Comprehensive Physiology Wiley Online Library

Pancreatic Secretion of Electrolytes and Water

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Pancreatic Structure
1.1 Ductal Tree
1.2 Ductal Cells
1.3 Acinar Cells
1.4 Intercellular Junctions
2 Methods of Study
2.1 Intact Gland Preparations
2.2 Ductal Perfusion
2.3 Micropuncture
2.4 Isolated Ductal Tissue
3 Species‐Dependent Patterns of Electrolyte Secretion
3.1 Dog, Cat, and Human
3.2 Rat and Mouse
3.3 Rabbit
3.4 Pig
3.5 Guinea Pig and Hamster
3.6 Sheep, Cow, and Horse
3.7 Primates
4 Sites of Electrolyte and Water Secretion
4.1 Secretin‐Stimulated Bicarbonate Secretion
4.2 Secretion Evoked by Cholecystokinin and Vagal Stimulation
5 Permeability of The Pancreatic Epithelium
5.1 Nonelectrolytes
5.2 Electrolytes
6 Cellular Mechanisms of Electrolyte Secretion from Acinar Cells
6.1 Ionic Requirements for Secretion
6.2 Electrophysiology and Stimulus‐Secretion Coupling
6.3 Cellular Models of Acinar Cell Secretion
7 Cellular Mechanisms of Electrolyte Secretion from Duct Cells
7.1 Stimulus‐Secretion Coupling
7.2 Ionic Requirements and Mechanisms of Secretion
8 Cellular Models of Duct Cell Secretion
9 Future Progress
Figure 1. Figure 1.

A: scanning electron‐microscopic view of rat pancreatic lobule treated with HCl to remove connective tissue. Arrowheads, long intercalated duct that connects with numerous acini (A). × 669. B: scanning electron‐microscopic overview of ductal system of rat pancreas after removal of most acini by ultrasonic vibration. Excretory duct (D) branches dichotomously. Arrows, intercalated ducts. × 220.

From Takahashi
Figure 2. Figure 2.

Low‐power micrograph of a segment from a humar, acinus. In acinar cells, polarized arrangement of rough endoplasmic reticulum, Golgi complex (G), and zymogen granules in relation to the nucleus (N) is evident. Parts of 2 centroacinar cells (cac) close to the lumen (L) are visible. Cp, capillary, × 6,000.

From Kern . In: The Exocrine Pancreas: Biology, Pathobiology, and Diseases, © 1986, Raven Press, New York
Figure 3. Figure 3.

A. Fine structural characteristics of intralobular ducts in human pancreas. Large number of mitochondria, especially in vicinity of Golgi complex (G). L, duct lumen, × 12,000. B. Fine structural characteristics of intralobular ducts in human pancreas. Interdigitations between lateral plasma membranes in adjacent cells, short microvilli on the luminal membrane, and cilia in the duct lumen. × 12,000.

From Kern . In: The Exocrine Pancreas: Biology, Pathobiology, and Diseases, © 1986, Raven Press, New York. From Kern . In: The Exocrine Pancreas: Biology, Pathobiology, and Diseases, © 1986, Raven Press, New York
Figure 4. Figure 4.

Fine structure of an interlobular duct in the human pancreas, nf, Nerve fibers in lamina propria. × 5,700.

From Kern . In: The Exocrine Pancreas: Biology, Pathobiology, and Diseases, © 1986, Raven Press, New York
Figure 5. Figure 5.

Schematic diagram of isolated rabbit pancreas and attached duodenum mounted on a polyvinylchloride frame, which is then suspended in a chamber filled with a physiological buffer solution. Stippled area, pancreatic tissue.

(Diagram courtesy of J. J. H. H. M. De Pont.)
Figure 6. Figure 6.

Vascular supply of cat pancreas. Stomach is shown separated from the duodenum, displaced anteriorly, and turned through 180°. A, aorta; CA, celiac axis; GDA, gastroduodenal artery; GSV, gastrosplenic vein; HA, hepatic artery; LA, lumbar artery; LGA, left gastric artery; PV, portal vein; RGA. right gastric artery; SA, splenic artery; SMA, superior mesenteric artery; SMV, superior mesenteric vein.

From Case et al.
Figure 7. Figure 7.

Electrolyte composition of pancreatic juice in anesthetized cat stimulated to secrete at different flow rates by continuous infusion of different doses of secretin. Flow rate is expressed per gram gland weight (cf. Fig. by assuming a gland weight of 10 g; this is only approximate.

From Case
Figure 8. Figure 8.

Electrolyte composition of pancreatic juice in the anesthetized rat stimulated by secretin (A), caerulein (an analogue of cholecystokinin) (B), and a combination of secretin and caerulein (C).

From Case ; data from Sewell and Young
Figure 9. Figure 9.

Electrolyte composition of pancreatic juice in the anesthetized rabbit stimulated with secretin (A) and cholecystokinin (CCK) (B).

Data from Seow and Young a)
Figure 10. Figure 10.

Electrolyte composition of pancreatic juice in anesthetized guinea pig stimulated with secretin (A) and cholecystokinin‐octapeptide (CCK‐8) (B). Broken line, residual anions (Na+ + K+ ‐ Cl), almost all bicarbonate. Flow rate is uncorrected for body weight or gland weight.

Data from Case et al. a)
Figure 11. Figure 11.

Effect of increasing doses of secretin on flow rate and electrolyte composition of pancreatic juice in anesthetized Syrian golden hamster. (Unpublished data of A. E. Ali, R. M. Case, and S. C. B. Rutishauser.)

Figure 12. Figure 12.

Effects of secretin on membrane potential and input resistance of rat pancreatic duct cell. An isolated interlobular duct that had been maintained in culture for 24 h was first deflated by sectioning an end. Deflated duct was then placed in a tissue bath and held steady by drawing each end into a micropipette. Because the lumen was not cannulated and drained in these experiments, accumulation of secreted fluid often caused an obvious dilatation of ducts. Bath (volume 1 ml) was perfused with a Krebs‐Ringer bicarbonate buffer ml/min) at 37°C. Potassium acetate‐filled microelectrode was then used to impale a duct cell across its basolateral membrane. A, lower of 2 traces shows duct cell membrane potential. Every 2 s a depolarizing square‐wave current pulse (.5 nA; 100 ms) was applied to the microelectrode. Upper trace, maximum change in membrane potential associated with these current pulses. X and Y, magnitude of injected current was varied to construct a current‐voltage plot. Z, single dose of secretin was injected into the bath to give an instantaneous hormone concentration of 10−8 M. B: input resistance calculated from deflections in membrane potential caused by injected current pulses.

From Gray et al. a)
Figure 13. Figure 13.

Tentative models for cellular mechanisms of NaCl secretion by acinar cells of pig (A) and perhaps human, mouse (B), and rat (C). Ach, acetylcholine; CCK, cholecystokinin; GRP, gastrin‐releasing peptide.

Figure 14. Figure 14.

Possible general mechanisms of active bicarbonate transport by pancreatic duct cell. Experimental evidence argues strongly in favor of either model B or C, which cannot be distinguished mechanistically. Both models could be written in many different ways. For example, hydration steps shown occurring in the lumen in B could equally take place inside the cell. In this case the bicarbonate ions generated would have to move across the apical membrane and into the lumen, possibly by diffusion down an electrochemical gradient. Bicarbonate ions might also enter the cell on an anion carrier located on the basolateral membrane. Once inside the cell they could then combine with protons being pumped back from the lumen to form carbon dioxide, which then diffuses across the apical membrane.

Figure 15. Figure 15.

Possible models for transport of weak organic acids and organic anions by pancreatic ductal epithelium. Such compounds can replace bicarbonate ions and support secretion. A: sulfamerazine. Secretion rates obtained with this compound depend on concentration of the lipid‐soluble undissociated buffer component on blood side of the cell. This indicates that the weak acid delivers protons to the lumen by diffusing across the epithelium. B: glycodiazine, actetate, butyrate, formate, and propionate. Secretion rates obtained with these compounds depend on the concentration of the buffer anion on the blood side of the cell. Presumably this means that the anions cross the basolateral membrane of the duct cell on a carrier. Once inside the cell they could combine with protons moving back from the lumen and then diffuse across the apical membrane as undissociated buffer components.

Figure 16. Figure 16.

Cellular model for duct cell electrolyte secretion. Forward (blood‐to‐lumen) transport of bicarbonate results from backward (lumen‐to‐blood) transport of protons. Primary (Na+‐H+‐ATPase) and secondary (Na+‐H+ exchange) active transport steps are located at luminal and basolateral membranes, respectively.

From Swanson and Solomon
Figure 17. Figure 17.

Cellular model for duct cell electrolyte secretion. Forward (blood‐to‐cell) transport of bicarbonate results from backward (cell‐to‐blood) transport of protons. Secondary active transport step (Na+‐H+ exchange) is located at the basolateral membrane, and bicarbonate leaves the cell down an electrochemical gradient across luminal membrane.

From De Pont et al. . In: Electrolyte and Water Transport Across Gastrointestinal Epithelia, © 1982, Raven Press, New York
Figure 18. Figure 18.

Cellular model for duct cell electrolyte secretion. Forward (blood‐to‐cell) transport of bicarbonate is driven by a secondary active transport process (Na+‐2HCO3‐Cl exchange) on basolateral membrane, and bicarbonate leaves the cell down an electrochemical gradient across the luminal membrane.

From Kuijpers et al.
Figure 19. Figure 19.

Small interlobular ducts dissected from pancreas of a copper‐deficient rat. A: collection of 10 ducts. Phase contrast. Bar, 250 μm. B: higher magnification of interlobular duct. Lumen (lu), epithelium (ep), and connective tissue layer (ct) are visible. Phase contrast. Bar, 25 μm. C: interlobular duct from which connective tissue layer has been microdissected. This low‐magnification view shows that duct was ∼1 mm in length. Bar, 100 μm. D: small projections visible on left side of duct are intralobular branches (ib) that have fractured close to their site of origin. Bar, 50 μm. E: apical region of individual cells can be seen clearly when the objective lens is focused on luminal membrane of epithelium. Bar, 10 μm. Bright‐field optics. Plates C‐E obtained from a videotape by photographing monitor screen.

From Arkle et al.
Figure 20. Figure 20.

Measurement of fluid secretion from isolated pancreatic duct using micropuncture techniques. Interlobular duct had been maintained in culture for 24 h, during which time the ends of the duct had sealed. Subsequent accumulation of fluid in closed luminal space has caused a dilatation of the lumen, a flattening of the epithelium against the connective tissue layer, and overall swelling of the duct. A: duct is first immobilized by applying a suction pipette (sp) to its outer connective tissue layer and then micropunctured using a beveled, oil‐filled collection pipette (cp). B: success is confirmed by injection of a small volume of colored oil into the lumen. C: duct is deflated by aspirating luminal fluid into the collection pipette. Pipette is then withdrawn from the lumen, the fluid ejected to waste in the tissue bath, and the duct immediately repunctured along the same entry track. Usually the collection period is then started by application of subatmospheric pressure to the pipette. D: if suction is not applied to the collection pipette, secreted fluid accumulates within the closed lumen of the duct, causing it to dilate. Photograph was taken 40 min after C, during which time the duct was perifused with Krebs‐Ringer bicarbonate buffer at 37°C. Phase contrast. Bars, 200 μm.

From Argent et al.
Figure 21. Figure 21.

Effects of secretin, bicarbonate ions, and dibutyryl cAMP on fluid secretion from isolated interlobular ducts that had been maintained in culture for 16–52 h. Individual ducts were micropunctured as shown in Fig. and secreted fluid was collected. At the end of the collection period the dimensions of the epithelium were measured and epithelial volume was calculated. Secretion rates, plotted as mean ± SE, are expressed as nl·h−1·nl−1 duct epithelium. •—•, Dose‐response curve for effect of secretin on fluid secretion from isolated ducts. Each point is mean of 4–12 observations on different ducts. ▵—▵, For comparison the dose‐response curve for secretin‐stimulated fluid secretion from the perfused copper‐replete rat pancreas is also shown. Redrawn from data in ref. 113 assuming that 1 clinical unit of secretin = 7.5 × 10−11 mol . ○, Effect of replacing bicarbonate ions in perifusion buffer with HEPES on secretin‐stimulated fluid secretion from isolated ducts. Mean of 6 observations on different ducts. □, Effect of 2 × 10−4 M dibutyryl cAMP on fluid secretion from isolated ducts. Mean of 7 observations on different ducts.

Data from Argent et al.


Figure 1.

A: scanning electron‐microscopic view of rat pancreatic lobule treated with HCl to remove connective tissue. Arrowheads, long intercalated duct that connects with numerous acini (A). × 669. B: scanning electron‐microscopic overview of ductal system of rat pancreas after removal of most acini by ultrasonic vibration. Excretory duct (D) branches dichotomously. Arrows, intercalated ducts. × 220.

From Takahashi


Figure 2.

Low‐power micrograph of a segment from a humar, acinus. In acinar cells, polarized arrangement of rough endoplasmic reticulum, Golgi complex (G), and zymogen granules in relation to the nucleus (N) is evident. Parts of 2 centroacinar cells (cac) close to the lumen (L) are visible. Cp, capillary, × 6,000.

From Kern . In: The Exocrine Pancreas: Biology, Pathobiology, and Diseases, © 1986, Raven Press, New York


Figure 3.

A. Fine structural characteristics of intralobular ducts in human pancreas. Large number of mitochondria, especially in vicinity of Golgi complex (G). L, duct lumen, × 12,000. B. Fine structural characteristics of intralobular ducts in human pancreas. Interdigitations between lateral plasma membranes in adjacent cells, short microvilli on the luminal membrane, and cilia in the duct lumen. × 12,000.

From Kern . In: The Exocrine Pancreas: Biology, Pathobiology, and Diseases, © 1986, Raven Press, New York. From Kern . In: The Exocrine Pancreas: Biology, Pathobiology, and Diseases, © 1986, Raven Press, New York


Figure 4.

Fine structure of an interlobular duct in the human pancreas, nf, Nerve fibers in lamina propria. × 5,700.

From Kern . In: The Exocrine Pancreas: Biology, Pathobiology, and Diseases, © 1986, Raven Press, New York


Figure 5.

Schematic diagram of isolated rabbit pancreas and attached duodenum mounted on a polyvinylchloride frame, which is then suspended in a chamber filled with a physiological buffer solution. Stippled area, pancreatic tissue.

(Diagram courtesy of J. J. H. H. M. De Pont.)


Figure 6.

Vascular supply of cat pancreas. Stomach is shown separated from the duodenum, displaced anteriorly, and turned through 180°. A, aorta; CA, celiac axis; GDA, gastroduodenal artery; GSV, gastrosplenic vein; HA, hepatic artery; LA, lumbar artery; LGA, left gastric artery; PV, portal vein; RGA. right gastric artery; SA, splenic artery; SMA, superior mesenteric artery; SMV, superior mesenteric vein.

From Case et al.


Figure 7.

Electrolyte composition of pancreatic juice in anesthetized cat stimulated to secrete at different flow rates by continuous infusion of different doses of secretin. Flow rate is expressed per gram gland weight (cf. Fig. by assuming a gland weight of 10 g; this is only approximate.

From Case


Figure 8.

Electrolyte composition of pancreatic juice in the anesthetized rat stimulated by secretin (A), caerulein (an analogue of cholecystokinin) (B), and a combination of secretin and caerulein (C).

From Case ; data from Sewell and Young


Figure 9.

Electrolyte composition of pancreatic juice in the anesthetized rabbit stimulated with secretin (A) and cholecystokinin (CCK) (B).

Data from Seow and Young a)


Figure 10.

Electrolyte composition of pancreatic juice in anesthetized guinea pig stimulated with secretin (A) and cholecystokinin‐octapeptide (CCK‐8) (B). Broken line, residual anions (Na+ + K+ ‐ Cl), almost all bicarbonate. Flow rate is uncorrected for body weight or gland weight.

Data from Case et al. a)


Figure 11.

Effect of increasing doses of secretin on flow rate and electrolyte composition of pancreatic juice in anesthetized Syrian golden hamster. (Unpublished data of A. E. Ali, R. M. Case, and S. C. B. Rutishauser.)



Figure 12.

Effects of secretin on membrane potential and input resistance of rat pancreatic duct cell. An isolated interlobular duct that had been maintained in culture for 24 h was first deflated by sectioning an end. Deflated duct was then placed in a tissue bath and held steady by drawing each end into a micropipette. Because the lumen was not cannulated and drained in these experiments, accumulation of secreted fluid often caused an obvious dilatation of ducts. Bath (volume 1 ml) was perfused with a Krebs‐Ringer bicarbonate buffer ml/min) at 37°C. Potassium acetate‐filled microelectrode was then used to impale a duct cell across its basolateral membrane. A, lower of 2 traces shows duct cell membrane potential. Every 2 s a depolarizing square‐wave current pulse (.5 nA; 100 ms) was applied to the microelectrode. Upper trace, maximum change in membrane potential associated with these current pulses. X and Y, magnitude of injected current was varied to construct a current‐voltage plot. Z, single dose of secretin was injected into the bath to give an instantaneous hormone concentration of 10−8 M. B: input resistance calculated from deflections in membrane potential caused by injected current pulses.

From Gray et al. a)


Figure 13.

Tentative models for cellular mechanisms of NaCl secretion by acinar cells of pig (A) and perhaps human, mouse (B), and rat (C). Ach, acetylcholine; CCK, cholecystokinin; GRP, gastrin‐releasing peptide.



Figure 14.

Possible general mechanisms of active bicarbonate transport by pancreatic duct cell. Experimental evidence argues strongly in favor of either model B or C, which cannot be distinguished mechanistically. Both models could be written in many different ways. For example, hydration steps shown occurring in the lumen in B could equally take place inside the cell. In this case the bicarbonate ions generated would have to move across the apical membrane and into the lumen, possibly by diffusion down an electrochemical gradient. Bicarbonate ions might also enter the cell on an anion carrier located on the basolateral membrane. Once inside the cell they could then combine with protons being pumped back from the lumen to form carbon dioxide, which then diffuses across the apical membrane.



Figure 15.

Possible models for transport of weak organic acids and organic anions by pancreatic ductal epithelium. Such compounds can replace bicarbonate ions and support secretion. A: sulfamerazine. Secretion rates obtained with this compound depend on concentration of the lipid‐soluble undissociated buffer component on blood side of the cell. This indicates that the weak acid delivers protons to the lumen by diffusing across the epithelium. B: glycodiazine, actetate, butyrate, formate, and propionate. Secretion rates obtained with these compounds depend on the concentration of the buffer anion on the blood side of the cell. Presumably this means that the anions cross the basolateral membrane of the duct cell on a carrier. Once inside the cell they could combine with protons moving back from the lumen and then diffuse across the apical membrane as undissociated buffer components.



Figure 16.

Cellular model for duct cell electrolyte secretion. Forward (blood‐to‐lumen) transport of bicarbonate results from backward (lumen‐to‐blood) transport of protons. Primary (Na+‐H+‐ATPase) and secondary (Na+‐H+ exchange) active transport steps are located at luminal and basolateral membranes, respectively.

From Swanson and Solomon


Figure 17.

Cellular model for duct cell electrolyte secretion. Forward (blood‐to‐cell) transport of bicarbonate results from backward (cell‐to‐blood) transport of protons. Secondary active transport step (Na+‐H+ exchange) is located at the basolateral membrane, and bicarbonate leaves the cell down an electrochemical gradient across luminal membrane.

From De Pont et al. . In: Electrolyte and Water Transport Across Gastrointestinal Epithelia, © 1982, Raven Press, New York


Figure 18.

Cellular model for duct cell electrolyte secretion. Forward (blood‐to‐cell) transport of bicarbonate is driven by a secondary active transport process (Na+‐2HCO3‐Cl exchange) on basolateral membrane, and bicarbonate leaves the cell down an electrochemical gradient across the luminal membrane.

From Kuijpers et al.


Figure 19.

Small interlobular ducts dissected from pancreas of a copper‐deficient rat. A: collection of 10 ducts. Phase contrast. Bar, 250 μm. B: higher magnification of interlobular duct. Lumen (lu), epithelium (ep), and connective tissue layer (ct) are visible. Phase contrast. Bar, 25 μm. C: interlobular duct from which connective tissue layer has been microdissected. This low‐magnification view shows that duct was ∼1 mm in length. Bar, 100 μm. D: small projections visible on left side of duct are intralobular branches (ib) that have fractured close to their site of origin. Bar, 50 μm. E: apical region of individual cells can be seen clearly when the objective lens is focused on luminal membrane of epithelium. Bar, 10 μm. Bright‐field optics. Plates C‐E obtained from a videotape by photographing monitor screen.

From Arkle et al.


Figure 20.

Measurement of fluid secretion from isolated pancreatic duct using micropuncture techniques. Interlobular duct had been maintained in culture for 24 h, during which time the ends of the duct had sealed. Subsequent accumulation of fluid in closed luminal space has caused a dilatation of the lumen, a flattening of the epithelium against the connective tissue layer, and overall swelling of the duct. A: duct is first immobilized by applying a suction pipette (sp) to its outer connective tissue layer and then micropunctured using a beveled, oil‐filled collection pipette (cp). B: success is confirmed by injection of a small volume of colored oil into the lumen. C: duct is deflated by aspirating luminal fluid into the collection pipette. Pipette is then withdrawn from the lumen, the fluid ejected to waste in the tissue bath, and the duct immediately repunctured along the same entry track. Usually the collection period is then started by application of subatmospheric pressure to the pipette. D: if suction is not applied to the collection pipette, secreted fluid accumulates within the closed lumen of the duct, causing it to dilate. Photograph was taken 40 min after C, during which time the duct was perifused with Krebs‐Ringer bicarbonate buffer at 37°C. Phase contrast. Bars, 200 μm.

From Argent et al.


Figure 21.

Effects of secretin, bicarbonate ions, and dibutyryl cAMP on fluid secretion from isolated interlobular ducts that had been maintained in culture for 16–52 h. Individual ducts were micropunctured as shown in Fig. and secreted fluid was collected. At the end of the collection period the dimensions of the epithelium were measured and epithelial volume was calculated. Secretion rates, plotted as mean ± SE, are expressed as nl·h−1·nl−1 duct epithelium. •—•, Dose‐response curve for effect of secretin on fluid secretion from isolated ducts. Each point is mean of 4–12 observations on different ducts. ▵—▵, For comparison the dose‐response curve for secretin‐stimulated fluid secretion from the perfused copper‐replete rat pancreas is also shown. Redrawn from data in ref. 113 assuming that 1 clinical unit of secretin = 7.5 × 10−11 mol . ○, Effect of replacing bicarbonate ions in perifusion buffer with HEPES on secretin‐stimulated fluid secretion from isolated ducts. Mean of 6 observations on different ducts. □, Effect of 2 × 10−4 M dibutyryl cAMP on fluid secretion from isolated ducts. Mean of 7 observations on different ducts.

Data from Argent et al.
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R. M. Case, B. E. Argent. Pancreatic Secretion of Electrolytes and Water. Compr Physiol 2011, Supplement 18: Handbook of Physiology, The Gastrointestinal System, Salivary, Gastric, Pancreatic, and Hepatobiliary Secretion: 383-417. First published in print 1989. doi: 10.1002/cphy.cp060320