110:(1995), conducted by the Hamer lab in collaboration with two groups of statistical experts in 1995, corroborated the original results for males with homosexual brothers sharing Xq28 at significantly elevated rates. This study also included heterosexual brothers, who showed significantly less than expected sharing of the Xq28 region, as expected for a genetic locus that in one form is associated with same-sex attraction and in another form is associated with opposite-sex attraction. In this study no link to Xq28 was found among homosexual females, indicating a different genetic pathway as for most sex-specific phenotypes.
242:(the researchers depended on their own judgement and sometimes based their judgement on a single question to the subject). They also lacked criteria "to select appropriate families for the study of a putative X-linked locus" — as they did not select families based on the presence of maternal transmission of homosexuality, the Xq28 contribution to male sexual orientation may have been hidden. In addition, the meta-analysis revealed that the family pedigree data of Rice
266:, overlapping with one of the regions detected in a previous genomewide linkage study by the Hamer lab. The authors concluded that "our findings, taken in context with previous work, suggest that genetic variation in each of these regions contributes to development of the important psychological trait of male sexual orientation." It was the largest study of the genetic basis of homosexuality to date and was published online in November 2014.
20:
155:. The junior researcher had assisted in the gene mapping in Hamer's 1993 study. Shortly after voicing her questions, she was summarily dismissed from her post-doctoral fellowship in Hamer's lab; who dismissed her could not be determined. Later, she was given another position in a different lab. Hamer stated that Crewdson's article was "seriously in error" and denied the allegations made against him. An official inquiry launched by the
404:
did not include the family pedigree data in their April 1999 report, which described only the genotyping results for a subset of 48 families. The realization that the family pedigree data actually supported X chromosome linkage came later when the data was analysed during the meta-analysis published
102:
of the X chromosome were analyzed for 40 families to see if a specific marker was shared by a disproportionate amount of brothers who were both gay. The results showed that among gay brothers, the concordance rate for markers from the Xq28 region were significantly greater than expected for random
273:
of 493,001 individuals concluded that hundreds or thousands of genetic variants underlie homosexual behavior in both sexes, with 5 variants in particular being significantly associated. They stated that in contrast to linkage studies that found substantial association of sexual orientation with
1193:
Zietsch, Brendan P.; Neale, Benjamin M.; Perry, John R. B.; Sanders, Alan R.; Martin, Eden R.; Beecham, Gary W.; Harris, Kathleen Mullan; Auton, Adam; Långström, Niklas; Lundström, Sebastian; Lichtenstein, Paul; Team16, Paul; Sathirapongsasuti, J. Fah; Guo, Shengru; Abdellaoui, Abdel; Busch,
199:. Hamer criticized the study for not selecting families for their study population based on maternal transmission as selecting only families that show an excess of maternal gay relatives is necessary to detect the Xq28 linkage. A meta-analysis of all data available at that time (i.e., Hamer
191:. Consequently, they concluded against the possibility of any gene in the Xq28 region having a large genetic influence on male sexual orientation (though they could not rule out the possibility of a gene in this region having a small influence). Rice
1135:
Sanders AR, Martin ER, Beecham GW, Guo S, Dawood K, Rieger G, Badner JA, Gershon ES, Krishnappa RS, Kolundzija AB, Duan J, Gejman PV, Bailey JM (November 2014). "Genome-wide scan demonstrates significant linkage for male sexual orientation".
103:
Mendelian segregation, indicating that a link did exist in that small sample. It was concluded that at least one form of male homosexuality is preferentially transmitted through the maternal side and is genetically linked to the Xq28 region.
93:
men in the United States and found increased rates of homosexuality among maternal uncles and cousins, but not among paternal relatives. This pattern of inheritance suggested that there might be linked genes on the
427:"Localization of loci for hypoxanthine phosphoribosyltransferase and glucose-6-phosphate dehydrogenase and biochemical evidence of nonrandom X chromosome expression from studies of a human X-autosome translocation"
253:
of male sexual orientation was conducted by several independent groups of researchers. The study population included 409 independent pairs of gay brothers from 384 families, who were analyzed with over 300,000
1331:
159:(ORI) to investigate the allegations of selective presentation of the data ended in December 1996. It determined that Hamer had not committed any scientific misconduct in his study.
258:
markers. The study confirmed the Xq28 linkage to homosexuality by two-point and multipoint (MERLIN) LOD score mapping. Significant linkage was also detected in the region near the
167:
Two further studies in the 1990s gave mixed results. One was an X chromosome linkage analysis of 54 pairs of gay brothers carried out by the independent research group of
Sanders
175:: both reported that the chromosomal location of maximum sharing was locus DXS1108 and both reported similar degrees of allele sharing (66% versus 67%). The second study by Rice
274:
variants on the X-chromosome, they found no excess of signal (and no individual genome-wide significant variants) on Xq28 or the rest of the X chromosome.
234:
were non-representative as they had an excess of paternal instead of maternal gay relatives thus obscuring the display of any X-chromosome linkage; the
1180:
Zeroing in on the “Gay Gene” The largest study yet of the genetic roots of homosexuality links sexual preference in men to two regions of the genome
149:
stated that an anonymous junior researcher in Hamer's laboratory alleged that Hamer selectively presented the data in his 1993 paper in the journal
366:, their linkage analysis needed to find that gay brothers share more than 50% of their alleles at the Xq28 region. In contrast, analyses by Hamer
1399:
Rogner, UC; Wilke, K; Steck, E; Korn, B; Poustka, A (1995). "The melanoma antigen gene (MAGE) family is clustered in the chromosomal band Xq28".
1251:
601:
Hu, Stella; Pattatucci, Angela M. L.; Patterson, Chavis; Li, Lin; Fulker, David W.; Cherny, Stacey S.; Kruglyak, Leonid; Hamer, Dean H. (1995).
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226:(1999) may have been unable to detect statistically significant linkage between Xq28 and male sexual orientation: the families
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Rice G; Anderson C; Risch N; Ebers G (Apr 1999). "Male homosexuality: absence of linkage to microsatellite markers at Xq28".
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with Xq28 (green) located at the tip of the long arm of the chromosome. Studies of the X chromosome as well as the entire
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of their sample was insufficient to adequately detect linkage and they lacked definite criteria for what constituted as
1352:
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Alexander S.; Wedow, Robbee; Maier, Robert; Nivard, Michel G.; Verweij, Karin J. H.; Ganna, Andrea (30 August 2019).
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also asserted that their results do not exclude the possibility of finding male homosexuality genes elsewhere in the
358:
report that gay brothers shared approximately 46% of their alleles at the Xq28 region. However, this result was not
1485:"Crh and Oprm1 mediate anxiety-related behavior and social approach in a mouse model of MECP2 duplication syndrome"
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did find greater than 50% allele sharing at Xq28 in gay brothers, thus yielding statistically significant results.
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which has been studied since at least 1980. The band contains three distinct regions, totaling about 8
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De Sario, Albertina; Geigl, Eva-Maria; Palmieri, Giuseppe; d'Urso, Michele; Bernardi, Giorgio (1996).
246:(1999), in contrast to the genotyping data, seemed to support X chromosome linkage for homosexuality.
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98:, since males always inherit their copy of the X chromosome from their mothers. Polymorphisms of
783:
Hamer, Dean; Simon LeVay (May 1994). "Evidence for a
Biological Influence in Male Homosexuality".
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Xq28 is a large, complex, and gene-dense region. Among its various genes are the 12 genes of the
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1196:"Large-scale GWAS reveals insights into the genetic architecture of same-sex sexual behavior"
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in 1998. The results of the study were indistinguishable from the results of the study by Hu
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studied 52 pairs of
Canadian gay brothers and found no statistically significant linkage in
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study, Rice, Risch and Ebers) presented several methodological reasons due to which Rice
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indicated that Xq28 has a significant but not exclusive role in male sexual orientation.
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because to show that male sexual orientation is influenced by a gene (or genes) at Xq28
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genes within Xq28 have been associated with phenotypes including anxiety and autism in
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603:"Linkage between sexual orientation and chromosome Xq28 in males but not in females"
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Samaco, RC; Mandel-Brehm, C; McGraw, CM; Shaw, CA; McGill, BE; Zoghbi, HY (2012).
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556:"A linkage between DNA markers on the X chromosome and male sexual orientation"
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Proceedings of the
National Academy of Sciences of the United States of America
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Proceedings of the
National Academy of Sciences of the United States of America
47:
1436:"Androgen receptor molecular biology and potential targets in prostate cancer"
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Ngun TC; Vilain E (2014). "The
Biological Basis of Human Sexual Orientation".
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828:"Study On 'Gay Gene' Challenged: Author Defends Findings Against Allegations"
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1281:"No 'gay gene': Massive study homes in on genetic basis of human sexuality"
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The authors of the meta-analysis (which included three authors of the Rice
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Wickelgren, Ingrid (23 April 1999). "Discovery of 'Gay Gene' Questioned".
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Epigenetic
Shaping of Sociosexual Interactions - from Plants to Humans
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would not detect Xq28's linkage with homosexuality simply by chance.
19:
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Hamer's findings were highlighted in scientific journals including
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Wickelgren I (23 Apr 1999). "Discovery of 'Gay Gene' Questioned".
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Byne, William (May 1994). "The
Biological Evidence Challenged".
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184:
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Hamer, D.; Hu, S; Magnuson, V.; Hu, N; Pattatucci, A. (1993).
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Pai, GS; Sprenkle, JA; Do, TT; Mareni, CE; Migeon, BR (1980).
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Kaiser J (28 Feb 1997). "No
Misconduct in 'Gay Gene' Study".
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of genetic information. The marker came to the public eye in
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1339:. Advances in Genetics. Vol. 86. pp. 167–84.
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Pool, R (Jul 1993). "Evidence for homosexuality gene".
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Genetic marker at the lower tip of human X chromosome
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486:"A Compositional Map of Human Chromosome Band Xq28"
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211:(1999), and the unpublished 1998 study by Sanders
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1178:Bob Grant for The Scientist. November 19, 2014
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693:"Human genetics. Sexual orientation and the X"
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1535:Molecular Genetic Study of Sexual Orientation
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290:has been identified as a coregulator for the
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1266:. geneticsexbehavior.info. 28 February 2018
1250:: CS1 maint: numeric names: authors list (
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1013:"Genetics and Male Sexual Orientation"
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374:(1995) and the 1998 study by Sanders
364:in a statistically significant manner
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125:and the topic of a mini-symposium in
1279:Lambert, Jonathan (29 August 2019).
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31:have linked Xq28 to the heredity of
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805:10.1038/scientificamerican0594-44
762:10.1038/scientificamerican0594-50
387:There was a 35% chance that Rice
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70:between the Xq28 marker and male
249:In 2012, a large, comprehensive
139:In June 1994, an article in the
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1440:Therapeutic Advances in Urology
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826:Crewdson, John (25 June 1995).
325:Daily Mail#Gay gene controversy
1558:Sexual orientation and science
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320:Biology and sexual orientation
256:single-nucleotide polymorphism
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1260:"Genetics of Sexual Behavior"
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350:The linkage analysis by Rice
271:genome-wide association study
1114:10.1126/science.284.5414.665
1063:10.1126/science.284.5414.571
959:10.1126/science.284.5414.571
157:Office of Research Integrity
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1563:Genes on human chromosome X
1264:Genetics of Sexual Behavior
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284:melanoma-associated antigen
50:situated at the tip of the
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1306:10.1038/d41586-019-02585-6
1011:Hamer, D. H (1999-08-06).
1150:10.1017/S0033291714002451
360:statistically significant
251:genome-wide linkage study
89:examined 114 families of
1452:10.1177/1756287210372380
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286:(MAGE) family, of which
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670:10.1126/science.8332894
580:10.1126/science.8332896
66:and others indicated a
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1138:Psychological Medicine
511:10.1073/pnas.93.3.1298
452:10.1073/pnas.77.5.2810
106:A follow-up study, Hu
36:
907:Finn R (8 Jan 1996).
331:The Science of Desire
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691:King MC (Jul 1993).
1434:Wilson, EM (2010).
1297:2019Natur.573...14L
1106:1999Sci...284..665R
834:, Washington Bureau
797:1994SciAm.270e..44L
785:Scientific American
754:1994SciAm.270e..50B
742:Scientific American
709:1993Natur.364..288K
662:1993Sci...261..291P
572:1993Sci...261..321H
502:1996PNAS...93.1298D
443:1980PNAS...77.2810P
128:Scientific American
1540:2010-03-02 at the
1206:(6456): eaat7693.
1023:(5429): 803a–803.
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269:In August 2019, a
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370:(1993), Hu
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260:centromere
189:haplotypes
64:Dean Hamer
23:The human
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925:0890-3670
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