Alpha 7 nicotinic acetylcholine receptors: Molecular pharmacology and role in neuroprotection

Current Anaesthesia & Critical Care - Tập 19 - Trang 202-214 - 2008
Ruan van Rensburg1, Paul L. Chazot1
1Durham Centre for Integrative Neuroscience, School of Biological and Biomedical Sciences, Durham University, South Road, Durham, DH1 3LE, UK

Tài liệu tham khảo

Zhao, 2004, Formation of amyloid fibers triggered by phosphatidylserine-containing membranes, Biochemistry, 43, 10302, 10.1021/bi049002c Patrick, 1987, Molecular biology of nicotinic acetylcholine receptors, Ann N Y Acad Sci, 505, 194, 10.1111/j.1749-6632.1987.tb51292.x Vijayaraghavan, 1992, Nicotinic receptors that bind alpha-bungarotoxin on neurons raise intracellular free Ca2+, Neuron, 8, 353, 10.1016/0896-6273(92)90301-S Alkondon, 2004, The nicotinic acetylcholine receptor subtypes and their function in the hippocampus and cerebral cortex, Prog Brain Res, 145, 109, 10.1016/S0079-6123(03)45007-3 Lukas, 1999, International Union of Pharmacology. XX. Current status of the nomenclature for nicotinic acetylcholine receptors and their subunits, Pharmacol Rev, 51, 397 Cooper, 1991, Pentameric structure and subunit stoichiometry of a neuronal nicotinic acetylcholine receptor, Nature, 350, 235, 10.1038/350235a0 Galzi, 1991, Functional architecture of the nicotinic acetylcholine receptor: from electric organ to brain, Annu Rev Pharmacol Toxicol, 31, 37, 10.1146/annurev.pa.31.040191.000345 Lukas, 1992, Heterogeneity and regulation of nicotinic acetylcholine receptors, Int Rev Neurobiol, 34, 25, 10.1016/S0074-7742(08)60097-5 Salvaterra, 1973, Binding of (125I)-alpha bungarotoxin to particulate fractions of rat and guinea pig brain, Biochem Biophys Res Commun, 55, 1311, 10.1016/S0006-291X(73)80037-3 Eterović, 1974, Nicotinic cholinergic receptor in brain detected by binding of alpha-(3H) bungarotoxin, Biochim Biophys Acta, 362, 346, 10.1016/0304-4165(74)90227-X McQuarrie, 1976, Studies on nicotinic acetylcholine receptors in mammalian brain. Preliminary characterization of membrane-bound alpha-bungarotoxin receptors in rat cerebral cortex, J Biol Chem, 251, 6335, 10.1016/S0021-9258(20)81864-X Oswald, 1981, Alpha-bungarotoxin binding and central nervous system nicotinic acetylcholine receptors, Neuroscience, 6, 1, 10.1016/0306-4522(81)90239-6 Elgoyhen, 2001, alpha10: a determinant of nicotinic cholinergic receptor function in mammalian vestibular and cochlear mechanosensory hair cells, Proc Natl Acad Sci USA, 98, 3501, 10.1073/pnas.051622798 Marks, 1986, Nicotinic binding sites in rat and mouse brain: comparison of acetylcholine, nicotine, and alpha-bungarotoxin, Mol Pharmacol, 30, 427 Wonnacott, 1986, alpha-Bungarotoxin binds to low-affinity nicotine binding sites in rat brain, J Neurochem, 47, 1706, 10.1111/j.1471-4159.1986.tb13078.x Goldman, 1987, Members of a nicotinic acetylcholine receptor gene family are expressed in different regions of the mammalian central nervous system, Cell, 48, 965, 10.1016/0092-8674(87)90705-7 Schoepfer, 1990, Brain alpha-bungarotoxin binding protein cDNAs and MAbs reveal subtypes of this branch of the ligand-gated ion channel gene superfamily, Neuron, 5, 35, 10.1016/0896-6273(90)90031-A Amar, 1993, Agonist pharmacology of the neuronal alpha7 nicotinic receptor expressed in Xenopus oocytes, FEBS Lett, 327, 284, 10.1016/0014-5793(93)81005-K Zhang, 1994, Neuronal acetylcholine receptors that bind alpha-bungarotoxin with high affinity function as ligand-gated ion channels, Neuron, 12, 167, 10.1016/0896-6273(94)90161-9 Palma, 1999, Nicotinic acetylcholine receptors assembled from the alpha7 and beta3 subunits, J Biol Chem, 274, 18335, 10.1074/jbc.274.26.18335 Khiroug, 2002, Rat nicotinic ACh receptor alpha7 and beta2 subunits co-assemble to form functional heteromeric nicotinic receptor channels, J Physiol, 540, 425, 10.1113/jphysiol.2001.013847 Clarke, 1985, Nicotinic binding in rat brain: autoradiographic comparison of [3H]acetylcholine, [3H]nicotine, and [125I]-alpha-bungarotoxin, J Neurosci, 5, 1307, 10.1523/JNEUROSCI.05-05-01307.1985 Gault, 1998, Genomic organization and partial duplication of the human alpha7 neuronal nicotinic acetylcholine receptor gene (CHRNA7), Genomics, 52, 173, 10.1006/geno.1998.5363 Belluardo, 2000, Central nicotinic receptors, neurotrophic factors and neuroprotection, Behav Brain Res, 113, 21, 10.1016/S0166-4328(00)00197-2 Elgoyhen, 1994, Alpha9: an acetylcholine receptor with novel pharmacological properties expressed in rat cochlear hair cells, Cell, 79, 705, 10.1016/0092-8674(94)90555-X Lustig, 2001, Molecular cloning and mapping of the human nicotinic acetylcholine receptor alpha10 (CHRNA10), Genomics, 73, 272, 10.1006/geno.2000.6503 Rubboli, 1994, Distribution of neuronal nicotinic receptor subunits in human brain, Neurochem Int, 25, 69, 10.1016/0197-0186(94)90055-8 Breese, 1997, Comparison of the regional expression of nicotinic acetylcholine receptor alpha7 mRNA and [125I]-alpha-bungarotoxin binding in human postmortem brain, J Comp Neurol, 387, 385, 10.1002/(SICI)1096-9861(19971027)387:3<385::AID-CNE5>3.0.CO;2-X Spurden, 1997, Nicotinic receptor distribution in the human thalamus: autoradiographical localization of [3H]nicotine and [125I] alpha-bungarotoxin binding, J Chem Neuroanat, 13, 105, 10.1016/S0891-0618(97)00038-0 Paterson, 2000, Neuronal nicotinic receptors in the human brain, Prog Neurobiol, 61, 75, 10.1016/S0301-0082(99)00045-3 Wonnacott, 1997, Presynaptic nicotinic ACh receptors, Trends Neurosci, 20, 92, 10.1016/S0166-2236(96)10073-4 Sher, 2004, Physiological roles of neuronal nicotinic receptor subtypes: new insights on the nicotinic modulation of neurotransmitter release, synaptic transmission and plasticity, Curr Top Med Chem, 4, 283, 10.2174/1568026043451393 Jensen, 2005, Neuronal nicotinic acetylcholine receptors: structural revelations, target identifications, and therapeutic inspirations, J Med Chem, 48, 4705, 10.1021/jm040219e Choi, 1987, Glutamate neurotoxicity in cortical cell culture, J Neurosci, 7, 357, 10.1523/JNEUROSCI.07-02-00357.1987 Séguéla, 1993, Molecular cloning, functional properties, and distribution of rat brain alpha7: a nicotinic cation channel highly permeable to calcium, J Neurosci, 13, 596, 10.1523/JNEUROSCI.13-02-00596.1993 Rothman, 1995, Excitotoxicity and the NMDA receptor—still lethal after eight years, Trends Neurosci, 18, 57, 10.1016/0166-2236(95)93869-Y Treinin, 1995, A mutated acetylcholine receptor subunit causes neuronal degeneration in C. elegans, Neuron, 14, 871, 10.1016/0896-6273(95)90231-7 Donnelly-Roberts, 1996, In vitro neuroprotective properties of the novel cholinergic channel activator (ChCA), ABT-418, Brain Res, 719, 36, 10.1016/0006-8993(96)00063-7 Gotti, 2004, Neuronal nicotinic receptors: from structure to pathology, Prog Neurobiol, 74, 363, 10.1016/j.pneurobio.2004.09.006 Kem, 2000, The brain alpha7 nicotinic receptor may be an important therapeutic target for the treatment of Alzheimer's disease: studies with DMXBA (GTS-21), Behav Brain Res, 113, 169, 10.1016/S0166-4328(00)00211-4 Picciotto, 2001, Neuronal nicotinic acetylcholine receptor subunit knockout mice: physiological and behavioral phenotypes and possible clinical implications, Pharmacol Ther, 92, 89, 10.1016/S0163-7258(01)00161-9 Martin, 2007, Schizophrenia and the alpha7 Nicotinic Acetylcholine Receptor, Int Rev Neurobiol, 78, 225, 10.1016/S0074-7742(06)78008-4 Morissette, 2007, Anxiety, anxiety disorders, tobacco use, and nicotine: a critical review of interrelationships, Psychol Bull, 133, 245, 10.1037/0033-2909.133.2.245 Orr-Urtreger, 1997, Mice deficient in the alpha7 neuronal nicotinic acetylcholine receptor lack alpha-bungarotoxin binding sites and hippocampal fast nicotinic currents, J Neurosci, 17, 9165, 10.1523/JNEUROSCI.17-23-09165.1997 Orr-Urtreger, 2000, Mice homozygous for the L250T mutation in the alpha7 nicotinic acetylcholine receptor show increased neuronal apoptosis and die within 1 day of birth, J Neurochem, 74, 2154, 10.1046/j.1471-4159.2000.0742154.x Peng, 1994, Human alpha7 acetylcholine receptor: cloning of the alpha7 subunit from the SH-SY5Y cell line and determination of pharmacological properties of native receptors and functional alpha7 homomers expressed in Xenopus oocytes, Mol Pharmacol, 45, 546 Matter-Sadzinski, 1992, Neuronal specificity of the alpha7 nicotinic acetylcholine receptor promoter develops during morphogenesis of the central nervous system, EMBO J, 11, 4529, 10.1002/j.1460-2075.1992.tb05554.x García-Guzmán, 1995, alpha-Bungarotoxin-sensitive nicotinic receptors on bovine chromaffin cells: molecular cloning, functional expression and alternative splicing of the alpha7 subunit, Eur J Neurosci, 7, 647, 10.1111/j.1460-9568.1995.tb00668.x Jones, 2005, Why doesn’t nicotinic ACh receptor immunoreactivity knock out?, Trends Neurosci, 28, 343, 10.1016/j.tins.2005.04.010 Moser, 2007, Evaluating the suitability of nicotinic acetylcholine receptor antibodies for standard immunodetection procedures, J Neurochem, 102, 479, 10.1111/j.1471-4159.2007.04498.x Herber, 2004, Biochemical and histochemical evidence of nonspecific binding of alpha7 nAChR antibodies to mouse brain tissue, J Histochem Cytochem, 52, 1367, 10.1177/002215540405201013 Nagavarapu, 2001, Characterization of a rat neuronal nicotinic acetylcholine receptor alpha7 promoter, J Biol Chem, 276, 16749, 10.1074/jbc.M009712200 Boisclair, 1993, Three clustered Sp1 sites are required for efficient transcription of the TATA-less promoter of the gene for insulin-like growth factor-binding protein-2 from the rat, J Biol Chem, 268, 24892, 10.1016/S0021-9258(19)74549-9 Ebert, 1995, Differential activation of the rat phenylethanolamine N-methyltransferase gene by Sp1 and Egr-1, J Biol Chem, 270, 17299, 10.1074/jbc.270.29.17299 Carrasco-Serrano, 1998, GC- and E-box motifs as regulatory elements in the proximal promoter region of the neuronal nicotinic receptor alpha7 subunit gene, J Biol Chem, 273, 20021, 10.1074/jbc.273.32.20021 Carrasco-Serrano, 2000, Phorbol ester activation of the neuronal nicotinic acetylcholine receptor alpha7 subunit gene: involvement of transcription factor Egr-1, J Neurochem, 74, 932, 10.1046/j.1471-4159.2000.0740932.x Carrasco-Serrano, 2004, Glucocorticoid activation of the neuronal nicotinic acetylcholine receptor alpha7 subunit gene: involvement of transcription factor Egr-1, FEBS Lett, 566, 247, 10.1016/j.febslet.2004.04.049 Tsai-Morris, 1988, 5’ flanking sequence and genomic structure of Egr-1, a murine mitogen inducible zinc finger encoding gene, Nucleic Acids Res, 16, 8835, 10.1093/nar/16.18.8835 Cao, 1993, Detection and characterization of cellular EGR-1 binding to its recognition site, J Biol Chem, 268, 16949, 10.1016/S0021-9258(19)85286-9 Sukhatme, 1988, A zinc finger-encoding gene coregulated with c-fos during growth and differentiation, and after cellular depolarization, Cell, 53, 37, 10.1016/0092-8674(88)90485-0 Enslen, 1994, Roles of calmodulin-dependent protein kinases and phosphatase in calcium-dependent transcription of immediate early genes, J Biol Chem, 269, 20872, 10.1016/S0021-9258(17)31903-8 Abe, 1991, Induction of the ‘zinc finger’ gene after transient focal ischemia in rat cerebral cortex, Neurosci Lett, 123, 248, 10.1016/0304-3940(91)90942-M Herdegen, 1993, The KROX-20 transcription factor in the rat central and peripheral nervous systems: novel expression pattern of an immediate early gene-encoded protein, Neuroscience, 57, 41, 10.1016/0306-4522(93)90110-2 Utsugisawa, 1999, Changes with aging and ischemia in nicotinic acetylcholine receptor subunit alpha7 mRNA expression in postmortem human frontal cortex and putamen, Neurosci Lett, 270, 145, 10.1016/S0304-3940(99)00473-5 Bessis, 1993, Negative regulatory elements upstream of a novel exon of the neuronal nicotinic acetylcholine receptor alpha2 subunit gene, Nucleic Acids Res, 21, 2185, 10.1093/nar/21.9.2185 Milton, 1995, The neuronal nicotinic acetylcholine receptor alpha2 subunit gene promoter is activated by the Brn-3b POU family transcription factor and not by Brn-3a or Brn-3c, J Biol Chem, 270, 15143, 10.1074/jbc.270.25.15143 Yang, 1995, Transcriptional analysis of acetylcholine receptor alpha3 gene promoter motifs that bind Sp1 and AP2, J Biol Chem, 270, 8514, 10.1074/jbc.270.15.8514 Liu, 1999, Cell type-specific activation of neuronal nicotinic acetylcholine receptor subunit genes by Sox10, J Neurosci, 19, 9747, 10.1523/JNEUROSCI.19-22-09747.1999 Du, 1998, Differential effects of heterogeneous nuclear ribonucleoprotein K on Sp1- and Sp3-mediated transcriptional activation of a neuronal nicotinic acetylcholine receptor promoter, J Biol Chem, 273, 19877, 10.1074/jbc.273.31.19877 Campos-Caro, 2001, Activity of the nicotinic acetylcholine receptor alpha5 and alpha7 subunit promoters in muscle cells, DNA Cell Biol, 20, 657, 10.1089/104454901753340640 Ebihara, 2002, Genomic organization and promoter analysis of the human nicotinic acetylcholine receptor alpha6 subunit (CHNRA6) gene: Alu and other elements direct transcriptional repression, Gene, 298, 101, 10.1016/S0378-1119(02)00925-3 Valor, 2003, Transcriptional regulation by activation and repression elements located at the 5’-noncoding region of the human alpha9 nicotinic receptor subunit gene, J Biol Chem, 278, 37249, 10.1074/jbc.M307043200 Lustig, 2002, Chromosome location and characterization of the human nicotinic acetylcholine receptor subunit alpha (alpha) 9 (CHRNA9) gene, Cytogenet Genome Res, 98, 154, 10.1159/000069804 Poser, 2003, Brain-derived neurotrophic factor protection of cortical neurons from serum withdrawal-induced apoptosis is inhibited by cAMP, J Neurosci, 23, 4420, 10.1523/JNEUROSCI.23-11-04420.2003 Avramopoulou, 2004, Soluble, oligomeric, and ligand-binding extracellular domain of the human alpha7 acetylcholine receptor expressed in yeast: replacement of the hydrophobic cysteine loop by the hydrophilic loop of the ACh-binding protein enhances protein solubility, J Biol Chem, 279, 38287, 10.1074/jbc.M402533200 Arias, 1997, Topology of ligand binding sites on the nicotinic acetylcholine receptor, Brain Res Brain Res Rev, 25, 133, 10.1016/S0165-0173(97)00020-9 Brejc, 2001, Crystal structure of an ACh-binding protein reveals the ligand-binding domain of nicotinic receptors, Nature, 411, 269, 10.1038/35077011 Celie, 2004, Nicotine and carbamylcholine binding to nicotinic acetylcholine receptors as studied in AChBP crystal structures, Neuron, 41, 907, 10.1016/S0896-6273(04)00115-1 Celie, 2005, Crystal structure of nicotinic acetylcholine receptor homolog AChBP in complex with an alpha-conotoxin PnIA variant, Nat Struct Mol Biol, 12, 582, 10.1038/nsmb951 Smit, 2001, A glia-derived acetylcholine-binding protein that modulates synaptic transmission, Nature, 411, 261, 10.1038/35077000 Smit, 2003, Structure and function of AChBP, homologue of the ligand-binding domain of the nicotinic acetylcholine receptor, Ann N Y Acad Sci, 998, 81, 10.1196/annals.1254.010 Marinou, 2003, Identification of regions involved in the binding of alpha-bungarotoxin to the human alpha7 neuronal nicotinic acetylcholine receptor using synthetic peptides, Biochem J, 372, 543, 10.1042/bj20021537 Kemp, 1990, Protein kinase recognition sequence motifs, Trends Biochem Sci, 15, 342, 10.1016/0968-0004(90)90073-K Unwin, 1988, Arrangement of the acetylcholine receptor subunits in the resting and desensitized states, determined by cryoelectron microscopy of crystallized Torpedo postsynaptic membranes, J Cell Biol, 107, 1123, 10.1083/jcb.107.3.1123 Unwin, 1993, Nicotinic acetylcholine receptor at 9Å resolution, J Mol Biol, 229, 1101, 10.1006/jmbi.1993.1107 Unwin, 1995, Acetylcholine receptor channel imaged in the open state, Nature, 373, 37, 10.1038/373037a0 Unwin, 1996, Projection structure of the nicotinic acetylcholine receptor: distinct conformations of the alpha subunits, J Mol Biol, 257, 586, 10.1006/jmbi.1996.0187 Bertrand, 1993, Stratification of the channel domain in neurotransmitter receptors, Curr Opin Cell Biol, 5, 688, 10.1016/0955-0674(93)90141-C Karlin, 1993, Structure of nicotinic acetylcholine receptors, Curr Opin Neurobiol, 3, 299, 10.1016/0959-4388(93)90121-E Ortells, 1996, A mixed helix-beta-sheet model of the transmembrane region of the nicotinic acetylcholine receptor, Protein Eng, 9, 51, 10.1093/protein/9.1.51 Görne-Tschelnokow, 1994, The transmembrane domains of the nicotinic acetylcholine receptor contain alpha-helical and beta structures, EMBO J, 13, 338, 10.1002/j.1460-2075.1994.tb06266.x Halevi, 2002, The C. elegans ric-3 gene is required for maturation of nicotinic acetylcholine receptors, EMBO J, 21, 1012, 10.1093/emboj/21.5.1012 Treinin, 1998, Two functionally dependent acetylcholine subunits are encoded in a single Caenorhabditis elegans operon, Proc Natl Acad Sci USA, 95, 15492, 10.1073/pnas.95.26.15492 Dominguez del Toro, 1994, Immunocytochemical localization of the alpha7 subunit of the nicotinic acetylcholine receptor in the rat central nervous system, J Comp Neurol, 349, 325, 10.1002/cne.903490302 Keller, 2001, Adjacent basic amino acid residues recognized by the COP I complex and ubiquitination govern endoplasmic reticulum to cell surface trafficking of the nicotinic acetylcholine receptor alpha-subunit, J Biol Chem, 276, 18384, 10.1074/jbc.M100691200 Lansdell, 2005, RIC-3 enhances functional expression of multiple nicotinic acetylcholine receptor subtypes in mammalian cells, Mol Pharmacol, 68, 1431, 10.1124/mol.105.017459 Williams, 2005, Ric-3 promotes functional expression of the nicotinic acetylcholine receptor alpha7 subunit in mammalian cells, J Biol Chem, 280, 1257, 10.1074/jbc.M410039200 Cooper, 1997, Host cell-specific folding and assembly of the neuronal nicotinic acetylcholine receptor alpha7 subunit, J Neurochem, 68, 2140, 10.1046/j.1471-4159.1997.68052140.x Sweileh, 2000, Multistep expression and assembly of neuronal nicotinic receptors is both host-cell- and receptor-subtype-dependent, Brain Res Mol Brain Res, 75, 293, 10.1016/S0169-328X(99)00302-2 Halevi, 2003, Conservation within the RIC-3 gene family. Effectors of mammalian nicotinic acetylcholine receptor expression, J Biol Chem, 278, 34411, 10.1074/jbc.M300170200 Blount, 1991, BIP associates with newly synthesized subunits of the mouse muscle nicotinic receptor, J Cell Biol, 113, 1125, 10.1083/jcb.113.5.1125 Chang, 1997, Calnexin-dependent enhancement of nicotinic acetylcholine receptor assembly and surface expression, J Biol Chem, 272, 28925, 10.1074/jbc.272.46.28925 Jeanclos, 2001, The chaperone protein 14-3-3eta interacts with the nicotinic acetylcholine receptor alpha4 subunit. Evidence for a dynamic role in subunit stabilization, J Biol Chem, 276, 28281, 10.1074/jbc.M011549200 Alexander JK, Jefford G, Criado M, Sagher D, Green WN. Ric-3 levels in the ER determine whether alpha-7 nicotinic receptors are expressed at the surface or retained in the ER. Society for Neuroscience Poster, 2007; 575.3/K3. Cheng, 2005, Cell surface expression of 5-hydroxytryptamine type 3 receptors is promoted by RIC-3, J Biol Chem, 280, 22502, 10.1074/jbc.M414341200 Benwell, 1988, Evidence that tobacco smoking increases the density of (−)-[3H]nicotine binding sites in human brain, J Neurochem, 50, 1243, 10.1111/j.1471-4159.1988.tb10600.x Nybäck, 1989, Attempts to visualize nicotinic receptors in the brain of monkey and man by positron emission tomography, Prog Brain Res, 79, 313, 10.1016/S0079-6123(08)62490-5 Romanelli, 1988, Subchronic treatment of rats with nicotine: interconversion of nicotinic receptor subtypes in brain, Eur J Pharmacol, 148, 289, 10.1016/0014-2999(88)90577-8 Wonnacott, 1990, The paradox of nicotinic acetylcholine receptor upregulation by nicotine, Trends Pharmacol Sci, 11, 216, 10.1016/0165-6147(90)90242-Z Flores, 1992, A subtype of nicotinic cholinergic receptor in rat brain is composed of alpha4 and beta2 subunits and is up-regulated by chronic nicotine treatment, Mol Pharmacol, 41, 31 Barrantes, 1995, alpha-Bungarotoxin binding sites in rat hippocampal and cortical cultures: initial characterisation, colocalisation with alpha7 subunits and up-regulation by chronic nicotine treatment, Brain Res, 672, 228, 10.1016/0006-8993(94)01386-V Rowell, 1997, Dose-response relationship for nicotine-induced up-regulation of rat brain nicotinic receptors, J Neurochem, 68, 1982, 10.1046/j.1471-4159.1997.68051982.x Gopalakrishnan, 1997, Regulation of human alpha4 beta2 neuronal nicotinic acetylcholine receptors by cholinergic channel ligands and second messenger pathways, Mol Pharmacol, 52, 524, 10.1124/mol.52.3.524 Zoli, 1998, Identification of four classes of brain nicotinic receptors using beta2 mutant mice, J Neurosci, 18, 4461, 10.1523/JNEUROSCI.18-12-04461.1998 Pauly, 1991, An autoradiographic analysis of cholinergic receptors in mouse brain after chronic nicotine treatment, J Pharmacol Exp Ther, 258, 1127 Peng, 1997, Chronic nicotine treatment up-regulates alpha3 and alpha7 acetylcholine receptor subtypes expressed by the human neuroblastoma cell line SH-SY5Y, Mol Pharmacol, 51, 776, 10.1124/mol.51.5.776 Quik, 1996, Similarity between rat brain nicotinic alpha-bungarotoxin receptors and stably expressed alpha-bungarotoxin binding sites, J Neurochem, 67, 145, 10.1046/j.1471-4159.1996.67010145.x Molinari, 1998, Up-regulation of human alpha7 nicotinic receptors by chronic treatment with activator and antagonist ligands, Eur J Pharmacol, 347, 131, 10.1016/S0014-2999(98)00084-3 Schwartz, 1985, In vivo regulation of [3H]acetylcholine recognition sites in brain by nicotinic cholinergic drugs, J Neurochem, 45, 427, 10.1111/j.1471-4159.1985.tb04005.x Olale, 1997, Chronic nicotine exposure differentially affects the function of human alpha3, alpha4, and alpha7 neuronal nicotinic receptor subtypes, J Pharmacol Exp Ther, 283, 675 Reitstetter, 1999, Dependence of nicotinic acetylcholine receptor recovery from desensitization on the duration of agonist exposure, J Pharmacol Exp Ther, 289, 656 Rowell, 1998, Long-lasting inactivation of nicotinic receptor function in vitro by treatment with high concentrations of nicotine, Neuropharmacology, 37, 103, 10.1016/S0028-3908(97)00193-7 Marks, 1992, Nicotine binding and nicotinic receptor subunit RNA after chronic nicotine treatment, J Neurosci, 12, 2765, 10.1523/JNEUROSCI.12-07-02765.1992 Bencherif, 1995, Mechanisms of up-regulation of neuronal nicotinic acetylcholine receptors in clonal cell lines and primary cultures of fetal rat brain, J Pharmacol Exp Ther, 275, 987 Zhang, 1995, Regulation of alpha4 beta2 nicotinic acetylcholine receptors in M10 cells following treatment with nicotinic agents, Neuroreport, 6, 313, 10.1097/00001756-199501000-00022 Madhok, 1995, Nicotine regulates nicotinic cholinergic receptors and subunit mRNAs in PC 12 cells through protein kinase A, Brain Res Mol Brain Res, 32, 143, 10.1016/0169-328X(95)00073-2 Hsu, 1997, Nicotine enhances the cyclic AMP-dependent protein kinase-mediated phosphorylation of alpha4 subunits of neuronal nicotinic receptors, J Neurochem, 69, 2427, 10.1046/j.1471-4159.1997.69062427.x Eilers, 1997, Functional deactivation of the major neuronal nicotinic receptor caused by nicotine and a protein kinase C-dependent mechanism, Mol Pharmacol, 52, 1105, 10.1124/mol.52.6.1105 Chazot, 2002, Spreading depression-induced preconditioning in the mouse cortex: differential changes in the protein expression of ionotropic nicotinic acetylcholine and glutamate receptors, J Neurochem, 83, 1235, 10.1046/j.1471-4159.2002.01240.x Tohgi, 1998, Alterations with aging and ischemia in nicotinic acetylcholine receptor subunits alpha4 and beta2 messenger RNA expression in postmortem human putamen. Implications for susceptibility to parkinsonism, Brain Res, 791, 186, 10.1016/S0006-8993(98)00093-6 De Koninck, 1995, Differential regulation of neuronal nicotinic ACh receptor subunit genes in cultured neonatal rat sympathetic neurons: specific induction of alpha7 by membrane depolarization through a Ca2+/calmodulin-dependent kinase pathway, J Neurosci, 15, 7966, 10.1523/JNEUROSCI.15-12-07966.1995 Ridley, 2001, Differential effects of chronic drug treatment on alpha3* and alpha7 nicotinic receptor binding sites, in hippocampal neurones and SH-SY5Y cells, Br J Pharmacol, 133, 1286, 10.1038/sj.bjp.0704207 Marin, 1994, Nicotine protects cultured striatal neurones against N-methyl-d-aspartate receptor-mediated neurotoxicity, Neuroreport, 5, 1977, 10.1097/00001756-199410000-00035 Semba, 1996, Nicotine protects against the dexamethasone potentiation of kainic acid-induced neurotoxicity in cultured hippocampal neurons, Brain Res, 735, 335, 10.1016/0006-8993(96)00926-2 Dajas-Bailador, 2000, The alpha7 nicotinic acetylcholine receptor subtype mediates nicotine protection against NMDA excitotoxicity in primary hippocampal cultures through a Ca(2+) dependent mechanism, Neuropharmacology, 39, 2799, 10.1016/S0028-3908(00)00127-1 Tohgi, 2000, Protective effect of nicotine through nicotinic acetylcholine receptor alpha7 on hypoxia-induced membrane disintegration and DNA fragmentation of cultured PC12 cells, Neurosci Lett, 285, 91, 10.1016/S0304-3940(00)01026-0 Hejmadi, 2003, Neuroprotection by nicotine against hypoxia-induced apoptosis in cortical cultures involves activation of multiple nicotinic acetylcholine receptor subtypes, Mol Cell Neurosci, 24, 779, 10.1016/S1044-7431(03)00244-6 Jonnala, 2001, Relationship between the increased cell surface alpha7 nicotinic receptor expression and neuroprotection induced by several nicotinic receptor agonists, J Neurosci Res, 66, 565, 10.1002/jnr.10022 Meyer, 1998, Neuroprotective and memory-related actions of novel alpha-7 nicotinic agents with different mixed agonist/antagonist properties, J Pharmacol Exp Ther, 284, 1026 Briggs, 1998, Activation and inhibition of the human alpha7 nicotinic acetylcholine receptor by agonists, Neuropharmacology, 37, 1095, 10.1016/S0028-3908(98)00110-5 Akaike, 1994, Nicotine-induced protection of cultured cortical neurons against N-methyl-d-aspartate receptor-mediated glutamate cytotoxicity, Brain Res, 644, 181, 10.1016/0006-8993(94)91678-0 Kaneko, 1997, Nicotine protects cultured cortical neurons against glutamate-induced cytotoxicity via alpha7-neuronal receptors and neuronal CNS receptors, Brain Res, 765, 135, 10.1016/S0006-8993(97)00556-8 Maggio, 1998, Nicotine prevents experimental parkinsonism in rodents and induces striatal increase of neurotrophic factors, J Neurochem, 71, 2439, 10.1046/j.1471-4159.1998.71062439.x O’Neill, 1998, Histological and behavioral protection by (-)-nicotine against quinolinic acid-induced neurodegeneration in the hippocampus, Neurobiol Learn Mem, 69, 46, 10.1006/nlme.1997.3803 Borlongan, 1995, (−)-Nicotine protects against systemic kainic acid-induced excitotoxic effects, Exp Neurol, 136, 261, 10.1006/exnr.1995.1103