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(Received for publication, April 3, 1996)
From the Departments of Human Physiology and
§ Biological Chemistry, University of California School of
Medicine, Davis, California 95616
Using a combination of data base searching,
polymerase chain reaction, and library screening, we have identified a
putative K-Cl cotransporter isoform (KCC2) in rat brain that is
specifically localized in neurons. A cDNA of 5566 bases was
obtained from overlapping clones and encoded a protein of 1116 amino
acids with a deduced molecular mass of 123.6 kDa. Over its full length,
the amino acid sequence of KCC2 is 67% identical to the widely
distributed K-Cl cotransporter isoform (KCC1) identified in rat brain
and rabbit kidney (Gillen, C., Brill, S., Payne, J.A., and Forbush, B.,
III (1996) J. Biol. Chem. 271, 16237-16244) but only
~25% identical to other members of the cation-chloride cotransporter
gene family, including ``loop'' diuretic-sensitive Na-K-Cl
cotransport and thiazide-sensitive Na-Cl cotransport. Based on analysis
of the primary structure as well as homology with other cation-chloride
cotransporters, we predict 12 transmembrane segments bounded by N- and
C-terminal cytoplasmic regions. Four sites for N-linked
glycosylation are predicted on an extracellular intermembrane loop
between putative transmembrane segments 5 and 6. Northern blot analysis
using a KCC2-specific cDNA probe revealed a very highly expressed
~5.6-kilobase transcript only in brain. Reverse
transcriptase-polymerase chain reaction revealed that KCC1 was present
in rat primary astrocytes and rat C6 glioma cells but that KCC2 was
completely absent from these cells, suggesting KCC2 was not of glial
cell origin. In situ hybridization studies demonstrated
that the KCC2 transcript was expressed at high levels in neurons
throughout the central nervous system, including CA1-CA4 pyramidal
neurons of the hippocampus, granular cells and Purkinje neurons of the
cerebellum, and many groups of neurons throughout the brainstem.
Volume 271, Number 27,
Issue of July 5, 1996
pp. 16245-16252
©1996 by The American Society for Biochemistry and Molecular Biology, Inc.
A NEURONAL-SPECIFIC ISOFORM
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J. M. Russell Sodium-Potassium-Chloride Cotransport Physiol Rev, January 1, 2000; 80(1): 211 - 276. [Abstract] [Full Text] [PDF]
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J. E. Race, F. N. Makhlouf, P. J. Logue, F. H. Wilson, P. B. Dunham, and E. J. Holtzman Molecular cloning and functional characterization of KCC3, a new K-Cl cotransporter Am J Physiol Cell Physiol, December 1, 1999; 277(6): C1210 - C1219. [Abstract] [Full Text] [PDF]
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W. Su, B. E. Shmukler, M. N. Chernova, A. K. Stuart-Tilley, L. de Franceschi, C. Brugnara, and S. L. Alper Mouse K-Cl cotransporter KCC1: cloning, mapping, pathological expression, and functional regulation Am J Physiol Cell Physiol, November 1, 1999; 277(5): C899 - C912. [Abstract] [Full Text] [PDF]
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I. Ehrlich, S. Lohrke, and E. Friauf Shift from depolarizing to hyperpolarizing glycine action in rat auditory neurones is due to age-dependent Cl- regulation J. Physiol., October 1, 1999; 520(1): 121 - 137. [Abstract] [Full Text] [PDF]
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M. Flagella, L. L. Clarke, M. L. Miller, L. C. Erway, R. A. Giannella, A. Andringa, L. R. Gawenis, J. Kramer, J. J. Duffy, T. Doetschman, et al. Mice Lacking the Basolateral Na-K-2Cl Cotransporter Have Impaired Epithelial Chloride Secretion and Are Profoundly Deaf J. Biol. Chem., September 17, 1999; 274(38): 26946 - 26955. [Abstract] [Full Text] [PDF]
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W. Jarolimek, A. Lewen, and U. Misgeld A Furosemide-Sensitive K+-Cl- Cotransporter Counteracts Intracellular Cl- Accumulation and Depletion in Cultured Rat Midbrain Neurons J. Neurosci., June 15, 1999; 19(12): 4695 - 4704. [Abstract] [Full Text] [PDF]
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D. B. Mount, A. Mercado, L. Song, J. Xu, A. L. George Jr., E. Delpire, and G. Gamba Cloning and Characterization of KCC3 and KCC4, New Members of the Cation-Chloride Cotransporter Gene Family J. Biol. Chem., June 4, 1999; 274(23): 16355 - 16362. [Abstract] [Full Text] [PDF]
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J. R. Williams, J. W. Sharp, V. G. Kumari, M. Wilson, and J. A. Payne The Neuron-specific K-Cl Cotransporter, KCC2. ANTIBODY DEVELOPMENT AND INITIAL CHARACTERIZATION OF THE PROTEIN J. Biol. Chem., April 30, 1999; 274(18): 12656 - 12664. [Abstract] [Full Text] [PDF]
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Y. Kakazu, N. Akaike, S. Komiyama, and J. Nabekura Regulation of Intracellular Chloride by Cotransporters in Developing Lateral Superior Olive Neurons J. Neurosci., April 15, 1999; 19(8): 2843 - 2851. [Abstract] [Full Text] [PDF]
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K. Hiki, R. J. D'Andrea, J. Furze, J. Crawford, E. Woollatt, G. R. Sutherland, M. A. Vadas, and J. R. Gamble Cloning, Characterization, and Chromosomal Location of a Novel Human K+-Cl- Cotransporter J. Biol. Chem., April 9, 1999; 274(15): 10661 - 10667. [Abstract] [Full Text] [PDF]
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 S. Haapa, S. Suomalainen, S. Eerikäinen, M. Airaksinen, L. Paulin, and H. Savilahti An Efficient DNA Sequencing Strategy Based on the Bacteriophage Mu in Vitro DNA Transposition Reaction Genome Res., March 1, 1999; 9(3): 308 - 315. [Abstract] [Full Text]
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S. N. Orlov, N. C. Adragna, V. A. Adarichev, and P. Hamet Genetic and biochemical determinants of abnormal monovalent ion transport in primary hypertension Am J Physiol Cell Physiol, March 1, 1999; 276(3): C511 - C536. [Abstract] [Full Text] [PDF]
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C. M. Gillen and B. Forbush III Functional interaction of the K-Cl cotransporter (KCC1) with the Na-K-Cl cotransporter in HEK-293 cells Am J Physiol Cell Physiol, February 1, 1999; 276(2): C328 - C336. [Abstract] [Full Text] [PDF]
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D. W. Hochman, R. D'Ambrosio, D. Janigro, and P. A. Schwartzkroin Extracellular Chloride and the Maintenance of Spontaneous Epileptiform Activity in Rat Hippocampal Slices J Neurophysiol, January 1, 1999; 81(1): 49 - 59. [Abstract] [Full Text] [PDF]
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H. Liapis, M. Nag, and D. M. Kaji K-Cl cotransporter expression in the human kidney Am J Physiol Cell Physiol, December 1, 1998; 275(6): C1432 - C1437. [Abstract] [Full Text] [PDF]
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E. J. Holtzman, S. Kumar, C. A. Faaland, F. Warner, P. J. Logue, S. J. Erickson, G. Ricken, J. Waldman, S. Kumar, and P. B. Dunham Cloning, characterization, and gene organization of K-Cl cotransporter from pig and human kidney and C. elegans Am J Physiol Renal Physiol, October 1, 1998; 275(4): F550 - F564. [Abstract] [Full Text] [PDF]
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P. K Lauf K+-Cl- cotransport: 'to be or not to be' oxygen sensitive J. Physiol., August 15, 1998; 511(1): 1 - 1. [Full Text] [PDF]
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P. Isenring, S. C. Jacoby, and B. Forbush III The role of transmembrane domain 2 in cation transport by the Na-K-Cl cotransporter PNAS, June 9, 1998; 95(12): 7179 - 7184. [Abstract] [Full Text] [PDF]
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P. Isenring, S. C. Jacoby, J. A. Payne, and B. Forbush III Comparison of Na-K-Cl Cotransporters. NKCC1, NKCC2, AND THE HEK CELL Na-K-Cl COTRANSPORTER J. Biol. Chem., May 1, 1998; 273(18): 11295 - 11301. [Abstract] [Full Text] [PDF]
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J. A. Payne Functional characterization of the neuronal-specific K-Cl cotransporter: implications for [K+]o regulation Am J Physiol Cell Physiol, November 1, 1997; 273(5): C1516 - C1525. [Abstract] [Full Text] [PDF]
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P. Isenring and B. Forbush III Ion and Bumetanide Binding by the Na-K-Cl Cotransporter. IMPORTANCE OF TRANSMEMBRANE DOMAINS J. Biol. Chem., September 26, 1997; 272(39): 24556 - 24562. [Abstract] [Full Text] [PDF]
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C. M. Gillen, S. Brill, J. A. Payne, and B. Forbush III Molecular Cloning and Functional Expression of the K-Cl Cotransporter from Rabbit, Rat, and Human. A NEW MEMBER OF THE CATION-CHLORIDE COTRANSPORTER FAMILY J. Biol. Chem., July 5, 1996; 271(27): 16237 - 16244. [Abstract] [Full Text] [PDF]
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L. Caron, F. Rousseau, E. Gagnon, and P. Isenring Cloning and Functional Characterization of a Cation-Cl- Cotransporter-interacting Protein J. Biol. Chem., October 6, 2000; 275(41): 32027 - 32036. [Abstract] [Full Text] [PDF]
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A. Mercado, L. Song, N. Vazquez, D. B. Mount, and G. Gamba Functional Comparison of the K+-Cl- Cotransporters KCC1 and KCC4 J. Biol. Chem., September 22, 2000; 275(39): 30326 - 30334. [Abstract] [Full Text] [PDF]
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P. Sangan, S. R. Brill, S. Sangan, B. Forbush III, and H. J. Binder Basolateral K-Cl Cotransporter Regulates Colonic Potassium Absorption in Potassium Depletion J. Biol. Chem., September 29, 2000; 275(40): 30813 - 30816. [Abstract] [Full Text] [PDF]
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M. Di Fulvio, T. M. Lincoln, P. K. Lauf, and N. C. Adragna Protein Kinase G Regulates Potassium Chloride Cotransporter-3 Expression in Primary Cultures of Rat Vascular Smooth Muscle Cells J. Biol. Chem., June 8, 2001; 276(24): 21046 - 21052. [Abstract] [Full Text] [PDF]
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N. Vazquez, A. Monroy, E. Dorantes, R. A. Munoz-Clares, and G. Gamba Functional differences between flounder and rat thiazide-sensitive Na-Cl cotransporter Am J Physiol Renal Physiol, April 1, 2002; 282(4): F599 - F607. [Abstract] [Full Text] [PDF]
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