The morphogenesis of the human Paneth cell. An immunocytochemical ultrastructural study

In human duodenal mucosa Paneth cells originate away from the base of crypts and migrate towards the base during maturation. The earliest cells in the Paneth cell lineage could be identified by labelling of lysozyme in the Golgi apparatus. Specific labelling for lysozyme was present in the rough endoplasmic reticulum, Golgi apparatus, condensing vacuoles, granules and many lysosomes of mature Paneth cells. The maturation of the Paneth cell is accompanied by an increase in the content of lysozyme in the secretory granules and with senescence lysozyme diffuses into the cytoplasm.     

The morphogenesis of the human Paneth cell

Summary In human duodenal mucosa Paneth cells originate away from the base of crypts and migrate towards the base during maturation The earliest cells in the Paneth cell lineage could be identified by labelling of lysozyme in the Golgi apparatus. Specific labelling for lysozyme was present in the rough endoplasmic reticulum, Golgi apparatus, condensing vacuoles, granules and many lysosomes of mature Paneth cells. The maturation of the Paneth cell is accompanied by an increase in the content of lysozyme in the secretory granules and with senescence lysozyme diffuses into the cytoplasm.     

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.     

Immunocytochemical localization of secretory component in Paneth cell secretory granules-rat Paneth cells participate in acquired immunity

Summary With the marker of Paneth cells-lysozyme, secretory component (SC) immunoreactivity was demonstrated exclusively in Paneth cells of rat small intestine. The other types of epithelial cells (columnar, goblet, endocrine) were negative. On electron microscopic level, many SC-positive colloidal gold particles were found in rough endoplasmic reticulum, Golgi complexes, basal membrane and secretory granules of Paneth cells. These results suggest that SC is not a component of ingested immune complex, but a membrane receptor on Paneth cell. It may function as receptor for polymeric IgA and mediate its transport across the mucosal epithelium. Thus, Paneth cells are responsible for SC synthesis and participate in IgA-mediated acquired immunity in rat small intestine.     

A study of the anatomy, histology and ultrastructure of the digestive tract of Orthrias angorae Steindachner, 1897

The anatomy, histology and ultrastructure of the digestive tract of Orthrias angorae (Steindachner, 1897) were investigated using light microscopy, scanning electron microscopy (SEM) and transmission electron microscopy (TEM). The histological structure consists of four layers: mucosa, submucosa, muscularis and serosa. The esophageal mucosa consists of undifferentiated basal epithelial cells, mucous cells and surface epithelial cells. It was observed that the J-shaped stomach had a meshwork of folds in the cardiac region, and longitudinal folds in the fundic and pyloric regions. A single layer of columnar cells, PAS positive only in their apical portions, forms the epithelium. The convoluted tube-shape intestine is lined by simple columnar epithelial cells, which have microvilli at the apical surface. The wall of the esophagus and stomach are thicker than that of the intestine because of the thick muscle layer. There were numerous goblet cells in the intestine. There were numerous gastric glands in the submucosa layer ofthe cardiac stomach, but none were present in the pyloric region of the stomach. There were no pyloric caeca between the stomach and intestine. The enterocytes with microvilli contained rough endoplasmic reticulum, ribosomes and rounded bodies, and the gastric cells contained a well-developed Golgi apparatus.     

A method of quantitating Paneth cell metaplasia of the stomach by image analysis

Paneth cells are one of the histologic components of intestinal metaplasia of the stomach, as are mucin-producing goblet cells. With the aid of an image quantifier, the distribution of Paneth cells histochemically labeled with acid fuchsin was analyzed for a gastrectomy specimen containing an adenocarcinoma of the intestinal type; the topographic distribution of goblet cells histochemically labeled with Alcian blue (pH 2.5) was also analyzed. The specimen was cut into 63 blocks (0.5 X 4.0 cm) in four zones; antrum (zone I), intermediate region (zone II) and fundus (zones III and IV). Paneth cells were found only in sections containing mucin-producing goblet cells. Paneth cells were found in 12.5% of the 16 sections from the antral zone I containing Alcian blue-positive goblet cells. The rates were 44.4% for the intermediate zone II and 55.5% for the distal fundic zone III. The total area occupied by Paneth cells was significantly lower in the gastric mucosa as compared to the duodenal mucosa. The "Paneth cell index" (total Paneth cell area/total goblet cell area) was highest in the duodenum, followed by the distal fundic zone III. This method of quantitating Paneth cell metaplasia of the stomach will be used to investigate the topographic distribution of those cells in populations with low and high incidences of intestinal metaplasia.     

Expression of duodenase-like protein in epitheliocytes of Brunner’s glands in human duodenal mucosa

A duodenase, a protease structurally related to human cathepsin G, was found earlier in bovine duodenal mucosa. It was demonstrated that under the influence of duodenase an enteropeptidase zymogen is activated in vitro showing the possible participation of duodenase in the cascade of activation of digestive enzymes. To identify a duodenase functional analog in human duodenum, an immunofluorescence study of duodenal mucosa was conducted by confocal microscopy using antibodies to human cathepsin G and to bovine duodenase. The previously unknown place of synthesis and secretion of cathepsin G — Paneth cells located at the bottom of Lieberkuhn crypts — was revealed. Binding of cathepsin G-specific antibodies in a rough endoplasmic reticulum zone and in the cryptal duct was observed. Duodenase-specific immunofluorescence but not that of cathepsin G was found in the epitheliocytes and secretory ducts of Brunner’s glands, which are characteristic sites of duodenase biosynthesis in cattle. Binding of CD14-specific antibodies in the Brunner’s glands, where the antibodies co-localized with the antibodies to duodenase, was also demonstrated. These data indicate the presence of a protein immunologically similar to duodenase in the human duodenal mucosa. Our study demonstrated the absence of its colocalization with cathepsin G in Brunner’s glands.     

Contribution of carbon monoxide-producing cells in the gastric mucosa of rat and monkey

 Recent studies have shown that carbon monoxide (CO) may function as a gaseous signaling molecule in a similar way to nitric oxide. In the gastrointestinal tract, immunoreactivity against a CO-producing enzyme, heme oxygenase-2 (HO-2), was reported in epithelial cells and neurons of submucosal and myenteric plexus. However, details of the epithelial cells in the gastric mucosa remain unknown. The aim of this study was to clarify if mRNA for HO-2 is expressed in the rat stomach, if HO-2 protein is present in the mucosa, and to define the cell types of the HO-2-immunoreactive cells. HO-2 mRNA and protein were detected in fundic and pyloric mucosa of rat stomach using an RNA protection assay and western blot analysis. Immunohistochemical study showed that HO-2 was localized in parietal cells of the fundic glands and gastrin cells of the pyloric glands of both rat and monkey. The results suggest that HO-2 enzyme is produced in the gastric mucosa, and that CO is released from parietal cells and gastrin cells. Accepted: 12 November 1997     

Light and electron microscopic identification of several types of endocrine cells in the gastrointestinal mucosa of the cat

Summary The following histological methods, previously proved to be useful in selective light microscopic detection of endocrine cells, were applied to the cat gastrointestinal mucosa: for the identification of biogenic amines, diazonium, ammoniacal silver and xanthydrol methods; for granules identification, methyl green-red acid dyes, toluidine blue, HCl-basic dye, lead-haematoxylin, phosphotungstic haematein and argyrophil methods. Results were compared with those of an extensive electron microscopic investigation.Five types of endocrine cells were identified in the gastric mucosa. Three types were found in the pyloric mucosa: the previously described 5-hydroxytryptamine-producing enterochromaffin cell, the gastrin producing G cell and a cell with an unknown function, labelled in this paper the X cell. Four types were found in the fundic mucosa: enterochromaffin cells (rarely observed), enterochromaffin-like cells secreting a 5-hydroxyindole but showing some ultrastructural and staining differences from true enterochromaffin cells (numerously present), A-like cells (few), resembling A cells of the pancreatic islets, and X cells, resembling those in the pyloric mucosa.In the intestinal mucosa, at least three endocrine cell types were distinguished in its duodenal part: enterochromaffin cells and two types of polypeptide-producing cells — some with smaller granules (S cells) and others with larger granules (L cells). Only two types were found in the mucosa of terminal ileum: enterochromaffin cells and numerously-occurring cells with large granules resembling in part duodenal L cells. The possibility of a relationship between S and L cells and the production respectively of the intestinal hormones secretin and cholecystokinin-pancreozymin was discussed.This investigation was supported by a grant N. 115/1139/0/4715 of the Italian Consiglio Nazionale delle Ricerche.     

Histological and histochemical observations in the stomach of the Senegal sole, Solea senegalensis

An histological and histochemical study was conducted on the stomach of adult Senegal sole, Solea senegalensis specimens. The stomach was made up of four distinct layers: mucosa, lamina propria-submucosa-, muscularis and serosa. Surface epithelial, glandular and rodlet cells were present in the mucosa. Cells of the columnar epithelium contained a basal nucleus. Numerous mitochondria, granular endoplasmic reticulum and Golgi apparatus consisting of several parallel cisternae and vesicles were observed in the cytoplasm of these cells. The lysosomes were small, round and dense. The gastric glands were numerous in the pyloric and fundic regions but absent in the cardiac stomach. These glands were formed by two cell-types: light and dark cells. The light cells were characterised by numerous mitochondria, while dark cells had slightly fewer mitochondria and a tubulo-vesicular system. Rodlet cells similar to those observed in other teleostean fish were present among the epithelial cells. Although the epithelial cells of the mucosa contained a weak presence of neutral and acid mucopolysaccharides/mucosubstances, these substances were abundant in the lamina propria-submucosa. Proteins rich in arginine, lysine, cysteine and cystine were rarely present in the mucosa and lamina propria-submucosa of stomach, while proteins rich in tyrosine were abundant in these layers. Acid phosphatase, and ATP-ase (pH 7.2 and 9.4) activities were detected in the mucosa and lamina propria-submucosa. Alkaline phosphatase activity was not detected.     

Effect of omeprazole on secretion,synthesis and the gene expression of pepsinogen in the guinea pig stomach mucosa

The effects of omeprazole, a proton pump inhibitor, on gene expression, protein synthesis, intracellular storage and secretion of pepsinogen in guinea pig stomach were investigated. After treatment with omeprazole for five days, acid and pepsinogen secretion into the gastric lumen was significantly reduced. Concomitant with this, there was an increase in intracellular pepsinogen as demonstrated by increased pepsin activity in the gastric mucosa, more intense immunohistochemical staining by antibodies specific of pepsinogen and accumulation of secretory granules in the cells producing pepsinogen. In these cells, the amount for pepsinogen mRNA was reduced as revealed by Northern blotting and in situ hybridization. Ultrastructurally the endoplasmic reticulum of these cells was poorly developed, the findings being consistent with a reduction in protein synthesis. It appears that omeprazole inhibits the secretion of pepsinogen, increasing the intracellular store and leading to the reduction in gene expression probably by a feedback mechanism and consequent reduction in pepsinogen synthesis. Since these changes were most evident in the acid-secreting fundic gland mucosa, as compared with other mucosae secreting only pepsinogen, namely pyloric and duodenal mucosa, it appears probable that these changes are linked with omeprazole-induced reduction in the acid secretion.     

The ultrastructure of mammary explants from virgin ovariectomized goats during hormone-induced lactogenesis

The effect of insulin (I), cortisol (F) and prolactin (P) on the ultrastructural morphology of epithelial cells of cultured mammary explants from virgin ovariectomized (OV-X) goats were studied. The epithelial cells showed little structural organization and were devoid of fat droplets and secretory protein granules at zero time of culture. The cytoplasm contained few profiles of smooth and rough endoplasmic reticulum and the Golgi apparatus was rudimentary. After being cultured in Waymouth's medium without added hormones the epithelial cells were indistinguishable from epithelial cells of uncultured explants. The addition of I induced changes mainly in the appearance of nucleoli. The nucleoli were enlarged and fibrillogranular areas with light spaces were observed. The most obvious cytological changes of epithelial cells of explants cultured in the presence of I and F are translocation of the nucleus into the basal cytoplasm, increase of rough endoplasmic reticulum, an increase in the size of the Golgi apparatus, presence of one or two lipid droplets and in some cells vacuoles with protein granules were present. Mitochondria were more abundant. The epithelial cells of explants cultured in the presence of I, F and P were characterized by the polarization of organelles within the cytoplasm and by the formation and release of protein granules and small and large fat droplets. The cell nucleus was in the basal cytoplasm, the Golgi apparatus was supranuclear. The rough endoplasmic reticulum was extensively developed and formed large sacs. Golgi vacuoles contained protein granules.(ABSTRACT TRUNCATED AT 250 WORDS)     

Localization of pepsinogen and cellular differentiation in the human gastric mucosa using lowicryl K4M

The localization of pepsinogens (PG A and PG C) was studied intracellularly in human gastric biopsies embedded in Lowicryl K4M, using affinity purified antibodies and protein A-gold. The homogeneous secretory granules of the chief cells contained both PG A and PG C, as was proved in serial sections. Identical reaction was seen in the core of the biphasic mocous neck cell granules, whereas the mantle did not label. Even the rough endoplasmic reticulum (RER) and Golgi complex of the chief- and mucous neck cells contained label. Transitional cells identified by the presence of granules of both chief- and mucous neck cells were seen. This type of mucous neck cell is thought to transform into a chief cell. However an increase of RER that could explain an increase of the pepsinogen production was not observed. A mixture of these granules were also found in morphologically characterized young parietal cells, suggesting a common precursor for these three cell-types. These observations makes the transformation from mucous neck- into chief cells questionable. In conclusion Lowicryl K4M appeared to be a significant improvement compared to the Epon 812. Its shows a better preservation of both cytoplasmic antigens and cellular fine structure. This improvement adds information on the transformation hypothesis. Lowicryl K4M enables us, firstly to distinguish PG A and C synthesizing RER in different types of cell and secondly to recognize immature cells with the characteristics of chief-, mucous neck-, and parietal cells in the fundic gland. Very likely these three cell-types all arise from a common precursor. It is questionable that in normal human gastric mucosa the mucous neck cells transform into chief cells.     

STUDIES ON SMALL INTESTINAL CRYPT EPITHELIUM : I. The Fine Structure of the Crypt Epithelium of the Proximal Small Intestine of Fasting Humans

Small intestinal crypt epithelium obtained from normal fasting humans by peroral biopsy of the mucosa was studied with the electron microscope. Paneth cells were identified at the base of the crypts by their elaborate highly organized endoplasmic reticulum, large secretory granules, and small lysosome-like dense bodies within the cytoplasm. Undifferentiated cells were characterized by smaller cytoplasmic membrane-bounded granules which were presumed to be secretory in nature, a less elaborate endoplasmic reticulum, many unattached ribosomes and, in some cells, the presence of glycogen. Some undifferentiated cells at the base of the crypts contained lobulated nuclei and striking paranuclear accumulations of mitochondria. Membrane-bounded cytoplasmic fragments, probably originating from undifferentiated and Paneth cells, were frequently apparent within crypt lumina. Of the goblet cells, some were seen actively secreting mucus. In these, apical mucus appeared to exude into the crypt lumen between gaps in the microvilli. The membrane formerly surrounding the apical mucus appeared to fuse with and become part of the plasma membrane of the cell, suggesting a merocrine secretory mechanism. Enterochromaffin cells were identified by their location between the basal regions of other crypt cells and by their unique intracytoplasmic granules.     

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.     

Importance of acidic mucin secretions by foveolar and mucous neck cells of rat fundic mucosa as the defence mechanisms against HCl as revealed by fasting.

The localization of neutral mucin and acidic mucins in both control and fasted rat gastric fundic mucosa were examined by microscopic and electron microscopic histochemical methods. By Carnoy's fixation, the surface mucous coat of the control rat gastric fundic mucosa was found to be composed of alternating layers of acidic mucins and neutral mucin, indicating the synchronous and cyclic secretions of them. In many gastric pits of the fundic glands, the acidic mucins were found to spring out from the deep foveolar regions like volcanoes. This phenomenon may suggest that the acidic mucins play a fundamental role in protecting the pit cells against HCl during its passage, and the layers of neutral mucin and acidic mucins in the surface coat is the safeguard against the HCl and digestive enzymes in the gastric lumen. In the fasting rat gastric fundic mucosa, the acidity and the amount of the gastric juice were markedly decreased, indicating the suppressed secretions of mucins and HCl. The decreased production of sulfomucin was directly demonstrated by 35SO4-autoradiography. Many mucous neck cells existing in close association with the parietal cells were ballooned due to accumulation of alcian blue (AB)-positive but high iron-diamine (HID)-negative sialomucin, which was not demonstrable in the control. The secretory granules of sialomucin contained in the ballooned mucous neck cells were positively stained ultrastructurally with cacodylate-ferric colloid to stain acid mucopolysaccharides.     

Morphological and histochemical observations on the duodenal glands of eight wild ungulate species native to North America

The duodenal glands of the species examined (Alces alces, Ovis canadensis, Cervus canadensis, Oreamnos americanus, Bison bison, Antilocapra americana, Odocoileus virginianas, Odocoileus heminous) are confined primarily to the submucosa of the small intestine. In one species, the moose, a significant population of secretory tubules also is observed in the mucosa. The ducts of the duodenal glands pierce the overlying muscularis mucosae to empty most often independently into the intestinal lumen. Those of the bison, unlike the other species examined, drain into intestinal glands. The duodenal glands consist primarily of a simple columnar epithelium, the cells of which contain basally positioned round or oval nuclei. The lumina of scattered duodenal glands in the pronghorn and to some extent those of the moose, white-tailed deer, and mule deer may be extremely dilated, and the surrounding epithelium thin and attenuated. Component cells of the duodenal glands of all the species examined show remarkably similar ultrastructural features. They exhibit scattered profiles of rough endoplasmic reticulum, dilated cisternae of which contain an electron-dense, amorphous material. Numerous well-developed Golgi complexes occupy the supranuclear region together with transport vesicles and forming secretory granules. Electron-dense, membrane-bound secretory granules generally are concentrated in the apical cytoplasm immediately subjacent to the cell membrane. The apical cell membrane exhibits short, scattered microvilli; and the basal cell membrane is smooth without apparent specialization. Histochemically, the duodenal glands of most species examined in this study consist of a heterogeneous population. The majority of the glands of the moose, elk, mountain goat, bison, pronghorn, and white-tailed deer elaborate a neutral mucin, whereas scattered individual glands, tubules or cells also produce acid mucins. Cells near the terminations of the ducts of the bighorn sheep are the only elements to produce acid mucins in the duodenal glands of this species. The duodenal glands of the bison are unusual in that only the peripheral portions of individual glands produce acid mucins. The remainder of the glands elaborate neutral mucins. Morphological differences between the two regions were not observed. The duodenal glands of the mule deer secrete both acid and neutral mucins. The structural and histochemical observations appear unrelated to the diet of individual species.     

Cytochemical features of the digestive tract mucosa of Hemisorubim platyrhynchos (Siluriformes: Pimelodidae)

Membranous organelles, acid glycoconjugates and lipids were characterized in the digestive tract mucosa of Hemisorubim platyrhynchos by cytochemistry techniques. Two types of mucous‐secreting cells were observed in the digestive tract epithelium: goblet cells in the oesophagus and intestine and epithelial cells in the stomach. These cells had a Golgi apparatus more developed than the other cell types. The cytochemical analysis revealed that secretory granules are reactive to acid glycoconjugates, varying in reaction intensity according to the region of the digestive tract. Acid glycoconjugate reactions were also observed in oesophageal epithelial cell microridges and in enterocyte microvilli. In the digestive tract, acid glycoconjugates act to protect the epithelial surface, increasing mucous viscosity, which facilitates the passage of food, prevents the binding of parasites and facilitates their removal. Through lipid staining, a coated membrane was observed around each secretory granule of the oesophageal and intestinal goblet cells, while gastric epithelial cells granules were fully reactive. Oxynticopeptic cells of the gastric glands showed lipid droplets in the cytoplasm and also in the mitochondrial matrix, which act as an energy reserve for these cells that have a high energy demand. Enterocytes showed a well‐developed smooth endoplasmic reticulum, especially in the apical region of the cell, being related to absorption and resynthesis of lipids.     

Immunocytochemical localization of rabbit gastric lipase and pepsinogen

Lipase and pepsin activities were determined in rabbit gastric biopsy specimens. Lipase activity was found to be restricted to a small part of the fundic mucosa, near the cardia, whereas pepsin activity spread over about two thirds of the total fundic area, overlapping that of lipase. The cells producing these two enzymes were labeled by immunofluorescence using polyclonal antibodies against rabbit gastric lipase (RGL) or antibodies against rabbit pepsinogen. The immunocytochemical localization showed unequivocally that RGL and pepsinogen, which were both present in the cardial area, were in fact located in different gastric cells. The cells producing pepsinogen were in the lower base of the gastric fundic glands, whereas the cells producing RGL were in the upper base of the same glands. The cells producing pepsinogen and RGL showed no significant morphological differences. In the part of the fundic area, where only pepsin activity was detected, cells producing pepsinogen covered both the lower and the upper base of the gastric glands. No chief cells were observed in the antral mucosa. RGL and pepsinogen could represent useful gastric enzyme markers for cellular differentiation studies.     

Observations on the anatomy of the stomach and duodenum of the bowhead whale, Balaena mysticetus

Gastric and cranial duodenal structure of the bowhead whale (Balaena mysticetus) was examined grossly and microscopically. The stomach was arranged in a series of four compartments. The first chamber, or forestomach, was a large nonglandular sac lined by a keratinized stratified squamous epithelium. It was followed by the fundic chamber, a large, somewhat globular and entirely glandular compartment. At the entrance of the fundic chamber, a narrow cardiac gland region could be defined. The remaining mucosa of the chamber contained the proper gastric glands. A narrow, tubular connecting channel, the third distinct gastric division, was lined by mucous glands and joined the fundic chamber with the final stomach compartment, or pyloric chamber. This fourth chamber was also tubular and lined by mucous glands but was of a diameter considerably larger than the connecting channel. The stomach terminated at the pyloric sphincter which consisted of a well-developed band of circular smooth-muscle bundles effecting a division between the pyloric chamber and small intestine. The small intestine began with the duodenal ampulla, a dilated sac considerably smaller than the fundic chamber of the stomach. The mucosa of this sac contained mucous glands throughout. The ampulla led without a separating sphincter into the duodenum proper which continued the intestine in a much more narrow tubular fashion. The mucosal lining of the duodenum was composed of villi and intestinal crypts. Although their occurrence varied among whales, enteroendocrine cells were identified within the mucous glands of the cardiac region, connecting channel, pyloric chamber, and cranial duodenum. The hepatopancreatic duct entered the wall of the duodenum shortly after the termination of the duodenal ampulla and continued intramurally along the intestine before finally joining the duodenal lumen.