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Functional Morphology of the Large Intestine

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Abstract

The sections in this article are:

1 Comparative Anatomy
1.1 Invertebrates
1.2 Chordata
2 Human Large Intestine
2.1 Gross Anatomy of Human Large Intestine
2.2 Development
3 Histology and Ultrastructure of Adult Mammalian Large Intestine
3.1 Cecum, Colon, Appendix, and Upper Rectum
3.2 Tunica Mucosa
3.3 Tela Submucosa
3.4 Tunica Muscularis
3.5 Lymphoid Tissue and Appendix
3.6 Tunica Serosa (Serosa)
3.7 Lower Rectum and Anal Canal
4 Restitution
5 Concluding Remarks
Figure 1. Figure 1.

A phylogeny of animal kingdom showing protostome and deuterostome divisions and their associated embryonic characteristics.

[From Barnes .]
Figure 2. Figure 2.

Diagram showing generalized arthropod body plan as viewed in a sagittal plane. Note that both foregut and hindgut have a portion of luminal surface covered by chitinous exoskeleton, illustrated here by a heavy dark line.

[From Barnes .]
Figure 3. Figure 3.

Diagrams illustrating relative positions of Malpighian tubules and rectum in relation to midgut and hindgut of insect. Specialization of rectum into rectal pads or papilla and ultrastructural anatomy of cells lining lumen are shown.

[Adapted from Berridge and Oschman .]
Figure 4. Figure 4.

Electron micrograph of a septate junction (desmosome) between epithelial cells lining cockroach rectum. Note ladderlike cross bridges spanning intercellular space.

(Courtesy of Dr. Peter Smith.)
Figure 5. Figure 5.

Diagram of digestive tracts of 6 representative aquatic vertebrates: lamprey (Petromyzon), shark (Squalus), chimaera (Callorhynchus), lungfish (Protopterus), sturgeon (Acipensus), and teleost (Perca).

[From Romer and Parsons .]
Figure 6. Figure 6.

Diagram of digestive tract and appendages of 4 vertebrates that are either amphibious or terrestrial. Note anatomic distinction between small and large intestine.

[Adapted from Romer and Parsons .]
Figure 7. Figure 7.

Diagram of large intestinal specializations found in birds. Arrows indicate retrograde flow of urine and feces mixture from cloaca into ceca.

[Adapted from Skadhauge .]
Figure 8. Figure 8.

Proportion of total body weight constituted by hindgut contents in various omnivores, carnivores, and herbivores.

[From von Engelhardt et al. after Stevens .]
Figure 9. Figure 9.

Diagram of gastrointestinal tract of a representative carnivore (A), omnivore (B), and 2 herbivores (C and D). Note extensive development of cecum and large intestine in herbivores.

[From von Engelhardt et al., after Stevens .]
Figure 10. Figure 10.

Diagram showing position and segments of adult human large intestine as it appears in body cavity. Greater omentum has been removed so transverse colon and coils of small intestine can be observed.

[From Basmajian .]
Figure 11. Figure 11.

Diagram of ileocaecal junction showing blood supply, lymph nodes, mesenteries, and orifice of ileum and appendix into cecum.

[From Snell .]
Figure 12. Figure 12.

Diagram of coronal section of adult terminal rectum, anal canal, anus, and adjacent structures.

[Adapted from Gray .]
Figure 13. Figure 13.

Diagram of adult human colon showing distribution of superior mesenteric artery and its branches that supply small intestine and appendix, cecum, and ascending and part of transverse colon (to arrow). Transverse colon has been elevated to show origin of artery.

[From Snell .]
Figure 14. Figure 14.

Diagram showing distribution of inferior mesenteric artery and its branches that supply distal one‐third of transverse, descending, and sigmoid colon, and rectum.

[From Hall‐Craggs .]
Figure 15. Figure 15.

Diagram showing arteries of rectum and anal canal from a frontal view. Note that superior rectal artery (Sup rectal a) gives a right and left branch, each of which anastomoses with branches of middle rectal artery, which is given off by internal pudendal artery (int pudendal a). Inferior rectal artery (inf rectal a) branches and anastomoses with branches from middle rectal artery.

[From Anderson .]
Figure 16. Figure 16.

Diagram showing distribution of lymphatic nodes and vessels of colon and upper rectum.

[From Jamieson and Dobson .]
Figure 17. Figure 17.

Diagrams showing various stages in development of gut tube as seen in sagittal midline section of human embryos. Arrows indicate extensions of amniotic cavity in presomite embryo (A), 7‐somite embryo (B), 14‐somite embryo (C), and end of month 1 (D).

[From Sadler .]
Figure 18. Figure 18.

Diagrams of transverse sections through human embryos at sequential stages of development from left to right. Effects of lateral folding on endoderm‐lined cavity are shown.

[From Sadler .]
Figure 19. Figure 19.

Diagram of human embryo to illustrate position of primitive intestinal loop and its continuity with vitelline duct. Note connection between allantois and cloaca.

[From Sadler .]
Figure 20. Figure 20.

Diagrams of primitive (prim) intestinal loop showing position of superior (sup) mesenteric artery (A) and position of cecal bud that designates division between small and large intestine after 180° rotation in primitive intestinal loop (B). Arrow in A indicates direction of counterclockwise rotation.

[From Sadler .]
Figure 21. Figure 21.

Diagram of 6‐wk‐old human embryo showing mesenteric connections with various parts of digestive tract.

[From Sadler .]
Figure 22. Figure 22.

Diagrams showing sequential development of anorectal canal and urogenital system in cloacal region. Arrow indicates path of descending urorectal septum.

[From Sadler .]
Figure 23. Figure 23.

Drawing of solid lumen phase of large intestinal development (A) and beginning of cavitation of lumen (B).

[From Sadler .]
Figure 24. Figure 24.

Light micrographs showing 4 stages in conversion of luminal epithelium from stratified to simple columnar in fetal rat colon. A: at 16 days, a stratified epithelium 2–3 cell layers thick surrounds a small lumen (L). × 600. B: at 17 days, main lumen (ML) projects into epithelium, forming deep clefts. In addition, secondary lumina (SL) are formed. × 600. C: by 19 days, secondary lumina (SL) are enlarged. Darkly stained and rounded apical cells (arrows) will be sloughed into main lumen. × 600. D: just before birth (22 days gestation), epithelium has changed to a simple columnar type that lines longitudinal ridges of connective tissue. Epithelium has become differentiated, as illustrated by goblet cells (GC). × 500.

[From Colony and Neutra .]
Figure 25. Figure 25.

Electron micrograph of nongoblet cell of 22‐day fetal rat proximal colon reacted for carbonic anhydrase activity, which is localized along lateral cell borders. Nu, nucleus. × 16,500.

[From Lacy and Colony .]
Figure 26. Figure 26.

Diagram of general structure and layers of various parts of gastrointestinal tract, including large intestine, as observed in cross section. Esophagus without submucosal glands (A), esophagus with submucosal glands (B), small intestine (C), and large intestine (D). Tunica Mucosa: E, epithelial layer (lamina epithelialis mucosae); F, lamina propria; G, lamina muscularis mucosa. Tunica submucosa: H, loose connective tissue and associated structures of submucosa. Tunica muscularis: I, circular muscle layer; K, longitudinal muscle layer. Tunica adventitia: L, adventitia; M, serosa.

[From Dellman after Krölling and Grau .]
Figure 27. Figure 27.

Scanning electron micrograph of luminal surface of monkey descending colon. Crypt openings are arranged in a regular array. × 510.

[From Specian and Neutra .]
Figure 28. Figure 28.

Light micrograph of human distal colon showing full extent of mucosa. Secretory granules of goblet cells stain darker in deeper crypt but lighter in foveola and surface. Principal cells are on surface and extend into crypt. MM, muscularis mucosa. × 315.

(Courtesy of Profs. Liliana Luciano and Enrico Reale.)
Figure 29. Figure 29.

Electron micrograph of guinea pig proximal colon showing surface principal cells (P), goblet cells (G), and vacuolated cells (V) in upper crypt. Freeze substitution, × 2,400.

(Courtesy of Profs. Liliana Luciano and Enrico Reale.)
Figure 30. Figure 30.

Freeze‐fracture replicas of guinea pig proximal (A) and distal (B) colonic principal cells showing strands of zonula occludens on P fracture face. Note density of intramembranous particles on microvilli (MV). Area of lateral plasma membrane subjacent to zonula occludens (L) is position of zonula adherens and has fewer intramembranous particles than other parts of lateral plasma membrane. Shadow direction is from bottom to top in both A and B. A, × 96,000; B, × 81,000.

[From Luciano et al. .]
Figure 31. Figure 31.

Electron micrograph of apical part of 2 adjacent surface principal cells from rat distal colon. Lateral plasma membranes are specialized into a junctional complex consisting of zonula occludens (ZO) and adjacent zonula adherens (ZA). A desmosome (D) is located more basally. Note distinct intermediate filaments on cytoplasmic side of desmosome. × 110,000.

Figure 32. Figure 32.

Electron micrograph of vacuolated cells lining crypt of guinea pig proximal colon. × 6,450.

(Courtesy of Profs. Liliana Luciano and Enrico Reale).
Figure 33. Figure 33.

High‐power electron micrograph of apical region of vacuolated cell in distal colonic crypt of guinea pig. Tannic acid added to fixative stains thick glycocalyx and fibrillar contents of vacuoles (V). × 30,800.

(Courtesy of Profs. Liliana Luciano and Enrico Reale).
Figure 34. Figure 34.

Electron micrographs of rat goblet cells along crypt wall. A: unstimulated cell with secretory granules (G) packed into apical cytoplasm. B: exocytotic pattern in which secretory granules have fused and expelled their contents after stimulation with acetylcholine eserine. A, × 6,300; B, × 3,670.

(Courtesy of Dr. Robert Specian.)
Figure 35. Figure 35.

Electron micrographs of 2 enterochromaffin (EC) cells obtained from human rectal biopsies. A: lightly staining cell, which has empty appearing granules (G) packed into basal cytoplasm. An occasional well‐formed secretory granule (SG) with more darkly stained contents is observed. Vacuolated dense bodies (arrows) are present in both EC cells and undifferentiated cells (UC). N, nucleus. × 19,750. B: EC cell showing localization of acid phosphatase. Reaction product is seen in supranuclear lysosomes (L), in some Golgi lamellae, and at periphery of some secretory granues. × 13,850.

[From Lorenzsson and Trier .]
Figure 36. Figure 36.

Electron micrograph of caveolated cell adjacent to vacuolated cells in rat distal colon. Note characteristic wide, straight microvilli from which bundles of long microfilaments project into apical cytoplasm. Arrowheads point to some caveolae. × 17,000.

(Courtesy of Profs. Liliana Luciano and Enrico Reale.)
Figure 37. Figure 37.

Diagram of external surface of large intestine showing aggregation of outer longitudinal muscle layers into 3 bands of taenae coli.

[From Gray .]
Figure 38. Figure 38.

Diagram showing distribution of lymphoid nodules in terminal small intestine and cecum of rabbit.

[Courtesy of Drs. Andreas Gebert and Helmut Bartels. Modified from Snipes .]
Figure 39. Figure 39.

Diagram of lymphoid follicle and associated tissue of rabbit appendix. See Table for description of cell types in each zone.

[From Waksman et al. .]
Figure 40. Figure 40.

Light micrograph of rabbit appendix showing luminal epithelium formed of lightly staining M cells covering clusters of lymphocytes (*) and more darkly staining columnar cells. Basal lamina (BL) separates epithelium from dome, which is rich in mixed blasts and some lymphocytes.

(Courtesy of Drs. Andreas Gebert and Helmut Bartels.)
Figure 41. Figure 41.

Freeze‐fracture replica of M cell of appendix showing extensive strands of zonula occludens extending along basolateral plasma membrane. × 2,500. Shadow direction is from bottom to top.

(Courtesy of Drs. Andreas Gebert and Helmut Bartels.)
Figure 42. Figure 42.

Freeze‐fracture replica of nondomed epithelial cell in cecal patch. Strands of tight junction (zonula occludens) occur on P fracture face as a honeycomb structure adjacent to the short wide microvilli.

(Courtesy of Drs. Andreas Gebert and Helmut Bartels).
Figure 43. Figure 43.

Schematic diagram illustrating distribution of epithelial cell types and zones within lower rectum and anal canal.

[Adapted from Fenger .]
Figure 44. Figure 44.

Light micrograph of histological section of colonic mucosa 30 min after damage by exposure to luminal HCl. Damage has been restricted to superficial epithelium on intercryptal surface. Epithelium has blistered into lumen, leaving basal lamina denuded (arrowheads) and forming subepithelial blisters. × 93.

[From Feil et al. .]
Figure 45. Figure 45.

Light micrograph of histological section of rabbit colon 5 h after luminal HCl exposure. Necrotic superficial epithelium has largely exfoliated into overlying mucus layer, forming a mucoid layer. Superficial surface and denuded basal lamina (arrowheads) have been recovered by migrating principal cells that now appear squamous to low cuboidal. × 62.

[From Feil et al. .]
Figure 46. Figure 46.

Transmission electron micrograph showing in rabbit colon a principal cell on left that has extended (arrow) a lamellipodium across denuded basal lamina (arrowhead) 30 min subsequent to luminal HCl exposure. Underlying capillary (Cap) remains patent. × 8,600.

[From Feil et al. .]


Figure 1.

A phylogeny of animal kingdom showing protostome and deuterostome divisions and their associated embryonic characteristics.

[From Barnes .]


Figure 2.

Diagram showing generalized arthropod body plan as viewed in a sagittal plane. Note that both foregut and hindgut have a portion of luminal surface covered by chitinous exoskeleton, illustrated here by a heavy dark line.

[From Barnes .]


Figure 3.

Diagrams illustrating relative positions of Malpighian tubules and rectum in relation to midgut and hindgut of insect. Specialization of rectum into rectal pads or papilla and ultrastructural anatomy of cells lining lumen are shown.

[Adapted from Berridge and Oschman .]


Figure 4.

Electron micrograph of a septate junction (desmosome) between epithelial cells lining cockroach rectum. Note ladderlike cross bridges spanning intercellular space.

(Courtesy of Dr. Peter Smith.)


Figure 5.

Diagram of digestive tracts of 6 representative aquatic vertebrates: lamprey (Petromyzon), shark (Squalus), chimaera (Callorhynchus), lungfish (Protopterus), sturgeon (Acipensus), and teleost (Perca).

[From Romer and Parsons .]


Figure 6.

Diagram of digestive tract and appendages of 4 vertebrates that are either amphibious or terrestrial. Note anatomic distinction between small and large intestine.

[Adapted from Romer and Parsons .]


Figure 7.

Diagram of large intestinal specializations found in birds. Arrows indicate retrograde flow of urine and feces mixture from cloaca into ceca.

[Adapted from Skadhauge .]


Figure 8.

Proportion of total body weight constituted by hindgut contents in various omnivores, carnivores, and herbivores.

[From von Engelhardt et al. after Stevens .]


Figure 9.

Diagram of gastrointestinal tract of a representative carnivore (A), omnivore (B), and 2 herbivores (C and D). Note extensive development of cecum and large intestine in herbivores.

[From von Engelhardt et al., after Stevens .]


Figure 10.

Diagram showing position and segments of adult human large intestine as it appears in body cavity. Greater omentum has been removed so transverse colon and coils of small intestine can be observed.

[From Basmajian .]


Figure 11.

Diagram of ileocaecal junction showing blood supply, lymph nodes, mesenteries, and orifice of ileum and appendix into cecum.

[From Snell .]


Figure 12.

Diagram of coronal section of adult terminal rectum, anal canal, anus, and adjacent structures.

[Adapted from Gray .]


Figure 13.

Diagram of adult human colon showing distribution of superior mesenteric artery and its branches that supply small intestine and appendix, cecum, and ascending and part of transverse colon (to arrow). Transverse colon has been elevated to show origin of artery.

[From Snell .]


Figure 14.

Diagram showing distribution of inferior mesenteric artery and its branches that supply distal one‐third of transverse, descending, and sigmoid colon, and rectum.

[From Hall‐Craggs .]


Figure 15.

Diagram showing arteries of rectum and anal canal from a frontal view. Note that superior rectal artery (Sup rectal a) gives a right and left branch, each of which anastomoses with branches of middle rectal artery, which is given off by internal pudendal artery (int pudendal a). Inferior rectal artery (inf rectal a) branches and anastomoses with branches from middle rectal artery.

[From Anderson .]


Figure 16.

Diagram showing distribution of lymphatic nodes and vessels of colon and upper rectum.

[From Jamieson and Dobson .]


Figure 17.

Diagrams showing various stages in development of gut tube as seen in sagittal midline section of human embryos. Arrows indicate extensions of amniotic cavity in presomite embryo (A), 7‐somite embryo (B), 14‐somite embryo (C), and end of month 1 (D).

[From Sadler .]


Figure 18.

Diagrams of transverse sections through human embryos at sequential stages of development from left to right. Effects of lateral folding on endoderm‐lined cavity are shown.

[From Sadler .]


Figure 19.

Diagram of human embryo to illustrate position of primitive intestinal loop and its continuity with vitelline duct. Note connection between allantois and cloaca.

[From Sadler .]


Figure 20.

Diagrams of primitive (prim) intestinal loop showing position of superior (sup) mesenteric artery (A) and position of cecal bud that designates division between small and large intestine after 180° rotation in primitive intestinal loop (B). Arrow in A indicates direction of counterclockwise rotation.

[From Sadler .]


Figure 21.

Diagram of 6‐wk‐old human embryo showing mesenteric connections with various parts of digestive tract.

[From Sadler .]


Figure 22.

Diagrams showing sequential development of anorectal canal and urogenital system in cloacal region. Arrow indicates path of descending urorectal septum.

[From Sadler .]


Figure 23.

Drawing of solid lumen phase of large intestinal development (A) and beginning of cavitation of lumen (B).

[From Sadler .]


Figure 24.

Light micrographs showing 4 stages in conversion of luminal epithelium from stratified to simple columnar in fetal rat colon. A: at 16 days, a stratified epithelium 2–3 cell layers thick surrounds a small lumen (L). × 600. B: at 17 days, main lumen (ML) projects into epithelium, forming deep clefts. In addition, secondary lumina (SL) are formed. × 600. C: by 19 days, secondary lumina (SL) are enlarged. Darkly stained and rounded apical cells (arrows) will be sloughed into main lumen. × 600. D: just before birth (22 days gestation), epithelium has changed to a simple columnar type that lines longitudinal ridges of connective tissue. Epithelium has become differentiated, as illustrated by goblet cells (GC). × 500.

[From Colony and Neutra .]


Figure 25.

Electron micrograph of nongoblet cell of 22‐day fetal rat proximal colon reacted for carbonic anhydrase activity, which is localized along lateral cell borders. Nu, nucleus. × 16,500.

[From Lacy and Colony .]


Figure 26.

Diagram of general structure and layers of various parts of gastrointestinal tract, including large intestine, as observed in cross section. Esophagus without submucosal glands (A), esophagus with submucosal glands (B), small intestine (C), and large intestine (D). Tunica Mucosa: E, epithelial layer (lamina epithelialis mucosae); F, lamina propria; G, lamina muscularis mucosa. Tunica submucosa: H, loose connective tissue and associated structures of submucosa. Tunica muscularis: I, circular muscle layer; K, longitudinal muscle layer. Tunica adventitia: L, adventitia; M, serosa.

[From Dellman after Krölling and Grau .]


Figure 27.

Scanning electron micrograph of luminal surface of monkey descending colon. Crypt openings are arranged in a regular array. × 510.

[From Specian and Neutra .]


Figure 28.

Light micrograph of human distal colon showing full extent of mucosa. Secretory granules of goblet cells stain darker in deeper crypt but lighter in foveola and surface. Principal cells are on surface and extend into crypt. MM, muscularis mucosa. × 315.

(Courtesy of Profs. Liliana Luciano and Enrico Reale.)


Figure 29.

Electron micrograph of guinea pig proximal colon showing surface principal cells (P), goblet cells (G), and vacuolated cells (V) in upper crypt. Freeze substitution, × 2,400.

(Courtesy of Profs. Liliana Luciano and Enrico Reale.)


Figure 30.

Freeze‐fracture replicas of guinea pig proximal (A) and distal (B) colonic principal cells showing strands of zonula occludens on P fracture face. Note density of intramembranous particles on microvilli (MV). Area of lateral plasma membrane subjacent to zonula occludens (L) is position of zonula adherens and has fewer intramembranous particles than other parts of lateral plasma membrane. Shadow direction is from bottom to top in both A and B. A, × 96,000; B, × 81,000.

[From Luciano et al. .]


Figure 31.

Electron micrograph of apical part of 2 adjacent surface principal cells from rat distal colon. Lateral plasma membranes are specialized into a junctional complex consisting of zonula occludens (ZO) and adjacent zonula adherens (ZA). A desmosome (D) is located more basally. Note distinct intermediate filaments on cytoplasmic side of desmosome. × 110,000.



Figure 32.

Electron micrograph of vacuolated cells lining crypt of guinea pig proximal colon. × 6,450.

(Courtesy of Profs. Liliana Luciano and Enrico Reale).


Figure 33.

High‐power electron micrograph of apical region of vacuolated cell in distal colonic crypt of guinea pig. Tannic acid added to fixative stains thick glycocalyx and fibrillar contents of vacuoles (V). × 30,800.

(Courtesy of Profs. Liliana Luciano and Enrico Reale).


Figure 34.

Electron micrographs of rat goblet cells along crypt wall. A: unstimulated cell with secretory granules (G) packed into apical cytoplasm. B: exocytotic pattern in which secretory granules have fused and expelled their contents after stimulation with acetylcholine eserine. A, × 6,300; B, × 3,670.

(Courtesy of Dr. Robert Specian.)


Figure 35.

Electron micrographs of 2 enterochromaffin (EC) cells obtained from human rectal biopsies. A: lightly staining cell, which has empty appearing granules (G) packed into basal cytoplasm. An occasional well‐formed secretory granule (SG) with more darkly stained contents is observed. Vacuolated dense bodies (arrows) are present in both EC cells and undifferentiated cells (UC). N, nucleus. × 19,750. B: EC cell showing localization of acid phosphatase. Reaction product is seen in supranuclear lysosomes (L), in some Golgi lamellae, and at periphery of some secretory granues. × 13,850.

[From Lorenzsson and Trier .]


Figure 36.

Electron micrograph of caveolated cell adjacent to vacuolated cells in rat distal colon. Note characteristic wide, straight microvilli from which bundles of long microfilaments project into apical cytoplasm. Arrowheads point to some caveolae. × 17,000.

(Courtesy of Profs. Liliana Luciano and Enrico Reale.)


Figure 37.

Diagram of external surface of large intestine showing aggregation of outer longitudinal muscle layers into 3 bands of taenae coli.

[From Gray .]


Figure 38.

Diagram showing distribution of lymphoid nodules in terminal small intestine and cecum of rabbit.

[Courtesy of Drs. Andreas Gebert and Helmut Bartels. Modified from Snipes .]


Figure 39.

Diagram of lymphoid follicle and associated tissue of rabbit appendix. See Table for description of cell types in each zone.

[From Waksman et al. .]


Figure 40.

Light micrograph of rabbit appendix showing luminal epithelium formed of lightly staining M cells covering clusters of lymphocytes (*) and more darkly staining columnar cells. Basal lamina (BL) separates epithelium from dome, which is rich in mixed blasts and some lymphocytes.

(Courtesy of Drs. Andreas Gebert and Helmut Bartels.)


Figure 41.

Freeze‐fracture replica of M cell of appendix showing extensive strands of zonula occludens extending along basolateral plasma membrane. × 2,500. Shadow direction is from bottom to top.

(Courtesy of Drs. Andreas Gebert and Helmut Bartels.)


Figure 42.

Freeze‐fracture replica of nondomed epithelial cell in cecal patch. Strands of tight junction (zonula occludens) occur on P fracture face as a honeycomb structure adjacent to the short wide microvilli.

(Courtesy of Drs. Andreas Gebert and Helmut Bartels).


Figure 43.

Schematic diagram illustrating distribution of epithelial cell types and zones within lower rectum and anal canal.

[Adapted from Fenger .]


Figure 44.

Light micrograph of histological section of colonic mucosa 30 min after damage by exposure to luminal HCl. Damage has been restricted to superficial epithelium on intercryptal surface. Epithelium has blistered into lumen, leaving basal lamina denuded (arrowheads) and forming subepithelial blisters. × 93.

[From Feil et al. .]


Figure 45.

Light micrograph of histological section of rabbit colon 5 h after luminal HCl exposure. Necrotic superficial epithelium has largely exfoliated into overlying mucus layer, forming a mucoid layer. Superficial surface and denuded basal lamina (arrowheads) have been recovered by migrating principal cells that now appear squamous to low cuboidal. × 62.

[From Feil et al. .]


Figure 46.

Transmission electron micrograph showing in rabbit colon a principal cell on left that has extended (arrow) a lamellipodium across denuded basal lamina (arrowhead) 30 min subsequent to luminal HCl exposure. Underlying capillary (Cap) remains patent. × 8,600.

[From Feil et al. .]
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Eric R. Lacy. Functional Morphology of the Large Intestine. Compr Physiol 2011, Supplement 19: Handbook of Physiology, The Gastrointestinal System, Intestinal Absorption and Secretion: 121-194. First published in print 1991. doi: 10.1002/cphy.cp060404