, . . 5 cases. 
Outer head supplied by ulnar and median, : : . SOR 
Outer head by median, inner head by ulnar, : : Seow 
Median giving twigs to both heads; inner head also with 

an ulnar nerve-supply, ; : . ; , i ar 


In some of these cases the twig from the deep branch of the 
ulnar nerve to the outer head was large; in others very small. 
The size of the twigs from the median was in inverse proportion 
to those from the ulnar. 

We are told in the text-books, Continental as well as English, 
that the outer head of flexor brevis pollicis is supplied by the 
median nerve, and the inner head by the deep branch of the 
ulnar. This seems at first sight a deviation from the funda- 
mental plan as observed in the foot, until we remember that 
dischoff! has shown that the muscle usually described as inner 
head of flexor brevis is in reality a part of the adductor, the 
true inner head lying concealed beneath it on the metacarpal 
bone of the thumb, and described by Henle as the interosseus 
primus volaris. I have not as yet been able to trace the 
nervous supply of this true deep head of the flexor brevis, but 
in the two cases in my table, in which the median gave twigs to 
the inner head, I believe some filaments passed through it to 
reach the interosseus primus volaris. | 

It may be well to note that the outer head of flexor brevis 
usually consists of two parts; a large mass of fibres from the 
annular ligament and os trapezium (this is the true outer head), 
and a fasciculus from the inner head, which crosses the deep 
surface of the long flexor tendon to join the more superficial 


1 Beitrage zur Anatomie des Hylobates leuciscus, 1870, ps 20, 


~ See ees 


NERVE SUPPLY OF FLEXOR BREVIS POLLICIS MUSCLE. 643 


part at ils insertion; it is the latter portion that one would 
naturally expect to receive an ulnar nerve-supply, but in every 
case both parts of the outer head received twigs. In the seven 
eases in which Vne outer head was supplied by the median alone, 
I found that in one case the contribution from the deep head 
was absent, in another it. was very small; in the remaining five 
this point was not noted. I found it absent in both hands of a 
young female Chimpanzee, which was kindly placed at my 
disposal by Professor Cunningham; the outer head was here 
.supplied by median alone. [ am inclined to think that the 
fasciculus of fibres from the deep to the superficial head has 
acted as a bridge, and, as it were, dragged the branch of the 
ulnar nerve across. In one of the cases given in the above table 
the ulnar had encroached in a very unusual manner on the 
domain of the median ; after giving twigs to the outer head of 
flexor brevis, it pierced the latter and supplied the opponens and 
abductor pollicis. The branches to the two latter muscles were 
as large as those which they normally receive from the median, 
and appear to have replaced the latter entirely. 

At first sight it appeared as if this trespassing of the ulnar 
nerve on median territory might be explained by a communica- 
tion between the median and ulnar in the arm, but in both arms 
of the Chimpanzee there was a very large communicating branch 
from the mediau to the ulnar in the fore-arm, and yet the outer 
head of flexor brevis was supplied by the median only. 

As the distribution of the ulvar nerve described above has 
apparently escaped the notice of anatomists, a few words on the 
method of exposing it may not be out of place. The nerve in 
question usually comes through the adductor pollicis near its 
radial margin, and, passing on the deep surface of the long flexor 
tendon, pierces the outer head of flexor brevis. Sometimes it is 
entirely concealed by the adductor, and may be brought into 
view by separating the heads of the flexor brevis. It is sur- 
rounded by parallel strands of connective tissue, which give it an 
intensely deceptive appearance. In looking for this nerve I 
begin by securing the branches of the median to the outer head; 
these I find sometimes close to the annular ligament in company 


} Thad the pleasure of showing this rare case to Professor Cunningham, and 
also to the University anatomist, Dr T. E. Little. 


644 NERVE SUPPLY OF FLEXOR BREVIS POLLICIS MUSCLE. 


with the nerves to abductor and opponens, often lower down 
from the first or second digital branches, sometimes in both 
these situations. I then rip up the sheath of the long flexor 
tendon, pull the latter aside, and separating the two heads of 
flexor brevis, by pushing without cutting, I see something 
stretching across which looks like a strand of connective tissue. 
I follow this into the muscle, and its nervous character becomes 
evident. I then take a note of the name of the student who is 
dissecting the part, and when he is doing the deep dissection of 
the palm he invites me to trace the nerve, and I verify its 
connection with the ulnar. 

The first example of this nerve I found last winter session 
(1884-85), while dissecting a hand in the Dissecting Rooms of 
Trinity College, Dublin, and, supposing it to be a very rare 
anomaly, I showed it to Professor Cunningham. I found that 
he had an entry of a similar case in his note-book, and also the 
record of the dissection of the hand of a negro from Sierra Leone, 
in which the median gave twigs to both heads of flexor brevis. 
[ must take this opportunity to express my thanks to Professor 
Cunningham for placing these two cases at my disposal. 

In the tive cases given in my table in which the radial head 
is said to be supplied by ulnar alone, the branch from the ulnar 
to the outer head was large; in two ont of the five the whole 
dissection was made by myself, and after a very careful search 
I could not find any twigs from the median to the outer head. 
In the other three cases the integument had been removed 
before I examined the subject. 

As in 19 out of 31 cases the outer head of flexor brevis had 
a double nerve-supply,! I think we should be justified in regarding 
this as the normal arrangement. 


1 From ulnar and median. 


ON THE MORPHOLOGY OF THE INTRINSIC MUSCLES 
OF THE LITTLE FINGER, WITH SOME OBSERVA- 
TIONS ON THE ULNAR HEAD OF THE SHORT 
FLEXOR OF THE THUMB. By H. Sr Joun Brooks, 
B.A. and M.B, (Dub.), Demonstrator of Anatomy in the 
University of Dublin. (PLATE XX1.) 


THE intrinsic muscles of the hand, more especially the 
marginal groups known in Human Anatomy as the “ thenar and 
hypothenar eminences,” have been frequently made the subject 
of investigation, but more from a descriptive point of view 
than with the object of elucidating their morphological value. 
In works on Human Anatomy great differences of opinion 
have prevailed as to the division and even the number of the 
short thumb muscles. Henle describes a double-headed flexor 
brevis distinct from the interosseus primus volaris, and also 
separate from the “outer head of flexor brevis” as ordinarily 
described in English text-books. The latter he terms a deep 
part of the abductor. Gegenbaur, on the other hand, ignores 
the presence of an ulnar or deep head altogether. Bischoff, first 
threw light on the true ulnar head of flexor brevis pollicis, 
which he shows to be identical with the interosseus primus 
volaris of Henle, but he does not appear to think that the 
muscles of the little finger call for any special attention. Pro- 
fessor Young,” of the Owens College, Manchester, has contributed 
some valuable papers on the muscles of the hand, and adopts 
(provisionally) the classification first proposed by Professor D. J. 
Cunningham,? of dividing the muscles of the manus and pes 
into three layers— 

1, A plantar layer of “ adductores.” 

2. An intermediate layer of “ flewores breves.” 

3. A dorsal layer of “ abductores.” 

Professor Cunningham has established this classification for the 
foot, and has shown that the origins of the marginal members 
1 Beitrige zur Anatomie des Hylobates leuciscus, 1870. 

2 Jour. of Anat. and Phys., vols. xiv. and xvi. 


3 Zoology of the Challenger, part xvi. pp. 19 and 48. 
VOL, XX. 2U 


646 MR H. ST JOHN BROOKS. 


have a tendency to wander towards the heel. The abductores 
hallucis and minimi digiti are thus the marginal and enlarged 
“dorsal interossei.” One of the most potent arguments by 
which Cunningham has established this theory is the fact, that 
in tetradactylous animals the origin of the (now marginal) 
abductor indicis is found farther back than the other interossei, 
unless confined by a rudimentary metatarsal. He adopts, with 
some reservation, the statement of Ruge,! that all the muscles 
which lie superficial to the deep division of the external plantar 
nerve are “contrahentes” or “adductores.” But, he remarks, 
“the great objection to this method of classifying the adductors, 
however, is that it is incapable of being extended to the hand. 
In the manus the deep division of the ulnar nerve passes inwards, 
under cover of the flexor brevis minimi digiti and also under (or 
through) the opponens minimi digiti, both of which therefore, 
according to this generalisation, would be looked upon as 
contrahentes or adductores.” ” 

The author believes that the facts detailed in this paper will 
show that this “flexor brevis minimi digiti” and the part of the 
opponens superficial to the deep ulnar nerve are really 
adductores. The true ulnar head of flexor brevis quinti in man 
is represented by that part of the opponens which lies beneath 
the deep ulnar nerve, and the radial head by the “ third palmar 
interosseous.” Further, the dissections described will tend to 
show that the true deep head of flexor brevis pollicis is very 
frequently indistinguishable ; it is difficult to say whether it is 
suppressed, or fused with the adductor obliquus, the valuable 
guide of nerve relation being absent on the radial side of the 
hand. 

With a view to establishing these points, I have examined 
the manus in Man, Chimpanzee (Zroglodytes) Orang (Pithecus), 
Cynocephalus anubis, Macacus nemestrinus, Colobus, Marmoset 
(Hapale), Cat (Felis domestica), Capybara (Hydrocherus), 
Virginian Opossum (Didelphys virginiana) and Australian 
Opossum (Phalangista vulpina). These animals were all most 
kindly placed at my disposal by Professor Cunningham from the 
stores of the Anatomical Department of Trinity College, Dublin. 


' Morph. Jahrbuch., bd. iv. p. 645. 
2_Op, cit., p. 136. 


MORPHOLOGY OF THE INTRINSIC MUSCLES. 647 


As I have to trace the wandering, both in origin and insertion, 
of the adductor or contrahentic! group of muscles, it will be well 
to mention what appears to be their typical condition. They 
are a group of flattened muscles, springing from a common 
pointed tendinous origin from the ligaments on the palmar 
aspect of the carpo-metacarpal articulation of the middle digit, 
and separated from the subjacent interossei by the deep branch 
of the ulnar nerve.2 They diverge to be inserted into the 
proximal phalanges in such a manner as to adduct the fingers 
to a line which usually passes through the middle digit. The 
origins of the two marginal members (a1,a°, fig. 2) show a great 
tendency to wander distally and thus overlie and dwarf the 
others ; not only this, but they also spread towards the sides of 
the carpus, creeping along the concavity of a curved plane, formed 
partly by the unciform ‘and os trapezium and partly by the 
anterior annular ligament. The insertion may also vary. 
Cunningham figures and describes a slip to the radial side of 
index finger in the hand of Cuscus, also a slip to the fibular side 
of the fourth digit in the foot of the Koala.? Ruge* describes 
and figures an adductor or contrahens passing to the fibular side 
of the fifth digit in the foot of Dasyurus hallucatus, while 
Cunningham figures a similar muscle in D. vivervinus, but 
describes it as “one of the abductors of the minimus,” regarding 
the nerve relation as exceptional.® 

In estimating what is a typical condition of adductores or 
contrahentes it appears to me that we should regard symmetry 
in size and direction of fibres rather than the presence of the 
full complement of muscles; the same remark applies to the 
flexores breves. The Cat affords a favourable example, It 
possesses two adductors for the index and little fingers, arising 


1 First described in the foot of Macacus by Halford of Melbourne, and named 
by him ‘‘ contrahentes digitorum.” He states that they are absent in the hand 
of the same animal (p. 12). In both of the specimens I dissected the contra- 
hentes were well represented. ‘‘ Not like man, bimanous and biped, nor yet 
quadrumanous, but cheiropodous.” Melbourne, 1863. 

2 Ruge (op. cit.) established this nerve relation for the contrahentes of the foot, 
and it is evidently applicable to the hand also. 

3 Zoology of the Challenger, part xvi., plate ii. fig. 3, a, and plate vii. fig. 2. 

4 Op. cit., p. 647. 

5 Op. cit., pp. 55 and 136; and plate xi. fig. 5, d® As far as I can judge 
from the figure the origin is close to the other adductors, 


648 MR H. ST JOHN BROOKS. 


from the ligamentous structures of the central part of the carpus, 
and lying superficial to the deep ulnar nerve (fig. 1, a,a°). There 
are four paired flexores breves for the four ulnar digits. These 
are equal in size, and each arises near the base of the correspond- 
ing metacarpal by a pointed origin, which soon bifurcates into 
two symmetrical fleshy bellies, to be inserted into the ulnar and 
radial sesamoids. These all lie beneath the deep ulnar nerve. 
There are two short thumb muscles, one (f1) representing the 
radial head or flexor brevis and the other (a!) probably the 
adductor pollicis or first contrahens, as its origin is associated 
with the other adductores. If this view be correct, there is no 
trace of the ulnar head of flexor brevis, There is an abductor 
minimi digiti, but the muscle known in Human Anatomy as 
“flexor brevis minimi digiti” 1 is entirely absent. 

In the Marmoset a very similar condition of the flexores 
breves is found, but we have here a different arrangement of the 
true flexor brevis minimi digiti (/); while the radial head 
(f°r) is inserted into the sesamoid as before, the ulnar head ( f*7) 
has no insertion into the phalanx either direct or indirect, but 
is inserted into the whole length of the metacarpal bone, form- 
ing an opponens. There is a “ flexor brevis” minimi digiti (a°a) 
arising from the hook of the unciform and from the anterior 
annular ligament and inserted with the abductor. It is 
relatively larger than in Man, and is separated by a small 
interval from the adductor quinti digiti; the latter and the 
adductor pollicis have spread in both directions in the manner 
indicated above. They thus unite in a median raphe as far as 
the metacarpo-phalangeal joint, and conceal the other two 
adductors (a* and a?); they also travel along the curved plane 
towards the little finger and thumb respectively. An abductor 
pollicis and a radial head of flexor brevis pollicis are also 
present, but the ulnar head of the latter is either suppressed or 
fused with the adductor. 

In Capybara we have chiefly to notice a remarkable case 
of suppression, while there are three powerful double-bellied 


As this name is confusing on account of the existence of a true flexor brevis 
of the little finger, I shall distinguish the ‘‘ flexor brevis” of Human Anatomy by 
inverted commas, and by the letters aa (adductor quinti aberrans). The true 


flexor brevis minimi digiti is indicated by the letters f°7=radial head and f®u= 
ulnar head, 


MORPHOLOGY OF THE INTRINSIC MUSCLES. 649 


flexores breves for the index, middle, and ring fingers, and the 
ulnar head of flexor brevis minimi (/*w), is well represented, 
the radial head (/°7) is not merely entirely wanting, but its 
place is not taken by any other muscle. The two adductors 
(to index and minimus) have the typical arrangement, and the 
adductor minimi remains comparatively remote from the vacant 
space where the radial belly of flexor brevis ought to be. We 
shall compare this with the Opossum later on. There is an 
abductor, but no sign of “flexor bevis” minimi digiti (aa). 
The abductor indicis shows, in a marked degree, the tendency 
of marginal muscles to take their origin from a point more 
proximal than the rest of the group (the thumb being ab- 
sent). 

In the Virginian Opossum there exists a very remarkable 
condition of the adductor quinti digiti' A large triangular 
sheet of muscle, like that in the Marmoset, is present, but this 
extends further ulnarwards and arises from the median raphe 
and the ligamentous structures at the base of the metacarpus 
(fig. 2, a). This origin is continuous with another muscular 
mass arising from the base of the hook of the unciform (a*a’), 
and diverging to be inserted into the ulnar sesamoid of the fifth 
finger. No sign of separation can be seen between these two 
muscles for at least a quarter of their length. Another muscle 
(a®a) lies quite parallel to the one last described, and arises from 
the tip of the unciform hook and from the annular ligament ; 
it is separated by a slight areolar interval from a°a’, but the 
origins and insertions of the two muscles are absolutely adjacent 
and their fibres are perfectly parallel. The muscle (a*a) is 
evidently the “flexor brevis” of Human Anatomy, as the 
origin, insertion, nerve-supply, and nerve relation are the same. 
There is a well-formed and typical abductor minimi having 
the same connections as in Man (fig. 2, abd*), and there is also a 
muscle arising from the annular ligament near the origin of aa 
and running parallel to the radial side of the latter to give the 
perforated tendon to the fifth digit,—this is a flexor brevis 
digitorum manus. The broad adductor pollicis extends its origin 
radial-wards along the curved plane, and has concealed and 
coalesced with, or suppressed, the true ulnar head of flexor 
brevis pollicis. On removing the muscles which lie superficial 


650 MR H. ST JOHN BROOKS. 


to the deep branch of the ulnar nerve a complete set of flexores 
breves come into view (fig. 3). On two of these, however, 
doubt may be thrown,—(1) the ulnar head of flexor brevis 
pollicis is probably only an artificially separated part of the 
adductor; (2) the minute radial head of flexor brevis minimi 
is partially blended with the fourth dorsal interosseous, and the 
existence of an arched tendon from the latter, “attached by one 
extremity to the distal end of the fifth metacarpal bone and by 
the other to the adjoining side of the first phalanx of the ring 
finger,” as described by Professor Young! renders the dis- 
crimination more difficult. In both hands, however, dissecting 
from both palmar and dorsal aspect, I was able to trace a small 
fasciculus to the radial sesamoid of the fifth finger.2 The 
ulnar belly (f/°w), a very small but perfectly distinct muscle, 
arises chiefly from the unciform but partly from the base of 
the fifth metacarpal, and is inserted into the ulnar sesamoid of 
the fifth finger. Both heads of the flexor brevis minimi are 
easily and naturally separable from a* and a*a’,even without the 


aid of the deep ulnar nerve, which passes between them and. 


makes “assurance doubly sure.” 

Professor Young has arrived at results somewhat different 
from the above.2 He describes the abductor minimi (fig. 2, 
abd°) as an opponens. (It arises from the pisiform bone, and 
is inserted into the ulnar side of the base of the preximal 
phalanx of the little finger. I could not find any fibres inserted 
into the metacarpal bone.) The muscle aa he calls the 
abductor, and aa’ the ulnar head of flexor brevis ; he has also 
a separation,t of which I could find no trace, in the adductor 
minimi, marking off the ulnar margin of it as the radial belly of 
the flexor brevis. 

It will be instructive at this stage to compare the foot of 
the same animal (fig. 4). Here we have an “annular ligament ” 
presenting a superficial resemblance to the anterior annular 
ligament of the hand, though its connections are very different ; 
but it has this property in common, that it affords a cwrved 

' Journal of Anat. and Phys., vol. xiv. p. 153. 3 

* It will be remembered that in Capybara this muscle #7 was entirely absent, 
though there could not be any fusion with another muscle in that case. 

3 Op. cit. 

* Op. cit., plate vii. fig. 1 ; a line between adm and f°. 


stead Mise bhi. ocd aaeiliien 


MORPHOLOGY OF THE INTRINSIC MUSCLES. 651 


plane along which the origin of a muscle can creep; and we 
have here a muscle (a°a) whose insertion and nerve relation is 
identical! with the muscle similarly lettered in the hand, and 
whose general position appears very similar. Between this and 
the adductor quinti (a), we have an interval in which the deep 
branch of the external plantar nerve appears lying superficial to 
a well-developed double-bellied flexor brevis quinti. We note 
the complete absence ef anything corresponding to a?a’ (fig. 2), 
also the slighter lateral extension of the origin of the adductor, 
quinti, therefore the lesser degree of functional replacement of 
and therefore the better development of flexor brevis minimi 
digiti. In the foot, unlike the hand, the fibular belly of flexor 
brevis hallucis (/1/°) is quite distinct, and arises in common with 
the tibial head considerably proximal to the adductor. I found 
an indication (not shown in the figure) of the separation into 
adductor obliquus and a. transversus described by Ruge? in 
Didelphys cancrivora. Ruge, however, gives no fibular head of 
flexor brevis hallucis for this species, and also omits all mention 
of the muscle a°a which is present in D. virginiana; the latter 
muscle is figured and described (after Young) by Cunningham,’ 
but he calls it an additional head of the abductor, and the nerve 
relation is not shown or described. 

The Australian Opossum or Vulpine Phalanger, while cor- 
responding in a general way with the Virginian Opossum, shows 
some instructive differences. The adductor indicis is on the 
same plane as the adductor pollicis, so that adductores pollicis 
and indicis arise from the radial side of the central raphe and 
adductor minimi from its ulnar side. The muscles aa and a’a’, 
which are separate in the Virginian Opossum (fig. 2), are here 
united to form one muscle, the deeper fibres of which (corre- 
sponding to aa’) are inserted into the metacarpal bone, and are 
therefore homologous to a*op (fig. 5 and fig. 8). The muscles 
corresponding to a°a’ and a in the Virginian Opossum, although 


1 Besides the phalangeal insertion a strong fasciculus passes to the base, and a 
weaker set of fibres to the head, of the metatarsal bone. This reminds us of the 
development of an opponens from the corresponding muscle in the hand, so fre- 
quently found in higher forms, and is also an argument against regarding this 
muscle as a flexor brevis digitorum. 

2 Op. cit., p. 648, and Taf. xxxv. fig. 49. 

3 Op. cit., p. 68, and plate vii. fig. 4, dt. 


652 MR H. ST JOHN BROOKS. 


still united at their origin, are more distinct than in the latter 


animal, as there is a trace of an aponeurotic septum between. 


them. The flexor brevis digitorum manus is absent, as the 
flexor sublimis supplies a perforated tendon to the fifth finger 
(while in Virginian Opossum the flexor sublimis has only three 
tendons, to the index, middle, and ring digits). On reflecting 
the layer of adductores, the deep branch of ulnar nerve is seen 
lying on the flexores breves. The fourth dorsal interosseous 
muscle shows the arched tendon to head of metacarpal and 
proximal phalanx of ring finger, as described by Cunningham 
in Cuscus, &c., and by Young in Virginian Opossum, but on the 
palmar surface of the latter muscle there is a_ beautifully 
distinct, fusiform, fleshy slip, inserted by a minute tendon into 
the radial sesamoid of the little finger; this is the radial belly 
of the true flexor brevis (f°). The ulnar belly of the same 
muscle appears to be absent, while in the Virginian Opossum it 
is well represented (fig. 3, fw). The radial border of the 
adductor pollicis functionally replaces the ulnar head of flexor 
brevis pollicis; while arising in common with the well- 
developed outer head of flexor brevis, and inserted into the ulnar 
sesamoid of the thumb, is a distinct fibrous band, which is 
evidently the rudiment of the true ulnar head. The presence 
of this band increases the probability that the real ulnar head is 
suppressed, not fused, in the Virginian Opossum, and therefore 
the muscle shown in fig. 3 is the result of an artificial division 
of the adductor. Professor Cunningham,! in his description of 
the hand of the Vulpine Phalanger, expresses the opinion which 
I have given above, viz., that the real ulnar belly of flexor 
brevis pollicis is suppressed ;-—he regards, however, the muscle 
corresponding to aa’ and a*a (fig. 2) as the ulnar head of flexor 
brevis minimi digiti. 

In Macacus nemestvinus there are four adductors, those to the 
index and ring digits being small. There is a broad triangular 
adductor pollicis inserted directly into the radial side of base of 
the first and second? phalanges of the thumb. From the part 

1 Op. cit., p. 25. 

* If this condition is constant in Macacus it shows a curious reappearance of 
a reptilian character in a high form of mammal. A similar condition (insertion 


nto second phalanx) of the adductor minimi digiti pedis is described by both 
Ruge and Cunningham in the Ornithorhynchus, ‘The latter also describes the 


4 Ww aa Be wal 7 


MORPHOLOGY OF THE INTRINSIC MUSCLES. 653 


arising from the carpus a fasciculus diverges to be inserted into 
the radial sesamoid (compare ala in Cynocephalus). These two 
parts are intimately connected for at least the proximal half of 
their length. On separating these two the deeper fibres of the 
large triangular portion are seen to be inserted into the ulnar 
sesamoid; as the origin of these latter fibres is from the 
ligaments at the base of the second and third metacarpal bone 
they evidently form a part of the real adductor, and cannot be 
regarded as a flexor brevis. Flexor brevis radial head and 
opponens pollicis are similar to the same in Man, but there 
appears to be no trace of an ulnar head of flexor brevis. The 
muscles of the little finger differ in only one point from those of 
the Marmoset; the radial head has become rather the larger, 
and more distinctly resembles the homologous muscle in Man 
(third palmar interosseous). The condition of paired flexores 
breves for the three middle digits, however, has vanished, and in 
its place we have an arrangement of “interossei” similar to 
that in Man. 

In Colobus the thumb is rudimentary, appearing only as a 
minute projection before the skin is reflected; it possesses, 
however, a metacarpal bone and phalanges, the distal phalanx 
being the size of a pin’s head. There is no long flexor tendon, 
but the short muscles are remarkably distinct and well-defined. 
Abductor pollicis is rather weak, but otherwise typical. The 
flexor brevis pollicis arises by a pointed tendinous process from 
the annular ligament near the os trapezium, and divides into 
two bellies, of which the radial is rather the larger; these are 
inserted with the abductor and adductor respectively. The 
sesamoid bones appear to be absent. A very few fibres of the 
radial head are inserted into the metacarpal bone, but there is a 
slip from the wlnar head which forms an opponens; it is 
inserted on the ulnar side of the middle line of the shaft. The 
broad, flat triangular adductor arises from the fascial structures 
over the middle metacarpal bone, and is separated by a compara- 
tively wide interval from the flexor brevis. 

There is a “flexor brevis” minimi digiti (aa), from the 
deeper part of the origin of which an opponens is formed. The 


insertion of the adductores into the ungual phalanges in the pes of Echidna 
also a slip to the ungual phalanx of the hallux in Cuscus (op. cit., p. 57). 


654 MR H. ST JOHN BROOKS. 


latter is inserted into the whole length of the metacarpal bone 
of the little finger. There is a true flexor brevis separated from. 
the two preceding muscles by the deep branch of the ulnar 
nerve; its radial belly forms the “third palmar interosseous ;’ 
its ulnar belly is very small and forms an opponens, a few fibres 
reaching as far as the head of the metacarpal bone; it is over- 
laid and dwarfed by the greatly developed opponens derived 
from the palmar layer of adductores. 

In Cynocephalus anubis (fig. 5) there is a large adductor pollicis 
arising from a strong fibrous band over the middle metacarpal, 
and from the ligaments over the bases of index and middle 
metacarpals, a slip (aa) passes from it to be strongly inserted 
into the radial sesamoid. There is a powerful radial head of 
flexor brevis (/17) arising from the anterior annular ligament and 
os trapezium ; from the deeper part of its origin the ulnar head 
springs. It is about one-fifth the size of the radial head, and is 
inserted along the middle line of shaft of metacarpal, its most 
distal fibres inclining to the ulnar side, to be inserted close to 
the ulnar sesamoid but not reaching it. It will be remembered 
that a similar muscle forms part of the ulnar head in Colobus; 
this muscle is concealed by aa in the figure; it is separated 
from the latter by a strong fibrous sheath. Turning to the 
little finger, we find a “flexor brevis” (a°a) arising from the 
annular ligament close to the tip of the very short unciform 
hook ; intimately blended with its deeper fibres of origin we 
find an opponens (a°op) inserted into the distal two-fifths of 
ulnar border of shaft of fifth metacarpal. The deep branch of 
ulnar nerve separates the foregoing from the true flexor brevis, 
which arises chiefly from unciform and diverges to form “ third 
palmar interosseous ” (f°r), and an opponens inserted into 
the whole length of the shaft of the metacarpal (f*op). The 
adductor (@°) is in this form rather widely separated from (a°a.) 

In the Chimpanzee the “flexor brevis” minimi digiti (fig. 7, 
aa) has the same origin and insertion as in Man; from its 
deeper fibres of origin an opponens (a°op) is developed, which 
is inserted into the distal four-fifths of the fifth metacarpal 
on its ulnar border. The deepest part of the origin of this 
opponens, from the base of the unciform hook, zs united with the 
pointed proximal extremity of the origin uf the adductor minimi 


MORPHOLOGY OF THE INTRINSIC MUSCLES. 655 


digitt; the latter has also a thin scattered extensive origin from 
the fascia over the ring metacarpal. The radial head of flexor 
brevis minimi digiti (f°) resembles the “third palmar inter- 
osseous ” in Man; a few fibres spring from the unciform. Closely 
associated with this origin, the ulnar head (f*w) arises from base 
of unciform hook, and is inserted into the proximal fifth of the 
ulnar border of shaft of fifth metacarpal ; it is slightly blended at 
its insertion with the other opponens (a*op), but quite separable 
at origin; the deep branch of the ulnar nerve passes between 
them. The adductor pollicis arises from a fibrous band, which 
passes from the head of the ring metacarpal to the base uf the 
middle metacarpal, and from the ligaments over the bases of the 
- middle and index metacarpals; it shows a slight division into 
adductor obliquus and a. transversus. There is no trace of any 
slip like ata in Cynocephalus. There is a well-developed 
radial head of flexor brevis pollicis arising from the annular 
ligament close to the os trapezium, and inserted with the 
abductor. The inner head is represented by a sharply-defined 
glistening fibrous band, which arises from the extreme base of 
the thumb metacarpal, and is inserted with the adductor in 
ulnar side of base of proximal phalanx of thumb. In the speci- 
men dissected (a young female) there were no sesamoid bones. 

In the Orang there is a condition of the muscles of the little 
finger more resembling Man than any other animal I have 
examined. The “flexor brevis” minimi digiti (°c) is relatively 
larger than in Man, and the opponens which arises in connection 
with it is inserted into the entire length of the shaft of the fifth 
metacarpal bone. The adductor of the thumb is the only 
adductor present, unless the others are represented by a very 
tough fibrous tissue which covers the interossei and the deep 
branch of ulnar nerve. Flexor brevis minimi digiti radial head is 
as in Chimpanzee, but the ulnar head appears to be entirely 
wanting. The adductor pollicis has the same connections as in 
man, but the slip which passes to the radial sesamoid (az) is 
wanting. The flexor brevis pollicis arises from the os trapezium 
and annular ligament, and divides into two fleshy bellies, a small 
ulnar, and a much larger radial, which are inserted into corre- 
sponding sides of the proximal phalanx, and separated by the 
slender long-flexor tendon (¢, fig. 6). 


656 MR H. ST JOHN BROOKS. 


In Man the “flexor brevis” minimi digiti (aa) is relatively 


smaller than in any of the other animals, and is frequently - 


wanting ; intimately connected with its deeper fibres of origin 
from the unciform hook we find an opponens, which is usually 
inserted into the distal four-fifths of the shaft of the metacarpal 
(as in the Chimpanzee), -but may be limited to the lower half or 
even less (fig. 8, a°op). Arising from the shaft of the fifth 
metacarpal, and in most cases from the unciform! is the radial 
head of the true flexor brevis (“third palmar interosseous,” / °7’) 
and closely associated with the unciform origin and inserted 
into the proximal fifth of the metacarpal bone there is a small 
muscle (/°op) separated by the deep branch of the ulnar nerve 
from the first mentioned opponens, and quite distinct from the 
latter at its origin. In the specimen from which fig. 8 was 
drawn this muscle was unusually large, and at its origin formed 
one muscle with the third palmar interosscous, In the thumb we 
find a very powerful adductor, whose origin is pierced by the 
deep palmar arch, the division into adductor transversus and a. 
obliquus being thus indicated ;? the a. obliquus is described in 
English text-books as the deep or inner head of flexor brevis. 
From the adductor obliquus a strong bundle of fibres * passes to 
the radial sesamoid of the thumb; this is identical with a'a in 
Cynocephalus. The radial head of flexor brevis is strong, and 
arises from the lower border of the annular ligament close to its 
attachment to the os trapezium. The true ulnar head (interosseus 
primus volaris) arises from the extreme base of the metacarpal 
bone of the thumb, and is inserted with the adductor, some fibres 
being often prolonged on to the dorsum of the phalanx to join the 
long extensor tendon, 

It is evident from the above descriptions that the so-called 
“flexor brevis” minimi digiti of the human hand is not 
homologous to the muscle of the same name in the foot; the 
flexor brevis minimi digiti of the foot lies close to the metatarsal 
bone and beneath the deep division of the external plantar nerve, 
while in the hand “flexor brevis” is superficial to the deep 
branch of ulnar nerve. Professor Cunningham‘ has shown that 


1 Henle, Afuskellehre, p. 246. 

2 Bischoff, op. cit., p. 20. 

3 See Quain’s Anatomy, 9th edition, vol. i. p. 226. 
STOpr Cite, pale, 


MORPHOLOGY OF THE INTRINSIC MUSCLES. 657 


the flexor brevis minimi digiti of the foot and the “third plantar 
interosseous muscle” are the fibular and tibial bellies of the 
same muscle. In the Opossum (fig. 4°), and in many of the 
mammalian feet figured by Professor Cunningham;? this condition 
is very evident, the two bellies being symmetrical and the 
insertion on each side phalangeal. In the Chimpanzee I found 
in both feet the flexor brevis minimi digiti and “third plantar 
interosseous” arising together by a pointed tendinous origin from 
the sheath of the peroneus longus tendon, as far back as the 
ridge on the cuboid bone; the “interosseous” had an additional 
origin from the shaft of the fifth metatarsal. ‘The fibular belly 
was inserted into the shaft of the metatarsal, its only connection 
with the phalanx being through the abductor tendon, into which 
a few fibres were inserted. A very similar condition to this is 
described in Professor Cunningham’s notes on the dissection of 
a negro foot from Sierra Leone ;? the insertion into the phalanx 
of the flexor brevis, however, was present. In European feet the 
following is a frequent condition of the flexor brevis minimi 
digiti:—The greater part of the muscle arises (more or less 
associated with the third plantar interosseous) from the sheath 
of peroneus longus, the remainder arising from the base of fifth 
metatarsal; the insertion is partly into the metatarsal, partly into 
abductor tendon, and the remainder directly into the phalanx. 

While in the foot the fibular belly of flexor brevis minimi 
shows a tendency to become reduced to an opponens, this 
tendency is more pronounced in the hand. In the following 
animals we find the insertion becoming more and more proximal, 
and the size of the muscle diminishing pari passu:—Cat, 
Marmoset, Macacus, Cynocephalus, Colobus, Man, Chimpanzee, 
Orang. In the latter it appears to be absent altogether. Inthe 
first three we find the opponens (a°op) (which lies superficial to 
the deep ulnar nerve) absent. In the others it shows a gradual 
increase in size, functionally replacing the deeper muscle. Again 
in the Cat the “flexor brevis” (of human anatomy) is absent 
but present in all the rest, and in the Cat alone (in this series) 
we find the true flexor brevis (ulnar head) with a phalangeal 
insertion. 


1 Op. cit., plate viii. fig. 1, f°; plate xi., figs. 1, 6, and 10, 
2 Unpublished as yet. 


658 MR H. ST JOHN BROOKS. 


Having now demonstrated the true flexor brevis, let us 


examine the so-called “flexor brevis” of man. It corresponds - 


in every way with (a5a) in Cynocephalus, Opossum, and Chim- 
panzee (figs. 5, 2, and 7). In the Opossum its origin is 
continuous with the adductor minimi digiti, the gap usually 
found between their origins being occupied by a muscle (a°a’), 
which is evidently only a deeper fasciculus of a®a. This deeper 
fasciculus, which is phalangeal in insertion, shifts its insertion to 
the metacarpal bone in higher forms, and appears as a@°op in 
Cynocephalus, Chimpanzee, and Man. We have thus in the 
Opossum what is practically one sheet of muscle @°, a°a’, and aa 
separated by the deep branch of ulnar nerve from the layer of 
flexores breves. It appears probable that the adductor minimi 
digiti extends both its origin and insertion towards the ulnar 
side of the hand, and that the pressure of the long flexor tendon 
(or tendons) afterwards divides the insertion into two slips, the 
deeper fibres of the ulnar part afterwards contracting an insertion 
into the metacarpal bone. The pressure of the tendon and the 
ereater development of the unciform process then separates the 
origin? also, so that the ulnar border of the adductor minimi 
digiti becomes divorced from the rest, and we call it “ flexor 
brevis” and opponens minimi digiti in Human Anatomy. In 
Man and in Orang the typical adductor minimi vanishes, and the 
aberrant part remains as its only representative. 

In Man, Macacus, and Cynocephalus, we find an indubitable 
part of the adductor pollicis (aa) passing to be inserted into the 
radial sesamoid; in some human hands, on raising the long 
flexor tendon, the adductor fibres are seen to pass in an almost 
continuous sheet across the base of the phalanx from the ulnar 
to the radial sesamoid, some small twigs from the arteria princeps 
pollicis being the only separation. The pressure of the single 
flexor tendon of the thumb has been insufficient to separate the 
insertions in some cases, or in any case to separate the origins. 

The travelling in a radial direction of the adductor pollicis 
causes it to overlie, to functionally replace, and therefore to 


1 In the more primitive types the muscles in the hand are powerful, while in 
higher forms there is a tendency for the muscular structures to retreat to the 
fore-arm, the hand being occupied by tendons. 

2 In Cynocephalus, in which the flexor tendons are enormously strong, we find 
a wide separation between a® and a®op (fig 5) 


PER pe 


al eg ayy 


MORPHOLOGY OF THE INTRINSIC MUSCLES. 659 


dwarf, the true ulnar head of flexor brevis. As the valuable 
guide of nerve relation is absent on the radial side of the hand, 
it is. difficult to say in most cases whether the ulnar head 
becomes fused with adductor pollicis or aborts entirely. I 
believe that fusion rarely if ever occurs, as we find either a 
ligamentous rudiment of the true ulnar head of flexor brevis (as 
in Chimpanzee and Vulpine Phalanger), or a complete absence of 
any fibres arising in common with the radial head or from the 
base of the meteesarpal bone of thamb (as in Macacus). It is 
curious that the true inner head should be better represented in 
some of the higher primates (Man, Orang) than in many of the 
lower monkeys. 

Bischoff! describes “two heads of the flexor brevis pollicis, 
the inner somewhat weaker and deeper than the outer,” in Cyno- 
cephalus maimon, Cercopithecus sabdus, Macacus cynomolgus, 
Pithecia hirsuta, and Hapale penicillata. I could not find any 
true inner head in Macacus nemestrinus or in Hapale, and 
from a careful inspection of his figures of Cynocephalus maimon 
and comparison with Cynocephalus anubis, 1 believe that what 
he calls the inner head of flexor brevis is identical with (aa fig. 
5), being a part of the adductor inserted into the radial sesamoid. 

While the dissections described above have shown that the 
so-called “ flexor brevis” minimi digiti of the hand of Man is a 
different muscle to the true flexor brevis, there are two possible 
views of its morphology besides the one advocated above :-— 


(a) That it is a part of the abductor. 
(>) That it is the division of the flexor brevis digitorum 
manus to the little finger. 


(a) In favour of the first view, it may be urged that the origin 
of the corresponding muscle on the radial side (abductor pollicis) 
has a great tendency to travel ulnarwards along the annular 
ligament, even passing the middle line in some cases. On the 
other hand, the nerve relation is a strong argument against this 
view. In Cynocephalus, where a second slip of the abductor 
takes origin from the annular ligament (fig. 5), the normal nerve 
relation is still preserved. The absolute continuity of origin of 
the “flexor brevis” (aa) with the normal adductor, which has 


1 Op. cit., pp. 90, 91. 


660 MR H. ST JOHN BROOKS. 


been shown in Virginian Opossum, Vulpine Phalanger, and even 
in so high a form as the Chimpanzee, is another reason against 
regarding “flexor brevis” as part of the abductor. 

(bh) In favour of the second view, the fact may be noticed 
that in the foot the tendons of the short flexor are not always 
perforated. In the Chimpanzee I found the tendon to the little 
fincer inserted into the floor of the flexor sheath, on the fibular 
side of the base of the prowimal phalanx of the little toe, and I 
have seen the same condition in Man. . In the toes of Ornitho- 
rhynchus, Cunningham has shown that none of the tendons of the 
short flexor are perforated. A strong argument against this view, 
however, is the presence of a flexor brevis digitorum manus to 
the little finger in the Opossum (fbdm, fig 2), coexisting with 
aa. Inthe foot of the Opossum there is a perforated tendon 
for the fifth toe coexisting with the muscle (a%a, fig. 4), and it 
appears probable that tue latier is the homologue of the “ flexor 
brevis” minimi digiti of the hand. 


EXPLANATION OF PLATE XXI. 


Fig. 1. Manus of Cat. jf! to /*, flexores breves,—/f! has only the 
radial head ; 2, deep branch of ulnar nerve passing under pisi-uncinate 
ligament, and then under two of the adductores, a and a”; a!, addue- 
tor pollicis ; abd®, abductor minimi digiti. 

Fig. 2. Manus of Virginian Opossum. a}, a?, at, a°, adductores; a°a 
and aba’, parts of adductor minimi digiti whose insertion has wandered 
to ulnar side of fifth digit; «, ulnar nerve, its deep branch passing 
under the adductores ; fbdm, flexor brevis digitorum manus reflected ; 
f'r, radial head of flexor brevis pollicis ; abd!, abductor pollicis. 

Fig. 3. Deeper dissection of manus of Opossum. jf! to 7°, flexores 
breves. The deep branch of ulnar nerve is shown passing superficial 
to all of these. 

Fig. 4. Pes of Virginian Opossum. epn, deep branch of external 
plantar nerve; f'¢ and ff, tibial and fibular heads of flexor brevis 
pollicis. 

Fig. 5. Manus of Cynocephalus anubis. f°r, radial head of flexor 
brevis minimi (“third palmar interosseous”) ; f°op, opponens minimi 
derived from true flexor brevis ; a°op opponens from layer of adduc- 
tores; aa, “ flexor brevis” minimi digiti ; op!, opponens pollicis ; a'a, 
part of adductor pollicis inserted into radial sesamoid ; ¢, tendon of 
flexor longus pollicis, 


Fig. 6. Thumb of Orang. /'7 and fw, radial and ulnar heads of 


MORPHOLOGY OF THE INTRINSIC MUSCLES. 661 


flexor brevis pollicis, having a common origin, but separated at their 
insertion by ¢, the slender tendon of flexor longus pollicis. 

Fig. 7. Diagram of the adductores of the manus of a Chimpanzee. 
d*, fourth dorsal interosseous ; a!o), adductor obliquus; a'tr, adductor 
transversus ; other letters as before. Observe that the origins of a 
and a@op are absolutely continuous for a short distance, and that a°op 
and a°a are united by half their length. 

Fig. 8. Short muscles of the little finger in Man, showing an 
unusually large part of the opponens minimi digiti derived from the 
true flexor brevis ; f°, “‘third palmar interosseous.” The “flexor 
brevis” minimi digiti (of human anatomy) was absent in this case. 


bo 
rs 


VOL. XX. 


ON THE NATURE OF THE RELATIONSHIP OF UREA 
FORMATION TO BILE SECRETION. By D. NoéL- 
Patron, M.D., B.Se., F.R.S.E., Biological Fellow of the 
University of Edinburgh. 

(From the Physiological Laboratory of the University of Edinburgh. ) 
(Continued from p. 531.) 
In confirmation of the conclusion that urea formation and bile 
secretion are related to one another through their inter-depend- 
ence upon destruction of blood-corpuscles, I here give a further 
experiment, in which toluylendiamin was used as the hemolytic 
agent. This experiment is especially valuable, because the 
disturbing influence of increased production of heemorytes does 
not appear to have obscured the extent of destruction. It was 
only some days after the destruction of corpuscles had reached 
its greatest extent that increased production fairly established 
itself. This is probably to be accounted for by the fact that a 


dog considerably older than those used in my previous experi- 
ments was employed. 


Exp. V1. 


For this experiment an old black retriever bitch, weighing 15°876 
kilos., was used. The subjoined table and fig. 5 show the influence 
of toluylendiamin on the urea production and on the number of blood- 
corpuscles. On the sixth day of the experiment 0-4 grm. of the drug 
was administered, while 0°3 grm. was given on the following day. 
On the seventh day the dog was distinctly jaundiced, and the urine was 
of a dark port wine colour and contained bile acids and pigments. On 
the eighth day the jaundice was more marked and general, but by the 
ninth day it had somewhat diminished. Throughout the experiment 
the dog seemed perfectly well and took its food greedily. Calculating 
as in the former experiments we find that a dog of this weight has 1221 
erms. of blood and 168 grms. of hemoglobin. Between the fifth and 
the eleventh day the corpuscles fell from 8,390,000 to 5,060,000—a fall 
of 39°69 per cent. Thus throughout the body 66-7 grms. of haemoglobin 
must have been set free, and, supposing this to be entirely broken up 
into urea, bilirubin, &c., we should expect to find a formation of 20:2 
erms. of urea above the normal. 

3efore the administration of the drug 6645 grms. of urea were daily 
excreted, and during the six days following its administration the 
production of urea in excess of the normal was 18°588. On the seventh 
day 1:251 erm. of urea in excess of the normal were excreted, making 
in all 19°839—only 0°361 grm. less than the amount calculated from 
the disintegration of heemocy tes. 


© 


ie 2) 


~~ 


a 


UREA FORMATION 


|Dayof' Urine Sp. G 


Exp. | in ¢.cs. 


{ 525 1013 
{ 525 1013 
520 1013 


j 460 1014 
) 460 1014 
6 500 1015 
7 | $490 | 1018 
8 ( 490 1018 


9 400 1018 
Oe |) 580 1015 
11 620 1014 
12, 470 1016 

Sue 520 1013 
i: ae 480 1015 
16° | 580 1013 
17 400 1015 


18 § 500 1014 
19 {500 1014 


20 600 | 1012 


Urea in 


Grms. 


OMNTIH MH DN 


« 


Leo) 


— 
bt pt 


10 


ago™ 


DAO. Dd 
ns sroe : 


t 


SONNaY 
SADROH 


| Hemocytes | 


‘741 
“741 
“D54 
596 
*596 
‘O70 
"156 
"156 
280 
“600 
626 
7°896 
"635 
566 


AND BILE SECRETION. 663 


temarks. 
per ec.mm,. | 


| Weight on 5th=15°87 kilos. 


Xf | Diet—Oatmeal=170 grms. 
,500,000 | Milk, . 320c.c. 


320,000 | 0:4 grm. Toluylendiamin. 
85,000 | 0°3 grm. Toluylendiamin. 


On En D> OD NICO CO 


bo 
On 
as 
S 
oO 


5,060,000 
5,510,000 | 
5,410,000 
5, 280,000 
6,030,000 
6,525,000 
6,690,000 
7,170,000 


Hzemocytes 
per c.wim. 


9,000,000 


8,000,000 


7,000,000 


6, 000,000 


’ 


5,000,000 


Exp. VI. —Infiuence of tolmyglendiamin on urea production and on number of 


hemocytes per c.mm, 


0°4 grms. given at a, and 0°3 grms. at e. 


664 DR D. NOEL-PATON. 


Even if we refuse to accept the hypothetical decomposition of 
the hemoglobin molecule given above, the very close relation’ 
ship between the extent of the destruction of blood-corpuscles 
and the increase in the urea excretion shown by these experi- 
ments clearly indicates the existence of a definite connection 
between these two processes. 


Direct ACTION ON THE HA&MOCYTES OF DRUGS INCREASING 
UREA PRODUCTION AND BILE SECRETION. 


As before stated, Afanassiew has connected the cholagogue 
action of toluylendiamin with its direct destructive influence 
upon the blood-corpuscles, while Paschkis has demonstrated 
that these most powerful hemolytic agents—the salts of the bile 
acids—stimulate bile secretion in a very marked degree. 

By observing the action of salicylate of soda, of benzoate of 
soda, of colchicine, and of mercury on the hemocytes, I have 
endeavoured to ascertain whether or not their double action 
on bile secretion and urea production can to any extent be 
explained by their primary direct influence on the blood- 
corpuscles. 

For this purpose the action of these substances outside the body, 
and in some cases in the living animal, has been studied. | 

Observations outside the body of the action of these drugs 
are absolutely essential, since changes produced in the cor- 
puscles after their introduction into the living body are not 
necessarily due to the direct action of the agents. Specially 
important is the demonstration of these changes in such a 
research as the present, when from the increased secretion of bile 
induced the possible action of bile acids and their salts must 
be carefully excluded. My object has been merely to ascertain 
if the drugs above mentioned have any hemolytic action, and 
I have not considered it expedient to describe at length the 
various changes which occur in the course of the disintegration 
of hzemocytes—a line of histological research which would, I 
believe, throw some important light on the minute structure of 
the red blood-corpuscle. 

In such a research various precautions are necessary. In the 
first place, the solution employed must have a neutral reaction, 


o 


UREA FORMATION AND BILE SECRETION. 665 


since any degree of acidity or alkalinity tends to destroy the 
corpuscles. Secondly, it is necessary to employ solution of the 
same specific gravity as the blood-serum—1025—or to use such 
a solvent as a 0°75 per cent. salt solution in which corpuscles 
remain unchanged. This salt solution is especially useful where 
dilute solutions of the reagent have to be employed. 

The action of these substances has been studied not only on 
the mammalian, but also on the amphibian blood ; but since their 
action on the latter is precisely similar to their influence on the 
former, I have not deemed it necessary here to describe it in 
detail, and have confined myself entirely to the changes ob- 
served in the human blood. 

The mode of experiment was as follows :—A drop of the solu- 
tion was placed on the finger, which was pricked through this. 
The blood was mixed with the solution, and the mixture was 
transferred to a shallow cell, covered and at once examined, 
usually under a Zeiss F lens. 


SALICYLATE OF SODA. 


On the action of salicylic acid and salicylate of soda on the 
red blood-corpuscles outsicle the body several observations are 
already recorded. 


Chirone, in a paper entitled “Acido Salicilico e. Salicilate” (Z/ 
Movemento med. e. chir., 11-12 maggio, quoted in Jahresb. for 1878, p. 
408,) states that the toxic action of this drug depends upon its destruc- 
tive action upon the hemocytes. 

Cotton, “‘ Act. de l’acide salicylique sur le Sang et en particulier sur 
les globules rouges” (Lyon Méd., t. }. p. 557, quoted by Prudden) 
shows that with a 3 per cent. solution the red corpuscles become 
globular and that the hemoglobin is decomposed. 

Prideaux (Practitioner, Sept. 1878, p. 171) states that it has no 
influence upon the hemocytes, but that the movements of the leuco- 
cytes are diminished. 

Thiersch (Volkmann’s Klin. Vortrdge, Nos. 84 and 85, p. 657) finds 
that salicylic acid destroys the red blood-corpuscles. 

Prudden (American Medical Journal for 1882, p. 64) describes 
very carefully the influence of salicylic acid dissolved in a 0°5 per cent. 
salt solution upon the hemocytes. He finds that a solution contain- 
ing 1 part in 500 causes a rapid disintegration of the corpuscles, but 
that in more dilute sclutions it acts merely like other diluteacids. He 
also notices that it acts more rapidly upon the corpuscles of the frog 
than upon the mammalian corpuscle. 


666 DR D. NOEL-PATON. 


A. Outside the body. 


Solutions of different strengths were employed, but with the 


stronger solutions disintegration of the corpuscles occurred so 
rapidly that it was impossible to study the various changes. 
A 1 per cent, solution in 0°75 per cent. salt solution may be 
employed with advantage. 

On allowing a drop of this solution to mix with a drop 
of blood the following changes occur:— 


1. Large blunt crenations appear round the periphery of the 

disc. 

. These crenations become sharper and longer, and the 
corpuscles shrink in size and assume a spherical form. 
The spines next become more numerous, entirely covering 
the surface of the corpuscle, at the same time they 

become shorter. 

4. The spines finally disappear, leaving a small highly refract- 
ing deeply pigmented sphere. 

5. The heemocyte now slowly increases in size, becomes paler, 
and on close examination it is seen to be granular—the 
cranules appearing to be in an active state of motion 
within the heemocyte. 

6. The granules become more and more apparent, and grow 
larger and larger and seem to collect the pigment around 
them, so that we have a colourless cell with a thin layer 
of pigment under the membrane and masses of pigment 
throughout the cell. 

7. The pigment is gradually dissolved out—colouring the 
blood-serum, and leaves within the cell colourless masses 
of the intracellular stroma. 

8. These masses become less and less distinct, and finally an 
almost homogeneous, colourless, transparent, spherical 
shadow is left, which requires for its detection very 
careful focusing. 


bo 


os 


Different corpuscles are seen to have a very different resisting 
power in regard to the action of salicylate of soda. Some 
corpuscles break down at once, while others hold out for long 
after the majority have been reduced to shadows. 


B. In the living body. 


Deiat alt as fate mis. 


? Cpe AG ND parte me? s 


UREA FORMATION AND BILE SECRETION. 667 


Exp. I. 

A female, aged 15 years, and weighing 40°8 kilos, who 
was suffering from favus but who was otherwise healthy, was 
employed in this and in the next experiment. The diet was as 
nearly as possible uniform, and at 12.30 each day the number 
of hemocytes per c.mm. were estimated with Gower’s hemocytometer. 

Between 2 p.m. on April 7th and 10 p.m. on April 8th, 13 grms. 
of salicylate of soda were administered in 20-grain doses. On the 
morning of the 8th well-marked symptoms were induced. The 
accompanying table shows, that under the influence of salicyate of soda 
a fall of 300,000 per c.mm. occurred in the number of corpuscles :— 


| Date. He of Hiemocyts Remarks. 
| 1885. | 
April 5th 4,825,000 

» 6th 4,900,000 
He eee 13 grms. salicylate of soda. 
» _9th 4,570,000 
;, 10th 4,650,000 | 
oe a Lith 4,720,000 
jae a PAA 
»» 18th 4,740,000 

ES ee ee oc | 

Exp. II. 


The same individual was used as in Exp. I, and the experiment was 
in every way, with the exception of the doses given, exactly the same 
as the last. The administration of the drug was commenced at 12 
noon on the 28th. On the evening of the 29th well-marked symptoms 
were induced. 

The following results were obtained :— 


Date. No. Esc ne Remarks. 
April 27th 4,850,000 
oP gth,.t*7) 4,850,000 
eat 29th | 4,695,000 | 7:2 grms. of sod. sal. in 24 hrs. 
», 30th nee 7°2 grms. of sod. sal. in 24 hrs. 
May Ist | 4,625,000 
«Sed 4,060,000 
51) ord 4,720,000 | Faint traces of salic. in urine. 
» bth 4,750,000 
Exp. II. 


A setter bitch, weighing 11°80 kilos. and recovering from the 
anemia induced by the administration of pyrogallic acid, was used. 
The diet was fixed. 


668 DR D. NOEL-PATON. 


er No. of Heemocytes Pa: 
Date. per e.mm. Remarks. 
May 14th 5,400,000 
») Lbthi 5,820,000 6 grms. salicylate of soda. 
se lGthi 5,000,000 6 gris. salicylate of soda. 
 liabi 4,555,000 
5. 18th 5,105,000 


These three experiments, yielding results so entirely uniform, 
clearly show that in the mammalian body the administration 
of even comparatively moderate doses of salicylate of soda is 
followed by a considerable destruction of blood-corpuscles. 


BENZOATE OF SODA. 


The benzoates like the salicylates belong to the carbolic acid 
vroup of pharmacological substances, most of which, as indicated 
by Harnack (Arzneimittellehre, s. 288), exert a destructive 
influence on the blood-corpuscles. I have been unable, however, 
to find any experiments on the direct action of the benzoates 
in the blood. 

When a 5°5 per cent. solution of benzoate of soda (sp gr. 1025) 
is mixed with a drop of human blood, the following changes 
occur in the corpuscles:— 


1. Some assume a globular form, and become larger and 
paler. 

A great number are enlarged and paler with a faintish more 
or less crenated outline. Their form becomes irregular, 
varying from almost spherical to fusiform. 

The largest number become somewhat reduced in size, lose 
their biconcave form, appear bounded by a strong dark 
border, and are markedly crenated. When a number 
of these are together a curious oscillatory movement may 
be seen, closely resembling Brownian movement. 

If such a preparation be placed for some time in a warm 
moist chamber a large number of the corpuscles are 
converted into shadows, while all stages between such 
shadows and the small crenated corpuscles may be 
studied. Under the cover-glass these changes go on 
much more slowly than without it. In this respect 


bo 


ae) 


UREA FORMATION AND BILE SECRETION, 669 


benzoate of soda appears to resemble toluylendiamin, which 
according to Afanassiew (Joc. cit.) requires the free ingress 
of air to the preparation to enable it to exert its 
destructive influence upon the hemocytes. 


In all these observations check experiments, with blood mixed 
with 0°75 per cent. salt solution, showed no destruction of cor- 
puscles. 

Benzoate of soda is therefore undoubtedly a hemolytic agent, 
but not of such power as the salicylate. 


COLCHICINE. 


I can find no observations on the direct action of colchicine 
on the blood-corpuscles. Schroff (Lehrbuch der Pharmacologie, p. 
615) describes the condition of animals poisoned with colchicine 
as follows :-— 

“Am constanten fand sich Enteritis bisweilen Gastritis und immer 
ein dickes, peschwarzes, theerartiges, schmieriges Blut, das die 
Hohlen des linken Herzens und die obere und untere Hohlader bis 
in ihre Verzweigungen, die dem Hirn, der Leber und der Niere 
angehoren erfiillte.’ 

All subsequent writers have merely quoted this observation of 
Schroff. 

A. Outside the body. 

A 5 per cent. solution of perfectly neutral colchicine dissolved 
in 0°75 per cent. salt solution was used. 

The following changes were observed :— 


1. The corpuscles become cupped, a convexity taking the 

place of the concavity on one side. 

The lips of the cup approach one another and become 

irregular. 

The corpuscles become spherical, some small and dark, 

others larger and paler. 

4, The large pale hemocytes become shadows simply by losing 
their colouring matter. 

Some of the smaller ones enlarge and throw out processes: 
these processes are either (1) long transparent filiform 
vibrating tails, sometimes markedly articulated, almost 
moniliform at other times with no apparent joints, but 
always with a somewhat club-shaped head ; or (2) round, 


bo 


Os 


670 DR D. NOEL-PATON. 


clear, and spherical. Both forms are to be seen in the 
one corpuscle. These processes often break off and cause. 
a granular appearance of the serum in which the cor- 
puscles float, at other times they remain attached. 
5. The pigment slowly dissolves out till only a layer inside 
the cell membrane is left. 
. This in turn, too, disappears, and nothing is left but a 
shadow, often with processes attached. 


B. Within the body. 

The method of enumerating the corpuscles per c.mm., em- 
ployed in my experiments on the action of salicylate of 
soda upon the blood in the living mammal, was unsuited for the 
investigation of the blood-changes induced by such a drug 
as colchicine, which by its well-known cathartic action causes 
a concentration of the blood as shown by Malassez. 

Another method of experiment had to be adopted :— 


Exp. I. 

A large male cat received no food for twenty-four hours. At 11 
o'clock on May 5th 01 grm. of colchicine, procured from Messrs 
Hopkin & Williams, London, dissolved in about 20 c.cs. of water was 
administered. 

During the afternoon the cat had several mucous evacuations of the 
bowels, and vomitted some glairy matter. In the evening it was dull ; 
it died on the morning of the 6th. 

The thorax was opened, and some blood taken from the right 
auricle was at once examined ; this contained a very large number of 
shadow corpuscles arranged in groups. Blood taken from the femoral 
vein, and from the lower part of the vena cava, also showed the 
presence of these shadows, but not in such large numbers as in the 
heart. 

The right side of the heart was full of fluid blood of a very dark 
colour, which coagulated on being placed in a glass vessel. 

This experiment clearly shows that colchicine within the animal 
body destroys the red blood-corpuscles. 


for) 


PERCHLORIDE OF MERCURY, 


On the action of the salts of mercury on the hemocytes 
outside the body, I can find only one investigation. This is by 
Polotebnow (Virchow’s <Avch., Bd. xxxi. s. 35), who employed a 
solution of albuminate of mercury, 1 c.mm. of which contained 
from 0-014 to 0015 orm. of perchloride of mereury, As a 
result of his investigations he concludes that :— 


UREA FORMATION AND BILE SECRETION. 671 


1. Under the influence of concentrated solution of albuminate 
of mercury, the blood-corpuscles after their round 
form is once altered have lost the capacity to resume 
their original shape, either through a loss of the elas- 
ticity of their membrane or in consequence of some 
other cause. 

2. In a solution of albuminate of mercury the corpuscles are 
very quickly destroyed, the rapidity being in proportion 
to the amount of mercury present, and being also more 
rapid when the blood is shaken up with air, or when the 
temperature is raised for thirty or forty minutes to 37° or 
38° C. 

3. Under the influence of mercury the corpuscles quickly 
lose their pigment, and the more concentrated the 
solution the more rapidly does this take place. 

4, The absorbent power of the blood for oxygen is also 
diminished. 

Of the many investigations on the action of mercury on the 

blood-corpuscles in the living body, I can find only one of any 
scientific value. The majority have been made on syphilitic 


Exp. IV. 


Date. | Hemocytes per c.mm. Weight of Rabbit. 
<< 
May 5th | 4,480,000 2°120 kilos. 


0°001 grm. of perchloride of mercury given daily hypodermically 
from 5th onwards. 


} 
| 
» 9th 


May 8th 4,200,000 | 2°010 kilos. 

4,000,000 1:900 ,, 

,, llth 4,000,000 | 1800 ,, 

Diarrhcea occurred on the 12th, and the corpuscles rose to— 
May 12th | 5,202,400 | 1°870 kilo. 

_ The treatment was stopped till the diarrhea ceased, and was then continued. 

May 15th | 3, 363, 800 1°736 kilo. 

»» 16th 3,200,000 1676 ,, 

The dose was progressively increased. 

,» 17th 3,140,000 1°660_,, 

,, 18th 3,000,000 1:650 ,, 

,, 19th 2,900,000 1500 ,, 

¥» 20th 2,800,000 | 1:460 ,, 

»» 2ist 2,700,000 1-444 ,, 

yy 22ne Animal died. 


672 DR D. NOEL-PATON. 


patients, with small doses of the drug extending over a 
lengthened period, so that as direct evidence of the action of 
the drug on the corpuscles they are of no value. 
Wilbowchewitch (Arch. de Physiologie, t. i. p. 530) records 
four experiments made upon rabbits, which clearly show that 
uuder large doses of perchloride of mercury a very marked 
diminution in the number of blood-corpuscles per ¢.mm., occurs. 
The preceding table gives the results of his fourth experiment. 


Such experiments are only of value when taken along with observa- 
tions in the hemolytic action of mercury outside the body, since 
without such observations it would not be fair to conclude that any 
increased destruction, and not merely a diminished production, had 
occurred, 

The chief difficulty in the way of studying the action of 
mercury upon the blood-corpuscles outside the body, is that 
nearly all the salts of this metal cause coagulation of albumin. 
The mercuric potassic iodide in dilute solution has not this 
action, and I have therefore employed it in the following obser- 
vations, 

I had previously ascertained from a number of experiments 
that the iodide of potassium has absolutely no action on the red 
corpuscles, and that the chloride of potassium is also tnert. 

I therefore consider it a fair deduction, that any action which 
the mercuric potassic iodide may exert is due to the mercury it 
contains. 

A solution composed of— 

1 germ. of mercuric chloride, 

3 grms. of potassic iodide, and 

60 c.c. of water, 
was prepared, and was diluted with 60 c.c. of a 0°75 per cent. salt 
solution, as its action without this dilution was so rapid as to 
prevent any observations on the nature of the changes induced. 

When this solution is added to a drop of blood, the heemocytes 
are seen to swell up and to lose their biconcave form, while the 
pigment disappears leaving the cell membrane enclosing the 
intranuclear network. Very frequently the membrane col- 
lapses and the corpuscle is represented by a shrivelled shadow. 

I have not thought it necessary to repeat Wilbowchewitch’s 
observations. In fact, the extremely poisonous nature of the 
salts of mercury render such observations difficult. 


UREA FORMATION AND BILE SECRETION. 673 


It would thus seem that the influence of these drugs on bile 
secretion and urea formation may be explained by their destruc- 
tive influence on red blood-corpuscles, hemoglobin being set free 
and excreted by the liver as already described. Such a direct 
influence on the blood-corpuscles is rendered all the more 
probable by their absorption into the very slow hepatic blood- 
stream, and also through their probable re-entrance into the 
liver, after excretion in the bile, through the entero-hepatic 
circulation. 


From these observations I would conclude :-—. 

1. That the destruction of blood-corpuscles must be considered 
as a powerful stimulant to the secretion of bile, the liver 
having as one of its functions the elimination from the 
system of effete haemoglobin. 

2. That urea production also is increased by the destruction 
of blood-corpuscles. 

3. That bile secretion and urea formation bear a direct 
relationship to one another, and that this relationship is 
due to the dependence of both upon destruction of 
heemocytes. 

4. That the hemolytic action of salicylate of soda, of 
benzoate of soda, of colchicine, and of perchloride of 
mercury, as well as their influence in increasing the pro- 
duction of urea, is due in great measure at least to their 
direct hzemolytic action. 

The connection of urea production with the destruction of 
hzemocytes naturally suggests the question--How much of the 
total urea produced in the animal body in its normal condition 
is derived from effete hemoglobin? This question I hope to 
discuss in a future paper. 


THE BLOOD-FORMING ORGANS AND _ BLOOD- 
FORMATION: AN EXPERIMENTAL RESEARCH. 
By Jown Locxnarrt Gipson, M.D., Formerly Senior 
Demonstrator of Physiology, University of Edinburgh. 


(Continued from p. 474, vol. xx.) 
Part III.—Own THE THYROID GLAND. 


We have now to consider that other organ which has occa- 
sionally been called a blood-forming one, namely, the thyroid 
gland: an organ of late brought specially into notice, on the one 
hand by the writings of Zesas and Credé as to its alleged blood- 
forming function, and on the other hand by the writings of 
Kocher and Reverdin as to its other functions, and as to the 
danger attending its total removal. Here, again, I have by ex- 
periment searched for evidence of a blood-forming function ; 
but this time the results have, in contrast with the positive ones 
got in the case of the bone-marrow, spleen, and lymphatic 
glands, been wholly negative, that is, do not show the existence 
of any such function. 

To the idea of a possible important blood-forming action of 
the thyroid, my attention was first directed by the two papers of 
Zesas already mentioned under extirpation of the spleen. In 
the last of these papers, Zesas, founding chiefly on an experiment 
on one dog, though further drawing support from the observa- 
tions of Credé, concludes that the thyroid is the principal organ 
which takes the place of.the spleen when the spleen is removed, 
and further says that when the thyroid too is removed then the 
other blood-forming organs—“lymph-glands and liver (?) ”-—are 
unable to make up for the loss of the spleen. (In this paper the 
researches of Neumann and Bizzozero on the bone-marrow are 
ignored.) 

On reading the above conclusion, I at once saw how insuffi- 
ciently it was based. For Zesas, in his single experiment, had 
removed both spleen and thyroid without first having ascertained 
the effect of removing the thyroid alone. 


BLOOD-FORMING ORGANS AND BLOOD-FORMATION, 675 


Accordingly I determined to make experiments as to a blood- 
forming action, and take care that operation was properly com- 
bined with blood-observation. Circumstances, however, for 
some time prevented this; and when at last I was able to begin 
I found that other observers were engaged on the subject. 
Nevertheless, none of their papers came to my notice till I had 
already excised the thyroid and obtained decided results. 

All observers are agreed as to the insufficiency of the grounds 
on which Zesas came to his conelusion, and all but one are 
agreed that the reason why the conclusion was so inaccurate 
was the fact that the removal of the thyroid is in the dog of itself 
always fatal. 


Before Credé and Zesas, the only writer who seems to have made a 
distinct positive statement as to a functional connection between the 
thyroid and the spleen is Tiedemann,! who said that the spleen is a 
blood-forming organ, and that after its removal its function is taken 
over by the thyroid. And with this statement may be contrasted 
Briicke’s assertion, in his “ Vorlesungen iiber Physiologie,”? that 
while we can refer the spleen to the lymphatic system we know of no 
connection between the thyroid and that system. 

Schiff, who seems to have been the first to pay particular attention 
to the subject of excision of the thyroid, and who began his experi- 
ments in 1856, has lately made further ones,* which are interesting 
and in many respects thorough, though they do not seem to have 
included observation of the blood, apparently because he assumed that 
the thyroid has nothing to do with blood-formation. 

He excised the gland from a number of dogs, and found that when 
both lobes (or rather, as there is in dogs no isthmus, both glands) were 
removed at one operation the animals, with but one exception, died 
between the third and the twenty-seventh day after. The animals 
became indifferent and melancholic; there was great itching of the 
skin; at intervals there were fibrillar muscular contractions, followed 
by clonic and tonic spasms ; many had actual tetanus-like attacks ; the 
“excitable zone” of the brain ceased to react to electrical stimulation ; 
and the blood-pressure fell, as a result of vaso-motor paralysis. In the 
exceptional case the animal had, at the time Schiff wrote, already 
survived nearly fifty days, and seemed likely to make a permanent 
recovery. 

Afterwards he found that if the lobes, instead of being both removed 
at one operation, were removed at two operations, separated by a con- 


1 Tiedemann, Zeitschr. f. Physiologie, Bd. v. Heft. 1 (1833).—Quoted by 
Tauber, in Virchow’s Archiv, Bd. xevi. p. 30. ; 

21. Aufl. (1874), Bd. i, p. 209; and repeated in 3. Aufl. (1881), Bd. i. p. 215, 

3 Schiff, ‘‘ Résumé d’une série d’expériences sur les effets de l’ablation des corps 
thyroides,” Revue médicale de la Suisse romande, 15. Février 1884; ‘Résumé 
d@ une nouvelle série.” Rev. Méd. de la Suisse rom., 15. Aout 1884. 


676 DR JOHN LOCKHART GIBSON. 


siderable interval (for adult dogs he considers twenty days quite 
enough), it was then possible to keep the animal alive ; and he explains. 
this by supposing that the function of the gland passes over to some 
other organ or organs, but does so only very slowly. Experimenting 
as to whether it is to the supra-renal capsules that the function passes, 
he finds that it is not. 

Further, having in a series of experiments placed the freshly-excised 
thyroid of a dog in the abdomen of another dog, waited from two to 
five weeks to let it become engrafted, and then removed both lobes of 
this dog’s thyroid at one operation, he found that the dog might, 
instead of dying like the unprepared dogs from which the whole gland 
was removed at one operation, survive like the dogs from which the 
lobes were removed at separate operations. 

From the facts obtained, he supposes that the gland may have the 
function of producing some substance which passes from it into the 
blood, and there serves as a necessary element in the nutrition of the 
nervous centres. 

Tauber,! in a recent paper, holds that Zesas is wrong in saying 
there is a relation between the spleen and the thyroid. But Tauber’s 
argument proceeds, at least in part, on an erroneous assumption made 
by him that the thyroid is in domestic animals (cats and dogs?) often 
entirely wanting. 

Another paper on excision of the thyroid is one by two Italian 
observers, Sanquirico and Canalis,? who likewise disagree with Zesas. 
They experimented on thirteen dogs. From seven both lobes were 
removed at the same time, from two both in the course of from six to 
twenty days, and in four a small portion of the gland was left unex- 
cised. 

Their results were—(1) In spite of primary healing, the animals 
from which both lobes were removed at the same time died in from 
six to twenty days. (2) The symptoms usually began after from 
twenty-four hours to three days (in one case only after thirteen days). 
The animals began to refuse food, swallowed badly, became tired and 
dyspnoeic, suffered from muscular trembling and from general or partial 
clonic cramps, groaned, and were generally restless. Walking and 
standing became more difficult, the animals often ran in circles, and 
they were very apt to fall. In running about they seemed to have a 
hindrance, as if they had lost control over their movements. On in- 
crease of these symptoms followed death. (3) In three cases death 
was preceded by a fall of temperature from 39° to 35°°5 C. (4) Arti- 
ficial nourishment failed to prevent death. (5) In some cases purulent 
conjunctivitis was developed, once with perforating corneal ulcer. 
(6) Examination of the blood revealed slight changes, which did not, 
however, overstep physiological limits. (7) Post mortem examination 


1 Tauber, ‘‘Zur Frage nach der physiologischen Beziehung der Schilddriise zur 
Milz,” Virchow’s Archiv, Bd, xevi. Heft 1 (Apr. 1884). 

2 Sanquirico and Canalis, ‘‘ Sulla estirpazione del cerpo tiroide,” Archivio per 
le scienze mediche, vol. viii. No. 10 (1884). Extracted in Centralbl. f. klin. Med., 
18 Oct. 1884. 


. 


BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 677 


gave nothing beyond distinct anemia of both brain substances, often 
hyperemia of the liver, and injection of the mesenteric vessels, once 
with extravasation into the intestinal mucous membrane. (8) Examin- 
ation of the urine yielded negative results. (9) Both absorption of 
septic matter and alteration of the vagi or of the recurrent laryngeal 
nerves could be excluded from consideration as possible causes of 
death. (10) The animals from which at first only one lobe was re- 
moved remained healthy till the other was removed, when the same 
symptoms appeared as where both were removed at the same time. 
(11) Two adult animals, from each of which one entire lobe and the 
upper two-thirds of the other were removed, died, the one after six, 
and the other after three days. (12) Two from which the spleen had 
first heen removed and one lobe and the lower two-thirds of the other 
lobe of the thyroid were afterwards removed remained perfectly 
healthy. (13) One of these two animals having after the extirpation 
of the spleen been artificially rendered anemic, an examination of its 
thyroid, extirpated during full reproduction of the blood, showed no 
sign of hematopoiesis. 

And from these facts they conclude—(1) The total extirpation of 
the thyroid is in dogs absolutely lethal, whether the spleen be removed 
or not. (2) Spleenless dogs bear partial removal of the thyroid with- 
out noteworthy effect. (3) There is no relationship between the spleen 
and the thyroid. (4) If a small part of the thyroid be left, this is 
enough to carry on the function of the gland. They cannotas yet say 
whether the position of this remaining part is of importance. (5) The 
lethal result cannot be ascribed to changes in the blood. (6) The 
thyroid has nothing to do with blood-formation. (7) It plays an 
important 7é/e in the animal economy, standing probably in relation to 
the nervous system. 

Two papers on excision of the thyroid by Dr Julius Wagner,! of 
Vienna, were published in June and July 1884, but came under my 
notice only after my own experiments were made. He obtained 
practically the same results as Schiff. His researches were specially 
directed to the relation of the thyroid to the nervous system, and he 
does not appear to have paid any attention to the supposition of a 
blood-forming action. He chose mostly very young animals, expecting 
that nervous symptoms would be more marked in them. 

Wagener draws attention to the fact that dogs are not perfectly suit- 
able for such experiments, inasmuch as they have, in the fat round the 
arch of the aorta, two small accessory thyroids, called by Wolfler,? who 
has described them, “aortic glands.” And it seems to me quite pos- 
sible that these glands may occasionally be very well developed, and 
may account for the case recorded by Bardeleben, where a dog survived 
after simultaneous removal of both lobes of the thyroid ; as well as for 
one of Zesas’ cases (to be afterwards mentioned), where, likewise, the 
animal seems to have survived, although much affected by the opera- 
tion. And they may also account for Schiff’s success in keeping dogs 


1 Wagner, Wiener med. Blatter, 1884, Nos. 25 and 30. 
2 Wolfler, Wiener med. Wochenschr., 1879, No. 8. 
VOL. XX. 2Y 


678 DR JOHN LOCKHART GIBSON, 


alive when he removed the second lobe at a considerable interval of 
time after the removal of the first, and thus gave the aortic glands 
time to hypertrophy. 

And, further, there has been another paper by Zesas,! in which 
he has somewhat modified his former conclusions. He experi- 
mented on ten dogs and three cats. In most of them the spleen was 
removed first, and the thyroid subsequently. After the second opera- 
tion, the animal invariably died. In two cases, however, it lived two 
months, and thus longer than any of the animals from which other 
observers had removed the thyroid alone—a fact certainly not in 
favour of the two organs having a compensatory function with refer- 
ence to each other. After removal of the thyroid alone, one cat and 
one dog lived four months, and one dog seems not to have died at all. 
As to the latter, I have already suggested that it may have been saved 
by the accessory thyroids (“‘aortic glands”); and in accordance with 
this suggestion is the fact, stated by Zesas, that it became very much 
lowered in condition and very miserable. 

In this paper, Zesas, though indeed speaking, on the one hand, of 
the very distinct symptoms which appear after removal of the thyroid, 
and, on the other, of the absence of symptoms after removal of the 
spleen, still holds that the thyroid has an important action in taking 
the place of the spleen when that organ has been removed. This 
opinion, however, seems entirely unsupported by his experiments. 
Certainly the fact that all the animals from which both thyroid and 
spleen were excised died is no proof of a compensatory function ; for 
it has been clearly shown, even by Zesas’ own experiments, that mere 
removal of the thyroid alone is quite sufficient to cause death, 

Zesas says that he found the same changes in the blood after ex- 
cision of the thyroid as after excision of the spleen, namely, an 
increase in the number of white corpuscles and a decrease in the num- 
ber of red; only that these changes appeared later and were less 
marked. And he further says that if both organs be removed the 
increase in the number of white corpuscles is enormous. What he 
means by “ enormous” is not stated. Certainly, as will further on be 
seen, there was no enormous increase of white corpuscles in the blood 
of a dog from which both organs were removed by myself. 

As to the general bearing of my own experiments on the foregoing, 
I have to say that while these experiments give, so far as I know, the 
only accurately recorded observations on the blood after excision of 
the thyroid, they seem to me to show that the removal of the gland 
produces no direct change in the number of corpuscles. In one case, 
indeed, there was, shortly before the death of the animal, a decided 
increase in the number of white corpuscles; but this can, I think, 
easily be explained in a way quite different from that given by Zesas. 
For, firstly, Zesas himself mentions that the changes in the blood ap- 
peared late; and, secondly, both his observations and -those of others 
show that shortly after the removal of the thyroid the animal refuses 


! Zesas, ‘Ist die Entfernung der Schilddriise ein physiologisch erlaubter Akt!” 
Arch. f. klin. Chir., Bd. xxx. (1884). 


- 


BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 679 


all nourishment; while, thirdly, it has been recorded by Bizzozero and 
various other observers that when an animal is starved the function 
of the blood-forming organs comes to a stand-still. So that the very 
natural explanation of the appearance of an increased number of white 
corpuscles and a diminished number of red after removal of the thyroid 
is simply that starvation has occurred, and has caused an inactivity of 
all the blood-forming organs as regards conversion of white corpuscles 
into red. To this explanation two of my cases, which were all in 
dogs, give an interesting support: one (Experiment IX.) showing a 
still important amount of food and practically no increase of white 
corpuscles, the other (X.) hardly any food and a decided increase. 

Where both spleen and thyroid have been excised, it would of 
course be quite natural, and in accordance with my explanation, that 
there should be a still greater increase in the number of white cor- 
puscles and decrease in the number of red than where only spleen or 
only thyroid has been excised. But, besides being able to say this, I 
can here, too, bring forward a case which does not even at first sight 
allow of Zesas’ view being taken, namely, one (XI.) where I excised 
the thyroid more than two months after the spleen had already been 
excised and yet found in the blood asa result of the excision of the 
thyroid no alteration whatever. In this case the animal took its food 
well until the last three or four days of its life. 

In this last paper, however, Zesas, while still supporting a blood- 
forming function of the thyroid, further says that the gland must 
have some other important function ; basing this conclusion on obser- 
vation of symptoms like those recorded by other writers. 


I now pass to a more particular description of my own experi- 
ments on excision of the thyroid, which were all, as I have said, 
on dogs. 


Antiseptic precautions were, as usual, observed during the opera- 
tion ; but the wound was not dressed. The wound was sewn up com- 
_ pletely, either with chromic-acid catgut or with horse-hair. In all, 
four operations (on three animals) were performed ; if one case be 
omitted where the animal died of hemorrhage a few hours after the 
operation.! In three out of the four there was healing by first inten- 
tion. 

The incision was made in the middle line, which enabled one easily 
to get down between the muscles ; and one then found the two lobes, 
or, rather, separate glands, lying, one on each side, under the sterno- 
thyroid muscles. Their mobility and slipperiness makes their removal 
somewhat difficult. As they are separate, the operation is a double 
one, by means of a single incision. 

In the removal of each lobe, first it was drawn out of the wound ; 
then the vessels passing to it were secured with three chromic-acid 
catgut ligatures, placed, one round those passing to its lower angle 
(with the tissues surrounding them), another round those passing to 


1 Here is also omitted the case of the animal of Experiment V. In this case, 
however, only one lobe (or gland) was ever removed. 


680 DR JOHN LOCKHART GIBSON. 


its upper angle, and the third round the remaining attachments ; and 
lastly all the attachments were divided on the distal side of the 
ligatures. 

From two dogs, A and B, both lobes were removed at the 
same time; and from the third dog, C, first one lobe, and then, 
after a month’s interval, the other lobe, was removed. 

I shall consider first the symptoms common to the two animals, 
both females, from which both lobes were removed at the 
same time. 

The first animal, A, or that of Experiment [X., was of the English- 
terrier type and about four months old, and weighed before the opera- 
tion 2 kilogrammes 80 decagrammes. The second, B, or that of 
Experiment X., was about eight months old, full-grown, and very 
strong and well-nourished, and weighed 5 kilogrammes 80 decagrammes. 

The first symptoms appeared in from twenty-four to forty- 
eight hours. The animals became distinctly apathetic and 
melancholic, showed disinclination to be disturbed, and seemed 
to have lost the desire for food, being only after considerable 
coaxing prevailed on to take even milk. Moreover, localised 
muscular spasms set in, which in A probably appeared first in 
the cesophagus, when, on the second day after the operation, 
it cried twice or thrice as if in sharp pain, especially after trying 
to swallow, and then put its paw into its mouth, as if to remove 
some offending substance. In both animals an apparent diffi- 
culty in swallowing was noticed. General trembling of the 
body set in very early, and was especially exaggerated when they 
were disturbed. Frequent fibrillar muscular twitches and a 
peculiar stiffness of the legs gave the uncertain gait spoken of 
by Sanquirico and Canalis. There seemed to be itching of 
the skin, more especially in the face; and pain, as evidenced 
sometimes by constant, and other times by intermittent 
groans, presumably caused by the localised cramp-like contrac- 
tions of the muscles. There seemed to be no pain in the 
wound; in fact, in A it healed by first intention. A, unlike B, 
would, after the appearance of symptoms, one day seem quite 
recovered, and take its food well, and then, next day, again show 
the symptoms, and appear uneasy and practically refuse all 
food. B, though much stronger to begin with, showed the 
symptoms earlier, and without any intermission. In A no general 
convulsions were observed, but in B they occurred pretty often. 


BLOOD-FORMING ORGANS AND BLOOD-FORMATION, 681 


Localised tonic and clonic spasms were common in both. Both 
seemed giddy when on their legs, and both showed the tendency 
to walk backwards observed by Wagner.! Both lost flesh, more 
especially B, which in a fortnight was reduced almost to a skele- 
ton. Both died, A thirteen, B seventeen days after the extir- 
pation. The weight of A had been reduced from 2 kilogrammes 
80 decagrammes to little more than 2 kilogrammes, and that of 
B from 5 kilogrammes 80 decagrammes to 3 kilogrammes 45 
decagrammes. During the last day or two both were in an 
almost comatose condition, from which, however, they could 
easily be roused. Both had inflammation of the conjunctiva ; 
and in one the inflammation was purulent, and before death 
implicated the cornea. It is interesting to note that Sanquirico 
and Canalis observed a similar affection of the conjunctiva. 

I now proceed to give the enumerations of the blood which 
were made in these two experiments. 


Experiment IX. (A). 


Proportion of 
Hzmocytes. Leucocytes. Leucocytes to 
| | Hzmocytes. 


Before Operation, . ‘ a 6,470,000 | 14,000 Tt 46223 


Operation on 24 September 1884, 
26 Sept., 2nd day after, . ‘ 6,080,000 19,000 1: 320 
. 2 ae . . | 6,980,000 15,000 1: 465°3 
2: cth  ,; . «| 6,880,000 38,000 1: 181 
i Gh |, . . | 6,950,000 12,000 1 :579 
Oct, 7th m : - 7,240,000 19,000 1:381 
Poe. oth, > “ex -¥ydgo;000 19,000 1: 393°6 


During the last three days of life no enumeration was made, 
‘as the animal was taking no food and was very miserable. Its 
condition, too, was such that any changes that might then have 
been found in the number of corpuscles could not have been 
ascribed to cessation of a blood-forming action of the thyroid. 

The results of the enumerations in this. case give no real 
support to the theory of Zesas; for the red corpuscles, after a 
decrease in number immediately following the operation, again 
increased, and, with slight irregularities, remained, until the last 
estimation, at any rate at their former number. The further 


IY Op: Cit. 


682 DR JOHN LOCKHART GIBSON. 


increase towards the end, I can ascribe to nothing unless to a 
diminution of the fluid contents of the vessels; but such a 
diminution may really have occurred, as the animal not only 
refused food, but also for the last few days would take neither — 
water nor any other liquid. The number of the white corpuscles 
cannot be said to have been influenced ; for, after the initial rise 
always observable after an operation, they resumed practically 
their former number and numerical relation to the red corpuscles. 
Of the exceptional number found one day, namely, 38,000 per 
e.m., I can give no explanation. 

On the whole case, then, it must be allowed that there was 
no change in the number of corpuscles sufficient to account for 
any of the symptoms, and that death must have been due to 
causes quite apart from the changes found in the blood. 

The changes in the blood of B seem to point to the same 
conclusion. 


Experiment X. (B). 


Proportion of 


Hemocytes. Leucocytes. Leucocytes to 
Hzemocytes. 
Before operation, ‘ 5 F 8,750,000 19,000 1: 460°5 
Operation on 18 October 1884. 
19 Oct., lst day after, . : 7,720,000 28,000 132757 
Ol eee ard = ; : 7,240,000 10,000 1: 724 
PR Sony Soetidn i : : 7,510,000 14,000 1:586°4 
2B hy Bhd re : 3 7,390,000 14,000 125248 
26 eee oul oe , ; 7,310,000 33,000 1 S225 
280 ees) el OGhines, E : 7,820,000 32,000 1: 22872 
SOs 12thee se, E 3 6,800,000 29,000 1:234°4 


In this experiment the usual fall in the number of corpuscles 
after the operation was never recovered from. But this cannot, 
I think, be ascribed to the removal of a blood-forming organ. It 
seems really to have been due to starvation, with consequent 
depression of the blood-forming functions. For the symptoms 
I have described set in very early, and continued without inter- 
mission ; and from the day after the operation the animal took 
hardly any food. And the same explanation will serve for the 
increase in the number of white corpuscles, and the further de- 
crease in the number of red corpuscles, towards the end of life. 
In other words, any changes observed in the blood, either in this 


BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 683 


experiment or in Experiment IX., could with perfect justice be 
ascribed to the effect of the removal of the thyroid on the 
general system; and there is nothing in them to support the 
idea of a direct blood-forming function of the organ. 

The case of the third animal, C, gives, likewise, no evidence in 
favour of a blood-forming function, and also otherwise gives 
very strong evidence against the supposition that the spleen 
and thyroid are related in function. The animal was the 
second one from which the spleen was excised (see Experi- 
ment II.). As may be remembered, it, after recovering from a 
slight ordinary initial fall in the number of red corpuscles, failed 
to show any other decrease in it until two months after the 
operation, when there was a distinct and pretty sudden one. It 
would have been interesting to watch the animal further without 
submitting it to another operation; but I wished to remove the 
thyroid from a spleenless animal, in order to watch the effect on 
the blood, and thought it an unusually good subject for such an 
experiment, as it had a very well developed thyroid. 

I decided to remove first one lobe, and then, after a month’s interval, 
the other. I should have liked to make the interval still longer, in 
hopes of keeping the animal alive after removal of both spleen and 


thyroid, as Schiff kept animals after removal of the thyroid alone ; 
but the time at my disposal did not allow me to do so. 


Experiment XT. (C). 


Proportion of 


Hemocytes. | Leucocytes. Leucocytes to 
| Hemocytes. 
Before operation, ‘ : : 6,590,000 17,000 1 :387°6 


One lobe excised on 6 January 1885. 


8 Jan., 2nd day after, . . | 5,590,000 19,000 1 : 294°5 
eee ath el, |.*. 6.210000 17,000 1 : 365-2 
1 ee 6th Ae : . | 6,210,000 16,000 1: 388-1 
Pie Oth -|,, Sebel We) 6;,190;060 22,000 1: 280-4 
eee 18th. \\,; . . | 7,230,000 13,000 1 :556°1 
oe) 20th sy, 7,330,000 13,000 1: 563°8 
eet 4th!) 5, 6,840,000 12,000 1 : 570 

6 Feb., 31st _,, 7,490,000 10,000 1:749 

Second lobe excised on 7 February. 

ee Dss? ¢ : : - : 6,910,000 | 15,000 1: 460°6 
ers 5 : ; - : 3 6,930,000 17,000 Ee 407-6 
Ge S : ; : : - 7,240,000 7,000 1': 1034°2 


ne) re a, 6,300,000 12,000 1: 525 


ts 


684 DR JOHN LOCKHART GIBSON. 


The results of the blood-enumerations will be found in the 
preceding table. 

The removal of the first lobe was followed by no symptom, 
and it will be noticed that the blood did not suffer. The irregu- 
larity in the number of red corpuscles, always, on the average, 
in the direction of increase, is such as I had already found in 
my animals after excision of the spleen; and must have been 
due to irregularity in the blood-forming activity of the bone- 
marrow and lymph-glands, in their attempts to make up for the 
loss of the spleen. Andit may here further be noticed, as against 
the supposition of a blood-forming action of the thyroid, that 
the loss of one lobe did not prevent the blood from being brought 
up to about the condition in which it was previous to the exci- 
sion of the spleen (7,520,000 red corpuscles per c.m.) three 
months before. 


Exactly a month after the removal of the first lobe the remaining 
one was removed, and it was found apparently not increased in size, 
its weight being the same as that of the first. Both after this opera- 
tion and after the first one the wound healed by first intention. 

For the first few days I hoped the animal would survive. It 
was quite lively, took its food well, and ran about as usual. Two 
days after, indeed, there seemed to be a little stiffness in putting 
down the feet; but one could not be sure, and next day none 
could be seen. Decided symptoms appeared only on the 
fourth day after the removal, viz, on the 11th of February. 
When trying to rise from its bed it at first failed, the hind 
legs, which had been under it, appearing to be quite stiff; 
and after it got up it fell once or twice in running across the 
room. It seemed, however, to recover, and then ran about as 
usual. But when taken up it trembled a good deal, and fibrillar 
muscular contractions were likewise evident. On running down 
stairs it had a slight convulsive attack, like a short epileptic fit. 
Krom this day the nervous symptoms gradually developed, 
although it took its food well until three or four days before its 
death, and consequently did not lose very much in weight. 

The nervous symptoms were very pronounced, and consisted, 
as in A and B, of almost constant trembling and fibrillar mus-- 
cular contractions and clonic and tonic convulsions. I saw it in at 
least two typical epileptic fits, each of which lasted some minutes, 


BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 685 


It had also attacks like those of tetanus, with well-marked opis- 
thotonus. The itching of the skin was not very marked. 
During the last few days of life the animal could hardly be 
got to come out of its bed, and slept most of the day. When 
it was disturbed, almost all the muscles became _ tonically 
contracted, the head being fixed and the eyes fixed and 
staring. It did not seem to suffer pain, except during some of 
the more severe tetanic convulsions, when it cried out. After 
the fourth or fifth day it had a constant and well-marked expres- 
sion of hebetude. On the 21st of February, fourteen days after 
the removal of the second lobe, having made a solution of fresh 
thyroid gland in a mixture of neutral salts, with the intention 
of now and then injecting a little into a vein of the animal, to 
see whether any evidence could be got in support of Schiff’s idea 
that when the substance of the thyroid is absorbed into the 
blood the animal is preserved in life, I was just preparing to 
make the first injection, when the animal died, in a strong con- 
vulsion. 

The animal, as has already been said, took its food well until 
three or four days before its death ; and accordingly the general 
condition was not reduced to the same extent as in A and P. 
Its weight on the day of death was 4 kilogrammes 10 deca- 
srammes, the original weight having been 5 kilogrammes. 

Post-mortem.—In A and B, from which the thyroid alone had been 
removed, the post-mortem appearances were entirely negative. Distinct 
anemia of the brain or spinal cord was not found. The vagi and 
recurrent laryngeal nerves were perfectly intact. Instead of the spleen 
showing any sign of blood-formation, it seemed smaller and drier than 
usual. Unfortunately, I did not notice whether the red bone-marrow 
had extended into the fatty marrow, 

In C, from which both spleen and thyroid had been removed, the 
post-mortem appearances were exactly the same as those in the dog of 
Experiment I., from which only the spleen had been removed ; with 
this exception, that the lymphatic glands were not enlarged. ‘The red 
marrow had extended into the shafts of the humerus and femur, and 
contained a considerable number of nucleated red cells, the marrow in 
the ribs and in the heads of the long bones being richest in them. 
Neither in this case nor in the two others could any increase in the 
size of the liver or excess in its vascularity be observed. : 

These experiments, taken together with the absence of any 
sign of haematopoiésis in the thyroid gland of the dogs of my 
other experiments, or in the second half of the thyroid of the 


686 DR JOHN LOCKHART GIBSON. 


dogs of Experiments V. and XI., warrant the assertion that the 
thyroid has apparently no blood-forming function, and that any 
such function it may really have can only be one extremely 
subordinate to some other and far more important function. Of 
any relationship between the spleen and the thyroid gland I 
have found no sign. 


Although the further consideration of the results of excision of the 
thyroid will be beyond the proposed object of this paper, yet I cannot 
here refrain from comparing with the observations on animals those 
which have recently been made on human subjects from whom the 
whole gland has been removed. For when the two sets of observa- 
tions are taken together they teach us a very important and most 
practical lesson, namely, that in the human subject the thyroid should 
never be removed entirely ; but that at any rate a small piece should 
always be left, if we are to prevent the appearance of symptoms which 
render the prolongation of life hardly desirable. 

Until within the last three or four years surgeons took no particular 
notice of the isolated cases of tetany which appeared after total excision 
of the thyroid ; and it was only when Weiss,! Wolfler,? and Falkson* 
showed that a considerable number of the cases of such excision in the 
cliniques of Professor Billroth, in Vienna, and Professor Schénborn, in 
Konigsberg, exhibited symptoms of tetany within a few weeks after 
the operation, that special attention was drawn to the connection 
between such symptoms and the particular operation they followed. 
Some of the patients, after having for days been kept under the influ- 
ence of narcotics, recovered ; and others died. Falkson supposed the 
tetany to be due to the division of the recurrent laryngeal nerves; but 
this cannot be the explanation, as the tetany symptoms are very like 
the convulsive symptoms in dogs after excision of the thyroid, where 
the recurrent nerves are quite intact. Weiss’s paper gives a good 
account of tetany, and graphically describes two cases of it after 
excision of the thyroid. Falkson, too, describes the symptoms observed 
by him, which are very like those in dogs. 

The tetany, however, which showed itself soon after the operation, 
as a rule soon passed off; and it was not until Kocher and Reverdin 
wrote on cachexia strumipriva that the really serious effect produced 
by the removal of the whole of the thyroid was pointed out. And 
even in spite of these writings surgeons are still excising the whole of 
the thyroid, as if in operating on the gland they need care merely for 
recovery from the surgical wound. 

Kocher took the trouble to ask all his cases of excision (total) of the 
thyroid to come back and report themselves, and has published the 


1 Weiss, “Ueber Tetanie,” Volkmann's Sammlung klinischer Vortrdge, No. 
189, (1881.) 

? Wilfler, “‘ Die Kropfexstirpationen an Hofrath Billroth’s Klinik von 1877- 
1881,” Wiener med. Wochenschr., 1882, No. 1. 

’ Falkson, ‘‘Zwei Fille von Tetanie nach Kropfexstirpation,” Berl. klin. 
Wochenschr., 1881, No. 12. 


* 


BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 687 


result.1_ He excised the whole gland thirty-four times. Five patients 
died, viz., three of the operation, and two from unknown causes after 
healing. Of the others, eighteen presented themselves in person, and 
six more wrote or were written about. The written accounts do not give 
evidence of much value. Four of the patients wrote that they were well. 
The husband of another wrote that his wife’s body was swollen and 
her limbs insensible, that she never felt warm, and that her menstrua- 
tion was irregular. And the husband of the last of the six wrote that 
his wife could not write herself, from nervous pain in the hands and 
feet, which, he said, were almost paralysed. Of the eighteen who 
presented themselves in person, only two showed no evil effects ; and 
even these two might be said to be only the exceptions which proved 
the rule. For in one of them there was a small accessory thyroid, 
which had become enlarged ; and in the other there was what Kocher 
calls a “ Strumarecidiv ” (relapse of goitre), a small piece of the gland 
having been left, and having enlarged to about the size of a pigeon’s 
egg. The others all showed more or less interference with the general 
health, and their symptoms were so much alike that Kocher describes 
them collectively under the name of “cachexia strumipriva.” 

The symptoms were as follows :—The patients, as a rule soon after 
leaving the hospital, but in some cases only four or five months after 
the operation (Reverdin quotes a case where the time was a year, and 
Martin one where it was six months), began to complain of tiredness, 
and especially of weakness and heaviness of the limbs. In many 
eases the feeling in the limbs amounted to pain. Also, pains in the 
neck, shoulder, and body were complained of. Then followed a feel- 
ing of cold in the extremities, with swelling of the hands and feet, 
which became of a bluish-red colour, and in winter were very cold and 
generally frost-bitten. Further, the intellectual activity became 
diminished, this being especially noticed in children at school, who 
went down in their class, and in whom their teachers found a constant 
decrease of mental capacity, even where they had before been the best 
pupils. Cerebration became very much delayed, a consequent slowness 
of speech became evident, and even the general movements of the 
body became slower. In some patients who were servants, the general 
increase of these symptoms made it necessary for them to leave their 
places, as they could not get through their work quickly enough. 
Some patients did not trouble themselves about the symptoms, while 
others complained of them very much, or had become very retired, 
because they were conscious of their slowness of cerebration, and that 
they were not like other people. Moreover, swellings appeared, 
affecting the face, hands, and feet, and being at first transient, lasting 
only a few hours, and then disappearing. In one case they appeared 
only after long pauses ; and in that case, while they lasted, the patient 
suffered from great breathlessness. In many patients the swelling re- 
mained, giving their faces a heavy, stupid look, and so making their 
acquaintances think they had become idiots. The whole face became 
thicker: the eyelids swollen and transparent, the nose thick, and the lips 


1 Kocher, ‘‘ Ueber Kropfexstirpation und ihre Folgen,” Arch. f. klin. Chir. ; 
Bd, xxix, 


688 DR JOHN LOCKHART GIBSON. 


likewise swollen. The hands and feet, too, and even the body, became 
permanently swollen ; and in two cases there was ascites. The skin over _ 
the swollen parts was thick and infiltrated and had lost its elasticity, the 
surface was dry, and the hair fell out. In advanced cases there 
was marked anemia, with a distinct decrease in the number of red 
corpuscles, and without, in the cases ‘ counted,” any distinct increase 
in the number of white. The minimum number of red corpuscles was 
2,168,000, four cases had 2,800,000, five had fewer than 3,500,000, 
three had fewer than 4,200,000, and two had fewer than 4,500,000. 
Two cases with slighter symptoms had a pretty normal number, viz., 
4,940,000 and 5,520,000. Only one patient with the symptoms at all 
marked had as many as 4,476,000. Where Kocher made enumerations 
in cases of partial excision, the number of red corpuscles was found 
normal. Where at the time of total excision the patient was still grow- 
ing, the growth was very much retarded. Kocher records that one 
case had distinct attacks of tetany, and, later, attacks of epilepsy; and, 
in connexion with this, remarks that many of the other cases had, in 
addition to the tired feeling already mentioned, also a stiffness of the 
limbs. The muscles were always well developed ; in fact, resembled 
those of pseudo-hypertrophic paralysis. Fibrillar and localised con- 
tractions of the muscles were observed. Kocher says that although 
the patients looked stupid they were not really idiots, but were capable 
of recognising and feeling the change in themselves. 

Kocher then proceeds to consider whether these symptoms could not 
be accounted for by the anzemia present, and thinks that the anemia 
perhaps gives some support to the idea of a blood-forming function of 
the thyroid. It seems to me, however, far more likely that the changes 
in the nervous system were caused directly by the removal of the 
thyroid, and instead of being mere consequences of the angemia were 
the causes of it, though it in turn, once developed, would of course 
intensify them. This explanation is quite borne out by my observa- 
tions on dogs, where the nervous symptoms made their appearance as 
the direct result of the excision, and could not be ascribed to anemia, 
which did not exist. Moreover, Kocher’s own blood-estimations show 
that in his cases the anemia made its appearance only after the other 
symptoms had become developed. And, indeed, Kocher admits that 
he cannot find proof of the thyroid being a blood-forming organ. If, 
he says, the thyroid replaces the spleen, why does not the spleen 
replace the thyroid? In none of his cases could he observe an increase 
in the size of the spleen. In his last case of total excision of the 
thyroid he estimated the blood for the first few weeks after the opera- 
tion, and found no change. And in cases of one-sided excision of the 
thyroid he was never able to find any subsequent swelling of the 
remaining lobe. 

Further, he tries to account for the anemia by changes in the trachea. 
He supposes that the trachea, receiving most of its blood-supply from 
the thyroideal vessels, is after excision of the thyroid not well 
nourished ; that consequently its walls become softer and its lumen 
narrower ; and that in this way the patients are prevented from getting 
enough of oxygen—in fact, suffer from an increasing deprivation of it. 


BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 689 


This explanation, however, cannot be accepted. In my dogs the 
trachea was examined particularly, and there was neither softening nor 
narrowing, and it seemed very fairly supplied with blood. Moreover, 
both in my dogs and in the animals experimented on by other ob- 
servers, the shortness of the time the animals survived is against the 
symptoms having been dependent on narrowing of the trachea. And, 
to return to human beings, many people go about with the larynx con- 
stricted by cicatrices, &c., to a much greater extent than in Kocher’s 
cases the trachea can have been constricted, and yet are able, by a 
slightly quicker respiration, to make up for the narrowness of the 
channel. 

Other explanations are: (a) the earlier one advanced by Schiff and 
Liebermeister, that the thyroid regulates the circulation in the brain 
(a function apparently first suggested by Maignien’) ; (4) the explana- 
tion more recently advanced by Schiff, that the thyroid produces some 
substance whose absorption into the blood is essential to life; and (c) 
the explanation that the thyroid secretes and excretes a substance 
injurious to the organism, an explanation which has received some 
support from the experiments of Victor Horsley (see next page). 

As to the last of these explanations, a great difficulty in the way of 
its acceptance is the fact that the thyroid has no excretory duct. And 
this difficulty does not seem got over by reference to the lymphatics, 
and their position in close relation to the gland vesicles. For to 
suppose that the injurious substance is taken up by the lymphatics 
would involve the improbable-looking further supposition of a passage 
from these back again into the blood, with a subsequent re-excretion 
by some other organ. 

Of Schiff’s two explanations the second seems much the more 
tenable, as being in accordance both with Schiff’s own experiments 
and with the observations of Sanquirico and Canalis, where only a 
very small piece of the thyroid was left. 

His earlier explanation, that the thyroid regulates the circulation in 
the brain, seems very insufficient. ‘The theory of sucha function is 
that during great muscular exertion, and consequent violent action of 
the heart, the pressure of contracted muscles on the jugular and thyr- 
oideal veins makes the thyroid become engorged with blood and swell 
up and thus press on the carotid arteries and prevent overfilling of the 
cerebral arteries. According to Guyon,? the pressure of the thyroid 
may even render the branches of the external carotid pulseless. 
Against this explanation, Schiff’s grafting-experiments, where placing 
the freshly-excised thyroid of a dog in the abdomen of another dog 
seemed in the case of subsequent excision of the thyroid of this dog to 
prevent the appearance of symptoms after such excision, would, if 
confirmed, be conclusive evidence. Moreover, the partial excisions of 


1 Quoted, in 1842, by Longet, in “Anatomie et Physiologie du Systéme 
Nerveux,” tome i. p. 807. 

2 Guyon, “ Note sur l’arrét de la circulation carotidienne pendant l’effort pro- 
longeé,” Arch. de Physiologie, t. 1. (1868).—Quoted by Nothnagel, in Ziemssen’s 
Handb. d. spec. Path. u. Ther., Bd. xi. 1, p. 6. 


699 DR JOHN LOCKHART GIBSON. 


Sanquirico and Canalis, where the leaving of one-third of one lobe 
sufficed for the preservation of life, are greatly against it. And here. 
it may be pointed out that a crucial experiment might be made on 
dogs. In these animals the vertebral arteries are so large that both 
carotids may be tied without material injury to the animal, and the 
experiment would consist in tying both carotids of a dog suffering 
from the effects of excision of the thyroid. If the symptoms following 
excision are really due to relief of the carotids from compression by 
the thyroid, then the ligature of the carotids ought at once to put a 
stop to them. I regret not having had time to make the experi- 
ment, but am very sure that no such effect would have been 
produced. 

In December 1884, in the ‘Brown Lectures,” Victor Horsley+ 
described some experiments on monkeys which yielded interesting 
results. By excising the whole thyroid he produced a condition and 
symptoms closely resembling those observed by Kocher in human 
beings, but in some respects intermediate between them and those 
observed in dogs. He found, moreover, an increased production of 
mucin in the body generally ; and in the case of the salivary glands 
found not only that those other than the parotid secreted far more 
than their normal amount of mucin, but also that even the parotid 
secreted it. He looks on the condition produced by his excisions as 
being that called myxoedema by Ord,? who applied the term to the 
general swelling in the cretinoid condition that occurs idiopathically 
in adult women; and he holds that myxoedema, cachexia strumipriva, 
and cretinism are processes identical in nature. As in cases of excision 
in human beings, so here too there was anzmia, which Horsley, con- 
sidering blood-formation one of the functions of the thyroid, attributed 
to its removal. The animals all died, usually in from five to seven 
weeks. Horsley’s conclusions are: that myxoedema and its allies are 
due to virtual or actual loss of the thyroid gland; that, however, the 
immediate cause of the changes may be vaso-motor and trophic lesions 
due to the loss; that the thyroid probably excretes a substance injuri- 
ous to the organism, and that this substance is of the nature of mucin ; 
that removal of one lobe causes enlargement of the other; and that 
subsequent removal of the other entails death, after production of the 
condition and symptoms described by him. 

Against the no doubt pretty strong arguments brought forward by 
Horsley in favour of an excretory function of the thyroid, I may here, 
besides referring back to the objection already made, of the absence of 
an excretory duct, also again refer to the apparent success of Schiff in 
securing dogs against a fatal result of excision of the thyroid by 
placing the freshly excised thyroid of another dog in the abdominal 
cavity some weeks before the operation. 

With reference to the surgical treatment of diseases of the thyroid, 
Horsley is reported (Lancet) to have condemned total removal of the 
thyroid, and to have done so on the ground that good results may be 
obtained by removal of a portion of it. 

1 Report of ‘ Brown Lectures,” Lancet, 27th Dec. 1884. 
2 Lancet, 27th Oct. 1877. 


BLOOD-FORMING ORGANS AND BLOOD-FORMATION. 691 


The conclusions to which my experiments and the observa- 
tions of others have led me are :— 


1. That the thyroid has, properly speaking, no blood-forming 
function. 

2. That any blood-forming action it may in some animals 
seem to have is due only to the presence of lymph-follicle- 
tissue in the thyroids of such animals. 

3. That its function is in no way compensatory to that of the 
spleen. 

4. That in dogs the total removal of the thyroid is always 
followed by death, after a definite train of nervous symptoms. 

5. That in dogs the presence of unusually well developed 
aortic thyroid glands may prevent the onset of the symptoms. 

6. That the removal of the whole thyroid from the human 
subject is unjustifiable, such removal being always, after a vary- 
ing interval, followed by the development of the very serious 
definite condition called “ cachexia strumipriva.” 

7, That in all excisions of the thyroid from the human sub- 
ject at least a small piece should be left, a small piece appearing 
to be sufficient for the carrying on of the essential function of the 
gland. 

8. That the function of the thyroid has special relation to the 
central nervous system, though what the true nature of such 
relation may be has yet to be definitely determined. 

9. That myxcedema, cachexia strumipriva, and cretinism are 
probably one and the same disease, and are due to virtual or 
actual absence of the thyroid. 


Anatomico-Pbysiological Jlotices. 


PLEXIFORM ARRANGEMENT OF THE CUTANEOUS NERVES 
IN THE GROIN. By Davin Hersurn, M.B., Senior Demon- 
strator of Anatomy, University of Edinburgh. 


Tue following notes were taken from a dissection performed by Mr 
R. J. Pope in the Practical Anatomy Rooms of the University of 
Edinburgh :— 

The external cutaneous nerve entered the thigh in its usual 
position, but soon gave off a branch, which, dividing into two portions, 
formed communications with the middle cutaneous nerve and the 
crural branch of the genito-crural nerve. 

The genito-crural nerve, in addition to the communication just 
mentioned, formed two junctions with the middle cutaneous nerve, and 
these were both found after the latter had pierced the sartorius 
muscle. 

The middle cutaneous nerve had the communications already men- 
tioned, namely, with the external cutaneous and crural branch of the 
genito-crural. When fully traced to its ultimate distribution, both its 
inner and outer divisions were seen to pierce the sartorius muscle and 
thereafter to form junctions with each other. 

The internal cutaneous nerve gave off the nerve to the pectineus 
muscle. In addition, it had three branches of some size distributed in 
the following manner :—one to the inner side of the thigh at its upper 
part ; another to the inner side of the thigh at its lower part ; and the 
third along the inner edge of the sartorius muscle. 

When Scarpa’s triangle was fully worked out, the presence of an 
accessory obturator nerve was revealed. It passed outwards over the 
brim of the pelvis, beneath Poupart’s ligament and the pectineus 
muscle, the latter receiving filaments of supply from the nerve in this 
position. Small twigs were also supplied to the hip-joint, and in 
addition it gave off a communicating branch which joined the super- 
ficial division of the obturator nerve, and thereafter, emerging from 
between the pectineus and adductor longus muscles, it passed upwards 
and outwards beneath the femoral artery and vein, and terminated by 
joining the internal saphenous nerve about 4 inches from Poupart’s 
ligament. 

As the dissection of the front of the thigh proceeded, a thin filament 
from the external cutaneous nerve was traced downwards over the 
surface of the sartorius muscle. It entered Hunter’s canal, and lay in 
the connective tissue surrounding the femoral artery. When dissected 


ANATOMICO-PHYSIOLOGICAL NOTICES. 693 


from this position it appeared to end in two fine filaments, which were 
traced into the substance of the vastus internus muscle. 

In the popliteal space the obturator nerve was carefully looked for, 
and it was found lying on the popliteal artery, by which it was carried 
to the posterior aspect of the knee-joint, which it entered by passing 
through the ligament of Winslow. From the popliteal space this 
nerve was traced upwards through the fibres of the adductor magnus 
muscle, and on the anterior surface of this muscle it was found to be 
continuous with the deep division of the obturator nerve. 


NOTE OF A CASE OF ABSENCE OF VAGINA, WITH UN- 
DEVELOPED UTERUS AND OVARIES. By J. A. Camp- 
BELL M.D., F.R.S.E. 


S. W., zt. 27, was admitted into the Carlisle Asylum in January 1877. 
She was stated to have been insane for seven years, and at intervals to 
have taken epileptic fits. She had never menstruated. Her mental 
state had alternated between excitement and depression, and she was 
said to have shown erotic tendencies during the periods of excitement, 
and to have been in the habit of exposing her person at such times. 
The doctor who attended her wrote to me to say that he thought she 
had an abdominal tumour, due to retained menses from imperfect 
hymen, and that I should look to this at once as a possible cause of 
her state. 

On admission, a full examination of her mental condition and phy- 
sical state was made and recorded. I quote the following :—“ She was 
undersized, had a low type of face, her lungs and heart were normal, 
and nothing abnormal was detected as regards abdominal viscera ; her 
mamme were well developed. She had much hair over pues ; her 
external genital organs were well developed, but no vagina could be 
found.” Owing to the expressed opinion of her former medical attend- 
ant, I asked Mr Page of Carlisle to examine her with me a few days 
after her admission. We agreed that no abdominal tumour existed ; 
that though the external genital organs were normal, there was an 
absence or occlusion of the vagina, a white glistening structure form- 
ing an impervious cul-de-sac where the opening of the vagina should 
have been. A catheter passed into the bladder could, by rectal 
examination, be distinctly felt without the intervention of anything 
corresponding to a uterus. 

The patient lived in the asylum for nine years, never menstruated, 
was examined on one or two occasions without anything further being 
made out. She had epileptic fits of a severe character at considerable 
intervals, and had attacks of excitement at short intervals of three, 
four, or five weeks, and at such periods she exhibited erotic tendencies, 
stripped off her clothes, and exposed her person. She ultimately de- 
veloped phthisis, and died of this disease. 

VOL. XX, 22 


694 ANATOMICO-PHYSIOLOGICAL NOTICES. 


Autopsy.—The viscera occupying the different cavities were care- 
fully examined and their state recorded. I need only mention that 
tubercular deposit of lungs and ulcers of intestines caused death. 

The external genital organs showed the appearances noted in the 
case. The internal organs of reproduction were almost undeveloped. 
The ovaries—very small, shrivelled, and indurated—occupied a normal 
position. The Fallopian tubes communicated with a rudimentary 
uterus about seven-eighths of an inch in length. So far as could be 
made out, it was devoid of an os, and lay very low in the pelvis. 
Examination failed to show any communication between uterus and 
external genital organs. 

Remarks.—Though I am quite aware such cases are not unknown, 
yet they are of sufficient rarity to make it worth while putting such a 
case on record. It is, I think, highly probable that the attacks of 
excitement occurred at what would, under other circumstances, have 
been monthly periods. A degree of sexual excitement was distinctly 
present in the case. 

If the ovaries shed ova, what became of them ? 


NOTES ON AN ABNORMAL ARRANGEMENT OF THE 
LARGE INTESTINE. By B. C. A. Winpuz, M.A, M.D.,, 
Professor of Anatomy in the Queen’s College, Birmingham. 


Tue following unusual arrangement of the large intestine was observed 
in a male subject recently examined in my dissecting room, and may 
be of interest when taken in conjunction with the papers which 
have appeared in the Journal of Anatomy and Physiology, and with 
Mr Treves’ recent lectures on the Anatomy of the Human Intestinal 
Canal and Peritoneum. 

The cecum was large and placed entirely above the margin of the 
ilium. It did not come into contact at all in its attachments with 
the posterior wall of the abdomen. The vermiform appendix lay on 
the iliacus muscle, and was directed downwards and outwards. The 
colon passed at first directly backward to reach the posterior wall, 
whieh it did just above the crest of the ilium. From this point it 
pursued its ordinary course until it arrived at the crest of the left 
ilium. Here it seemed to sink into a kind of peritoneal pocket, 
formed by a process passing from the anterior lateral wall to the anterior 
and inferior surface of the colon, which turned upon itself here, and 
ascended along the posterior wall of the abdomen on the right side of 
the descending colon. It was included in the same fold of peritoneum, 
and into the neck of the fold passed the sigmoid artery, which broke 
up into two sets of branches to supply the two tubes. Arrived at the 
splenic flexure, the ascending tube was held firmly in position by a 
process of peritoneum passing from the sustentaculum lienis, which 
was exceptionally strong, including the splenic flexure of the trans- 


ae 


ANATOMICO-PHYSIOLOGICAL NOTICES. 695 


verse and descending colon, and passing on to the second tube. The 
free edge of this process formed the margin of a deep and capacious 
pouch, whose concavity was directed to the right side of the abdomen. 
From this point the colon again descended, and was again, but to a 
much smaller extent, looped up by a band of peritoneum passing to it 
from in front of the left side of the vertebral column close to the duo- 
deno-jejunal fold. The free margin of this band also marked off a 
pouch of less dimension than the first mentioned, the direction of whose 
concavity was similar to that of the larger, in which it was included. 
The colon after this small loop passed down in a nearly straight line 
to the rectum. In this course it was bound to the posterior wall close 
to the vertebral column by a fold of peritoneum, the line of reflection 
of which on either side was nearly straight. There were thus three 
tubes of large intestine throughout the entire length of the left side of 
the abdomen, two descending and one ascending. The peritoneum 
on the posterior wall was somewhat roughened, and there were a few 
evidently morbid adhesions between the coils of intestine. These 
were, however, quite different from the true process, in which the first 
mentioned two tubes were included. 


A NEW METHOD FOR THE QUANTITATIVE ESTIMATION 
OF URIC ACID. By Joun B. Haycrart, M.B., B.Sc, 
E.R.S.E., Professor of Physioloyy, Mason and Queen’s Colleges, 
Birmingham. 


Wisuine to continue a research upon albuminoid metabolism, and 
having been much disheartened by the unsatisfactory nature of the 
processes used for the estimation of uric acid, I determined, if possible, 
to introduce one which should be at the same time exact and easy of 
application. 

The method of Heintz is certainly not very reliable, and is not 
applicable when small quantities of the acid are present in the urine. 
Salkowski’s modification, although certainly more reliable, is far too 
laborious, and is not free from other objections. The same may be 
said of a somewhat similar process introduced by Fokker. Other 
methods, depending on the power which the acid has of reducing 
certain metallic salts, or upon the decomposition of the uric acid, and 
the collection of its decomposition products, I tried, but soon had 
to give them up. Inasmuch as uric acid forms some salts more in- 
soluble than itself, and as some of these are with metals easy of estima- 
tion, it occurred to me that it might be possible to precipitate it as a 
salt, say of lead, barium, or silver; and then, by estimating the metal 
in the precipitate, to calculate the uric acid in combination with it. 

Many experiments were undertaken with the uric acid salts of 
barium, mercury, and lead; their solubilities in various acids, &c., 
were tested; and endeavours were made to separate (in these forms) 


696 ANATOMICO-PHYSIOLOGICAL NOTICES. 


the acid from solutions of mixed salts and other compounds found in 
urine, and likely to interfere with the process. One or two rough 
methods were devised, but were soon discarded from their want of 
accuracy. An attempt to separate out uric acid asa silver salt was 
more successful. 

Urate of silver is, as is well known, very insoluble in water. I find 
it to be very soluble in 20 per cent. nitric acid, and in somewhat 
stronger sulphuric acid. It is insoluble in ammonia, and almost in- 
soluble in strong acetic acid. 

After a futile attempt to separate out the uric acid from the phos- 
phates of the urine, after acidulation with acetic acid, I succeeded in 
elaborating the following process :— 

The fact that urate of silver is insoluble in ammonia. water is a very 
curious one, inasmuch as this is a solvent for nearly all the silver salts 
—-for example, the chlorides, phosphates, and oxalates. On adding a 
solution of silver nitrate to a solution of acid urate of sodium, I found 
that, instead of a precipitation of silver urate, an immediate reduction 
occurred, the black precipitate not re-dissolving in ammonia water. 
If, however, the solutions be previously rendered ammoniacal, a white 
gelatinous precipitate of the urate at once forms. The silver, however, 
becomes partially reduced before it is possible to collect and wash it. 
I find, however, that the previous addition of bicarbonate of sodium 
prevents this reduction. The same obtains with the acid urate of 
sodium normally present in urine, the chlorides and phosphates re- 
maining in solution, the urate of silver alone falling on the addition 
of the ammoniacal nitrate. In order to estimate the silver in this 
precipitate of the urate, the latter was collected in a filter, and washed 
with distilled water, and taking advantage of its ready solubility im 
nitric acid, it was dissolved in that reagent. 

A method not long introduced by Volhard,! and one which is avail- 
able for the estimation of silver in an acid medium, was used as the 
final step of the process. 

The only difficulty that occurs is in filtering and washing the pre- 
cipitate of urate of silver. It is gelatinous, and clogs up the filter. It 
is imperative to use a Sprengel’s pump, and a properly made asbestos 
filter is advisable. In this case, the whole process may be completed 
in little more than half an hour. The filter should be prepared in the 
following way :—A small funnel is half filled with broken glass. A 
sufficient quantity of asbestos is shaken in a flask with water until all 
the fibres have separated, forming an uniform pulp. This is poured 
on the glass, and should form an uniform felt, one quarter of an inch 
thick. The filter can be used again and again for several analyses. 


Description oF MeEtHop. 


Solutions Required.—1. Centinormal ammonic sulphocyanate. Dis- 
solve about 8 grammes of crystals in a litre of water, and adjust it to 


' “Die Anwendung des Schwefeleyammoniums in der Maasanalyse,” Liebig’s 
Annalen, Band exe. p. 1. See also Sutton’s Volumetric Analysis. 


ANATOMICO-PHYSIOLOGICAL NOTICES. 697 


decinormal silver solution. Dilute with 9 volumes of water. One 
cubic centimetre is equivalent to 0-00168 of uric acid. 

2. A saturated solution of iron alum. 

3. Pure nitric acid (20-30 per cent.). Dilute the commercial acid, 
boil and preserve from light in a blackened flask. 

4, Strong ammonia. 

5. Ammoniacal silver solution. Dissolve 5 grammes of nitrate in 
100 cubic centimetres water, and add ammonia, until the solution be- 
comes clear. 

Process.—Measure off 25 cubic centimetres of urine in a pipette, 
and place it in a small beaker, with about 1 gramme of bicarbonate of 
sodium. Add 2 or 3 cubic centimetres of ammonia, which will pro- 
duce a precipitate of ammonia-magnesium phosphate. On adding 1 to 
2 cubic centimetres of the ammoniacal silver solution, the uric acid 
falls as a white gelatinous precipitate of urate of silver. 

This is collected on the asbestos filter, and carefully washed, until 
the washings give no trace of silver, with a drop of salt solution. The 
urate is then washed through the filter by the aid of a few cubic 
centimetres of the nitric acid, and the silver in this solution estimated 
by Volhard’s method, 

Add a few drops of the saturated solution of iron alum, which is 
the indicator, and drop in the centinormal solution of ammonic sulpho- 
cyanate. A white precipitate will form, together with a transient 
reddish coloration, which latter becomes permanent when the process 
is at an end. 

It is easy to calculate the uric acid, which will be the number of 
cubic centimetres of the sulphocyanate used multiplied by 0:00168. 

If the urine contain albumen, this should previously be removed. 
If uric acid or urates be present in such quantity as to cause turbidity, 
the secretion should be warmed and diluted. 

In order to test the accuracy of the process, I prepared several solu- 
tions of acid urate of sodium of known strength. To these I added 
various quantities of common salt, sulphate of magnesia, and phos- 
phate of soda, in order to imitate, as far as possible, the urinary 
secretion. On estimating the uric acid in these solutions, I obtained 
wonderfully correct results, as the subjoined tables will show. In all 
cases a quantity not much more than a milligramme was lost during 
the process, and may be simply accounted for by the fact that no salt 
of uric acid is absolutely insoluble. 


Experiment 1. Experiment 2. Experiment 3. 


Uric acid present as urate, . 0°0486 0°0459 0°0211 
Uric acid found, ; , 0°0470 0°0437 0:0198 
Loss during process, . : 0°0016 00022 0:0013 


3 percent. 5 percent. 6 per cent. loss. 


In order further to test its accuracy, 50 cubic centimetres of urine 
were divided into two equal portions; to the first, 25 cubic centimetres 
of a solution of acid urate of sodium of known strength were added; 
to the second, 25 cubic centimetres of water were added. 

When estimated, the two fluids should show a difference equal to 
the quantity of the salt added, 


698 ANATOMICO-PHYSIOLOGICAL NOTICES. 


Experiment 1. Experiment 2. Experiment 3. 


Uric acid found (a), . : 0°034 0026 0°041 

Uric acid found (0), . : 0°0144 0°0105 0°0254 
Difference, : : 3 0°0196 0°0155 0°0156 
Uric acid added, : ; 0021 0°016 0°0160 


The above figures show how delicate the process is, and what 
accurate results may be obtained from its use. This depends, in the 
first case, on the great insolubility of the silver urate; and secondly, 
on the great delicacy of the final estimation of the silver by the 
ammonic sulphocyanate. It may be objected to this process that there 
may be other substances present in the urine which may form com- 
pounds with the silver nitrate, thus vitiating the result. Xanthin, as 
is well known, combines with silver. This substance may be present 
in small quantities, but its silver compound is, I find, insoluble in 
nitric acid. I am aware of no substance present in urine normally, 
or likely to appear as an abnormality, which forms a combination 
with silver both insoluble in ammonia and soluble in nitric acid. 

I should probably have tried much sooner the separation of the 
urie acid as a silver-salt, but for a statement of Salkowski’s.! This 
distinguished chemist analysed the precipitate which forms after 
adding ammoniacal nitrate of silver to urine. He found a variable 
quantity of both silver and magnesia, and concluded from this that 
the uric acid precipitates out, both as an urate of magnesium and an 
urate of silver. In this case, I could not, of course, use the silver 
process for its estimation. His facts may otherwise be explained. As 
is well known, ammonio-magnesium phosphate separates out slowly. 
Salkowski, immediately after the addition of ammonia, filtered off the 
ammonio-magnesium phosphate, and then added silver-nitrate. Pro- 
bably, with the urate of silver, a small and variable quantity of the 
double phosphate came down, for this would probably be not yet 
entirely precipitated. As to the variable quantity of silver, this is 
easy to account for. In my own investigations, I attempted at one 
time to estimate the uric acid by its reducing action on silver (a 
method which was quite unsatisfactory), and I found that the same 
quantity of urate reduced a very variable quantity of silver, depend- 
ing upon two factors—time and temperature. With prolonged boiling, 
as many as four parts of silver are reduced by one of the urate. Now, 
in Salkowski’s investigations, the reduction of the silver was disre- 
garded (for he speaks of the black precipitate on the filter-paper). 
The amount of the reduction would not be the same in any two ex- 
periments. In my own method, all reduction is prevented by the 
addition of the bicarbonate of sodium. I have since added to urine 
large quantities of magnesium-sulphate, without interfering with the 
accuracy of the results. 

This method was elaborated during the year 1883. I have much 
pleasure in expressing my indebtedness to the Scientific Grants Com- 
mittee of the British Medical Association for assistance in the shape of 
a grant to cover my expenses. 


1 Pfliiger’s Archiv., B. 5, p. 210. 


INDEX. 


Abnormal Arteries, 26. 
Accommodation of Eyes, 475, 563. 
Alimentary Canal in Ganoids, 602. 
Alligator, Heart of, 547. 

Animals, Wild, Vitality of, 594. 
Arterial System, Morphology of, 193. 
Arteries, Abnormalities of, 26, 31, 32. 
Axis and Os Odontoideum, 238. 


Berling, Gilbert, on Hypertrophy of Leg, 
58. 


Biceps, Bursa connected with, 30. 

Bicipital Rib, 405. 

Bile Secretion and Urea Formation, 114, 
267, 520, 562. 

Birds’ Eggs, Pigment of, 225. 

Blood-Forming Organs, 100, 324, 456, 674. 

Brooks, St John, Abnormal Coronary Ar- 
tery, 26; Variations in Nerve Supply 
of Thumb Muscle, 641; Muscles of 
Hand, 645. 

Bursa connected with Biceps, 30. 


Campbell, J. A.,on Absence of Vagina, 693. 
Carapax of Tortoises, 220. 

Collins, A. W., on Rare Abnormalities, 30. 
Conurus carolinensis, Skeleton of, 407. 
Coronary Artery, Abnormal, 26. 

pel John, on Pentadactylous Pes, 


Cunningham, Professor, Axis and Os Odon- 
toideum, 238; on Spines of Cervical 
Vertebre, 637. 

Curran, William, Vitality of Wild Ani- 
mals, 594. 

Cysts, Origin of, 432. 


Dentition of Shrews, 359. 

Die Chirurgische Anatomie, You Pro- 
fessor Max Schiiller, 548. 

Dobson, G. E., on Dentition of Shrews, 359. 

Dwight, Thomas, Frozen Sections of a 
Child, 192. 


Eggs of Birds, Pigment of, 225. 

Eye, Convergence and Accommodation of, 
475, 563. 

Eye, Suspensory Ligaments of, 1. 


Floating Kidney, 544. 

Fowl, Dorking, Pes in, 591. 

Fox, R. H., on Function of Tonsils, 557. 

Fraser, J. W., on Infused Beverages and 
Peptic Digestion, 361. 


Freeman, R. A., on Anatomy of Mole, 
201. 


Frog, Supernumerary Leg in, 516. 


| Gadow, Hans, on Reproduction of Cara- 


pax, 220. 
Ganoids, Alimentary Canal in, 602; 
Pancreas in, 629. 


| Geococcyx, Skeleton of, 244. 
Gibson, J. Lockhart, on Blood-Forming 


Organs, 100, 324, 456, 674. 


Hand, Nerves and Muscles of, 641, 645. 

Hare, A. W., on Micro-organisms, 76. 

Hay craft, Professor, Estimation of Uric 
Acid, 6 

Heart of ” Alligator, 547. 

Hepburn, David, Plexiform Arrangement 
of Nerves, 692. 

Histological Methods, 307. 

Humphry, Prof., on Old Age and Changes 
Incidental to it, 191; on Spina Bifida, 
546, 583. 

Hunter, Wm., on Histological Methods, 
307. 

Hypertrophy of Leg, 358. 


Ichthyosaurus, Hind Limb of, 532. 

Index of Pelvic Brim as Basis of Classifi- 
cation, 125. 

Infused Beverages, 361. 

Inter-clavicular Muscle, 544, 

Intestine, Abnormal Arrangement of, 694. 


Kidney, Floating, 544. 


Lane, W. Arbuthnot, on Variations in 
Skeleton, 388; on Floating Kidney, 
544; on Inter-clavicular Muscle, 544. 

Leg, Hypertrophy of, 358; supernumer- 
ary, 516. 

Ligaments, Nature of, 39. 

Limb of Ichthyosaurus, 532. 

Lumbar Curve in Man, 536. 

Lung, Abnormal Lobe, 32, 35. 

Lockwood, C. B., on Anatomy of Orbit, 1. 


Macalister, Professor, on Morphology of 
Arterial System, 193. 

Macallum, A. B., on Alimentary Canal in 
Ganoids, 602. 

Macdonnell, R. L., on Bicipital Rib, 405. 

Maddox, U. E., on Convergence and Ac- 
commodation, 475, 563. 


770 


Maylard, A. E., on Abnormalities of 
Lung, 34. 

M‘Aldowie, Alex., on Pigment of Birds 
Eggs, 225. 

Micro-organisms, 76. 

Mills, T. ”W., on Heart of Alligator, 547. 

Mole, Anatomy of, 201. 

Morphology of Arterial System, 193. 

Muscles, Abnormalities of, 356, 544. 

Muscles’ of Hand, Nerve Supply, 641; 
Morphology of, 645. 


Nevus, Diffuse Venous, 358. 

Navajo Skull, 426, 430. 

Neural Spines of Cervical Vertebree, 637. 

Nerves, Plexiform Arrangement of, 692. 

Noél-Paton, D., on Urea For mation and 
Bile Secretion, 114, 267, 520, 662. 


Orbit, Muscles, Ligaments, and Fascia 
of, 1. 
Os Odontoideum and Axis, 238. 


Palmer, John, Abnormalities of Pelvis, 354. 
Pancreas in Ganoids, 629. 

Pelvic Index, 125. 

Pelvis, Abnormalities of, 354. 
Pentadactylous Pes, 591. 

Peptic Digestion, 361. 

Pes, Pentadactylous, 591. 

Pigment of Birds’ Eggs, 225. 


Rana palustris, Supernumerary Leg in, 616. 

Rennie, G. E., on Sterno- Petrosoph ar- 
yngeus Muscle, 356. 

repr of Committee on Spina Bifida, 546. 
Reproduction of Carapax, 220. 

Revye 2 Anthropologie, 546. 

Rib, Bicipital, 405. 

Robertson, Robert, on Splenic Histology, 
509, 


Sacral Index, 317. : 
Shrews, Dentition of, 359. 


PRINTED BY 


INDEX. 


Shufeldt, R. W., on Geococcyx, 244; on 
Conurus carolinensis, 407; on Navajo 
Skull, 426. 

Skeleton, Geococcyx, 244; Conurus curo- 
linensis, 407; Variations in Human, 244. 

Skull, Navajo, 426, 430. 

Sowerby’s Whale, 144. 

Spina Bifida, 546, 583. 

Splenic Histology, 509. 

Sterno-Petrosopharyngeus Muscle, 356. 

Sutton, J. Bland, on Ligaments, 39; on 
Origin of Cysts, 432. 


Talpa Europea, Anatomy of, 201. 
_Tenon, Capsule of, 1. 

Thompson, Professor D’Arcy, on Limb of 
Ichthyosaurus, 532. 

Tonsils, Function of, 557. 

Tortoises, Carapax of, 220. 

Treves, K., Anatomy of Intestine and 
Peritoneum in Man, 189 

Tuckerman, Fred.,7on Supernumerary Leg 
in a Frog, 516, 

| Turner, Sir W., on Pelvic Index, 125; on 
Sowerby’s Whale, 144 ; on Sacral Index, 
317; on Navajo Skull, 430; Note on 
Supernumerary Limb, 519; on Lumbar 
Curve in Man, 536. 


Urea Formation and Bile Secretion, 114, 
267, 520, 662. 

Uric Acid, Estimation of, 695. 

Uterus and Ovaries, Undeveloped, 693. 


Vagina, Absence of, 693. 

Vertebrze, Cervical, Neural Spines of, 637. 
| Vitality under Fire, 594. 

Vital Relations of Micro-Organisms, 76, 


Windle, Professor B., Abnormal Large 


NEILL AND COMPANY, 


Intestine, 694. 

Whale, Sowerby’s, 144. 

Woodhead, G. 5., on Micro-Organisms, 
76 


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