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p. [3l7]-330

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MKtocorr mowTiON tmt chart

(ANSI ond ISO TtST CHA»T No 2|

^ APPLIED IfVHGE In

gar. '6^J Cast Main <^tre«t

S"-£ HochMttf. f«c. York 14609 USA

■Wg f^'6} *82 - 03U0 - Phon.

^S ('!«) 2M- 5989 -Fo«

UNIVERSITY OF TORONTO
ST i; DIES

PHVsioi.ociifAi. si:ries

No. VV OBSERVATIONS ON THE GLVCOGEN CONTENT
OF CERTAIN INVERTEBRATES AND FISHES, uv L. G.

KlUlORN AND J. J. R. MACLEOD

iRtPHIMKl) HIDM IIIE Ul <IEBLV JuLBNAL .^^ E.M'EmMENUL PlIVsluLI^V, \uL. Xll)

THE UNIVERSITY LIBRARY: PUBLISHED BY
THE LIBRARIAN, lyjo

COMMlTTtt or MANAOEMINT

I

p,„,PMnii W. J. A««AW«. Pn-D

f EuiMW } }• Utaamm, ».a., MB.

TuMtwmfm y. f- McMotwcb, P11.D.
B«o.^»..C.H.Ji«c«U,B.A.Sc..C.B..C.M.G..D.S.O.

PaoMMW G. H. NHom. Tu.V.
PtanMOU G«o«» M. Wwm». MA.
emumi Minor: H. H. Umwoii, M-A. .^_.

ai

(»HSKKVATI(>\S ON IHK (iLV((HiKN (((NTKNT nh CllilAIN
INVKK'IKIiHA'I'Ks ANI> Kl ilKS. hy I. (i Kii.i...iix nnd
J. .1, II. M.v I.Kuli. ( Ftuiii till' Miiiiiif Hi<ili>;;i<iil Stiiticir Nniiaitno,
B.C., and ttii- DffMiitiiifiit of l'liyHi<>l<>;;y 'I'lii' l'iii\.-rsit_v "I' TMroiito.)

(l!r,tivt'i fnr f.M 11, thiu, ••lOi llrr.iuhr, 11(111 )

Tllltui'iiiliM T llic lUiiinul kiii;;>|i>iii from \\\f .\iiiu Im lo Man ■■iitli<ili\ilral.'
is known to Im' an important i-onNtitin'nt of tlic or;,Mni>m. It is mo.st
cliunu'teristirally rf|ir.'>ifnti-il witliin ihr iiIIm l.y \\-- |io|\ Hi^'i-imiide
jflyootrrn, anil in the i-iicnlatiny; tliiiils hy ;;luco^' In tli.' Iii'dirr
vtTtpbrati'.H till- |KTr(;!itiii,'c iimoiitit of ;c'y''"K''" '" <'"■ 'I'lli'ii-nt oi;;ans
tiiul tissues xaiicH (ionHi(K-iulil\', Ikmiijj usually ^iicutt'st in iIih liver. 'I'he
amount is. however, elosely related to the time of feeding; and the nature
of tli»! fiKMl. In the eireuiatin;; fluids of tie' animal, on the other hand,
.here is apparently a toieralily constant peic<>tita;;e of ;jlucose, if the
temporary increase which follow.s immt-diately u)H)n the in;jesiion of foo<l
(the postpraixlial rise) lie discounted.

Here and there in the animal kin;;dom. howiver. it has Iteen assi^rted
that very little, if any. ;;lyeoi;en (certain moiluscH) or ;;lucoHe (Selachians)
is present. If this slumld prove to he th<- ease, it Would mean either
that .som« other carUjIiydrate is substituted for jjlueose and its |m)1v-
sMccharidi — such as a pentose,— or that inetaliolism proi'eeds in these
iinimals in the ab.sence of any of the lii^^her earlfohydrates. .Serutinv <if
the published re.searehes upon which the.se generaliuations depend shows
that the methods employed have l)een very unecjual in value lM)th (|ualittt-
tively and ipiantitatively. For the detection of <,'lyeo;;en tin- microeliemical
reaction with io<line has been extensively used, and for the i|uantitative
determination of this subst'iuce usually the somewhat uncertain method of
liriicke-Kiilz : it is only here and ther.' that the more certain methixl of
I'fliijjer has b.ien employed. For the detection .nd measurement (jf the
t;lucose it has been usual to utilise the reducin;; power after precipitation
of the proteins by various metlio<ls.

In consideration of these facts, it was tliuu^rht advisable to seize the
n])portunity atlbrded by several weeks' residence at the .Marine iJioloj^'ical
Station (situated on the ea.st coa.st of Vancouver Island, at Nanaimo, B.C.)
to determine by standard nietluxls the rel.'itivi- ainoutits of the ubove-
nientioned carboliydrates in selected varieties of marine animals. In
planning' such an investijjation it was recognised that it is decidedly risky

t

i

SIH

KillMini (tmi MurliMMi

-■f ..

to ftwiiiim' itmt iiiatlKNii for th" j««>li»ti'>n lunl <litiTiiiiimtion of glycogi-n
snit •"ilulih- u'lliiciti;; '•(»rlK>liy'li»t<' vilmh liavf Ummi nhown to \x' r»«liii*>l>
fur hij(ln'r lanil uminiiiiiU iitii"! u\m< ik-ii-hxiu ily Iw ho in the vnriiMl rotulitioii«
iiH't with in tlii« lowi-r iiiurirn- imimalt. Tin- t-lifinicai »truftur«> of tli*-
ciiniliitiii^' fliii<lH iiiul of thf tixNiK'H mijjlil fi>r rxanipltv !>.• no fur tliUVrfiit
AH to intttfrn- with idi- |iic)|Hr iciiiuvai of tli-- | roti-in>« prior to I'^tiiiialion
of lliu j{'y'^'*t{''" '"■ k''"'"*'' NotwilliHtiiiidin;; tliin |M«itilil<i nourcf nf f«lla('y
it wan thoujjlit timt tlir lir-t ntt-p in llif invi'««tij,'(itioii Mhoijhl ponnisl of
d<t<riiiiiintiiin ot lli- iil)«)Vf ".uluitiuu'fH l«y tin- iinvpt'd Htundiinl nii-tlioflH
n-wrviiij} for tutun rt««'uicli a nmri' .l.-tiiilfil inv«Mtij{ation of tli« caumiM
for (irviiitiun in tin- roHiiltH.

A j,'.ii.inl r«'viuw of thf wurk U'lirinjj on the diNtrihution of j{lyco(»iMi
in thf iiiiiriiiil kinji'lniii in ;{iv.'n l.y I'flujj.T (I) (up to I!t0:i) and hy
HirdiTniunn CJt (up to l!M I )

Murh ot tht! work rcfi'rn'tl to in then.' ri-vieww in niirriM-hemicftI in
niiturc. lUnI nuiy for tli.- pr.»rnt U' (iiHrei;iud«-d. Of the work in which
j{lycojj<»n Will isolutfil hy ( li.iiiiiul nLthiHls. th*' foliowinj: rrffri'iu»>»» liav.-
more or Ij-nm iH-arinj;.

AninnjjHl thu I'roto/.uH. )»lyioj{t'n hai Itei-n inolatpil liy too Hrticke
Kulz Hiithotl in Ii'*>iH"riu (Hiirfurth (3)), jMirticularly in a culture of
tJiiiUfonia Hfinti tns The iioliitid ^lyfo>ji'n jjavo the (harncterixti"'
rtwtioii with iinJine. and yifld.d a rfdi.<ii);; Huhstann- aftt-r hydrolysis
with mineral acid. The starch-likf >{ranuicn. called para^lycojjfn. which
have iH'fH dcHcrilwd in tln' prutuplasm of ccrtrtin of the InfuHoriu
((Ji „ -ina). ari- upptin-ntly not ordinary >{lyeoj{en, for altliouRh they
give a brown n-actinn with io4line. they are insoluble in cold water, and
an- not hsdrolyscd to riducinj; HU^ur by naliva. Hy prolon;;e<i hydrolysis
with weak acid, however leducinjj subntance is produced (Fiiitschli (4).
see also Maupas (.'))(.

The Mo-calltd (ilan/.ktirper, found particularly in I'elouiyxa, are also
often eonsideretl as iMMog closely relaterl to ;;lyco>;en. They are readily
formed in the <!ell when the animal i^ f<'d with {M)lysaccharideH (starch.
celluioNC), they ;;ive a reaction with iixline like that j;iven by ){lycoj{en.
and reducti')n f)ccurs after hydrolysis of the IxKlies of animals containing;
thtin. Apart from these observations, there aj)pears to l>e no chemical
evidence that the Ixxlies are really }jlyco;ien (Creeff (ti)).

Hiedermann. in sumniinjj up the work on the I'roto/.oa, states that
the digestion of starch pnK-eeds in these animals much ' the same way
as in the Meto/ou. and that stora;;e of the ditjested starch iti the form
of lAycoff'U certainly CK'curs in I'elomyxa and often in the Ciliata and
(ireijarina (loc. cit., p. :{S(I).

In Kuijleim. which represents the hijjhest of the Flajjellata, a substance
called paramyloii 1ms Ijeen extensively studied by microchemical methods,
and has been found to vary <;reatly in amount accordin;; to the pabulum
upon which tlit- Ku>;lena is jjrown. and whether or not it is as.sociated with

The tilyn^n CamIbuI of c«rUm InvertrbfAU.. «n<l Ki«h»« ai»

.•hl.,r.,i.h>ll Wh«ii chlorophyll .h ,.rr«..„t ... th- r..|l |Nir».i.> I.,,, ,n
..hu...l.4..ily formal «,„| ran I... .|iM.,lv«l out by w.*k Hikali «n.| ,,r.
.■.I,.Ut«l hy ..n.l alcohol It ^iveH „., colour with ..kIj,,.. i, i„*,|„h|^ i..
vvat«r. »..<! wh.,n hy.lroly^ul hy ^.-i.l yi.1,1, « r.-iunng M.»«t*...-.. which
r..rm...U« w.t:. y,.«,t, l»i«MU«. i« »«id „ot lo .li^w. thi. i... . -i^l. how.vrr
(Hiitni'hl. (7)).

It i<* «t*t.-.l l,v |'flu^.,r (o,,. .it.) that jfjycoKM, l.a« »„.,.„ .l«mri»»H| u
I.rH»«nt ... .■^.U.nof th., F>hi.uKl..r.„ata (HHtvrui.l. holothuri«..«, a,.,| .„

MK.IIK.-.. It h«« h...... .|«t.ct«l hy ...irr.Kh...ni<.»| .n.th-U i.i tl... ..jfij. ,,f

......■.•tH ,u„l ...nlluH,-, Am ..n,.h tt. 2 p. rt.nt. of |tlv.-o«.„. apmr-ntly

Mlt..,r,,.al with that pr..,.a.e.l fro.,. r»l,f,it liver and fm,,, oy.t,.ra, hm molateil
hy HardBii him] Vom.j; (H) fn»i„ wit^htMl |.r.Hw»i| y,n»t.

TakiMK tl... ol««rvatio..H a., a whole, it in eerUin that |H)ly«cchari.le
M.a...r,Hl, wh..-l. .. .„or.. or 1«.,h clom..ly relat.'d t-, -ly.o^en. is a .om.non

.oiiHUtUfiitof th«celUof the low«Ht ar.iu.alM a* well .h of su.-h plant-, an
yeiiMt. wh.ch .lo not 8y„theti»e .ugarM through th.- a<a.o,. of .hlorophyll

Kro... c.-rtain AHc.iri.hf a.i.i aU. from Tii-nia. Kohort (!») wan ahle to
IMvjMire a Btn.i.Kly opalH.Hc.,nt wat.'ry w.lution reactin;; with i.xlirw Ou
•.tiiiuliriK the glycojr..,, |«ca.i.t. .lijjeMte.l, alth.,„i{h the extract did not cauM
d.««Nt..m ..f ad.led starch. Thin would s«en, to ,»,i.,t to the exigence of a
spciHc jflycoKciaHe \V..i„|a...| (10) fouu.l that alx)Ut otie-third of the
weight of drie,! Asi-ariH consisted of glyco^f.-n , in 'r...nia even a higher
-iuiount. namely, one-half of the dry weight, .•onmste.l of thia material. In
Dmton.a hepaticun. alao a laige amount of glycogen was prenent It is
interesting to t.ote in passing that this glycogen was fou.,d -luickly to dis
app.-«r when th.- worn.a w.-n; kept under anaerobic conditions. (Jlycogin
IS also present in considerable amount in the earthworm (Lesser (IDi
In the Hocalled chlonjgoger. cells which surroun.i the alimentary canal
very large amounts of glycogen have been descril^d: the leucocytes in
the body Hu.ds also eonUin it. but it is said to be absent from the muscles
(Cuenot (12)).

A considerable aniount of attention has been paid to the carlwhydrate
present m the digestive gland (liver) of various Molluica Biedermann
k'lves an ade.juate review of the work. Of particular imporUnce in the
present connexi.m is the work of Barfurth (cf. p. :U2. op. cit), who found
m a.r-breathing gastropods after feeding for one to three days with bread
from 3 4 to (5 4 per cent, in the liver and -.Hi per cent, in the f.wt He
concludes that the liver of gast.ojKxIs plays almost as imporUnt a part
m storing glycogen as does that of mammals.

In those snails (Helix) in which there is relatively a large amount of
connective tissue, the glycogen was found to be stored mainly in the
plasma cells that are present in this tis.sne, whereas in snails (Lin.ax) in
winch tli.iv IS little connective tissn.- the plasma cells .sckh. l)ecoiiu. Hlled
an.i the glycogen is present mainly in tlie columnar epithelial c.-lls Iwth
"t the intestine and of the biliary passages (Barfurth, op. cit i> :i->H)

VOL. XII., NO. 4.-1920. r B , y .i_,^p. j.n,.

sto

KiliMtrr. aihI MtM'txiil

Tht-Mf olMi-rvittiuiix HIP of ifiiciMiJ Ml r.niiii>«iiMi with III- .>li<t«'r%«« llrM

m*<l>- *>) <'l.iin|p Hrriinr-I mil »iil»w'i|urntly •' iitlriii.'.M.y llttifiutli ,u,.|
Hii •l>-iiii nil. l'> ••»'• <tl»'t tlii»t tli» r<Hiffiit"« •'{ til'- itoitiiM'h "tii.l u|>j» I

pirtiiMi of llip int. -.tiiKv ulu-r i\u<y havp U-pii |wrli»ll>' •lij,'.-^«'-<l w>\

•ml -ml ..i' itir lisr |Mi«H ),'.'» .\li»«ir}»ii<m .>.-rur» IIiikmi^Ii Hi*- "II'' l><>tl,
til ill'- iiiif»liii<' mnl liiltiiry |hi»x.i^i"' uikI wlnti it i» <oMi|il«<tiH| ihr ilml
left ill lh<' >»t<»iii«('li i» roluiirl'-'t lli'iii;,'li »t Hr-tt it wi»» ••oLiitriMJ hy th.
•mmTiUkii "if tli«> liviT It m iiii|Miitatit ti> iKttH tli^of fwti ill ftmiM'^i'Mi
Willi tli« [(r'-'-iil iii\i«tli;{'ili'ili l>.<nii»i. tiny in<li<)ltf lliut w- liiitnl \»

f«rrlful not t imIii'I- thut mi) i(ly<oj;fh I'oiiimI in tin- liviT of iii'illu'" •

ix niii'x'Mirilv |»r«'««'iil in tlii>« or;;i»ii iit •^ucli. At n iimll«T of li»'"t wvii,.!
inv.«««ti«i4lor«. (Ki.-nt>!i'l. Koliiniuiii ilU), Hott»!'.*i ( U)> Iwvi. U<i-n iuii»l-l'
to <li"lirt iiiiv nly>"i{.ii .itli.r l>y ii(i<riH'hiiiu<-n! or tiii«'liiiiiieiil iM>'tli<M|ii iii
thi* livriH of Aplytiii mill Anon Tin' two olwrvi'M Imtt iiniilioniil liiiv-
intlt'iKiHlfUtly foiin.l in Vplyxi* tth«' «« liar«». in plwf of jjlyroKi-n >i
p«-nfo»>-vi''l'linu ••iilmtmiif ( ixiitiwMHi). tli.' prociw i<l«Mitity of wliirli i-.
uiKvrtkiii KolimiMin tliounlil tliit |«'iitiwan to \m iltriviHl from nn
i|i|;i')«ti-<l ffjiiilui' of till' fo-xl of till' Kiiiiiinl, wlmli coti-iHt'. of mi Alj^a il'lv i
Uctticai rttnl wliii'h contikint |»'ntoi»mi ait wi-ll itn xtmcli Tlif n-.tiilii.'
tiiuU \t> way into llu' biliary iHimaK'"* *" <l'"«'rilx*<l iilwvi-. Hott«/./i
%ifri'»'H tliat it in ilerixiil from llic pi-ntonan of JMvii liilt iN'lifVi-i that it
haa Iwc-oiuf partly lir. ki-n up •*.. nt to form mi mul »ul)Ht«ii<e (a<i<i.'
pflntt>Nti|U)>) which talcfH tli.' plm-f of luiinnil aciil. which in ahieiit m
them' niolluHCH.

\I. H.'ii/,*' (lo) <'oul<l not timl a trace of jflycojji-n or of waler-iwliihl.'
carlnihyilrate in lh«' inuwh'" or liv«-r of <Vto|HHl!*. Some iviilt-ncK wnt
ohtain.'il, howmer, that a jjliicoprotiiii may ivpivwnt thf larlHihydrut.-
atura;;e mati-rial in tlit-se aniniaU.

Further attention Iuim Imimi puiil hy lli'ii/.f mnl MirkHiittcin (hi)
to till- Hup|K(Hfil alxsmt'f of ;,'ly<"o^i»ii in tli.' li%»!rof moilum-!*. 'lln-y |«>in'
out that if this foni'lusion in i-ontiruifii it >voiiiil inilifate that, in so far »i
thi-ir ciirliohvilnite mctalHjIisni in coiii'irn. i|. ■i-rtain nmriiiH moliuHcs oocup\
a pi-culiar |)o.sitioii in tlu' animal >cali', ;.'lv<'ojCfn Im iiij; apiHirmtly prcHent
in all othtTH. Thi'y show that a ofrtaiii ♦■rror may l»' inciirroil in isti
mating jjlycoj;<>n in the oijjans ot H.a aniniuis on utcount of the preHcme
of the Halts of soil wat.;r partic-iilariy M«. Tin- .Mt,'(<)ll». whiih h
formed with Koll ailsorhs som« of the j;lyL'oj;fii. ami wln-n alcohol ih
subseiini'iitiv atlilfil a pr-eipitut-' is formi<i from whuli Koiliii^' wati-r il.« s
not dissolve out all tiie ^^lyeo^'.ii Tiiest; authors also jioint oi.' that un
error of anotli<r iiaturi' may Im- iiicMincd l)coausf of tiie pr-seiicf of ;;hicn
samin derivatives and of pentoHans, unless the tissue lie treateil with tln'
stronjj alkali for a -urtieient lenjjth of time to ensure the entire destrurtion
of these suhstaiiees. 'I'lii^ matter will he r. fi-ned to at;aiii later. It is
reeommended hy .StarkenstiMii and Heiize that the atjove-mentioned
Hources of inaccuracy be eireumveiiled by prolonijed heatinj; with KOH

• don «<l<iir)<^ iirily i>iiK volimm HMC'tnl ni Im<i of Mlri>li»l to ttrtnipiUt*
lli«- i{lyi'.>i;.fi nii>l lliiiilly liyi|ii.ly«iiii{ th« rthohol |irif-i|>i(iitii liint'lly
witliiMil tif'.t ■>' nil .|ivH.lviiii{ It III w.»I.T 'rti«> o»rtr li<> ••x|h rim.-iiUl

|tr.Hif ltitwi>v*-r lliiit tln'ir iiiM(h<Hl i.< iiny iMi>r>> (••■••itntl.- ili.-m ti ri;;iiiat

iiitfth<H| of I'fliii,'. r I'll, y f'MiiKl .11 iiiui'li nn I 2^ jjriM. i^lynij^. II n, fUv
livfr it( A|ity«iit Irtipxitiiii

Amoii); lliK t'riHtaiH'it it witx foiiiiii In li<)|)|i>- Si-y|«r (17) tlmt ihrr*
WM -(tiii luiiiM' j{lvct>i»^ii III ill.' \\\,.r of •» «i yinrul.l finvti^d (Khfik
r.liiiii) aft.i altttviilioii CUinJ.' Ki'itutKl liint |(i.vi..imly foiiiMl the
aiiioitnt to vary iiMirK or N-m in rp|ittioii>thi{i t.> tlif tim« of iii.iiiltiii;j. lM>iiit{
;,'r'rtl<'xf jiiJit j.rior to ilii!« |»'ri>xi At tlm tiiiii of iiioiiltiii^' Kirwi-li ( |H)

t'liiiil ll eilirf Unly loli tli fn-Hli wUii iriiytlKli) to iMiitmii It H2 jmt niit ,

whrn-an four iiioiitln Ufoiv llii^ |M.ri>i<l it runittiiiiMj only 0(W |»t t*iit.

Until ri'i-eiitly very little iiiforiiiiitioti rxiittMl rotir«riiiny thp |iri>m>iir«
of j{lyro}j«>ri ill llif HnIii'k 'rimt tiiiii' iit U'unt \n pr- -(I'lil in tlu' tmmi.ncif
Miarih.' tisli linH Iw.ti dliowii liy (I |trriii(r<l I'livy Miinkf ami otliwrx.
it vm% ittftt'il by lii-riianl that tliit ;,'lyi'oji»Mi in unusually rfsintanl to the
iiit1u«>tici! of ixmt-niortcni i'lian;{i''». «nii tliiit it (lo*-* not ri»<lily iii-<a[ip-«r
■liirin;; liuiijjAr Diiriin; u.»|iliyxin, Iiow«-v*t ||i>- glypoyt-n rapiiUy
ili.ia|i[H'iirH.

Schoiidorff nnil Warliliohl.ir (l!l| (nl«ti|uuiely review all tlie»>'
■ i|il«»r inv<'»tij{utio.i>t. ittnl coiitriliutf nunitroiiM ot>>u'rviition» of tln>ir own
III whifli till* ftiiiount of y:lypo;,'i'M, iJi't<'riiiin<'<l by riliii,'.»rn iik lli.xl. in n
i.ir^p varit'ty of ti^lif-. <iiu;;lit in fiv^h \vHt«T in ;;iv<'n. 'I'll'- |Mrit'iitiij{«
iiiiunint v(irit'<l Ix'twofii 2 .'i iiiul I2!U. I'lolonjjml liiinj{»T was* found to
r.iuMj roiisiderublt? reduction in tin- aiitount of jt'v'o^fi'n iu hih-Ii tixli m
til'' pikt« (Istox lufiiiN), wliii'li renmin iictivt- hut to causn only NJiylit
r.diiKion in tin- carp (("yprinus i«rpi«t) and other ti«li wliidi hide
thi'iiiselvtH away in the mud durin;; winter. I'oHt-uinrtem jflycojjenolyMM
did not appi'ar to prix-ied as rapidly a« in Maniiiialia Th.- •>Htiiiiiition»
were iiijuh- hy iii.mimiii iii^ Ixjth t lit- i-.-diiciii;; and fhf rolatiiij; {»<>wer of
tin- hydrulyHi'd },'ly<'o;;eii prt-cipitat-H. the cIohc corrt'HjMinih'nre of the
i-'MultM olitum.'d hy the two ni.thixN indicatiii;; that tli<' niat>'rial is
ilunnically identical with that present in nianiinaliiiii tissues.

Mktiuids.

(;iycoj;..n.

The orjjaii or lissiU' was cut into hihuU pieces, pressi-d lietween

tiltiT paper. wci;;hei|, and dropped into !>.") per cent, ali'oliol, in wliii-h

It was shipped frcjin the station to 'I'oronto, the journey 'M-eiipyiii;; aljout

.1 week.

It is pi)-<>iilili' in the rjisc of oil'- or two specimens ut niiiscle (e.jj. siphon
•iHiscle of elaiiu that tle-r.' wis rel.itiveiy tmt small a .piaiitity of alcohol
"iitirely to prevent a oitain uiiiounl of ^^lycojjenolysis, 'I'he inu.scle in

II

. .^..M^kT^l'l^j^JjT'M^M'^.L f :i.*

822

KillHirii and MhcIoo'I

tlii'so cAHflH wii." (It'cidtKlly coinpart, hiuI may not have Item sufficiently cut
up to PUHure inniu-diiitc penetration hy the alculiol. I'hiH |)OHNil)lt> Houroe
of error wa.s, however, not incurred in the vaHt majority of caNcH. The
alcohol-preserved material wa.s pressed bef>'"en tiller paper and again
weij;hed. It was then lieateil for three hoi with ♦!() percent. KOH and
th.! ^;lyc<)j;en determined hy l'llilj;er's methiHl.

An already uieiitioni'd. .Starkenstein and Heii/.e state that j;lycoj;en
determination hy the I'Hii^er nietluMl is inaccurate in the prenence of
«ea water Ix-ciiuse of adsorption of the }jlycuj;en hy the Mjj(OH)^ and
Fe(()H)3 that are formed when KOll is added. This adsorU'd ^jlycojjen
after precipitation with alcohol does nut iK'conie completely dissolved in
boiling water.

Hy dissolving; ei|ual amounts of |)ure glynijen in sea water and in
distilled water aiici then cirrying out the usual f'lluger procesa with both,
we have contirnied tlu- observations of the abt)ve authors. We do not
l)el'eve, however, that the organs and tissues of si a animals contain a
Rutficiency of salts preci|iital)le by KOll to make »ny signiticant error in
the glycogen determinatioiLs. This conclusion is Itased on the following
observations : —

The liver o' it rabl)it was cut in snuill pieces and ijuantities of 20 grin,
each were added to eipial voliunes of (a) sea water, (b) 0"!> per cent, sodium
chloride solution, in which they were thoroughly ground in a mortar. To
each of the resulting suspensions ei|ual volumes of 60 per cent. KOll
solution were added and the gly<'ogen content deterniined by the PHiiger
process. The following results were obtained : in (a) 067r>, (b) 0784 per
cent. Kven in the presence of a very large excess of sea water — very
much more than could l)o present even in tissue which has not been
pressed lH>tween Hlter paper — the error incurred is not excessive.

In order to see whether any large yield of glycogen would be obtained
when the alcohol precipitate was directly hydrolysed, 2S grm. of oy.ster
(from which the excess of sea water had been removed by pressing
between Hlter papers) was treated with KOH and the glycogen precipitated
with alcohol and divided into two portions, >i and h. The precipitate in
(t was dissolved in l)oiling water in the usual manner (i.e. on the filter
pajHjr, and the filtrate hydroly.sed), whereas that i:i '< was removed from
the filter to a flask by a fine stream of water and directly hydrolyse<l.
The following results were obtained : in a. 0"5(i0 grm. glucose, in h, •()})4
grin, glucose. There is therefore evidence that it is necessary to mo<lify
the I'fliiger ])roce«.s to the extent that the glycogen precipitates are
washed from the filter paper into a flask and directly hydrolysed. This
procedure has not been followed in the present investigation, partly
because Starkenstein and Henze's work did not come to our notice
until the work was nearly completed, and partly because the precipitates
in many cases weri' of such a nature that their removal from the filter

:»^._^.!^ W *i :2tJlr

-'■ ..-J jyr,

Tl... (

ily<'()i»eii roiiti'iit .if i-.Ttniii Invrrti'hriiti's ami Fislu's :j-2:j

papi'i- woiiM liiivf Ix'.'ii |iriicticiilly iiiiiin.Hsililr. It is (•unsii|,.r.Ml ilmt tl

le

■rnir iiiiMirr<'(

II this lU'codiit is iiiHiiriiilit'

tiiiniiiciiiil, rs|(i'ciiill_v siiire very littli'
of tin- Miilts of st-ii wiiliT could liiivo Ih'.'ii |ii(-'Si'iit ill t'.>tiH.su.( used for
till' e.stiiiiutioiiN.

Hksii.ts.

( Jly co^jcii

Tln'si> im- given in ptTcciitujjfM liotli of flip original hiiiI tlip iilcohol-
tliii'd raatciiiil. Althoui;li iioitlur result in itself is inoie tliiiii upproxi-
iiiiite because of the iinpns.-iijility of ri'inoviiig all tlie aillierent moisture
or alcohol by nieiins of tilter paper, they are siitliciently closo to he u.sed
for the purjHJse in view, namely, to determine the relative ami>unt8 of
jflycojjen in the tissues of ditlerent animals.

The observations were made on speeimens f oiii the following itouds
01 animals: —

The Kchinoderinata ( Asteroidea).

The Mollusca (I.ainellibranchiata).

The Arthropoda (Crustacea).

Tlii Fishes (Elasmobraiu-hii and Teleo.stomi).

I. THE ASTKUOIDKA.

TaBI.K I.— 'PrKCBNTAOE ok (ilAlOliK.N (AS MkXTKOBK) IN THK HkPATIC Cwa

OK Stakkishes.

( ilyoof;en i)ercoiit)ige.H

Weight

No.

Species.

of
material
(gnu.).

M

Pisaster oelira«r

41!)

•r

,1 II

14-2

*:>

Iti'ti

H.

I'ixanter brevixpinus

H

L

PycnoinKha lieliantoiilcs .

34.1

1,

„ ,)

24 ti

s

Kvaatariii.M Tnwchelli

11 '9o

t'l

Liiidia fuliata .

•>•>

«.

'♦»».-•

18

Calculatcil

for original

material.

1-2.T

Oli2

Calculated

fur aU'ohol-

pres«'rve<l

material.

1-52
0268
0-47H
0-232

UD3
O-tiO
0-90
0-4(il
0403

Kemark*.

1 large .s|)ecimen.

2 liiiiall sjK'cimens.

3 ., It

1 8()ecimen ; liver

very small.
From 1 apeciiiieD.

Mud .'(tar-lisli.

'\

The variable amount of glycogen found in these aninialH is probably
dependent upon whether or not they have recently been feeding, the per-
centage of glycogen being plainly proportional to the size of tlie caeca.
The glycogen precipitate in M was further examined. After redissolving
in lioiling water it wa.s found to give a characteristic reaction with iodine,
and on hydrolysis to yield a sugar which fermented readily with yeast.

f^...^:

•♦^HIBhii^i'-* *

884

KilU>rn and Macleixi

ri. THE M(»l,I-l'SCA.

TaIILK II. PERCENTAliK OK (i[.Y('lMIEN IN VaBIOIR I'aRTH OK THE lioflY OK THE
IIuHSE Cl.AM (Scill/llTHOtlllH NlTTM.I.I).

tllvro^'en perrentafji's

. No. OrgHii or tiwiue.

A Dif^eNtive glan<l
*' 1* ^»

F MuFcle of siphon

Q i .,

C j Musde of foot

Pi,.

B I Adductor inu.iclca

O i

R 1 Posterior adductor

Weight of

iiintcriul Calculuted ] Culculaterl for

(grill.). for oriniiiiil alcohol-pn-servcd

iiiHtrnal, material.

31-2

225

.'.6 1

30

54-2.-.

19 4

ear.

20-«

171

(a) 0-31

(b) 1-56"
(a) 0077
(1.) (1-982

(a) 0-46

(b) 1-70

(a) 0-40

(b) 2-67 I
(b)2-76?

* Hydrolysed glycdgeii fermented with yeast.

The estimations niiirkid (a) wire in ule <>ii eUms that had lieeii kept fur some time ( 1 -'.i weeks)
in a ^ack imiueised in the sea at the wharf Those marked (h) were made on clams that had been
kept only a day or so after diRjjinf; them up.

Tlie.se ob.servati')n.s wt'ie carried out on two batelie.s of clams. Att:r
being dug from a suiidy beach at low tide the clani.s were placed in a sack
which was then immersed in the sea at the end of the wharf, .specinii-iis
being removed from time to time for analysis. The clam.s of the first
batch a, although leceiving no food, remained alive in the sack for over
two weeks, but. those of batch '- did not survive in the .sack for more tluiii
a few days, and they di.siiitegrated very rapidly after death. The e.stinm-
tions, the results of which are recorded above, of group n were made after
one week in the sack ; those of group //, on the other hand, were made
within a day or two after the clams were collected. It is po.ssible that
variations in the .state of the material accounts for tlie persistently higlier
percentages of glycogen in group /< than in group d. In order to make
a comparison of the glycogen in the various organs and tissues, it is neces-
sary, therefore, to take the results of each batch, a or h, separately.

In batch <i the largest amount of glycogen was found in the muscle of
the foot, with very nearly the .same amount in the adductor muscles.
Decidedly less was present in the liver, and only a trace in the muscle of
the .siphon. More interesting resuli.-i are those of batch '*. In this case
the adductor muscles contained a liigher percentage of glycogen than we
can find recorded for muscle. (The highest given by Pfliiger in "Das
Glykogen" is 2-44 percent This was found by Aldehoff in the glutjeus
maximus of the horse.) We are, however, not certain that it is typical
glycogen that is responsible for all the high percentage found in the case
of the adductor muscles. There are several reasons for this doubt : first.

» ^■l-»:..*>4ili-*W'

.JC'^iWiU.'

JilA'.

S">''flBiS?""n|*'j (t ^z

Tilt' Olyidgen ('ontent of certain Invertebrates and F'ishes 32.'5

thi- procipitiitt! prixlucfd by alcoliol did not m-ttle hh urdinarily, hut re-
i|uired the a<ldition of a conHiili-raijIe amount of scMJiuni chloride to cause
it to do ho; second, on noiitraiisiiij; the hydrolysed solution of thf pri'
cipitatc and fi-rmcntin;,' with yt-ast only u Ninall (pmntity of ;;ii!s collected,
whereas the precipitates from the liver, when similarly treated, jjive a
much larger amount of ;,'as. VVe had not sufficient material with which
to investi;;ate this (|ueHtion further. Tln-se muscles are entirely ditFerent
in structure from the muscle of the foot or siphon, heiti}^ com{>ose<l of
larjje bundh's of very pale substance.

There is also a hi<,'h percentaj;e of ;;lycoi;eii iti the inuscle of the f<K)t,
although this muscle can be used but seldom alter the clam liiis assumed
its more or less porman<'nt position in the sand. The glycogen in the
siphon mu.sde is decidedly less (0!tr> per cent.); this muscle is used to
hold the tul)es open as well as to retract the siphon on the approach
of danger.

It will be observed that a considerable amount of glycogen was found
in the liver. This is of interest because i,\' tlie belief already referred
to (p. 320), that there is no glycogen in the liver of certain molluscs
(Aplysia and Arion). That the material found by us was glycogen, as
ordinarily understood, was shown by its general behaviour towards strong
alkali, iodine, etc.. and by the fact that the hydrolysed glycogen readily
fermented with yeast and gave typical glycosazone crystals.

It will be recalled that several observers (Frentzel, K(dimann,
Hotta/zi) have averred that tliere is no glycogen in the liver of certain
other species of the .Mollusca. These workers found evidence of the
presence of pentoses and niethylpentcses, and they ha»e suggested that
[)olymerised forms of these may replace glycogen. Since they did not
consider the likelihood that the pento.ses were derived from inosinic acid
or guanylic acid, the conclusions cannot be given much weight. As a
matter of fact, it has been shown by Mr i'.. K. Berkeley, workin" in the
biological station that the pentose present in the tissues of closely related
species is derived from one or other of the above-mentioned nucleotids.

The percentage of gljcogen is notably different in the small crab
(Cancer productu.s) and in the lobster. The difference is probably
dependent upon feeding conditions. The lobsters were shipped from
.St Andrews, on the New Brunswick coast, to Toronto in moist seaweed,
tlie first batch in September, when the weather was warm, and the second
liatcli in November, when it was very cold. The difference in temperature
does not appear to influence the results. The larger crabs were caught
at St Andrews in \oveml)er and transported to Toronto. They were
almost dead when received. Although they contain much less glycogen
in proportion than the smaller crabs, it will be noted that there is decidedly
more in the liver and in mu.scle than in those tissues in the loKster.

A comparison of the glycogen content of different muscles in the lobster
is of interest. In the muscles of the back and tail the average for four

.-■"If

i

■■^t

;!

.■''J^>^^&:±^.:.

326

Killtorii uiid Macleod

5^,.,?

111. TIIK UMMKOI'ODA.

TaIILK 111 l'KR.K.NT,\.,K „t < JlV. .MiK.V IN VaHI.M m PaKM "t TIIK M..1.V OK
I UIHTACKA (CaMKR I'HODUCTIJK AND lloMAHl .S AMKRI. AM H).

No. I Sj>ecie

Oigitii or ti-i»ii«.

D
E

Cancer prtxiiii'tiin Liver

MuDclen

p. 34 Homaruii .
„38

„ 40
.. 41
.. 44

., 48

I (p. 3) Humane I.
(p. 4)

Homarus II.

j

Honiai'u* III. .
i Cancer irrotatiis

Liver

I. Heart

II. „

1. Muacle (tail)
„ (claw)
II. Mu«le (claw)

n. „ (t« 1)

i an (of >\x lob-
sters) t
Liver

Muscle (liai'k)
.> (claw)

Liver

Muscle (red claw)

,, (white claw)

„ (back)

Muscle (liack)

Heart J
Liver

Weight of

material
(g"").

7 37<

UK,

.•JfttU
2 37
194

ftft

Si)

4ri7
4S'fi

7'32

20
20

20
20
20
20

20

277
24-47

()lycof{ei> ,«rcentageii

<'.l.ulHte.l /''»'"''*l«<|
fro.u oriK.nal '""""'"^"h'.l
n.ateriil. I'r«»«"e.l

material.

139

0-.J7
0M7

0-7H
OUI
142
036
<I17
OKI
0-3!

0M5

OO.''.
0-31
Trace

0 13
017

017 i

"•30 I

0-30

O-ftl
100

• Weight after preservation in alcohol.

t The hearU of six 1 il-ters were used for this determination. The weight of the animals and
of the heart m each spemmen was as follows: (l)2000Brm. and IS.'i (,rm , (2) 1660 urni and
rSS rm., (3) 1150 grra. and l-QOgmi., (4) 1150 grm. and lO'J grm. (5) 1100 gnu. and 1 -00 grm
(8) heart I'OOgrm. b •

X The hearts of five specimens were used.

loKsters was 032 per cent., being remarkably close to this figure for all of
the animals. The pale muscle of the claw gave an average for three
lobsters of about 015, only a trace being found in that of a fourth animal.
There is therefore decidedly more glycogen in the tail muscle. Of greater
interest are the results obtained for the heart. Being very small (weighinj;
only 1-95 grm. in a lob.ster weighing 200 grm.), it wa.s necessary to collect
the hearts of t ?ral animals to make the analysis with any precision. The
average of thri determinations on different collections of material wa.s
106 per cent., the lowe.st value being 085 per cent. The relatively high

,,^ <:*%

The (ilyp.);,'.'!! C.tnt.iit ..f cTtnin Invertehrat.s aii<l Fi^h.'s :i27

perceiitii^'f of ){lyc.i;r,.,i in this piimitiv.. f„n„ „f l.-art i-* iiit.'it-stii.;: in
view of the fact that u similar r.-siilt was ohUin..! in th.' h.'.irt of fi."h.-M.
Further refeienc to the si<;niHcuiire of tliin ..I.H..rvati..ri will t«; f„un.l in

ronnexion with the lattt'i

WitI
(■i inside
tlie muse

th regard to the liver, the poreenta^e amounts of ;;lv<-ojr..n varied
srably ( viz. lK.'tween 005 and 0-7« per rent), thus eontrastinK with
iscles, where the amount-* were tolerably constant. Xo doubt" hh in
mammals, the glycogen eontent of tliis vispus cjependi priniarilv on the
activity of digeHti(.n. which it i.s presumed must have varied forditferfut
individuals. Tlie largest result was obtained in a specimen caught in
September.

IV. THK FISH EH.
Table IV. -Pkkck.ntvuk ok (ii.vecxiKx in Varioch Paiits or thk Body ok

FiSHKS CAUIIHT DUBI»<i JfLV .VXD AUliUHT.

i

( ilyc'ugeu percentaKes

Weight of
material
, (grm.).

No.

•S|iecies.

< >igan ur ti. -ue.

Calculated
imiiiuriKinal

Calculated
from alcohol-

i

materi'il.

preserved

1

material.

H

Klasniobraiicliii (Sijii*lu<
Sucklii), dog-tish

Liver

44-2

1

O-ttST

(»0«I9

I

ti

,,

1 20

0-16

0-209

V

..

..♦

198

Xoue

None

K

Muscle (body

wall)

50

0018

0-025

W

Miucle*

IK'S

None

None

(t

Heart

, 7-3

0-447

0847

IT

•

44

...

0172

Z

Chimaira (rat-flith) .

Liver t

16-7

None

A,

162

..

Teleoatiinii (( 'ypritius

Mu.scle

10-3

carpio), rarp

p. 9'i

(a) Liver

344

(a) Trace

96

(a) Muscle

50

(a) ,.

99

CO „

50

(b) 0-021

(1)) 0 028

23

Liver

10

(i-60

24

MiLscle

■20

0-2W

25

Liver

10

5-60

26

Muscle I

20

Trace

98

Christivoiiier Numiycush
(lake trout)

(a) Liver

26-6

None

98

(b) „

45

0-055

97'

(a*Muscle

1391

Trace

97

(b) „

U-70 1

"

* Dnglish caught in nets and kept in .small Unk for some days. Otherwise the dog-fish
freshly caught by line from the end of the wharf.

t Dead Kome time. * Fish dead at Iea>t -24 hours.

".'

328

Kil)H)rii iiikI Mac'li'txl

• r.K

I

Tilt' oiitMtHhilini; iVatun- of ihomI of tin'**" rcHnlts js tlit- iilimMic' <>r t}n>
•<inttll niiioiiiit of K'.vojffii. The or;;an« wcri' reiiiovcd for clicinical investi
jtntion imni('iliat»'ly after the MmIi wt-n- ciitiylit ; cxn-pt in the ciihp of
rirtniii of the clojj-tisli niid thi' nit-tixli. in wliii-li tin- tl>tli. rntrniiiMil in intH
weru brought to tht-Ntntion in n lmlf<lin<l condition in the tank of the UhiI.
Thi" rc'Hults Ntftml nut in sharp pontraNt with thone of Sdiondorff ami
Wachholdir ainady rt-fcrri'd to, in which the iiwr and niuwlcs of a hui;;
HhI of frcMh-watcr ( Khiiu') ti.Hhfs show in j;t'n«Tal as hi<{h perrentaj^ns of
;.'lyco;,'i'n as havf lieen found in any animal Tin- oidy fundauif-nlal
ditfirenco iM'twcun tin- two strics of obMer\ationH in with roj^ard to tlif
time of year at which the Hsh were caujfht. The ditft-rcncc cannot At all
dcjj«!nd on whether the tish wa.i cauj^dit in fresh or in sea water, since in
our «eries tish from lK)th were examined. The carp numlHjred 21-2ti
inclusive were olitained in N'ovemher from a local deulei'. Two were ali\e
when brought to the lalwralory. It will l.e noted that decidedly more
glycojjen was found than in the MsheH examined in August (Nos. it5-(t!tt.
Although not ponclusive. this supports the view that the ylycoj;en content
of tishes is very low in the Nuninier months and high in winter.

The very low glycogen content of tish caught in Humincr, as compared
with the decidedly high percentage in winter, i.s in conformity with
similar observations in Amphibia and hibernating inammals. Thus
Athanasiu (op. cit.t found in the liver of Tusca 277 per cent, and
4."J5 per ci-nt. in duly, and S-21 per cent, and 7.') 2 per cent, in October
and Xovember. Lesser (20) and Hleibtreu (21) have also demon-
strated a great decrease of glycogen in tlie liver and other tissues of frog-
during the spring and sinnnier ni' s (accompaniefi by a steady increase
ill the ovaries) and its great abundance in the late autumn and early
'•■inter.

One other fact deserves attention in connexion with these results,
namely, the relatively high percentage of glycogen in the heart of the dog-
fish. In hearts from two recently caught dog Jish 0-447 per cent, (of fresh
weight) of glycogen was found, when the livers oi ihesame tishes contained
only 00.")" per cent, and OIti per cent., and the muscles of tlie l)ody wall
only 00 1 H per cent. In hearts from other dog-fishes that had been caught
for some time ami were partially asphyxiated, 0172 per cent, (of alcohol-
dried tissue) givcogen was foinid when no trace could be detected in the
liver or muscles.

The presence of a large amount of glycogen in heart muscle has been
known for some time(cf. I'fliiger, loc. c it.), and has recently been confirmed
by Cruickshank (2:J), who found in the heart of the dog from 0:)00-
0(i;jl per cent. It has furtlier been shown by histo-chemical methods that
the conducting tissues (the A-V node and A-V bundle) are especially rich
in glycogen, a fact which is of particidar interest in tiie light of our
observation that there is a /ery high percentage in the primitive heart
of the dog-fish, and al.so in that of the lobster. We were unfortunate

I

r

iiii :

Till' (!lycu)(eii Content «>|' ('••rtnin Iiivi«rtehr(»t«'H and Fi^lifH '.\i\)

111 f»iliii;j to Ht'cure more iiintfrial ujioii whii-h to follow up tliis iiitert-tttiH;;

i>l«ervation,

L'(»V(Lf,srnN,s.

|{y Pflilj^cr'n iiD'tiiixl the folldwiiii; |»!rceiitiip' niiiuuiitx of jjlycu^'fii
wero fuuiiii present in tin- ili;;4'><livi' ;;liiiid (lic|iatii paiu'iens) nf rfpn.'-
Hentative npecifH from various ai|iiiilir phyla: Afitoroiijiii, 0 2H2 to T'l'i ;
ijiinflliliriiULliiiUii. OH! to ISO, (,'ruNtacfa. OO.'j to l.tit. KluHiiioliraiicliii,
iiono to 02 1 ; 'I'l'leostoiiii, iioiif to tio.

The varying aiiiountN arc appitrt-iitly (iepciKloiit partly on fuolin^
cdiiditionH and partly on thf seuMoii.

In the niUHclcM the followin;; percentajji' anioiintH were found: -
l^inellihranchiata. ()'()77 to 2'ti7 (the latter value wan ol)taiiied in tlie
adductor iiiuscIoh); CVuHtacca, truce to O'Uti ; Kla.HmoliruiK-hii. none to
OOIH; TelfOHtomi, none to 0 2!».

In all casi's where it was posNihlc to Hfcure ii 8>itti('ient uniotiiit of heart
muscle the jjlycojjen content was fctund to he several times ;;reater than
that of the other muscles, and sometimes ;;reater than that of the liver;
thus in the l<jlj8ter from OH") to 1'42 |»er cent, was founil in the heart,
compared with a maximum of ();<l) per cent, in the muscles.

In several cases the jjlycojjen was found to yield a yeast-fermentable
su^jar after hydrolysis, hut in others there is some evidence that n certain
proportion of the " reducinij material " was due to other suhstances.

LITKKAi'UHK KEKKUKKD Id.

(1) rn.CiiBR, E. F. \V., "l>as (ilykogeii," Arch. f. .1. ^es. I'hysiol., 1903,
scvi. 1.

(2) iSiRUERMANN, W., ill W i II te IS t e i ri 's HiU!ill)iicli iler vei'^'leiclieiidHii Physii)!.,
ii. 1 Halfte, 1910-1'Jll.

(3) Uarkuktii, " Vergleicli. Iii.stiiilicmisrhe Uiitersuihunj;eii uhenlagCHykiiHe'ii
Arch. f. laikr. Aimt., I8f<6. x\v. 259.

(4) lii'TsciIi.l, ()., " Bciiu'rkuii^'cii ulwr eiin-n der (flykiij,'('ii verwandteii Kciriier
ill (leii Gregar'iien," Zeitschr. f. Hiol., 18(S5, .x.\i. 611.

(5) Maupas, K., "Siir le >;lycogeiie rhez les Iiifusoires cilies," Compt. lleiui.
Acad, des .Sciences, l8cS5. t. 101, 15(i4.

(6) UiiEKFK, I!., " I'fliiMiy.'ca pahi.<tiis, eiii aiiiiileiiarti^'ei Orgaiiisniiis di'.s
Suszwasser.*!," Arch. f. liiikr. Anal., liS74, x. .'il.

(7) HCtsciii-I, O., " lieitriii^e zur Ki-untnis des Parainyloii.s," Arcli. f. Piotisten-
kunde, 1906, vii. 197.

(8) Hauubn, a., and W.J. Young, "(ilycogeii from Yeast.' Trans. Clieiii. Soc.,

1902, l-K-xxi. 1224.

(9) KouKKi, K., " I'ber die Kiiiyiue wiilielloser Ticre," Arch. f. d. i;es. I'liysinl.,

1903, xcix. 174.

ill

:mt Til.- (ii

\r<H{.ii CotiU'tit of certain lnv<'rt«>hriit«'M itrul KiMlien

(III) Wkimi.am., •• n*r .|.Mi «)lvk..vTiiuM'l..ilt

Zuimclir. f. Hi..|., 1901, «li H9 , uU.. |»0'.'. xl

iiKi'ii.ilt piniKtr lumitiiichcr Wuriuei

III. .'.«.

Ml) I.K-KH, K. I.. "(I

l»0!>.

ifliiNi In- l'r"we"»« l..i Kn>{>-iiwiirtiii>rii," /•>it««lir. f. Bn

(I'.') Cl'r>'i L, " Ktu.leii |itiy><ii>|<>f{tr)iiv« Mir leu (UiKiMhiuti'H, ' ,Vrt!li. lU llml

IM»«, XV. Cf. iiiH.l^r

inunn, ii|i. cit.

(13) K'XIMamn, R, •• KiiiiK'* l!.'nl«ii|iiiiiim.|i iil».r(lif Vfriliiuiiii« ■l«r Knliinliyilrnt.

Iwi A|.lyniiMi," (Viitrull.j. f. I'lijiinl., IS'J'J, xiii. 465

(14) UoTUWi, K, "<'oMtril)iitiiiii« .1 III |.liy»ioli>Kif c.
Anh. ital. »Ih l.iol., IKOl, x\xv, 317.

>lil|inri't' lie la (JlKentinn,

(IS) Krn'xk, M., •• lifitni^'f mr Miixki'lrliiMiile ilffi- ()ct<>|H)di>ti,"/<>it.4chr. f. phyoioi

C'licm., lyi»4, xliii. 47

(16) .Stahkknstkin, K., Hill M. Mkn/b "n»T.I.-ii N»cli
.M«ere»iiiol)iiMkHii. ■ Zeu,rlii. f. pIivhihI. Cli.-ni., lUI:.', Ixxxii. 41

wfis von (ilykoKfli \m

(17) lloii'B Sk.ti.kh, K, "ri.pr riiterwliit<le im ihetiiiiHiheii Bau uikI in ,l.r
VenlmiuiiK Imlipier iind iiuilerer Tliipr«, " Airli. f. .1. jjpr. |'hyiiii)l., liv. 395.

(18) KiH.- II, 1. It, cf. llJH.Ur

III a II II, 0)1. lit.

(19) SciM.NiPOHKK, », aii.l K. NVachhoi.i.ich, " UUr lien (}|yk«Kmii.t-iH'w.-fl,„|
<|pr FiHche," Anh. f. .1. «?». I'liyHiol.. 1914, civil. 147.

(20) Lmnbr, K. .1.. '• |)a« Verhalteii .le« (JlykoKeii* <|er KriJiiche \m Amixyhi.we
uii.l Re«tiluti.>ii,"2fitN.lir. f. I'.iol., 191.1, Ix. 388.

(21) HJ.CIBTRIU, M., "\Veit«re Uiiter»ucliiiii({eii iiber lUa Verhalten <ie^
(JlykoKMn im Kierttock Ht-r H. fuH.a." Arrli f. -1. k'". I'liy»i<>l., 1911, cxii. 328.

(23) Cruukshank, E. VV. M., "On the rro<liicti,.n uii.l UtilinHtioM of Ulyfoj{fii
in Normal luiil hialietic AnimaU," .lourii. I'liysiol., 1913, xWii. I.

<!^^S

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UNIVtMlTT or TOKONTO tTVDllS

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4» TN dwnlMrfer nrkMl fi«tM» kjr O. O. NAimni. . .
$i 1W yalMwdwuhiii rf tfcg mam, by A. B. MMAUtm
6i Tte alMeiptfM of fitt is tte iaUitiM* by G, E. WtuioM

7: TlM4iMiibllt{M«rftt,lM«ridM.plMMaiMtM.B0UMilNB

Md inm IB ttriatid •««:»}. I7 Mm» L. Mnrrm

8: S<w<«CT t— kHi —4 i>ilri |»hw wnw, »y A. B. MwMUW
9: Oa tkt 'AUribulfiMi of potMriim la rami eoil^ by
C. P. Buntnt 77. '

Hfc 10: On tte pntbBM* M^iira of iIm ■obMwM prometiait

Erowth in ymmg wrinwla, by CMUint Fuw umI A. Aiua
I4CMXVM

Mo. 1 1 s Tb« eoomorativ* vkIho of kiri nd butter io nowth.

by Casihib FwiK «ad A. tbrnea tiMMXxm

No. la : The fMm oTjrt^ fracttope ott tk« frowtli of rats, by

Casimm ^mc Md A. tmvat Macauwm

No. 1.1: A oew Mtoee^tion o* tlM tHomtviv function, by

T. G.B11001S

On duuifM ki Um ftouwrult and ttdMil*^ irf tiw
kklooy Micompw^nBf aetiv%. by T. G. Wttuemmi I. J.

MAOcmxtt

No. 14: Purther <ri>Mrv«tiaiMi on tho diffiervaOBl Mtion of\

aOrma&a, by Fkamk A. HaAhmah and Lms McI^mnMJUf .
No. 15: TIm oicdMiiiain for vModibrtatioo from •drmotin, by

FsAMK A. Hastman aod LoM MelMBituM Fiunm

No. 16 : AlfaciMlin VModitetor niccliiai«m w ^m cat at dtffamit

affH. by Pranii A. iiMni.ti and U«u« 6. Kiuonf . . . .
No. ITS Location of^^^^ adrMalia VMoiUator mccMoisaM, by

FtaAmc A. HMWfwm, L.G. Kiukmw and Lots Frasbs..
No. it: VaseiAtfeba<igMprDdBe«lby«lrciia8ainv«rtebntM,

1^ Frank A. HAantAM. Lbsu* 0. KiuKHtN and Ross S.

Ljuio

No. 19: Sioiptifiad fas aat^fyik^ t^ j- J. R. Macmod . .

No. so: Adrena&i vasodilator mechanisBis, by Frank A.
Hartmam. LttiUR G. KiLacHtN aad L(m Fraser

No. ai:^ Coo«tn^kN>^tN»ad»sa{ia ae^g^ obor igwmtbetic
aad dorsa! root g ai^is, by Frank A. Hartmam, Lbslib
G. KiUMiN asd Lora Frasbr

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S4< ji The «poiilit)i*i>it« dfv ■*l>>pi>irnt of an li ulo«i« vOnUititfn

m »li-viMi-br.»ii.- c.ilo. h\ | | K M \n M>l« 035

S- .1 I lu' ili.iynn'.i'. i>l .ulilnsl^. h\ I J tt MxiiMiK o 3S

V>> j(. Siiiiplili.'il K I- •">•''*"<''»• ''^ 1 J K M A< I I i>l> 035

\i». 35: Dh^crvalii'ic I'li ilccrrcbrate cats, by l.<>i-> l-Nx-m, K

s l.vM. ,in»l I I K Mm iu>i> o 35

No .Ml l>t .ith pi.Klii.ol h\ t\iin; lU* aJreii.il vtiii< hv !■ A

II \Ml\i\N .iiul \V I, Hi m/ o 3S
Ni) j; A.ii.'ii.ii \ilffiialiii i<ii tlie opU'vii, b> I \ H\kimvn

.iikI K.<»' S I \m. "35

N.>. JH; rill' .uiiKii 1)1 \ilrriialin mi tli» ki»lii»\. by Ikank A.

llvKiMW .iiul Ki'«» S l.wi. <i 35

So. 31); Siittic r#i.i'Ml \»oik on the ..oiitrol ot the respiratory

..Hire, by | | K. Mm iii>i< . . "35

\,i v>: Sliulics III lln? ri-i;iMicr.ilii>ri el ilciici \ .iKil iii,iHHH.ili.in

iiiii>iU, bv K A n \KiM\N. W K Hi M/ aiiU I. ti Kn homn •) 5"
No •,! lln- luiKtu'ii.il patlioUiyy of klir^'iciil shoik, by J. J. R.

M M 1 MM. " 35

Nil \J i»n ^fiihl.iiioii, h\ I I K Mmium' ii.'S

No, ;v Ob'.erv .iluiii- on the i;l\i:omMi conU'iit ot cm. ini in-

viiubrali* anil li^lu•s. by I,, li. Kli liOKN .nul ). |. K.

M \i 1 Kon " -'>

No -,4: 'I lie bchaviotir of the respiration* alter detcrebration

III the.. It. by J. J. K. M \i 1 l-.oi> . '<^S

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