Chapter 15: Sex Determination

Where possible, references have been linked to PubMed, the National Center for Biotechnology Information’s online database of journal article citations. Citations that do not include links to PubMed are either referring to papers that are too old to be included in PubMed, or to books, which are not listed on PubMed.

Agate, R. J. and 7 others. 2002. Neural, not gonadal, origin of brain sex differences in a gynandromorphic finch. Proc. Natl. Acad. Sci. USA 100: 4873–4878.
PubMed Link

Ahmad, S. M. and B. S. Baker. 2002. Sex-specific deployment of FGF signaling in Drosophila recruits mesodermal cells into the male genital imaginal disc. Cell 109: 651–661.
PubMed Link

Andersson, S., D. M. Berman, E. P. Jenkins and D. W. Russell. 1991. Deletion of steroid 5a-reductase 2 gene in male pseudohermaphroditism. Nature 354: 159–161.
PubMed Link

Arango, N. A., R. Lovell-Badge and R. R. Behringer. 1999. Targeted mutagenesis of the endogenous mouse Mis gene promoter: In vivo definition of genetic pathways of vertebrate sexual development. Cell 99: 409–419.
PubMed Link

Arnold, A. P. and P. S. Burgoyne. 2004. Are XX and XY brain cells intrinsically different? Trends Endocr. Metab. 15: 6–11.
PubMed Link

Austin, J., A. Tamar-Mattis, T. Mazur, M. J. Henwood and W. C. Rossi. 2011  Disorders of sex development: When and how to tell the patient. Pediatr. Endocrinol. Rev. 83: 213–217.
PubMed Link

Avila, F. W. and J. W. Erickson. 2007. Drosophila JAK/STAT pathway reveals distinct initiation and reinforcement steps in early transcription of Sxl. Curr. Biol. 17: 643–648.
PubMed Link

Bailey, J. M., M. P. Dunne and N. G. Martin. 2000. Genetic and environmental influences on sexual orientation and its correlates in an Australian twin sample. J. Pers. Soc. Psychol. 78: 524–536.
PubMed Link

Baker, B. S. 1989. Sex in flies: The splice of life. Nature 340: 521–524.
PubMed Link

Baker, B. S., B. J. Taylor and J. C. Hall. 2001. Are complex behaviors specified by directed regulatory genes? Reasoning from Drosophila. Cell 105: 13–24.
PubMed Link

Baker, B. S., R. N. Nagoshi and K. C. Burtis. 1987. Molecular genetic aspects of sex determination in Drosophila. BioEssays 6: 66–70.
PubMed Link

Bakker, J. and M. J. Baum. 2008. Role for estradiol in female-typical brain and behavioral sexual differentiation. Front. Neuroendocrinol. 29: 1–16.
PubMed Link

Bakker, J., C. De Mees, Q. Douhard, J. Balthazart, P. Gabant, J. Szpirer and C. Szpirer. 2006. a-fetoprotein protects the developing female mouse brain from masculinization and defeminization by estrogens. Nature Neurosci. 9: 220–226.
PubMed Link

Baltzer, F. 1914. Die Bestimmung und der Dimorphismus des Geschlechtes bei Bonellia. Sber. Phys.-Med. Ges. Würzb. 43: 1–4.

Barbash, J. E., G. P. Thelin, D. W. Kolpin and R. J. Gilliom. 2001. Major herbicides in ground water: Results from the National Water-Quality Assessment. J. Environ. Qual. 30: 831–845.
PubMed Link

Barraclough, C. A. and R. A. Gorski. 1962. Studies on mating behavior in the androgen-sterilized female rat in relation to the hypothalamic regulation of sexual behavior. J. Endocrinol. 25: 175–182.
PubMed Link

Barrionuevo, F., S. Bagheri-Fam, J. Klattig, R. Kist, M. M. Taketo, C. Englert and G. Scherer. 2006. Homozygous inactivation of Sox9 causes complete XY sex reversal in mice. Biol. Reprod. 74: 195–201.
PubMed Link

Barrios, F. and 9 others. 2010. Opposing effects of retinoic acid and Fgf9 on Nanos2 expression and meiotic entry of mouse germ cells. J. Cell Sci. 123: 871–880.
PubMed Link

Barske, L. A. and B. Capel. 2010. Sex determination: An avian sexual revolution. Nature 464: 171–172.
PubMed Link

Battaglin, W. A., E. T. Furlong, M. R. Burkhardt and C. J. Peter. 2000. Occurrence of sulfonylurea, sulfonamide, imidazolinone, and other herbicides in rivers, reservoirs, and ground water in the midwestern United States 1998. Sci. Total Environ. 248: 123–133.
PubMed Link

Baum, M. J. 2009. Sexual differentiation of pheromone processing: Links to male-typical mating behavior and partner preference. Horm. Behav. 55: 579–588.
PubMed Link

Beckman, K., M. Grskovic, F. Gebauer and M. W. Hentze. 2005. A dual inhibitory mechanism restricts Msl-2 mRNA translation for dosage compensation in Drosophila. Cell 122: 529–540.
PubMed Link

Bell, L. R., J. I. Horabin, P. Schedl and T. W. Cline. 1991. Positive autoregulation of Sex-lethal by alternative splicing maintains the female determined state in Drosophila. Cell 65: 229–239.
PubMed Link

Bergeron, J. M., D. Crews and J. A. MacLachlan. 1994. PCBs as environmental estrogens: Turtle sex determination as a biomarker of environmental contamination. Environ. Health Perspect. 102: 780–781.
PubMed Link

Bergeron, J. M., E. Willingham, C. T. Osborn III, T. Rhen and D. Crews. 1999. Developmental synergism of steroidal estrogens in sex determination. Environ. Health Perspect. 107: 93–97.
PubMed Link

Berkovitz, G. D. and 7 others. 1992. The role of the sex-determining region of the Y chromosome (SRY) in the etiology of 46,XX true hermaphrodites. Hum. Genet. 88: 411–416.
PubMed Link

Bernard, P. and 6 others. 2012. Wnt signaling in ovarian development inhibits Sf1 activation of Sox9 via the Tesco enhancer. Endocrinology 153: 901–912.
PubMed Link

Bernard, P., H. Sim, K. Knower, E. Vilain and V. Harley. 2008. Human SRY inhibits b-catenin-mediated transcription. Int. J. Biochem. Cell Biol. 40: 2889–2900.
PubMed Link

Beukeboom, L. W. 1995. Sex determination in Hymenoptera: A need for genetic and molecular studies. Bioessays 17: 813–817.
PubMed Link

Beyer, C., C. Pilgrim and I. Reisert. 1991. Dopamine content and metabolism in mesencephalic and diencephalic cell cultures: Sex differences and effects of sex steroids. J. Neurosci. 11: 1325–1333.
PubMed Link

Billeter, J. C., J. Atallah, J. J. Krupp, J. G. Millar and J. D. Levine. 2009. Specialized cells tag sexual and species identity in Drosophila melanogaster. Nature 461: 987–991.
PubMed Link

Billeter, J.-C. and S. F. Goodwin. 2004. Characterization of Drosophila fruitless-gal4 transgenes reveals expression in male-specific fruitless neurons and innervation of male reproductive structures. J. Comp. Neurol. 475: 270–287.
PubMed Link

Birk, O. S. and 11 others. 2000. The LIM homeobox gene LHX9 is essential for mouse gonad formation. Nature 403: 909–913.
PubMed Link

Black, M. P., M. Baillien, J. Balthazart and M. S. Grober. 2005. Socially induced and rapid increases in aggression are inversely related to brain aromatase activity in a sex-changing fish, Lythrypnus dalli. Proc. Biol. Sci. 272: 2435–2440.
PubMed Link

Blackless, M., A. Charuvastra, A. Derryck, A. Fausto-Sterling, K. Lauzanne and E. Lee. 2000. How sexually dimorphic are we? Review and synthesis. Am. J. Hum. Biol. 12: 151–166.
PubMed Link

Bleier, R. 1984. Science and Gender. Pergamon, New York.

Blumenstyk, G. 2003. The story of Syngenta and Tyrone Hayes at UC Berkeley: A Berkeley scientist says a corporate sponsor tried to bury his unwelcome findings and then buy his silence. Chron. Higher Ed. 50 (10) October. http: //www.mindfully.org/Pesticide/2003/Syngenta-Tyrone-Hayes31oct03.htm.

Boggs, R. T., P. Gregor, S. Idriss, J. M. Belote and M. McKeown. 1987. Regulation of sexual differentiation in D. melanogaster via alternative splicing of RNA from the transformer gene. Cell 50: 739–747.
PubMed Link

Bowles, J., C. W. Feng, C. Spiller, T. L. Davidson, A. Jackson and P. Koopman. 2010. Fgf9 suppresses meiosis and promotes male germ cell fate in mice. Dev. Cell. 19: 440–449.
PubMed Link

Bradford, S. T. and 10 others. 2009a. The cerebellin 4 precursor gene is a direct target of SRY and SOX9 in mice. Biol. Reprod. 80: 1178–1188.
PubMed Link

Bradford, S. T., D. Wilhelm, R. Bandiera, V. Vidal, A. Schedl and P. Koopman. 2009b. A cell-autonomous role for WT1 in regulating Sry in vivo. Hum. Mol. Genet. 18: 3429–3438.
PubMed Link

Breedlove, S. M. 1997. Sex on the brain. Nature 389: 801.
PubMed Link

Brennan, J. and B. Capel. 2004. One tissue, two fates: Molecular genetics that underlie testis versus ovary development. Nature Reviews Genet. 5: 509–521.
PubMed Link

Brennan, J., J. Karl and B. Capel. 2002. Divergent vascular mechanisms downstream of Sry establish the arterial system in the XY gonad. Dev. Biol. 244: 418–428.
PubMed Link

Bridges, C. B. 1925. Sex in relation to chromosomes and genes. Am. Nat. 59: 127–137.

Bull, J. J. 1980. Sex determination in reptiles. Q. Rev. Biol. 55: 3–21.

Bull, J. J., W. H. N. Gutzke and D. Crews. 1988. Sex reversal by estradiol in three reptilian orders. Gen. Comp. Endocrinol. 70: 425–428.
PubMed Link

Bullejos, M. and P. Koopman. 2005. Delayed Sry and Sox9 expression in developing mouse gonads underlie B6-YDOM sex reversal. Dev. Biol. 278: 473–481.
PubMed Link

Cain, D. A., L. J. Guillette Jr., A. A. Rooney and D. P. Pickford. 1997. Alterations in steroidogenesis in alligators (Alligator mississippiensis) exposed naturally and experimentally to environmental contaminants. Env. Health Persp. 105: 528–531.

Cameron, N., A. Del Corpo, J. Diorio, K. McAllister, S. Sharma and M. J. Meaney. 2008. Maternal programming of sexual behavior and hypothalamic-pituitary-gonadal function in the female rat. PLoS One. 3(5): e2210
PubMed Link

Capel, B., K. H. Albrecht, L. L. Washburn and E. M. Eicher. 1999. Migration of mesonephric cells into the mammalian gonad depends on Sry. Mech. Dev. 84: 127–131.
PubMed Link

Cate, R. L. and 18 others. 1986. Isolation of the bovine and human genes for Müllerian inhibiting substance and expression of the gene in animal cells. Cell 45: 685–698.
PubMed Link

Certel, S. J., M. G. Savella, D. C. Schlegel and E. A. Kravitz. 2007. Modulation of Drosophila male behavioral choice. Proc. Natl. Acad. Sci. USA 104: 4706–4711.
PubMed Link

Champagne, F. A. 2008. Epigenetic mechanisms and the transgenerational effects of maternal care. Front. Neuroendocrinol. 29: 386–397.
PubMed Link

Champagne, F. A., I. C. Weaver, J. Diorio, S. Dymov, M. Szyf and M. J. Meaney. 2006. Maternal care associated with methylation of the estrogen receptor-alpha1b promoter and estrogen receptor-alpha expression in the medial preoptic area of female offspring. Endocrinology. 147: 2909–2915.
PubMed Link

Christiansen, A. E., E. L. Keisman, S. M. Ahmad and B. S. Baker. 2002. Sex comes in from the cold: The integration of sex and pattern. Trends Genet. 18: 510–516.
PubMed Link

Coe, W. R. 1936. Sexual phases in Crepidula. J. Exp. Zool. 72: 455–477.

Collin, R. 1995. Sex, size, and position: A test of models predicting size at sex change in the protandrous gastropod, Crepidula fornicata. Am. Nat. 146: 815–831.

Colvin, J., R. P. Green, J. Schmal, B. Capel and D. M. Ornitz. 2001. Male-to-female sex reversal in mice lacking fibroblast growth factor 9. Cell 104: 875–889.
PubMed Link

Combes, A. N., D. Wilhelm, T. Davidson, E. Dejana, V. Harley, A. Sinclair and P. Koopman. 2009. Endothelial cell migration directs testis cord formation. Dev. Biol. 326: 112–120.
PubMed Link

Cool, J. and B. Capel. 2009. Mixed signals: Development of the testis. Semin. Reprod. Med. 27: 5–13.
PubMed Link

Cool, J. and B. Capel. 2009. Mixed signals: Development of the testis. Semin. Reprod. Med. 27: 5–3.
PubMed Link

Couse, J. F. and K. S. Korach. 2001. Contrasting phenotypes in reproductive tissues of female estrogen receptor null mice. Ann. N.Y. Acad. Sci. 948: 1–8.
PubMed Link

Couse, J. F., S. C. Hewitt, D. O. Bunch, M. Sar, V. R. Walker, B. J. Davis and K. S. Korach. 1999. Postnatal sex reversal of the ovaries in mice lacking estrogen receptors alpha and beta. Science 286: 2328–2331.
PubMed Link

Crain, D. A. and L. J. Guillette, Jr. 1998. Reptiles as models of contaminant-induced endocrine disruption. Anim. Reprod. Sci. 53: 77–86.
PubMed Link

Crews, D. 2003. Sex determination: Where environment and genetics meet. Evol. Dev. 5: 50–55.
PubMed Link

Crews, D. and J. J. Bull. 2009. Mode and tempo in environmental sex determination in vertebrates. Semin. Cell Dev. Biol. 20: 251–255.
PubMed Link

Dalton, R. 2002. Frogs put in the gender blender by America’s favourite herbicide. Nature 416: 665–666.
PubMed Link

De Mees, C. and 8 others. 2006. a-fetoprotein controls female fertility and prenatal development of the gonadotropin-releasing hormone pathway through an antiestrogenic action. Mol. Cell Biol. 26: 2012–2018.
PubMed Link

De Santa Barbara, P., B. Moniot, F. Poulat and P. Berta. 2000. Expression and subcellular localization of SF-1, SOX9, WT1, and AMH proteins during early human testicular development. Dev. Dynam. 217: 293–298.
PubMed Link

Demir, E. and B. J. Dickson. 2005. Fruitless splicing specifies male courtship behavior in Drosophila. Cell 121: 785–794.
PubMed Link

Desvages, G., M. Girondot and C. Pieau. 1993. Sensitive stages for the effects of temperature on gonadal aromatase activity in embryos of the marine turtle Dermochelys coriacea. Gen. Comp. Endocrinol. 92: 54–61.
PubMed Link

Dewing, P. and 10 others. 2006. Direct regulation of adult brain function by the male-specific factor SRY. Curr. Biol. 16: 415–420.
PubMed Link

Dewing, P., T. Shi, S. Horvath and E. Vilain. 2003. Sexually dimorphic gene expression in mouse brain precedes gonadal differentiation. Brain Res: Mol. Brain Res. 118: 82–90.
PubMed Link

Ditch, L. M. and 7 others. 2005. Drosophila retained/dead ringer is necessary for neuronal pathfinding, female receptivity, and repression of fruitless-independent male courtship behaviors. Development 132: 155–164.
PubMed Link

Dorizzi, M., G. Richard-Mercier, G. Desvages and C. Pieau. 1994. Masculinization of gonads by aromatase inhibitors in a turtle with temperature-dependent sex determination. Differentiation 58: 1–8.

Eicher, E. M. and 8 others. 1996. Sex-determining genes on mouse autosomes identified by linkage analysis of C57BL/6J-Y-POS sex reversal. Nature Genet. 14: 206–209.
PubMed Link

Eicher, E. M. and L. L. Washburn. 1983. Inherited sex reversal in mice: Identification of a new sex-determining gene. J. Exp. Zool. 228: 297–304.
PubMed Link

Erickson, J. W. and J. J. Quintero. 2007. Indirect effects of ploidy suggest X chromosome dose, not the X:A ratio, signals sex in Drosophila. PLoS Biol. 5: e332.
PubMed Link

Fan, W. and 10 others. 2007. Atrazine-induced aromatase expression is SF-1 dependent: Implications for endocrine disruption in wildlife and resproductive cancers in humans. Environ. Health Persp. 115: 720–727.
PubMed Link

Fausto-Sterling, A. 1992. Myths of Gender. Basic Books, New York.

Fisher, C. R., K. H. Graves, A. F. Parlow and E. R. Simpson. 1998. Characterization of mice deficient in aromatase (ArKO) because of a targeted disruption of the cyp19 gene. Proc. Natl. Acad. Sci. USA 95: 6965–6970.
PubMed Link

Ford, C. E., K. W. Jones and P. E. Polani. 1959. A sex chromosomal anomaly in a case of gonadal dysgenesis (Turner's syndrome). Lancet 1: 771–713.
PubMed Link

Freiman, R. N., S. R. Albright, S. Zheng, W. C. Sha, R. E. Hammer and R. Tjian. 2002. Requirement of tissue-specific TBP-associated factor TAF11105 in ovarian development. Science 293: 2084–2087.
PubMed Link

Geddes, P. and J. A. Thomson. 1890. The Evolution of Sex. Walter Scott, London.

Geissler, W. M. and 9 others. 1994. Male pseudohermaphroditism caused by mutations of testicular 17b-hydroxysteroid dehydrogenase 3. Nature Genet. 7: 34–39.
PubMed Link

Gempe, T., M. Hasselmann, M. Schiøtt, G. Hause, M. Otte and M. Beye. 2009. Sex determination in honeybees: Two separate mechanisms induce and maintain the female pathway. PLoS Biol. 7: e1000222.
PubMed Link

Gilbert, S. F.1978. The embryological origins of the gene theory.J. Hist. Biol.11:307–351.
PubMed Link

Gilbert, S. G., A. Tyler and E. Zackin. 2005. Bioethics and the New Embryology: Springboards for Debate. Sinauer Associates, Sunderland, MA.

Godwin, J. 2009. Social determination of sex in reef fishes: Environmental sex determination in vertebrates. Semin. Cell Dev. Biol. 20: 264–270.
PubMed Link

Godwin, J., J. A. Luckenbach and R. J. Borski. 2003. Ecology meets endocrinology: Environmental sex determination in fishes. Evol. Dev. 5: 40–49.
PubMed Link

González, A. N., H. Lu and J. W. Erickson. 2008. A shared enhancer controls a temporal switch between promoters during Drosophila primary sex determination. Proc. Natl. Acad. Sci. USA 105: 18436–18441.
PubMed Link

Gubbay, J. and 8 others. 1990. A gene mapping to the sex-determining region of the mouse Y chromosome is a member of a novel family of embryonically expressed genes. Nature 346: 245–250.
PubMed Link

Gutzke, W. H. N. and D. B. Chymiy. 1988. Sensitive periods during embryology for hormonally induced sex determination in turtles. Gen. Comp. Endocrinol. 71: 265–267.
PubMed Link

Hacker, A., B. Capel, P. Goodfellow and R. Lovell-Badge. 1995. Expression of Sry, the mouse sex determining gene. Development 121: 1603–1614.
PubMed Link

Handa, N. and 7 others. 1999. Structural basis for recognition of the tra mRNA precursor by the Sex-lethal protein. Nature 398: 579–585.
PubMed Link

Hayes, T. 2005. Welcome to the revolution. Integrative biology and assessing the impact of endocrine disruptors on environmental and public health. Integr. Comp. Biol. 45: 321–329.

Hayes, T. B. and 10 others. 2010. Atrazine induces complete feminization and chemical castration in male African clawed frogs (Xenopus laevis). Proc. Natl. Acad. Sci. USA 107: 4612–4617.
PubMed Link

Hayes, T. B., A. Collins, M. Lee, M. Mendoza, N. Noriega, A. Stuart and A. Vonk. 2002a. Hermaphroditic, demasculinized frogs after exposure to the herbicide atrazine at low ecologically relevant doses. Proc. Natl. Acad. Sci. USA 99: 5476–5480.
PubMed Link

Hayes, T., K. Haston, M. Tsui, A. Hoang, C. Haeffele and A. Vonk. 2002b. Herbicides: Feminization of male frogs in the wild. Nature 419: 895–896.
PubMed Link

Hayes, T., K. Haston, M. Tsui, A. Hoang, C. Haeffele and A. Vonk. 2003. Atrazine-induced hermaphroditism at 0.1 ppb in American leopard frogs (Rana pipiens): Laboratory and field evidence. Envir. Health Perspec. 111: 568–575.
PubMed Link

Heikkilä, M. and 7 others. 2005. The partial female-to-male sex reversal in Wnt-4-deficient females involves induced expression of testosterone biosynthetic genes and testosterone production, and depends on estrogen action. Endocrinology 146: 4016–4023.
PubMed Link

Heinrichs, V., L. C. Ryner and B. S. Baker. 1998. Regulation of sex-specific selection of fruitless 5′ splicing sites by transformer and transformer-2. Mol. Cell Biol. 18: 450–458.
PubMed Link

Hess, R. A., D. Bunick, K.-H. Lee, J. Bahr, J. A. Taylor, K. S. Korach and D. B. Lubahn. 1997. A role for oestrogens in the male reproductive system. Nature 390: 509–512.
PubMed Link

Hiramatsu, R. and 9 others. 2009. A critical time window of Sry action in gonadal sex determination in mice. Development 136: 129–138.
PubMed Link

Holderegger, C. and D. Keefer. 1986. The ontogeny of the mouse estrogen receptor: The pelvic region. Am. J. Anat. 177: 285–297.
PubMed Link

Huang, B., S. Wang, Y. Ning, A. N. Lamb and J. Bartley. 1999. Autosomal XX sex reversal caused by duplication of SOX9. Am. J. Med. Genet. 87: 349–353.
PubMed Link

Hughes, I. A., C. Houk, S. F. Ahmed and  P. A. Lee. 2006. Consensus statement on management of intersex disorders. J. Pediatr. Urol. 2:148–162
PubMed Link

Hull, C. L. 2003. Letter to the editor: How sexually dimorphic are we? Am. J. Hum. Biol. 15: 112–114.

Imperato-McGinley, J., L. Guerrero, T. Gautier and R. E. Peterson. 1974. Steroid 5a-reductase deficiency in man: An inherited form of male pseudohermaphroditism. Science 186: 1213–1215.
PubMed Link

Ito, H. and 6 others. 2012. Fruitless recruits two antagonistic chromatin factors to establish single-neuron sexual dimorphism. Cell 149: 1327–1338.
PubMed Link

Jääskeläinen, J. 2012. Molecular biology of androgen insensitivity. Mol. Cell Endocrinol. 352: 4–12.
PubMed Link

Jacklin, D. 1981. Methodological issues in the study of sex-related differences. Dev. Rev. 1: 266–273.

Jacobs, P. A. and J. A. Strong. 1959. A case of human intersexuality having a possible XXY sex-determining mechanism. Nature 183: 302–303.
PubMed Link

Jameson, S. A., Y-T. Lin, and B. Capel. 2012. Testis development requires  the repression of Wnt4 by Fgf signaling. Dev. Bio. 370: 24–32.
PubMed Link

Jeyasuria, P., W. M. Roosenburg and A. R. Place. 1994. Role of P-450 aromatase in sex determination of the diamondback terrapin, Malaclemys terrapin. J. Exp. Zool. 270: 95–111.
PubMed Link

Johnson, M. L., A. A. Nagengast and H. K. Salz. 2010. PPS, a large multidomain protein, functions with sex-lethal to regulate alternative splicing in Drosophila. PLoS Genet. 6(3): e1000872.
PubMed Link

Jordan-Young, R. M. 2010. Brainstorm: The Flaws in the Science of Sex Differences. Harvard University Press, Cambridge, MA.

Jost, A. 1947. Recerches sur la différenciation sexuelle de l’embryon de lapin. Arch. Anat. Microsc. Morphol. 36: 271–315.

Jost, A. 1953. Problems of fetal endocrinology: The gonadal and hypophyseal hormones. Recent Prog. Horm. Res. 8: 379–418.

Kandel, E. R., J. H. Schwartz and T. M. Jessell. 1995. Essentials of Neural Science and Behavior. Appleton and Lange, Norwalk, CT.

Kashimada, K. and P. Koopman. 2010. Sry: The master switch in mammalian sex determination. Development. 137: 3921–3930.
PubMed Link

Kashimada, K., E. Pelosi, H. Chen, D. Schlessinger, D. Wilhelm and P. Koopman. 2011. FOXL2 and BMP2 act cooperatively to regulate follistatin gene expression during ovarian development. Endocrinology 152: 272–280.
PubMed Link

Keyes, L. N., T. W. Cline and P. Schedl. 1992. The primary sex determination signal of Drosophila acts at the level of transcription. Cell 68: 933–943.
PubMed Link

Kim, S., V. J. Bardell and D. Zarkower. 2007. Cell type-autonomous and non-autonomous requirements for Dmrt1 in postnatal testis differentiation. Dev. Biol. 307: 314–325.
PubMed Link

Kimchi, T., J. Xu and C. Dulac. 2007. A functional circuit underlying male sexual behaviour in the female mouse brain. Nature 448: 1009–1014.
PubMed Link

Kimura, K. I., M. Ote, T. Tazawa and D. Yamamoto. 2005. Fruitless specifies sexually dimorphic neural circuitry in the Drosophila brain. Nature 438: 229–233.
PubMed Link

Koopman, P., A. Münsterberg, B. Capel, N. Vivian and A. Lovell-Badge. 1990. Expression of a candidate sex-determining gene during mouse testis differentiation. Nature 348: 450–452.
PubMed Link

Koopman, P., J. Gubbay, N. Vivian, P. Goodfellow and R. Lovell-Badge. 1991. Male development of chromosomally female mice transgenic for Sry. Nature 351: 117–121.
PubMed Link

Kudwa, A. E., C. Bodo, J.-A. Gustafsson and E. F. Rissman. 2005. A previously uncharacterized role for estrogen receptor b: Defeminization of male brain and behavior. Proc. Natl. Acad. Sci. USA 102: 4608–4612.
PubMed Link

Lahr, G., S. C. Maxson, A. Mayer, W. Just, C. Pilgrim and I. Reisert. 1995. Transcription of the Y chromosomal gene Sry in adult mouse brain. Brain Res: Mol. Brain Res. 33: 179–182.
PubMed Link

Langlois, V. S., A. C. Carew, B. D. Pauli, M. G. Wade, G. M. Cooke and V. L. Trudeau. 2010. Low levels of the herbicide atrazine alter sex ratios and reduce metamorphic success in Rana pipiens tadpoles raised in outdoor mesocosms. Environ. Health Perspect. 118: 552–557.
PubMed Link

Langman, J. 1981. Medical Embryology, 4th Ed. Williams & Wilkins, Baltimore.

Lau, Y. F. and Y. Li. 2009. The human and mouse sex-determining SRY genes repress the Rspol/b-catenin signaling. J. Genet. Genomics 36: 193–202.
PubMed Link

Lee, G., M. Foss, S. F. Goodwin, T. Carlo, B. J. Taylor and J. C. Hall. 2000. Spatial, temporal, and sexually dimorphic expression patterns of the fruitless gene in the Drosophila central nervous system. J. Neurobiol. 43: 404–426.
PubMed Link

Lee, Y. H., W. S. Yueh, J. L. Du, L. T. Sun and C. F. Chang. 2002. Aromatase inhibitors block natural sex change and induce male function in the protandrous black porgy, Acanthopagrus schlegeli Bleeker: Possible mechanism of natural sex change. Biol. Reprod. 66: 1749–1754.
PubMed Link

Lillie, F. R. 1917. The freemartin: A study of the actions of sex hormones in foetal life of cattle. J. Exp. Zool. 23: 371–452.

Maatouk, D. M., L. DiNapoli, A. Alvers, K. L. Parker, M. M. Taketo and B. Capel. 2008. Stabilization of b-catenin in XY gonads causes male-to-female sex-reversal. Hum. Mol. Genet. 17: 2949–2955.
PubMed Link

MacBride, S. M. and 7 others. 1999. Mushroom body ablation impairs short-term memory and long-term memory of courtship conditioning in Drosophila melanogaster. Neuron 24: 967–977.
PubMed Link

Margarit, E., M. D. Coll, R. Olivo, D. Gómez, A. Soler and F. Ballesta. 2000. SRY gene transferred to the long arm of the X chromosome in a Y-positive XX true hermaphrodite. Am. J. Med. Genet. 90: 25–28.
PubMed Link

Marx, J. 1995. Mammalian sex determination: Snaring the genes that divide sexes for mammals. Science 269: 1824–1825.
PubMed Link

Mathews, G. A. and A. P. Arnold. 1990. Antiestrogens fail to prevent the masculine ontogeny of zebra finch song system. Gen. Comp. Endocrin. 80: 48–58.
PubMed Link

Mayer, A., G. Lahr, D. F. Swaab, C. Pilgrim and I. Reisert. 1998. The Y-chromosomal genes SRY and ZFY are transcribed in adult human brain. Neurogenetics. 1: 281–288.
PubMed Link

Mayer, A., G. Mosler, W. Just, C. Pilgrim and I. Reisert. 2000. Developmental profile of Sry transcripts in the mouse brain. Neurogenetics 3: 25–30.
PubMed Link

Mazahery, A. R. and 7 others. 2013. Functional analysis of ectodermal b-catenin during external genitalia formation. Congen. Anom. 53: 34–41.
PubMed Link

McCarthy, M. M. and A. P. Arnold. 2011. Reframing sexual differentiation of the brain. Nature Neurosci. 14: 677–683.
PubMed Link

McClelland, K., J. Bowles and P. Koopman. 2012. Male sex determination: Insights into molecular mechanisms. Asian J. Androl. 14: 164–171.
PubMed Link

McEwen, B. S., I. Leiberburg, C. Chaptal and L. C. Krey. 1977. Aromatization: Important for sexual differentiation of the neonatal rat brain. Horm. Behav. 9: 249–263.
PubMed Link

McLaren, A. 1991. Development of the mammalian gonad: The fate of the supporting cell lineage. Bioessays 13: 151–156.
PubMed Link

McLaren, A. 1995. Germ cells and germ cell sex. Phil. Trans. Roy. Soc. Lond. B 350: 229–233.
PubMed Link

Meaney, M. J. and M. Szyf. 2005. Maternal care as a model for experience-dependent chromatin plasticity. Trends Neurosci. 28: 456–463.
PubMed Link

Merke, D. P., S. R. Bornstein, N. A. Avila and G. P. Chrousos. 2002. Future directions in the study and management of congenital adrenal hyperplasia due to 21-hydroxylase deficiency. Ann. Intern. Med. 136: 320–334.
PubMed Link

Meyer, W. J., B. R. Migeon and C. J. Migeon. 1975. Locus on human X chromosome for dihydrotestosterone receptor and androgen insensitivity. Proc. Natl. Acad. Sci. USA 72: 1469–1472.
PubMed Link

Migeon, C. J. and A. B. Wisniewski. 2000. Human sex differentiation: From transcription factors to gender. Horm. Res. 53: 111–119.
PubMed Link

Miyagawa, S. and 10 others. 2009. Genetic interactions of the androgen and Wnt/b-catenin pathways for the masculinization of external genitalia. Mol. Endocrinol. 23: 871–880.
PubMed Link

Morgan, T. H. and C. B. Bridges. 1919. The origin of gynandromorphs. In Contributions to the Study of Drosophila. Publication no. 278. Carnegie Institution of Washington, Washington, DC, pp. 1–122.

Mourabit, S., M. Edenbrow, D. P. Croft and T. Kudoh. 2011. Embryonic development of the self-fertilizing mangrove killifish Kryptolebias marmoratus. Dev. Dyn. 240: 1694–1704.
PubMed Link

Murdock, C. and T. Wibbels. 2006. Dmrt1 expression in response to estrogen treatment in a reptile with temperature-dependent sex determination. J. Exp. Zool. B Mol. Dev. Evol. 306: 134–139.
PubMed Link

Nagel, S. C. and F. S. vom Saal. 2003. Endocrine control of sexual differentiation: Effects of maternal-fetal environment and endocrine-disrupting chemicals. Adv. Mol. Cell Biol. 34: 15–38.

Nagoshi, R. N., M. McKeown, K. C. Burtis, J. M. Belote and B. S. Baker. 1988. The control of alternative splicing at genes regulating sexual differentiation in D. melanogaster. Cell 53: 229–236.
PubMed Link

Ngun, T. C., N. Ghahramani, F. J. Sanche, S. Bocklandt and E. Vilain. 2011. The genetics of sex differences in brain and behavior. Frontiers Neuroendocrinol. 32: 227–246.
PubMed Link

Ogawa, S. and 7 others. 2000. Abolition of male sexual behaviors in mice lacking estrogen receptors alpha and beta (ab-ERKO). Proc. Natl. Acad. Sci. USA 97: 14737–14741.
PubMed Link

Ottolenghi C. and 7 others. 2005. Foxl2 is required for commitment to ovary differentiation. Hum. Mol. Genet. 14: 2053–2062.
PubMed Link

Palmer, S. J. and S. Burgoyne. 1991. XY follicle cells in the ovaries of XO/XY and XO/XY/XY mosaic mice. Development 111: 1017–1019.
PubMed Link

Parma, P. and 8 others. 2006. R-spondin1 is essential in sex determination, skin differentiation and malignancy. Nature Genet. 38: 1304–1309.
PubMed Link

Pask, A. and J. A. Graves. 1999. Sex chromosomes and sex-determining genes: Insights from marsupials and monotremes. Cell. Mol. Life Sci. 55: 864–875.
PubMed Link

Pieau, C., N. Girondot, G. Richard-Mercier, M. Desvages, P. Dorizzi and P. Zaborski. 1994. Temperature sensitivity of sexual differentiation of gonads in the European pond turtle. J. Exp. Zool. 270: 86–93.

Pisarska, M. D., G. Barlow and F. T. Kuo. 2011. Minireview: Roles of the forkhead transcription factor FOXL2 in granulosa cell biology and pathology. Endocrinology. 152: 1199–1208.
PubMed Link

Pontiggia, A., R. Rimini, V. R. Harley, P. N. Goodfellow, R. Lovell-Badge and M. E. Bianchi. 1994. Sex-reversing mutations affect the architecture of Sry/DNA complexes. EMBO J. 13: 6115–6124.
PubMed Link

Qin, Y. and C. E. Bishop. 2005. Sox9 is sufficient for functional testis development producing fertile male mice in the absence of Sry. Hum. Mol. Genet. 14: 1221–1229.
PubMed Link

Radder, R. S., A. E. Quinn, A. Georges, S. D. Sarre and R. Shine. 2008. Genetic evidence for co-occurrence of chromosomal and thermal sex-determining systems in a lizard. Biol. Lett. 4: 176–178.
PubMed Link

Reddy, V. R., F. Naftolin and K. J. Ryan. 1974. Conversion of androstenedione to estrone by neural tissues from fetal and neonatal rats. Endocrinology 94: 117–121.
PubMed Link

Rhen, T. and J. W. Lang. 1994. Temperature-dependent sex determination in the snapping turtle: Manipulation of the embryonic sex steroid environment. Gen. Comp. Endocrinol. 96: 243–254.
PubMed Link

Roberts, L. M., J. A. Visser and H. A. Ingraham. 2002. Involvement of a matrix metalloproteinase in MIS-induced cell death during urogenital development. Development 129: 1487–1496.
PubMed Link

Roberts, L. M., Y. Hirokawa, M. W. Nachtigal and H. A. Ingraham. 1999. Paracrine-mediated apoptosis in reproductive tract development. Dev. Biol. 208: 110–122.
PubMed Link

Ruta V., S. R. Datta, M. L. Vasconcelos, J. Freeland, L. L. Looger and R. Axel. 2010. A dimorphic pheromone circuit in Drosophila from sensory input to descending output. Nature 468: 686–690.
PubMed Link

Ryner, L. C. and B. S. Baker. 1991. Regulation of doublesex pre-mRNA processing occurs by 3′ splice site activation. Genes Dev. 5: 2071–2085.
PubMed Link

Salz, H. K. 2011. Sex determination in insects: A binary decision based on alternative splicing. Curr. Opin. Genet. Dev. 21: 395–400.
PubMed Link

Sanderson, J. T., W. Seinen, J. P. Giesy and M. van den Berg. 2000. 2-Chloro-s-triazine herbicides induce aromatase (CYP19) activity in H295 human adrenocortical carcinoma cells: A novel mechanism of estrogenicity? Toxicol. Sci. 54: 121–127.

Sariola, H. and M. Saarma. 1999. GDNF and its receptors in the regulation of the ureteric branching. Int. J. Dev. Biol. 43: 413–418.
PubMed Link

Schiebinger, L. 1989. The Mind Has No Sex? Harvard University Press, Cambridge, MA.

Schiebinger, L. 1993. Nature’s Body: Gender in the Making of Modern Science. Beacon Press, Boston.

Schmahl, J., Y. Kim, J. S. Colvin, D. M. Ornitz and B. Capel. 2004. Fgf9 induces proliferation and nuclear localization of FgfR2 in Sertoli precursors during male sex determination. Development 131: 3627–3636.
PubMed Link

Sefton, L., J. R. Timmer, Y. Zhang, F. Béranger and T. W. Cline. 2000. An extracellular activator of the Drosophila JAK/STAT pathway is a sex-determination signal element. Nature 405: 970–973.
PubMed Link

Sekido, R. and R. Lovell-Badge 2009. Sex determination and SRY: Down to a wink and a nudge? Trends Genet. 25: 19–29.
PubMed Link

Sekido, R. and R. Lovell-Badge. 2008. Sex determination involves synergistic action of Sry and Sf1 on a specific Sox9 enhancer. Nature 453: 930–934.
PubMed Link

Sekido, R., I. Bar, V. Narvárez, G. Penny and R. Lovell-Badge. 2004. SOX9 is up-regulated by the transient expression of SRY specifically in Sertoli cell precursors. Dev. Biol. 274: 271–279.
PubMed Link

Shen, W.-H., C. C. D. Moore, Y. Ikeda, K. L. Parker and H. A. Ingraham. 1994. Nuclear receptor steroidogenic factor 1 regulates the Müllerian-inhibiting substance gene: A link to the sex determination cascade. Cell 77: 651–661.
PubMed Link

Shoemaker, C., M. Ramsey, J. Queen and D. Crew. 2007. Expression of Sox9, Mis, and Dmrt1 in the gonad of a species with temperature-dependent sex determination. Dev. Dyn. 236: 1055–1063.
PubMed Link

Siegel, R. W. and J. C. Hall. 1979. Conditioned responses in courtship behavior of normal and mutant Drosophila. Proc. Natl. Acad. Sci. USA 76: 3430–3434.
PubMed Link

Siiteri, P. K. and J. D. Wilson. 1974. Testosterone formation and metabolism during male sexual differentiation in the human embryo. J. Clin. Endocrinol. Metab. 38: 113–125.
PubMed Link

Sinclair, A. H. and 9 others. 1990. A gene from the human sex-determining region encodes a protein with homology to a conserved DNA-binding motif. Nature 346: 240–244.
PubMed Link

Smith, C. A. and A. H. Sinclair. 2001. Sex determination in the chicken embryo. J. Exp. Zool. 290: 691–699.
PubMed Link

Smith, C. A., C. M. Shoemaker, K. N. Roeszler, J. Queen, D. Crews and A. H. Sinclair. 2008. Cloning and expression of R-Spondin1 in different vertebrates suggests a conserved role in ovarian development. BMC Dev. Biol. 8: 72.
PubMed Link

Smith, C. A., K. N. Roeszler, T. Ohnesorg, D. M. Cummins, P. G. Farlie, T. J. Doran and A. H. Sinclair. 2009. The avian Z-linked gene DMRT1 is required for male sex determination in the chicken. Nature 461: 267–271.
PubMed Link

Stevens, N. M. 1905. Studies in Spermatogenesis with Especial Reference to the “Accessory Chromosome.” Publication 36. Carnegie Institution of Washington, Washington, DC.

Stockinger, P. D. Kvitsiani, S. Rotkopf, L. Tirián and B. J. Dickson. 2005. Neural circuitry that governs Drosophila male courtship behavior. Cell 121: 795–807.
PubMed Link

Thigpen, A. E., D. L. Davis, T. Gautier, J. Imperato-McGinley and D. W Russell. 1992. The molecular basis of steroid 5a-reductase deficiency in a large Dominican kindred. N. Engl. J. Med. 327: 1216–1219.
PubMed Link

Tran, D., N. Meusy-Dessolle and N. Josso. 1977. Anti-Müllerian hormone is a functional marker of foetal Sertoli cells. Nature 269: 411–412.
PubMed Link

Trelstad, R. L., A. Hayashi, K. Hayashi and P. K. Donahoe. 1982. The epithelial-mesenchymal interface of the male rate Müllerian duct: Loss of basement membrane integrity and ductal regression. Dev. Biol. 92: 27–40.
PubMed Link

Vainio, S., M. Heikkilä, A. Kispert, A. Chin and A. P. McMahon. 1999. Female development in mammals is regulated by Wnt4 signalling. Nature 397: 405–409.
PubMed Link

Valcarcel, J., R. Singh, P. D. Zamore and M. R. Green. 1993. The protein Sex-lethal antagonizes the splicing factor U2AF to regulate alternative splicing of transformer pre-mRNA. Nature 362: 171–175.
PubMed Link

Valenzuela, N., J. Neuwald and R. Literman. 2013. Transcriptional evolution underlying vertebrate sexual development. Dev. Dyn. 242: 307–319.
PubMed Link

Vidal, V. P. I., M.-C. Chaboissier, D. G. de Rooij and A. Schedl. 2001. Sox9 induces testis development in XX transgenic mice. Nature Genet. 28: 216–217.
PubMed Link

Vrontou, E., S. P. Nilsen, E. Demir, E. A. Kravitz and B. J. Dickson. 2006. fruitless regulates aggression and dominance in Drosophila. Nature Neurosci. 9: 1469–1471.
PubMed Link

Wang, L. and D. J. Anderson. 2010. Identification of an aggression-promoting pheremone and its receptor neurons in Drosophila. Nature 463: 227–231.
PubMed Link

Warner, R. R., D. L. Fitch and J. D. Standish. 1996. Social control of sex change in the shelf limpet Crepidula norrisiarum: Size-specific responses to local group composition. J. Exp. Marine Biol. Ecol. 204: 155–167.

Washburn. L. L. and E. M. Eicher. 1989. Normal testis determination in the mouse depends on genetic interaction of a locus on chromosome 17 and the Y chromosome. Genetics 123: 173–179.
PubMed Link

Werner, M. H., J. R. Huth, A. M. Groneborn and G. M. Clore. 1995. Molecular basis of human 46X,Y sex reversal revealed from the three-dimensional solution structure of the human SRY–DNA complex. Cell 81: 705–714.
PubMed Link

Wibbels, T. and, D. Crews. 1994. Putative aromatase inhibitor induces male sex determination in a female unisexual lizard and in a turtle with temperature-dependent sex determination. J. Endocrinol. 141: 295–299.
PubMed Link

Wilhelm, D., L. L. Washburn, V. Truong, M. Fellous, E. M. Eicher and P. Koopman. 2009. Antagonism of the testis- and ovary-determining pathways during ovotestis development in mice. Mech. Dev. 126: 324–336.
PubMed Link

Wilhelm, D., L. L. Washburn, V. Truonga, M. Fellous, E. M. Eicher and P. Koopman. 2009. Antagonism of the testis- and ovary-determining pathways during ovotestis development in mice. Mech. Dev. 126: 324–336.
PubMed Link

Willerton, L., R. A. Smith, D. Russell and S. Mackay. 2004 Effects of FGF9 on embryonic Sertoli cell proliferation and testicular cord formation in the mouse. Int. J. Dev. Biol. 48: 637–643.
PubMed Link

Wilson, E. B. 1905. The chromosomes in relation to the determination of sex in insects. Science 22: 500–502.

Yao, H. H. and 6 others. 2004. Follistatin operates downstream of Wnt4 in mammalian ovary organogenesis. Dev. Dyn. 230: 210–215.
PubMed Link

Yu, J. Y., M. I. Kanai, E. Demir, G. S. Jefferis and B. J. Dickson. 2010. Cellular organization of the neural circuit that drives Drosophila courtship behavior. Curr. Biol. 20: 1602–1614.
PubMed Link

Zhao, D. and 8 others. 2010. Somatic sex identity is cell autonomous in the chicken. Nature 464: 237–242.
PubMed Link

 

Citation for the BioWeb Conference Links
Barresi, M. J., and Lee, K. 2005–2010. Biology Web Conferences. Smith College, Northampton, MA.

© All the material on this website is protected by copyright. It may not be reproduced in any form without permission from the copyright holder.
Home Link