{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,6]],"date-time":"2026-03-06T07:22:40Z","timestamp":1772781760218,"version":"3.50.1"},"reference-count":36,"publisher":"Proceedings of the National Academy of Sciences","issue":"1","content-domain":{"domain":["www.pnas.org"],"crossmark-restriction":true},"short-container-title":["Proc. Natl. Acad. Sci. U.S.A."],"published-print":{"date-parts":[[2002,1,8]]},"abstract":"<jats:p>\n                    Associations have been reported of the seven-repeat\n (7R) allele of the human dopamine receptor D4 (\n                    <jats:italic>DRD4<\/jats:italic>\n                    ) gene\n with both attention-deficit\/hyperactivity disorder and the\n personality trait of novelty seeking. This polymorphism occurs in a\n 48-bp tandem repeat in the coding region of\n                    <jats:italic>DRD4<\/jats:italic>\n                    ,\n with the most common allele containing four repeats (4R) and\n rarer variants containing 2\u201311. Here we show by DNA\n resequencing\/haplotyping of 600\n                    <jats:italic>DRD4<\/jats:italic>\n                    alleles,\n representing a worldwide population sample, that the origin of 2R\u20136R\n alleles can be explained by simple one-step recombination\/mutation\n events. In contrast, the 7R allele is not simply related to the other\n common alleles, differing by greater than six\n recombinations\/mutations. Strong linkage disequilibrium was found\n between the 7R allele and surrounding\n                    <jats:italic>DRD4<\/jats:italic>\n                    polymorphisms,\n suggesting that this allele is at least 5\u201310-fold \u201cyounger\u201d than\n the common 4R allele. Based on an observed bias toward nonsynonymous\n amino acid changes, the unusual DNA sequence organization, and the\n strong linkage disequilibrium surrounding the\n                    <jats:italic>DRD4<\/jats:italic>\n                    7R allele, we propose that this allele originated as a rare\n mutational event that nevertheless increased to high frequency in human\n populations by positive selection.\n                  <\/jats:p>","DOI":"10.1073\/pnas.012464099","type":"journal-article","created":{"date-parts":[[2002,7,26]],"date-time":"2002-07-26T10:36:44Z","timestamp":1027679804000},"page":"309-314","update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":383,"title":["Evidence of positive selection acting at the human dopamine receptor D4 gene locus"],"prefix":"10.1073","volume":"99","author":[{"given":"Yuan-Chun","family":"Ding","sequence":"first","affiliation":[{"name":"Department of Biological Chemistry, College of Medicine,\r University of California, Irvine, CA 92697; Laboratory of\r Cellular and Molecular Evolution, Kunming Institute of Zoology, Chinese\r Academy of Sciences, Kunming 650223, China; Department\r of Genetics, School of Medicine, Yale University, New Haven, CT\r 06520; Department of Pediatrics, Medical Center,\r University of California, Orange, CA 92868; and Child\r Development Center, University of California, Irvine, CA 92715"}]},{"given":"Han-Chang","family":"Chi","sequence":"additional","affiliation":[{"name":"Department of Biological Chemistry, College of Medicine,\r University of California, Irvine, CA 92697; Laboratory of\r Cellular and Molecular Evolution, Kunming Institute of Zoology, Chinese\r Academy of Sciences, Kunming 650223, China; Department\r of Genetics, School of Medicine, Yale University, New Haven, CT\r 06520; Department of Pediatrics, Medical Center,\r University of California, Orange, CA 92868; and Child\r Development Center, University of California, Irvine, CA 92715"}]},{"given":"Deborah L.","family":"Grady","sequence":"additional","affiliation":[{"name":"Department of Biological Chemistry, College of Medicine,\r University of California, Irvine, CA 92697; Laboratory of\r Cellular and Molecular Evolution, Kunming Institute of Zoology, Chinese\r Academy of Sciences, Kunming 650223, China; Department\r of Genetics, School of Medicine, Yale University, New Haven, CT\r 06520; Department of Pediatrics, Medical Center,\r University of California, Orange, CA 92868; and Child\r Development Center, University of California, Irvine, CA 92715"}]},{"given":"Atsuyuki","family":"Morishima","sequence":"additional","affiliation":[{"name":"Department of Biological Chemistry, College of Medicine,\r University of California, Irvine, CA 92697; Laboratory of\r Cellular and Molecular Evolution, Kunming Institute of Zoology, Chinese\r Academy of Sciences, Kunming 650223, China; Department\r of Genetics, School of Medicine, Yale University, New Haven, CT\r 06520; Department of Pediatrics, Medical Center,\r University of California, Orange, CA 92868; and Child\r Development Center, University of California, Irvine, CA 92715"}]},{"given":"Judith R.","family":"Kidd","sequence":"additional","affiliation":[{"name":"Department of Biological Chemistry, College of Medicine,\r University of California, Irvine, CA 92697; Laboratory of\r Cellular and Molecular Evolution, Kunming Institute of Zoology, Chinese\r Academy of Sciences, Kunming 650223, China; Department\r of Genetics, School of Medicine, Yale University, New Haven, CT\r 06520; Department of Pediatrics, Medical Center,\r University of California, Orange, CA 92868; and Child\r Development Center, University of California, Irvine, CA 92715"}]},{"given":"Kenneth K.","family":"Kidd","sequence":"additional","affiliation":[{"name":"Department of Biological Chemistry, College of Medicine,\r University of California, Irvine, CA 92697; Laboratory of\r Cellular and Molecular Evolution, Kunming Institute of Zoology, Chinese\r Academy of Sciences, Kunming 650223, China; Department\r of Genetics, School of Medicine, Yale University, New Haven, CT\r 06520; Department of Pediatrics, Medical Center,\r University of California, Orange, CA 92868; and Child\r Development Center, University of California, Irvine, CA 92715"}]},{"given":"Pamela","family":"Flodman","sequence":"additional","affiliation":[{"name":"Department of Biological Chemistry, College of Medicine,\r University of California, Irvine, CA 92697; Laboratory of\r Cellular and Molecular Evolution, Kunming Institute of Zoology, Chinese\r Academy of Sciences, Kunming 650223, China; Department\r of Genetics, School of Medicine, Yale University, New Haven, CT\r 06520; Department of Pediatrics, Medical Center,\r University of California, Orange, CA 92868; and Child\r Development Center, University of California, Irvine, CA 92715"}]},{"given":"M. Anne","family":"Spence","sequence":"additional","affiliation":[{"name":"Department of Biological Chemistry, College of Medicine,\r University of California, Irvine, CA 92697; Laboratory of\r Cellular and Molecular Evolution, Kunming Institute of Zoology, Chinese\r Academy of Sciences, Kunming 650223, China; Department\r of Genetics, School of Medicine, Yale University, New Haven, CT\r 06520; Department of Pediatrics, Medical Center,\r University of California, Orange, CA 92868; and Child\r Development Center, University of California, Irvine, CA 92715"}]},{"given":"Sabrina","family":"Schuck","sequence":"additional","affiliation":[{"name":"Department of Biological Chemistry, College of Medicine,\r University of California, Irvine, CA 92697; Laboratory of\r Cellular and Molecular Evolution, Kunming Institute of Zoology, Chinese\r Academy of Sciences, Kunming 650223, China; Department\r of Genetics, School of Medicine, Yale University, New Haven, CT\r 06520; Department of Pediatrics, Medical Center,\r University of California, Orange, CA 92868; and Child\r Development Center, University of California, Irvine, CA 92715"}]},{"given":"James M.","family":"Swanson","sequence":"additional","affiliation":[{"name":"Department of Biological Chemistry, College of Medicine,\r University of California, Irvine, CA 92697; Laboratory of\r Cellular and Molecular Evolution, Kunming Institute of Zoology, Chinese\r Academy of Sciences, Kunming 650223, China; Department\r of Genetics, School of Medicine, Yale University, New Haven, CT\r 06520; Department of Pediatrics, Medical Center,\r University of California, Orange, CA 92868; and Child\r Development Center, University of California, Irvine, CA 92715"}]},{"given":"Ya-Ping","family":"Zhang","sequence":"additional","affiliation":[{"name":"Department of Biological Chemistry, College of Medicine,\r University of California, Irvine, CA 92697; Laboratory of\r Cellular and Molecular Evolution, Kunming Institute of Zoology, Chinese\r Academy of Sciences, Kunming 650223, China; Department\r of Genetics, School of Medicine, Yale University, New Haven, CT\r 06520; Department of Pediatrics, Medical Center,\r University of California, Orange, CA 92868; and Child\r Development Center, University of California, Irvine, CA 92715"}]},{"given":"Robert K.","family":"Moyzis","sequence":"additional","affiliation":[{"name":"Department of Biological Chemistry, College of Medicine,\r University of California, Irvine, CA 92697; Laboratory of\r Cellular and Molecular Evolution, Kunming Institute of Zoology, Chinese\r Academy of Sciences, Kunming 650223, China; Department\r of Genetics, School of Medicine, Yale University, New Haven, CT\r 06520; Department of Pediatrics, Medical Center,\r University of California, Orange, CA 92868; and Child\r Development Center, University of California, Irvine, CA 92715"}]}],"member":"341","published-online":{"date-parts":[[2001,12,26]]},"reference":[{"key":"e_1_3_4_1_2","doi-asserted-by":"crossref","first-page":"610","DOI":"10.1038\/350610a0","volume":"350","author":"Van Tol H H M","year":"1991","unstructured":"H H M Van Tol, J R Bunzow, H-C Guan, R K Sunahara, P Seeman, H B Niznik, O Civelli Nature (London) 350, 610\u2013614 (1991).","journal-title":"Nature (London)"},{"key":"e_1_3_4_2_2","doi-asserted-by":"crossref","first-page":"767","DOI":"10.1093\/hmg\/2.6.767","volume":"2","author":"Lichter J B","year":"1993","unstructured":"J B Lichter, C L Barr, J L Kennedy, H H M Van Tol, K K Kidd, K J Livak Hum Mol Genet 2, 767\u2013773 (1993).","journal-title":"Hum Mol Genet"},{"key":"e_1_3_4_3_2","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1007\/s004390050166","volume":"98","author":"Chang F-M","year":"1996","unstructured":"F-M Chang, J R Kidd, K J Livak, A J Pakstis, K K Kidd Hum Genet 98, 91\u2013101 (1996).","journal-title":"Hum Genet"},{"key":"e_1_3_4_4_2","first-page":"21","volume":"1","author":"La Hoste G J","year":"1996","unstructured":"G J La Hoste, J M Swanson, S B Wigal, C Glabe, T Wigal, N King, J L Kennedy Mol Psychiatry 1, 21\u201324 (1996).","journal-title":"Mol Psychiatry"},{"key":"e_1_3_4_5_2","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1016\/S0149-7634(99)00062-7","volume":"24","author":"Swanson J M","year":"2000","unstructured":"J M Swanson, P Flodman, J Kennedy, M A Spence, R Moyzis, S Schuck, M Murias, J Moriarity, C Barr, M Smith, M Posner Neurosci Biobehav Rev 24, 21\u201325 (2000).","journal-title":"Neurosci Biobehav Rev"},{"key":"e_1_3_4_6_2","doi-asserted-by":"crossref","first-page":"4754","DOI":"10.1073\/pnas.080070897","volume":"97","author":"Swanson J","year":"2000","unstructured":"J Swanson, J Oosterlaan, M Murias, S Schuck, Flodman, M A Spence, M Wasdell, Y Ding, H-C Chi, M Smith, et al. Proc Natl Acad Sci USA 97, 4754\u20134759, . (First Published April 18, 2000; 10.1073\/pnas.080070897). (2000).","journal-title":"Proc Natl Acad Sci USA"},{"key":"e_1_3_4_7_2","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1038\/ng0196-78","volume":"12","author":"Ebstein R P","year":"1996","unstructured":"R P Ebstein, O Novick, R Umansky, B Priel, Y Osher, D Blaine, E R Bennett, L Nemanov, M Katz, R H Belmaker Nat Genet 12, 78\u201380 (1996).","journal-title":"Nat Genet"},{"key":"e_1_3_4_8_2","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1038\/ng0196-81","volume":"12","author":"Benjamin J","year":"1996","unstructured":"J Benjamin, L Li, C Patterson, B D Greenberg, D L Murphy, D H Hamer Nat Genet 12, 81\u201384 (1996).","journal-title":"Nat Genet"},{"key":"e_1_3_4_9_2","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1016\/S1566-2772(01)00007-X","volume":"1","author":"Swanson J","year":"2001","unstructured":"J Swanson, C Deutsch, D Cantwell, M Posner, J Kennedy, C Barr, R Moyzis, S Schuck, P Flodman, M A Spence Clin Neurosci Res 1, 207\u2013216 (2001).","journal-title":"Clin Neurosci Res"},{"key":"e_1_3_4_10_2","volume-title":"DSM-IV: Diagnostic and Statistical Manual of Mental Disorders","year":"1994","unstructured":"DSM-IV: Diagnostic and Statistical Manual of Mental Disorders (Am. 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