Figure 15—1. Schematic structure of a portion of the digestive tract with various components and their functions. (Redrawn and reproduced, with permission, from Bevelander G: Outline of Histology, 7th ed. Mosby, 1971.) Diagram of general organization of the alimentary canal. Basic mucosal forms in the gastrointestinal tract H & E Ч 100 Figure 15—2. Surface of the tongue on the region close to its V-shaped boundary, between the anterior and posterior portions. Note the lymphoid nodules (lingual tonsil), glands, and papillae. enis kunTovani qsovili Tongue, muscle, mid Tongue, muscle, high Features of the human tongue and distribution of papillae. a-d, X45 Tongue, filiform and fungiform papillae, with taste buds, high Diagram and photomicrograph (X640) of a taste bud. Diagram of a section through an incisor tooth and surrounding bony and mucosal structures. Diagram showing the basic organization and structure of enamel rods. EM of young enamel shows enamel rods cut obliquely. Arrows, boundaries between adjacent rods. X14,700. Parts of two adjacent enamel rods are seen at higher magnification. Arrows, boundary between the two rods. X60,000. Photomicrograph of a decalcified tooth showing dental pulp and structure of dentin. X120. Left inset. Longitudinal profiles of the dentinal tubules. X240. Right inset. Cross-sectional profiles of dentinal tubules. X240. Diagram comparing the components of the salivon in the three major salivary glands. Parotid gland, high EM of the apical portion of parotid gland serous cells. SV, secretory vesicles; L, lumen; rER, rough endoplasmic reticulum; G, Golgi apparatus profiles; IV, immature secretory vesicles; JC, junctional complexes; IC, intercellular space; M, mitochondria. X15,000. Low-magnification electron micrograph of a mucous acinus. L, lumen; MyC, myoepithelial cell processes. Low magnification EM of mixed acini. X6,000. EM of the sublingual gland prepared by traditional fixation in formaldehyde. X15,000. Submaxillary gland, high saylapavi Masson's trichrome Ч 9 saylapavi Masson's trichrome Ч 9 Photomicrograph of the esophagus. X8. Photomicrograph of the esophageal mucosa. X240 Photomicrograph of an esophageal submucosal gland. X110. Esophagus - stomach junction, low kuWi SEM of mucosal surface of the stomach. Arrows, mucus. X1,000. Higher magnification SEM showing the apical surface of the surface mucous cells that line the stomach and gastric pits. X3,000. Body of the stomach H & E Ч 12 Body of stomach: structure of glands Photomicrograph (x 320) and diagram of gastric glands Figure 15—11. Photomicrograph of a section of the gastric glands in the fundus of the stomach. Note the superficial mucus-secreting epithelium. Parietal cells (light-stained) predominate in the mid and upper regions of the glands; chief (zymogenic) cells (dark-stained) predominate in the lower region of the gland. MM, muscularis mucosae. PT stain. Low magnification. Figure 15—12. Photomicrograph of a mucus-secreting surface epithelium (A) and mucous neck cells intercalated between oxyntic (parietal) cells located in the mid portion of the gastric gland (B). Abundant capillaries can be seen. PT stain. Medium magnification. Stomach, corpus, high Stomach, corpus, high Stomach, corpus, high Figure 15—13. Photomicrograph of the basal portion of the gastric gland in the fundus. This section shows parietal cells rich in mitochondria and their characteristic intracellular canaliculi (arrowheads). Chief cells show red secretory granules in their cytoplasm. PT stain. Gastric body mucosa PAS/haematoxylin/orange G TS Ч 200 Figure 15—14. Electron micrograph of an active parietal cell. Note the microvilli (MV) protruding into the intracellular canaliculi and the abundant mitochondria (M). x10,200. (Courtesy of S Ito.) Diagram of a chief cell. Diagram of a parietal cell Figure 15—16. Diagram of a parietal cell, showing the main steps in the synthesis of hydrochloric acid. Active transport by ATPase is indicated by arrows and diffusion is indicated by dotted arrows. Under the action of carbonic anhydrase, blood CO2 produces carbonic acid. Carbonic acid dissociates into a bicarbonate ion and a proton H+, which is pumped into the stomach lumen in exchange for K+. A high concentration of intracellular K+ is maintained by the Na+,K+ ATPase, while HCO3— is exchanged for Cl— by an antiport. The tubulovesicles of the cell apex are seen to be related to hydrochloric acid secretion, because their number decreases after parietal cell stimulation. The bicarbonate ion returns to the blood and is responsible for a measurable increase in blood pH during digestion. Diagram of parietal cell HCl synthesis. Figure 15—15. Composite diagram of a parietal cell, showing the ultrastructural differences between a resting cell (left) and an active cell (right). Note that the tubulovesicles (TV) in the cytoplasm of the resting cell fuse to form microvilli (MV) that fill up the intracellular canaliculi (IC). G, Golgi complex; M, mitochondria. (Based on the work of Ito S, Schofield GC. J Cell Biol 1974;63:364.) Peptic cell - rat EM Ч 7200 Figure 15—17. Electron micrograph of a section of gastric gland in the fundus of the stomach. Note the lumen and the parietal cells, containing abundant mitochondria; chief cells, with extensive rough endoplasmic reticulum; and enteroendocrine cells (closed type), with basal secretory granules. x5300. Pyloric tomach H & E Ч 75 s Stomach, pylorus, high Photomicrograph of dividing cells in the isthmus of a pyloric gland. X240. Inset. High magnification of the area indicated by the rectangle showing a dividing cell in the isthmus. X580. Gastroduodenal junction - monkey H & E Ч 12 Pyloric - duodenal junction, low Pyloric - duodenal junction, high Pyloric - duodenal junction, high Figure 15—35. Photomicrograph of the duodenum, showing villi and duodenal glands in the submucosa. H&E stain. Low magnification. Duodenum with Brunner's glands, low Brunner's glands, high Duodenum Alcian blue Ч 150 Duodenum H & E Ч 15 Duodenal epithelium EM Ч 14 500 Jejunum, low Jejunum, high Jejunum with Paneth cells, high Jejunum with Paneth cells, high mucosal surface of small intestine openings located between the bases of the villi that lead into the intestinal glands (crypts of Lieberkühn). Figure 15—41. Regeneration of the epithelial lining of the stomach and small intestine. Note differences in the location of stem cells. Figure 15—20. Photomicrograph of the small intestine. Note the villi, intestinal glands, submucosa, muscle layers, and serosa. PT stain. Low magnification. Enterocytes EM Ч 56 000 wvrili nawlavi Figure 15—24. Structure of a microvillus. A cytoskeleton of actin filaments, associated with other proteins, keeps the shape of the microvillus. The actin filaments are continuous with the microfilaments of the terminal web (see Chapter 4), which also contains intermediate filaments. Note that in this location actin filaments have a structural role and are not related to movement, as is usually the case when these microfilaments are present. To fulfill its supportive role, actin is associated with other proteins that link the microfilaments to one another, to fibrin, and to the cell membrane and a specific protein–-villin–-in its tip. Small intestine - monkey Duodenum: H & E Ч 20 Small intestine - monkey Ileum: H & E Ч 16 Figure 15—32. Scanning electron micrograph of the intestinal surface after removal of the mucosal epithelium, showing the basement membrane. Note that this layer is continuous when covering the remnants of the intestinal villi but assumes the structure of a sieve when covering the lymphoid follicles above Peyer’s patches. This configuration permits an easier means for immunogenic materials to reach underlying lymphoid tissues. Figure 15—34. Blood circulation (left), lymphatic circulation (center), and innervation (right) of the small intestine. The smooth muscle system for contracting the villi is illustrated in the villus on the right. Intestinal villi and crypts SEM Ч 100 Intestinal villi and crypts H & E Ч 150 Intestinal villi H & E: LS Ч 100 Gastric body mucosa PAS/haematoxylin/orange G TS Ч 200 Crypts of Lieberkuhn H & E: TS Ч 400 Colon TS, H & E Ч 200 Intestinal villi H & E, TS Ч 300 Intestinal villi Enzyme histochemical method for alkaline phosphatase Ч 128 Intestinal villi Carmine perfused Ч 10 Tip of intestinal villus Thin section: toluidine blue Ч 320 Enterocytes EM Ч 4540 Figure 15—22. Electron micrograph of an absorptive epithelial cell of the small intestine. Note the accumulation of mitochondria in its apex. The luminal surface is covered with microvilli (shown in transverse section in the inset). Actin filaments, sectioned transversely, constitute the principal structural feature in the core of the microvilli. x6300. (Courtesy of KR Porter.) Figure 15—23. Electron micrograph of epithelium of the small intestine. Abundant microvilli at the cell apex can be seen to form the brush border. At the left are 2 lymphocytes migrating in the epithelium. In the center is an enteroendocrine cell (E) with its basal secretory granules. x1850. Enterocytes EM Ч 22 000 Figure 15—25. Lipid absorption in the small intestine. Lipase promotes the hydrolysis of lipids to monoglycerides and fatty acids in the intestinal lumen. These compounds are stabilized in an emulsion by the action of bile acids. The products of hydrolysis cross the microvilli membranes passively and are collected in the cisternae of the smooth endoplasmic reticulum (SER), where they are resynthesized to triglycerides. These triglycerides are surrounded by a thin layer of proteins that form particles called chylomicrons (0.2—1 micrometers in diameter). Chylomicrons are transferred to the Golgi complex and then migrate to the lateral membrane, cross it by a process of membrane fusion (exocytosis), and flow into the extracellular space in the direction of the blood and lymphatic vessels. Most chylomicrons go to the lymph; a few go to the blood vessels. The long-chain lipids (>C12) go mainly to the lymphatic vessels. Fatty acids of fewer than 10—12 carbon atoms are not reesterified to triglycerides but leave the cell directly and enter the blood vessels. RER, rough endoplasmic reticulum. (Based on results of Friedman HI, Cardell RR Jr: Anat Rec 1977;188:77.) Ileum with Peyer's patches, low Photomicrograph of Peyer's patches. X40. Diagrams of an enterocyte in different phases of absorption. Figure 15—30. Photomicrograph of the basal portion of 2 glands (crypts) of the small intestine. Note the enteroendocrine cell, Paneth cell, goblet cell, and a cell in mitosis. Diagram of an enteroendocrine (APUD) cell. Figure 15—18. Electron micrograph of an enteroendocrine cell (open type) of the human duodenum. Note the microvilli in its apex. x6900. (Courtesy of AGE Pearse.) Schematic diagram showing the distribution of gastrointestinal peptide hormones produced by enteroendocrine cells in the alimentary canal. CCK, cholecystokinin; VIP, vasoactive intestinal peptide; GIP, gastric inhibitory peptide. Figure 15—31. Electron micrograph from a region of intestine where a lymphoid nodule is covered by the intestinal mucosa. Note the presence of M cells that form a special compartment containing lymphoid cells. A macrophage (an antigen-presenting cell) is also in the compartment. (Courtesy of M Neutra.) Figure 15—33. Some aspects of immunologic protection of the intestine. A: A condition that is more frequent in the upper tract, such as in the jejunum. There are many IgA-secreting plasma cells, scattered lymphocytes, and some macrophages. Note that the lymphocytes in the epithelial lining are located outside the epithelial cells, and below the tight junctions. B: A condition that is more frequent in the ileum, where aggregates of lymphocytes are located under M cells. The M cells transfer foreign material (microorganisms and macromolecules) to lymphocytes located deep in the cavities of the M cells. Lymphocytes spread the information received from this foreign material to other regions of the digestive tract, and probably to other organs, through blood and lymph. Ileum, high Ileocaecal junction H&EЧ5 Colon H&EЧ4 Colon, high Colon H & E Ч 100
LS: Alcian blue/van Gieson Ч 80
Colon TS, Alcian blue/van Gieson Ч 320 Colon TS, H & E Ч 200 EM of an enteroendocrine cell of the colon. Arrowheads, boundary between the enteroendocrine cell and the adjacent epithelial cells; BL, basal lamina; . G, secretory granules; arrows, direction of secretion the across the basal lamina; CT, connective tissue; C, capillary; M, mitochondria; rER, rough endoplasmic reticulum; sER, smooth endoplasmic reticulum. Appendix H &EЧ5 Photomicrograph of a cross section through the vermiform appendix. X10. Appendix H & E Ч 75 Figure 15—38. Photomicrograph of a section of large intestine with its various layers. Note the absence of villi. M, mucosa; MM, muscularis mucosae; SM, submucosa; ME, muscularis externa. PT stain. Low magnification. SEM of the human mucosal surface of the large intestine. X140 Figure 15—40. Electron micrograph of epithelial cells of the large intestine. Note the microvilli at the luminal surface, the well-developed Golgi complex, and dilated intercellular spaces filled by interdigitating membrane leaflets, a sign of active water transport. x3900. EM of dividing goblet cells. GC, goblet cells; M, mucinogen granules; C, chromosomes; N, nuclei of nondividing intestinal epithelial cells; L, lumen of the gland;. CT, connective tissue; E, eosinophil. X5000. Rectum, high Rectum, high Recto-anal junction H & E Ч 60 Drawing of the rectum and anal canal. Photomicrographs of anal canal, X10 and X200. SQZ,squamous zone; ATZ, anal transitional zone; CRZ, colo-rectal zone. Diagram and SEM of M cells in a lymphatic nodule of the intestine. Figure 15—37. Photomicrograph of a group of neurons (with large nuclei) and satellite cells (with small nuclei) constituting a component of the myenteric plexus between 2 smooth muscle layers. Note the red stained collagen fibers. Picrosirius-hematoxylin. Medium magnification. Comparison of structure of parts of the gastrointestinal tract H & E Ч 15 Components of the wall of the gastrointestinal tract H & E Ч 480 EM of the myenteric (Auerbach's) plexus. SM, smooth muscle; CB, nerve cell bodies; N, nerve fibers; SC, Schwann cell; BV, blood vessel. X3,800. Photomicrograph of intestinal glands showing Paneth cells. X240. Inset. High magnification of the area indicated by the rectangle. X680.