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Mechanisms of Fluid Transport Across Renal Tubules

James A. Schafer, W. Brian Reeves, Thomas E. Andreoli

10.1002/cphy.cp080115

Source: Supplement 25: Handbook of Physiology, Renal Physiology

Originally published: 1992

Published online: January 2011

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Theoretical Foundations of Water Transport in Epithelia
2 Volume Absorption in the Proximal Tubule
3 Water Permeability of the Proximal Tubule
3.1 Measurement of Water Permeability
3.2 Importance of the Osmotic Water Permeability
4 Location of the Osmotic Gradient
4.1 Luminal Hypotonicity Produced by Solute Absorption
4.2 Absorbate Hypertonicity
5 “Passive” Driving Forces for Volume Absorption
5.1 NaCl Diffusion in Volume Absorption
5.2 Reflection Coefficient Differences
6 Models of Solute‐Solvent Coupling in Proximal Volume Absorption
7 Routes of Volume Movement in the Proximal Nephron
7.1 Measurement of Cell Membrane Osmotic Water Permeabilities
8 Consequences of Transcellular Volume Flow
9 The ADH‐Sensitive Distal Nephron
9.1 Measurement of Permeability Changes Produced by ADH
9.2 Wafer Permeability of ADH‐Sensitive Nephron Segments in Vivo and in Vitro
10 Site of the Change in Water Permeability with ADH
11 Evaluation of the Pf/PDw Ratio
11.1 Large Pore Hypothesis
11.2 Unstirred Layer Effects
12 The Narrow Channel Hypothesis: Single‐File Diffusion Through Small Aqueous Channels
13 ADH Increases the Number of Narrow Aqueous Channels in Apical Plasma Membranes
14 The Apparent EA for Water Transport in Cortical Collecting Tubules
14.1 The Raw Data
14.2 Correction for Diffusion Constraints in Series with Apical Membranes
14.3 The “True” EA for Water Transport
15 Pseudo‐“Breaks” in EA Measurements
16 Comparison of ADH‐Dependent Apical Membrane Water Channels with Gramicidin A Channels
17 Parallel Paths for Water and Solute Permeation
18 Morphologic Studies
19 Intracellular Mediators of ADH Action
19.1 Modulation of ADH Action—α‐Adrenergic Agents
19.2 Atrial Natriuretic Peptide
20 Prostaglandins
20.1 Calcium
20.2 Protein Kinase C
21 Summary
Figure 1. Figure 1.

Estimate of osmotic water permeability in rabbit proximal convoluted tubule by extrapolation to infinite perfusion rate. Tubule segments were bathed in isosmotic bicarbonate buffer at 25° C to block active transport. They were perfused with a solution that was hypoosmotic to the bathing solution by 20 mOsm/kg H2O. A. Rate of volume absorption (Jv, nl/min, not normalized per tubule length) plotted as function of perfusion rate (V0, nl/min). B. Inverse of volume absorption plotted as function of inverse of perfusion rate. Data points were fitted with a straight line. Y‐intercept is predicted inverse of volume flow that would occur at an infinite perfusion rate when the whole tubule length would be exposed to uniform transepithelial osmolality difference of 20 mOsm/kg H2O. Volume flow rate and osmolality difference were used to compute osmotic water permeability of 2,420 μm/s.

Adapted from Andreoli et al. 10 with permission
Figure 2. Figure 2.

Effect of osmotic water permeability on osmotic gradient required to produce normal rates of volume absorption. Transepithelial osmolality differences that would be required to produce volume reabsorption rates in the normal range observed in the mammalian proximal tubule are calculated using the relationship: Jv = PfA′w ΔC, where Jv is volume flow, Pf is osmotic water permeability coefficient, A′ is area of apparent luminal surface area per millimeter of tubule length, w is partial molal volume of water, and ΔC is required osmolality difference. Relationship between Jv and ΔC plotted for four values of osmotic water permeability that span the range of commonly reported values.
Figure 3. Figure 3.

Demonstration of development of luminal hypotonicity during volume absorption in proximal tubule. Segments of in vivo rat proximal convoluted tubules, isolated between oil blocks, were perfused at two different rates with 154 mM NaCl. Adjacent peritubular capillary network was perfused simultaneously with the same solution and rate of volume absorption and osmolality of perfused and collected samples was measured so that transepithelial osmolality difference could be determined. Both volume absorption rate and luminal hypotonicity increase as perfusion rate is increased, and both are not significantly different from zero when active transport is inhibited by NaCN.

Adapted from the data of Green and Giebisch 108 and reprinted with permission from Kidney International 245
Figure 4. Figure 4.

Method of collection of absorbate droplets from proximal tubules. Isolated segment of rabbit proximal convoluted or straight tubule is perfused (left to right in the figure) while bathed in light mineral oil. Absorbate appears on peritubular surface as torus‐shaped droplets surrounding tubule. Absorbate can be collected and analyzed at regular intervals using volumetric collection pipet.

Reprinted with permission from American Journal of Physiology 16
Figure 5. Figure 5.

Hyperosmolality and composition of absorbate from (A) rabbit proximal convoluted tubule and (B) rabbit proximal straight tubule. Tubule segments are perfused under oil and absorbate forming on peritubular surface is sampled for analysis as shown in Figure 4. The composition of the perfusate (left) is average of measured perfusate and collectate concentrations. Concentrations indicated are in mM for Na+ and glucose. Osmolalities are in mOsm/kg H2O. P values indicate significance of paired difference in average perfusate and absorbate compositions.

Data from Barfuss and Schafer 16,18. Reproduced with permission from Kidney International 245
Figure 6. Figure 6.

Changes in transepithelial voltage with rapid cooling in isolated perfused Ambystoma proximal tubule segments. A. Rapid cooling with symmetrical solutions. B. Rapid cooling with simultaneous 8% increase in NaCl concentration of bathing solution. Phase of slow depolarization at low temperature (A) is attributed to relaxation of paracellular diffusion potential (ΔEpar) by 0.27 mV, resulting from diffusion of NaCl out of interspace, in which NaCl concentration is 3 mM higher than in bathing solution. Repolarization in B is attributed to hyperpolarization by ΔEpar −0.43 mV resulting from equilibration of higher NaCl concentration in bathing solution with that in interspace. Absence of either slow depolarization or repolarization with cooling would indicate a match between interspace and bathing solution NaCl concentration.

Reprinted from the work of Sackin 239 with permission from author and American Journal of Physiology
Figure 7. Figure 7.

Flow dependence of volume absorption in rabbit proximal straight tubules. For upper two sets of data, tubules were perfused with high Cl−, low perfusate resembling late proximal tubule fluid and bathed with the normal Cl− concentration bathing solution. Lowest group of data points was obtained when Cl− and HCO3 concentrations were equal in the perfusate and bathing solutions.

Data from Schafer et al. 255. Figure reprinted from 245 with permission from Kidney International
Figure 8. Figure 8.

Time course of ADH effect on water permeability in mammalian cortical collecting tubule. A. Rabbit cortical collecting tubule perfused at 25° C 113. B. Rat cortical collecting tubule perfused at 37° C 232.
Figure 9. Figure 9.

Rate‐limiting site for water permeation. Segment of cortical collecting tubule was perfused with 125 mOsm Krebs‐Ringer‐phosphate (KRP) buffer and bathed in 290 mOsm Krebs‐Ringer‐bicarbonate (KRB) buffer at 25° C. Time shown below each photomicrograph is that elapsed after rabbit was sacrificed. At 200 min, ADH was added to bath. Interval between small divisions in photomicrograph scale is 2.13 μm.

Adapted from 247
Figure 10. Figure 10.

Rate‐limiting site for urea permeation in cortical collecting tubule. Perfusing solution was 125 mOsm KRP buffer plus 165 mOsm urea throughout experiment. Bath was initially 290 mOsm KRB buffer, but was changed to 125 mOsm KRP plus 165 mOsm urea at 250 min after rabbit was sacrificed. Bath contained 250 μU ml−1 ADH throughout. Interval between small divisions in photomicrograph scale is 2.13 μm.

Adapted from 248
Figure 11. Figure 11.

Series resistance elements for water diffusion across membrane bordered by two unstirred layers. Total resistance of unstirred layer Runs is sum of and . Total resistance to water diffusion across membrane and unstirred layers is given by equation 6.

From 126
Figure 12. Figure 12.

Plot of relationship between Pf and PDw according to equation 14. Values of n used at various temperatures were free solution viscosity values 124.
Figure 13. Figure 13.

Arrhenius plots of relation between In Pf and 1/T in cortical collecting tubules in absence (closed circles, ± SEM) and presence (open circles, ± SEM) of ADH. EA values calculated according to equation 16.

Adapted from 1,124
Figure 14. Figure 14.

Arrhenius plots of relation between In PDw and 1/T in cortical collecting tubules in the absence (closed circles, ± SEM) and the presence (open circles, ± SEM) of ADH. The EA values calculated according to equation 16.

Adapted from 124
Figure 15. Figure 15.

Arrhenius plot of relation between In PDn−butanol and 1/T between 6° and 38° C, in cortical collecting tubules. Open circles are mean experimental values with indicated SEM. EA values calculated according to equation 16.

Adapted from 124
Figure 16. Figure 16.

Schematic illustration of shuttle mechanism for moving water channels into apical cell membrane. In resting state (left), water channels (heavy dots) reside in walls of small tubular vesicles in cytoplasm. Upon stimulation by ADH, these vesicles fuse with apical membrane (middle). Termination of ADH effect involves removal of channel from apical membrane and transport to multivesicular bodies (MVB).
Figure 17. Figure 17.

Flux inhibition in toad urinary bladder. Time course of ADH‐induced hydroosmotic response was measured at various transepithelial osmotic gradients. In presence of large osmotic gradients, ADH effect decays rapidly but is relatively stable in presence of small osmotic gradients 119.
Figure 18. Figure 18.

Schematic illustration of hormone‐receptor‐adenylate cyclase system. Rs and Ri denote hormone receptors for stimulatory and inhibitory hormones, respectively. Gs and Gi are stimulatory and inhibitory guanine‐nucleotide binding regulatory proteins; C is the catalytic subunit of adenylate cyclase 127.


Figure 1.

Estimate of osmotic water permeability in rabbit proximal convoluted tubule by extrapolation to infinite perfusion rate. Tubule segments were bathed in isosmotic bicarbonate buffer at 25° C to block active transport. They were perfused with a solution that was hypoosmotic to the bathing solution by 20 mOsm/kg H2O. A. Rate of volume absorption (Jv, nl/min, not normalized per tubule length) plotted as function of perfusion rate (V0, nl/min). B. Inverse of volume absorption plotted as function of inverse of perfusion rate. Data points were fitted with a straight line. Y‐intercept is predicted inverse of volume flow that would occur at an infinite perfusion rate when the whole tubule length would be exposed to uniform transepithelial osmolality difference of 20 mOsm/kg H2O. Volume flow rate and osmolality difference were used to compute osmotic water permeability of 2,420 μm/s.

Adapted from Andreoli et al. 10 with permission


Figure 2.

Effect of osmotic water permeability on osmotic gradient required to produce normal rates of volume absorption. Transepithelial osmolality differences that would be required to produce volume reabsorption rates in the normal range observed in the mammalian proximal tubule are calculated using the relationship: Jv = PfA′w ΔC, where Jv is volume flow, Pf is osmotic water permeability coefficient, A′ is area of apparent luminal surface area per millimeter of tubule length, w is partial molal volume of water, and ΔC is required osmolality difference. Relationship between Jv and ΔC plotted for four values of osmotic water permeability that span the range of commonly reported values.



Figure 3.

Demonstration of development of luminal hypotonicity during volume absorption in proximal tubule. Segments of in vivo rat proximal convoluted tubules, isolated between oil blocks, were perfused at two different rates with 154 mM NaCl. Adjacent peritubular capillary network was perfused simultaneously with the same solution and rate of volume absorption and osmolality of perfused and collected samples was measured so that transepithelial osmolality difference could be determined. Both volume absorption rate and luminal hypotonicity increase as perfusion rate is increased, and both are not significantly different from zero when active transport is inhibited by NaCN.

Adapted from the data of Green and Giebisch 108 and reprinted with permission from Kidney International 245


Figure 4.

Method of collection of absorbate droplets from proximal tubules. Isolated segment of rabbit proximal convoluted or straight tubule is perfused (left to right in the figure) while bathed in light mineral oil. Absorbate appears on peritubular surface as torus‐shaped droplets surrounding tubule. Absorbate can be collected and analyzed at regular intervals using volumetric collection pipet.

Reprinted with permission from American Journal of Physiology 16


Figure 5.

Hyperosmolality and composition of absorbate from (A) rabbit proximal convoluted tubule and (B) rabbit proximal straight tubule. Tubule segments are perfused under oil and absorbate forming on peritubular surface is sampled for analysis as shown in Figure 4. The composition of the perfusate (left) is average of measured perfusate and collectate concentrations. Concentrations indicated are in mM for Na+ and glucose. Osmolalities are in mOsm/kg H2O. P values indicate significance of paired difference in average perfusate and absorbate compositions.

Data from Barfuss and Schafer 16,18. Reproduced with permission from Kidney International 245


Figure 6.

Changes in transepithelial voltage with rapid cooling in isolated perfused Ambystoma proximal tubule segments. A. Rapid cooling with symmetrical solutions. B. Rapid cooling with simultaneous 8% increase in NaCl concentration of bathing solution. Phase of slow depolarization at low temperature (A) is attributed to relaxation of paracellular diffusion potential (ΔEpar) by 0.27 mV, resulting from diffusion of NaCl out of interspace, in which NaCl concentration is 3 mM higher than in bathing solution. Repolarization in B is attributed to hyperpolarization by ΔEpar −0.43 mV resulting from equilibration of higher NaCl concentration in bathing solution with that in interspace. Absence of either slow depolarization or repolarization with cooling would indicate a match between interspace and bathing solution NaCl concentration.

Reprinted from the work of Sackin 239 with permission from author and American Journal of Physiology


Figure 7.

Flow dependence of volume absorption in rabbit proximal straight tubules. For upper two sets of data, tubules were perfused with high Cl−, low perfusate resembling late proximal tubule fluid and bathed with the normal Cl− concentration bathing solution. Lowest group of data points was obtained when Cl− and HCO3 concentrations were equal in the perfusate and bathing solutions.

Data from Schafer et al. 255. Figure reprinted from 245 with permission from Kidney International


Figure 8.

Time course of ADH effect on water permeability in mammalian cortical collecting tubule. A. Rabbit cortical collecting tubule perfused at 25° C 113. B. Rat cortical collecting tubule perfused at 37° C 232.



Figure 9.

Rate‐limiting site for water permeation. Segment of cortical collecting tubule was perfused with 125 mOsm Krebs‐Ringer‐phosphate (KRP) buffer and bathed in 290 mOsm Krebs‐Ringer‐bicarbonate (KRB) buffer at 25° C. Time shown below each photomicrograph is that elapsed after rabbit was sacrificed. At 200 min, ADH was added to bath. Interval between small divisions in photomicrograph scale is 2.13 μm.

Adapted from 247


Figure 10.

Rate‐limiting site for urea permeation in cortical collecting tubule. Perfusing solution was 125 mOsm KRP buffer plus 165 mOsm urea throughout experiment. Bath was initially 290 mOsm KRB buffer, but was changed to 125 mOsm KRP plus 165 mOsm urea at 250 min after rabbit was sacrificed. Bath contained 250 μU ml−1 ADH throughout. Interval between small divisions in photomicrograph scale is 2.13 μm.

Adapted from 248


Figure 11.

Series resistance elements for water diffusion across membrane bordered by two unstirred layers. Total resistance of unstirred layer Runs is sum of and . Total resistance to water diffusion across membrane and unstirred layers is given by equation 6.

From 126


Figure 12.

Plot of relationship between Pf and PDw according to equation 14. Values of n used at various temperatures were free solution viscosity values 124.



Figure 13.

Arrhenius plots of relation between In Pf and 1/T in cortical collecting tubules in absence (closed circles, ± SEM) and presence (open circles, ± SEM) of ADH. EA values calculated according to equation 16.

Adapted from 1,124


Figure 14.

Arrhenius plots of relation between In PDw and 1/T in cortical collecting tubules in the absence (closed circles, ± SEM) and the presence (open circles, ± SEM) of ADH. The EA values calculated according to equation 16.

Adapted from 124


Figure 15.

Arrhenius plot of relation between In PDn−butanol and 1/T between 6° and 38° C, in cortical collecting tubules. Open circles are mean experimental values with indicated SEM. EA values calculated according to equation 16.

Adapted from 124


Figure 16.

Schematic illustration of shuttle mechanism for moving water channels into apical cell membrane. In resting state (left), water channels (heavy dots) reside in walls of small tubular vesicles in cytoplasm. Upon stimulation by ADH, these vesicles fuse with apical membrane (middle). Termination of ADH effect involves removal of channel from apical membrane and transport to multivesicular bodies (MVB).



Figure 17.

Flux inhibition in toad urinary bladder. Time course of ADH‐induced hydroosmotic response was measured at various transepithelial osmotic gradients. In presence of large osmotic gradients, ADH effect decays rapidly but is relatively stable in presence of small osmotic gradients 119.



Figure 18.

Schematic illustration of hormone‐receptor‐adenylate cyclase system. Rs and Ri denote hormone receptors for stimulatory and inhibitory hormones, respectively. Gs and Gi are stimulatory and inhibitory guanine‐nucleotide binding regulatory proteins; C is the catalytic subunit of adenylate cyclase 127.

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Article Title: Mechanisms of Fluid Transport Across Renal Tubules
 

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James A. Schafer, W. Brian Reeves, Thomas E. Andreoli. Mechanisms of Fluid Transport Across Renal Tubules. Compr Physiol 2011, Supplement 25: Handbook of Physiology, Renal Physiology: 659-713. First published in print 1992. doi: 10.1002/cphy.cp080115