Zebrafish 17beta-hydroxysteroid dehydrogenases: An evolutionary perspective

Molecular and Cellular Endocrinology

Volumn 301, Issue 1-2, 2009, Pages 20-26

  Mindnich, R ^a   Adamski, J ^b  

^a UNIVERSITY OF PENNSYLVANIA   (United States)

^b INSTITUTE OF EPIDEMIOLOGY   (Germany)

Author keywords

17beta hydroxysteroid dehydrogenase (17beta HSD); Evolution; Fatty acids; Steroids; Zebrafish

Indexed keywords

ACYL COENZYME A DEHYDROGENASE; LONG CHAIN FATTY ACID; STEROID HORMONE; TESTOSTERONE 17BETA DEHYDROGENASE;

ARTICLE; CELL FUNCTION; ENZYME SPECIFICITY; EVOLUTION; FATTY ACID METABOLISM; FLY; HUMAN; MAMMAL; NONHUMAN; NUCLEOTIDE SEQUENCE; PRIORITY JOURNAL; PROTEIN EXPRESSION; SEQUENCE HOMOLOGY; STEREOSPECIFICITY; VERTEBRATE; WORM; YEAST;

17-HYDROXYSTEROID DEHYDROGENASES; ANIMALS; EVOLUTION, MOLECULAR; HUMANS; SEQUENCE HOMOLOGY, AMINO ACID; ZEBRAFISH;

EID: 60249101573     PISSN: 03037207     EISSN: None     Source Type: Journal    
DOI: 10.1016/j.mce.2008.12.002     Document Type: Article

References (85)

1. Adamski J., Carstensen J., Husen B., Kaufmann M., de Launoit Y., Leenders F., Markus M., and Jungblut P.W. New 17 beta-hydroxysteroid dehydrogenases. Molecular and cell biology of the type IV porcine and human enzymes. Ann. N. Y. Acad. Sci. 784 (1996) 124-136
2. Adamski J., Husen B., Marks F., and Jungblut P.W. Purification and properties of oestradiol 17 beta-dehydrogenase extracted from cytoplasmic vesicles of porcine endometrial cells. Biochem. J. 288 Pt 2 (1992) 375-381
3. Adamski J., Leenders F., Carstensen J.F., Kaufmann M., Markus M.M., Husen B., Tesdorpf J.G., Seedorf U., de Launoit Y., and Jakob F. Steroids, fatty acyl-CoA, and sterols are substrates of 80-kDa multifunctional protein. Steroids 62 (1997) 159-163
4. Akinola L.A., Poutanen M., and Vihko R. Cloning of rat 17 beta-hydroxysteroid dehydrogenase type 2 and characterization of tissue distribution and catalytic activity of rat type 1 and type 2 enzymes. Endocrinology 137 (1996) 1572-1579
5. Ando A., Kikuti Y.Y., Shigenari A., Kawata H., Okamoto N., Shiina T., Chen L., Ikemura T., Abe K., Kimura M., et al. cDNA cloning of the human homologues of the mouse Ke4 and Ke6 genes at the centromeric end of the human MHC region. Genomics 35 (1996) 600-602
6. Aziz N., Maxwell M.M., St Jacques B., and Brenner B.M. Downregulation of Ke 6, a novel gene encoded within the major histocompatibility complex, in murine polycystic kidney disease. Mol. Cell. Biol. 13 (1993) 1847-1853
7. Baker M.E. Co-evolution of steroidogenic and steroid-inactivating enzymes and adrenal and sex steroid receptors. Mol. Cell. Endocrinol. 215 (2004) 55-62
8. Barbaglio A., Sugni M., Di Benedetto C., Bonasoro F., Schnell S., Lavado R., Porte C., and Candia Carnevali D.M. Gametogenesis correlated with steroid levels during the gonadal cycle of the sea urchin Paracentrotus lividus (Echinodermata: Echinoidea). Comp. Biochem. Physiol. A: Mol. Integr. Physiol. 147 (2007) 466-474
9. Beaudoin F., Gable K., Sayanova O., Dunn T., and Napier J.A. A Saccharomyces cerevisiae gene required for heterologous fatty acid elongase activity encodes a microsomal beta-keto-reductase. J. Biol. Chem. 277 (2002) 11481-11488
10. Biswas M.G., and Russell D.W. Expression cloning and characterization of oxidative 17beta- and 3alpha-hydroxysteroid dehydrogenases from rat and human prostate. J. Biol. Chem. 272 (1997) 15959-15966
11. Blomquist C.H., Lindemann N.J., and Hakanson E.Y. Inhibition of 17 beta-hydroxysteroid dehydrogenase (17 beta-HSD) activities of human placenta by steroids and non-steroidal hormone agonists and antagonists. Steroids 43 (1984) 571-586
12. Breitling R., Krazeisen A., Moller G., and Adamski J. 17beta-hydroxysteroid dehydrogenase type 7-an ancient 3-ketosteroid reductase of cholesterogenesis. Mol. Cell. Endocrinol. 171 (2001) 199-204
13. Breitling R., Marijanovic Z., Perovic D., and Adamski J. Evolution of 17beta-HSD type 4, a multifunctional protein of beta-oxidation. Mol. Cell. Endocrinol. 171 (2001) 205-210
14. Day J.M., Foster P.A., Tutill H.J., Parsons M.F., Newman S.P., Chander S.K., Allan G.M., Lawrence H.R., Vicker N., Potter B.V., et al. 17beta-hydroxysteroid dehydrogenase Type 1, and not Type 12, is a target for endocrine therapy of hormone-dependent breast cancer. Int. J. Cancer 122 (2008) 1931-1940
15. Desnoyers S., Blanchard P.G., St-Laurent J.F., Gagnon S.N., Baillie D.L., and Luu-The V. Caenorhabditis elegans LET-767 is able to metabolize androgens and estrogens and likely shares common ancestor with human types 3 and 12 17beta-hydroxysteroid dehydrogenases. J. Endocrinol. 195 (2007) 271-279
16. Deyashiki Y., Ogasawara A., Nakayama T., Nakanishi M., Miyabe Y., Sato K., and Hara A. Molecular cloning of two human liver 3 alpha-hydroxysteroid/dihydrodiol dehydrogenase isoenzymes that are identical with chlordecone reductase and bile-acid binder. Biochem. J. 299 Pt 2 (1994) 545-552
17. Dufort I., Rheault P., Huang X.F., Soucy P., and Luu-The V. Characteristics of a highly labile human type 5 17beta-hydroxysteroid dehydrogenase. Endocrinology 140 (1999) 568-574
18. Dufort I., Soucy P., Labrie F., and Luu-The V. Molecular cloning of human type 3 3 alpha-hydroxysteroid dehydrogenase that differs from 20 alpha-hydroxysteroid dehydrogenase by seven amino acids. Biochem. Biophys. Res. Commun. 228 (1996) 474-479
19. Entchev E.V., Schwudke D., Zagoriy V., Matyash V., Bogdanova A., Habermann B., Zhu L., Shevchenko A., and Kurzchalia T.V. LET-767 is required for the production of branched chain and long chain fatty acids in Caenorhabditis elegans. J. Biol. Chem. 283 (2008) 17550-17560
20. Fomitcheva J., Baker M.E., Anderson E., Lee G.Y., and Aziz N. Characterization of Ke 6, a new 17beta-hydroxysteroid dehydrogenase, and its expression in gonadal tissues. J. Biol. Chem. 273 (1998) 22664-22671
21. Geissler W.M., Davis D.L., Wu L., Bradshaw K.D., Patel S., Mendonca B.B., Elliston K.O., Wilson J.D., Russell D.W., and Andersson S. Male pseudohermaphroditism caused by mutations of testicular 17 beta-hydroxysteroid dehydrogenase 3. Nat. Genet. 7 (1994) 34-39
22. Han G., Gable K., Kohlwein S.D., Beaudoin F., Napier J.A., and Dunn T.M. The Saccharomyces cerevisiae YBR159w gene encodes the 3-ketoreductase of the microsomal fatty acid elongase. J. Biol. Chem. 277 (2002) 35440-35449
23. He X.Y., Merz G., Mehta P., Schulz H., and Yang S.Y. Human brain short chain L-3-hydroxyacyl coenzyme A dehydrogenase is a single-domain multifunctional enzyme. Characterization of a novel 17beta-hydroxysteroid dehydrogenase. J. Biol. Chem. 274 (1999) 15014-15019
24. He X.Y., Merz G., Yang Y.Z., Pullakart R., Mehta P., Schulz H., and Yang S.Y. Function of human brain short chain L-3-hydroxyacyl coenzyme A dehydrogenase in androgen metabolism. Biochim. Biophys. Acta 1484 (2000) 267-277
25. Huang X.F., and Luu-The V. Molecular characterization of a first human 3(alpha→beta)-hydroxysteroid epimerase. J. Biol. Chem. 275 (2000) 29452-29457
26. Huang X.F., and Luu-The V. Gene structure, chromosomal localization and analysis of 3-ketosteroid reductase activity of the human 3(alpha→beta)-hydroxysteroid epimerase. Biochim. Biophys. Acta 1520 (2001) 124-130
27. Huyghe S., Mannaerts G.P., Baes M., and Van Veldhoven P.P. Peroxisomal multifunctional protein-2: the enzyme, the patients and the knockout mouse model. Biochim. Biophys. Acta 1761 (2006) 973-994
28. Ikeda S., Sasazuki S., Natsukawa S., Shaura K., Koizumi Y., Kasuga Y., Ohnami S., Sakamoto H., Yoshida T., Iwasaki M., et al. Screening of 214 single nucleotide polymorphisms in 44 candidate cancer susceptibility genes: a case-control study on gastric and colorectal cancers in the Japanese population. Am. J. Gastroenterol. 103 (2008) 1476-1487
29. Janer G., and Porte C. Sex steroids and potential mechanisms of non-genomic endocrine disruption in invertebrates. Ecotoxicology 16 (2007) 145-160
30. Jansson A.K., Gunnarsson C., Cohen M., Sivik T., and Stal O. 17beta-hydroxysteroid dehydrogenase 14 affects estradiol levels in breast cancer cells and is a prognostic marker in estrogen receptor-positive breast cancer. Cancer Res. 66 (2006) 11471-11477
31. Kazeto Y., Ijiri S., Matsubara H., Adachi S., and Yamauchi K. Cloning of 17beta-hydroxysteroid dehydrogenase-I cDNAs from Japanese eel ovary. Biochem. Biophys. Res. Commun. 279 (2000) 451-456
32. Keay J., Bridgham J.T., and Thornton J.W. The Octopus vulgaris estrogen receptor is a constitutive transcriptional activator: evolutionary and functional implications. Endocrinology 147 (2006) 3861-3869
33. Kho K.H., and Inaba K. Expression of 17beta-hydroxysteroid dehydrogenase in testis of the ascidian Ciona intestinalis [corrected]. Mol. Cell 18 (2004) 171-176
34. Kohler H.R., Kloas W., Schirling M., Lutz I., Reye A.L., Langen J.S., Triebskorn R., Nagel R., and Schonfelder G. Sex steroid receptor evolution and signalling in aquatic invertebrates. Ecotoxicology 16 (2007) 131-143
35. Kuervers L.M., Jones C.L., O'Neil N.J., and Baillie D.L. The sterol modifying enzyme LET-767 is essential for growth, reproduction and development in Caenorhabditis elegans. Mol. Genet. Genomics 270 (2003) 121-131
36. Le Lain R., Nicholls P.J., Smith H.J., and Maharlouie F.H. Inhibitors of human and rat testes microsomal 17beta-hydroxysteroid dehydrogenase (17beta-HSD) as potential agents for prostatic cancer. J. Enzyme Inhib. 16 (2001) 35-45
37. Leenders F., Husen B., Thole H.H., and Adamski J. The sequence of porcine 80 kDa 17 beta-estradiol dehydrogenase reveals similarities to the short chain alcohol dehydrogenase family, to actin binding motifs and to sterol carrier protein 2. Mol. Cell. Endocrinol. 104 (1994) 127-131
38. Lin H.K., Jez J.M., Schlegel B.P., Peehl D.M., Pachter J.A., and Penning T.M. Expression and characterization of recombinant type 2 3 alpha-hydroxysteroid dehydrogenase (HSD) from human prostate: demonstration of bifunctional 3 alpha/17 beta-HSD activity and cellular distribution. Mol. Endocrinol. 11 (1997) 1971-1984
39. Lukacik P., Kavanagh K.L., and Oppermann U. Structure and function of human 17beta-hydroxysteroid dehydrogenases. Mol. Cell. Endocrinol. 248 (2006) 61-71
40. Lukacik P., Kller B., Bunkoczi G., Kavanagh K., Hwa Lee W., Adamski J., and Opermann U. Structural and biochemical characterization of human orphan DHRS10 reveals a novel cytosolic enzyme with steroid dehydrogenase activity. Biochem. J. 402 (2007) 419-427
41. Luu-The V., Tremblay P., and Labrie F. Characterization of type 12 17beta-hydroxysteroid dehydrogenase, an isoform of type 3 17beta-hydroxysteroid dehydrogenase responsible for estradiol formation in women. Mol. Endocrinol. 20 (2006) 437-443
42. Marijanovic Z., Laubner D., Moller G., Gege C., Husen B., Adamski J., and Breitling R. Closing the gap: identification of human 3-ketosteroid reductase, the last unknown enzyme of mammalian cholesterol biosynthesis. Mol. Endocrinol. 17 (2003) 1715-1725
43. Markus M., Husen B., Leenders F., Jungblut P.W., Hall P.F., and Adamski J. The organelles containing porcine 17 beta-estradiol dehydrogenase are peroxisomes. Eur. J. Cell Biol. 68 (1995) 263-267
44. Marques A.T., Antunes A., Fernandes P.A., and Ramos M.J. Comparative evolutionary genomics of the HADH2 gene encoding Abeta-binding alcohol dehydrogenase/17beta-hydroxysteroid dehydrogenase type 10 (ABAD/HSD10). BMC Genomics 7 (2006) 202
45. Mindnich R., and Adamski J. Functional aspects of 17beta-hydroxysteroid dehydrogenase 1 determined by comparison to a closely related retinol dehydrogenase. J. Steroid Biochem. Mol. Biol. 104 (2007) 334-339
46. Mindnich R., Deluca D., and Adamski J. Identification and characterization of 17beta-hydroxysteroid dehydrogenases in the zebrafish, Danio rerio. Mol. Cell. Endocrinol. 215 (2004) 19-30
47. Mindnich R., Haller F., Halbach F., Moeller G., Hrabe de Angelis M., and Adamski J. Androgen metabolism via 17beta-hydroxysteroid dehydrogenase type 3 in mammalian and non-mammalian vertebrates: comparison of the human and the zebrafish enzyme. J. Mol. Endocrinol. 35 (2005) 305-316
48. Mindnich R., Hrabe de Angelis M., and Adamski J. Functional genome analysis indicates loss of 17beta-hydroxysteroid dehydrogenase type 2 enzyme in the zebrafish. J. Steroid Biochem. Mol. Biol. 103 (2007) 35-43
49. Mizuta T., and Kubokawa K. Presence of sex steroids and cytochrome P450 genes in amphioxus. Endocrinology 148 (2007) 3554-3565
50. Moeller G., and Adamski J. Multifunctionality of human 17beta-hydroxysteroid dehydrogenases. Mol. Cell. Endocrinol. 248 (2006) 47-55
51. Moon Y.A., and Horton J.D. Identification of two mammalian reductases involved in the two-carbon fatty acyl elongation cascade. J. Biol. Chem. 278 (2003) 7335-7343
52. Mustonen M.V., Poutanen M.H., Isomaa V.V., Vihko P.T., and Vihko R.K. Cloning of mouse 17beta-hydroxysteroid dehydrogenase type 2, and analysing expression of the mRNAs for types 1, 2, 3, 4 and 5 in mouse embryos and adult tissues. Biochem. J. 325 Pt 1 (1997) 199-205
53. Nokelainen P., Puranen T., Peltoketo H., Orava M., Vihko P., and Vihko R. Molecular cloning of mouse 17 beta-hydroxysteroid dehydrogenase type 1 and characterization of enzyme activity. Eur. J. Biochem. 236 (1996) 482-490
54. Ohnesorg T., Keller B., Hrabe de Angelis M., and Adamski J. Transcriptional regulation of human and murine 17beta-hydroxysteroid dehydrogenase type-7 confers its participation in cholesterol biosynthesis. J. Mol. Endocrinol. 37 (2006) 185-197
55. Peltoketo H., Luu-The V., Simard J., and Adamski J. 17beta-hydroxysteroid dehydrogenase (HSD)/17-ketosteroid reductase (KSR) family; nomenclature and main characteristics of the 17HSD/KSR enzymes. J. Mol. Endocrinol. 23 (1999) 1-11
56. Penning T.M., Burczynski M.E., Jez J.M., Hung C.F., Lin H.K., Ma H., Moore M., Palackal N., and Ratnam K. Human 3alpha-hydroxysteroid dehydrogenase isoforms (AKR1C1-AKR1C4) of the aldo-keto reductase superfamily: functional plasticity and tissue distribution reveals roles in the inactivation and formation of male and female sex hormones. Biochem. J. 351 (2000) 67-77
57. Penning T.M., Burczynski M.E., Jez J.M., Lin H.K., Ma H., Moore M., Ratnam K., and Palackal N. Structure-function aspects and inhibitor design of type 5 17beta-hydroxysteroid dehydrogenase (AKR1C3). Mol. Cell. Endocrinol. 171 (2001) 137-149
58. Pletnev V.Z., and Duax W.L. Rational proteomics IV: modeling the primary function of the mammalian 17beta-hydroxysteroid dehydrogenase type 8. J. Steroid Biochem. Mol. Biol. 94 (2005) 327-335
59. Plourde M., Manhes C., Leblanc G., Durocher F., Dumont M., Sinilnikova O., and Simard J. Mutation analysis and characterization of HSD17B2 sequence variants in breast cancer cases from French Canadian families with high risk of breast and ovarian cancer. J. Mol. Endocrinol. 40 (2008) 161-172
60. Poutanen M., Miettinen M., and Vihko R. Differential estrogen substrate specificities for transiently expressed human placental 17 beta-hydroxysteroid dehydrogenase and an endogenous enzyme expressed in cultured COS-m6 cells. Endocrinology 133 (1993) 2639-2644
61. Puranen T., Poutanen M., Ghosh D., Vihko P., and Vihko R. Characterization of structural and functional properties of human 17 beta-hydroxysteroid dehydrogenase type 1 using recombinant enzymes and site-directed mutagenesis. Mol. Endocrinol. 11 (1997) 77-86
62. Puranen T., Poutanen M., Ghosh D., Vihko R., and Vihko P. Origin of substrate specificity of human and rat 17beta-hydroxysteroid dehydrogenase type 1, using chimeric enzymes and site-directed substitutions. Endocrinology 138 (1997) 3532-3539
63. Qin K.N., and Rosenfield R.L. Expression of 17 beta-hydroxysteroid dehydrogenase type 5 in human ovary: a pilot study. J. Soc. Gynecol. Investig. 7 (2000) 61-64
64. Rantakari P., Strauss L., Kiviranta R., Lagerbohm H., Paviala J., Holopainen I., Vainio S., Pakarinen P., and Poutanen M. Placenta defects and embryonic lethality resulting from disruption of mouse hydroxysteroid (17-beta) dehydrogenase 2 gene. Mol. Endocrinol. 22 (2008) 665-675
65. Ryan K.J., and Engel L.L. The interconversion of estrone and estradiol-17beta by rat liver slices. Endocrinology 52 (1953) 277-286
66. Ryan K.J., and Engel L.L. The interconversion of estrone and estradiol by human tissue slices. Endocrinology 52 (1953) 287-291
67. Seth G., McIvor R.S., and Hu W.S. 17beta-hydroxysteroid dehydrogenase type 7 (Hsd17b7) reverts cholesterol auxotrophy in NS0 cells. J. Biotechnol. 121 (2006) 241-252
68. Sha J.A., Dudley K., Rajapaksha W.R., and O'Shaughnessy P.J. Sequence of mouse 17beta-hydroxysteroid dehydrogenase type 3 cDNA and tissue distribution of the type 1 and type 3 isoform mRNAs. J. Steroid Biochem. Mol. Biol. 60 (1997) 19-24
69. Shafqat N., Marschall H.U., Filling C., Nordling E., Wu X.Q., Bjork L., Thyberg J., Martensson E., Salim S., Jornvall H., et al. Expanded substrate screenings of human and Drosophila type 10 17beta-hydroxysteroid dehydrogenases (HSDs) reveal multiple specificities in bile acid and steroid hormone metabolism: characterization of multifunctional 3alpha/7alpha/7beta/17beta/20beta/21-HSD. Biochem. J. 376 (2003) 49-60
70. Shen Z., Peng Z., Sun Y., Vaananen H.K., and Poutanen M. Overexpression of human hydroxysteroid (17beta) dehydrogenase 2 induces disturbance in skeletal development in young male mice. J. Bone Miner. Res. 23 (2008) 1217-1226
71. Song D., Liu G., Luu-The V., Zhao D., Wang L., Zhang H., Xueling G., Li S., Desy L., Labrie F., et al. Expression of aromatase and 17beta-hydroxysteroid dehydrogenase types 1, 7 and 12 in breast cancer. An immunocytochemical study. J. Steroid Biochem. Mol. Biol. 101 (2006) 136-144
72. Sternberg R.M., Hotchkiss A.K., and Leblanc G.A. The contribution of steroidal androgens and estrogens to reproductive maturation of the eastern mud snail Ilyanassa obsoleta. Gen. Comp. Endocrinol. 156 (2008) 15-26
73. Sultmann H., Murray B.W., and Klein J. Identification of seven genes in the major histocompatibility complex class I region of the zebrafish. Scand. J. Immunol. 51 (2000) 577-585
74. Talalay P., Dobson M.M., and Tapley D.F. Oxidative degradation of testosterone by adaptive enzymes. Nature 170 (1952) 620-621
75. Tarrant A.M. Hormonal signaling in cnidarians: do we understand the pathways well enough to know whether they are being disrupted?. Ecotoxicology 16 (2007) 5-13
76. Thisse B., and Thisse C. Fast Release Clones: A High Throughput Expression Analysis. ZFIN Direct Data Submission (2004). http://zfin.org/ http://zfin.org
77. Torroja L., Ortuno-Sahagun D., Ferrus A., Hammerle B., and Barbas J.A. scully, an essential gene of Drosophila, is homologous to mammalian mitochondrial type II L-3-hydroxyacyl-CoA dehydrogenase/amyloid-beta peptide-binding protein. J. Cell Biol. 141 (1998) 1009-1017
78. Twan W.H., Hwang J.S., Lee Y.H., Wu H.F., Tung Y.H., and Chang C.F. Hormones and reproduction in scleractinian corals. Comp. Biochem. Physiol. A: Mol. Integr. Physiol. 144 (2006) 247-253
79. Vihko P., Harkonen P., Oduwole O., Torn S., Kurkela R., Porvari K., Pulkka A., and Isomaa V. 17 beta-hydroxysteroid dehydrogenases and cancers. J. Steroid Biochem. Mol. Biol. 83 (2002) 119-122
80. Wu L., Einstein M., Geissler W.M., Chan H.K., Elliston K.O., and Andersson S. Expression cloning and characterization of human 17 beta-hydroxysteroid dehydrogenase type 2, a microsomal enzyme possessing 20 alpha-hydroxysteroid dehydrogenase activity. J. Biol. Chem. 268 (1993) 12964-12969
81. Yaeger R., Nolan P.C., Su T., Avila-Bront A., Wang X., Cheung Y.K., Liu H., Rangarajan V.S., Hibshoosh H., Powell C.A., et al. 17HSD 2 may be higher in African-American breast cancer and is associated with estrogen receptor-negative tumors. Cancer Genomics Proteomics 4 (2007) 341-348
82. Yang S.Y., He X.Y., and Miller D. HSD17B10: a gene involved in cognitive function through metabolism of isoleucine and neuroactive steroids. Mol. Genet. Metab. 92 (2007) 36-42
83. Zhang Y., Word R.A., Fesmire S., Carr B.R., and Rainey W.E. Human ovarian expression of 17 beta-hydroxysteroid dehydrogenase types 1, 2, and 3. J. Clin. Endocrinol. Metab. 81 (1996) 3594-3598
84. Zhongyi S., Rantakari P., Lamminen T., Toppari J., and Poutanen M. Transgenic male mice expressing human hydroxysteroid dehydrogenase 2 indicate a role for the enzyme independent of its action on sex steroids. Endocrinology 148 (2007) 3827-3836
85. Zhou L.Y., Wang D.S., Senthilkumaran B., Yoshikuni M., Shibata Y., Kobayashi T., Sudhakumari C.C., and Nagahama Y. Cloning, expression and characterization of three types of 17beta-hydroxysteroid dehydrogenases from the Nile tilapia, Oreochromis niloticus. J. Mol. Endocrinol. 35 (2005) 103-116