Reviews in Fish Biology and Fisheries

, Volume 20, Issue 1, pp 101–121 | Cite as

Sexual determination and differentiation in teleost fish



The present work reviews the latest information on the cellular, molecular and physiological aspects of sexual determination and differentiation in teleost fish. The group exhibits a large variety of mechanisms of sexual determination. These may be genetic, or depend on environmental conditions such as temperature, pH, and social factors, all of which can influence the proportion of the sexes. Additionally, sex steroids play an important role in the regulation of sexual differentiation. The patterns of gonadal sexual differentiation are diverse, and species may be hermaphroditic or gonochoristic, some of the latter displaying juvenile hermaphroditism. In recent years, several genes involved in the sexual determination and differentiation pathways in vertebrates, particularly in mammals, have also been characterized in teleosts. Conserved as well as diversified functions have been proposed.


Sexual determination Sexual differentiation Gonadal sexual differentiation Sex determining genes Teleosts 



The authors thank Isabel Pérez Montfort for English revision. This work was supported by grants from SEP—CONACyT (Sol. 81269) and PAPIIT of DGAPA, UNAM, No. IN218908.


  1. Alfaqih MA, Brunelli JP, Drew RE, Thorgaard GH (2009) Mapping of five candidate sex-determining loci in rainbow trout (Oncorhynchus mykiss). BMC Genet 10:2PubMedGoogle Scholar
  2. Almeida-Toledo LF, Foresti F, Daniel MF, Toledo-Filho SA (2000) Sex chromosome evolution in fish: the formation of the neo-Y chromosome in Eigenmannia (Gymnotiformes). Chromosoma 109(3):197–200PubMedGoogle Scholar
  3. Artieri CG, Mitchell LA, Ng SH, Parisotto SE, Danzmann RG, Hoyheim B, Phillips RB, Morasch M, Koop BF, Davidson WS (2006) Identification of the sex-determining locus of Atlantic salmon (Salmo salar) on chromosome 2. Cytogenet Genome Res 112(1–2):152–159PubMedGoogle Scholar
  4. Asoh K, Shapiro DY (1997) Bisexual juvenile gonad and gonochorism in the fairy basslet, Gramma loreto. Copeia 1997:22–31Google Scholar
  5. Baras E, Prignon C, Gohoungo G, Melard C (2000) Phenotypic sex differentiation of blue tilapia under constant and fluctuating thermal regimes and its adaptive and evolutionary implications. J Fish Biol 57:210–223Google Scholar
  6. Baroiller JF, D’Cotta H (2001) Environment and sex determination in farmed fish. Comp Biochem Physiol C Toxicol Pharmacol 30(4):399–409Google Scholar
  7. Baroiller JF, Guiguen Y (2001) Endocrine and environmental aspects of sex differentiation in gonochoristic fish. EXS 91:177–201PubMedGoogle Scholar
  8. Baroiller JF, Guiguen Y, Fostier A (1999) Endocrine and environmental aspects of sex differentiation in fish. Cell Mol Life Sci 55:910–931Google Scholar
  9. Baron D, Guiguen Y (2003) Gene expression during gonadal sex differentiation in rainbow trout (Oncorhynchus mykiss): from candidates genes studies to high throughout genomic approach. Fish Physiol Biochem 28:119–123Google Scholar
  10. Baron B, Buckle F, Espina S (2002) Environmental factors and sexual differentiation in Poecilia sphenops (Pisces: Poeciliidae). Aquac Res 33:615–619Google Scholar
  11. Baron D, Cocquet J, Xia X, Fellous M, Guiguen Y, Veitia RA (2004) An evolutionary and functional analysis of Foxl2 in rainbow trout gonad differentiation. J Mol Endocrinol 33(3):705–715PubMedGoogle Scholar
  12. Baron D, Houlgatte R, Fostier A, Guiguen Y (2005a) Large-scale temporal gene expression profiling during gonadal differentiation and early gametogenesis in rainbow trout. Biol Reprod 73:959–966PubMedGoogle Scholar
  13. Baron D, Fostier A, Brenton B, Guiguen Y (2005b) Androgen and estrogen treatment alter steady state messenger (mRNA) levels of testicular steroidogenic enzymes in the rainbow trout, Oncorhynchus mykiss. Mol Reprod Dev 71:471–479PubMedGoogle Scholar
  14. Bertollo LAC, Fontes MS, Fenocchio AS, Cano J (1997) The X1X2Y sex chromosomes system in the fish Hoplias malabaricus I. G-, C and chromosome replication banding. Chromosome Res 5:493–499PubMedGoogle Scholar
  15. Beullens K, Eding EH, Gilson P, Ollevier F, Komen J, Richter CJJ (1997) Gonadal differentiation, intersexuality and sex ratios of European eel (Anguilla anguilla L.) maintained in captivity. Aquaculture 153:135–150Google Scholar
  16. Blaser H, Eisenbeiss S, Neumann M, Reichman-Fried M, Thisse B, Thisse C, Raz E (2005) Transition from non-motile behaviour to directed migration during early PGC development in zebrafish. J Cell Sci 118:4027–4038PubMedGoogle Scholar
  17. Blázquez M, Piferrer F (2004) Cloning, sequence analysis, tissue distribution, and sex-specific expression of the neural form of P450 aromatase in juvenile sea bass (Dicentrarchus labrax). Mol Cell Endocrinol 219(1–2):83–94PubMedGoogle Scholar
  18. Blázquez M, Zanuy S, Carillo M, Piferrer F (1998) Structural and functional effects of early exposure to estradiol-17 beta and 17 alpha-ethynylestradiol on the gonads of the gonochoristic teleost Dicentrarchus labrax. Fish Physiol Biochem 18:37–47Google Scholar
  19. Bogart MH (1987) Sex determination: a hypothesis based on steroid ratios. J Theor Biol 128(3):349–357PubMedGoogle Scholar
  20. Bollig F, Mehringer R, Perner B, Hartung C, Schäfer M, Schartl M, Volff JN, Winkler C, Englert C (2006) Identification and comparative expression analysis of a second wt1 gene in zebrafish. Dev Dyn 235(2):554–561PubMedGoogle Scholar
  21. Bull JJ (1985) Sex determining mechanisms: an evolutionary perspective. Experientia 41:1285–1296PubMedGoogle Scholar
  22. Callard GV, Tchoudakova AV, Kishida M, Wood E (2001) Differential tissue distribution, developmental programming, estrogen regulation and promoter characteristics of Cyp19 genes in teleost fish. J Steroid Biochem Mol Biol 79(1–5):305–314PubMedGoogle Scholar
  23. Chai C, Chan WK (2000) Developmental expression of a novel Ftz-F1 homologue, ff1b (NR5A4), in the zebrafish Danio rerio. Mech Dev 91:421–426PubMedGoogle Scholar
  24. Chang CF, Lin BY, Lau EL, Lee MF, Yueh WS, Lee YH, Chang CN, Huang JD, Tacon P, Lee FY, Du JL, Sun LT (1997) The endocrine mechanism of sex reversal in the protandrous black porgy, Acanthopagrus schlegeli: a review. Chin J Physiol 40(4):197–205PubMedGoogle Scholar
  25. Chang X, Kobayashi T, Senthilkumaran B, Kobayashi-Kajura H, Sudhakumari CC, Nagahama Y (2005) Two types of aromatase with different encoding genes, tissue distribution and developmental expression in Nile tilapia (Oreochromis niloticus). Gen Comp Endocrinol 141(2):101–115PubMedGoogle Scholar
  26. Chen TR, Reisman HM (1970) A comparative chromosome study of the North American species of sticklebacks (Teleostei: Gasterosteidae). Cytogenetics 9:321–332PubMedGoogle Scholar
  27. Chen J, Fu Y, Xiang D, Zhao G, Long H, Liu J, Yu Q (2008) XX/XY heteromorphic sex chromosome systems in two bullhead catfish species, Liobagrus marginatus and L. styani (Amblycipitidae, Siluriformes). Cytogenet Genome Res 122(2):169–174PubMedGoogle Scholar
  28. Chiang EF, Pai CI, Wyatt M, Yan YL, Postlethwait J, Chung B (2001) Two Sox9 genes on duplicated zebrafish chromosomes: expression of similar transcription activators in distinct sites. Dev Biol 231:149–163PubMedGoogle Scholar
  29. Cocquet J, Pailhoux E, Jaubert F, Servel N, Xia X, Pannetier M, De Baere E, Messiaen L, Cotinot C, Fellous M, Veitia RA (2002) Evolution and expression of FOXL2. J Med Genet 39(12):916–921PubMedGoogle Scholar
  30. Colombo G, Grandi G (1995) Sex differentiation in the European eel: histological analysis of the effects of sex steroids on the gonad. J Fish Biol 47:394–413Google Scholar
  31. Conover DO (1984) Adaptive significance of temperature-dependent sex determination in a fish. Am Nat 123:297–313Google Scholar
  32. Conover DO, Daemond SB (1991) Absence of temperature-dependent sex determination in northern populations of two cyprinodontid fishes. Can J Zool 69:530–533Google Scholar
  33. Conover DO, Kynard BE (1981) Environmental sex determination: interaction of temperature and genotype in a fish. Science 213:577–579PubMedGoogle Scholar
  34. Crews D (1996) Temperature-dependent sex determination: the interplay of steroid hormones and temperature. Zool Sci 3(1):1–13Google Scholar
  35. Crews D, Bergeron JM (1994) Role of reductase and aromatase in sex determination in the red-eared slider (Trachemys scripta), a turtle with temperature-dependent sex determination. J Endocrinol 143(2):279–289PubMedGoogle Scholar
  36. D′Cotta H, Fostier A, Guiguen Y, Govoroun M, Baroiller JF (2001) Aromatase plays a key role during normal and temperature induced sex differentiation of tilapia Oreochromis niloticus. Mol Reprod Dev 59:265–276PubMedGoogle Scholar
  37. Desprez D, Melard C (1998) Effect of ambient water temperature on sex determinism in the blue tilapia Oreochromis aureus. Aquaculture 162:1–2Google Scholar
  38. Devlin RH, Nagahama Y (2002) Sex determination and sex differentiation in fish: an overview of genetic, physiological, and environmental influences. Aquaculture 208:191–364Google Scholar
  39. Du QY, Wang FY, Hua HY, Chang ZJ (2007) Cloning and study of adult-tissue-specific expression of Sox9 in Cyprinus carpio. J Genet 86(2):85–91PubMedGoogle Scholar
  40. Fenske M, Segner H (2004) Aromatase modulation alters gonadal differentiation in developing zebrafish (Danio rerio). Aquat Toxicol 67:105–126PubMedGoogle Scholar
  41. Filby AL, Thorpe KL, Maack G, Tyler CR (2007) Gene expression profiles revealing the mechanisms of anti-androgen- and estrogen-induced feminization in fish. Aquat Toxicol 81(2):219–231PubMedGoogle Scholar
  42. Francis RC, Barlow GW (1993) Social control of primary sex differentiation in the Midas cichlid. Proc Natl Acad Sci USA 90:10673–10675PubMedGoogle Scholar
  43. Führer D (2002) Lessons from studies of complex genetic disorders: identification of FOXL2-a novel transcription factor on the wing to fertility. Eur J Endocrinol 146(1):15–18PubMedGoogle Scholar
  44. Giuili G, Shen WH, Ingraham HA (1997) The nuclear receptor SF-1 mediates sexually dimorphic expression of Müllerian inhibiting substance, in vivo. Development 124:1799–1807PubMedGoogle Scholar
  45. Godwin J (2009) Social determination of sex in reef fishes. Semin Cell Dev Biol 20(3):264–270PubMedGoogle Scholar
  46. Govoroun M, McMeek OM, D′Cotta H, Ricordel MJ, Smith T, Fostier A, Guiguen Y (2001) Steroid enzyme gene expression during natural and androgen-induced gonadal differentiation in the rainbow trout, Oncorhynchus mykiss. J Exp Zool 290:558–566PubMedGoogle Scholar
  47. Govoroun MS, Pannetier M, Pailhoux E, Cocquet J, Brillard JP, Couty I, Batellier F, Cotinot C (2004) Isolation of chicken homolog of the FOXL2 gene and comparison of its expression patterns with those of aromatase during ovarian development. Dev Dyn 231(4):859–870PubMedGoogle Scholar
  48. Grandi G, Chicca M (2008) Histological and ultrastructural investigation of early gonad development and sex differentiation in Adriatic sturgeon (Acipenser naccarii, Acipenseriformes, Chondrostei). J Morphol 269(10):1238–1262PubMedGoogle Scholar
  49. Grier H (2000) Ovarian germinal epithelium and folliculogenesis in the common snook, Centropomus undecimalis (Teleostei: centropomidae). J Morphol 243(3):265–281PubMedGoogle Scholar
  50. Guan G, Kobayashi T, Nagahama Y (2000) Sexually dimorphic expression of two types of DM (Doublesex/Mab-3)-domain genes in a teleost fish, the Tilapia (Oreochromis niloticus). Biochem Biophys Res Commun 272:662–666PubMedGoogle Scholar
  51. Guiguen Y, Baroiller JF, Ricordel MJ, Iseki K, Mcmeel OM, Martin SA, Fostier A (1999) Involvement of estrogens in the process of sex differentiation in two fish species: the rainbow trout (Oncorhynchus mykiss) and a tilapia (Oreochromis niloticus). Mol Reprod Dev 54(2):154–162PubMedGoogle Scholar
  52. Guo Y, Cheng H, Huang X, Gao S, Yu H, Zhou R (2005) Gene structure, multiple alternative splicing, and expression in gonads of zebrafish Dmrt1. Biochem Biophys Res Commun 330:950–957PubMedGoogle Scholar
  53. Hamaguchi S (1982) A light- and electron-microscopic study on the migration of primordial germ cells in the teleost, Oryzias latipes. Cell Tissue Res 227:139–151PubMedGoogle Scholar
  54. Harrington RW, Crossman RA (1976) Effects of temperature and sex genotype on meristic counts of the gonochoristic cyprinodontid fish Rivulus cylindraceus Poey. Can J Zool 54:245–254PubMedGoogle Scholar
  55. Hayes TB (1998) Sex determination and primary sex differentiation in amphibian: genetic and developmental mechanisms. J Exp Zool 281:373–399PubMedGoogle Scholar
  56. Herpin A, Rohr S, Riedel D, Kluever N, Raz E, Schartl M (2007) Specification of primordial germ cells in medaka (Oryzias latipes). BMC Dev Biol 7:3PubMedGoogle Scholar
  57. Herpin A, Fischer P, Liedtke D, Kluever N, Neuner C, Raz E, Schartl M (2008) Sequential SDF1a and b-induced mobility guides Medaka PGC migration. Dev Biol 320(2):319–327PubMedGoogle Scholar
  58. Higa M, Kanda H, Kitahashi T, Ito M, Shiba T, Ando H (2000) Quantitative analysis of fushi tarazu factor 1 homolog messenger ribonucleic acids in the pituitary of salmon at different prespawning stages. Biol Reprod 63:1756–1763PubMedGoogle Scholar
  59. Hoar WS (1969) Reproduction. In: Hoar WS, Randall DJ (eds) Fish physiology, vol 3. Academic Press, New York, pp 1–72Google Scholar
  60. Hobbs JP, Munday PL, Jones GP (2004) Social induction of maturation and sex determination in a coral reef fish. Proc Biol Sci 271(1553):2109–2114PubMedGoogle Scholar
  61. Hossain MS, Larsson A, Scherbak N, Olsson PE, Orban L (2008) Zebrafish androgen receptor: isolation, molecular, and biochemical characterization. Biol Reprod 78(2):361–369PubMedGoogle Scholar
  62. Ijiri S, Kaneko H, Kobayashi T, Wang DS, Sakai F, Paul-Prasanth B, Nakamura M, Nagahama Y (2008) Sexual dimorphic expression of genes in gonads during early differentiation of a teleost fish, the Nile tilapia Oreochromis niloticus. Biol Reprod 78(2):333–341PubMedGoogle Scholar
  63. Iwata E, Nagai Y, Hyoudou M, Sasaki H (2008) Social environment and sex differentiation in the false clown anemonefish, Amphiprion ocellaris. Zool Sci 25(2):123–128PubMedGoogle Scholar
  64. Jeong HB, Park JG, Park YJ, Takemura A, Hur SP, Lee YD, Kim SJ (2009) Isolation and characterization of DMRT1 and its putative regulatory region in the protogynous wrasse, Halichoeres tenuispinis. Gene 438(1–2):8–16PubMedGoogle Scholar
  65. Johnson AD, Drum M, Bachvarova RF, Masi T, White ME, Crother BI (2003) Evolution of predetermined germ cells in vertebrate embryos: implications for macroevolution. Evol Dev 5(4):414–431PubMedGoogle Scholar
  66. Johnston CM, Barnett M, Sharpe PT (1995) The molecular biology of temperature-dependent sex determination. Philos Trans R Soc Lond B Biol Sci 350(1333):297–303PubMedGoogle Scholar
  67. Josso N, Racine C, di Clemente N, Rey R, Xavier F (1998) The role of anti-Mullerian hormone in gonadal development. Moll Cell Endocrinol 145:3–7Google Scholar
  68. Karube M, Fernandino JI, Strobl-Mazzulla P, Strüssmann CA, Yoshizaki G, Somoza GM, Patiño R (2007) Characterization and expression profile of the ovarian cytochrome P-450 aromatase (cyp19A1) gene during thermolabile sex determination in pejerrey, Odontesthes bonariensis. J Exp Zool Part A Ecol Genet Physiol 307(11):625–636Google Scholar
  69. Kent J, Wheatley SC, Andrews JE, Sinclair AH, Koopman P (1996) A male-specific role for SOX9 in vertebrate sex determination. Development 122:2813–2822PubMedGoogle Scholar
  70. Kitano T, Takamune K, Kobayashi T, Nagahama Y, Abe SI (1999) Suppression of P450 aromatase gene expression in sex reversed males produced by rearing genetically female larvae at a high water temperature during a period of sex differentiation in the Japanese flounder (Paralichthys olivaceus). J Mol Endocrinol 23:167–176PubMedGoogle Scholar
  71. Kitano T, Takamune K, Nagahama Y (2000) Aromatase inhibitor and 17α-methyltestosterone cause sex reversal from genetic females to phenotypic males and suppression of P450 aromatase gene expression in Japanese flounder (Paralichthys olivaceus). Mol Reprod Dev 56:1–5PubMedGoogle Scholar
  72. Kitano T, Yoshinaga N, Shiraishi E, Koyanagi T, Abe S (2007) Tamoxifen induces masculinization of genetic females and regulates P450 aromatase and Müllerian inhibiting substance mRNA expression in Japanese flounder (Paralichthys olivaceus). Mol Reprod Dev 74(9):1171–1177PubMedGoogle Scholar
  73. Klüver N, Herpin A, Braasch I, Driessle J, Schartl M (2009) Regulatory back-up circuit of medaka Wt1 co-orthologs ensures PGC maintenance. Dev Biol 325(1):179–188PubMedGoogle Scholar
  74. Knaut H, Pelegri F, Bohmann K, Schwarz H, Nusslein-Volhard C (2000) Zebrafish vasa RNA but not its protein is a component of the germ plasm and segregates asymmetrically before germline specification. J Cell Biol 149:875–888PubMedGoogle Scholar
  75. Knaut H, Werz C, Geisler R, Nüsslein-Volhard C, Tübingen Screen Consortium (2003) A zebrafish homologue of the chemokine receptor Cxcr4 is a germ-cell guidance receptor. Nature 421(6920):279–282PubMedGoogle Scholar
  76. Kobayashi T, Kajiura-Kobayashi H, Nagahama Y (2003) Induction of XY sex reversal by estrogen involves altered gene expression in a teleost, tilapia. Cytogenet Genome Res 101:289–294PubMedGoogle Scholar
  77. Kobayashi T, Matsuda M, Kajiura-Kobayashi H, Suzuki A, Saito N, Nakamoto M, Shibata N, Nagahama Y (2004) Two DM domain genes, DMY and DMRT1, involved in testicular differentiation and development in the medaka, Oryzias latipes. Dev Dyn 231(3):518–526PubMedGoogle Scholar
  78. Komen J, de Boer P, Richter CJJ (1992) Male sex reversal in gynogenetic XX females of common carp Cyprinus carpio L. by a recessive mutation in a sex-determining gene. J Hered 83:431–434Google Scholar
  79. Kondo M, Nanda I, Hornung U, Asakawa S, Shimizu N, Mitani H, Schmid M, Shima A, Schartl M (2003) Absence of the candidate male sex-determining gene dmrt1b(Y) of medaka from other fish species. Curr Biol 13:416–420PubMedGoogle Scholar
  80. Kondo M, Nanda I, Hornung U, Schmid M, Schartl M (2004) Evolutionary origin of the medaka Y chromosome. Curr Biol 14:1664–1669PubMedGoogle Scholar
  81. Kondo M, Hornung U, Nanda I, Imai S, Sasaki T, Shimizu A, Asakawa S, Hori H, Schmid M, Shimizu N, Schartl M (2006) Genomic organization of the sex-determining and adjacent regions of the sex chromosomes of medaka. Genome Res 16(7):815–826PubMedGoogle Scholar
  82. Koopman P (2001) The genetics and biology of vertebrate sex determination. Cell 105:843–847PubMedGoogle Scholar
  83. Koopman P, Münsterberg A, Capel B, Vivian N, Lovell-Badge R (1990) Expression of a candidate sex determining gene during mouse testis differentiation. Nature 348:450–452PubMedGoogle Scholar
  84. Koprunner M, Thisse C, Thisse B, Raz E (2001) A zebrafish nanos-related gene is essential for the development of primordial germ cells. Genes Dev 15:2877–2885PubMedGoogle Scholar
  85. Koshimizu E, Strüssmann CA, Okamoto N, Fukuda H, Sakamoto T (2009) Construction of a genetic map and development of DNA markers linked to the sex-determining locus in the Patagonian Pejerrey (Odontesthes hatcheri). Mar Biotechnol (NY). doi: 10.1007/s10126-009-9194-1
  86. Koya Y, Fujita A, Niki F, Ishihara E (2003) Sex differentiation and pubertal development of gonads in the viviparous mosquitofish, Gambusia affinis. Zool Sci 20:1231–1242PubMedGoogle Scholar
  87. Kurokawa H, Saito D, Nakamura S, Katoh-Fukui Y, Ohta K, Baba T, Morohashi K, Tanaka M (2007) Germ cells are essential for sexual dimorphism in the medaka gonad. Proc Natl Acad Sci USA 104(43):16958–16963PubMedGoogle Scholar
  88. Kwon JY, McAndrew BJ, Penman DJ (2001) Cloning of brain aromatase gene and expression of brain and ovarian aromatase genes during sexual differentiation in genetic male and female Nile tilapia Oreochromis niloticus. Mol Reprod Dev 59(4):359–370PubMedGoogle Scholar
  89. Larsson SH, Charlieu JP, Miyagawa K, Engelkamp D, Rassoulzadegan M, Ross A, Cuzin F, van Heyningen V, Hastie ND (1995) Subnuclear localization of WT1 in splicing or transcription factor domains is regulated by alternative splicing. Cell 81:391–401PubMedGoogle Scholar
  90. Lee BY, Kocher TD (2007) Exclusion of Wilms tumour (WT1b) and ovarian cytochrome P450 aromatase (CYP19A1) as candidates for sex determination genes in Nile tilapia (Oreochromis niloticus). Anim Genet 38(1):85–86PubMedGoogle Scholar
  91. Lewis ZR, McClellan MC, Postlethwait JH, Cresko WA, Kaplan RH (2008) Female-specific increase in primordial germ cells marks sex differentiation in threespine stickleback (Gasterosteus aculeatus). J Morphol 269(8):909–921PubMedGoogle Scholar
  92. Li JZ, Zhou YP, Zhen Y, Xu Y, Cheng PX, Wang HN, Deng FJ (2009) Cloning and characterization of the SSB-1 and SSB-4 genes expressed in zebrafish gonads. Biochem Genet 47(3–4):179–190PubMedGoogle Scholar
  93. Liu Z, Wu F, Jiao B, Zhang X, Hu C, Huang B, Zhou L, Huang X, Wang Z, Zhang Y, Nagahama Y, Cheng CH, Wang D (2007) Molecular cloning of doublesex and mab-3-related transcription factor 1, forkhead transcription factor gene 2, and two types of cytochrome P450 aromatase in Southern catfish and their possible roles in sex differentiation. J Endocrinol 194(1):223–241PubMedGoogle Scholar
  94. Loffler KA, Zarkower D, Koopman P (2003) Etiology of ovarian failure in blepharophimosis ptosis epicanthus inversus syndrome: FOXL2 is a conserved, early-acting gene in vertebrate ovarian development. Endocrinology 144(7):3237–3243PubMedGoogle Scholar
  95. Maack G, Segner H (2003) Morphological development of the gonads in zebrafish. J Fish Physiol 62:895–906Google Scholar
  96. Mair GC, Scott AG, Penman DJ, Skibinski DOF, Beardmore JA (1991) Sex determination in the genus Oreochromis 2. Sex reversal, hybridisation, gynogenesis and triploidy in O. aureus Steindachner. Theor Appl Genet 82:153–160Google Scholar
  97. Marchand O, Govoroun M, D’Cotta H, McMeel O, Lareyre J, Bernot A, Laudet V, Guiguen Y (2000) DMRT1 expression during gonadal differentiation and spermatogenesis in the rainbow trout, Oncorhynchus mykiss. Biochem Biophys Acta 1493:180–187PubMedGoogle Scholar
  98. Martins RS, Deloffre LA, Mylonas CC, Power DM, Canário AV (2007) Developmental expression of DAX1 in the European sea bass, Dicentrarchus labrax: lack of evidence for sexual dimorphism during sex differentiation. Reprod Biol Endocrinol 5:19PubMedGoogle Scholar
  99. Matsuda M (2005) Sex determination in the teleost medaka, Oryzias latipes. Annu Rev Genet 39:293–307PubMedGoogle Scholar
  100. Matsuda M, Matsuda C, Hamaguchi S, Sakaizumi M (1998) Identification of the sex chromosomes of the medaka, Oryzias latipes, by fluorescence in situ hybridization. Cytogenet Cell Genet 82:257–262PubMedGoogle Scholar
  101. Matsuda M, Nagahama Y, Shinomiya A, Sato T, Matsuda C, Kobayashi T, Morrey CE, Shibata N, Asakawa S, Shimizu N, Hori H, Hamaguchi S, Sakaizumi M (2002) DMY is a Y-specific DM-domain gene required for male development in the medaka fish. Nature 417:559–563PubMedGoogle Scholar
  102. Matsuda M, Sato T, Toyazaki Y, Nagahama Y, Hamaguchi S, Sakaizumi M (2003) Oryzias curvinotus has DMY, a gene that is required for male development in the medaka, O. latipes. Zool Sci 20:159–161PubMedGoogle Scholar
  103. Matsuda M, Shinomiya A, Kinoshita M, Suzuki A, Kobayashi T, Paul-Prasanth B, Lau EL, Hamaguchi S, Sakaizumi M, Nagahama Y (2007) DMY gene induces male development in genetically female (XX) medaka fish. Proc Natl Acad Sci USA 104(10):3865–3870PubMedGoogle Scholar
  104. Matthews J, Gustafsson JA (2003) Estrogen signaling: a subtle balance between ERα and ERβ. Mol Interv 3:281–292PubMedGoogle Scholar
  105. Meijide FJ, Lo Nostro FL, Guerrero GA (2005) Gonadal development and sex differentiation in the cichlid fish Cichlasoma dimerus (Teleostei, Perciformes): a light and electron-microscopy study. J Morphol 264:191–210PubMedGoogle Scholar
  106. Menuet A, Pellegrini E, Anglade I, Blaise O, Laudet V, Kah O, Pakdel F (2002) Molecular characterization of three estrogen receptor forms in zebrafish: binding characteristics, transactivation properties, and tissue distributions. Biol Reprod 66(6):1881–1892PubMedGoogle Scholar
  107. Miles CG, Rankin L, Smith SI, Niksic M, Elgar G, Hastie ND (2003) Faithful expression of a tagged Fugu WT1 protein from a genomic transgene in zebrafish: efficient splicing of pufferfish genes in zebrafish but not mice. Nucleic Acids Res 31(11):2795–2802PubMedGoogle Scholar
  108. Miller D, Summers J, Silber S (2004) Environmental versus genetic sex determination: a possible factor in dinosaur extinction? Fertil Steril 81(4):954–964PubMedGoogle Scholar
  109. Miura T, Miura C, Konda Y, Yamauchi K (2002) Spermatogenesis-preventing substance in Japanese eel. Development 129:2689–2697PubMedGoogle Scholar
  110. Molyneaux KA, Wylie C (2004) Primordial germ cell migration. Int J Dev Biol 48:537–544PubMedGoogle Scholar
  111. Morais da Silva S, Hacker A, Harley V, Goodfellow P, Swain A, Lovell-badge R (1996) Sox9 expression during gonadal development implies a conserved role for the genes in testis differentiation in mammals and birds. Nat Genet 14:62–68PubMedGoogle Scholar
  112. Munday PL, Buston PM, Warner RR (2006a) Diversity and flexibility of sex changes strategies in animals. Trends Ecol Evol 21:89–95PubMedGoogle Scholar
  113. Munday PL, White JW, Warner RR (2006b) A social basis for the development of primary males in a sex changing fish. Proc R Soc B 273:2845–2851PubMedGoogle Scholar
  114. Nakamoto M, Suzuki A, Matsuda M, Nagahama Y, Shibata N (2005) Testicular type Sox9 is not involved in sex determination but might be in the development of testicular structures in the medaka, Oryzias latipes. Biochem Biophys Res Commun 333(3):729–736PubMedGoogle Scholar
  115. Nakamoto M, Matsuda M, Wang DS, Nagahama Y, Shibata N (2006) Molecular cloning and analysis of gonadal expression of Foxl2 in the medaka, Oryzias latipes. Biochem Biophys Res Commun 344(1):353–361PubMedGoogle Scholar
  116. Nakamoto M, Wang DS, Suzuki A, Matsuda M, Nagahama Y, Shibata N (2007) Dax1 suppresses P450arom expression in medaka ovarian follicles. Mol Reprod Dev 74(10):1239–1246PubMedGoogle Scholar
  117. Nakamura M (1984) Effects of estradiol-17 beta on gonadal sex differentiation in two species of salmonids, the masu salmon, Oncorhynchus masou, and the chum salmon, O. keta. Aquaculture 43:1–3Google Scholar
  118. Nakamura M, Nagahama Y (1989) Differentiation and development of Leydig cells, and change of testosterone levels during testicular differentiation in tilapia Oreochromis niloticus. Fish Physiol Biochem 7:211–219Google Scholar
  119. Nakamura M, Takahashi H (1973) Gonadal sex differentiation in tilapia, with special regard to the time of estrogen treatment effective in inducing complete feminization of genetic males. Bull Fac Fish Hokkaido Univ 24:1–13Google Scholar
  120. Nakamura M, Kobayashi T, Chang XT, Nagahama Y (1998) Gonadal sex differentiation in teleost fish. J Exp Zool 281:362–372Google Scholar
  121. Nakamura S, Kobayashi D, Aoki Y, Yokoi H, Ebe Y, Wittbrodt J, Tanaka M (2006) Identification and lineage tracing of two populations of somatic gonadal precursors in medaka embryos. Dev Biol 295(2):678–688PubMedGoogle Scholar
  122. Nakamura S, Aoki Y, Saito D, Kuroki Y, Fujiyama A, Naruse K, Tanaka M (2008) Sox9b/sox9a2-EGFP transgenic medaka reveals the morphological reorganization of the gonads and a common precursor of both the female and male supporting cells. Mol Reprod Dev 75(3):472–476PubMedGoogle Scholar
  123. Nanda I, Kondo M, Hornung U, Asakawa S, Winkler C (2002) A duplicated copy of DMRT1 in the sex-determining region of the Y chromosome of the medaka, Oryzias latipes. Proc Natl Acad Sci USA 99:11778–11783PubMedGoogle Scholar
  124. Nelson JS (1994) Fishes of the world. Wiley, New YorkGoogle Scholar
  125. Ospina-Alvarez N, Piferrer F (2008) Temperature-dependent sex determination in fish revisited: prevalence, a single sex ratio response pattern, and possible effects of climate change. PLoS ONE 3(7):e2837PubMedGoogle Scholar
  126. Ota K, Tateno Y, Gojobori T (2003) Highly differentiated and conserved sex chromosome in fish species (Aulopus japonicus: Teleostei, Aulopidae). Gene 317:187–193PubMedGoogle Scholar
  127. Parenti LR, Grier H (2004) Evolution and phylogeny of gonad morphology in bony fishes. Integr Comp Biol 44:333–348Google Scholar
  128. Park SY, Jameson L (2005) Minireview: transcriptional regulation of gonadal development and differentiation. Endocrinology 146(3):1035–1042PubMedGoogle Scholar
  129. Parker KL, Schimmer BP, Schedl A (1999) Genes essential for early events in gonadal development. Cell Mol Life Sci 55:831–838PubMedGoogle Scholar
  130. Parker KL, Rice DA, Lala DS, Ikeda Y, Luo X, Wong M, Bakke M, Zhao L, Frigeri C, Hanley NA, Stallings N, Schimmer BP (2002) Steroidogenic factor 1: an essential mediator of endocrine development. Recent Prog Horm Res 57:19–36PubMedGoogle Scholar
  131. Patiño R, Takashima F (1995) Gonads. In: Takashima F, Hibiya T (eds) An atlas of fish histology, normal and pathological features. Kodanska/Gustav Fisher, Tokyo, pp 128–153Google Scholar
  132. Pavlidis M, Koumoundouros G, Sterioti A, Somarakis S, Divanach P, Kentouri M (2000) Evidence of temperature-dependent sex determination in the European sea bass (Dicentrarchus labrax L.). J Exp Zool 287(3):225–332PubMedGoogle Scholar
  133. Peichel CL, Ross JA, Matson CK, Dickson M, Grimwood J et al (2004) The master sex determination locus in threespine sticklebacks is on a nascent Y chromosome. Curr Biol 14:1416–1424PubMedGoogle Scholar
  134. Perner B, Englert C, Bollig F (2007) The Wilms tumor genes wt1a and wt1b control different steps during formation of the zebrafish pronephros. Dev Biol 309(1):87–96PubMedGoogle Scholar
  135. Pieau C, Dorizzi M (2004) Oestrogens and temperature-dependent sex determination in reptiles: all is in the gonads. J Endocrinol 181(3):367–377PubMedGoogle Scholar
  136. Pieau C, Dorizzi M, Richard-Mercier N (2001) Temperature-dependent sex determination and gonadal differentiation in reptiles. EXS 91:117–141PubMedGoogle Scholar
  137. Piferrer F, Donaldson EM (1989) Gonadal differentiation in coho salmon, Oncorhynchus kisutch, after a single treatment with androgen or estrogen at different stages during ontogenesis. Aquaculture 77:2–3Google Scholar
  138. Piferrer F, Zanuy S, Carrillo M, Solar I, Devlin RH (1994) Brief treatment with aromatase inhibitor during sex differentiation causes chromosomally female salmon to develop as normal functional males. J Exp Zool 270:255–262Google Scholar
  139. Rasmussen TH, Jespersen A, Korsgaard B (2006) Gonadal morphogenesis and sex differentiation intraovarian embryos of the viviparous fish Zoarces viviparus (Teleostei, Perciformes, Zoarcidae): a histological and ultrastructural study. J Morphol 267:1032–1047PubMedGoogle Scholar
  140. Raymond CS, Kettlewell JR, Hirsch B, Bardwell VJ, Zarkower D (1999) Expression of Dmrt1 in the genital ridge of mouse and chicken embryos suggests a role in vertebrate sexual development. Dev Biol 215(2):208–220PubMedGoogle Scholar
  141. Raz E (2003) Primordial germ-cell development: the zebrafish perspective. Nat Rev Genet 4:690–700PubMedGoogle Scholar
  142. Raz E (2004) Guidance of primordial germ cell migration. Curr Opin Cell Biol 16:169–173PubMedGoogle Scholar
  143. Rey R, Lukas-Croisier C, Lasala C, Bedecarras P (2003) AMH/MIS: what we know already about the gene, the protein and its regulation. Mol Cell Endocrinol 211:21–31PubMedGoogle Scholar
  144. Rodríguez-Marí A, Yan YL, Bremiller RA, Wilson C, Canestro C, Postlethwait JH (2005) Characterization and expression pattern of zebrafish Anti-Mullerian hormone (Amh) relative to sox9a, sox9b, and cyp19a1a, during gonad development. Gene Expr Patterns 5:655–667PubMedGoogle Scholar
  145. Römer U, Beisenherz W (1996) Environmental determination of sex in Apistogramma (Cichlidae) and two other freshwater fishes (Teleostei). J Fish Biol 48:714–725Google Scholar
  146. Ross RM (1990) The evolution of sex-change mechanisms in fishes. Environ Biol Fishes 29:81–93Google Scholar
  147. Ross JA, Urton JR, Boland J, Shapiro MD, Peichel CL (2009) Turnover of sex chromosomes in the stickleback fishes (gasterosteidae). PLoS Genet 5(2):e1000391PubMedGoogle Scholar
  148. Rougeot C, Krim A, Mandiki SN, Kestemont P, Mélard C (2007) Sex steroid dynamics during embryogenesis and sexual differentiation in Eurasian perch, Perca fluviatilis. Theriogenology 67(5):1046–1052PubMedGoogle Scholar
  149. Rubin DA (1985) Effect of pH on sex ratio in cichlids and a poeciliid (Teleostei). Copeia 1985:233–235Google Scholar
  150. Sadovy Y, Colin PL (1995) Sexual development and sexuality in the Nassau grouper. J Fish Biol 46:961–976Google Scholar
  151. Sadovy Y, Shapiro DY (1987) Criteria for the diagnosis of hermaphroditism in fishes. Copeia 1987:36–156Google Scholar
  152. Saito D, Morinaga C, Aoki Y, Nakamura S, Mitani H, Furutani-Seiki M, Kondoh H, Tanaka M (2007) Proliferation of germ cells during gonadal sex differentiation in medaka: insights from germ cell-depleted mutant zenzai. Dev Biol 310(2):280–290PubMedGoogle Scholar
  153. Sawyer SJ, Gerstner KA, Callard GV (2006) Real-time PCR analysis of cytochrome P450 aromatase expression in zebrafish: gene specific tissue distribution, sex differences, developmental programming, and estrogen regulation. Gen Comp Endocrinol 147(2):108–117PubMedGoogle Scholar
  154. Schartl M (2004a) A comparative view on sex determination in medaka. Mech Dev 121(7–8):639–645PubMedGoogle Scholar
  155. Schartl M (2004b) Sex chromosome evolution in non-mammalian vertebrates. Curr Opin Genet Dev 14:634–641PubMedGoogle Scholar
  156. Schlueter PJ, Sang X, Duan C, Wood AW (2007) Insulin-like growth factor receptor 1b is required for zebrafish primordial germ cell migration and survival. Dev Biol 305(1):377–387PubMedGoogle Scholar
  157. Schultz RJ (1993) Genetic regulation of temperature-mediated sex ratios in the livebearing fish Poeciliopsis lucida. Copeia 1993:1148–1151Google Scholar
  158. Scott AG, Penman DJ, Beardmore JA, Skibinski DOF (1989) The ‘‘YY’’ supermale in Oreochromis niloticus (L.) and its potential in aquaculture. Aquaculture 78:3–4Google Scholar
  159. Shapiro D (1981) Size, maturation and the social control of sex reversal in the coral reef fish Anthias squamipinnis. J Zool 193:105–128Google Scholar
  160. Shapiro DY (1990) Sex-changing fish as a manipulable system for the study of the determination, differentiation, and stability of sex in vertebrates. J Exp Zool Suppl 4:132–136PubMedGoogle Scholar
  161. Shapiro DY, Rasotto MB (1993) Sex differentiation and gonadal development in the diandric, protogynous wrasse, Thalassoma bifasciatum (Pisces, Labridae). J Zool 230:231–245Google Scholar
  162. Shen XY, Cui JZ, Gong QL, Nagahama Y (2007) Cloning of the full-length coding sequence and expression analysis of Sox9b in guppy (Poecilia reticulata). Fish Physiol Biochem 33(3):195–202Google Scholar
  163. Shinomiya A, Tanaka M, Kobayashi T, Nagahama Y, Hamaguchi S (2000) The vasa-like gene, olvas, identifies the migration path of primordial germ cells during embryonic body formation stage in the medaka, Oryzias latipes. Dev Growth Differ 42:317–326PubMedGoogle Scholar
  164. Shinomiya A, Hamaguchi S, Shibata N (2001) Sexual differentiation of germ cell deficient gonads in the medaka, Oryzias latipes. J Exp Zool 290:402–410PubMedGoogle Scholar
  165. Shinomiya A, Shibata N, Sakaizumi M, Hamaguchi S (2002) Sex reversal of genetic females (XX) induced by the transplantation of XY somatic cells in the medaka, Oryzias latipes. Int J Dev Biol 46:711–717PubMedGoogle Scholar
  166. Shoemaker C, Ramsey M, Queen J, Crews D (2007) Expression of Sox9, Mis, and Dmrt1 in the gonad of a species with temperature-dependent sex determination. Dev Dyn 236(4):1055–1063PubMedGoogle Scholar
  167. Siegfried KR, Nüsslein-Volhard C (2008) Germ line control of female sex determination in zebrafish. Dev Biol 324(2):277–287PubMedGoogle Scholar
  168. Simpson ER, Mahendroo MS, Means GD (1994) Aromatase cytochrome P450, the enzyme responsible for estrogen synthesis. Endocr Rev 15:342–355PubMedGoogle Scholar
  169. Sinclair AH, Berta P, Palmer MS, Hawkins JR, Griffiths BL, Smith MJ, Foster JW, Frischauf AM, Lovell-Badge R, Goodfellow PN (1990) A gene from the human sex-determining region encodes a protein with homology to a conserved DNA-binding motif. Nature 346(6281):240–244PubMedGoogle Scholar
  170. Smith CA, Smith MJ, Sinclair AH (1999) Gene expression during gonadogenesis in the chicken embryo. Gene 234(2):395–402PubMedGoogle Scholar
  171. Stocco C (2008) Aromatase expression in the ovary: hormonal and molecular regulation. Steroids 73(5):4734–4787Google Scholar
  172. Strüssmann CA, Ito LS (2005) Where does gonadal sex differentiation begin? Gradient of histological sex differentiation in the gonads of pejerrey, Odontesthes bonariensis (Pisces, Atherinidae). J Morphol 265(2):190–196PubMedGoogle Scholar
  173. Strüssmann CA, Moriyama S, Hanke EF, Calsina Cota JC, Takashima F (1996a) Evidence of thermolabile sex determination in pejerrey. J Fish Biol 48:643–651Google Scholar
  174. Strüssmann CA, Cota JCC, Phonlor G, Higuchi H, Takashima F (1996b) Temperature effects on sex differentiation of two South American atherinids, Odontesthes argentinensis and Patagonina hatcheri. Environ Biol Fishes 47:143–154Google Scholar
  175. Strüssmann CA, Saito T, Usui M, Yamada H, Takashima F (1997) Thermal thresholds and critical period of thermolabile sex determination in two atherinid fishes, Odontesthes bonariensis and Patagonina hatcheri. J Exp Zool 278:167–177Google Scholar
  176. Suzuky Y, Nijhout F (2006) Evolution of a polyphenism by genetic accommodation. Science 311:650–652Google Scholar
  177. Takahashi H (1974) Modification of the development of female reproductive organs in the guppy, Poecilia reticulata, following androgen treatment in their juvenile period. Bull Fac Fish Hokkaido Univ 25:174–199Google Scholar
  178. Takahashi H, Shimizu M (1983) Juvenile intersexuality in a cyprinid fish, the Sumatra barb, Barbus tetrazona tetrazona. Bull Fac Fish Hokkaido Univ 34:69–78Google Scholar
  179. Takamatsu N, Kanda H, Ito M, Yamashita A, Yamashita S, Shiba T (1997) Rainbow trout SOX9: cDNA cloning, gene structure and expression. Gene 202:167–170PubMedGoogle Scholar
  180. Tanaka M, Kinoshita M, Kobayashi D, Nagahama Y (2001) Establishment of medaka (Oryzias latipes) transgenic lines with the expression of green fluorescent protein exclusively in germ cells: a useful model to monitor germ cells in a live vertebrate. Proc Natl Acad Sci USA 98:2544–2549PubMedGoogle Scholar
  181. Tanaka K, Takehana Y, Naruse K, Hamaguchi S, Sakaizumi M (2007) Evidence for different origins of sex chromosomes in closely related Oryzias fishes: substitution of the master sex-determining gene. Genetics 177:2075–2081PubMedGoogle Scholar
  182. Thorgaard GH (1977) Heteromorphic sex chromosomes in male rainbow trout. Science 196(4292):900–902PubMedGoogle Scholar
  183. Trant JM, Gavasso S, Ackers J, Chung BC, Place AR (2001) Developmental expression of cytochrome P450 aromatase genes (CYP19a and CYP19b) in zebrafish fry (Danio rerio). J Exp Zool 290(5):475–483PubMedGoogle Scholar
  184. Uyeno T, Miller RR (1971) Multiple sex chromosomes in a Mexican Cyprinodontidae fish. Nature 231:452–453PubMedGoogle Scholar
  185. Valenzuela N, LeClere A, Shikano T (2006) Comparative gene expression of steroidogenic factor 1 in Chrysemys picta and Apalone mutica turtles with temperature-dependent and genotypic sex determination. Evol Dev 8(5):424–432PubMedGoogle Scholar
  186. Van Doorn GS, Kirkpatrick M (2007) Turnover of sex chromosomes induced by sexual conflict. Nature 449(7164):909–912PubMedGoogle Scholar
  187. Vandeputte M, Dupont-Nivet M, Chavanne H, Chatain B (2007) A polygenic hypothesis for sex determination in the European sea bass Dicentrarchus labrax. Genetics 176(2):1049–1057PubMedGoogle Scholar
  188. Veith AM, Froschauer A, Korting C, Nanda I, Hanel R, Scmid M, Schartl M, Volff JN (2003) Cloning dmrt1 gen of Xiphophorus maculatus: dmY/dmrt1Y is not the master sex-determining gene in the platyfish. Gene 317:59–66PubMedGoogle Scholar
  189. Vidal VP, Chaboissier MC, de Rooij DGH, Schedk A (2001) Sox9 induces testis development in XX transgenic mice. Nat Genet 28(3):216–217PubMedGoogle Scholar
  190. Vigier B, Forest MG, Eychenne B, Bézard J, Garrigou O, Robel P, Josso N (1989) Anti-Mullerian hormone produces endocrine sex reversal of fetal ovaries. Proc Nat Acad Sci USA 86:3684–3688PubMedGoogle Scholar
  191. Vizziano D, Randuineau G, Baron D, Cauty C, Guiguen Y (2007) Characterization of early molecular sex differentiation in rainbow trout, Oncorhynchus mykiss. Dev Dyn 236:2198–2206PubMedGoogle Scholar
  192. Volff JN, Schartl M (2001) Variability of genetic sex determination in poeciliid fishes. Genetica 111(1–3):101–110PubMedGoogle Scholar
  193. Von Hofsten J, Olsson PE (2005) Zebrafish sex determination and differentiation: involvement of FTZ-F1 genes. Rep Biol Endocrinol 3:63Google Scholar
  194. Von Hofsten J, Jones I, Karlsson J, Olsson PE (2001) Developmental expression patterns of FTZ-F1 homologues in zebrafish (Danio rerio). Gen Comp Endocrinol 121(2):146–155Google Scholar
  195. Von Hofsten J, Larsson J, Jones I, Olsson PE (2002) Expression and regulation of fushi tarazu factor-1 and steroidogenic genes during reproduction in Arctic charr (Salvelinus alpinus). Biol Reprod 67:1297–1304Google Scholar
  196. Von Hofsten J, Larsson A, Olsson PE (2005) Novel steroidogenic factor-1 homolog (ff1d) is coexpressed with anti-Mullerian hormone (AMH) in zebrafish. Dev Dyn 233:595–604Google Scholar
  197. Wagner T, Wirth J, Meyer J, Zabel B, Held M, Zimmer J, Pasantes J, Bricarelli FD, Keutel J, Hustert E, Wolf U, Tommerup N, Schempp W, Scherer G (1994) Autosomal sex reversal and campomelic dysplasia are caused by mutations in and around the SRY-related gene Sox9. Cell 79:1111–1120PubMedGoogle Scholar
  198. Wang LH, Tsai CL (2000) Effects of temperature on the deformity and sex differentiation of tilapia, Oreochromis mossambicus. J Exp Zool 286:534–537PubMedGoogle Scholar
  199. Wang DS, Kobayashi T, Senthilkumaran B, Sakai F, Sudhakumari CC, Suzuki T, Yoshikuni M, Matsuda M, Morohashi K, Nagahama Y (2002) Molecular cloning of DAX1 and SHP cDNAs and their expression patterns in the Nile tilapia, Oreochromis niloticus. Biochem Biophys Res Commun 297(3):632–640PubMedGoogle Scholar
  200. Wang D, Kobayashi T, Zhou L, Nagahama Y (2004) Molecular cloning and gene expression of Foxl2 in the Nile tilapia, Oreochromis niloticus. Biochem Biophys Res Commun 320(1):83–89PubMedGoogle Scholar
  201. Wang DS, Kobayashi T, Zhou LY, Paul-Prasanth B, Ijiri S, Sakai F, Okubo K, Morohashi K, Nagahama Y (2007) Foxl2 up-regulates aromatase gene transcription in a female-specific manner by binding to the promoter as well as interacting with ad4 binding protein/steroidogenic factor 1. Mol Endocrinol 21(3):712–725PubMedGoogle Scholar
  202. Watanabe M, Tanaka M, Kobayashi D, Yoshiura Y, Oba Y, Nagahama Y (1999) Medaka (Oryzias latipes) FTZ-F1 potentially regulates the transcription of P-450 aromatase in ovarian follicles: cDNA cloning and functional characterization. Mol Cell Endocrinol 149:221–228PubMedGoogle Scholar
  203. Weidinger G, Stebler J, Slanchev K, Dumstrei K, Wise C, Lovell-Badge R, Thisse C, Thisse B, Raz E (2003) Dead end, a novel vertebrate germ plasm component, is required for zebrafish primordial germ cell migration and survival. Curr Biol 13:1429–1434PubMedGoogle Scholar
  204. Western PS, Sinclair AH (2001) Sex, genes, and heat: triggers of diversity. J Exp Zool 290(6):624–631PubMedGoogle Scholar
  205. Wilhelm D, Palmer S, Koopman P (2007) Sex determination and gonadal development in mammals. Physiol Rev 87:1–28PubMedGoogle Scholar
  206. Wourms JP (1981) Viviparity: the maternal-fetal relationships in fishes. Am Zool 21(2):473–515Google Scholar
  207. Yamaguchi T, Yamaguchi S, Hirai T, Kitano T (2007) Follicle-stimulating hormone signaling and Foxl2 are involved in transcriptional regulation of aromatase gene during gonadal sex differentiation in Japanese flounder, Paralichthys olivaceus. Biochem Biophys Res Commun 359(4):935–940PubMedGoogle Scholar
  208. Yamamoto T (1969) Sex differentiation. In: Hoar WS, Randall DJ (eds) Fish physiology, vol 3. Academic Press, London, pp 117–175Google Scholar
  209. Yamazaki F (1976) Application of hormones in fish culture. J Fish Res Board Can 33:948–958Google Scholar
  210. Yan YL, Miller CT, Nissen RM, Singer A, Liu D, Kirn A, Draper B, Willoughby J, Morcos PA, Amsterdam A, Chung BC, Westerfield M, Haffter P, Hopkins N, Kimmel C, Postlethwait JH, Nissen R (2002) A zebrafish sox9 gene required for cartilage morphogenesis. Development 129:5065–5079PubMedGoogle Scholar
  211. Yokoi H, Kobayashi T, Tanaka M, Nagahama Y, Wakamatsu Y, Takeda H, Araki K, Morohashi KI, Ozato K (2002) Sox9 in a teleost fish, medaka (Oryzias latipes): evidence for diversified function of Sox9 in gonad differentiation. Mol Reprod Dev 63:5–16PubMedGoogle Scholar
  212. Yoon C, Kawakami K, Hopkins N (1997) Zebrafish vasa homologue RNA is localized to the cleavage planes of 2- and 4-cell-stage embryos and is expressed in the primordial germ cells. Development 124:3157–3166PubMedGoogle Scholar
  213. Yoshinaga N, Shiraishi E, Yamamoto T, Iguchi T, Abe S, Kitano T (2004) Sexually dimorphic expression of a teleost homologue of Mullerian inhibiting substance during gonadal sex differentiation in Japanese flounder, Paralichthys olivaceus. Biochem Biophys Res Commun 322:508–513PubMedGoogle Scholar
  214. Yoshizaki G, Takeuchi Y, Sakatani S, Takeuchi T (2000) Germ cell-specific expression of green fluorescent protein in transgenic rainbow trout under control of the rainbow trout vasa-like gene promoter. Int J Dev Biol 44:323–326PubMedGoogle Scholar
  215. Zanaria E, Muscatelli F, Bardoni B, Strom TM, Guioli S, Guo W, Lalli E, Moser C, Walker AP, McCabe ERB, Meltinger T, Monaco AP, Sassone-Corsi P, Camerino G (1994) An unusual member of the nuclear hormone receptor superfamily responsible for X linked adrenal hypoplasia congenita. Nature 372:635–641PubMedGoogle Scholar
  216. Zarkower D (2001) Establishing sexual dimorphism: conservation amidst diversity? Nat Rev Genet 2:175–185PubMedGoogle Scholar
  217. Zhou RJ, Liu L, Guo LQ, Yu HS, Cheng H, Huang X, Tiersch TR, Berta P (2003) Similar gene structure of two Sox9a genes and their expression patterns during gonadal differentiation in a teleost fish, rice field eel (Monopterus albus). Mol Reprod Dev 66:211–217PubMedGoogle Scholar

Copyright information

© Springer Science+Business Media B.V. 2009

Authors and Affiliations

  • Guerrero-Estévez Sandra
    • 1
  • Moreno-Mendoza Norma
    • 1
  1. 1.Instituto de Investigaciones BiomedicasUniversidad Nacional Autonoma de Mexico – UNAMMexicoMexico

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