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Physiol. Rev. 80: 211-276, 2000;
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Physiological Reviews, Vol. 80, No. 1, January 2000, pp. 211-276
Copyright ©2000 by the American Physiological Society

Sodium-Potassium-Chloride Cotransport

John M. Russell

Department of Biology, Biological Research Laboratories, Syracuse, New York

Russell, John M. Sodium-Potassium-Chloride Cotransport. Physiol. Rev. 80: 211-276, 2000.Obligatory, coupled cotransport of Na+, K+, and Cl- by cell membranes has been reported in nearly every animal cell type. This review examines the current status of our knowledge about this ion transport mechanism. Two isoforms of the Na+-K+-Cl- cotransporter (NKCC) protein (~120-130 kDa, unglycosylated) are currently known. One isoform (NKCC2) has at least three alternatively spliced variants and is found exclusively in the kidney. The other (NKCC1) is found in nearly all cell types. The NKCC maintains intracellular Cl- concentration ([Cl-]i) at levels above the predicted electrochemical equilibrium. The high [Cl-]i is used by epithelial tissues to promote net salt transport and by neural cells to set synaptic potentials; its function in other cells is unknown. There is substantial evidence in some cells that the NKCC functions to offset osmotically induced cell shrinkage by mediating the net influx of osmotically active ions. Whether it serves to maintain cell volume under euvolemic conditons is less clear. The NKCC may play an important role in the cell cycle. Evidence that each cotransport cycle of the NKCC is electrically silent is discussed along with evidence for the electrically neutral stoichiometries of 1 Na+:1 K+:2 Cl- (for most cells) and 2 Na+:1 K+:3 Cl- (in squid axon). Evidence that the absolute dependence on ATP of the NKCC is the result of regulatory phosphorylation/dephosphorylation mechanisms is decribed. Interestingly, the presumed protein kinase(s) responsible has not been identified. An unusual form of NKCC regulation is by [Cl-]i. [Cl-]i in the physiological range and above strongly inhibits the NKCC. This effect may be mediated by a decrease of protein phosphorylation. Although the NKCC has been studied for ~20 years, we are only beginning to frame the broad outlines of the structure, function, and regulation of this ubiquitous ion transport mechanism.




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L. N. Nejsum, J. Praetorius, and S. Nielsen
NKCC1 and NHE1 are abundantly expressed in the basolateral plasma membrane of secretory coil cells in rat, mouse, and human sweat glands
Am J Physiol Cell Physiol, August 1, 2005; 289(2): C333 - C340.
[Abstract] [Full Text] [PDF]

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 J. Physiol.Home page
D. Street, J.-J. Nielsen, J. Bangsbo, and C. Juel
Metabolic alkalosis reduces exercise-induced acidosis and potassium accumulation in human skeletal muscle interstitium
J. Physiol., July 15, 2005; 566(2): 481 - 489.
[Abstract] [Full Text] [PDF]

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 J. Biol. Chem.Home page
C. M. Liedtke, X. Wang, and N. D. Smallwood
Role for Protein Phosphatase 2A in the Regulation of Calu-3 Epithelial Na+-K+-2Cl-, Type 1 Co-transport Function
J. Biol. Chem., July 8, 2005; 280(27): 25491 - 25498.
[Abstract] [Full Text] [PDF]

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 Am. J. Physiol. Cell Physiol.Home page
C. D. Ciano-Oliveira, M. Lodyga, L. Fan, K. Szaszi, H. Hosoya, O. D. Rotstein, and A. Kapus
Is myosin light-chain phosphorylation a regulatory signal for the osmotic activation of the Na+-K+-2Cl- cotransporter?
Am J Physiol Cell Physiol, July 1, 2005; 289(1): C68 - C81.
[Abstract] [Full Text] [PDF]

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 J. Neurosci.Home page
C. Marchetti, J. Tabak, N. Chub, M. J. O'Donovan, and J. Rinzel
Modeling Spontaneous Activity in the Developing Spinal Cord Using Activity-Dependent Variations of Intracellular Chloride
J. Neurosci., April 6, 2005; 25(14): 3601 - 3612.
[Abstract] [Full Text] [PDF]

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 J. Neurophysiol.Home page
X. Jin, J. R. Huguenard, and D. A. Prince
Impaired Cl- Extrusion in Layer V Pyramidal Neurons of Chronically Injured Epileptogenic Neocortex
J Neurophysiol, April 1, 2005; 93(4): 2117 - 2126.
[Abstract] [Full Text] [PDF]

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 Physiol. Rev.Home page
G. Gamba
Molecular Physiology and Pathophysiology of Electroneutral Cation-Chloride Cotransporters
Physiol Rev, April 1, 2005; 85(2): 423 - 493.
[Abstract] [Full Text] [PDF]

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 QJMHome page
J.H.M. Groeneveld, Y.W.J. Sijpkens, S.-H. Lin, M.R. Davids, and M.L. Halperin
An approach to the patient with severe hypokalaemia: the potassium quiz
QJM, April 1, 2005; 98(4): 305 - 316.
[Abstract] [Full Text] [PDF]

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 J. Physiol.Home page
J. A Fraser, C. E. J Rang, J. A Usher-Smith, and C. L.-H Huang
Slow volume transients in amphibian skeletal muscle fibres studied in hypotonic solutions
J. Physiol., April 1, 2005; 564(1): 51 - 63.
[Abstract] [Full Text] [PDF]

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 Am. J. Physiol. Cell Physiol.Home page
N. D. Smallwood, B. S. Hausman, X. Wang, and C. M. Liedtke
Involvement of NH2 terminus of PKC-{delta} in binding to F-actin during activation of Calu-3 airway epithelial NKCC1
Am J Physiol Cell Physiol, April 1, 2005; 288(4): C906 - C912.
[Abstract] [Full Text] [PDF]

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 J. Physiol.Home page
R. Liu and A. E. G Persson
Simultaneous changes of cell volume and cytosolic calcium concentration in macula densa cells caused by alterations of luminal NaCl concentration
J. Physiol., March 15, 2005; 563(3): 895 - 901.
[Abstract] [Full Text] [PDF]

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 HypertensionHome page
A. Delgado-Almeida, A. Aviv, N. K. Hollenberg, and A. Weder
Reinterpreting Sodium-Potassium Data in Salt-Sensitivity Hypertension: A Prospective Debate
Hypertension, February 1, 2005; 45(2): e4 - e5.
[Full Text] [PDF]

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 JGPHome page
J. Denton, K. Nehrke, X. Yin, R. Morrison, and K. Strange
GCK-3, a Newly Identified Ste20 Kinase, Binds To and Regulates the Activity of a Cell Cycle-dependent ClC Anion Channel
J. Gen. Physiol., January 31, 2005; 125(2): 113 - 125.
[Abstract] [Full Text] [PDF]

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 Am. J. Physiol. Heart Circ. Physiol.Home page
B. G. Zani and H. G. Bohlen
Sodium channels are required during in vivo sodium chloride hyperosmolarity to stimulate increase in intestinal endothelial nitric oxide production
Am J Physiol Heart Circ Physiol, January 1, 2005; 288(1): H89 - H95.
[Abstract] [Full Text] [PDF]

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 J. Neurosci.Home page
C. Sheldon, A. Diarra, Y. M. Cheng, and J. Church
Sodium Influx Pathways during and after Anoxia in Rat Hippocampal Neurons
J. Neurosci., December 8, 2004; 24(49): 11057 - 11069.
[Abstract] [Full Text] [PDF]

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 J. Pharmacol. Exp. Ther.Home page
P. A. Sonalker, S. P. Tofovic, and E. K. Jackson
Increased Expression of the Sodium Transporter BSC-1 in Spontaneously Hypertensive Rats
J. Pharmacol. Exp. Ther., December 1, 2004; 311(3): 1052 - 1061.
[Abstract] [Full Text] [PDF]

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 J. Physiol.Home page
M. G. van Emst, S. Klarenbeek, A. Schot, J. J. Plomp, A. Doornenbal, and M. E. Everts
Reducing chloride conductance prevents hyperkalaemia-induced loss of twitch force in rat slow-twitch muscle
J. Physiol., November 15, 2004; 561(1): 169 - 181.
[Abstract] [Full Text] [PDF]

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 J. Biol. Chem.Home page
C. F. Simard, N. D. Daigle, M. J. Bergeron, G. M. Brunet, L. Caron, M. Noel, V. Montminy, and P. Isenring
Characterization of a Novel Interaction between the Secretory Na+-K+-Cl- Cotransporter and the Chaperone hsp90
J. Biol. Chem., November 12, 2004; 279(46): 48449 - 48456.
[Abstract] [Full Text] [PDF]

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 J. Neurosci.Home page
B. Lenart, D. B. Kintner, G. E. Shull, and D. Sun
Na-K-Cl Cotransporter-Mediated Intracellular Na+ Accumulation Affects Ca2+ Signaling in Astrocytes in an In Vitro Ischemic Model
J. Neurosci., October 27, 2004; 24(43): 9585 - 9597.
[Abstract] [Full Text] [PDF]

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 QJMHome page
N. Zalunardo, M. Lemaire, M.R. Davids, and M.L. Halperin
Acidosis in a patient with cholera: a need to redefine concepts
QJM, October 1, 2004; 97(10): 681 - 696.
[Abstract] [Full Text] [PDF]

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 J. Exp. Biol.Home page
J. P. Ianowski, R. J. Christensen, and M. J. O'Donnell
Na+ competes with K+ in bumetanide-sensitive transport by Malpighian tubules of Rhodnius prolixus
J. Exp. Biol., October 1, 2004; 207(21): 3707 - 3716.
[Abstract] [Full Text] [PDF]

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 J. Biol. Chem.Home page
C. F. Simard, G. M. Brunet, N. D. Daigle, V. Montminy, L. Caron, and P. Isenring
Self-interacting Domains in the C Terminus of a Cation-Cl- Cotransporter Described for the First Time
J. Biol. Chem., September 24, 2004; 279(39): 40769 - 40777.
[Abstract] [Full Text] [PDF]

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 J. Neurosci.Home page
H. Kaneko, I. Putzier, S. Frings, U. B. Kaupp, and T. Gensch
Chloride Accumulation in Mammalian Olfactory Sensory Neurons
J. Neurosci., September 8, 2004; 24(36): 7931 - 7938.
[Abstract] [Full Text] [PDF]

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 J. Physiol.Home page
J. A. Fraser and C. L.-H. Huang
A quantitative analysis of cell volume and resting potential determination and regulation in excitable cells
J. Physiol., September 1, 2004; 559(2): 459 - 478.
[Abstract] [Full Text] [PDF]

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 JGPHome page
M. Sugita, C. Hirono, and Y. Shiba
Gramicidin-perforated Patch Recording Revealed the Oscillatory Nature of Secretory Cl- Movements in Salivary Acinar Cells
J. Gen. Physiol., June 28, 2004; 124(1): 59 - 69.
[Abstract] [Full Text] [PDF]

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 Am. J. Physiol. Cell Physiol.Home page
L. M. Maglova, W. E. Crowe, and J. M. Russell
Perinuclear localization of Na-K-Cl-cotransporter protein after human cytomegalovirus infection
Am J Physiol Cell Physiol, June 1, 2004; 286(6): C1324 - C1334.
[Abstract] [Full Text] [PDF]

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 J. Physiol.Home page
T. Schilling, F. Lehmann, B. Ruckert, and C. Eder
Physiological mechanisms of lysophosphatidylcholine-induced de-ramification of murine microglia
J. Physiol., May 15, 2004; 557(1): 105 - 120.
[Abstract] [Full Text] [PDF]

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 J. Biol. Chem.Home page
J.-H. Shin, W. Namkung, J. Y. Choi, J.-H. Yoon, and M. G. Lee
Purinergic Stimulation Induces Ca2+-dependent Activation of Na+-K+-2Cl- Cotransporter in Human Nasal Epithelia
J. Biol. Chem., April 30, 2004; 279(18): 18567 - 18574.
[Abstract] [Full Text] [PDF]

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 Am. J. Physiol. Heart Circ. Physiol.Home page
G. O. Andersen, T. Skomedal, M. Enger, A. Fidjeland, T. Brattelid, F. O. Levy, and J.-B. Osnes
{alpha}1-AR-mediated activation of NKCC in rat cardiomyocytes involves ERK-dependent phosphorylation of the cotransporter
Am J Physiol Heart Circ Physiol, April 1, 2004; 286(4): H1354 - H1360.
[Abstract] [Full Text] [PDF]

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 J. Biol. Chem.Home page
H. Guizouarn, N. Gabillat, and F. Borgese
Evidence for Up-regulation of the Endogenous Na-K-2Cl Co-transporter by Molecular Interactions with the Anion Exchanger tAE1 Expressed in Xenopus Oocyte
J. Biol. Chem., March 19, 2004; 279(12): 11513 - 11520.
[Abstract] [Full Text] [PDF]

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 Am. J. Physiol. Cell Physiol.Home page
A. R. Gosmanov, Z. Fan, X. Mi, E. G. Schneider, and D. B. Thomason
ATP-sensitive potassium channels mediate hyperosmotic stimulation of NKCC in slow-twitch muscle
Am J Physiol Cell Physiol, March 1, 2004; 286(3): C586 - C595.
[Abstract] [Full Text]

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 J. Biol. Chem.Home page
E. Gagnon, M. J. Bergeron, G. M. Brunet, N. D. Daigle, C. F. Simard, and P. Isenring
Molecular Mechanisms of Cl- Transport by the Renal Na+-K+-Cl- Cotransporter: IDENTIFICATION OF AN INTRACELLULAR LOCUS THAT MAY FORM PART OF A HIGH AFFINITY Cl--BINDING SITE
J. Biol. Chem., February 13, 2004; 279(7): 5648 - 5654.
[Abstract] [Full Text] [PDF]

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 J. Biol. Chem.Home page
K. Piechotta, N. Garbarini, R. England, and E. Delpire
Characterization of the Interaction of the Stress Kinase SPAK with the Na+-K+-2Cl- Cotransporter in the Nervous System: EVIDENCE FOR A SCAFFOLDING ROLE OF THE KINASE
J. Biol. Chem., December 26, 2003; 278(52): 52848 - 52856.
[Abstract] [Full Text] [PDF]

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 Proc. Natl. Acad. Sci. USAHome page
K. E. Gavrikov, A. V. Dmitriev, K. T. Keyser, and S. C. Mangel
Cation-chloride cotransporters mediate neural computation in the retina
PNAS, December 23, 2003; 100(26): 16047 - 16052.
[Abstract] [Full Text] [PDF]


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