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14-September-2008 11:27:24 - Androgen receptor Androgen receptor Structure of the ligand binding domain of the androgen receptor rainbow cartoon complexed with testosterone white sticks based on PDB 2AM9. Available structures: 1e3g, 1gs4, 1i37, 1i38, 1r4i, 1t5z, 1t63, 1t65, 1t73, 1t74, 1t76, 1t79, 1t7f, 1t7m, 1t7r, 1t7t, 1xj7, 1xnn, 1xow, 1xq3, 1z95, 2am9, 2ama, 2amb, 2ao6, 2ax6, 2ax7, 2ax8, 2ax9, 2axa, 2ihq, 2nw4, 2oz7 Identifiers Symbols AR; AIS; DHTR; HUMARA; KD; NR3C4; SBMA; SMAX1; TFM External IDs OMIM: 313700 MGI: 88064 HomoloGene: 28 Gene ontology Molecular function: transcription factor activity receptor activity androgen receptor activity steroid binding androgen binding zinc ion binding lipid binding sequence-specific DNA binding metal ion binding protein dimerization activity Cellular component: nucleus cytoplasm Biological process: in utero embryonic development transcription regulation of transcription, DNA-dependent transport signal transduction cell-cell signaling sex differentiation cell proliferation male gonad development cell growth male somatic sex determination androgen receptor signaling pathway prostate gland development RNA expression pattern More reference expression data Orthologs Human Mouse Entrez 367 11835 Ensembl ENSG00000169083 ENSMUSG00000046532 Uniprot P10275 P19091 Refseq NM_000044 mRNA NP_000035 protein NM_013476 mRNA NP_038504 protein Location Chr X: 66.68 - 66.87 Mb Chr X: 94.35 - 94.52 Mb Pubmed search 1 2 The androgen receptor AR, also known as NR3C4 nuclear receptor subfamily 3, group C, member 4, is a type of nuclear receptor1 which is activated by binding of either of the androgenic hormones testosterone or dihydrotestosterone.2 The androgen receptor is most closely related to the progesterone receptor, and progestins in higher dosages can block the androgen receptor.34 The main function of the androgen receptor is as a DNA binding transcription factor which regulates gene expression;5 however, the androgen receptor has other functions as well.6 Androgen regulated genes are critical for the development and maintenance of the male sexual phenotype. Contents 1 Function 1.1 Androgenic hormones and their effect on development 1.2 Mechanism of androgen receptor action 1.2.1 Genomic 1.2.2 Non-genomic 2 Genetics 2.1 Gene 2.2 AR deficiencies 3 Structure 3.1 Isoforms 3.2 Domains 4 References 5 See also 6 External links Function Androgenic hormones and their effect on development In some cell types testosterone interacts directly with androgen receptors while in others testosterone is converted by 5-alpha-reductase to dihydrotestosterone, an even more potent agonist for androgen receptor activation.7 Testosterone appears to be the primary androgen receptor activating hormone in the Wolffian duct while dihydrotestosterone is the main androgenic hormone in the urogenital sinus, urogenital tubercle, and hair follicles.8 Hence testosterone is primarily responsible for the development of male primary sexual characteristics while dihydrotestosterone is responsible for secondary male characteristics. Androgens cause slow epiphysis, or maturation of the bones, but more of the potent epiphysis effect comes from the estrogen produced by aromatization of androgens. Steroid users of teen age may find that their growth had been stunted by androgen and/or estrogen excess. People with too little sex hormones can be short during puberty but end up taller as adults as in androgen insensitivity syndrome or estrogen insensitivity syndrome.9 Mechanism of androgen receptor action Genomic The primary mechanism of action for androgen receptors is direct regulation of gene transcription. The binding of an androgen to the androgen receptor results in a conformational change in the receptor which in turn causes dissociation of heat shock proteins, transport from the cytosol into the cell nucleus, and dimerization. The androgen receptor dimer binds to a specific sequence of DNA known as a hormone response element. Androgen receptors interact with other proteins in the nucleus resulting in up or down regulation of specific gene transcription.10 Up-regulation or activation of transcription results in increased synthesis of messenger RNA which in turn is transcribed by ribosomes to produce specific proteins. One of the known target genes of androgen receptor activation is insulin-like growth factor I IGF-1.11 Thus, changes in levels of specific proteins in cells is one way that androgen receptors control cell behavior. Non-genomic More recently, androgen receptors have been shown to have a second mode of action. As has been also found for other steroid hormone receptors such as estrogen receptors, androgen receptors can have actions that are independent of their interactions with DNA.612 Androgen receptors interact with certain signal transduction proteins in the cytoplasm. Androgen binding to cytoplasmic androgen receptors can cause rapid changes in cell function independent of changes in gene transcription, such as changes in ion transport. Regulation of signal transduction pathways by cytoplasmic androgen receptors can indirectly lead to changes in gene transcription, for example, by leading to phosphorylation of other transcription factors. One function of androgen receptor that is independent of direct binding to its target DNA sequence, is facilitated by recruitment via other DNA binding proteins. One example is serum response factor, a protein which activates several genes that cause muscle growth.13 Genetics Gene The AR gene for the androgen receptor is located on the X chromosome at Xq11-12.1415 AR deficiencies The androgen insensitivity syndrome, formerly known as testicular feminization, is caused by a mutation of the Androgen Receptor gene located on the X chromosome locus:Xq11-Xq12.16 The androgen receptor seems to affect neuron physiology and is defective in Kennedy disease.1718 In addition, point mutations and trinucleotide repeat polymorphisms has been linked to a number of additional disorders.19 Structure Structural domains of the two isoforms AR-A and AR-B of the human androgen receptor. Numbers above the bars refer to the amino acid residues which separate the domains starting from the N-terminus left to C-terminus right. NTD = N-terminal domain, DBD = DNA binding domain. LBD = ligand binding domain. AF = activation function. Structural domains of the two isoforms AR-A and AR-B of the human androgen receptor. Numbers above the bars refer to the amino acid residues which separate the domains starting from the N-terminus left to C-terminus right. NTD = N-terminal domain, DBD = DNA binding domain. LBD = ligand binding domain. AF = activation function. Isoforms Two isoforms of the androgen receptor A and B have been identified:20 AR-A - 87 kDa - N-terminus truncated lacks the first 187 amino acids AR-B - 110 kDa - full length Domains Like other nuclear receptors, the androgen receptor is modular in structure and is comprised of the following functional domains labeled A through F:21 A/B - N-terminal regulatory domain contains:22 activation function 1 AF-1 between residues 101 and 370 required for full ligand activated transcriptional activity activation function 5 AF-5 between residues 360-485 is responsible for the constitutive activity activity without bound ligand dimerization surface involving residues 1-36 containing the FXXLF motif where F = phenylalanine, L = leucine, and X = any amino acid residue and 370-494 which both interact with the LBD in an intramolecular232425 head-to-tail interaction262728 C - DNA binding domain DBD D - Hinge region - flexible region that connects the DBD with the LBD; along with the DBD, contains a ligand dependent nuclear localization signal29 E - Ligand binding domain LBD containing activation function 2 AF-2, responsible for agonist induced activity activity in the presence of bound agonist AF-2 binds either the N-terminal FXXFL motif intramolecularly or coactivator proteins containing the LXXLL or preferably FXXFL motifs28 A ligand dependent nuclear export signal30 F - C-terminal domain References ^ Lu NZ, Wardell SE, Burnstein KL, Defranco D, Fuller PJ, Giguere V, Hochberg RB, McKay L, Renoir JM, Weigel NL, Wilson EM, McDonnell DP, Cidlowski JA December 2006. International Union of Pharmacology. LXV. The pharmacology and classification of the nuclear receptor superfamily: glucocorticoid, mineralocorticoid, progesterone, and androgen receptors. Pharmacol. Rev. 58 4: 782-97. doi:10.1124/pr.58.4.9. PMID 17132855. ^ Roy AK, Lavrovsky Y, Song CS, Chen S, Jung MH, Velu NK, Bi BY, Chatterjee B 1999. Regulation of androgen action. Vitam. Horm. 55: 309-52. PMID 9949684. ^ Bardin CW, Brown T, Isomaa VV, Jänne OA 1983. Progestins can mimic, inhibit and potentiate the actions of androgens. Pharmacol. Ther. 23 3: 443-59. PMID 6371845. ^ Raudrant D, Rabe T 2003. Progestogens with antiandrogenic properties. Drugs 63 5: 463-92. PMID 12600226. ^ Mooradian AD, Morley JE, Korenman SG 1987. Biological actions of androgens. Endocr. Rev. 8 1: 1-28. PMID 3549275. ^ a b Heinlein CA, Chang C 2002. The roles of androgen receptors and androgen-binding proteins in nongenomic androgen actions. Mol. Endocrinol. 16 10: 2181-7. doi:10.1210/me.2002-0070. PMID 12351684. ^ Davison SL, Bell R April 2006. Androgen physiology. Semin. Reprod. Med. 24 2: 71-7. doi:10.1055/s-2006-939565. PMID 16633980. ^ Sinisi AA, Pasquali D, Notaro A, Bellastella A 2003. Sexual differentiation. J. Endocrinol. Invest. 26 3 Suppl: 23-8. PMID 12834017. ^ Frank GR September 2003. Role of estrogen and androgen in pubertal skeletal physiology. Med. Pediatr. Oncol. 41 3: 217-21. doi:10.1002/mpo.10340. PMID 12868122. ^ Heemers HV, Tindall DJ December 2007. Androgen receptor AR coregulators: a diversity of functions converging on and regulating the AR transcriptional complex. Endocr. Rev. 28 7: 778-808. doi:10.1210/er.2007-0019. PMID 17940184. ^ Pandini G, Mineo R, Frasca F, Roberts CT Jr, Marcelli M, Vigneri R, Belfiore A March 2005. Androgens up-regulate the insulin-like growth factor-I receptor in prostate cancer cells. Cancer Res. 65 5: 1849-57. doi:10.1158/0008-5472.CAN-04-1837. PMID 15753383. ^ Fix C, Jordan C, Cano P, Walker WH 2004. Testosterone activates mitogen-activated protein kinase and the cAMP response element binding protein transcription factor in Sertoli cells. Proc Natl Acad Sci U S A 101 30: 10919-24. doi:10.1073/pnas.0404278101. PMID 15263086. ^ Vlahopoulos S, Zimmer WE, Jenster G, Belaguli NS, Balk SP, Brinkmann AO, Lanz RB, Zoumpourlis VC, Schwartz RJ 2005. Recruitment of the androgen receptor via serum response factor facilitates expression of a myogenic gene. J. Biol. Chem. 280 9: 7786-92. doi:10.1074/jbc.M413992200. PMID 15623502. ^ Chang CS, Kokontis J, Liao ST 1988. Molecular cloning of human and rat complementary DNA encoding androgen receptors. Science 240 4850: 324-6. doi:10.1126/science.3353726. PMID 3353726. ^ Trapman J, Klaassen P, Kuiper GG, van der Korput JA, Faber PW, van Rooij HC, Geurts van Kessel A, Voorhorst MM, Mulder E, Brinkmann AO 1988. Cloning, structure and expression of a cDNA encoding the human androgen receptor. Biochem. Biophys. Res. Commun. 153 1: 241-8. PMID 3377788. ^ Brown TR 1995. Human androgen insensitivity syndrome abstract. J. Androl. 16 4: 299-303. PMID 8537246. ^ Kennedy WR, Alter M, Sung JH 1968. Progressive proximal spinal and bulbar muscular atrophy of late onset. A sex-linked recessive trait. Neurology 18 7: 671-80. PMID 4233749. ^ Yu Z, Dadgar N, Albertelli M, Gruis K, Jordan C, Robins DM, Lieberman AP 2006. Androgen-dependent pathology demonstrates myopathic contribution to the Kennedy disease phenotype in a mouse knock-in model. J. Clin. Invest. 116 10: 2663-72. doi:10.1172/JCI28773. PMID 16981011. ^ Rajender S, Singh L, Thangaraj K 2007. Phenotypic heterogeneity of mutations in androgen receptor gene. Asian J. Androl. 9 2: 147-79. doi:10.1111/j.1745-7262.2007.00250.x. PMID 17334586. ^ Wilson CM, McPhaul MJ 1994. A and B forms of the androgen receptor are present in human genital skin fibroblasts. Proc. Natl. Acad. Sci. U.S.A. 91 4: 1234-8. doi:10.1073/pnas.91.4.1234. PMID 8108393. ^ Brinkmann AO, Klaasen P, Kuiper GG, van der Korput JA, Bolt J, de Boer W, Smit A, Faber PW, van Rooij HC, Geurts van Kessel A, Voorhorst MM, Mulder E, Trapman J 1989. Structure and function of the androgen receptor. Urol. Res. 17 2: 87-93. PMID 2734982. ^ Jenster G, van der Korput HA, Trapman J, Brinkmann AO 1995. Identification of two transcription activation units in the N-terminal domain of the human androgen receptor. J. Biol. Chem. 270 13: 7341-6. doi:10.1074/jbc.270.13.7341. PMID 7706276. ^ Schaufele F, Carbonell X, Guerbadot M, Borngraeber S, Chapman MS, Ma AA, Miner JN, Diamond MI July 2005. The structural basis of androgen receptor activation: intramolecular and intermolecular amino-carboxy interactions. Proc. Natl. Acad. Sci. U.S.A. 102 28: 9802-7. doi:10.1073/pnas.0408819102. PMID 15994236. ^ Klokk TI, Kurys P, Elbi C, Nagaich AK, Hendarwanto A, Slagsvold T, Chang CY, Hager GL, Saatcioglu F March 2007. Ligand-specific dynamics of the androgen receptor at its response element in living cells. Mol. Cell. Biol. 27 5: 1823-43. doi:10.1128/MCB.01297-06. PMID 17189428. ^ van Royen ME, Cunha SM, Brink MC, Mattern KA, Nigg AL, Dubbink HJ, Verschure PJ, Trapman J, Houtsmuller AB April 2007. Compartmentalization of androgen receptor protein-protein interactions in living cells. J. Cell Biol. 177 1: 63-72. doi:10.1083/jcb.200609178. PMID 17420290. ^ Langley E, Zhou ZX, Wilson EM 1995. Evidence for an anti-parallel orientation of the ligand-activated human androgen receptor dimer. J. Biol. Chem. 270 50: 29983-90. doi:10.1074/jbc.270.50.29983. PMID 8530400. ^ Berrevoets CA, Doesburg P, Steketee K, Trapman J, Brinkmann AO 1998. Functional interactions of the AF-2 activation domain core region of the human androgen receptor with the amino-terminal domain and with the transcriptional coactivator TIF2 transcriptional intermediary factor2. Mol. Endocrinol. 12 8: 1172-83. doi:10.1210/me.12.8.1172. PMID 9717843. ^ a b Dubbink HJ, Hersmus R, Verma CS, van der Korput HA, Berrevoets CA, van Tol J, Ziel-van der Made AC, Brinkmann AO, Pike AC, Trapman J 2004. Distinct recognition modes of FXXLF and LXXLL motifs by the androgen receptor. Mol. Endocrinol. 18 9: 2132-50. doi:10.1210/me.2003-0375. PMID 15178743. ^ Kaku N, Matsuda KI, Tsujimura A, Kawata M April 2008. Characterization of nuclear import of the domain-specific androgen receptor in association with the importin α/β and Ran-GTP systems. Endocrinology. doi:10.1210/en.2008-0137. PMID 18420738. ^ Saporita AJ, Zhang Q, Navai N, Dincer Z, Hahn J, Cai X, Wang Z October 2003. Identification and characterization of a ligand-regulated nuclear export signal in androgen receptor. J. Biol. Chem. 278 43: 41998-2005. doi:10.1074/jbc.M302460200. PMID 12923188. See also Androgen External links MeSH Androgen+Receptors Brinkmann AO. Androgen physiology: receptor and metabolic disorders, in Robert McLachlan, or: Endocrinology of Male Reproduction. Endotext.org. Retrieved on 2008-04-29. Gottlieb B 2007-07-24. The Androgen Receptor Gene Mutations Database Server. McGill University. Retrieved on 2008-04-29. Thompson J 2006-09-30. Molecular Mechanisms of Androgen Receptor Interactions. Helsinki University Biomedical Dissertations No. 80. University of Helsinki. Retrieved on 2008-04-29. v d e Transcription factors and intracellular receptors 1 Basic domains 1.1 Basic leucine zipper bZIP Activating transcription factor AATF, 1, 2, 3, 4, 5, 6, 7 AP-1 c-Fos, FOSB, FOSL1, FOSL2, c-Jun, JUNB, JUND BACH 1, 2 BATF BLZF1 C/EBP α, β, γ, δ, ε, ζ CREB 1, 3, L1 CREM DBP DDIT3 GABPA HLF MAF B, F, G, K NFE 2, L1, L2 NRL NRF1 XBP1 1.2 Basic helix-loop-helix bHLH ATOH1 AhR AHRR ARNT ASCL1 BHLHB2 BMAL ARNTL, ARNTL2 CLOCK EPAS1 HAND 1, 2 HES 5, 6 HEY 1, 2, L HES1 HIF 1A, 3A ID 1, 2, 3, 4 LYL1 MXD4 MYCL1 MYCN Myogenic regulatory factors MyoD, Myogenin, MYF5, MYF6 Neurogenins NeuroD 1, 2 NPAS 1, 2, 3 OLIG 1, 2 Scleraxis TAL1 Twist USF1 1.3 bHLH-ZIP AP-4 MAX MITF MNT MLX MXI1 Myc SREBP 1, 2 1.4 NF-1 NFIC 1.5 RF-X NFX1 1.6 Basic helix-span-helix bHSH AP-2 α, β, γ, δ, ε 2 Zinc finger DNA-binding domains 2.1 Nuclear receptor Cys4 subfamily 1 Thyroid hormone α, β, CAR, FXR, LXR α, β, PPAR α, β/δ, γ, PXR, RAR α, β, γ, ROR α, β, γ, Rev-ErbA α, β, VDR subfamily 2 COUP-TF I, II, Ear-2, HNF4 α, γ, PNR, RXR α, β, γ, Testicular receptor 2, 4, TLX subfamily 3 Steroid hormone Androgen, Estrogen α, β, Glucocorticoid, Mineralocorticoid, Progesterone, Estrogen related α, β, γ subfamily 4 NUR NGFIB, NOR1, NURR1 subfamily 5 LRH-1, SF1 subfamily 6 GCNF subfamily 0 DAX1, SHP 2.2 Other Cys4 GATA 1, 2, 3, 4, 5, 6 MTA 1, 2, 3 2.3 Cys2His2 General transcription factors TFIIA, TFIIB, TFIID, TFIIE, TFIIF 1, 2, TFIIH 1, 2, 4, 2I, 3A, 3C1, 3C2 ATBF1 BCL 6, 11A, 11B CTCF E4F1 EGR 2, 3 ERV3 GFI1 GLI-Krüppel family 1, 2, 3, YY1 HIC 1, 2 HIVEP 1, 2, 3 IKZF 1, 2, 3 ILF 2, 3 KLF 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 17 MTF1 MYT1 OSR1 SP 1, 2, 4, 7 Zbtb7 7A, 7B ZBTB 16, 17, 20, 32, 33, 40 zinc finger 3, 7, 9, 10, 19, 22, 24, 33B, 34, 35, 41, 43, 44, 51, 74, 143, 146, 148, 165, 202, 217, 219, 238, 239, 259, 267, 268, 281, 295, 318, 330, 346, 350, 365, 366, 384, 423, 451, 452, 471, 593, 638, 649, 655 2.4 Cys6 HIVEP1 2.5 Alternating composition AIRE DIDO1 GRLF1 ING 1, 2, 4 JARID 1A, 1B, 1C, 1D, 2 JMJD1B 3 Helix-turn-helix domains 3.1 Homeo domain ARX CDX 1, 2 CRX CUTL1 DLX 3, 4, 5 EMX2 EN 1, 2 FHL 1, 2, 3 HESX1 HHEX HLX Homeobox A1, A3, A4, A5, A7, A9, A10, A11, A13, B1, B2, B3, B4, B5, B6, B7, B8, B9, B13, C4, C5, C6, C8, C9, C10, C11, C13, D1, D3, D4, D8, D9, D10, D11, D12, D13 HOPX MEIS 1, 2 MEOX2 MNX1 MSX 1, 2 NANOG NKX 2-1, 2-2, 2-3, 2-5, 3-1, 3-2 PBX 1, 2, 3 PHF 1, 3, 6, 8, 10, 16, 17, 20, 21A PITX 1, 2, 3 POU domain PIT-1, BRN-3: A, B, C, Octamer transcription factor: 1, 2, 3/4, 6, 7, 11 OTX 1, 2 PDX1 3.2 Paired box PAX 1, 2, 3, 4, 5, 6, 7, 8, 9 3.3 Fork head / winged helix E2F 1, 2, 3, 4, 5 FOX proteins C1, C2, E1, G1, H1, K2, L2, M1, N3, O1A, O3A, O4, P1, P2, P3 3.4 Heat Shock Factors HSF 1, 2, 4 3.5 Tryptophan clusters ELF 4, 5 EGF ELK 1, 3, 4 ERF ERG ETS 1, 2, SPIB ETV 1, 4, 5, 6 FLI1 Interferon regulatory factors 1, 2, 3, 4, 5, 6, 7, 8 MYB MYBL2 3.6 TEA domain transcriptional enhancer factor 1, 2 4 β-Scaffold factors with minor groove contacts 4.1 Rel homology region NF-κB NFKB1, NFKB2, REL, RELA, RELB NFAT C1, C2, C3, C4, 5 4.2 STAT STAT 1, 2, 3, 4, 5, 6 4.3 p53 p53 TBX 1, 2, 3, 5, 19, 21, 22 4.4 MADS box Mef2 A, B, C, D SRF 4.6 TATA binding proteins TBP TBPL1 4.7 High mobility group HMGB 1, 2, 3 HNF 1A, 1B LEF1 SOX 2, 3, 4, 5, 6, 8, 9, 10, 11, 12, 13, 14, 15, 18, 21 SRY SSRP1 TOX 1, 2, 3, 4 4.10 Cold-shock domain CSDA, YBX1 4.11 Runt CBF CBFA2T2, CBFA2T3, RUNX1, RUNX2, RUNX3, RUNX1T1 0 Other transcription factors 0.2 HMGIY HMGA 1, 2 HBP1 0.3 Pocket domain Rb RBL1 RBL2 0.5 AP-2/EREBP-related factors Apetala 2 EREBP B3 0.6 Miscellaneous ARID 1A, 1B, 2, 3A, 3B, 4A CAP IFI 16, 35 MLL 2, 3, T1 MNDA NFI A, B, C, X NFY A, B, C Rho/Sigma R-SMAD v d e Sex hormones and related agents primarily G03, also L02, H01C - human endogenous in CAPS Progestogens: receptor Agonist PROGESTERONE, Dienogest, Desogestrel, Drospirenone, Dydrogesterone, Ethisterone, Etonogestrel, Ethynodiol diacetate, Gestodene, Gestonorone, Levonorgestrel, Lynestrenol, Medroxyprogesterone, Megestrol, Norelgestromin, Norethisterone, Norethynodrel, Norgestimate, Norgestrel, Norgestrienone, Tibolone SPRM Asoprisnil, CDB-4124 Antiprogestogen Mifepristone Androgens: receptor Agonist TESTOSTERONE, Androstanolone, Fluoxymesterone, Mesterolone, Methyltestosterone, see also anabolic steroids Antiandrogen Bicalutamide, Cyproterone, Dienogest, Flutamide, Nilutamide, Spironolactone Estrogens: receptor Agonist ESTRADIOL, ESTRIOL, ESTRONE, Chlorotrianisene, Dienestrol, Diethylstilbestrol, Ethinylestradiol, Fosfestrol, Mestranol, Polyestradiol phosphate SERM Afimoxifene, Arzoxifene, Bazedoxifene, Clomifene, Fulvestrant, Lasofoxifene, Raloxifene, Tamoxifen, Toremifene AI Aminoglutethimide, Anastrozole, Atamestane, Exemestane, Fadrozole, Formestane, Letrozole, Vorozole Antiestrogen Mepitiostane Gonadotropins: FSHR/LHCGR ovulation stim. Clomifene, Urofollitropin Antigonadotropin Danazol, Gestrinone GnRH: receptor Agonist Buserelin, Goserelin, Histrelin, Leuprorelin, Nafarelin, Triptorelin Antagonist Abarelix, Cetrorelix, Ganirelix Retrieved from http://en..org/wiki/Androgen_receptor Categories: Genes on chromosome X | Human proteins | Intracellular receptors | Transcription factors Views Article Discussion this page History Personal tools Log in / create account Navigation Main page Contents Featured content Current events Random article Search Go Search Interaction Community portal Recent changes Contact Donate to Help Toolbox What links here Related changes Upload file Special pages Printable version Permanent link Cite this page Languages Español Français This page was last modified on 28 July 2008, at 19:39
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