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Vol. 50, Issue 2, 291-314, June 1998
-Aminobutyric AcidA Receptors: Classification on
the Basis of Subunit Structure and Receptor Function
Molecular Neurobiology Unit (E.A.B.), Royal Free Hospital
School of Medicine, London, England;
Neuroscience Discovery (P.S.),
Lilly Research Laboratories, Indianapolis, Indiana;
Department of
Pharmacology (R.W.O.), U.C.L.A. School of Medicine, Los Angeles,
California;
Institute of Pharmacology (H.M.), ETH and University of
Zurich, Zurich, Switzerland;
Department of Biochemical Psychiatry
(W.S.), University of Vienna, Vienna, Austria;
Department of
Experimental Biology (G.B.), University of Cagliari, Cagliari,
Italy;
Pharmaceutical Division (C.B.), Schering A.G., Berlin,
Germany;
Department of Pharmacology, University of Alberta
(A.N.B.), Edmonton, Canada;
Synthélabo Recherche (L.E.R.S.)
(S.Z.L.), Bagneux, France
I. Introduction
A. Earlier Classifications of 
-Aminobutyric Acid Receptors
1. -Aminobutyric acidA and -aminobutyric
acidB receptors.
2. -Aminobutyric acidC receptors.
3. Benzodiazepine receptors.
4. Excitatory -aminobutyric acidA receptors.
B. Conclusion on -Aminobutyric Acid Receptor Types
II. Approaches to the Classification of the
-Aminobutyric AcidA Receptors
A. Transductional Criteria
B. Operational Criteria
C. Structural Criteria
III. The Structures of the -Aminobutyric AcidA
Receptors
A. The Repertoire of Subunit Types
B. The Subunit Number per Receptor Molecule
C. The Subunit Isoforms in One Receptor
D. Possibilities for Subunit Stoichiometry
IV. Principles of the Classification
A. Application of Selectivities at the Binding Site for
Benzodiazepines and Their Functional Analogs
1. The choice of a classification system.
2. Benzodiazepine-responsive -aminobutyric acid receptor
subtypes.
3. Notation.
B. Advantages and Possible Modification of the Classification
C. Nomenclature for the Site Binding Benzodiazepines and Modulators
with Similar Activity
D. Benzodiazepine Insensitivity
V. Chromosomal Localization of -Aminobutyric AcidA
Receptor Genes
VI. Other Binding Sites in Relation to the Receptor Classification
A. Other Modulatory Sites
B. The -Aminobutyric Acid Recognition Site
VII. Conclusions
Acknowledgments
References
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 A. C. Hall, K. C. Rowan, R. J. N. Stevens, J. C. Kelley, and N. L. Harrison The Effects of Isoflurane on Desensitized Wild-Type and {alpha}1(S270H) {gamma}-Aminobutyric Acid Type A Receptors Anesth. Analg., May 1, 2004; 98(5): 1297 - 1304. [Abstract] [Full Text] [PDF]
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D. Berezhnoy, Y. Nyfeler, A. Gonthier, H. Schwob, M. Goeldner, and E. Sigel On the Benzodiazepine Binding Pocket in GABAA Receptors J. Biol. Chem., January 30, 2004; 279(5): 3160 - 3168. [Abstract] [Full Text] [PDF]
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C. T. Moore, C. G. Wilson, C. A. Mayer, S. S. Acquah, V. J. Massari, and M. A. Haxhiu A GABAergic inhibitory microcircuit controlling cholinergic outflow to the airways J Appl Physiol, January 1, 2004; 96(1): 260 - 270. [Abstract] [Full Text] [PDF]
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E. Sanna, M. C. Mostallino, F. Busonero, G. Talani, S. Tranquilli, M. Mameli, S. Spiga, P. Follesa, and G. Biggio Changes in GABAA Receptor Gene Expression Associated with Selective Alterations in Receptor Function and Pharmacology after Ethanol Withdrawal J. Neurosci., December 17, 2003; 23(37): 11711 - 11724. [Abstract] [Full Text] [PDF]
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S. W. Baumann, R. Baur, and E. Sigel Individual Properties of the Two Functional Agonist Sites in GABAA Receptors J. Neurosci., December 3, 2003; 23(35): 11158 - 11166. [Abstract] [Full Text] [PDF]
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 O. A. Weisz and J. P. Johnson Noncoordinate regulation of ENaC: paradigm lost? Am J Physiol Renal Physiol, November 1, 2003; 285(5): F833 - F842. [Abstract] [Full Text] [PDF]
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F.-C. Hsu, G.-J. Zhang, Y. S. H. Raol, R. J. Valentino, D. A. Coulter, and A. R. Brooks-Kayal Repeated neonatal handling with maternal separation permanently alters hippocampal GABAA receptors and behavioral stress responses PNAS, October 14, 2003; 100(21): 12213 - 12218. [Abstract] [Full Text] [PDF]
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 R. A. Stone, J. Liu, R. Sugimoto, C. Capehart, X. Zhu, and K. Pendrak GABA, Experimental Myopia, and Ocular Growth in Chick Invest. Ophthalmol. Vis. Sci., September 1, 2003; 44(9): 3933 - 3946. [Abstract] [Full Text] [PDF]
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S. T. Sinkkonen, S. Mansikkamaki, T. Moykkynen, H. Luddens, M. Uusi-Oukari, and E. R. Korpi Receptor Subtype-Dependent Positive and Negative Modulation of GABAA Receptor Function by Niflumic Acid, a Nonsteroidal Anti-Inflammatory Drug Mol. Pharmacol., September 1, 2003; 64(3): 753 - 763. [Abstract] [Full Text] [PDF]
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K. Gamel-Didelon, L. Kunz, K. J. Fohr, M. Gratzl, and A. Mayerhofer Molecular and Physiological Evidence for Functional {gamma}-Aminobutyric Acid (GABA)-C Receptors in Growth Hormone-secreting Cells J. Biol. Chem., May 23, 2003; 278(22): 20192 - 20195. [Abstract] [Full Text] [PDF]
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A. C. Engblom, F. F. Johansen, and U. Kristiansen Actions and Interactions of Extracellular Potassium and Kainate on Expression of 13 gamma -Aminobutyric Acid Type A Receptor Subunits in Cultured Mouse Cerebellar Granule Neurons J. Biol. Chem., May 2, 2003; 278(19): 16543 - 16550. [Abstract] [Full Text] [PDF]
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P. Follesa, L. Mancuso, F. Biggio, M. C. Mostallino, A. Manca, M. P. Mascia, F. Busonero, G. Talani, E. Sanna, and G. Biggio gamma -Hydroxybutyric Acid and Diazepam Antagonize a Rapid Increase in GABAA Receptors alpha 4 Subunit mRNA Abundance Induced by Ethanol Withdrawal in Cerebellar Granule Cells Mol. Pharmacol., April 1, 2003; 63(4): 896 - 907. [Abstract] [Full Text] [PDF]
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 G. Tamas, A. L. A. Simon, and J. Szabadics Identified Sources and Targets of Slow Inhibition in the Neocortex Science, March 21, 2003; 299(5614): 1902 - 1905. [Abstract] [Full Text] [PDF]
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S. A. G. Visser, F. L. C. Wolters, P. H. van der Graaf, L. A. Peletier, and M. Danhof Dose-Dependent EEG Effects of Zolpidem Provide Evidence for GABAA Receptor Subtype Selectivity in Vivo J. Pharmacol. Exp. Ther., March 1, 2003; 304(3): 1251 - 1257. [Abstract] [Full Text] [PDF]
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F.-C. Hsu and S. S. Smith Progesterone Withdrawal Reduces Paired-Pulse Inhibition in Rat Hippocampus: Dependence on GABAA Receptor alpha 4 Subunit Upregulation J Neurophysiol, January 1, 2003; 89(1): 186 - 198. [Abstract] [Full Text] [PDF]
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Y. A. Blednov, S. Jung, H. Alva, D. Wallace, T. Rosahl, P.-J. Whiting, and R. A. Harris Deletion of the alpha 1 or beta 2 Subunit of GABAA Receptors Reduces Actions of Alcohol and Other Drugs J. Pharmacol. Exp. Ther., January 1, 2003; 304(1): 30 - 36. [Abstract] [Full Text] [PDF]
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R. Riquelme, C. P. Miralles, and A. L. De Blas Bergmann Glia GABAA Receptors Concentrate on the Glial Processes That Wrap Inhibitory Synapses J. Neurosci., December 15, 2002; 22(24): 10720 - 10730. [Abstract] [Full Text] [PDF]
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G. W. Sawyer, D. C. Chiara, R. W. Olsen, and J. B. Cohen Identification of the Bovine gamma -Aminobutyric Acid Type A Receptor alpha Subunit Residues Photolabeled by the Imidazobenzodiazepine [3H]Ro15-4513 J. Biol. Chem., December 13, 2002; 277(51): 50036 - 50045. [Abstract] [Full Text] [PDF]
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P. A. Goldstein, F. P. Elsen, S.-W. Ying, C. Ferguson, G. E. Homanics, and N. L. Harrison Prolongation of Hippocampal Miniature Inhibitory Postsynaptic Currents in Mice Lacking the GABAA Receptor alpha 1 Subunit J Neurophysiol, December 1, 2002; 88(6): 3208 - 3217. [Abstract] [Full Text] [PDF]
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S. W. Baumann, R. Baur, and E. Sigel Forced Subunit Assembly in alpha 1beta 2gamma 2 GABAA Receptors. INSIGHT INTO THE ABSOLUTE ARRANGEMENT J. Biol. Chem., November 22, 2002; 277(48): 46020 - 46025. [Abstract] [Full Text] [PDF]
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D. N. Bowser, D. A. Wagner, C. Czajkowski, B. A. Cromer, M. W. Parker, R. H. Wallace, L. A. Harkin, J. C. Mulley, C. Marini, S. F. Berkovic, et al. Altered kinetics and benzodiazepine sensitivity of a GABAA receptor subunit mutation [gamma 2(R43Q)] found in human epilepsy PNAS, November 12, 2002; 99(23): 15170 - 15175. [Abstract] [Full Text] [PDF]
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M. Scheller and S. A. Forman Coupled and Uncoupled Gating and Desensitization Effects by Pore Domain Mutations in GABAA Receptors J. Neurosci., October 1, 2002; 22(19): 8411 - 8421. [Abstract] [Full Text] [PDF]
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J. E. Kralic, E. R. Korpi, T. K. O'Buckley, G. E. Homanics, and A. L. Morrow Molecular and Pharmacological Characterization of GABAA Receptor alpha 1 Subunit Knockout Mice J. Pharmacol. Exp. Ther., September 1, 2002; 302(3): 1037 - 1045. [Abstract] [Full Text] [PDF]
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I. Sarto, T. Klausberger, N. Ehya, B. Mayer, K. Fuchs, and W. Sieghart A Novel Site on gamma 3 Subunits Important for Assembly of GABAA Receptors J. Biol. Chem., August 16, 2002; 277(34): 30656 - 30664. [Abstract] [Full Text] [PDF]
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M. W. Fleck Molecular Actions of (S)-Desmethylzopiclone (SEP-174559), an Anxiolytic Metabolite of Zopiclone J. Pharmacol. Exp. Ther., August 1, 2002; 302(2): 612 - 618. [Abstract] [Full Text] [PDF]
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N. Collinson, F. M. Kuenzi, W. Jarolimek, K. A. Maubach, R. Cothliff, C. Sur, A. Smith, F. M. Otu, O. Howell, J. R. Atack, et al. Enhanced Learning and Memory and Altered GABAergic Synaptic Transmission in Mice Lacking the alpha 5 Subunit of the GABAA Receptor J. Neurosci., July 1, 2002; 22(13): 5572 - 5580. [Abstract] [Full Text] [PDF]
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J. G. Newell and S. M. J. Dunn Functional Consequences of the Loss of High Affinity Agonist Binding to gamma -Aminobutyric Acid Type A Receptors. IMPLICATIONS FOR RECEPTOR DESENSITIZATION J. Biol. Chem., June 7, 2002; 277(24): 21423 - 21430. [Abstract] [Full Text] [PDF]
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J. H. Holden and C. Czajkowski Different Residues in the GABAA Receptor alpha 1T60-alpha 1K70 Region Mediate GABA and SR-95531 Actions J. Biol. Chem., May 17, 2002; 277(21): 18785 - 18792. [Abstract] [Full Text] [PDF]
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T. Klausberger, J. D. B. Roberts, and P. Somogyi Cell Type- and Input-Specific Differences in the Number and Subtypes of Synaptic GABAA Receptors in the Hippocampus J. Neurosci., April 1, 2002; 22(7): 2513 - 2521. [Abstract] [Full Text] [PDF]
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S.-A. Thompson, P. B. Wingrove, L. Connelly, P. J. Whiting, and K. A. Wafford Tracazolate Reveals a Novel Type of Allosteric Interaction with Recombinant gamma -Aminobutyric AcidA Receptors Mol. Pharmacol., April 1, 2002; 61(4): 861 - 869. [Abstract] [Full Text] [PDF]
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K. M. Wohlfarth, M. T. Bianchi, and R. L. Macdonald Enhanced Neurosteroid Potentiation of Ternary GABAA Receptors Containing the delta Subunit J. Neurosci., March 1, 2002; 22(5): 1541 - 1549. [Abstract] [Full Text] [PDF]
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S. Boehm and H. Kubista Fine Tuning of Sympathetic Transmitter Release via Ionotropic and Metabotropic Presynaptic Receptors Pharmacol. Rev., March 1, 2002; 54(1): 43 - 99. [Abstract] [Full Text] [PDF]
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L. R. McMahon, L. R. Gerak, L. Carter, C. Ma, J. M. Cook, and C. P. France Discriminative Stimulus Effects of Benzodiazepine (BZ)1 Receptor-Selective Ligands in Rhesus Monkeys J. Pharmacol. Exp. Ther., February 1, 2002; 300(2): 505 - 512. [Abstract] [Full Text] [PDF]
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S. B. Christie, C. P. Miralles, and A. L. De Blas GABAergic Innervation Organizes Synaptic and Extrasynaptic GABAA Receptor Clustering in Cultured Hippocampal Neurons J. Neurosci., February 1, 2002; 22(3): 684 - 697. [Abstract] [Full Text] [PDF]
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