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Microscopic changes in the pancreatic gland of the cat produced by sympathetic and parasympathetic stimulation.

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Departments of Ph?lsiology and Anatomy, YcGiTZ L'niuarsily,
Xontreal, Canada
A combined physiological and histological investigation of
the pancreatic gland of the cat was undertaken with the objcct of studying some of the effects produced h p the action
of the sympathetic and parasympathetic secretory nerves and
also of certain sl-mpatho- and parasympatliomimetic, drugs.
The liistological part of the worli was confined t o study of the
discharge of the zymogen granules from the acinous cells
when the gland was stimulated t o secretory activity hy various methods. The more apparent clianges in the islands of
Langerhans were also notcd.
A similar investigatioii of the histological changes in the
acinous cells of the dog's pancreas was previously performed
by Babkin, Rubttslikin and Savitch ( '09). However, the conditions regulating pancreatic sec reliori in tlie cat are somewhat different from those obtaining in tlie clog, especially as
regards the effect of vagal stimulation. In the dog vagal
stimulation elicits a small amount of pancreatic secretion, but
in decerebrate cats it has been found impossible to obtain a
visible secretion of pancreatic juicc under prolonged rhythmic
stimulation of the vagus nerves or under pilocarpine (1Cor.ovitsky, '23). Hence it was thought desirable to study the
above-mentioned changes in the pancreatic gland of tlie cat,
including the more apparent changes in the islands of
Physiological .methods
The cats were fasted for 24 hours before experirnent. During brief ether anesthesia a mixture of urethane and chloralose (0.5 gm. arid 0.05 gm. respectively per kilogram body
weight) was given intravenously. Tracheotomy was then
performed, and the vagus nerves were cut in the neck and
placed on electrodes. The stomach and duodenum were separated by tying of the mucous and submncous membranes, the
muscular layer being left intact. When necessary, the pancreatic duct was cannulated, arid a cannula was inserted also
into the duodenum (in the latter case, in order to introduce
HCI). I n one control experiment, the spinal cord was cut
below the medulla during brief ether anesthesia, artificial
respiration was applied, and the vagus nerves were stimulated. In experiments involving splanchnic stimulation, the
splanchnic nerves were tightly ligatured and placed on shield
electrodes, and the blood pressure was recoided in order to
test the efficacy of each stimulatioil. In some of these experiments, the suprarenal glands were removed.
The vagus or the splanchnic nerves ~7erestimulated rhvtlimically with an induction current. I n either case thc nerves
were stimulated alternately for &minute periods, with rest
intervals of 2 o r 3 minutes, the stimulations Ineing continued
for 1; to 6 hours. During this time the current was gradually
increased from 14 cm. to 7 or 8 em. Adreiialin (Parlie, Davis
& Compariy's) and choline chloride solution, when used as
stimuli, were introduced intravenously.
Hist o 1o gicnl m c t 720 ds
At the beginning of each experiment a control sample of
tissue was taken from the pancreas and placed in freshly prepared Zenker-formol solution. Latterly a modified form of
this solution (Baley, ' 3 7 ) was fouiid to be more suitable, riz. :
3% inereuric chloride
3% potassium bichroinate
40%' formalin (added immediately before using)
45 ce.
45 ec.
10 cc.
At inten-als during the espcrimc~iit pieces of I)ailclae:ls w e r ~
escihe(1 and piacwI in this solutioii, a11d at Ilie concluiioii of
t 11c c s p e cn t the gla 11il
fixed by in tril-a1.t( L rial intiection
of the samc soliition. Tlir ti\sue l as c~m-hctltledin paraffin.
Sections w e i ~cut, 4 or 5 p thick, w i d stained with neutral
gentian (crl\-stlal-\iolct-or*angc U ) . F o r cwrilrol ~ U I - ~ J ~ I Stwo
cats, -\?rhich had lwen fasted l o r 24 hours, w c i ~aiiostlietizetl
wit11 nembutal arid then injected intm arterially u.jtli h i k e r formol solution. RimiIarlJ- two citts, which had hwn fed 3
houi.s before, w w e anesthetized with nernhiit a1 and injected
with the fixing fluid.
t:x P t; I\' I hl E N '1'A I,
t cst iwg
The aciiioiis cells of the pancreatic. gland of a eat, f'astd
for 34 hoiirs, on microsco1)e cxarnination are swii t o he in a
qniclsceiit state. The cells a r e filled w i t h zymogen graniil~s,
which, niide~-neutral gentian staining, show up d e ~ pviolet
against a background of pale gi.ayish-vio1et cytoplasm. The
ducts are empt?, The 1r ~ i n s p a r ~ nareas
of the islands nf
Langerhans stand out in marked coiitrast to the iwinon::
tissue ; orily tlie P-cvJl cords are visible, their granules being
stained deep blnc. Vost of the islarids are in w r y close coil
tact with the aciuous tissue; there appears to be 110 capsulP
separating them.
The acinous w l l h i n the imrncdiate vicinity of the islands
a r e more dciiisely packed with zymogen granules than a r e
those in the more rcmote parts of the lohule, so tliat a kind
of peri-insular ciwle, 01' 'halo,' of uned~austtedacinons cells
appears around each islancl. This phenomenon aras forrncrlv
o b s e r r c ~ lby Jarotsky (1899), Dale ( 'M), Rehaler ( '20). and
Dolley ( '25). (Figs. 1 and 2.)
When in a fasting animal it l-iappcned tliat tlic pancreatic
gland had heeii in a state of moclex-ttc secretory activity at the
heginning of thc experiment, owing to the presenre of' some
food residncs i r r the stomach, the acinous cells surrouiiding
the islands and forming the 'halos' were more distinctly
risihle thau in the quicsceiit gland. T n thew modcratcly acti\.c
glands, secretion conld also be s c ~ nin the ducts ; whrii there
were no food residues in t l i ~stornacti or iiitestitie the paiicreatic ducts were always empty. 111 auimals fed :3 liours
beforc experiment, the inici-oscopic pictur*thwas similar to tlic>
above, though tlie changes W‘CI*C I Z ~ O I T protiouiiced ; thrl ‘ lialos ’
a-ere more riumcli*ousand morc strongly riia~ked,ant7 the dncts
\vei*e in many instauces filled with scbcretioii.
S t im itlnf io E )
LC ith
h ydrorli I(, I-ir (ivid
Introduction into the duodeiium of a h o u t I0 cc. of 0.2% HC’1
at i n t e i ~ a l sof 3 hour, during a pcriod of 3 to .i lionrs, elicited
a flow of paiicrcatic juice, thc volume o f w h i c h varied in different experirnents. Iii sections of paiiciw~staken f’iwrn a n i mals so treated, it was observed that most of tlie aciiious cclll\
were well fillctl with xvmogcn granules. The gmiiiilcs o c c ~ i pied the apical zone of the cells, but werc not so closely I)RCI~CCI
a \ after . r ~ q ystimulation
( s e e b e l o ~ v ) . They ~ e r soniewliat
smaller. tiiaii the granule:, in resting cellb. ‘ l ’ h ~ ductules WPIT
1)ractically I’ree from stainable matei.ial, so tliat, a s tlicy liad
probably contaiiicrl some scciwtion, this milst liavr l w n of a
watc1.5- coiisistcncy. The duchies were riot clilatcd--as is to
bc cixpectecl ’\~rlic11a watery swrCtioii, wliicdli escapcs 1-apiclly.
is heiiig procluced (fig. 3 ) .
T’(iga1 d i m iilutioii
The rilgus nerves x e r c stirnuluted i*hj-thmicallj-in thc ~ C C B
wit11 a11 incluction current (coil 14 to i mi.) foi. a pcJriod of
1) l o 6 lionrs. Sniall samplcs of paricreiih f o r histological
examinatioii ~vei*etakeii at intervals during t h c ~c~sperinic~it.
I n some of tlie cxpei-irnents thcl panci*eatic duct W R S caiitii1kited, h i t it was a striking f a c t that, i n spite of strong arid
prolonged stimulation of the vagus i i ~ w e s tliercl
was i ~ i + ~ c t i cally 110 visi1-h secretion of pancreatic jnirc. One control
experiment was performed in, in order to a\ oid the
possible unfavorable efTect of anesthetics 011 tlie fiiiictioiiiiig
of the p a n c ~ c a sthe
, spinal cord had been cut below the medulla
and all the blood vessels going to the head tied. Although the
effect of the small amount of ether used at the outset undoubtedly soon passed off, the effect of vagal stimulation in
this experiment was also practically negative. Thus, during
4$ hours of vagal stimulation the gland secreted only 0.4 cc.
of juice, whereas the introduction of 25 cc. of 10/N HC1 into
the duodenum provoked a flow of juice amounting to 1.6 cc. in
23 minutes.
Notwithstanding the absence of a flow of juice, prolonged
vagal stimulation produced extremely great morphological
changes in the gland (figs. 4 and 5). Most of the acinous cells
were found to have become entirely depleted of their zymogen
granules. Where zymogen granules still remained in the cells,
it was observed that they were congregated in the apical region. The cytoplasm of the acinous cells had become yellow,
the nuclei being only faintly visible. The ducts were filled
with material which took the same stain as the granules, and
they had the appearance o f a tracery of branches and twigs.
About 70% of all the islands, a s seen under low power magnification, were surrounded with ‘ l i a l o ~ . ~However, in the
aciiious cclls forming these ‘halos’ the granules were much
less numerous than in the cclls o f the resting gland. The
Hood vessels were dilated and filled with erythrocytes. If the
period of vagal stimulation was shorter, the above-described
changes in the gland were less pronounced.
I n later stages of vagal stimulation the islands underwent
marlred changes. The outlines of the cell cords had become
indistinct, vacuoles had appeared in them, and the capillaries
were dilated aiid filled with red blood corpuscles. Some of
the islands appeared to be greatly distorted. I n many places
between the islands and the acinous tissue, groups of cells
could he discerned which iii their apical part contained some
zymogen granules and in their basal part were stained the
same blue color as the granules of the P-cells (fig. 6).
u 1at i o I I with c+ uliii c
C’holine chloride iu equal doses of 2.5 mg. to 5 ing. \$“ils
introduced intravenously at intervals of 5 or 10 minutcs dui-ing a period of 2 to 4 hours ( a total of 160 mg. to 3:37 mg. of
choliric being atlrninisteretl). TTlirn tlie pancreatic gland is
stimulatetl with cholincl, the histological picture is similar to
that ohtained w7l11311 the gland is stimulated throngh the vagns
nerves. Uiidcr low powt’r mrtg~iifiration, 80 to 907( of the
islarids a r e seeii to he encirclt4 by ‘halos’ (fig. 7 ) .
01 t7ir
splnwlL1tic. Jtcri‘cs with flic suprcirmcil
gla M d.7 i l l t a ctt
Rhptlimic stimulation of the hplanclinic n e t ~ c swith an irrdnction cnrrent (coil 11 to iem.) during a prriod of 2 t o 6
hours also p roduced cliariges in tlir acinous aiid thci ihland
tissue, although no secretion appeared in tlie oariiiula which
had been introduced into tlie paiicrcatic duct. These changes,
horn-ev~r,were less ilrilstic than those which resultccl from
vagwl stimulation. The zymogeri granules had been piwtl?;
discharged from tlie cells and thc secretion could be w e n in
the clucts. The capillaries contained many crytlirocytes and
u-err dilated, but less so tliari after vagal stimulation. Tt
coiild he observed that about 19 to 3 0 9 of tlie islands mire
encircled by ‘Iialos. ’ C‘ertain changes liacl oocui.rcd in the
islarids themselves. T h e cell cords sliowetl many irregular+
ties arid contairied vacuoles. In niaiiy places they sccriiecl to
he continuous with the acirious tissiie, especially i i i the experimerits where t h e w was prolonged stimulation (figs.8 and 9 ) .
The capil1ai.y network was dilated and contained red blood
co r p US el e s .
141(1tio11(if f 1 1 ~spla
s u p ra retin 1 gla I
a f t r r r p m o r c x l o f the
L ~ S
h f t c r the suprarenwls had been removed, the histological
picture obtained wlien the pancreatic gland was suhjec~tctlto
the effect of rhythmic splanchnic stimulation during a period
of lfrto 5 hours resemlrled that described in tlie previous section, hut the changes were much more pronounced (fig. 10).
The acinous cells in the center of the lobule, particularly
around the islands, were depleted of zymogen granules ; those
a t tlie periphery of the lobule coritaiiied fair numbers of
granules. There was secretory material in tlie ducts. ‘Halos’
of unexhausted acinous cells round the islands were rarely
present. I n this type of experiment the ‘halos’ had begun to
disappear very early, i.e., a t the end of the first l+ to 2 hours
of splanchnic stimulation. The islands themselves were
hardly visible under low magnification, appearing as transparent areas. Under high power these areas were seen to
contain colorless cords of island cells and dilated capillaries,
and the island cells w7ere in very close contact with the surrounding acinous tissue. M7hatever explanation may be given
in regard to all these changes t o be observed in the insular
cells, it is clear that, with tlie suprarenals removed, even brief
stimulation of the splanchnic nerves causes profound modifications t o occur.
Stinttilation with adre9talin
I n response to intravenous injection of various amounts of
adrenalin no visible secretion emanated from a cannula inserted into the pancreatic duet, nor was there any marked
discharge of zymogen granules from the acinous cells. Rdrenalin solution was injected during a period of 4 t o 5 hours,
samples of the pancreatic tissue being taken f o r histological
investigation a t intervals during the experiment. Massive
doses of adrenalin corresponded to 1 cc. of 1:10,000 solution
administered every 5 minutes. Less intciise stimulation of
the gland was produced by the injection of adrenalin solution
of the same concentration at 15-minute intervals. The impression obtained from these experiments wits that during the
first 14 to 2 hours of adrenalin administration there was a
slight diminution in the numbers of granules in the acinous
cells. Subsequently up lo tlie fourth o r fifth hour of aclrenalin
administration thcre was no further diminution and all the
acinous cells still held fair quantit,ies of grtmnles (fig. 11).
Thus the ‘halos’ do not stand out so distinctly as in glands
that have been subjected to other types of stimulation. The
islands are clearly discernible on account of the deep blue
granules of the P-cells, and have tlie appearance of being
somewhat hypertrophied. 1t has been found by other inrestigators that prolonged treatment of animals with adrenalin
(injected over periods of np t o 30 days) produced great
changes in most of tlie orgaris and in many instances resulted
in degeneratij-e processes setting in. The data concei*niiig
the effect produced on tlie islands by prolonged adrenalin
administration awe, Iiowever, conflicting. Satwornitzkaia, Sirnnitzky and Spassky (’32) arid Rernabeo (’35) noted hyperplastic phenomena in the island cells; Igura (’27) found the
islands to he atrophied, and Samson (’32) described them as
The cat has proved to he an exceptionally suitable animal
in which t o study the more intimate mechanism regulating the
secretory activity of the pancreatic gland. Whereas in the
dog stimulation of the v a p s nerves produces a definite though
scanty secretion of juice rich in organic material, in our experiments on the cat only 0.25 to 0.5 ec. of secretion could be
obtained from the pancreatic duct during 4 o r 5 hours of
vagal stimulation. Nevertheless the microscopic changes in
the cells of the eat’s pancreas under these circumstances were
as great as in those of the dog’s. The large zymogen granules had disappeared almost entirely from the acinous cells
and small ones remained, a condition which would indicate
that, under the influence of the vagus, certain intimate processes occurred in the acinous cells which clianged the appearance of the secretory material. Many of the acinous cells
were entirely depleted of their granules, and wlicre granules
still remained these had accumulated in the apical region of
the cell. The ductules were filled with material stainable with
tlie same C i p as tlie granules. Thus we have here an exceptionally good illustration of tlie type of nerve effect which
Heidenhain (1878) designated ‘trophic.’ Since the organic
colloidal material was evacuated from the acinous cells probably quite unaccompanied by water, it remained in the ductules, where it was coagulated by the fixing reagents. Therefore in the cat the vagus supply to the pancreatic aciiious
tissue is purely ‘trophic. ’ Again the hnmoral stimulus (i.e.,
secretin, formed in the duodenum in response l o HC1 administra tion) produces what Heidenhain termed a ‘secretory’
effect. Under its influence water and salts pass through the
cells in abundance, only a small amount of organic colloidal
material being discharged from them. As already observed
by previous investigators, the pancreatic juice under such
circumstances is poor in organic material and enzymes.
Stimulation of the vagus nerves seems also to produce
another effect on the acinous tissue. Mention has already
been made several limes under ‘Experimeiital results’ of
‘halos ’ of unexliausted acinoiis cells, surrounding the islands
and containing fair numbers of secrctory granules. These
were observed in the pancreas of control animals and of animals that had recently been fed, o r had received EICI. The
preseiiee of these ‘halos ’ is one of the most characteristie
histological effects of prolonged vagal stimulation o r of choline administration. On the other hand, a diminution in the
number o f ‘halos’ is in most cases characteristic of the histological picture following splancliriic stimulation, especially
when the supi-arena1 glands have been removed. No definite
cxplanatiori of this phenomcrioii can now he offered. However, a n attractive hypotlicsis I\-oulcl be that the acinous tissue
in the close vicinity of the islands is subjected to the action
of insulin diffused from the islands. Miss Hcbb ( ’ 3 7 ) ,working in our laboratory, has shown that insulin inhibits the
activity of the acinous cells and diminishes, or even reverses,
the action of the vag-us nerve. There a r e good grounds for
believing that the vagus is a secretory nerve for the islands
when the vagus is stimu(compare La Barre, ’ 3 3 ) . The~*efore,
lated, its ‘trophic’ effect might be diminished in those cells
which are subjected to the immediate action of the insulin
secreted by the islands. Selye ( ’37) observed that in the pancreas of rats, in which an ‘alarm reaction’ had been produced
hy the administration of adrenalin, formaldehyde o r morphine
sulfate, the acinous cells in the immediate vicinity of the
islands retained their secretory granules much longer than
those in the regions more remote from them. I n the present
investigation peri-insular ‘halos ’ were observed in the pancreas of control animals and were diminished or increased
according to the type of stimulation to which the gland was
subjected, but it is very doubtful that they were the result
of the general, unspecific traumatic stimulus of an ‘alarm
reaction.’ I n special control experiments on cats, which were
subjected to the usual preliminary operative procedure here
employed (the vagi being cut in the neck arid the pylorus
ligatured, etc.) but were not further interfered with, no diminution in the number of zymogen granules in tlie acinous cells
of the pancreas was noticeable; thP ‘halos’ were no more
marked in samples of pancreas obtained 4 hours after the
operation than in the control pieces.
The different effects produced on tlie pancreas by splanchnic stimulation (with the suprarenal glands either intact or
removed) and when adrenalin was administclred is probably
attributable to the antagonistic effects of the sympathetic
nerve and adrenalin on the acinous cells. TT’hereas splnnchnic
stimulation with the suprarenals intact, and more especially
with tlie suprarenals removed, causes a discharge of zymogen
granules (as does vagal stirnulation), administration of adrenalin seems to inhibit this entirely. Since an intimate relationship exists between carbohydrate metabolism and the
secretory activity of the pancreatic gland (Still, Bennett and
Scott, ’33; Hebb, ’35, ’37, ’38)’ the discrepancy between the
effects of splanchnic nerve stimulation and adrenalin administration might perhaps be accounted f o r by the peculiar effect
which adrenalin exerts o n carbohydrate metabolism. Bccording to Cori (’31) tlie carbohydrate metabolism is eventually
inhibited by adrenalin, which diminishes the ntilization of
sugar in the tissues.
The changes brought about by vagal and splanchnic stimulation respectively in the histological picture of the pancreatic
gland are not entirely similar. The presence of ‘halos’ of
uneshausted acinous cells around the islands of Langerhans
was found to be very characteristic of tlie vagus effect, as well
as of choline administration. A decrease in the number of
‘halos ’ was a typical phenomenon in the splanchnic experiments. This difference in the effects of the vagw and the
splanchnic nerve may perhaps be explained by the theory that
the island tissue is acted upon by the vagus but not by the
splanchnic. The histological studies of De Castro (’23) on
the mouse’s pancreas lend support to this theory. He came
to the conclusion that the vagus innervates the acinoiis tissue
of the pancreas and also the islands of Tmgerhans, while the
sympathetic nerve fibers supply only the blood vessels to the
gland. If this view is accepted, then it is clear why the stimulation of the splanchnic nerves does not increase the output
of insulin, which mould check the discharge of zymogen granules from the near-by acinous cells.
1. Prolonged stimulation with an induction current of the
vagus nerves in the cut did not elicit any visible secretion
which could be collected from the cnniiulated main cliict of thc
pancreatic gland.
2. Microscopic investigation of the pancreatic gland, after
it had been subjected t o prolonged action of the vagus nerves,
showed that a ) the acinous cells were almost entirely depleted
of secretory graniiles, and b) the small ducts were distended
with a material stainable with the same dyes as tlie granules.
3. The introduction of a solution of HC1 into the duodenum
produced a flow of pancreatic juice, hut histological examination showed that the discharge of secretory granules from the
acinons cells mas very slight.
4. Intravenous injection of choline chloride produced
changes in the pancreatic gland similar to those brought about
by ~ a g a stimulation.
5. There was no discharge of secretory granules from the
cells of the pancreas in response t o intravenous administration of moderate or of massive doses of adrenalin.
6. The effect of splanchnic stimulation on the acinous tissue
of the pancreas was similar to that produced by ~ a g a stimul
lation; it mas much more pronounced when the suprarenal
glands had been removed than when they were left intact.
7. Under raga1 stimulation o r choline administration the
discharge of granules from the acinous cells surrounding the
islands of Larigerhans was apparently slower than from the
cells in the other parts of the pancreatic lobules, as shown by
the presence of ‘halos’ of unexhausted acinous cells around
many of the islands.
8. Changes were observcd in the structure of the island
tissue as a result of stimulation of the secretory nerves of the
pancreas as well as after administration of choline.
This investigation was made possible by a grant from the
Ella Sachs Plotz Foundation, which is gratefully acknowledged. The writer is greatly indebted to Prof. B. P. Babkin
for suggesting this problem arid for providing laboratory
facilities in addition to much helpful criticism and advice in
connection with the work. She is grateful to Prof. C. P.
Martin for kindly providing the necessary accommodation and
facilities for this research in the department of anatomy,
histology and embryology. She desires to acknowledge the
interest shown by Dr. D. J. Bowie and Di-. Hans Selye in this
work. She also wishes t o thank Nr. D. J. NcKinnon and
Mr. G. Bresnahan f o r their valuable technical assistance, and
Miss J. F. Oswald f o r aid in the preparation of the manuscript.
33 1
AVI) W. W. ~ A V I T C T I 1909 Quoted bp B. P.
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p. 617. J. Springer, Berlin.
BALEY,J. H. 1937 Staining methods f o r thc islets of Langcrhanq. J. Path.
and Hact., vol. 44, p. 272.
E. 193.5 1,esioni anatorniche ed alterazioni cndocrinc da adrenalina.
Riv. P a t . sper., N.S., vol. 4, p. 121.
CORI, C. F. 1931 Mammalian carbohydrate metabolism. Plrysiol. Rev., vol. 11,
p. 143.
DALE, I€. H. 1905 O n the islets of Langerhans ' in the pancreas. Phil. Trans.
Koy. 800. London, Ser. B, vol. 197, p. 25.
DE Casrrto, F. 1023 Contribution B la connaisswnce de l'innerration du pancr6as.
Rech. Biol. Univ. de Madrid, T . 21, p. 423.
DOLLEY,D. H. 1926 The general morphology of pancreatic cell function in
terms of the nucleo-cytoplasmic relation. Am. J. Anat., vol. 35, p. 153.
HFER,C. 0. 1935 Eelation between the blood sugar concentration and the exocrine function of the pancreas. Arch. Internat. Pha~r~iacotiyn.
vo1. 52, p. 33.
1937 The effect of insulin administration on the response of the
pancreas to parasympathetic stirnulation. Quart. .J. Exp. Physiol.,
vol. 26, p. 339.
____1938 The effect of injected insulin on the storage of glycogen in
tho pancreas and liver. Quart. J . 32x1). Physiol., vol. 27, p. 237.
HEIDENHAIN, R. 1878 Ucber Rekretorische und trophische I)riisennprven. PflUger's Arch., Bd. 1 7 , S. 1.
8. 1927 Einfluss des Adrenalins auf die iiinersekretorischeii Organe.
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A. J. 1899 Uebcr die Verkndernngen i n die Grbsse nnd im Bau der
Pankreaszellcn bei cinigcn h r t e n der Inanition. Virchow 's Arch.,
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L. K. 1923 The part played by the ducts in the pancreatic secretion. J. Physiol., vol. 57, p. 215.
J. 1933 DiabGte et insulin8niie. Masson ct Cic, Paris.
P. C. 1932 Tissue changes following continuous intravenous injection
of epinephrine hydrochloride into dogs. Arrh. Path., vol. 13, p. 745.
A44h'D A. J . SPASSKY1932 Iler Einflusa
chronischcr ddrenaline inspritzungrn auf den endokrincn Apparat.
I. Der Einfluss der ~~drcIialininjekti[inen
auf die morphologische Beschaffenheit dcs endokrincn Parrnchyms der Knuchspeicheldruse. Z.
mikrosk.-anat. Forsch., Bd. 28, S. 427.
ntisls of Histology, 11th ed., p. 371
Longmans, Green bk Po., London.
SELYE,H. 1937 Studies on adaptation. Reprinted from Endocrinology, vol. 21,
p. 169.
AND V. B. S c m 1933 A study of the metabolic
activity of the pancreas. a m . J. Physiol., 1701. 106, p. 509.
1 Pancreas of normal eat (fastcd 24 hours). The ‘halos’ of acinous cells
containing undischargcd zymogcn granulcs staid out distinctly around the islands
of Langcrhans. X 75.
2 Pnnercas of normal cat (fasted 24 honrs). The acinons cells a r e filled with
qniogen granulcs. X 400.
3 Effect of stirnulation with HC1. The numbers of zymogen granules in the
aeinous cells arc not appreciably diminishcd. No sceretion is visible in the ducts.
X 400.
4 Effect of vagal stimulation. Xccretion may be seen in t h e ducts. The numbers of granules in the cells are much diminished; some of tlie cells a r e almost
cmptp. x 400.
5 Effect of vagal stimulation. With the exception of those forming ‘halos’
around the islands, the acinous cells contain practically no granulcs. X 75.
6 Effect of vagal stimulation. One of the islands. Note the close contact
bctween the island and the acinous tissue. The island is distorted; it contains a
number of vacuoles, and many erythrocytes a r e present in tlie capillaries. x 300.
7 EITwt 01’ cholinc admiii
Thr aciiious cells, cxccptiiig ;hose f o r m i n g
halo^ ’ :ironncl tlic islands, illtic;illy empty :,f gr:i~iules. X $ 3 .
8 Eflect of ~plnnc.liiiicstimulation. with tlic supmrvri;rl g1;tndln i u t a c t . Kate
that. t h c aciiions eclls iii tlic: irnmrdinto viciiiity of tlir isl:intls coillain 1mictic:illy
PIO z p i o p e t i griiiiules.
The is1:incls stand oil: rlearly. X 75.
9 E f f w t of sploncliiiic stimulation, w i t h t h c suyr:irrn:’I glands int:rct. The
gr:inulea of tlir p-cell C(JIYXS stand out distinctlp. In many plact’s tlle 1)mnd:iry
hettnecri the p re11 c o r d s :tiid tlre sorroiioding :Lclnoiw timile is qtiitcl indistiiig~iidable. X 300.
10 F2ffc:rt of s~iI:iiicl~rii(~
stirnulatitin, with the siipixn:na I gl:t?ids rrmoved. The
eells at the c e n t e r of tlir 1ol)iiles c*ont:iin f c w z~-n10gc~n
grumilrs : tliose a t t h e
priphe1.y still retain sonic granulvs. ‘I‘hcrc arc :ilniost no i s l a n d s :ipp:irPent in
this ~ ~ O U FofJ lobulcs. X 7 3 .
11 Effect, of adrenalin administration.
The ncinous cells are fillet1 with
granules. Tile iskinds stand o u t p r o r r i i i ~ c r t t l ~ .X 7 3 .
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cat, change, microscopy, gland, pancreaticum, producer, parasympathetic, sympathetic, stimulating
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