Skip to main content
SpringerLink
Log in

Organizational Effects of Gonadal Hormones on Human Sexual Orientation

  • Research
  • Published:
Adaptive Human Behavior and Physiology Aims and scope Submit manuscript

Abstract

Purpose

Sexual attraction to males or females is perhaps the largest behavioral sex difference across animal species. Experiments in laboratory mammals show that prenatal androgens mediate this sex difference, but ethical considerations preclude such experimentation in humans. Multiple lines of converging correlational evidence are therefore needed to demonstrate such mediation in humans.

Methods

We review available data linking human sexual orientation to endocrine action, including research on endocrine disorders and biomarkers of early sex hormones. We also perform a meta-analysis across 13 studies comprising 56,804 individuals to investigate a possible link between non-heterosexuality and polycystic ovary syndrome (PCOS), an endocrine condition associated with elevated androgens in females.

Results and conclusions

We find converging evidence that prenatal gonadal hormones influence the development of human sexual orientation and orchestrate its sexual differentiation primarily by regulating patterns of gene expression in the developing brain. Evidence is particularly strong that androgens increase sexual attraction to females. In our meta-analysis, PCOS was more common in non-heterosexual females (r = 0.18, p < 0.001). Some evidence also indicates that estrogens increase sexual attraction to males. We discuss why data may be less clear regarding variation in sexual orientation among males, including the possible existence of subgroups characterized by distinct biological pathways that contribute to same-sex sexual orientation. Moving forward, we propose that multiple measures and/or markers be considered together to better characterize early hormonal action on human sexual orientation.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Log in via an institution

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Institutional subscriptions

Fig. 1
Fig. 2
Fig. 3
Fig. 4
Fig. 5

Similar content being viewed by others

Data Availability

Data for meta-analysis are available at https://osf.io/fzeu7/.

References

  • Abbott, D. H., Barnett, D. K., Bruns, C. M., & Dumesic, D. A. (2005). Androgen excess fetal programming of female reproduction: A developmental aetiology for polycystic ovary syndrome? Human Reproduction Update, 11(4), 357–374. https://doi.org/10.1093/humupd/dmi013.

    Article  Google Scholar 

  • Agate, R. J., Grisham, W., Wade, J., Mann, S., Wingfield, J., Schanen, C., & Arnold, A. P. (2003). Neural, not gonadal, origin of brain sex differences in a gynandromorphic finch. Proceedings of the National Academy of Sciences, 100(8), 4873–4878.

  • Aguilar, R. (2023). Finger-length ratios and sexual orientation in Southern Spain: A large sample study on sex-linked personality and cognitive traits. Archives of Sexual Behavior. https://doi.org/10.1007/s10508-023-02670-y.

    Article  Google Scholar 

  • Arnold, A. P. (2004). Sex chromosomes and brain gender. Nature Reviews Neuroscience, 5(9), 701–708.

    Article  Google Scholar 

  • Arnold, A. P. (2012). The end of gonad-centric sex determination in mammals. Trends in Genetics, 28(2), 55–61. https://doi.org/10.1016/j.tig.2011.10.004.

    Article  Google Scholar 

  • Arnold, A. P., & Chen, X. (2009). What does the four core genotypes mouse model tell us about sex differences in the brain and other tissues? Frontiers in Neuroendocrinology, 30(1), 1–9.

    Article  Google Scholar 

  • Bailey, J. M., Dunne, M. P., & Martin, N. G. (2000). Genetic and environmental influences on sexual orientation and its correlates in an australian twin sample. J Pers Soc Psychol, 78(3), 524–536. https://doi.org/10.1037/0022-3514.78.3.524.

    Article  Google Scholar 

  • Bailey, J. M., Vasey, P. L., Diamond, L. M., Breedlove, S. M., Vilain, E., & Epprecht, M. (2016). Sexual orientation, controversy, and science. Psychological Science in the Public Interest, 17(2), 45–101.

    Article  Google Scholar 

  • Bakker, J., & Baum, M. J. (2008). Role for estradiol in female-typical brain and behavioral sexual differentiation. Frontiers in Neuroendocrinology, 29(1), 1–16. https://doi.org/10.1016/j.yfrne.2007.06.001.

    Article  Google Scholar 

  • Bakker, J., De Mees, C., Douhard, Q., Balthazart, J., Gabant, P., Szpirer, J., & Szpirer, C. (2006). Alpha-fetoprotein protects the developing female mouse brain from masculinization and defeminization by estrogens. Nature Neuroscience, 9(2), 220–226.

    Article  Google Scholar 

  • Balasubramanian, R., & Crowley, W. F. (2011). Isolated GnRH deficiency: A disease model serving as a unique prism into the systems biology of the GnRH neuronal network. Molecular and Cellular Endocrinology, 346(1), 4–12. https://doi.org/10.1016/j.mce.2011.07.012.

    Article  Google Scholar 

  • Balthazart, J. (2011). Minireview: Hormones and human sexual orientation. Endocrinology, 152(8), 2937–2947.

    Article  Google Scholar 

  • Balthazart, J. (2020). Sexual partner preference in animals and humans. Neuroscience & Biobehavioral Reviews, 115, 34–47. https://doi.org/10.1016/j.neubiorev.2020.03.024.

    Article  Google Scholar 

  • Balthazart, J., & Court, L. (2017). Human sexual orientation: The importance of evidentiary convergence. Archives of Sexual Behavior, 46(6), 1595–1600.

    Article  Google Scholar 

  • Barahal, H. S. (1940). Testosterone in psychotic male homosexuals. Psychiatric Quarterly.

  • Barry, J. A., Kay, A. R., Navaratnarajah, R., Iqbal, S., Bamfo, J. E. A. K., David, A. L., & Hardiman, P. J. (2010). Umbilical vein testosterone in female infants born to mothers with polycystic ovary syndrome is elevated to male levels. Journal of Obstetrics and Gynaecology, 30(5), 444–446. https://doi.org/10.3109/01443615.2010.485254.

    Article  Google Scholar 

  • Blanchard, R. (2001). Fraternal birth order and the maternal immune hypothesis of male homosexuality. Hormones and Behavior, 40(2), 105–114. https://doi.org/10.1006/hbeh.2001.1681.

    Article  Google Scholar 

  • Bogaert, A. F. (2006). Biological versus nonbiological older brothers and men’s sexual orientation. Proc Natl Acad Sci U S A, 103(28), 10771–10774. https://doi.org/10.1073/pnas.0511152103.

    Article  Google Scholar 

  • Bogaert, A. F. (2010). Physical development and sexual orientation in men and women: An analysis of NATSAL-2000. Archives of Sexual Behavior, 39(1), 110–116. https://doi.org/10.1007/s10508-008-9398-x.

    Article  Google Scholar 

  • Bogaert, A. F., & Hershberger, S. (1999). The relation between sexual orientation and penile size. Archives of Sexual Behavior, 28(3), 213–221. https://doi.org/10.1023/A:1018780108597.

    Article  Google Scholar 

  • Bogaert, A. F., Skorska, M. N., Wang, C., Gabrie, J., MacNeil, A. J., Hoffarth, M. R., & Blanchard, R. (2018). Male homosexuality and maternal immune responsivity to the Y-linked protein NLGN4Y. Proceedings of the National Academy of Sciences, 115(2), 302–306.

  • Bouvattier, C., Mignot, B., Lefèvre, H., Morel, Y., & Bougnères, P. (2006). Impaired sexual activity in male adults with partial androgen insensitivity. The Journal of Clinical Endocrinology & Metabolism, 91(9), 3310–3315.

    Article  Google Scholar 

  • Boyd, R., Richerson, P. J., & Henrich, J. (2011). The cultural niche: Why social learning is essential for human adaptation. Proceedings of the National Academy of Sciences, 108(supplement_2), 10918–10925.

    Article  Google Scholar 

  • Braunstein, G. D., & Hershman, J. M. (1976). Comparison of serum pituitary thyrotropin and chorionic gonadotropin concentrations throughout pregnancy. The Journal of Clinical Endocrinology & Metabolism, 42(6), 1123–1126.

    Article  Google Scholar 

  • Breedlove, S. M. (2010). Minireview: Organizational hypothesis: Instances of the fingerpost. Endocrinology, 151(9), 4116–4122.

    Article  Google Scholar 

  • Breedlove, S. M. (2017). Prenatal influences on human sexual orientation: Expectations versus data. Archives of Sexual Behavior, 46, 1583–1592.

    Article  Google Scholar 

  • Bronstein, J., Tawdekar, S., Liu, Y., Pawelczak, M., David, R., & Shah, B. (2011). Age of onset of polycystic ovarian syndrome in girls may be earlier than previously thought. Journal of Pediatric and Adolescent Gynecology, 24(1), 15–20. https://doi.org/10.1016/j.jpag.2010.06.003.

    Article  Google Scholar 

  • Brunner, F., Fliegner, M., Krupp, K., Rall, K., Brucker, S., & Richter-Appelt, H. (2016). Gender role, gender identity and sexual orientation in CAIS (XY-Women) compared with subfertile and infertile 46,XX women. The Journal of Sex Research, 53(1), 109–124. https://doi.org/10.1080/00224499.2014.1002124.

    Article  Google Scholar 

  • Bulun, S. E. (2016). Chapter 17 - Physiology and Pathology of the Female Reproductive Axis. In S. Melmed, K. S. Polonsky, P. R. Larsen, & H. M. Kronenberg (Eds.), Williams Textbook of Endocrinology (Thirteenth Edition) (pp. 589–663). Elsevier.

  • Burke, F. F., Hinks, M., Salia, S., Sparkes, K. M., & Swift-Gallant, A. (2022). Using animal models to study the interplay between the Biodevelopmental Pathways underlying human sexual orientation. Archives of Sexual Behavior, 1–6.

  • Calkin, J. (2023). 25 August 2023). A boy named Sophie. The Telegraph. Retrieved from https://www.telegraph.co.uk/family/life/sophie-ottaway-rare-birth-defect-gender-surgery/.

  • Cappelletti, M., & Wallen, K. (2016). Increasing women’s sexual desire: The comparative effectiveness of estrogens and androgens. Hormones and Behavior, 78, 178–193. https://doi.org/10.1016/j.yhbeh.2015.11.003.

    Article  Google Scholar 

  • Carani, C., Qin, K., Simoni, M., Faustini-Fustini, M., Serpente, S., Boyd, J., & Simpson, E. R. (1997). Effect of Testosterone and Estradiol in a man with Aromatase Deficiency. New England Journal of Medicine, 337(2), 91–95. https://doi.org/10.1056/nejm199707103370204.

    Article  Google Scholar 

  • Cardona Attard, C., Cameron-Pimblett, A., Puri, D., La Rosa, C., Talaulikar, V. S., Davies, M. C., & Conway, G. S. (2020). Relationship and sexual experiences in women with early-onset oestrogen deficiency: Comparison between women with Turner syndrome and premature ovarian insufficiency. Clinical Endocrinology, 93(4), 473–481. https://doi.org/10.1111/cen.14271.

    Article  Google Scholar 

  • Carter, M. J. (2014). Gender socialization and identity theory. Social Sciences, 3(2), 242–263.

    Article  Google Scholar 

  • Chen, C. H., Wang, P. H., Hsieh, M. T., Tzeng, C. R., Wu, Y. H., Lee, C. S., & Chang, H. Y. (2014). Sexual orientations of women with polycystic ovary syndrome: Clinical observation in Taiwan. Taiwanese Journal of Obstetrics and Gynecology, 53(4), 542–546. https://doi.org/10.1016/j.tjog.2014.09.002.

  • Choi, J., & Smitz, J. (2014). Luteinizing hormone and human chorionic gonadotropin: Distinguishing unique physiologic roles. Gynecological Endocrinology, 30(3), 174–181. https://doi.org/10.3109/09513590.2013.859670.

    Article  Google Scholar 

  • Clemens, L. G. (2012). 155The Aromatization Hypothesis, 1970–1990. In J. Balthazart, & G. Ball (Eds.), Brain aromatase, Estrogens, and Behavior (p. 0). Oxford University Press.

  • Cooke, P. S., Nanjappa, M. K., Ko, C., Prins, G. S., & Hess, R. A. (2017). Estrogens in male physiology. Physiological Reviews, 97(3), 995–1043. https://doi.org/10.1152/physrev.00018.2016.

    Article  Google Scholar 

  • Cornil, C. A., & Bakker, J. (2019). Alternative views on the role of sex steroid hormones on the emergence of phenotypic diversity in female sexual orientation. Archives of Sexual Behavior, 48(5), 1309–1313.

    Article  Google Scholar 

  • Daae, E., Feragen, K. B., Waehre, A., Nermoen, I., & Falhammar, H. (2020). Sexual orientation in individuals with congenital adrenal hyperplasia: A systematic review. Frontiers in Behavioral Neuroscience, 38.

  • De Vries, G. J., Rissman, E. F., Simerly, R. B., Yang, L. Y., Scordalakes, E. M., Auger, C. J., & Arnold, A. P. (2002). A model system for study of sex chromosome effects on sexually dimorphic neural and behavioral traits. Journal of Neuroscience, 22(20), 9005–9014.

    Article  Google Scholar 

  • Ehrhardt, A. A., Meyer-Bahlburg, H. F., Rosen, L. R., Feldman, J. F., Veridiano, N. P., Zimmerman, I., & McEwen, B. S. (1985). Sexual orientation after prenatal exposure to exogenous estrogen. Archives of Sexual Behavior, 14(1), 57–77.

    Article  Google Scholar 

  • Forastieri, V., Andrade, C. P., SOUZA, A. L. V., Silva, M. S., El-Hani, C. N., de Azevedo Moreira, L. M., & Flores, R. Z. (2002). Evidence against a relationship between dermatoglyphic asymmetry and male sexual orientation. Human Biology, 861–870.

  • Foresti, M. (2015). Adulthood with Turner Syndrome: Quality of life, psychosocial adjustment and clinical management in 70 Italian women (Ph.D.), Studiorum Università di Bologna. Dottorato di ricerca in Scienze psicologiche, University of Bologna Digital Library.

  • Forest, M. G. (1985). Pitfalls in prenatal diagnosis of 21-Hydroxylase Deficiency by Amniotic Fluid Steroid Analysis? A six years experience in 102 pregnancies at risk a. Annals of the New York Academy of Sciences, 458(1), 130–147.

    Article  Google Scholar 

  • Galis, F., Broek, T., Van Dongen, C. M., S., & Wijnaendts, L. C. (2010). Sexual dimorphism in the prenatal digit ratio (2D: 4D). Archives of Sexual Behavior, 39(1), 57–62.

    Article  Google Scholar 

  • Gangestad, S. W., Bailey, J. M., & Martin, N. G. (2000). Taxometric analyses of sexual orientation and gender identity. J Pers Soc Psychol, 78(6), 1109–1121. https://doi.org/10.1037//0022-3514.78.6.1109.

    Article  Google Scholar 

  • Goodarzi, M. O., Guo, X., Yildiz, B. O., Stanczyk, F. Z., & Azziz, R. (2007). Correlation of adrenocorticotropin steroid levels between women with polycystic ovary syndrome and their sisters. American Journal of Obstetrics and Gynecology, 196(4), https://doi.org/10.1016/j.ajog.2006.12.009.

  • Goto, M., Hanley, K. P., Marcos, J., Wood, P. J., Wright, S., Postle, A. D., & Hanley, N. A. (2006). In humans, early cortisol biosynthesis provides a mechanism to safeguard female sexual development. The Journal of Clinical Investigation, 116(4), 953–960.

    Article  Google Scholar 

  • Grimbos, T., Dawood, K., Burriss, R. P., Zucker, K. J., & Puts, D. A. (2010). Sexual orientation and the second to fourth finger length ratio: A meta-analysis in men and women. Behavioral Neuroscience, 124(2), 278.

    Article  Google Scholar 

  • Gur, E. B., Karadeniz, M., & Turan, G. A. (2015). Fetal programming of polycystic ovary syndrome. World Journal of Diabetes, 6(7), 936.

    Article  Google Scholar 

  • Hall, C. M., Jones, J. A., Meyer-Bahlburg, H. F., Dolezal, C., Coleman, M., Foster, P., & Clayton, P. E. (2004). Behavioral and physical masculinization are related to genotype in girls with congenital adrenal hyperplasia. The Journal of Clinical Endocrinology & Metabolism, 89(1), 419–424.

    Article  Google Scholar 

  • Hall, J. A. Y., & Kimura, D. (1994). Dermatoglyphic asymmetry and sexual orientation in men. Behavioral Neuroscience, 108(6), 1203.

    Article  Google Scholar 

  • Handy, A. B., Jackowich, R. A., Wibowo, E., Johnson, T. W., & Wassersug, R. J. (2016). Gender preference in the sexual attractions, fantasies, and relationships of voluntarily castrated men. Sexual Medicine, 4(1), e51–e59.

    Article  Google Scholar 

  • Han, T. S., & Bouloux, P. M. (2012). Kallmann syndrome and other causes of hypothalamic hypogonadism and related development disorders. Handbook of neuroendocrinology (pp. 597–617). Elsevier.

  • Harris, A., Bewley, S., & Meads, C. (2020). Sex hormone levels in lesbian, bisexual, and Heterosexual Women: Systematic review and exploratory Meta-analysis. Archives of Sexual Behavior, 49(7), 2405–2420. https://doi.org/10.1007/s10508-020-01717-8.

    Article  Google Scholar 

  • Heemers, H. V., & Tindall, D. J. (2007). Androgen receptor (AR) coregulators: A diversity of functions converging on and regulating the AR transcriptional complex. Endocrine Reviews, 28(7), 778–808.

    Article  Google Scholar 

  • Henrich, J. (2016). The secret of our success: How culture is driving human evolution, domesticating our species, and making us smarter. princeton University press.

  • Hettmer, E., Hoepffner, W., Keller, E., & Brähler, E. (1995). [Studies on sexual development, sexual behavior and ability to experience sex of young women with Ullrich-Turner syndrome]. Therapeutische Umschau, 52(2), 146–149.

    Google Scholar 

  • Hier, D. B., & Crowley, W. F. Jr. (1982). Spatial ability in androgen-deficient men. New England Journal of Medicine, 306(20), 1202–1205.

    Article  Google Scholar 

  • Hill, A. K., Dawood, K., & Puts, D. A. (2013). Biological foundations of sexual orientation. In A. R. D’Augelli, & C. J. Patterson (Eds.), Handbook of psychology and sexual orientation (pp. 55–68). Oxford University Press.

  • Hines, M. (2011). Prenatal endocrine influences on sexual orientation and on sexually differentiated childhood behavior. Frontiers in Neuroendocrinology, 32(2), 170–182. https://doi.org/10.1016/j.yfrne.2011.02.006.

    Article  Google Scholar 

  • Hines, M., Ahmed, S. F., & Hughes, I. A. (2003). Psychological outcomes and gender-related development in complete androgen insensitivity syndrome. Archives of Sexual Behavior, 32, 93–101.

    Article  Google Scholar 

  • Hiraishi, K., Sasaki, S., Shikishima, C., & Ando, J. (2012). The second to fourth digit ratio (2D: 4D) in a japanese twin sample: Heritability, prenatal hormone transfer, and association with sexual orientation. Archives of Sexual Behavior, 41, 711–724.

    Article  Google Scholar 

  • Holliday, J. (2015). 10 August 2015). The agony of learning you’ve been raised the wrong sex: A rare condition meant doctors weren’t sure what sex Joe was born. With support from Princess Diana, he was finally accepted as a girl. Then came a shattering discovery. Dailiy Mail.com. Retrieved from https://www.dailymail.co.uk/femail/article-3191600/The-agony-learning-ve-raised-wrong-sex-rare-condition-meant-doctors-weren-t-sure-sex-Joe-born-support-Princess-Diana-finally-accepted-girl-came-shattering-discovery.html.

  • Holmes, L., Watts-Overall, T. M., Slettevold, E., Gruia, D. C., Raines, J., & Rieger, G. (2021). Sexual orientation, sexual Arousal, and finger length ratios in women. Archives of Sexual Behavior, 50, 3419–3432.

    Article  Google Scholar 

  • Homburg, R. (2009). Androgen circle of polycystic ovary syndrome. Human Reproduction, 24(7), 1548–1555. https://doi.org/10.1093/humrep/dep049.

    Article  Google Scholar 

  • Homburg, R., Gudi, A., Shah, A., & Layton, M., A (2017). A novel method to demonstrate that pregnant women with polycystic ovary syndrome hyper-expose their fetus to androgens as a possible stepping stone for the developmental theory of PCOS. A pilot study. Reproductive Biology and Endocrinology, 15, 1–4.

    Article  Google Scholar 

  • Hong, H., Branham, W. S., Ng, H. W., Moland, C. L., Dial, S. L., Fang, H., & Tong, W. (2015). Human sex hormone-binding globulin binding affinities of 125 structurally diverse chemicals and comparison with their binding to androgen receptor, estrogen receptor, and alpha-fetoprotein. Toxicological Sciences, 143(2), 333–348. https://doi.org/10.1093/toxsci/kfu231.

    Article  Google Scholar 

  • Hughes, I. A., & Deeb, A. (2006). Androgen resistance. Best Practice & Research Clinical Endocrinology & Metabolism, 20(4), 577–598. https://doi.org/10.1016/j.beem.2006.11.003.

    Article  Google Scholar 

  • Jabbour, J., Holmes, L., Sylva, D., Hsu, K. J., Semon, T. L., Rosenthal, A. M., & Savin-Williams, R. C. (2020). Robust evidence for bisexual orientation among men. Proceedings of the National Academy of Sciences, 117(31), 18369–18377.

  • Jägetoft, Z., Unenge Hallerbäck, M., Julin, M., Bornehag, C. G., & Wikström, S. (2022). Anthropometric measures do not explain the 2D:4D ratio sexual dimorphism in 7-year-old children. Am J Hum Biol, 34(9), e23776. https://doi.org/10.1002/ajhb.23776.

    Article  Google Scholar 

  • Jones, B. C., Hahn, A. C., Fisher, C. I., Wang, H., Kandrik, M., & DeBruine, L. M. (2018). General sexual desire, but not desire for uncommitted sexual relationships, tracks changes in women’s hormonal status. Psychoneuroendocrinology, 88, 153–157.

    Article  Google Scholar 

  • Khorashad, B. S., Aghili, Z., Kreukels, B. P. C., Hiradfar, M., Roshan, G. M., Afkhamizadeh, M., & Cohen-Kettenis, P. T. (2016). Psychosexual outcome among iranian individuals with 5α-Reductase Deficiency Type 2 and its relationship with parental sexism. The Journal of Sexual Medicine, 13(11), 1629–1641. https://doi.org/10.1016/j.jsxm.2016.09.011.

    Article  Google Scholar 

  • Kim, G. W., Kim, S. K., & Jeong, G. W. (2016). Neural activation-based sexual orientation and its correlation with free testosterone level in postoperative female-to-male transsexuals: Preliminary study with 3.0-T fMRI. Surgical and Radiologic Anatomy, 38(2), 245–252. https://doi.org/10.1007/s00276-015-1547-z.

    Article  Google Scholar 

  • Kratochvíl, L., & Flegr, J. (2009). Differences in the 2nd to 4th digit length ratio in humans reflect shifts along the common allometric line. Biology Letters, 5(5), 643–646. https://doi.org/10.1098/rsbl.2009.0346.

    Article  Google Scholar 

  • Kronawitter, D., Gooren, L. J., Zollver, H., Oppelt, P. G., Beckmann, M. W., Dittrich, R., & Mueller, A. (2009). Effects of transdermal testosterone or oral dydrogesterone on hypoactive sexual desire disorder in transsexual women: Results of a pilot study. European Journal of Endocrinology, 161(2), 363–368.

    Article  Google Scholar 

  • Lalumiere, M. L., Blanchard, R., & Zucker, K. J. (2000). Sexual orientation and handedness in men and women: A meta-analysis. Psychological Bulletin, 126(4), 575.

    Article  Google Scholar 

  • Lew-Starowicz, Z., Jeż, W., Irzyniec, T., Kabzińska, M., & Boćkowska, E. (2003). Sexual aspects of women with Turner’s Syndrome. Sexuality and Disability, 21(4), 241–248. https://doi.org/10.1023/B:SEDI.0000010066.89406.0b.

    Article  Google Scholar 

  • Lippa, R., & Arad, S. (1997). The structure of sexual orientation and its relation to masculinity, femininity, and gender diagnosticity: Different for men and women. Sex Roles, 37, 187–208.

    Article  Google Scholar 

  • Lolli, L., Batterham, A. M., Kratochvíl, L., Flegr, J., Weston, K. L., & Atkinson, G. (2017). A comprehensive allometric analysis of 2nd digit length to 4th digit length in humans. Proceedings of the Royal Society B: Biological Sciences, 284(1857), 20170356. doi:https://doi.org/10.1098/rspb.2017.0356.

  • Luoto, S., & Rantala, M. J. (2017). On estrogenic masculinization of the human brain and behavior. Hormones and Behavior.

  • Lupu, D. C., Monedero, I., Rodriguez-Ruiz, C., Pita, M., & Turiegano, E. (2023). In support of 2D: 4D: More data exploring its conflicting results on handedness, sexual orientation and sex differences. Plos One, 18(8), e0280514.

  • Malas, M. A., Dogan, S., Evcil, E. H., & Desdicioglu, K. (2006). Fetal development of the hand, digits and digit ratio (2D: 4D). Early Human Development, 82(7), 469–475.

    Article  Google Scholar 

  • Manning, J. T. (2010). Digit ratio (2D:4D), sex differences, allometry, and finger length of 12–30-year olds: Evidence from the British Broadcasting Corporation (BBC) internet study. American Journal of Human Biology, 22(5), 604–608. https://doi.org/10.1002/ajhb.21051.

    Article  Google Scholar 

  • Martin, J. T., & Nguyen, D. H. (2004). Anthropometric analysis of homosexuals and heterosexuals: Implications for early hormone exposure. Hormones and Behavior, 45(1), 31–39. https://doi.org/10.1016/j.yhbeh.2003.07.003.

    Article  Google Scholar 

  • Martin, J. T., Puts, D. A., & Breedlove, S. M. (2008). Hand asymmetry in heterosexual and homosexual men and women: Relationship to 2D:4D digit ratios and other sexually dimorphic anatomical traits. Archives of Sexual Behavior, 37(1), 119–132. https://doi.org/10.1007/s10508-007-9279-8.

    Article  Google Scholar 

  • McCarthy, M. M. (2020). A new view of sexual differentiation of mammalian brain. Journal of Comparative Physiology A, 206(3), 369–378. https://doi.org/10.1007/s00359-019-01376-8.

    Article  Google Scholar 

  • McCarthy, M. M., Auger, A. P., Bale, T. L., Vries, G. J. D., Dunn, G. A., Forger, N. G., & Wilson, M. E. (2009). The Epigenetics of Sex differences in the brain. The Journal of Neuroscience, 29(41), 12815–12823. https://doi.org/10.1523/jneurosci.3331-09.2009.

    Article  Google Scholar 

  • McCormick, C. M., & Carré, J. M. (2020). Facing off with the phalangeal phenomenon and editorial policies: A commentary on Swift-Gallant, Johnson. Di Rita and Breedlove (2020) Hormones and Behavior, 120, 104710. https://doi.org/10.1016/j.yhbeh.2020.104710.

    Article  Google Scholar 

  • McEwen, B. S., Lieberburg, I., Chaptal, C., & Krey, L. C. (1977). Aromatization: Important for sexual differentiation of the neonatal rat brain. Hormones and Behavior, 9(3), 249–263.

    Article  Google Scholar 

  • McEwen, B. S., Plapinger, L., Chaptal, C., Gerlach, J., & Wallach, G. (1975). Role of fetoneonatal estrogen binding proteins in the associations of estrogen with neonatal brain cell nuclear receptors. Brain Research, 96(2), 400–406. https://doi.org/10.1016/0006-8993(75)90755-6.

    Article  Google Scholar 

  • McFadden, D., & Champlin, C. A. (2000). Comparison of auditory evoked potentials in heterosexual, homosexual, and bisexual males and females. Journal of the Association for Research in Otolaryngology, 1, 89–99.

    Article  Google Scholar 

  • McFadden, D., Loehlin, J. C., & Pasanen, E. G. (1996). Additional findings on heritability and prenatal masculinization of cochlear mechanisms: Click-evoked otoacoustic emissions. Hearing Research, 97(1), 102–119. https://doi.org/10.1016/S0378-5955(96)80012-8.

    Article  Google Scholar 

  • McFadden, D., & Pasanen, E. G. (1998). Comparison of the auditory systems of heterosexuals and homosexuals: Click-evoked otoacoustic emissions. Proceedings of the National Academy of Sciences, 95(5), 2709–2713.

  • Merke, D. P., & Bornstein, S. R. (2005). Congenital adrenal hyperplasia. The Lancet, 365(9477), 2125–2136. https://doi.org/10.1016/S0140-6736(05)66736-0.

    Article  Google Scholar 

  • Meyer-Bahlburg, H. F. (2001). Gender and sexuality in classic congenital adrenal hyperplasia. Endocrinology and Metabolism Clinics, 30(1), 155–171.

    Article  Google Scholar 

  • Meyer-Bahlburg, H. F., Dolezal, C., Baker, S. W., & New, M. I. (2008). Sexual orientation in women with classical or non-classical congenital adrenal hyperplasia as a function of degree of prenatal androgen excess. Archives of Sexual Behavior, 37(1), 85–99. https://doi.org/10.1007/s10508-007-9265-1.

    Article  Google Scholar 

  • Meyer-Bahlburg, H. F. L. (1984). Psychoendocrine Research on sexual orientation. Current Status and Future Options. In De G. J. Vries, De J. P. C. Bruin, H. B. M. Uylings, & M. A. Corner (Eds.), Progress in Brain Research (61 vol., pp. 375–398). Elsevier.

  • Meyer-Bahlburg, H. F. L., Ehrhardt, A. A., Rosen, L. R., Gruen, R. S., Veridiano, N. P., Vann, F. H., & Neuwalder, H. F. (1995). Prenatal estrogens and the development of homosexual orientation. Developmental Psychology, 31(1), 12.

    Article  Google Scholar 

  • Money, J., & Mittenthal, S. (1970). Lack of personality pathology in Turner’s syndrome: Relation to cytogenetics, hormones and physique. Behavior Genetics, 1(1), 43–56. https://doi.org/10.1007/BF01067370.

    Article  Google Scholar 

  • Money, J., Schwartz, M., & Lewis, V. G. (1984). Adult erotosexual status and fetal hormonal masculinization and demasculinization: 46, XX congenital virilizing adrenal hyperplasia and 46, XY androgen-insensitivity syndrome compared. Psychoneuroendocrinology, 9(4), 405–414.

    Article  Google Scholar 

  • Morris, J. A., Jordan, C. L., & Breedlove, S. M. (2004). Sexual differentiation of the vertebrate nervous system. Nature Neuroscience, 7(10), 1034.

    Article  Google Scholar 

  • Motta-Mena, N. V., & Puts, A. (2017). Endocrinology of human female sexuality, mating, and reproductive behavior. Hormones and Behavior, 91, 19–35. https://doi.org/10.1016/j.yhbeh.2016.11.012.

    Article  Google Scholar 

  • Mustanski, B. S., Bailey, J. M., & Kaspar, S. (2002). Dermatoglyphics, handedness, sex, and sexual orientation. Archives of Sexual Behavior, 31, 113–122.

    Article  Google Scholar 

  • Naftolin, F., Ryan, K., Davies, I., Reddy, V., Flores, F., Petro, Z., & Wolin, L. (1975). The formation of estrogens by central neuroendocrine tissues Paper presented at the Proceedings of the 1974 Laurentian Hormone Conference.

  • Nascimento, R. L. P., de Andrade Mesquita, I. M., Gondim, R., dos Apóstolos, R. A. A. C., Toralles, M. B., de Oliveira, L. B., & Barroso, U. (2018). Gender identity in patients with 5-alpha reductase deficiency raised as females. Journal of Pediatric Urology, 14(5), 419. .e411-419.e416.

    Article  Google Scholar 

  • Newbold, R. R. (1993). Gender-related behavior in women exposed prenatally to diethylstilbestrol. Environmental Health Perspectives, 101(3), 208–213.

    Article  Google Scholar 

  • New, M. I., Lekarev, O., Jacob, M., Macdonald, A., Parsa, A., & Yuen, T. T. (2023). Congenital adrenal hyperplasia owing to 21-hydroxylase deficiency. Genetic steroid disorders (pp. 35–61). Elsevier.

  • Oakes, M. B., Eyvazzadeh, A. D., Quint, E., & Smith, Y. R. (2008). Complete androgen insensitivity syndrome—a review. Journal of Pediatric and Adolescent Gynecology, 21(6), 305–310.

    Article  Google Scholar 

  • Page, M. J., McKenzie, J. E., Bossuyt, P. M., Boutron, I., Hoffmann, T. C., Mulrow, C. D., & Brennan, S. E. (2021). The PRISMA 2020 statement: An updated guideline for reporting systematic reviews. International Journal of Surgery, 88, 105906.

    Article  Google Scholar 

  • Pasterski, V. L., Geffner, M. E., Brain, C., Hindmarsh, P., Brook, C., & Hines, M. (2005). Prenatal hormones and postnatal socialization by parents as determinants of male-typical toy play in girls with congenital adrenal hyperplasia. Child Development, 76(1), 264–278.

    Article  Google Scholar 

  • Pasterski, V., Zucker, K. J., Hindmarsh, P. C., Hughes, I. A., Acerini, C., Spencer, D., & Hines, M. (2015). Increased cross-gender identification independent of gender role behavior in girls with congenital adrenal hyperplasia: Results from a standardized assessment of 4-to 11-year-old children. Archives of Sexual Behavior, 44, 1363–1375.

    Article  Google Scholar 

  • Pearcey, S. M., Docherty, K. J., & Dabbs, J. M. Jr. (1996). Testosterone and sex role identification in lesbian couples. Physiology & Behavior, 60(3), 1033–1035.

    Article  Google Scholar 

  • Phoenix, C., Goy, H., Gerall, R. W., A. A., & Young, W. C. (1959). Organizing action of prenatally administered testosterone propionate on the tissues mediating mating behavior in the female Guinea pig. Endocrinology, 65(3), 369–382. https://doi.org/10.1210/endo-65-3-369.

    Article  Google Scholar 

  • Pourquet, A., Teoli, J., Bouty, A., Renault, L., Roucher, F., Mallet, D., & Plotton, I. (2022). Steroid profiling in the amniotic fluid: Reference range for 12 steroids and interest in 21-Hydroxylase Deficiency. The Journal of Clinical Endocrinology & Metabolism, 108(5), e129–e138. https://doi.org/10.1210/clinem/dgac656.

    Article  Google Scholar 

  • Puts, D. A., Gaulin, S. J., Sporter, R. J., & McBurney, D. H. (2004). Sex hormones and finger length: What does 2D: 4D indicate? Evolution and Human Behavior, 25(3), 182–199.

    Article  Google Scholar 

  • Puts, D. A., Jordan, C. L., & Breedlove, S. M. (2006a). Defending the brain from estrogen. Nature Neuroscience, 9(2), 155–156.

    Article  Google Scholar 

  • Puts, D. A., Jordan, C. L., & Breedlove, S. M. (2006b). O brother, where art thou? The fraternal birth-order effect on male sexual orientation. Proceedings of the National Academy of Sciences, 103(28), 10531–10532.

  • Puts, D., & Motta-Mena, N. (2017). Is human brain masculinization estrogen receptor-mediated? Reply to Luoto and Rantala. Hormones and Behavior. https://doi.org/10.1016/j.yhbeh.2017.07.018.

    Article  Google Scholar 

  • Quigley, C. A. (2002). The postnatal gonadotropin and sex steroid surge—insights from the androgen insensitivity syndrome. The Journal of Clinical Endocrinology & Metabolism, 87(1), 24–28.

    Google Scholar 

  • Rahman, Q., Xu, Y., Lippa, R. A., & Vasey, P. L. (2020). Prevalence of sexual orientation across 28 nations and its association with gender Equality, Economic Development, and Individualism. Archives of Sexual Behavior, 49(2), 595–606. https://doi.org/10.1007/s10508-019-01590-0.

    Article  Google Scholar 

  • Rice, W. R., Friberg, U., & Gavrilets, S. (2012). Homosexuality as a consequence of epigenetically canalized sexual development. The Quarterly Review of Biology, 87(4), 343–368.

    Article  Google Scholar 

  • Rice, W. R., Friberg, U., & Gavrilets, S. (2013). Homosexuality via canalized sexual development: A testing protocol for a new epigenetic model. Bioessays, 35(9), 764–770.

    Article  Google Scholar 

  • Richards, G., Browne, W. V., Aydin, E., Constantinescu, M., Nave, G., Kim, M. S., & Watson, S. J. (2020). Digit ratio (2D:4D) and congenital adrenal hyperplasia (CAH): Systematic literature review and meta-analysis. Hormones and Behavior, 126, 104867. https://doi.org/10.1016/j.yhbeh.2020.104867.

    Article  Google Scholar 

  • Robinson, K., Galloway, K., Bewley, S., & Meads, C. (2017). Lesbian and bisexual women’s gynaecological conditions: A systematic review and exploratory meta-analysis. BJOG: An International Journal of Obstetrics & Gynaecology, 124(3), 381–392. https://doi.org/10.1111/1471-0528.14414.

    Article  Google Scholar 

  • Roney, J. R., & Simmons, Z. L. (2013). Hormonal predictors of sexual motivation in natural menstrual cycles. Hormones and Behavior, 63(4), 636–645. https://doi.org/10.1016/j.yhbeh.2013.02.013.

    Article  Google Scholar 

  • Rosenfield, K., Dawood, K., & Puts, D. (2019). Organizational effects of hormones on sexual orientation. In O. C. Schultheiss, & P. H. Mehta (Eds.), Routledge International Handbook of Social Neuroendocrinology (1 ed., pp. 256–280). Routledge/Psychology Press.

  • Safron, A., Barch, B., Bailey, J. M., Gitelman, D. R., Parrish, T. B., & Reber, P. J. (2007). Neural correlates of sexual arousal in homosexual and heterosexual men. Behavioral Neuroscience, 121(2), 237–248. https://doi.org/10.1037/0735-7044.121.2.237.

    Article  Google Scholar 

  • Safron, A., Sylva, D., Klimaj, V., Rosenthal, A. M., & Bailey, J. M. (2020). Neural responses to sexual Stimuli in Heterosexual and homosexual men and women: Men’s responses are more specific. Archives of Sexual Behavior, 49(2), 433–445. https://doi.org/10.1007/s10508-019-01521-z.

    Article  Google Scholar 

  • Sanders, A. R., Martin, E. R., Beecham, G. W., Guo, S., Dawood, K., Rieger, G., & Kolundzija, A. B. (2015). Genome-wide scan demonstrates significant linkage for male sexual orientation. Psychological Medicine, 45(7), 1379–1388.

    Article  Google Scholar 

  • Schrager, S., & Potter, B. E. (2004). Diethylstilbestrol exposure. American Family Physician, 69(10), 2395–2400.

    Google Scholar 

  • Seth, A. (2019). Optimizing stature in congenital adrenal hyperplasia: Challenges and Solutions. The Indian Journal of Pediatrics, 86(6), 489–491. https://doi.org/10.1007/s12098-019-02961-0.

    Article  Google Scholar 

  • Shirazi, T. N., Self, H., Dawood, K., Rosenfield, K. A., Penke, L., Carré, J. M., & Puts, D. A. (2019). Hormonal predictors of women’s sexual motivation. Evolution and Human Behavior, 40(3), 336–344.

    Article  Google Scholar 

  • Shirazi, T. N., Self, H., Dawood, K., Welling, L. L., Cárdenas, R., Rosenfield, K. A., & Breedlove, S. M. (2021). Evidence that perinatal ovarian hormones promote women’s sexual attraction to men. Psychoneuroendocrinology, 134, 105431.

    Article  Google Scholar 

  • Shirazi, T. N., Self, H., Rosenfield, K. A., Dawood, K., Welling, L. L., Cárdenas, R., & Breedlove, S. M. (2022). Low perinatal androgens predict recalled childhood gender nonconformity in men. Psychological Science, 33(3), 343–353.

    Article  Google Scholar 

  • Silfen, M. E., Denburg, M. R., Manibo, A. M., Lobo, R. A., Jaffe, R., Ferin, M., & Oberfield, S. E. (2003). Early endocrine, metabolic, and Sonographic characteristics of polycystic ovary syndrome (PCOS): Comparison between nonobese and obese adolescents. The Journal of Clinical Endocrinology & Metabolism, 88(10), 4682–4688. https://doi.org/10.1210/jc.2003-030617.

    Article  Google Scholar 

  • Singh, D., Vidaurri, M., Zambarano, R. J., & Dabbs, J. M. Jr. (1999). Lesbian erotic role identification: Behavioral, morphological, and hormonal correlates. J Pers Soc Psychol, 76(6), 1035.

    Article  Google Scholar 

  • Skaletsky, H., Kuroda-Kawaguchi, T., Minx, P. J., Cordum, H. S., Hillier, L., Brown, L. G., & Bieri, T. (2003). The male-specific region of the human Y chromosome is a mosaic of discrete sequence classes. Nature, 423(6942), 825–837.

    Article  Google Scholar 

  • Skorska, M. N., Coome, L. A., Peragine, D. E., Aitken, M., & VanderLaan, D. P. (2021). An anthropometric study of sexual orientation and gender identity in Thailand. Scientific Reports, 11(1), 18432. https://doi.org/10.1038/s41598-021-97845-9.

    Article  Google Scholar 

  • Skorska, M. N., Geniole, S. N., Vrysen, B. M., McCormick, C. M., & Bogaert, A. F. (2015). Facial structure predicts sexual orientation in both men and women. Archives of Sexual Behavior, 44(5), 1377–1394. https://doi.org/10.1007/s10508-014-0454-4.

    Article  Google Scholar 

  • Slabbekoorn, D., Van Goozen, S. H., Sanders, G., Gooren, L. J., & Cohen-Kettenis, P. T. (2000). The dermatoglyphic characteristics of transsexuals: Is there evidence for an organizing effect of sex hormones. Psychoneuroendocrinology, 25(4), 365–375.

    Article  Google Scholar 

  • Smith, E. P., Boyd, J., Frank, G. R., Takahashi, H., Cohen, R. M., Specker, B., & Korach, K. S. (1994). Estrogen resistance caused by a mutation in the estrogen-receptor gene in a man. New England Journal of Medicine, 331(16), 1056–1061. https://doi.org/10.1056/nejm199410203311604.

    Article  Google Scholar 

  • Speiser, P. W., & White, P. C. (2003). Congenital adrenal hyperplasia. New England Journal of Medicine, 349(8), 776–788.

    Article  Google Scholar 

  • Swartz, S. K., & Soloff, M. S. (1974). The lack of estrogen binding by human alpha-fetoprotein. Journal of Clinical Endocrinology and Metabolism, 39(3), 589–591. https://doi.org/10.1210/jcem-39-3-589.

    Article  Google Scholar 

  • Swift-Gallant, A., Coome, L. A., Aitken, M., Monks, D. A., & VanderLaan, D. P. (2019). Evidence for distinct biodevelopmental influences on male sexual orientation. Proceedings of the National Academy of Sciences, 116(26), 12787–12792.

  • Swift-Gallant, A., Coome, L. A., Monks, D. A., & VanderLaan, D. P. (2017). Handedness is a biomarker of variation in anal sex role behavior and recalled childhood gender nonconformity among gay men. Plos One, 12(2), e0170241.

  • Swift-Gallant, A., Coome, L. A., Monks, D. A., & VanderLaan, D. P. (2018). Gender nonconformity and birth order in relation to anal sex role among gay men. Archives of Sexual Behavior, 47, 1041–1052.

    Article  Google Scholar 

  • Swift-Gallant, A., Coome, L. A., Ramzan, F., & Monks, D. A. (2016). Nonneural androgen receptors affect sexual differentiation of Brain and Behavior. Endocrinology, 157(2), 788–798. https://doi.org/10.1210/en.2015-1355.

    Article  Google Scholar 

  • Swift-Gallant, A., Johnson, B. A., Di Rita, V., & Breedlove, S. M. (2020). Through a glass, darkly: Human digit ratios reflect prenatal androgens, imperfectly. Hormones and Behavior, 120, 104686.

    Article  Google Scholar 

  • Swift-Gallant, A., Shirazi, T., Puts, D. A., & Breedlove, S. M. (2022). Evidence for Perinatal Steroid influence on human sexual orientation and gendered behavior. Cold Spring Harbor Perspectives in Biology, 14(8), https://doi.org/10.1101/cshperspect.a039123.

  • Titus-Ernstoff, L., Perez, K., Hatch, E. E., Troisi, R., Palmer, J. R., Hartge, P., & Hoover, R. (2003). Psychosexual characteristics of men and women exposed prenatally to diethylstilbestrol. Epidemiology (Cambridge, Mass.), 14(2), 155–160. https://doi.org/10.1097/01.ede.0000039059.38824.b2.

    Article  Google Scholar 

  • Trakakis, E., Loghis, C., & Kassanos, D. (2009). Congenital adrenal hyperplasia because of 21-hydroxylase deficiency: A genetic disorder of interest to obstetricians and gynecologists. Obstetrical & Gynecological Survey, 64(3), 177–189.

    Article  Google Scholar 

  • Valentova, J. V., & Havlíček, J. (2013). Perceived sexual orientation based on vocal and facial stimuli is linked to self-rated sexual orientation in czech men. Plos One, 8(12), e82417.

  • Valentova, J. V., Kleisner, K., Havlíček, J., & Neustupa, J. (2014). Shape differences between the faces of homosexual and heterosexual men. Archives of Sexual Behavior, 43, 353–361.

    Article  Google Scholar 

  • Vink, J. M., Sadrzadeh, S., Lambalk, C. B., & Boomsma, D. I. (2006). Heritability of polycystic ovary syndrome in a dutch twin-family study. The Journal of Clinical Endocrinology & Metabolism, 91(6), 2100–2104. https://doi.org/10.1210/jc.2005-1494.

    Article  Google Scholar 

  • Wallen, K. (2005). Hormonal influences on sexually differentiated behavior in nonhuman primates. Front Neuroendocrinol, 26(1), 7–26. https://doi.org/10.1016/j.yfrne.2005.02.001.

    Article  Google Scholar 

  • Wallen, K. (2009). The organizational hypothesis: reflections on the 50th anniversary of the publication of Phoenix, Goy, Gerall, and Young (1959). Hormones and behavior, 55(5), 561–565. https://doi.org/10.1016/j.yhbeh.2009.03.009.

  • Wampold, C. H. (2018). The association between fraternal birth order and anal-erotic roles of men who have sex with men. Archives of Sexual Behavior, 47, 2445–2454.

    Article  Google Scholar 

  • Wassersug, R. J., Westle, A., & Dowsett, G. W. (2017). Men’s sexual and relational adaptations to erectile dysfunction after prostate cancer treatment. International Journal of Sexual Health, 29(1), 69–79. https://doi.org/10.1080/19317611.2016.1204403.

    Article  Google Scholar 

  • Watts, T. M., Holmes, L., Raines, J., Orbell, S., & Rieger, G. (2018). Finger length ratios of identical twins with discordant sexual orientations. Archives of Sexual Behavior, 47, 2435–2444.

    Article  Google Scholar 

  • Wisniewski, A. B., Migeon, C. J., Meyer-Bahlburg, H. F., Gearhart, J. P., Berkovitz, G. D., Brown, T. R., & Money, J. (2000). Complete androgen insensitivity syndrome: Long-term medical, surgical, and psychosexual outcome. The Journal of Clinical Endocrinology & Metabolism, 85(8), 2664–2669.

    Google Scholar 

  • Witchel, S. F. (2017). Congenital adrenal hyperplasia. Journal of Pediatric and Adolescent Gynecology, 30(5), 520–534.

    Article  Google Scholar 

  • Wong, W. I., Pasterski, V., Hindmarsh, P. C., Geffner, M. E., & Hines, M. (2013). Are there parental socialization Effects on the sex-typed behavior of individuals with congenital adrenal hyperplasia? Archives of Sexual Behavior, 42(3), 381–391. https://doi.org/10.1007/s10508-012-9997-4.

    Article  Google Scholar 

  • Wudy, S. A., Dörr, H. G., Solleder, C., Djalali, M., & Homoki, J. (1999). Profiling steroid hormones in amniotic fluid of midpregnancy by routine stable isotope Dilution/Gas Chromatography-Mass Spectrometry: Reference values and concentrations in fetuses at risk for 21-Hydroxylase Deficiency1. The Journal of Clinical Endocrinology & Metabolism, 84(8), 2724–2728. https://doi.org/10.1210/jcem.84.8.5870.

    Article  Google Scholar 

  • Wu, M. V., Manoli, D. S., Fraser, E. J., Coats, J. K., Tollkuhn, J., Honda, S. I., & Shah, N. M. (2009). Estrogen masculinizes neural pathways and sex-specific behaviors. Cell, 139(1), 61–72.

    Article  Google Scholar 

  • Xita, N., & Tsatsoulis, A. (2006). Fetal programming of polycystic ovary syndrome by androgen excess: Evidence from experimental, clinical, and genetic association studies. The Journal of Clinical Endocrinology & Metabolism, 91(5), 1660–1666.

    Article  Google Scholar 

  • Xu, X., Coats, J. K., Yang, C. F., Wang, A., Ahmed, O. M., Alvarado, M., & Shah, N. M. (2012). Modular genetic control of sexually dimorphic behaviors. Cell, 148(3), 596–607.

    Article  Google Scholar 

  • Zheng, L., Hart, T. A., & Zheng, Y. (2012). The relationship between intercourse preference positions and personality traits among gay men in China. Archives of Sexual Behavior, 41, 683–689.

    Article  Google Scholar 

  • Zietsch, B. P., & Sidari, M. J. (2020). The Kinsey scale is ill-suited to most sexuality research because it does not measure a single construct. Proceedings of the National Academy of Sciences, 117(44), 27080–27080.

  • Zuloaga, D. G., Puts, D. A., Jordan, C. L., & Breedlove, S. M. (2008). The role of androgen receptors in the masculinization of brain and behavior: What we’ve learned from the testicular feminization mutation. Hormones and Behavior, 53(5), 613–626. https://doi.org/10.1016/j.yhbeh.2008.01.013.

    Article  Google Scholar 

Download references

Acknowledgements

We thank Mike Bailey for helpful feedback on a previous draft.

Funding

No funding was received to assist with the preparation of this manuscript.

Author information

Authors and Affiliations

  1. Department of Psychology, Memorial University of Newfoundland and Labrador, St. John’s, NL, Canada

    Ashlyn Swift-Gallant

  2. Department of Psychology and Counseling, Immaculata University, Immaculata, PA, USA

    Toe Aung

  3. Department of Anthropology, Pennsylvania State University, University Park, PA, USA

    Kevin Rosenfield & David Puts

  4. Department of Psychology, Pennsylvania State University, University Park, PA, USA

    Khytam Dawood

Authors
  1. Ashlyn Swift-Gallant
    View author publications

    You can also search for this author in PubMed Google Scholar

  2. Toe Aung
    View author publications

    You can also search for this author in PubMed Google Scholar

  3. Kevin Rosenfield
    View author publications

    You can also search for this author in PubMed Google Scholar

  4. Khytam Dawood
    View author publications

    You can also search for this author in PubMed Google Scholar

  5. David Puts
    View author publications

    You can also search for this author in PubMed Google Scholar

Contributions

A.S.G., K.R. and D.P. co-wrote the first draft of the manuscript, A.S.G. collected and coded meta-analysis data, T.A. conducted statistical analyses, D.P. produced Figs. 1 and 2, and 5; A.S.G. produced Fig. 3, TA produced Fig. 4, and all authors reviewed the manuscript.

Corresponding author

Correspondence to David Puts.

Ethics declarations

Competing Interests

The corresponding author (D.P.) is an Associate Editor of this journal; the authors declare that they have no other conflicts of interest.

Ethical Approval

No ethical approval was sought for this research; it is a review of published literature.

Financial Interests

The authors declare they have no financial interests.

Non-financial Interests

None.

Additional information

Publisher’s Note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Rights and permissions

Springer Nature or its licensor (e.g. a society or other partner) holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law.

Reprints and permissions

About this article

Check for updates. Verify currency and authenticity via CrossMark

Cite this article

Swift-Gallant, A., Aung, T., Rosenfield, K. et al. Organizational Effects of Gonadal Hormones on Human Sexual Orientation. Adaptive Human Behavior and Physiology 9, 344–370 (2023). https://doi.org/10.1007/s40750-023-00226-x

Download citation

  • Received:

  • Revised:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s40750-023-00226-x

Keywords

Search

Navigation