Glycoconjugate histochemistry of the rat fundic gland using Griffonia simplicifolia agglutinin-II during the development

The development and maturation of fundic glands of Wistar rats were studied using Griffonia simplicifolia agglutinin-II (GSA-II) histochemistry at the light microscopic and electron microscopic levels. In adult rats, mucous neck cells and cells intermediate between mucous neck cells and chief cells were specifically labeled with GSA-II, whereas other fundic gland cells were virtually negative. Ontogenetic studies revealed that GSA-II positive cells appeared at the bottom of the gland by 21 days of gestation. With differentiation and aging, the elongation of the fundic gland continued, and the labeling intensity of the mucous neck cells increased by 3 weeks after birth. Cells intermediate between mucous neck cells and chief cells were discernible from 3 days after birth. Typical mucous neck cells appeared at 3 weeks after birth, when their labeling intensity with colloidal gold (CG) particles approximated that of adults. On the other hand, the reactive cell population gradually moved from the bottom toward the middle portion of the gland. Finally, the reactive cells were localized at the neck portion of the fundic gland. These results suggest that GSA-II is a valuable marker for studying mucous neck cells and both their precursor cells and their derivatives.     

Glycoconjugate cytochemistry of the rat fundic gland using lectin/colloidal-gold conjugates and Lowicryl K4M

Summary The fundic gland of the rat stomach was studied using the low-temperature embedding resin Lowicryl K4M and postembedding staining with lectin/colloidal-gold (CG) conjugates. Intense labeling with Ricinus communis agglutinin I was observed not only in mucous-producing cells but also in parietal cells. In contrast, Helix pomatia agglutinin (HPA) only labeled mucous neck cells and intermediate cells between mucous neck cells and chief cells. The other epithelial cells present in the rat fundic gland showed virtually no reaction with this lectin. Our results indicate that HPA might be a marker lectin of mucous neck cells and their derivatives. The combination of embedding in the hydrophilic resin Lowicryl K4M and postembedding staining with lectin-CG conjugates provided satisfactory staining results, and made it possible to visualize the precise distribution of terminal glycoconjugates in intracellular components as well as on the plasma membrane.     

Glycoconjugate cytochemistry of the rat fundic gland using lectin/colloidal-gold conjugates and Lowicryl K4M. Helix pomatia lectin is a specific marker for mucous neck cells in fundic glands of the rat gastric mucosa

The fundic gland of the rat stomach was studied using the low-temperature embedding resin Lowicryl K4M and postembedding staining with lectin/colloidal-gold (CG) conjugates. Intense labeling with Ricinus communis agglutinin I was observed not only in mucous-producing cells but also in parietal cells. In contrast, Helix pomatia agglutinin (HPA) only labeled mucous neck cells and intermediate cells between mucous neck cells and chief cells. The other epithelial cells present in the rat fundic gland showed virtually no reaction with this lectin. Our results indicate that HPA might be a marker lectin of mucous neck cells and their derivatives. The combination of embedding in the hydrophilic resin Lowicryl K4M and postembedding staining with lectin-CG conjugates provided satisfactory staining results, and made it possible to visualize the precise distribution of terminal glycoconjugates in intracellular components as well as on the plasma membrane.     

Immunocytochemical study of pepsinogen 1-producing cells in the fundic mucosa of the stomach in developing mice

Summary Development and maturation of pepsinogen 1-producing cells were studied in the gastric fundic mucosa of the mouse by means of light- and electron-microscopic immunocytochemistry using rabbit anti-rat pepsinogen 1-serum. In the adult mouse, secretory granules in mucous neck cells, transitional mucous neck/chief cells and chief cells are immunolabeled. The numerical density of gold particles on zymogen granules is not significantly altered among different stages of maturation of chief cells. In addition, rough endoplasmic reticulum and Golgi complex of these cell types show a weak labeling. In mice from day 16 of gestation to postnatal day 14 mucous neck cells and chief cells cannot be distinguished, but only one type of pepsinogen 1-producing cell, called primitive chief cell, is identified in the fundic gland. The intensity of immunoreactivity of secretory granules in primitive chief cells is uniform within an individual cell but varies greatly among different cells. The majority of primitive chief cells contains weakly labeled granules regardless of the maturation stage of cells or of animals. On postnatal day 21, mucous neck, transitional and chief cells are distinguishable, and secretory granules in these cells are intensely immunolabeled as in the adult. These results suggest that pepsinogen 1-production rapidly increases with differentiation of mucouse neck and chief cells.     

Fine structure of the oxynticopeptic cells in the gastric glands of the ruin lizard, Podarcis sicula campestris De Betta, 1857

The results of an ultrastructural investigation of the gastric glands of the ruin lizard are reported. In this reptile the stomach can be divided into a larger fundus and a smaller pars pilorica. Fundic glands are characterized by three main kinds of cells: mucous, endocrine, and oxynticopeptic; the latter were not observed in the pyloric glands. The morphological features of the oxynticopeptic cells change from the proximal to the distal region of the fundic mucosa. In the proximal region, numerous electron-dense secretory granules, a well-developed granular endoplasmic reticulum, an evident Golgi complex, and a reduced system of smooth-surfaced vesicles and tubules in the apical cytoplasm characterize these cells. In the distal fundic region, oxynticopeptic cells possessed numerous mitochondria and a well-developed smooth-surfaced endoplasmic reticulum, but secretory granules were rare. These data suggest the existence of a gradient in the production of proteolytic enzymes, and perhaps also of hydrochloric acid, along the oral-aboral axis of the stomach. The results are discussed with regard to the evolution of the gastric glands and of the digestive mechanism in vertebrates.     

Ultrastructure of the cardiac and pyloric glands of the gastric mucosa of the South African hedgehog,Atelerix frontalis

The cardiac and pyloric glands in the gastric mucosa of the South African hedgehog, Atelerix frontalis, are described. The cardiac area of the stomach contains proper cardiac glands and lacks undifferentiated fundic glands. The cardiac glands are simple tubular, coiled, and lined with columnar cells ultrastructurally similar to those of the gastric surface epithelium. Secretory granules with varying electron densities fill the apical cytoplasm of these cells. In contrast to other mammals, these glands lack mucous neck cells. The neck of the pyloric glands contains only a single cell type, whereas the basal regions of these glands contain “light” and “dark” cells. The secretory granules in the “dark” cells and the pyloric neck cells have a moderate electron density and often contain an electron dense core. An electron-lucent cytoplasm with numerous polysomes is characteristic of the “light” cells. Some “light” cells contain electron-dense granules in the apical cytoplasm. The presence of only neutral mucins in the cardiac gland cells denotes the absence of mucous neck cells. The acidic mucins within the pyloric neck cells seem to indicate that these cells are mucous neck cells, whereas the neutral mucins within the basally located pyloric gland cells show at least a partial functional difference from the pyloric neck cells. © 1993 Wiley-Liss, Inc.     

Immunocytochemistry and in situ hybridization studies of pepsinogen C-producing cells in developing rat fundic glands

The ontogeny of pepsinogen C-producing cells in rat fundic glands was studied by means of light and electron microscopy using an antiserum raised against a synthetic peptide based on rat pepsinogen C. To confirm the immunocytochemistry results, the expression of rat pepsinogen C messenger RNA (mRNA) in the fundic gland was also examined by in situ hybridization using a digoxigenin-labeled RNA probe. In adult rats, pepsinogen C was produced by chief cells, mucous neck cells, and intermediate mucopeptic cells. Pepsinogen C-producing cells appeared in embryos as early as 18.5 days’ gestation. The development of these cells could be classified into four stages: (1) 18.5 days’ gestation to 0.5 days after birth; (2) 0.5 days to 2 weeks after birth; (3) 3–4 weeks after birth; (4) 4–8 weeks after birth. In embryos and young animals, pepsinogen C-producing cells were mucopeptic cells. By 4 weeks after birth, mucous neck cells could be distinguished morphologically. The maturation stages of the chief cells could be traced by electron microscopy along the longitudinal axis of the rat fundic gland by double-staining with anti-pepsinogen C antibody and periodic acid-thiocarbohydrazide-silver proteinate. Positive reactions for pepsinogen C and pepsinogen C mRNA expression were detected in mucous neck cells. Therefore, we conclude that mucous neck cells are precursor cells of chief cells. Mucous neck cells, intermediate cells, and chief cells are in the same differentiating cell lineage.     

Self-renewal of the human gastric epithelium: new insights from expression profiling using laser microdissection

The gastric mucosa is subject to continual bidirectional renewal by differentiation from stem and transit amplifying cells. It was the aim of this study to characterize the self-renewal of the human gastric mucosa and its two major types of glands in the fundus and antrum, respectively. Three characteristic regions (pit, proliferative, and lower neck regions) were isolated from fundic and antral units by the use of laser microdissection, and expression profiles concerning 15 marker genes were generated by RT-PCR analysis. The surface mucous cells (SMCs) of fundic and antral units differed in their expression of at least four secretory genes, i.e., gastric lipase, TFF3, FCGBP, and lysozyme. The maturation of mucous neck cells was shown to occur stepwise, first towards a mucous phenotype followed by a serous differentiation step. Also, a stepwise maturation of both the antral SMCs and antral gland cells was observed. Additionally, the presence of gastric lipase was also demonstrated for the first time in antral gland cells. In conclusion, the different expression profiles of SMCs of the fundic and antral units could be the basis for the different self-renewal rates of fundic and antral SMCs and could influence the spatial organization of the bacterial microbiota within the various parts of the gastric mucosa.     

Expression of N-acetylglucosamine residues in developing rat fundic gland cells

The development of rat fundic gland was studied by immunohistochemistry using a recently developed monoclonal antibody, HIK 1083, at both light and electron microscope levels. Antibody HIK 1083 recognized oligosaccharides with a non-reducing terminal -linked N-acetylglucosamine (GlcNAc) residue. In the developing rat fundic gland, cells expressing -GlcNAc residues were discernible from day 19.5 of gestation and continued to exist till adult. The distribution of the -GlcNAc expressing cells was consistent with that described previously for cells reacting to Griffonia simplicifolia lectin (GSA-II) in all developmental stages. These cells were located at the bottom of the fundic gland when they first appeared. With the elongation and maturation of the gland, these cells moved upwards and were finally restricted in the neck region of the gland. Combining previous reports and the present electron microscopical observations, HIK 1083-positive cells in the adult rat fundic gland are mucous neck cells. The interaction between antibody HIK 1083 and GSA-II lectin was investigated. GSA-II prevented the subsequent binding of HIK 1083, while HIK 1083 did not prevent GSA-II binding to mucous neck cells. Our results suggested that -GlcNAc residues exist in rat fundic gland from day 19.5 of gestation and continue to exist till adult. Cells expressing -GlcNAc residues appeared as typical mucous neck cells from postnatal four weeks.     

Immunolocalization patterns of cytokeratins during salivary acinar cell development in mice

Embryonic development of the mouse salivary glands begins with epithelial thickening and continues with sequential changes from the pre-bud to terminal bud stages. After birth, morphogenesis proceeds, and the glands develop into a highly branched epithelial structure that terminates with saliva-producing acinar cells at the adult stage. Acinar cells derived from the epithelium are differentiated into serous, mucous, and seromucous types. During differentiation, cytokeratins, intermediate filaments found in most epithelial cells, play vital roles. Although the localization patterns and developmental roles of cytokeratins in different epithelial organs, including the mammary glands, circumvallate papilla, and sweat glands, have been well studied, their stage-specific localization and morphogenetic roles during salivary gland development have yet to be elucidated. Therefore, the aim of this study was to determine the stage and acinar cell type-specific localization pattern of cytokeratins 4, 5, 7, 8, 13, 14, 18, and 19 in the major salivary glands (submandibular, sublingual, and parotid glands) of the mouse at the E15.5, PN0, PN10, and adult stages. In addition, cell physiology, including cell proliferation, was examined during development via immunostaining for Ki67 to understand the cellular mechanisms that govern acinar cell differentiation during salivary gland morphogenesis. The distinct localization patterns of cytokeratins in conjunction with cell physiology will reveal the roles of epithelial cells in salivary gland formation during the differentiation of serous, mucous or seromucous salivary glands.     

Glycoconjugate distribution in the human fundic mucosa revealed by lectin- and glycoprotein-gold cytochemistry

Summary The glycoconjugates of the human fundic mucosa were characterized at the ultrastructural level by means of direct (Helix pomatia agglutinin-gold complex) and indirect lectin techniques (Concanavalin A and horseradish peroxidase-gold complex; wheat germ agglutinin and ovomucoid-gold complex). Surface mucous cells and mucous neck cells secreted O-glycoproteins with N-acetylgalactosamine and N-acetylglucosamine residues at the non reducing terminus of the saccharidic chain. The secretory granules of the mucous neck cells showed condensed areas slightly reactive to ConA. The results obtained in the chief cells suggest that these cells secrete N-glycoproteins rich in mannose and/or glucose residues. Transitional cells, presenting both morphological characteristics and lectin binding pattern intermediate to the mucous neck and chief cells have been observed. The surface of the intracellular canaliculi of the parietal cell was labelled by HPA, WGA and ConA. In the neck region of the gastric glands, immature parietal cells containing abundant mucous granules reactive to HPA, WGA and ConA were observed. The present results further corroborate the existence of a common cell precursor for surface mucous, mucous neck and parietal cells. In a further step, mucous neck cells gradually differentiate into chief cells the transitional cells being an intermediate stage.     

Lysozyme localization in human gastric and duodenal epithelium

Summary The distribution of lysozyme in normal gastric and duodenal mucosa was studied by light- and electronmicroscopic immunocytochemical techniques (direct enzyme-labeled antibody method).In the duodenal mucosa, lysozyme was found in the Paneth cells and the epithelial cells of Brunner's glands. Electron-microscopically, lysozyme was found in rough endoplasmic reticulum and perinuclear spaces, which were assumed to be protein-synthesizing organelles, and also in the secretory granules of Paneth cells. Additionally, lysozyme was detected in the stomach in mucinous granules and in some parts of the rough endoplasmic reticulum within the epithelial cells of the pyloric glands, the mucous neck cells of the fundic glands, and in several surface epithelial cells of the plyoric and fundic regions.This suggests that some quantity of lysozyme in gastrointestinal secretion originates from the gastric and duodenal glands, and that it acts as a defense mechanism in the gastrointestinal tract.     

Glycoconjugate distribution in the human fundic mucosa revealed by lectin- and glycoprotein-gold cytochemistry

The glycoconjugates of the human fundic mucosa were characterized at the ultrastructural level by means of direct (Helix pomatia agglutinin-gold complex) and indirect lectin techniques (Concanavalin A and horseradish peroxidase-gold complex; wheat germ agglutinin and ovomucoid-gold complex). Surface mucous cells and mucous neck cells secreted O-glycoproteins with N-acetylgalactosamine and N-acetylglucosamine residues at the non reducing terminus of the saccharidic chain. The secretory granules of the mucous neck cells showed condensed areas slightly reactive to ConA. The results obtained in the chief cells suggest that these cells secrete N-glycoproteins rich in mannose and/or glucose residues. "Transitional cells", presenting both morphological characteristics and lectin binding pattern intermediate to the mucous neck and chief cells have been observed. The surface of the intracellular canaliculi of the parietal cell was labelled by HPA, WGA and ConA. In the neck region of the gastric glands, immature parietal cells containing abundant mucous granules reactive to HPA, WGA and ConA were observed. The present results further corroborate the existence of a common cell precursor for surface mucous, mucous neck and parietal cells. In a further step, mucous neck cells gradually differentiate into chief cells the transitional cells being an intermediate stage.     

The morphology of the mucous gland and its responses to prolactin in the skin of the red-spotted newt

The mucous gland of the red-spotted newt, Notophthalamus viridescens viridescens, Rafinesque was examined by histochemical and ultrastructural techniques and its cytological responses to various hormonal conditions were studied. Its secretory epithelial cells produce and release in merocrine fashion a neutral, unsulphated mucosubstance. The secretory epithelium is bounded peripherally by a thin, but apparent non-functional, myo-epithelium. The duct of this mucous gland consists of a single keratinized tubular cell that extends from the neck region of the gland to the surface of the epidermis. Mucous secretion is absent or greatly reduced on the skins of newts maintained under laboratory conditions for a few weeks but reappears after injection of ovine prolactin. Mucous glands in laboratory conditioned animals show a 4-fold increase in volume brought about by the engorgement of their epithelial cells with secretory granules. Ovine prolactin reduces the volume of the glands to unconditioned levels with a corresponding reduction in granular content, suggesting that prolactin functions in the release of the granules. This view is reinforced by the findings that autotransplantation of the pituitary gland prevents the conditioning effect and that glandular volume increases in auto-transplanted animals given ergocornine. Granular accumulation begins also in hypophysectomized newts but ceases after a week, indicating the need for some hypophyseal factor in the synthesis as well as the release of the granules. Ovine prolactin restores mucous glands of hypophysectomized newts to the unconditioned state. Contrary to earlier findings, ovine prolactin induces a reduction in the volume of the mucous gland in thyroidectomized newts.     

Modification of cell evagination and cell differentiation in quail oviduct hyperstimulated by progesterone

Quail oviduct development is controlled by sex steroid hormones. Estrogen (E) induce cell proliferation, formation of tubular glands by epithelial cell evagination and cell differentiation. Progesterone (P) strongly increases the secretory process in E-treated quails, but inhibits cell proliferation, cell evagination and differentiation of ciliated cells. The balance between E and P is critical for harmonious development of the oviduct. After 6 daily injections of two doses of estradiol benzoate (10 or 20 micrograms/d) and high doses of P (4 mg/d), tubular gland formation by epithelial cell evagination was inhibited, while epithelial cell proliferation occurred, as shown by the height of the villi and the increase in DNA. Secretory processes were strongly stimulated. Ovalbumin, a tubular gland cell marker and avidin, a mucous cell marker, were localized by immunofluorescence and immunogold labeling. Ovalbumin was localized only in the rudimentary tubular glands, whereas avidin was dispersed throughout the secretory cells. High doses of progesterone inhibited tubular gland cell proliferation, disturbed the distribution of avidin and inhibited differentiation of ciliated cells. Ovalbumin synthesis occurred only in epithelial cells which were evaginated despite the hyperstimulation. Ovalbumin gene expression appeared highly dependent upon the cell position.     

Lectin cytochemistry in the gastrointestinal tract with special reference to glycosylation in the Golgi apparatus of Brunner's gland cells.

Two hydrophilic, low temperature-embedding resins, Lowicryl K4M and LR White, were compared in lectin cytochemistry. Post-embedding staining of colloidal gold-labeled Griffonia symplicifolia agglutinin II (GSA-II) resulted in staining of the Golgi apparatus and mucous granules of mucous neck cells in the gastric fundic gland, pylorocytes, and Brunner's gland cells embedded in either resin, although it was much easier to make ultra-thin sections with LR White-embedded material than with the other. Post-fixation with uranyl acetate followed by LR White embedding improved general ultrastructure so that lectin binding sites were identified precisely. All examined lectins, soybean agglutinin (SBA), Maclura pomifera agglutinin (MPA), GSA-II, and Ulex europaeus agglutinin I (UEA-I), stained mucous granules and the Golgi apparatus, in which the staining pattern was characteristic of each lectin: cis cisternae were labeled with SBA and MPA, intermediate cisternae with GSA-II, and trans cisternae and mucous granules with SBA, GSA-II, UEA-I, and lightly with MPA. No labeling was observed in the rough endoplasmic reticulum with any lectin. These findings suggest that the Golgi apparatus is the site of O-linked glycosylation and can be divided into at least three distinct compartments with regard to the glycosylation.     

Distribution of the glio-interstitial system in molluscs

Summary Ten hamsters received repeated injections of ³H-thymidine for 4 days and were allowed to survive for 7, 28, 42 and 100 days. Changes in spatial distribution of the labelled cells and in labelling indices of each cell line in the gastric glands were studied at various days after ³H-thymidine injections, and the fate of the mucous neck cell, the replacement of the chief cell and the mode of cell migration were discussed.After 4 days of repeated injections of ³H-thymidine, the labelled parietal cells and the mucous neck cells were concentrated at the neck area. Starting from the neck area, they migrated an average of 3 micra downwards per day. By 42 days, they reached the middle level of the glands, where the labelled mucous neck cells decreased but the labelled chief cells increased in number. The differentiation of the chief cell then appears to take place at the middle level of the glands through transformation of the migratory mucous neck cells. After 4 days of the labelling, about 1.8% of the chief cells located in the lower part of the glands was found to undergo in situ replication. This indicates that the renewal of this cell type is partly assured by its own mitotic activity.The foveolar cell — the future surface epithelium — seems to migrate upwards along the long axis of the glandular tubule in the pipe line system, which means first produced, first migrates. After migrating out from the neck area, the parietal cell and the mucous neck cell (the future chief cell) take an average of 200 days to reach the lower end of the glands. In the process of migration, however, the cells produced contemporaneously at the neck area became scatteringly spread from the neck towards the bottom of the gland. The time required for the newly-formed cells to reach the lower end of the gland varied between 100 and 300 days. In the gastric glands the cells first produced at the neck area do not first reach the lower end of the glands. This mode of random migration is referred to as the stochastic flow system. As one of the probable factors which disturb the pipe line flow of downward cell migration, cellular movements perpendicular to the long axis of the glandular tubule were suggested to occur at random at an any level of the gastric glands.Supported by a Grant-in-Aid for Cancer Research from the Ministry of Education, Science and Culture, Japan     

The origin of pre-neoplastic metaplasia in the stomach: chief cells emerge from the Mist

The digestive-enzyme secreting, gastric epithelial chief (zymogenic) cell is remarkable and underappreciated. Here, we discuss how all available evidence suggests that mature chief cells in the adult, mammalian stomach are postmitotic, slowly turning over cells that arise via a relatively long-lived progenitor, the mucous neck cell, The differentiation of chief cells from neck cells does not involve cell division, and the neck cell has its own distinct pattern of gene expression and putative physiological function. Thus, the ontogeny of the normal chief cell lineage exemplifies transdifferentiation. Furthermore, under pathophysiogical loss of acid-secreting parietal cell, the chief cell lineage can itself trasndifferentiate into a mucous cell metaplasia designated Spasmolytic Polypeptide Expressing Metaplasia (SPEM). Especially in the presence of inflammation, this metaplastic lineage can regain proliferative capacity and, in humans may also further differentiate into intestinal metaplasia. The results indicate that gastric fundic lineages display remarkable plasticity in both physiological ontogeny and pathophysiological pre-neoplastic metaplasia.     

Transdifferentiation of mucous neck cells into chief cells in fundic gastric glands shown by GNA lectin histochemistry

The epithelium of the gastric mucosa and its glands in the corpus of rat stomach contains mucous surface cells (MSCs), parietal cells, mucous neck cells (MNCs), zymogenic or chief cells (ZCs), several types of enteroendocrine cells, and intermediate cells with characteristics between MNCs and ZCs also called transitional or prezymogenic cells (pre-ZCs).The aim of our work was to analyze the expression of Mannose (Man) in the rat gastric glands by means of Galanthus nivalis lectin (GNA) histochemistry to identify the differences between MNC, pre-ZCs and ZCs and to establish the relationships between these cells. Most of the cytoplasm of MNCs was negative for GNA histochemistry. Intensity of GNA labeling in the gastric gland showed a graduation from pre-ZCs (weak labeling) to ZCs (moderate labeling). Labeling of ZCs was stronger at the perinuclear and apical cytoplasm.In the last years, strong evidence has been reported supporting that ZCs differentiate from MNCs. Our work also supports the origin of ZCs from MNCs, because the GNA labeling graduation might be due to oligosaccharides which are not expressed in MNCs, start to express in pre-ZCs and are more abundant in ZCs, indicating that differentiation from MNCs to ZCs is a process in which glycans with Man moieties are synthesized.     

The development of the stomach in Clarias lazera and the intestinal absorption of protein macromolecules

Summary The development of the stomach of the teleost, Clarias lazera, during the early posthatching period, is described, and the developing stomach is compared with that of adult Clarias.The stomach develops in two distinct parts: the corpus, which differentiates first, and the pylorus. The corpus contains a mucous surface epithelium, arranged in folds, and a tubular gland system containing only one type of gland cell, to which the secretion of pepsinogen and HCl is attributed. The pyloric region does not contain tubular glands.From the ultrastructure of the gland cells, the ³H-thymidine labeling index, and the onset of acid production (as determined with pH indicators) it is concluded that a functional stomach is present in juveniles with a standard length of ± 11 mm (approximately 12 days after fertilization at 23–24° C).The ultrastructure of the intestinal epithelium has also been studied. The intestine consists of three segments, similar to those described for stomachless teleosts and a number of fish larvae. In larvae as well as in juveniles, the enterocytes of the second segment show pinocytosis of horseradish peroxidase, although in the juveniles the stomach has already developed. This second segment has the same relative length in all studied larvae and juveniles and is also present in adult Clarias.It is therefore concluded that the capacity to absorb protein macromolecules is not specifically related to the absence of a functional stomach in this teleost species.