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Comparative Physiology of Colonic Electrolyte Transport

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Abstract

The sections in this article are:

1 Comparative Structure of Large Intestine
2 Absorptive Load
3 Electrolyte Concentration Patterns
4 Electrolyte Transport Pattern in vivo
5 Basic Mechanisms of Electrolyte Transport
6 Absorption of Volatile Fatty Acid
7 Conclusions
Figure 1. Figure 1.

Digestive tracts of pig, dog, pony, and sheep. Arrows indicate termination of ascending colon. Major portion of colon of ruminants and pigs is arranged in 2 spirals. Centripetal spiral reverses direction to form centrifugal spiral, as indicated by slight upward bend near center of colon. Large (ascending) colon of pony is arranged in right and left ventral colons and left and right dorsal colons, which terminate at origin of small colon (transverse colon).

[From Stevens 62.]
Figure 2. Figure 2.

Volatile fatty acid (VFA) content, net VFA production, volume of water, and net transmural flux of water determined for individual segments of pony large intestine as a function of time. Negative values designate net absorption of VFA or water. Rates of flow determined by marker techniques 6 allowed volume and total VFA changes to be analyzed in terms of inflow and outflow to a segment and by difference, the rate of net addition or removal of VFA or water.

[From Argenzio 4.]
Figure 3. Figure 3.

Digesta osmolality and ion concentrations along gastrointestinal tract of pony. Data represent mean values (n = 24, where n is the number of ponies) obtained over 12‐h period between meals for ponies fed either conventional diet or high‐fiber diet 12. Concentration of PO43− was calculated under assumption that pKa of NaH2PO4 was 6.8 and utilizing mean pH of contents in each segment. At pH of large intestinal contents, NH3+ and organic acids (OA) exist primarily in their ionized forms. Concentration of HCO3 calculated as difference in concentration of measured cations and anions. S, stomach; SI1, SI2, SI3, 3 equal segments of small intestine; C, cecum; VC, ventral colon; DC, dorsal colon; SC, small colon.

[From Argenzio 4.]
Figure 4. Figure 4.

Mean concentrations (n = 9, where n is the number of dogs) of cations and anions along digestive tract of dog. Animals were killed 4–6 h after meat meal. S, stomach; J, jejunum; I, ileum; C, colon.

[Data from Alexander 2.]
Figure 5. Figure 5.

Possible mechanisms to account for results obtained from Ringer's and theophylline Ringer's solutions in proximal colon of pig. For simplicity, the models assume a single barrier to transport from lumen (L) to blood (B). In all likelihood, H ions and HCO3 ions are produced by intracellular hydration of CO2 36, so barrier shown probably represents mucosal membrane. All values expressed in μeq · cm−2 · h−1. Values for short‐circuit current (Isc) obtained from parallel in vitro experiments are included and are in reasonably good agreement with current flows calculated from in vivo experiments. Arrows between barriers represent net passive ion flows traversing tissue in response to transmural potential difference shown at bottom.

[From Argenzio and Whipp 14.]
Figure 6. Figure 6.

Net water movement in intestine of diarrhetic (shaded bars) and control (open bars) calves. Points examined were 15%, 40%, 65%, and 90% of distance from duodenum (D) to ileocecal valve (I). C, proximal spiral colon. Data are means ± SE of 10 animals in each group.

[From Bywater and Logan 22.]
Figure 7. Figure 7.

Partial pressure of CO2 initially present in luminal solution and after perfusion through proximal colon of pig. Initial was set to approximate that of plasma. Solutions consisted of isotonic NaCl, mannitol, or sodium acetate buffered with HCO3. The final of mannitol solution was not significantly different from plasma, whereas final of NaCl and NaAc solutions were, respectively, greater and less than plasma concentration (P < 0.001). [Modified from Argenzio and Whipp 13.]

Figure 8. Figure 8.

Relationship of net solute transport to net water transport in experiments with sodium acetate solution. Regression lines are shown over range of values encountered in these in vivo perfusion experiments of pig proximal colon. The 95% confidence limits are shown. Intercepts for HCO3, Na+, and acetate were −0.96 ± 0.06, 0.25 ± 0.06, and 1.4 ± 0.09 meq/min, respectively, at zero net water movement.

[From Argenzio and Whipp 13.]
Figure 9. Figure 9.

Possible mechanism to explain absorption of undissociated acid (HAc) and its partial dependence on Na+ absorption. A Na+‐H+ exchange mechanism on mucosal membrane provides additional source of luminal H ions to those resulting from consumption of luminal CO2, as originally postulated by Ash and Dobson 16. Thus these 2 sources of H ions, in an unstirred layer adjacent to mucosal membranes, could continuously regenerate HAc for absorption into cell, despite alkalinization of contents in main lumen. Driving force for overall process would be rapid removal of HAc from cell by metabolism or transport across serosal membranes 10. Increased steady‐state production of intracellular H ion due to HAc absorption and dissociation inside cell could result in increased uptake of Na+ observed in presence of acetate. When net acetate absorption exceeds net Na+ absorption, a net increase in luminal HCO3 and a decrease in would be observed. In this case excess of intracellular H ions would have to leave cell via serosal membranes (not shown). For equal rates of HAc absorption and Na+‐H+ exchange, net alkalinization of lumen would not occur and a net consumption of luminal CO2 would not be present.



Figure 1.

Digestive tracts of pig, dog, pony, and sheep. Arrows indicate termination of ascending colon. Major portion of colon of ruminants and pigs is arranged in 2 spirals. Centripetal spiral reverses direction to form centrifugal spiral, as indicated by slight upward bend near center of colon. Large (ascending) colon of pony is arranged in right and left ventral colons and left and right dorsal colons, which terminate at origin of small colon (transverse colon).

[From Stevens 62.]


Figure 2.

Volatile fatty acid (VFA) content, net VFA production, volume of water, and net transmural flux of water determined for individual segments of pony large intestine as a function of time. Negative values designate net absorption of VFA or water. Rates of flow determined by marker techniques 6 allowed volume and total VFA changes to be analyzed in terms of inflow and outflow to a segment and by difference, the rate of net addition or removal of VFA or water.

[From Argenzio 4.]


Figure 3.

Digesta osmolality and ion concentrations along gastrointestinal tract of pony. Data represent mean values (n = 24, where n is the number of ponies) obtained over 12‐h period between meals for ponies fed either conventional diet or high‐fiber diet 12. Concentration of PO43− was calculated under assumption that pKa of NaH2PO4 was 6.8 and utilizing mean pH of contents in each segment. At pH of large intestinal contents, NH3+ and organic acids (OA) exist primarily in their ionized forms. Concentration of HCO3 calculated as difference in concentration of measured cations and anions. S, stomach; SI1, SI2, SI3, 3 equal segments of small intestine; C, cecum; VC, ventral colon; DC, dorsal colon; SC, small colon.

[From Argenzio 4.]


Figure 4.

Mean concentrations (n = 9, where n is the number of dogs) of cations and anions along digestive tract of dog. Animals were killed 4–6 h after meat meal. S, stomach; J, jejunum; I, ileum; C, colon.

[Data from Alexander 2.]


Figure 5.

Possible mechanisms to account for results obtained from Ringer's and theophylline Ringer's solutions in proximal colon of pig. For simplicity, the models assume a single barrier to transport from lumen (L) to blood (B). In all likelihood, H ions and HCO3 ions are produced by intracellular hydration of CO2 36, so barrier shown probably represents mucosal membrane. All values expressed in μeq · cm−2 · h−1. Values for short‐circuit current (Isc) obtained from parallel in vitro experiments are included and are in reasonably good agreement with current flows calculated from in vivo experiments. Arrows between barriers represent net passive ion flows traversing tissue in response to transmural potential difference shown at bottom.

[From Argenzio and Whipp 14.]


Figure 6.

Net water movement in intestine of diarrhetic (shaded bars) and control (open bars) calves. Points examined were 15%, 40%, 65%, and 90% of distance from duodenum (D) to ileocecal valve (I). C, proximal spiral colon. Data are means ± SE of 10 animals in each group.

[From Bywater and Logan 22.]


Figure 7.

Partial pressure of CO2 initially present in luminal solution and after perfusion through proximal colon of pig. Initial was set to approximate that of plasma. Solutions consisted of isotonic NaCl, mannitol, or sodium acetate buffered with HCO3. The final of mannitol solution was not significantly different from plasma, whereas final of NaCl and NaAc solutions were, respectively, greater and less than plasma concentration (P < 0.001). [Modified from Argenzio and Whipp 13.]



Figure 8.

Relationship of net solute transport to net water transport in experiments with sodium acetate solution. Regression lines are shown over range of values encountered in these in vivo perfusion experiments of pig proximal colon. The 95% confidence limits are shown. Intercepts for HCO3, Na+, and acetate were −0.96 ± 0.06, 0.25 ± 0.06, and 1.4 ± 0.09 meq/min, respectively, at zero net water movement.

[From Argenzio and Whipp 13.]


Figure 9.

Possible mechanism to explain absorption of undissociated acid (HAc) and its partial dependence on Na+ absorption. A Na+‐H+ exchange mechanism on mucosal membrane provides additional source of luminal H ions to those resulting from consumption of luminal CO2, as originally postulated by Ash and Dobson 16. Thus these 2 sources of H ions, in an unstirred layer adjacent to mucosal membranes, could continuously regenerate HAc for absorption into cell, despite alkalinization of contents in main lumen. Driving force for overall process would be rapid removal of HAc from cell by metabolism or transport across serosal membranes 10. Increased steady‐state production of intracellular H ion due to HAc absorption and dissociation inside cell could result in increased uptake of Na+ observed in presence of acetate. When net acetate absorption exceeds net Na+ absorption, a net increase in luminal HCO3 and a decrease in would be observed. In this case excess of intracellular H ions would have to leave cell via serosal membranes (not shown). For equal rates of HAc absorption and Na+‐H+ exchange, net alkalinization of lumen would not occur and a net consumption of luminal CO2 would not be present.

References
 1. Alexander, F. The concentration of certain electrolytes in the digestive tract of the horse and pig. Res. Vet. Sci. 3: 78–83, 1962.
 2. Alexander, F. The concentration of electrolytes in the alimentary tract of the rabbit, guinea pig, dog, and cat. Res. Vet. Sci. 6: 238–244, 1965.
 3. Alexander, F., and J. C. D. Hickson. The salivary and pancreatic secretions of the horse. In: Physiology of Digestion and Metabolism in the Ruminant, edited by A. T. Phillipson. Newcastle upon Tyne, UK: Oriel, 1969, p. 375–389.
 4. Argenzio, R. A. Functions of the equine large intestine and their interrelationship in disease. Cornell Vet. 65: 303–330, 1975.
 5. Argenzio, R. A., and D. Lebo. Ion transport by the pig colon: effects of theophylline and dietary sodium restriction. Can. J. Physiol. Pharmacol. 60: 929–935, 1982.
 6. Argenzio, R. A., J. E. Lowe, D. W. Pickard, and C. E. Stevens. Digesta passage and water exchange in the equine large intestine. Am. J. Physiol. 226: 1035–1042, 1974.
 7. Argenzio, R. A., N. Miller, and W. von Englehardt. Effect of volatile fatty acids on water and ion absorption from the goat colon. Am. J. Physiol. 229: 997–1002, 1975.
 8. Argenzio, R. A., H. W. Moon, L. J. Kemeny, and S. C. Whipp. Colonic compensation in transmissible gastroenteritis of swine. Gastroenterology 86: 1501–1509, 1984.
 9. Argenzio, R. A., and M. Southworth. Sites of organic acid production and absorption in gastrointestinal tract of the pig. Am. J. Physiol. 228: 454–460, 1975.
 10. Argenzio, R. A., M. Southworth, J. E. Lowe, and C. E. Stevens. Interrelationship of Na, HCO3, and volatile fatty acid transport by equine large intestine. Am. J. Physiol. 233 (Endocrinol. Metab. Gastrointest. Physiol. 2): E469–E478, 1977.
 11. Argenzio, R. A., M. Southworth, and C. E. Stevens. Sites of organic acid production and absorption in the equine gastrointestinal tract. Am. J. Physiol. 226: 1043–1050, 1974.
 12. Argenzio, R. A., and C. E. Stevens. Cyclic changes in ionic composition of digesta in the equine intestinal tract. Am. J. Physiol. 228: 1224–1230, 1975.
 13. Argenzio, R. A., and S. C. Whipp. Interrelationship of sodium, chloride, bicarbonate, and acetate transport by the colon of the pig. J. Physiol. Lond. 295: 365–381, 1979.
 14. Argenzio, R. A., and S. C. Whipp. Effect of Escherichia coli heat‐stable enterotoxin, cholera toxin, and theophylline on ion transport in porcine colon. J. Physiol. Lond. 320: 469–487, 1981.
 15. Argenzio, R. A., and S. C. Whipp. Effect of theophylline and heat‐stable enterotoxin of Escherichia coli on transcellular and paracellular ion movement across isolated porcine colon. Can. J. Physiol. Pharmacol. 61: 1138–1148, 1983.
 16. Ash, R. W., and A. Dobson. The effect of absorption on the acidity of rumen contents. J. Physiol. Lond. 169: 39–61, 1963.
 17. Banta, C. A., E. T. Clemens, M. M. Krinsy, and C. E. Stevens. Sites of organic acid production and patterns of digesta movement in the gastrointestinal tract of dogs. J. Nutr. 109: 1592–1600, 1979.
 18. Bergman, E. N., R. S. Reid, M. G. Murray, J. M. Brockway, and F. G. Whitlaw. Interconversions and production of volatile fatty acids in the sheep rumen. Biochem. J. 97: 53–58, 1965.
 19. Binder, H. J., and C. L. Rawlins. The effect of conjugated dihydroxy bile salts on electrolyte transport in the rat colon. J. Clin. Invest. 52: 1460–1466, 1973.
 20. Binder, H. J., and C. L. Rawlins. Electrolyte transport across isolated large intestinal mucosa. Am. J. Physiol. 225: 1232–1239, 1973.
 21. Blair‐West, J. R., J. P. Coghlan, D. A. Denton, and R. D. Wright. Effect of endocrines on salivary glands. In: Handbook of Physiology. Alimentary Canal. Secretion, edited by C. F. Code and W. Heidel. Washington, DC: Am. Physiol. Soc., 1968, sect. 6, vol. II, chapt. 38, p. 633–664.
 22. Bywater, R. J., and E. F. Logan. The site and characteristics of intestinal water and electrolyte loss in Escherichia coli‐induced diarrhea in calves. J. Comp. Pathol. 84: 599–610, 1974.
 23. Chien, W.‐J., and C. E. Stevens. Coupled active transport of Na and Cl across forestomach epithelium. Am. J. Physiol. 223: 997–1003, 1972.
 24. Clarke, L., J. Moore, and H. Garner. Diurnal variation of plasma aldosterone in the horse: dependence on feeding schedule (Abstract). Physiologist 21: 21, 1978.
 25. Clauss, W. Circadian rhythms in Na transport. In: Intestinal Absorption and Secretion, edited by E. Skadhauge and K. Heintze, Lancaster, UK: MTP, 1984, p. 273–283.
 26. Clauss, W., J. Durr, and G. Rechkemmer. Characterization of conductive pathways in guinea pig distal colon in vitro. Am. J. Physiol. 248 (Gastrointest. Liver Physiol. 11): G176–G183, 1985.
 27. Clemens, E. T., C. E. Stevens, and M. Southworth. Sites of organic acid production and pattern of digesta movement in the gastrointestinal tract of swine. J. Nutr. 105: 759–768, 1975.
 28. Cremaschi, D., D. R. Ferguson, S. Henin, P. S. James, G. Meyer, and M. W. Smith. Post‐natal development of amiloride sensitive sodium transport in pig distal colon. J. Physiol. Lond. 292: 481–494, 1979.
 29. Crump, M. H., R. A. Argenzio, and S. C. Whipp. Effect of acetate on solute and water absorption from the pig colon. Am. J. Vet. Res. 41: 1565–1568, 1980.
 30. Donowitz, M. Ca2+ in the control of active intestinal Na and Cl transport: involvement in neurohumoral action. Am. J. Physiol. 245 (Gastrointest. Liver Physiol. 8): G165–G177, 1983.
 31. Edmonds, C. J., and J. Marriott. Factors influencing the electrical potential across the mucosa of rat colon. J. Physiol. Lond. 194: 437–450, 1968.
 32. Engelhardt, W. von, and G. Rechkemmer. Colonic transport of short‐chain fatty acids and the importance of the microclimate. In: Intestinal Absorption and Secretion, edited by E. Skadhauge and K. Heintze, Lancaster, UK: MTP, 1984, p. 93–101.
 33. Fan, C. C., R. G. Faust, and D. W. Powell. Coupled sodium chloride transport by rabbit ileal brush‐border membrane vesicles. Am. J. Physiol. 244 (Gastrointest. Liver Physiol. 7): G375–G385, 1983.
 34. Field, M. Regulation of small intestinal ion transport by cyclic nucleotides and calcium. In: Secretory Diarrhea, edited by M. Field, J. S. Fordtran, and S. G. Schultz. Bethesda, MD: Am. Physiol. Soc., 1980, chapt. 3, p. 21–30.
 35. Frizzell, R. A., K. Heintze, and C. P. Stewart. Mechanism of intestinal chloride secretion. In: Secretory Diarrhea, edited by M. Field, J. S. Fordtran, and S. G. Schultz. Bethesda, MD: Am. Physiol. Soc., 1980, chapt. 2, p. 11–19.
 36. Frizzell, R. A., M. J. Koch, and S. G. Schultz. Ion transport by rabbit colon. I. Active and passive components. J. Membr. Biol. 27: 297–316, 1976.
 37. Frizzell, R. A., and S. G. Schultz. Effect of aldosterone on ion transport by rabbit colon in vitro. J. Membr. Biol. 39: 1–26, 1978.
 38. Fromm, M., and U. Hegel. Segmental heterogeneity of epithelial transport in rat large intestine. Pfluegers Arch. 378: 71–83, 1978.
 39. Fromm, M., and U. Hegel. Aldosterone action in different segments of large intestine. In: Intestinal Absorption and Secretion, edited by E. Skadhauge and K. Heintze, Lancaster, UK: MTP, 1984, p. 233–249.
 40. Heintze, K., and K.‐U. Petersen. Na/H and Cl/HCO3 exchange as a mechanism for HCO3‐stimulated NaCl absorption by gall bladder. In: Hydrogen Ion Transport in Epithelia, edited by I. Schultz, G. Sachs, J. G. Forte, and K. J. Ullrich. Amsterdam: Elsevier/North‐Holland, 1980, p. 345–354.
 41. Herschel, D. A., R. A. Argenzio, M. Southworth, and C. E. Stevens. Absorption of volatile fatty acid, Na, and H2O by the colon of the dog. Am. J. Vet. Res. 42: 1118–1124, 1981.
 42. Hubel, K. A. Bicarbonate secretion in rat ileum and its dependence on intraluminal chloride. Am. J. Physiol. 213: 1409–1413, 1967.
 43. Hubel, K. A. Effect of luminal chloride concentration on bicarbonate secretion in rat ileum. Am. J. Physiol. 217: 40–45, 1969.
 44. Hungate, R. E. Ruminal fermentation. In: Handbook of Physiology. Alimentary Canal. Bile; Digestion; Ruminal Physiology, edited by C. F. Code. Washington, DC: Am. Physiol. Soc., 1968, sect. 6, vol. V, chapt. 130, p. 2725–2745.
 45. Hyden, S. The use of reference substances and the measurement of flow in the alimentary tract. In: Digestive Physiology and Nutrition of the Ruminant, edited by D. Lewis, London: Butterworths, 1961, p. 35–47.
 46. Kenelly, J. J., F. X. Aherne, and W. C. Sauer. Volatile fatty acid production in the hindgut of swine. Can. J. Anim. Sci. 61: 349–358, 1981.
 47. Knickelbein, R., P. S. Aronson, W. Atherton, and J. W. Dobbins. Sodium and chloride transport across rabbit ileal brush border. I. Evidence for Na‐H exchange. Am. J. Physiol. 245 (Gastrointest. Liver Physiol. 8): G504–G510, 1983.
 48. Knickelbein, R., P. S. Aronson, C. M. Schron, J. Seifter, and J. W. Dobbins. Sodium and chloride transport across rabbit ileal brush border. II. Evidence for Cl−‐HCO3− exchange and mechanism of coupling. Am. J. Physiol. 249 (Gastrointest. Liver Physiol. 12): G236–G245, 1985.
 49. Low, A. G., I. G. Partridge, and I. E. Sambrook. Studies on digestion and absorption in the intestines of growing pigs. 2. Measurements of the flow of dry matter, ash, and water. Br. J. Nutr. 39: 515–526, 1978.
 50. Mylrea, P. J. Digestion of milk in young calves. II. The absorption of nutrients from the small intestine. Res. Vet. Sci. 7: 394–406, 1966.
 51. Nellans, H. N., R. A. Frizzell, and S. G. Schultz. Coupled sodium‐chloride influx across the brush border of rabbit ileum. Am. J. Physiol. 225: 467–475, 1973.
 52. Podestra, R. B., and D. F. Mettrick. HCO3 transport in rat jejunum: relationship to NaCl and H2O transport in vivo. Am. J. Physiol. 232 (Endocrinol. Metab. Gastrointest. Physiol. 1): E62–E68, 1977.
 53. Powell, D. W., and C.‐C. Fan. Coupled NaCl transport: cotransport or parallel ion exchange? In: Mechanisms of Intestinal Electrolyte Transport and Regulation by Calcium, edited by M. Donowitz and G. W. G. Sharp. New York: Liss, 1984, p. 13–26.
 54. Rechkemmer, G., and W. von Englehardt. Absorptive processes in different colonic segments of the guinea pig and the effects of short chain fatty acids. In: Colon and Nutrition, edited by W. Kasper and H. Goebell, Lancaster, UK: MTP, 1982, p. 61–68.
 55. Robinson, C. S., H. Luckey, and H. Mills. Factors affecting the hydrogen ion concentration of the contents of the small intestine. J. Biol. Chem. 147: 175–181, 1943.
 56. Robinson, J. W. L. Inhibition of transport processes in the dog colon. In: Intestinal Ion Transport, edited by J. W. L. Robinson. Lancaster, UK: MTP, 1976, p. 287–299.
 57. Rubsamen, K., and W. von Englehardt. Absorption of Na, H ions, and short chain fatty acids from the sheep colon. Pfluegers Arch. 391: 141–146, 1981.
 58. Ruppin, H., S. Bar‐Meir, and K. H. Soergel. Absorption of short‐chain fatty acids by the colon. Gastroenterology 78: 1500–1507, 1980.
 59. Schmall, L. M., R. A. Argenzio, and S. C. Whipp. Effects of intravenous endotoxin in gastrointestinal function in the equine (Abstract). Proc. Conf. Res. Workers Anim. Dis., Chicago, 1982.
 60. Sellin, J. H., and R. De Soignie. Rabbit proximal colon: a distinct transport epithelium. Am. J. Physiol. 246 (Gastrointest. Liver Physiol. 9): G603–G610, 1984.
 61. Stevens, C. E. Transport across rumen epithelium. In: Transport Mechanisms in Epithelia, edited by H. H. Ussing and N. A. Thorn. Copenhagen: Munksgaard, 1973, p. 10–27. (Alfred Benzon Symp. 5.)
 62. Stevens, C. E. Comparative physiology of the digestive system. In: Dukes' Physiology of Domestic Animals (9th ed.), edited by M. J. Swenson. Ithaca, NY: Cornell Univ. Press, 1977, p. 216–232.
 63. Tapper, E. J. Local modulation of intestinal ion transport by enteric neurons. Am. J. Physiol. 244 (Gastrointest. Liver Physiol. 7): G457–G468, 1983.
 64. Turnberg, L. A., F. A. Bieberdorf, S. G. Morawski, and J. S. Fordtran. Interrelationships of chloride, bicarbonate, sodium, and hydrogen transport in the human ileum. J. Clin. Invest. 49: 557–567, 1970.
 65. Turnberg, L. A., J. S. Fordtran, N. W. Carter, and F. C. Rector. Mechanism of bicarbonate absorption and its relationship to sodium transport in the human jejunum. J. Clin. Invest. 49: 548–556, 1970.
 66. Umesaki, Y., T. Yajima, T. Yokokura, and M. Mutai. Effect of organic acid absorption on bicarbonate transport in rat colon. Pfluegers Arch. 379: 43–47, 1979.
 67. Ussing, H. H. The distinction by means of tracers between active transport and diffusion. Acta Physiol. Scand. 19: 43–56, 1949.
 68. Welsh, M. J., P. L. Smith, M. Fromm, and R. A. Frizzell. Crypts are the site of intestinal fluid and electrolyte secretion. Science Wash. DC 218: 1219–1221, 1982.
 69. Will, P. C., J. L. Lebowitz, and V. Hopfer. Induction of amiloride‐sensitive sodium transport in the rat colon by mineralocorticoids. Am. J. Physiol. 238 (Renal Fluid Electrolyte Physiol. 7): F261–F268, 1980.

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How to Cite

Robert A. Argenzio. Comparative Physiology of Colonic Electrolyte Transport. Compr Physiol 2011, Supplement 19: Handbook of Physiology, The Gastrointestinal System, Intestinal Absorption and Secretion: 275-285. First published in print 1991. doi: 10.1002/cphy.cp060409