Fluid and Electrolyte Secretion by Salivary Glands

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Fluid and Electrolyte Secretion by Salivary Glands

D. I. Cook, J. A. Young

10.1002/cphy.cp060301

Source: Supplement 18: Handbook of Physiology, The Gastrointestinal System, Salivary, Gastric, Pancreatic, and Hepatobiliary Secretion

Originally published: 1989

Published online: January 2011

Full Article on Wiley Online Library

Abstract
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References

Abstract

The sections in this article are:

1 Anatomical Nomenclature

2 Two‐Stage Hypothesis

3 Mechanism Of Formation of Primary Fluid

3.1 Secretion Models

3.2 Elements of Secretory Mechanism

3.3 Overview of Secretion Control

4 Ductal Electrolyte Transport

4.1 Elements of Absorptive Mechanism

4.2 Plasticity of Ductal Transport Properties

4.3 Water Permeability

4.4 Control of Ductal Transport

4.5 Overview of Ductal Electrolyte Transport

5 Appendix

5.1 null

	Figure 1. Model showing secretory system dependent on active Na⁺ transport pump in luminal plasma membrane. In this example, Na⁺ is actively transported from cytosol across luminal membrane, but a system based on a luminal membrane Cl^‐ pump, or even a NaCl pump, is also feasible. Cl^‐ is shown moving passively across tight junctions, but it would be equally possible for the passively transported ion to take a transcellular route. Basolateral Na⁺‐K⁺‐ATPase, which exchanges 3Na⁺ for 2K⁺, can be seen to be working against luminal membrane pump.
	Figure 2. Model showing secretory system dependent on active transport of K⁺ by basolateral Na⁺‐K⁺‐ATPase. Potassium is concentrated in cytosol by Na⁺‐K⁺‐ATPase and enters saliva passively across luminal membrane. In this example, K⁺ is shown entering saliva via a K⁺‐Cl^‐ symport, but it might as readily take a conductive pathway, and Cl^‐ could then take either a transcellular or a para‐cellular route. Number 2 on the K⁺‐Cl^‐ symport indicates that 2 molecules of KCl are secreted for each cycle of Na⁺‐K⁺‐ATPase.
	Figure 3. Model showing secretory system dependent on active transport of Cl^‐ by basolateral Na⁺‐K⁺‐2Cl^‐ symport. Cl^‐ is concentrated in cytosol and enters saliva passively via a conductive pathway in luminal membrane. Na⁺ enters saliva passively across tight junctions. Circuit is completed by current flow across basolateral membrane, carried by K⁺ through K⁺ channels and via electrogenic Na⁺‐K⁺‐ATPase. Number 3 on symport indicates that 3 cycles of symport deliver 6Cl^‐ into cytosol for each cycle of Na⁺‐K⁺‐ATPase.
	Figure 4. Model showing secretory system dependent on active transport of HCO^‐₃ by basolateral Na⁺‐H⁺ antiport. Protons are expelled from cytosol by antiport leading to intracellular accumulation of OH^‐, which reacts with CO₂ to form HCO^‐₃. HCO^‐₃ enters saliva passively via a conductive pathway in luminal membrane and Na⁺ enters passively across tight junctions. Circuit is completed by current flow across basolateral membrane, carried by K⁺ through K⁺ channels and via electrogenic Na⁺‐K⁺‐ATPase. Number 3 on antiport indicates that 3 cycles of antiport generate 3HCO^‐₃ in cytosol for each cycle of Na⁺‐K⁺‐ATPase.
	Figure 5. Model showing secretory system dependent on active transport of Cl^‐ by paired basolateral Na⁺‐H⁺ and Cl^‐‐HCO^‐₃ antiports. HCO^‐₃ is first concentrated in cytosol by Na⁺‐H⁺ antiport, and Cl^‐‐HCO^‐₃ antiport utilizes the HCO^‐₃ gradient energy to concentrate Cl^‐ in cytosol. Cl^‐ enters saliva passively via a conductive pathway in luminal membrane and Na⁺ enters passively across tight junctions. Circuit is completed by current flow across basolateral membrane, carried by K⁺ ions through K⁺ channels and via electrogenic Na⁺‐K⁺‐ATPase. Number 3 on each antiport indicates that 3 cycles of each antiport deliver 3 Cl^‐ ions into cytosol for each cycle of Na⁺‐K⁺‐ATPase.
	Figure 6. Current‐voltage relation for channel from luminal membrane of cultured cell from continuous mouse mandibular cell line. Recordings are from a cell‐attached patch in which pipette contained K⁺ 140 mM) and Cl^‐ 25 mM). Reversal potential is close to −10 mV and slope conductance is 25 pS. Right, representative channel events at each pipette potential employed. Data from Cook et al. 26
	Figure 7. Model for electrolyte transport by excretory duct epithelium of rat mandibular gland. Na⁺ enters cytosol from lumen either via conductive pathway or a nonconductive Na⁺‐H⁺ antiport and is expelled to interstitium by Na⁺ pump. K⁺ is transported from interstitium to cytosol by Na⁺ pump and enters saliva via a luminal K⁺‐H⁺ antiport. Cl^‐ is absorbed across conductive pathways in both plasma membranes, as well as via a Cl^‐‐HCO^‐₃ antiport in luminal membrane. HCO^‐₃ is concentrated in cytosol by basolateral and luminal Na⁺‐H⁺ antiports and can be secreted into lumen by luminal Cl^‐‐HCO^‐₃ and K⁺‐H⁺ antiports. Adapted from Knauf et al. 74

Figure 1.

Model showing secretory system dependent on active Na⁺ transport pump in luminal plasma membrane. In this example, Na⁺ is actively transported from cytosol across luminal membrane, but a system based on a luminal membrane Cl^‐ pump, or even a NaCl pump, is also feasible. Cl^‐ is shown moving passively across tight junctions, but it would be equally possible for the passively transported ion to take a transcellular route. Basolateral Na⁺‐K⁺‐ATPase, which exchanges 3Na⁺ for 2K⁺, can be seen to be working against luminal membrane pump.

Figure 2.

Model showing secretory system dependent on active transport of K⁺ by basolateral Na⁺‐K⁺‐ATPase. Potassium is concentrated in cytosol by Na⁺‐K⁺‐ATPase and enters saliva passively across luminal membrane. In this example, K⁺ is shown entering saliva via a K⁺‐Cl^‐ symport, but it might as readily take a conductive pathway, and Cl^‐ could then take either a transcellular or a para‐cellular route. Number 2 on the K⁺‐Cl^‐ symport indicates that 2 molecules of KCl are secreted for each cycle of Na⁺‐K⁺‐ATPase.

Figure 3.

Model showing secretory system dependent on active transport of Cl^‐ by basolateral Na⁺‐K⁺‐2Cl^‐ symport. Cl^‐ is concentrated in cytosol and enters saliva passively via a conductive pathway in luminal membrane. Na⁺ enters saliva passively across tight junctions. Circuit is completed by current flow across basolateral membrane, carried by K⁺ through K⁺ channels and via electrogenic Na⁺‐K⁺‐ATPase. Number 3 on symport indicates that 3 cycles of symport deliver 6Cl^‐ into cytosol for each cycle of Na⁺‐K⁺‐ATPase.

Figure 4.

Model showing secretory system dependent on active transport of HCO^‐₃ by basolateral Na⁺‐H⁺ antiport. Protons are expelled from cytosol by antiport leading to intracellular accumulation of OH^‐, which reacts with CO₂ to form HCO^‐₃. HCO^‐₃ enters saliva passively via a conductive pathway in luminal membrane and Na⁺ enters passively across tight junctions. Circuit is completed by current flow across basolateral membrane, carried by K⁺ through K⁺ channels and via electrogenic Na⁺‐K⁺‐ATPase. Number 3 on antiport indicates that 3 cycles of antiport generate 3HCO^‐₃ in cytosol for each cycle of Na⁺‐K⁺‐ATPase.

Figure 5.

Model showing secretory system dependent on active transport of Cl^‐ by paired basolateral Na⁺‐H⁺ and Cl^‐‐HCO^‐₃ antiports. HCO^‐₃ is first concentrated in cytosol by Na⁺‐H⁺ antiport, and Cl^‐‐HCO^‐₃ antiport utilizes the HCO^‐₃ gradient energy to concentrate Cl^‐ in cytosol. Cl^‐ enters saliva passively via a conductive pathway in luminal membrane and Na⁺ enters passively across tight junctions. Circuit is completed by current flow across basolateral membrane, carried by K⁺ ions through K⁺ channels and via electrogenic Na⁺‐K⁺‐ATPase. Number 3 on each antiport indicates that 3 cycles of each antiport deliver 3 Cl^‐ ions into cytosol for each cycle of Na⁺‐K⁺‐ATPase.

Figure 6.

Current‐voltage relation for channel from luminal membrane of cultured cell from continuous mouse mandibular cell line. Recordings are from a cell‐attached patch in which pipette contained K⁺ 140 mM) and Cl^‐ 25 mM). Reversal potential is close to −10 mV and slope conductance is 25 pS. Right, representative channel events at each pipette potential employed.

Data from Cook et al. 26

Figure 7.

Model for electrolyte transport by excretory duct epithelium of rat mandibular gland. Na⁺ enters cytosol from lumen either via conductive pathway or a nonconductive Na⁺‐H⁺ antiport and is expelled to interstitium by Na⁺ pump. K⁺ is transported from interstitium to cytosol by Na⁺ pump and enters saliva via a luminal K⁺‐H⁺ antiport. Cl^‐ is absorbed across conductive pathways in both plasma membranes, as well as via a Cl^‐‐HCO^‐₃ antiport in luminal membrane. HCO^‐₃ is concentrated in cytosol by basolateral and luminal Na⁺‐H⁺ antiports and can be secreted into lumen by luminal Cl^‐‐HCO^‐₃ and K⁺‐H⁺ antiports.

Adapted from Knauf et al. 74

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D. I. Cook, J. A. Young. Fluid and Electrolyte Secretion by Salivary Glands. Compr Physiol 2011, Supplement 18: Handbook of Physiology, The Gastrointestinal System, Salivary, Gastric, Pancreatic, and Hepatobiliary Secretion: 1-23. First published in print 1989. doi: 10.1002/cphy.cp060301

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