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Notes on the cytology of the parietal cells of the stomach of the rat.

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NOTES ON T H E CYTOLOGY O F T H E PARIETAL
CELLS O F THE STOMACH O F THE RAT
H. W. BEAMS AND R. L. KING
Department of Zoology, State University of Iowa
TWO PLATES ( N I N E FIGURES)
Intracellular secretory canaliculi were demonstrated in the
parietal cells of the stomach of mammals long ago by the
beautiful researches of Miiller ( '98), Bensley ( '98), and
Zimmermann ( '98). I n their work these investigators employed the chrome-silver technique, which remains to-day in
many respects the best and most desirable method for the
study of intracellular secretory canaliculi. It is gratifying
to note that the essential facts recorded by the above-named
investigators as regards intracellular canaliculi have been
confirmed by the more recent investigations on living tissue
by Harvey and Bensley ( '12). Accordingly, it is our purpose
in this brief note to simply reemphasize the presence of intracellular canaliculi in the parietal cells of the stomach and to
record the topographical relationships of the mitochondria
and Golgi material to them. Furthermore, the suggestion is
made that the occurrence of intracellular canaliculi in gland
cells is directly correlated with a more or less specific type
of secretion.
MATERIAL AND METHODS
The material for this study consists of portions of the
pyloric region of the stomach of the rat, prepared according
to the method of Regaud for mitochondria, Mann-Kopsch
(Weigl) and Nassonov for Golgi apparatus, and the rapid
chrome-silver method of Golgi f o r the demonstration of intracellular secretory canaliculi. Tissue impregmated by the
31
THE ANATOMICAL RECORD, VOL. 53, NO. 1
32
H. W. BEAMS AND R. L. KING
chrome-silver method of Golgi was subsequently stained with
Delafield’s hematoxylin and eosin or by the method of
Delaney ( ’27).
It was found important t o stain only a very short time in
Delafield’s hematoxylin, as it seemed to have a marked
destructive effect on the silver impregnation.
Vital studies were also made on the parietal cells by use
of neutral red and Janus green B.
DESCRIPTION
Since the general histology of the gastric glands is so well
known, it is not necessary for us to enter into a detailed
description here, It will suffice to point out that, in general,
the parietal cells are found distributed throughout the whole
length of the gastric glands, but are more numerous in the
neck and body than in the fundus. They are for the most
part larger than the zymogenic cells and usually possess a
spherical or triangular form, with the base of the cell resting
directly on the basement membrane of the gland.
I n an unstained supravital preparation the parietal cells
appear slightly granular (mitochondria), but the mitochondria here are not so prominent as the secretion granules in
the chief cells. However, following routine histological technique the parietal cells often appear more or less homogeneous with a strikingly acidophilic cytoplasm. I n successful
preparations of the Golgi rapid chrome-silver method, one
finds that perhaps the most marked feature of the parietal
cells is the rather complicated system of secretory canaliculi
(figs. 1, 2, 6, and 7). These canaliculi form a basket-like
closed rete of very fine tubules which occupies a position in
the cytoplasm between the nucleus and the cell periphery. At
the inner or luminal end of the cell the basket-like network
of intracellular canaliculi penetrate the wall of the cell and
become continuous with one or more efferent ductules which
extend between the clefts of adjacent zymogenic cells to connect with the main lumen of the gland. Hence, it is through
this system of canaliculi that the secretion of the parietal
cells passes into the lumen of the gland.
PARIETAL
CELLS OF THE RAT'S STOMACH
33
It is the general opinion of some investigators that metallic
impregnation methods are quite generally unreliable and so
give rise to artifacts. Accordingly, they have viewed the
intracellular canaliculi with considerable skepticism until the
important work of Fitzgerald ('lo) and Harvey and Bensley
('la) on living tissue. We have confirmed Harvey and
Bensley's results by use of vital methods and, like them, have
found the intracellular canaliculi to stain readily with neutral
red, in which case, their general character is found to exactly
coincide with that demonstrated by the chrome-silver method
of Golgi. This evidence on the study of the intracellular
canaliculi of the parietal cells leads us to think that they
compose a permanent system of secretory canaliculi, and are
not transitory structures as has been claimed by some.
Furthermore, since the work of Harvey and Bensley has
shown the secretion of the parietal cells to be of a highly
viscid nature and since this is also the case for the secretion
of certain other cells which contain intracellular secretory
canaliculi (St. Hilaire, '27), the suggestion seems plausible
that the presence of intracellular canaliculi is in some way
associated with a highly viscid form of secretion of low
diffusibility.
Recently the presence of intracellular canaliculi has been
described in the hepatic cells of amphibians by Pollister ( '31)
and by Dawson ('31). These investigators have shown that
the Golgi apparatus lies in close juxtaposition to the intraand intercellular canaliculi-a relationship they hold to be
circumstantial evidence that this cell component is in some
way concerned with the synthesis of secretory products.
I n preparations of the parietal cells prepared according to
the osmic-acid impregnation methods of Nassonov and MannKopsch (Weigl), one finds that the intracellular canaliculi
appear clear or light brown in contrast to the dull gray cytoplasm. I n those cases where the canaliculi are filled with
secretion they appear light brown which is comparable to the
'Pollister has recently (Am. J. Anat., vol. 50, no. 2) decided that the canaliculi
previously reported by him are not intracellular.
34
H. W. BEAMS AND R. L. KING
appearance of the secretion in the lumen of the gland. While
the intracellular secretory canaliculi cannot be studied in
great detail following osmification, they are nevertheless
sufficiently clear to make a careful study of their relationship
to the Golgi apparatus. The Golgi material in the parietal
cells is composed chiefly of isolated filaments, which rarely
form a network, and are scattered fairly generally throughout the cytoplasm with perhaps a slight concentration in some
instances around the nucleus (fig. 4 ) . I n general we were
never able to find as complete a network of the Golgi material
as that reported by Kolster ( ’13). However, we are not in
agreement with Siraska (’29) and Shirasaka (’30) that the
Golgi material does not exist in the parietal cells. Furthermore, Siraska ’s assumption that the absence of Golgi material
in the parietal cells might be associated with the secretion of
hydrochloric acid is not in harmony with the findings of
Harvey and Bensley that the whole system of intracellular
canaliculi when stained with neutral red “is without question
on the alkaline side of the reaction.”
Unlike the Golgi apparatus in the liver cells, no concentration around the intracellular secretory canaliculi is apparent.
This condition agrees with the recent findings of Beams and
King (’32 b) for the parietal cells of the salivary glands of
certain Orthoptera. I n addition, there seems to be no welldefined secretogenous zone in the parietal cells comparable
to that concerned with the secretion of bile in the liver cell
(Pollister, Dawson). Nevertheless, this fact does not remove
the possibility that the close topographical relationship such
as occurs between the Golgi material and the intracellular
bile canaliculi may not be of general significance.
It is of considerable interest to note that in the parietal
cells of the stomach the walls of the intracellular canaliculi
are not usually blackened by osmic-acid treatment. This
condition is in contrast with that which exists in certain
Protozoa (contractile vacuole, Nassonov, ’24) and in the
parietal cells of the salivary glands of the grasshopper
(Beams and King, ’32 b). Furthermore, the membrane sur-
PARIETAL CELLS OF THE RAT'S
STOMACH
35
rounding the intracellular canaliculi of the parietal cells of
the stomach cannot be differentiated so clearly as that in the
parietal cells of the grasshopper (Beams and King, '32 b).
The apparent presence here of two different membranes
separating the cell from its surroundings seems to be comparable to the view of Bayliss ('20) that if "gland cells are
possessed of a membrane on the end next to the blood vessel
of such kind as to be impermeable to some substances produced in the cell, while on the ends next the duct the membrane is permeable to those substances, we can account for
a flow of water as long as the osmotically active substances
are being formed. ''
When the parietal cells are examined in the fresh supravital preparations, one finds them loaded with small bodies
of spherical or rectangular form, and of low refractive index,
which stain readily with Janus green B. These granules
were accurately described long ago by Altmann ( '94) whose
excellent figures, as pointed out by Bensley ( '28)' have not
been improved upon by the various mitochondria1 researches
which have followed. Other investigators, notably No11 and
Sokoloff ( '05), Eklof ( '14), Hoven ( '12), and Lim and Ma
( '26) agree that these granules represent mitochondria.
I n preparations made according to the technique of Regaud
f o r mitochondria one finds the intracellular canaliculi appearing as clear spaces (which might easily be mistaken for the
canalicular apparatus of Bensley and Cowdry or the Saftkanalchen of Holmgren) with the mitochondria deeply stained
(figs. 5, 8, and 9). The size and number of the mitochondria
seem somewhat variable, but they appear not to be concentrated around the intracellular canaliculi. Their form seems
to vary from a short rod-like to a spherical condition.
Whether or not this variation in form of the mitochondria
can be interpreted to have a functional significance, such as
the formation of secretion, is not definitely known. No direct
morphological relationship between the mitochondria and
Golgi material as held by Parat ( '29) and Hosselet ('31) was
ever observed by us.
36
H. W. BEAMS A N D R. L. KING
CONCLUSIONS
1. Intracellular secretory canaliculi similar to those described by Muller, Bensley, and Zimmermann have been
demonstrated in the parietal cells of the stomach of the rat.
They appear as a complicated network of canaliculi which
surround the nucleus and penetrate the greater part of the
cytoplasm. The intracellular canaliculi seem to be permanent
structures in the parietal cells and communicate directly with
the lumen of the gland by one or more efferent ductules.
2. The Golgi material appears as robust filaments, which
seldom form a network and are scattered throughout the
cytoplasm of the cell. There is no marked concentration of
the Golgi material around the intracellular secretory canaliculi. This seems to indicate that the close topographical relationship reported to exist between the Golgi apparatus and
intracellular bile canaliculi ( Pollister, Dawson) is not generally significant from the point of view of secretion.
3. The mitochondria appear generally as coarse spherical
bodies distributed abundantly throughout the cytoplasm.
They show no obvious relation to the intracellular canaliculi.
Furthermore, there is no evidence of the hypertrophy of mitochondria into Golgi material (‘inactive’ as opposed to ‘active’
mitochondria) in the manner as urged by Parat and others.
LITERATURE CITED
ALTMANN,R. 1894 Die Elementarorganismen und ihre Beziehungen zu den
Zelleu. Leipzig: Verlag Von Veit & Comp.
BAYLISS,W. M. 1920 Principles of general physiology. Longmans, Green L%
Co., N. Y.
BEAMS,H. W., AND R. L. KING 1932a Cytoplasmic structures in the ganglion
cells of certain Orthoptera, with special reference to the intracellular
canaliculi, Golgi bodies, and mitochondria. J. Morph., vol. 53.
1932 b The architecture of the parietal cells of the salivary glands
of the grasshopper, with special ref erence t o the intracellular canaliculi,
Golgi bodies, and mitochondria. J. Morph., vol. 53.
BENSLEY,R. R. 1898 The structure of the mammalian gastric glands. Quart.
J. Micr. Sci., vol. 41.
1928 The gastric glands. Special Cytology, vol. 1. New York.
DAWSON,A. B. 1931 The hepatic cells of Neeturns. Anat. Rec., vol. 51
(abstract).
DELANEY,P. A. 1927 Reliable methods for the fixation and staining of Nissl
substance. Anat. Rec., vol. 36.
PARIETAL
EKLOF, H.
CELLS OF THE RAT’S
STOMACH
37
1914 Chondriosomenstudien a n den Epithel und Driisenzellen bei
Saugetieren. Anat. Hefte, Bd. 51.
FITZOW, M. 1’. 1910 The origin of hydrochloric acid i n the gastric tubules.
Proc. Roy. Soc., B, vol. 83.
HARVEY,
B. C. H., AND 1z. R. BENSLEY1912 Upon the formation of the hydrochloric acid in the foveolae and on the surface of the gastric mucous
membrane, and the non-acid character of the gland cells a n d lumina.
Biol. Bull., vol. 23.
HOSSELET,C. 1931 Contribution A 1’6tude du chondriome chez les insectes
(Culieides et Phryganides). Arch. de Zool. exp6r. et gBnBr., T. 72.
HOVEN,
H. 1912 Contribution 3i 1’6tude du fonctionnement des cellules glandulaires. Du rble du chondriome dans la secretion. Arch. f . Zellf., Bd. 8.
KOLSTER,R. 1913 Uber die durch Golgi’s Arsenik- und Cajal’s UrannitratSilbermethode darstellbaren Zellstrukturen. Anat. Anz., Bd. 44 (supplement).
LIM, R. K. S., AND W. C. MA 1926 Mitochondria1 changes in the cells of the
gastric glands in relation to activity. Quart. J. Exper. Physiol.,
vol. 16.
11. Uber die Fundusdriisen des Magens.
MULLER, E. 1898 Drusenstudien.
Ztschr. f. wiss. Zool., Bd. 64.
NASSONOV,D. N. 1924 Das Exkretionsapparat (kontraktilc Vakuole) der
Protozoa als Homologen des Golgischen Apparats der Metazoazellen.
Arch. f . mikr. Anat., Bd. 103.
1905 Zur Histologie der ruhenden und thatigen
NOLL, A., AND A. SOKOLOFF
Fundusdriisen des Magens. Arch. f . Anat. u. Physiol., Bd. 94.
PAL4T, M. 1928 Contribution A 1’Btude morphologique e t physiologique du
cytoplasme ; chondriome, vacuome (appareil de Golgi) , enclaves, etc.
Arch. d’Anat. Micr., T. 24.
1929 Le chondriome actif de la cellule animale et le phBnoni6ne
de pachynese. C. R. Acad. Sc., T. 188.
POLLISTER,
A. W. 1931 The architecture of the hepatic cells of Amphiuma.
The Collecting Net, vol. 6.
SAINT-HILAIAIRE,
K. 1927 Histo-Physiologische Studieii iiber die Spinndriisen der
Tenthredinideiilarven. Zeit. f . Zellf. u. mikr. Anat., Bd. 5 .
SHIRASAKA,M. 1930 Studien iiber die morphologisehen Veranderungen des
Golgischen Apparatus in den Zellen des Verdauungssystems des
Kaninchens infolge der Darreichung von K-, Ca-, und Mg Salzen.
Folia Anat. Japonica, Bd. 8.
SIRASKA,S . 1929 The existence or non-existence of Golgi7s network in delomorphous cells of the gastric glands (in Japanese).
Nippon
Shokwakibyogakukmai Zasshi, vol. 28. (Taken from Biol. Absts.,
vol. 5, no. 1.)
ZIXHERMANN, Ti. W. 1898 Beitrage zur Eenntnis einiger Drusen und Epithelien.
Arch. f. mikr. Anat., Bd. 52.
PLATE 1
EXPLANATION OF FIGURES
1 Cross-section of gastric gland showing intracellular canaliculi connceted by
ductules to the lumen. (Golgi’s rapid method.)
2 Longitudinal section of gastric gland, as above.
3 Zymogenic cells showing Golgi apparatus and secretion granules in luminal
end of cell. (Modified Mann-Kopsch.)
4 Parietal cell with intracellular canaliculi clear and Golgi apparatus impregnated. (Modified Mann-Kopsch.)
5 Longitudinal section of gastric gland showing granular mitochondria and
clear intracellular canaliculi. (Regaud.)
38
PLATE 1
PARIETBL CELLS O F THE RAT'S STOMACH
1%.w. BEAXIS .<NU R . L. KING
39
PLATE 2
EXPLANATION O F FIGURES
6 Photomicrograph
7 Photomicrograph
8 Photomicrograph
9 Photomicrograph
ehondria. X 2750.
X 375.
as in figure 1. x 375.
as in figure 5. X 750.
highly magnified to show clear canaliculi among the mito
as in figure 2.
40
PARIETAL CELLS O F T H E RAT’S STOMACH
1%. W. BEAJIS AND R . L. ICING
41
PLATE 2
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