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Microscopy of the living pancreas in situ.

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Microscopy of the Living Pancreas in Situ'
D e p n r t n i e i i t o,f Aiicitoiniy, College 0.f Meflicine, Uizivet-sit;] of C i n c i n n f l l i ,
Ciizci~ciitrti.Ohio 45219
The purpose of this investigation was to study the microscopic anatomy of the
living pancreas i i i s i t u . The results indicate that: (1) it is possible to study cellular detail in the
living pancreas with resolution closely approaching the limit of resolution of the light microscope; (2) blood flow through individual capillaries in both acinar and islet tissue is intermittent;
(3) local blood flow through acinar capillaries is regulated by smooth muscle precapillary
sphincters and by endothelial sphincters, while flow through capillaries in islets is regulated
locally by endothelial sphincters only; (4) insular-acinar capillary anastomoses exist but are not
frequent; (5) secretory canaliculi between adjacent acinar cells exist in life and pass between
centro-acinar cells to reach the lumen of the ducts; (6) processes of beta cells may pass between
two adjacent cells to provide additional surface area for transcapillary exchange; (7) the
formation and release of zymogen granules occurs within 45-90 minutes in acinar cells stimulated with pancreozymin and the formation and release of beta granules occurs within 20-60
minutes in beta cells stimulated with tolbutamide; and (8) cytological changes during the
secretory cycle in acinar and beta cells are consistent with the current ultrastructural theory of
Recent improvements in methods used of the organ in situ and its alteration by
to examine living tissue and organs in situ physiologic, pharmacologic, and pathohave resulted in a better understanding of logic stimuli.
the dynamic structure and function of sevMATERIALS A N D METHODS
eral organs, and also have permitted rapid
sequential quantitative measurements of
Three hundred white mice (ICR) were
dynamic events in living material (Bloch, used. Each animal was anesthetized by
'65, '66, '68; Bloch and McCuskey, '66; injecting 20% ethyl carbamate (Urethane)
McCuskey, '66, '68a'b). Using such subcutaneously or sodium pentobarbital
methods in conjunction with routine intraperitoneally (Nembutal, 5 mgiml). To
histologic procedures, this paper reports expose the pancreas a left subcostal incithe results of a study with the following sion was made and the pancreas and
objectives: (1) to elucidate further the spleen exteriorized and placed on a supmicroscopic anatomy of the pancreas in porting ring covered with Saran Warpa.
situ in the living animal; ( 2 ) to establish The lobules of the exteriorized pancreas
the necessary morphologic and physiologic were carefully teased apart under a stereocriteria for quantifying the response of the binocular microscope avoiding hemorrhorgan to various physiologic and phar- age and tissue damage, and the preparamacodynamic stimuli at the cellular level; tion covered with a small piece of Saran
(3) to apply these criteria in evaluating the Wrap (McCuskey and Chapman '67a, b,
process of secretion in acinar and beta '68). Homeostasis was maintained by concells; and (4) to compare and contrast this stant irrigation of the preparation with
histology observed in life with that Ringer's solution, the temperature of
obtained in fixed preparations. While previ- which was maintained automatically at
ous studies of the living pancreas have the body temperature of the animal. This
been reported (Kuhne and Lea, 1882; was accomplished by heaters regulated by
Mathews, 1899; Covell, '28; O'Leary, '30; a proportional temperature controller, the
Berg, '30; Beck and Berg, '31; Ruangsiri, control thermistor probe of which was
'49; Brunfeldt et al, '58; Palmer, '59; placed on the surface of the pancreas. The
Palmer and Wilkins, '60; Bunnag et nl., '63, reference temperature (animal body tem'66, '67; Heisig, '67a, b, '68) only a few
Various portiuns of this work were presented in motion picform at the 80th Meeting of the American AssMiation of'
(Palmer, '59; Palmer and Wilkins, '60; Hei- ture
Anatomists. Kansas City, April 5, 1967; at the Midwest Anatosig, '67a, b, '68) have provided the resolu- mists' Meeting. November 1 1 , 1967; and at the 16th MicrocirConference, Atlantic City, April 14, 1968 (McCuskey
tion of cellular detail necessary for careful culatory
a n d Chapman, '67a. b, '68).
Dow Chemical, Midland. Michigan
analysis of the living microscopic anatomy
AM. J. ANAT,I26 395-408
perature) was established by monitoring
the rectal temperature of the animal with
another thermistor probe connected to an
indicating thermistor thermometer and
then setting the controller to maintain this
temperature at the surface of the pancreas
automatically. Using this system the surface temperature of the pancreas was
maintained at the rectal temperature of
the animal 2 0.3"C. Body temperature was
maintained at 37.5"C 2 1°Cby surrounding
the animal with non-absorbent cotton.
Under these conditions blood pH, pOz,
pCOn and red and white blood cell counts
remain within normal limits for a minimum of four hours (McCuskey and Chapman, '67a).
Microscopy of the living pancreas was
accomplished by placing the tray containing the animal and its exposed pancreas on
the stage of a modified Leitz Panphot
microscope, transilluminating the organ
with various wave-lengths of monochromatic light (400 to 700 mp), and examining
it at magnifications between 225 x and
1500 x using Leitz water immersion objectives (22 X, 55 X, 75 X, 80 X and 90 X with
appropriate oculars (McCuskey and Chapman '67a, b, '68).
Optical images were recorded directly by
cinkphotomicrography using a 16-mm
Arrifiex 16-S motion picture camera or
they were projected onto the photocathode
of a 8134A vidicon tube contained in an
RCA PK-301 television system. The resulting televised images were kinkrecorded at
30 fps from the video monitor using the
Arriflex equipped with a special motor' to
synchronize the framing of the video and
photographic images (Bloch, '65, '66;
McCuskey, '68a, b; McCuskey and Chapman '67a, b, '68). Tri-X film was used.
Quantitative information from the optical images was obtained by direct measurement with a calibrated filar micrometer
ocular while that from the video images by
the line selector method and/or stopmotion analysis of the motion picture film
(Bloch and McCuskey, '66).
To examine the secretory process in acinar and beta cells, the cells were stimulated by intraperitoneal administration of 5
units of pancreozymin and 5 mg tolbutamide 5 , respectively, and the response
studied for up to six hours.
Histologic sections were made from the
pancreases studied in life using routine
histologic methods. Immediately after
measurements were made in vivo the
organ was fixed in Bouin's, Zenker's, Carnoy-Lebrun fluids, or in 10% neutral
formalin; i t was then dehydrated, cleared
and embedded in paraffin. Sections, 3 p to
10 p thick, were cut on a Minot-type microtome and stained with hematoxylin and
eosin, Gomori's chrome-hematoxylinphloxine, Masson or with periodic acidSchiff (PAS) using a diastase control
(Humanson, '67). The stained sections
were studied by light microscopy and compared with observations made in life.
Measurements were made with a calibrated filar micrometer ocular and the
same objectives used in vivo.
Images were secured of living pancreatic acinar cells, centro-acinar cells, the
cells of inter- and intralobular ducts; of
alpha and beta cells of the pancreatic
islets; and of the blood vessels supplying
these cells. Figure 1 is a diagrammatic
illustration of the morphology of the mouse
pancreas as observed in life. In general the
resolution of cellular detail was limited
only by the optics employed and was found
to be adequate for measuring quantitatively the dynamic cellular response to
various physiologic and pharmacologic
stimuli (McCuskey and Chapman, '67a, b,
'68). The maximum combined optical-electronic resolution was 0.5 p.
Upon focusing the microscope on the
surface of the living pancreas, the differences in the vascularity and cellular organization between the acinar and islet tissue
were apparent immediately. The islets
were round or ovoid structures located
adjacent to inter- or intralobular ducts,
arterioles and venules, and were highly
vascular structures isolated from the surrounding acinar tissue by a distinct connective tissue capsule (figs. 1,2,5). At low
magnifications the extensive insular capillary network resembled that of a renal
glomerulus . Occasion ally capillary an ast omoses were seen between the islet and the
adjacent acinar tissue (figs. 1, 2, 6). Each
islet was composed of cords or small
'Eastman Kodak Company, Rochester, New York.
4 Sigma Chemical Coinpany, St. Louis, Missouri.
Upjohn Company, Kalamazoo, Michigan.
Fig. 1 Schematic drawing of the relationships between the acinar tissue and pancreatic islet in the living
pancreas. ILA, interlobular arteriole; ILD, interlobular duct; ILV, interlobular venule; 1, islet cell; IC, insular
capillary; IAA, insular-acinar anastomosis; CT, connective tissue capsule of islet; A, acinar cell; AC, acinar
capillary; CA, centro-acinar cell; SC, intercellular secretory canaliculus; PDC, periductular capillary; IA,
arteriole; and IV. venule in vascular pedicle of the islet
groups of cells. The cells were smaller in
size than the acinar cells with each cell
bordered on at least two sides by a capillary. A s a result, capillaries were closely
spaced with only one or two parenchymal
cells intervening (figs. 1, 2, 5). In contrast,
the acinar cells were larger and grouped in
acini, at the periphery of which was
located a capillary network with its vessels
being separated more widely than those in
the islet (figs. 1, 2, 7). Blood entered the
capillaries surrounding an acinus from
intralobular arterioles by flowing through a
precapillary smooth muscle sphincter.
These sphincters locally controlled the
volume and rate of blood flow being delivered to the acinus. In contrast, blood
entered the insular capillaries via muscular afferent arterioles which arose from
interlobular or intralobular arterioles.
Upon entering the islet the arteriole(s)
terminated abruptly in the extensive capillary network. No muscular precapillary
sphincters were observed at the origins of
Fig. 2 Insular-acinar anastomosis in the living
mouse pancreas, as traced from single frames of 16-mm
motion picture; see also Figures 5 and 6. IAA, insularacinar anastomosis; AC, acinar capillary; IC, insular
capillary; CT, connective tissue capsule of islet. Arrows
indicate direction of blood flow.
the insular capillaries from the afferent
Bulging endothelial cells within acinar
and insular capillaries further influenced
the linear velocity and volume of blood
flow through individual capillaries (figs. 8,
9a, b). These cells frequently were located
at the junctions of the capillary network
and were independently contractile. While
these cells rarely occluded the capillary
lumen completely, they were effective in
altering the flow of blood through the individual capillaries; red blood cells
frequently were able to squeeze past a
bulging endothelial cell with considerable
distortion but the less plastic white blood
cells became impacted at those locations
and plugged the capillary for several
minutes to several hours. Blood flow
resumed upon relaxation of the endothelial
The flow of blood in the interlobular and
intralobular vessels was rapid (no cellular
detail seen). In contrast, flow of blood
through the insular and acinar capillaries
was variable and intermittent. The linear
velocity of blood flow in the capillaries
varied from rapid to zero and rarely was
the same in any two adjacent Capillaries.
Intermittency and reversal of flow was
common, but occurred with no definable
rhythm. This variability in the flow of blood
through the capillaries appeared to be the
result of the independently contractile
endothelial cells, the periodic relaxation
and contraction of which modified the flow
and pressure characteristics within the
capillary network.
Exocrine Pancreus Living acinar cells
were pyramidal. The following cytological
structures were discernible in the acinar
cells: zymogen granules, mitochondria,
nuclei and nucleoli (fig. lo). In well fed
animals zymogen granules filled the entire
cell but were more concentrated toward its
apex. The nuclei and their enclosed
nucleoli were round and located basally.
Within a given acinus the distribution of
zymogen granules within individual cells
varied from cell to cell. In addition to zymogen granules a variable number of less
dense vacuole-like structures were
observed which often contained one or
more granules (fig. 11). These were most
prominent near the nucleus but sometimes
filled the entire cell, and were related to the
secretory cycle of the cell (discussed
below). Mitochondria were filamentous,
basally located structures, the majority of
which were perpendicular to the basal
membrane of the cell. Intravital staining
with Janus green B (Ruangsiri, '49)
confirmed the identity of mitochondria.
The dynamic cellular events observed
during the formation and release of zymogen granules in individual acinar cells,
following the administration of pancreozymin, were the following: (1) apical release
of existing zymogen granules; (2) formation of new granules and small vacuoles in
the basal, ergastoplasmic region of the
cell; ( 3 ) formation of large vacuoles containing granule(s) in the Golgi region of
the cell; (4) maturation of these granulecontaining vacuoles so that the size of
each vacuole decreased and the granule/
vacuole ratio increased; (5) apical release of newly formed granules which
appeared to dissolve at the membrane of
the cell; they were not seen in the lumen.
The formation and release of zymogen
granules occurred within 45 to 90 minutes
(fig. 3).
Cells of the terminal portions of the
intralobular ducts (centro-acinar cells)
were recognized by the central position
within the acini, the size being smaller
than surrounding acinar cells, and by the
small number of zymogen granules. The
lumen of the intralobular ductule formed
by these cells often passed between and
behind the centro-acinar cells to form
intercellular canaliculi bordering directly
on the acinar cells (fig. 1).
Intralobular ducts were composed of
cuboidal cells and were paralleled by intralobular arterioles and venules from which
the ducts derived a periductular network of
capillaries (fig. 1). The smallest intralobular ducts, however, did not have an extensive plexus, being followed a short distance
into the lobule by intralobular arterioles
which rapidly terminated in the acinar
capillary network which encased each acinus. Only a few capillaries formed the
blood supply to these ducts.
Endocrine Pancreas Cells of the pancreatic islets, in contrast to the acinar cells,
exhibited no consistent polarity or shape.
While differentiation between alpha and
beta cells was difficult, the following
characteristics were fairly constant. Beta
Fig. 3 Secretory cycle in a living acinar cell in s i t u , as traced from single frames of a motion picture (McCuskey
and Chapman, '67b). A, apex of acinar cell; Z, zymogen granules: N, nucleus; BG, basal granules and vacuoles;
PZ,prozymogen granules. 0', initial appearance of cell; a', eight minutes after the administration of pancreozymin
many zymogen granules ( Z ) have been released and small granules and vacuoles (BG) are forming in the basal
ergastoplasrnic region of the cell; 2 O ' , after twenty minutes all zymogen granules have been released and
prozyrnogen granules (PZ) are being formed in the nuclear (Golgi?) region of the cell; 30',after thirty minutes
extensive prozymogen granule formation is continuing; 45',new zymogen granules ( Z ) are being formed and
released; 6 O ' , more new zymogen granules (Z) are being formed and released, while synthesis of prozymogen
granules (PZ) is diminishing.
cells were located predominantly in the
central portions of the islet and generally
had a more spherical nucleus than the
alpha cells, their nuclei generally being
ovoid (fig. 12). Frequently, cytoplasmic
processes of the beta cells extended
between adjacent cells to abutt on a nearby
capillary. The majority of alpha cells were
confined to the periphery of the islet. Topical application of diphenylthiocarbazone,
an orange-colored, zinc-chelating agent,
helped to confirm the identification of beta
cells (Brunfeldt et al., '58) as did their
degranulation after the administration of
alloxan or tolbutamide (McCuskey and
Chapman, '67b). Alpha cells, however,
failed to respond to alloxan or tolbutamide
administration and did not stain regularly
or as intensely with diphenylthiocarbazone.
The sequence for the formation and
release of granules within beta cells is illustrated in figure 4. A complete cycle occurred within 20 to 60 minutes in beta cells
stimulated with tolbutamide. The release
of beta granules exhibited no polarity in
contrast to secretion of zymogen granules
in acinar cells; granules dissolved and
were released at the cell membrane on
several sides of the cell that were adjacent
to capillaries.
Cells were located at the periphery of the
islet which were neither alpha nor beta
cells. They were spindle-shaped and had
the appearance of modified ductular epithelial cells. Generally a cord of' these cells
Fig. 4 Secretory cycle in living beta cells in situ, as traced from single frames of a motion picture
(McCuskey and Chapman, '67b). B, beta granules; N, nucleus; PB, pre-beta granules and vacuoles.
0', initial appearance of cell prior to administration of tolbutamide; 5', five minutes later many of the
mature beta granules (B) have been released and small granules and vacuoles (PB) are forming in the
perinuclear (Golgi?) region; 25', after twenty-five minutes there are many pre-beta granules and
vacuoles (PB) and a few new beta granules (B) are formed; 40', by forty minutes many new beta
granules (B) have formed and are being released while the number of pre-beta granules and vacuoles
formed a pedicle from the islet to an adjacent duct. Occasional granules resembling
those found in beta cells were visible. The
pedicles were accompanied by the afferent
and efferent blood vessels to the islet.
Comparative measurements between
the structures observed in life and those in
selected, well preserved static histologic
preparations revealed that the major
fixation artifact observed in the fixed
tissue was shrinkage of blood vessels
(table 1).
This study illustrates the feasibility of
studying and recording photographically
cellular detail and secretory processes in
the living pancreas while maintaining
with resolution closely
approaching that of the light microscope.
In the mouse the flow of blood through
individual capillaries of the acinar and
islet tissue is intermittent. Total flow
through an islet, however, is not intermittent. While Heisig ('68) also reported intermittent flow in the acinar capillaries of the
rabbit, Bunnag et al. ('63) did not observe
intermittency of flow in the capillaries of
the islets. The failure of this latter group of
investigators to observe intermittent flow
in vessels of the islet may be due to the use
of the stereo-binocular microscope which
is not adequate to resolve individual
capillaries in the highly vascular islets.
Endothelial cells in capillaries of both
acini and islets appear to be responsible for
Comparutive measurements between blood vessels
"in viuo" and in fixed preparations
Intralobular arterioles
Acinar capillaries
Intralobular venules
Insular arterioles
Insular Capillaries
Insular venules
Insular-acinar anastomoses
In uiuo2
9 & 2.5
6 ? 1.5
13 t 4.3
10 ? 2.0
5 t 1.2
1 3 & 3.1
7 -c 1.1
6 +- 3.0
4 3 1.3
9 ? 4.0
-to. 392)
6 2 2.2
3 +- 1.5
8 +- 3.2
IN,number of measurements Tturly animals were used
'Mean values in micra 2 S D
'S E of differences between means
None were tound In histologic sections
locally regulating the flow of blood through
individual capillaries. The contraction or
bulging of the nuclear regions into the
lumens of capillaries alters pressure/flow
gradients leading to reversal of flow in a
manner similar to that reported in the living liver (McCuskey, '66, '68a) and spleen
(Bloch, '68). As in the above organs, endothelial cells of the pancreatic capillaries
are responsive to various vasoactive hormones and metabolites (McCuskey and
Chapman, '68). While in acinar tissue,
precapillary smooth muscle sphincters
regulate the flow of blood from arterioles
into capillaries, these sphincters have not
been observed at the arteriolar-capillary
junction in islets; more precise control of
flow through individual capillaries in the
islets is provided by the endothelial
sphincters. Other evidence that endothelial cells may be contractile is suggested by the presence of fibrils within the
cytoplasm (Zelander et al., '62; Puchtler et
al., '68) and the immunofluorescent
demonstration of contractile protein in
such cells (Becker and Murphy, '69).
Several morphological features of the
pancreas, the existence of which have
been questioned in the past, have been
confirmed by this investigation. (1) Occasional capillary anastomoses occur
between the capillary bed of the islet and
that of the acinar tissue, as reported by
Kuhne and Lea (1882), Berg ('30), Beck
and Berg ('31), Wharton ('32), and Ferner
('52). The infrequency with which they are
seen and their small size may explain why
their presence was not detected in viuo by
Brunfeldt et al. ('58) and Bunnag et al.
('63). Also, the failure to identify them in
plastic injected or fixed material by Brun-
feldt et al. ('58) may be due to the tendency
of the small vessels of the pancreas to
shrink appreciably during fixation. (2)
Intercellular secretory canaliculi (:Bensley,
'11; Opie, '32; Ito, '66) between adjacent
acinar cells exist in life and are not fixation
artifacts. These pass between ce:ntro-acinar cells to reach the lumen of the intralobular ducts and provide the anatomic
pathway for acinar secretory products to
reach the lumen. (3) Processes of beta cells
may extend between two adjacent cells.
These appear to provide additional. surface
area for transcapillary exchange since
beta granules are present in these areas.
(4) A vascular pedicle connects islets to the
ducts and contains spindle-shaped cells,
the cytology of which somewhat resembles
the duct cells; some, however, contain
granules similar to those in beta cells.
Such cells appear similar to those described by Bunnag ('66) who reported them
to be precursors of islet cells.
The morphologic and temporal sequence
of events during the formation and release
of zymogen granules in individual acinar
cells corresponded well with those reported
by Warshawsky et al. ('63) using autoradiographic methods, and with electron
microscopic studies (Palade, '59; Palade et
al., '61; Sjostrand, '61). In contrast to the
report of Covell('28), no zymogen granules
were seen outside the ceil; these disappeared upon contract with the cell membrane as reported by Kiihne and Lea (1882)
and Junqueira and Hirsch ('56). The formation and release of granules in beta
cells was more rapid than that in acinar
cells and followed a sequence similar to
that described by Lacy ('67). In agreement
with the report by Lacy ('67),
no secretory
granules were found outside the cell; the
granules seemed to disappear at the membrane of the cells as has been reported
from other electron microscopic studies
(Lacy et al., '59; Williamson et al., '61;
Lazarus and Volk, '62; Volk and Lazarus,
Humanson, G. L. 1967 Animal TissueTechniques. W.
H. Freeman, San Francisco, pp, 301-340.
Ito, S. 1966 Pancreas. In: Histology. R. 0. Greep, ed.
2nd edition, McGraw-Hill, New York, pp. 554-567.
Junqueira, L. C. U. and G. C. Hirsch 1956 Cell secretion: A study of pancreas and salivary glands. Intern.
Rev. Cytol., 5: 323-364.
Lacy, P. E. 1967 The pancreatic beta cell. New Eng.
J. Med., 276: 187-194.
Lacy, P. E., A. F. Cardez and W. D. Wilson 1959
Electron microscopy of the rat pancreas -effects of
glucagon administration. Diabetes, 8: 36-44.
The authors wish to thank Marian L.
S. S. and €3. W. Volk 1962 Ultramicroscopic
Miller and Rosemary Schulte for their Lazarus,
and histochemical studies on pancreatic beta cells
illustrative and technical assistance,
stimulated by tolbutamide. Diabetes, I I : 2-1 1.
Mathews, A. 1899 The changes in the structure of
some aspects of cell metabolism. J. Morph., 15:
(Suppl.) 171-222.
McCuskey, R. S. 1966 A dynamic and static study of
hepatic arterioles and hepatic sphincters. Amer J.
Beck, J. S. P. and B. N. Berg 1931 The circulatory
Anat., 119: 455-478.
pattern in the islands of Langerhans. Amer. J. Path.,
__ 1968a Microscopy of the living liver in situ.
7: 31-36.
Anat. Rec., 160: 521.
Becker, C. G., and G. E. Murphy 1969 Demonstration
__ 1968b Dynamic microscopic anatomy of the
of contractile protein in endothelium and cells of the
fetal liver. 111. Erythropoiesis. Anat. Rec., 161:
heart valves, endocardium, intima, arteriosclerotic
plaques and Aschoff bodies of rheumatic heart disease. Amer. J. Path., 55: 1-38.
McCuskey, R. S., andT. M. Chapman 1967a MicrosBerg, B. N. 1930 A study of the islands of Langerhans
copic anatomy of the living pancreas in situ. Anat.
in viuu with observations on the circulation. Amer. J.
Rec., 157: 407-408.
Physiol., 95: 186-189.
- 1967b Microscopic observations on the secreBloch, E. H. 1965 The dynamic histologyof organs i n
tory process in cells of the living pancreas in situ.
situ. 11. The lung, liver and kidney. Anat. Rec., 151:
Midwest Anatomists Meeting, Ann Arbor, Nov. 11.
(Motion Picture).
__ 1966 Television microphotometry of organs __ 1968 Microcirculation of the pancreas. XVI
Microcirculation Conference, Atlantic City, April 14.
in situ.Methods in Med. Res., 1 1 : 228-238.
(Motion Picture).
- 1968 Microanatomy of living mammalian
O'Leary, J. L. 1930 An experimental study of the islet
spleens in situ. I. Endothelium. Anat. Rec., 160: 520.
cells of the pancreas in uivo. Anat. Rec., 45: 27-58.
Bloch, E. H., and R. S. McCuskey 1966 Sequential
Opie, E. L. 1932 Cytology of the pancreas. In: Special
quantitative measurements in microseconds of living
Cytology. E. V. Cowdry, ed. 2nd edition, Hoeber, New
tissue. Anat. Rec., 154: 507-508.
York, Vol. 1, pp. 241-271.
Brunfeldt, K., K. Hunhammer and A. P. Skoupy 1958
Studies on the vascular system of the islets of Lan- Palade, G. E. 1959 Subcellulur Particles. P. Hayashi,
ed. Ronald Press, New York, pp. 6468.
gerhans in mice. Acta Endocr., 29: 47S480.
Palade, G. E., P. Siekevitz and L. G. Car0 1961 StrucBunnag, S. C. 1966 Postnatal neogenesis of islets of
ture, chemistry and function of the pancreatic exoLangerhans in the mouse. Diabetes, 15: 480-491.
crine cell. In: The Exocrine Pancreas. A. V. S. de
Bunnag, S. C., S. Bunnag and N. E. Warner 1963
Reuck and M. P. Cameron, ed. Little, Brown and Co.,
Microcirculation in the islets of Langerhans of the
Boston, pp. 23-49.
mouse. Anat. Rec., 146: 117-124.
Bunnag, S. C., N. E. Warner and S. Bunnag 1966 Palmer, A. A. 1959 A study of blood flow in minute
of the pancreatic region of the rat with referEffect of tolbutamide on postnatal neogenesis of the
ence to intermittent corpuscular flow in individual
islets of Langerhans in mouse. Diabetes, 15:
capillaries. Quart. J. Exper. Physiol., 44: 149-159.
- 1967 Effect of alloxan on the mouse pancreas Palmer, A. A., and A. G. Wilkins 1960 A closed
method for the microscopic study of the living rat
during and after recovery from diabetes. Diabetes,
pancreas in situ. Austral. J. Exp. Biol. Med., 38:
16: 83-89.
Covell, W. P. 1928 A microscopic study of pancreatic
secretions in the living animal. Anat. Rec., 40: Puchtler, H., F. Sweat, M. S. Terry and H. M. Conner
1968 Investigation of staining, polarization and
fluorescence - microscopic properties of myoendoFerner, H. 1952 Das Inselsystem des Pankreas. Stuttthelial cells. J. Microscopy, 89: 95-104.
gart, George Thieme Verlag.
Heisig, N. 1967a Neue Methode zur Intravitalmikros- Ruangsiri, C. 1949 Changes in islets of Langerhans
in living mice after alloxan administration. Anat.
kopie der terminalen Strombahn des Pankreas mit
Rec., 105: 399-419.
einer optischen Kammer. Z. ges. exper. Med., 142:
Sjostrand, F. S. 1961 The fine structure of the exo- 1967b Pancreatic microcirculation under the crine pancreas cells. In: The Exocrine Pancreas. A.
influence of adequate secretory stimulation. BiblioV. S. de Reuck and M. P. Cameron, ed. Little, Brown
theca Anat., 9: 176-180.
and Co., Boston, pp. 1-19.
__ 1968 Functional analysis of the microcircula- Volk, B. W., and S. S. Lazarus 1963 Ultramicroscopic
tion in the exocrine pancreas. Adv. Microcirc.. 1:
studies of rabbit aancreas durina cortisone treat89-151.
ment. Diabetes. 12: 162-173.
Warshawsky, H., C. P. Leblond and B. Droz 1963
Synthesis and migration of proteins in the cells of the
exocrine pancreas as revealed by specific activity
determination from radioautographs. J. Cell Biol., 16:
Williamson, J. R., P. E. Lacy and J. W. Grisham 1961
Ultrastructural changes in the islets of' the rat
produced by tolbutamide. Diabetes 10: 460-469.
Zelander, R., R. Ekholm and Y. Edlund 19682 Ultrastructural organization of the rat exocrine pancreas.
111. Intralobular vessels and nerves. Ultrastruct.
Res., 7:84-101.
Zimmermann, K. W. 1927 Die Speicheldrussen der
Mundhohle und die Bauchspeicheldrusse. In: Handbuch microscop. Anat. Menschen. W. Vnn Mollendorff, ed. Val. 5, pt. 1, Springer, Berlin, pp. 215-228.
5 Low power view of acinar tissue (A) and an islet (I) transilluminated with
light a t 550 mp illustrating the numerous capillary loops in the islet and the
gross difference in appearance between the acinar and islet tissue. C, capillary;
ct, connective tissue capsule of islet. From single frame of motion picture.
6 Same as figure 5 except the tissue was transilluminated with light at 414 m p
(absorption peak for hemoglobin) to demonstrate the insular-acinar anastomosis
(IAA). From single frame of motion picture.
7 Acinus (A) with adjacent capillary (c) and nucleus of endothelial cell (e). In
contrast to the islet, capillaries are separated widely, being confined to the
periphery of the acinus. From single frame of motion picture.
8 Bulging endothelial cell (e) in wall of a capillary (C) in a pancreatic islet. B,
beta cell. From single frame of motion picture.
Capillary (C) in acinar tissue illustrating the contraction of endothelial cells
(E, and EJ: (a) initial appearance; (b) 30 seconds after topical application of
1 y norepinephrine. From single frames of motion picture.
R. S. McCuskcy and T. M. Chapman
10 Binucleated acinar cell and adjacent capillary. Note zymogen granules, 2 ;
nucleus, N ; mitochondrion, m; capillary, c; endothelial cell, e. From single
frame of motion picture.
Acinar cell with prozymogen granules (PZ). Z, zymogen granules; A, apex of
cell. From single frame of motion picture.
12 Beta cells (B) in pancreatic islet. Note nucleus, N; beta granules, g; capillary,
C ; endothelial cell, e. From single frame of motion picture.
K. S McCuskev and T. M Chapman
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living, microscopy, pancreas, situ
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