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Renal Ion Channels

Lawrence G. Palmer

10.1002/cphy.cp080116

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 Historical Overview
2 What is an Ion Channel?
3 Techniques for Studying Ion Channels
3.1 Fluctuation (Noise) Analysis
3.2 Reconstitution in Lipid Bilayers
3.3 Patch‐Clamp
3.4 Application of the Patch‐Clamp to Renal Tubules
3.5 Parameters Derived from Patch‐Clamp Measurements
4 Ion Channels Along the Nephron
4.1 Proximal Tubule
4.2 Thick Ascending Limb/Diluting Segment
4.3 Collecting Tubule/Urinary Bladder
4.4 Channels in Cell Volume Homeostasis
5 Summary
Figure 1. Figure 1.

Schematic of patch‐clamp technique. First step is formation of cell‐attached patch, in which single‐channel currents can be studied in situ. Withdrawal of pipette cell from results in inside‐out patch, which is useful for establishing ion selectivity and for testing effects of changing composition of medium bathing cytoplasmic side of patch. Applying suction or voltage to pipette ruptures the patch, forming a low‐resistance connection between inside of pipette and cytoplasm. In this whole‐cell configuration, currents across entire cell membrane can be measured under voltage‐clamp conditions, and cell can be dialyzed with pipette solution. Withdrawal of pipette from cell leads to outside‐out configuration, similar to inside‐out patch except that extracellular side of membrane faces out.
Figure 2. Figure 2.

Photomicrograph of split tubule. Rat cortical collecting tubule was isolated and partially split by hand using sharp forceps and dissecting needle. Lower portion of tubule has been opened, and luminal membrane of epithelial cells is accessible to patch‐clamp pipette.

Courtesy of Dr. Gustavo Frindt
Figure 3. Figure 3.

Model of ion transport in proximal tubule. Na+ enters cells by cotransport with organic substrates (sugars, amino acids and organic anions), by exchange with H+, and, at least in late proximal tubule, through Na+‐selective channels. K+‐selective and nonselective cation channels are also found in apical membrane, but are not illustrated. Na+ exits cells in exchange for K+, which in turn is recycled across basolateral membrane through K+ channels. For simplicity, postulated stoichiometries not shown.
Figure 4. Figure 4.

Apical membrane Na+ channels in perfused rabbit proximal straight tubule. Current recordings are from a cell‐attached patch with NaCl solution in pipette. Current levels at which channels are closed are marked with arrows. Downward current deflections represent channel openings that give rise to inward currents (Movement of Na+ from pipette to cell). Voltages correspond to those of bath relative to pipette. Slope conductance, obtained from the i–V curve illustrated below, is 12 pS.

From Gögelein and Greger 23
Figure 5. Figure 5.

Basolateral membrane K+ channels in isolated rabbit proximal tubule treated with collagenase to remove basement membrane. Current recordings (A) are from cell‐attached patch with 200 mM KCl in pipette and 145 mM KCl in bath. Current levels at which channels are closed are at bottom of small arrows. Upward current deflections at positive voltages represent channel openings with positive charge (K+) moving from cell to pipette. Downward current deflections, observed at negative voltages, represent K+ moving from pipette to cell. Voltages correspond to those of bath relative to those of pipette. B. Slope conductance, obtained from i–V curve, is 54 pS for inward current. Reversal potential under these conditions is near zero. However, in presence of bathing solution with normal K+ concentrations, reversal potential shifts to positive voltages because of presence of normal cell resting potential.

From Parent et al. 91
Figure 6. Figure 6.

Model of ion transport by thick ascending limb. Na+, Cl−, and K+ enter cell together through triple cotransporter. K+ recycles across apical membrane through channels, while Cl− exits basolateral membrane through anion‐selective channels. Na+ leaves cell across basolateral membrane in exchange for K+. There may also be K+‐selective channels on basolateral membrane, although these have not been identified at single‐channel level.
Figure 7. Figure 7.

K+ channels from apical membrane of rabbit thick ascending limb. Current recordings (A) are from cell‐attached patch with K+ gluconate solution in pipette and in bath. Current levels at which channels are closed are marked with lines. Downward current deflections represent channel openings that give rise to inward currents (K+ movement from pipette to cell). Upward current deflections, observed at +110 mV, represent movement of cations from cell to pipette. Voltages correspond to those of bath relative to pipette. In all traces, channel is usually open, as shown by (B) plot of open probability (Po) versus voltage. Maximal slope conductance, obtained from i–V curve, is 22 pS. Distribution of channel open times follows two exponentials with time constants of 33 and 1.5 ms. Closed times are distributed with one exponential, with mean closed time of 1.0 ms.

From Wang et al. 119
Figure 8. Figure 8.

Cl− channels from basolateral membrane of the isolated rabbit thick ascending limb. Current recordings (A) are from cell‐attached patch with NaCl solution in pipette and in bath. Current levels at which channels are closed are marked with dashed lines. Downward current deflections at negative voltages represent channel openings that give rise to inward currents (Cl− movement from cell to pipette). Upward current deflections, observed at positive voltages, represent movement of outward currents or movement of Cl− from pipette to bath. Voltages correspond to those of bath relative to pipette. There are at least two channels of similar conductance in patch. (B) Slope conductance, obtained from the i–V curve, is 42 pS.

From Paulais and Teulon 93
Figure 9. Figure 9.

Model of ion transport by principal cells of cortical collecting tubule. Na+ enters cells through amiloride‐sensitive channels in apical membrane and exits across basolateral membrane in exchange for K+. The K+ is partly recycled across basolateral membrane through K+ channels and partly secreted across apical membrane, also through channels. Basolateral membrane also contains Cl channels, although these cells do not contribute to transepithelial Cl− transport.
Figure 10. Figure 10.

Na+ channels from apical membrane of rat cortical collecting tubule. Current recordings (A) are from cell‐attached patch with NaCl solution in pipette and in bath. In lower trace, pipette solution also contains 0.5 μM amiloride. Current levels at which channels are closed are marked with lines. Downward current deflections represent channel openings that give rise to inward currents (Na+ movement from pipette to cell). Voltages correspond to those of pipette relative to bath. Maximal slope conductance (B), obtained from the i–V curve, is 5 pS. This value is similar for control solutions (open circles) and those with amiloride (filled circles), as well as for excised patches with NaCl solution on both sides of patch (solid squares.) When Na+ gradient is applied across patch by replacing two‐thirds of Na+ in bath with K+, reversal potential shifts with ENa, the Nernst potential for Na+. Open and closed times of channels (C) are distributed with one exponential under control conditions, corresponding to mean open and closed times of 1.0 and 0.8 s, respectively.

From Palmer and Frindt 85
Figure 11. Figure 11.

Cl− channels from basolateral membrane of isolated rabbit cortical collecting tubules. Current recordings are from cell‐attached patch KCl solution in pipette and NaCl solution in bath. Current levels at which channels are closed are marked as Co. Downward current deflections at negative voltages represent channel openings that give rise to inward currents (Cl− movement from pipette to cell). Upward current deflections, observed at positive voltages, represent movement of inward currents or movements of Cl− from cell to pipette. Channels appeared to be “double‐barrelled,” having two conductance states one of which was twice the other. Thus three current levels at each voltage correspond to closed state, half‐open (or one‐barrel open) state, and fully open (two‐barrel) state. Slope conductance, obtained from i–V curve, is 46 pS for the fully open state.

From Sansom et al. 102
Figure 12. Figure 12.

Model of volume regulatory decrease in generic epithelial cell. Cell swelling is thought to increase activity of stretch‐sensitive K+ channels, in basolateral membrane and Ca2+ channels in apical membrane. Influx of Ca2+ through stretch‐activated channels will open Ca2+‐activated K+ channels in apical membrane. K+ will leave cell through one or both pathways. In addition, basolateral Cl channels, activated by stretch, Ca2+, or unknown third mechanism will mediate net Cl− efflux.
Figure 13. Figure 13.

Stretch‐ and volume‐activated K+ channels in basolateral membrane of Necturus proximal tubule. Channel activity recorded from cell‐attached patch with high K+ in pipette. Upward current deflections represent channel openings with inward current carried by K+ from pipette to cell. Application of suction (6 cm H2O) to pipette or superfusion of tubule with hypotonic solution (100 mOsm) activated channels. Open time histograms followed single exponential with mean open time 0.7 ms. Closed times distributed with two exponentials having time constants of 1.7 and 43 ms under control conditions. Effect of suction and swelling was to decrease longer closed‐time constant by factor of ∼3.

From Sackin 99
Figure 14. Figure 14.

Volume‐activated K+ channels in apical membrane of thick ascending limb cells in culture. Channel activity was recorded from cell‐attached patch (A‐C) with high K+ in pipette. Downward current deflections correspond to channel openings with inward current carried by K+ from pipette to cell. Cell was exposed to normal osmolality (293 mOsM, A), lowered osmolality (220 mOsM, B), and then back to normal solution (C). D‐F: activation of channel by 10 μM Ca2+ in excised, inside‐out patch. Effect of osmolality required presence of Ca2+ in external bathing medium as shown by plot of fractional open probability versus time.

From Taniguchi and Guggino 90


Figure 1.

Schematic of patch‐clamp technique. First step is formation of cell‐attached patch, in which single‐channel currents can be studied in situ. Withdrawal of pipette cell from results in inside‐out patch, which is useful for establishing ion selectivity and for testing effects of changing composition of medium bathing cytoplasmic side of patch. Applying suction or voltage to pipette ruptures the patch, forming a low‐resistance connection between inside of pipette and cytoplasm. In this whole‐cell configuration, currents across entire cell membrane can be measured under voltage‐clamp conditions, and cell can be dialyzed with pipette solution. Withdrawal of pipette from cell leads to outside‐out configuration, similar to inside‐out patch except that extracellular side of membrane faces out.



Figure 2.

Photomicrograph of split tubule. Rat cortical collecting tubule was isolated and partially split by hand using sharp forceps and dissecting needle. Lower portion of tubule has been opened, and luminal membrane of epithelial cells is accessible to patch‐clamp pipette.

Courtesy of Dr. Gustavo Frindt


Figure 3.

Model of ion transport in proximal tubule. Na+ enters cells by cotransport with organic substrates (sugars, amino acids and organic anions), by exchange with H+, and, at least in late proximal tubule, through Na+‐selective channels. K+‐selective and nonselective cation channels are also found in apical membrane, but are not illustrated. Na+ exits cells in exchange for K+, which in turn is recycled across basolateral membrane through K+ channels. For simplicity, postulated stoichiometries not shown.



Figure 4.

Apical membrane Na+ channels in perfused rabbit proximal straight tubule. Current recordings are from a cell‐attached patch with NaCl solution in pipette. Current levels at which channels are closed are marked with arrows. Downward current deflections represent channel openings that give rise to inward currents (Movement of Na+ from pipette to cell). Voltages correspond to those of bath relative to pipette. Slope conductance, obtained from the i–V curve illustrated below, is 12 pS.

From Gögelein and Greger 23


Figure 5.

Basolateral membrane K+ channels in isolated rabbit proximal tubule treated with collagenase to remove basement membrane. Current recordings (A) are from cell‐attached patch with 200 mM KCl in pipette and 145 mM KCl in bath. Current levels at which channels are closed are at bottom of small arrows. Upward current deflections at positive voltages represent channel openings with positive charge (K+) moving from cell to pipette. Downward current deflections, observed at negative voltages, represent K+ moving from pipette to cell. Voltages correspond to those of bath relative to those of pipette. B. Slope conductance, obtained from i–V curve, is 54 pS for inward current. Reversal potential under these conditions is near zero. However, in presence of bathing solution with normal K+ concentrations, reversal potential shifts to positive voltages because of presence of normal cell resting potential.

From Parent et al. 91


Figure 6.

Model of ion transport by thick ascending limb. Na+, Cl−, and K+ enter cell together through triple cotransporter. K+ recycles across apical membrane through channels, while Cl− exits basolateral membrane through anion‐selective channels. Na+ leaves cell across basolateral membrane in exchange for K+. There may also be K+‐selective channels on basolateral membrane, although these have not been identified at single‐channel level.



Figure 7.

K+ channels from apical membrane of rabbit thick ascending limb. Current recordings (A) are from cell‐attached patch with K+ gluconate solution in pipette and in bath. Current levels at which channels are closed are marked with lines. Downward current deflections represent channel openings that give rise to inward currents (K+ movement from pipette to cell). Upward current deflections, observed at +110 mV, represent movement of cations from cell to pipette. Voltages correspond to those of bath relative to pipette. In all traces, channel is usually open, as shown by (B) plot of open probability (Po) versus voltage. Maximal slope conductance, obtained from i–V curve, is 22 pS. Distribution of channel open times follows two exponentials with time constants of 33 and 1.5 ms. Closed times are distributed with one exponential, with mean closed time of 1.0 ms.

From Wang et al. 119


Figure 8.

Cl− channels from basolateral membrane of the isolated rabbit thick ascending limb. Current recordings (A) are from cell‐attached patch with NaCl solution in pipette and in bath. Current levels at which channels are closed are marked with dashed lines. Downward current deflections at negative voltages represent channel openings that give rise to inward currents (Cl− movement from cell to pipette). Upward current deflections, observed at positive voltages, represent movement of outward currents or movement of Cl− from pipette to bath. Voltages correspond to those of bath relative to pipette. There are at least two channels of similar conductance in patch. (B) Slope conductance, obtained from the i–V curve, is 42 pS.

From Paulais and Teulon 93


Figure 9.

Model of ion transport by principal cells of cortical collecting tubule. Na+ enters cells through amiloride‐sensitive channels in apical membrane and exits across basolateral membrane in exchange for K+. The K+ is partly recycled across basolateral membrane through K+ channels and partly secreted across apical membrane, also through channels. Basolateral membrane also contains Cl channels, although these cells do not contribute to transepithelial Cl− transport.



Figure 10.

Na+ channels from apical membrane of rat cortical collecting tubule. Current recordings (A) are from cell‐attached patch with NaCl solution in pipette and in bath. In lower trace, pipette solution also contains 0.5 μM amiloride. Current levels at which channels are closed are marked with lines. Downward current deflections represent channel openings that give rise to inward currents (Na+ movement from pipette to cell). Voltages correspond to those of pipette relative to bath. Maximal slope conductance (B), obtained from the i–V curve, is 5 pS. This value is similar for control solutions (open circles) and those with amiloride (filled circles), as well as for excised patches with NaCl solution on both sides of patch (solid squares.) When Na+ gradient is applied across patch by replacing two‐thirds of Na+ in bath with K+, reversal potential shifts with ENa, the Nernst potential for Na+. Open and closed times of channels (C) are distributed with one exponential under control conditions, corresponding to mean open and closed times of 1.0 and 0.8 s, respectively.

From Palmer and Frindt 85


Figure 11.

Cl− channels from basolateral membrane of isolated rabbit cortical collecting tubules. Current recordings are from cell‐attached patch KCl solution in pipette and NaCl solution in bath. Current levels at which channels are closed are marked as Co. Downward current deflections at negative voltages represent channel openings that give rise to inward currents (Cl− movement from pipette to cell). Upward current deflections, observed at positive voltages, represent movement of inward currents or movements of Cl− from cell to pipette. Channels appeared to be “double‐barrelled,” having two conductance states one of which was twice the other. Thus three current levels at each voltage correspond to closed state, half‐open (or one‐barrel open) state, and fully open (two‐barrel) state. Slope conductance, obtained from i–V curve, is 46 pS for the fully open state.

From Sansom et al. 102


Figure 12.

Model of volume regulatory decrease in generic epithelial cell. Cell swelling is thought to increase activity of stretch‐sensitive K+ channels, in basolateral membrane and Ca2+ channels in apical membrane. Influx of Ca2+ through stretch‐activated channels will open Ca2+‐activated K+ channels in apical membrane. K+ will leave cell through one or both pathways. In addition, basolateral Cl channels, activated by stretch, Ca2+, or unknown third mechanism will mediate net Cl− efflux.



Figure 13.

Stretch‐ and volume‐activated K+ channels in basolateral membrane of Necturus proximal tubule. Channel activity recorded from cell‐attached patch with high K+ in pipette. Upward current deflections represent channel openings with inward current carried by K+ from pipette to cell. Application of suction (6 cm H2O) to pipette or superfusion of tubule with hypotonic solution (100 mOsm) activated channels. Open time histograms followed single exponential with mean open time 0.7 ms. Closed times distributed with two exponentials having time constants of 1.7 and 43 ms under control conditions. Effect of suction and swelling was to decrease longer closed‐time constant by factor of ∼3.

From Sackin 99


Figure 14.

Volume‐activated K+ channels in apical membrane of thick ascending limb cells in culture. Channel activity was recorded from cell‐attached patch (A‐C) with high K+ in pipette. Downward current deflections correspond to channel openings with inward current carried by K+ from pipette to cell. Cell was exposed to normal osmolality (293 mOsM, A), lowered osmolality (220 mOsM, B), and then back to normal solution (C). D‐F: activation of channel by 10 μM Ca2+ in excised, inside‐out patch. Effect of osmolality required presence of Ca2+ in external bathing medium as shown by plot of fractional open probability versus time.

From Taniguchi and Guggino 90
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Article Title: Renal Ion Channels
 

How to Cite

Lawrence G. Palmer. Renal Ion Channels. Compr Physiol 2011, Supplement 25: Handbook of Physiology, Renal Physiology: 715-738. First published in print 1992. doi: 10.1002/cphy.cp080116