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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 .]
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 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 .]
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 . 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 .]
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 .]
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 , 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 .]
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 .]
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 .]

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 .]
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 . 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 . 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 .]


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 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 .]


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 . 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 .]


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 .]


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 , 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 .]


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 .]


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 .]



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 .]


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 . 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 . 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.

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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