The brain-pituitary-gonad axis in male teleosts, with special emphasis on flatfish (Pleuronectiformes)

Comparative Biochemistry and Physiology - A Molecular and Integrative Physiology

Volumn 137, Issue 3, 2004, Pages 447-477

  Weltzien, Finn Arne ^a,b,d   Andersson, Eva ^a   Andersen, Øivind ^c   Shalchian Tabrizi, Kamran ^b   Norberg, Birgitta ^a  

^a INSTITUTE OF MARINE RESEARCH   (Norway)

^b UNIVERSITY OF OSLO   (Norway)

^c Inst Aquaculture Res (AKVAFORSK)   (Norway)

^d MUSÉUM NATIONAL D'HISTOIRE NATURELLE   (France)

Author keywords

Follicle stimulating hormone; Gonadotropin releasing hormone; Luteinizing hormone; Phylogeny; Puberty; Receptors; Reproduction; Spermatogenesis; Steroids; Testis

Indexed keywords

11 OXOTESTOSTERONE; ACTIVIN; CHORIONIC GONADOTROPIN; DOPAMINE; FOLLITROPIN; FOLLITROPIN RECEPTOR; GONADORELIN; GONADORELIN RECEPTOR; GONADOTROPIN; GROWTH FACTOR; GROWTH HORMONE; GROWTH HORMONE RECEPTOR; HYDROXYPROGESTERONE; LUTEINIZING HORMONE; NEUROHORMONE; NEUROTRANSMITTER; PROTEIN SUBUNIT; SEX HORMONE; SOMATOMEDIN C;

AQUACULTURE; BLOOD LEVEL; BRAIN; CELL PROLIFERATION; CHEMICAL STRUCTURE; EMBRYOLOGY; ENZYME ACTIVITY; FLATFISH; GERM CELL; HORMONE STRUCTURE; HORMONE SYNTHESIS; HYPOPHYSIS GONAD SYSTEM; MALE; MAMMAL; MEIOSIS; MORPHOLOGY; NERVOUS TISSUE; NEUROENDOCRINE SYSTEM; NONHUMAN; NUCLEOTIDE SEQUENCE; NUTRITIONAL STATUS; PHYLOGENY; PLEURONECTIFORMES; PROTEIN EXPRESSION; RAINBOW TROUT; REAL TIME POLYMERASE CHAIN REACTION; RECEPTOR BINDING; REPRODUCTION; REVERSE TRANSCRIPTION POLYMERASE CHAIN REACTION; REVIEW; SERTOLI CELL; SEXUAL MATURATION; SPERMATOGENESIS; SPERMATOZOON; SPERMATOZOON COUNT; STEROIDOGENESIS; TELEOST; TESTIS; VERTEBRATE;

ANGUILLIFORMES; CATFISH; CYPRINIDAE; PLEURONECTIFORMES; SALMONIDAE; TELEOSTEI; VERTEBRATA;

EID: 1442302287     PISSN: 10956433     EISSN: None     Source Type: Journal    
DOI: 10.1016/j.cbpb.2003.11.007     Document Type: Review

References (305)

1. Adams B.A., Vickers E.D., Warby C., et al. Three forms of gonadotropin-releasing hormone, including a novel form, in a basal salmonid, Coregonus clupeaformis. Biol. Reprod. 67:2002;232-239.
2. Alok D., Hassin S., Kumar R.S., Trant J.M., Yu K., Zohar Y. Characterization of a pituitary GnRH-receptor from a perciform fish, Morone saxatilis: functional expression in a fish cell line. Mol. Cell. Endocrinol. 168:2000;65-75.
3. Amano M., Hyodo S., Urano A., et al. Activation of salmon gonadotropin-releasing hormone synthesis by 17α-methyltestosterone administration inyearling masu salmon, Oncorhynchus masou. Gen. Comp. Endocrinol. 95:1994;374-380.
4. Amano M., Takahashi A., Yamanome T., Okubo K., Aida K., Yamamori K. Molecular cloning of three cDNAs encoding different GnRHs in the brain of barfin flounder. Gen. Comp. Endocrinol. 126:2002;325-333.
5. Amano M., Oka Y., Yamanome T., Okuzawa K., Yamamori K. Three GnRH systems in the brain and pituitary of a pleuronectiform fish, the barfin flounder Verasper moseri. Cell Tissue Res. 309:2002;323-329.
6. Andersson E., Borg B., de Leeuw R. Characterization of gonadotropin-releasing hormone binding sites in the pituitary of the three-spined stickleback, Gasterosteus aculeatus. Gen. Comp. Endocrinol. 76:1989;41-45.
7. Andersson E., Fjelldal P.G., Klenke U., et al. Three forms of GnRH in the brain and pituitary of the turbot, Scophthalmus maximus: immunological characterization and seasonal variation. Comp. Biochem. Physiol. B. 129:2001;551-558.
8. Anglade I., Zandbergen T., Kah O. Origin of the pituitary innervation of the goldfish. Cell Tissue Res. 273:1993;345-355.
9. Ball J.N., Baker B.I. The pituitary gland: anatomy and histophysiology. Hoar W.S., Randall D.J. Fish Physiology, vol. II. The Endocrine System. 1969;1-110 Academic Press, New York.
10. Ball J.N. Hypothalamic control of the pars distalis in fishes, amphibians, and reptiles. Gen. Comp. Endocrinol. 44:1981;135-170.
11. Barnett C.W., Pankhurst N.W. Reproductive biology and endocrinology of greenback flounder Rhombosolea tapirina (Gunther 1862). Mar. Freshw. Res. 50:1999;35-42.
12. Bass A. Dimorphic male brains and alternative reproductive tactics in a vocalizing fish. Trends Neurosci. 15:1992;139-145.
13. Berlinsky D.L., King W., Smith T.I.J., Hamilton R.D., Holloway J., Sullivan C.V. Induced ovulation of Southern flounder Paralichthys lethostigma using gonadotropin releasing hormone analogue implants. J. World Aquacult. Soc. 27:1996;143-152.
14. Berlinsky D.L., William K., Hodson R.G., Sullivan C.V. Hormone induced spawning of summer flounder Paralichthys dentatus. J. World Aquacult. Soc. 28:1997;79-86.
15. Billard R., Jalabert B., Breton B. Les cellules de Sertoli des poissons téléostéens. I - étude ultrastructural. Ann. Biol. Anim. Bioch. Biophys. 12:1972;19-32.
16. Billard R., Fostier A., Weil C., Breton B. Endocrine control of spermatogenesis in teleost fish. Can. J. Fish. Aquat. Sci. 39:1982;65-79.
17. Bogerd J., Li K.W., Janssen-Dommerholt J., Goos H.J.Th. Two gonadotropin-releasing hormones from African catfish (Clarias gariepinus). Biochem. Biophys. Res. Commmun. 187:1992;127-134.
18. Bogerd J., Blomenröhr M., Andersson E., et al. Discrepancy between molecular structure and ligand selectivity of a testicular follicle-stimulating hormone receptor of the African catfish (Clarias gariepinus). Biol. Reprod. 64:2001;1633-1643.
19. Borg B. Androgens in teleost fishes. Comp. Biochem. Physiol. C. 109:1994;219-245.
20. Bosma P.T., Rebers F.E.M., van Dijk W., Willems P.H.G.M., Goos H.J.Th., Schulz R.W. Inhibitory and stimulatory interactions between endogenous gonadotropin-releasing hormones in the African catfish (Clarias gariepinus). Biol. Reprod. 62:2000;731-738.
21. Braun T., Schofield P.R., Sprengel R. Amino-terminal leucine-rich repeats in gonadotropin receptors determine hormone selectivity. EMBO J. 10:1991;1885-1890.
22. Breton B., Govoroun M., Mikolajczyk T. GTH I and GTH II secretion profiles during the reproductive cycle in female rainbow trout: relationship with pituitary responsiveness to GnRH-A stimulation. Gen. Comp. Endocrinol. 111:1998;38-50.
23. Bromage N.R., Porter M.J.R., Randall C.F. The environmental regulation of maturation in farmed finfish with special reference to the role of photoperiod and melatonin. Aquaculture. 197:2001;63-98.
24. Burton M.P., Idler D.R., Ng T.B. The immunofluorescent location of teleost gonadotropins and thyrotropins in flounder pituitary. Gen. Comp. Endocrinol. 43:1981;135-147.
25. Burton M.P., Idler D.R. The reproductive cycle in winter flounder, Pseudopleuronectes americanus (Walbaum). Can. J. Zool. 62:1984;2563-2567.
26. Burzawa-Gérard E. Chemical data on pituitary gonadotropins and their implication to evolution. Can. J. Fish. Aquat. Sci. 39:1982;80-91.
27. Campbell C.M., Walsh J.M., Idler D.R. Steroids in the plasma of the winter flounder (Pseudopleuronectes americanus Walbaum). A seasonal study and investigation of steroids involvement in oocyte maturation. Gen. Comp. Endocrinol. 29:1976;14-20.
28. Canario A.V., Scott A.P. Effects of steroids and human chorionic gonadotrophin on in vitro oocyte final maturation in two marine flatfish: the dab, Limanda limanda, and the plaice, Pleuronectes platessa. Gen. Comp. Endocrinol. 77:1990;161-176.
29. Carolsfeld J., Powell J.F., Park M., et al. Primary structure and function of three gonadotropin-releasing hormones, including a novel form, from an ancient teleost, herring. Endocrinology. 141:2000;505-512.
30. Cavaco, J.E.B., 1998. Sexual steroid hormones participate in the control of puberty in male African catfish, Clarias gariepinus. Ph.D. Thesis, Utrecht University, The Netherlands (ISBN 90-3931655-4).
31. Cavaco J.E.B., Vischer H.F., Lambert J.G.D., Goos H.J.Th., Schulz R.W. Mismatch between patterns of circulating and testicular androgens in African catfish, Clarias gariepinus. Fish Physiol. Biochem. 17:1997;155-162.
32. Cavaco J.E.B., Schulz R.W., Goos H.J.Th. Sex steroids have diverse effects on pituitary gland and testis during puberty in African catfish Clarias gariepinus. Ann. NY Acad. Sci. 839:1998;584-585.
33. Cavaco J.E., van Baal J., van Dijk W., Hassing G.A., Goos H.J., Schulz R.W. Steroid hormones stimulate gonadotrophs in juvenile male African catfish (Clarias gariepinus). Biol. Reprod. 64:2001;1358-1365.
34. Chang J.P., Yu K.L., Wong A.O.L., Peter R.E. Differential actions of dopamine receptor subunits on gonadotropin and growth hormone release in vitro in goldfish. Neuroendocrinology. 51:1990;664-674.
35. Childs G.V., Hazum E., Amsterdam A., Limor R., Naor Z. Cytochemical evidence for different routes of gonadotropin-releasing hormone processing by large gonadotropes and granulosa cells. Endocrinology. 119:1986;1329-1338.
36. Clearwater S.J., Crim L.M. Gonadotropin-releasing hormone-analogue treatment increase sperm motility, seminal plasma pH and sperm production in yellowtail flounder, Pleuronectes ferrugineus. Fish Physiol. Biochem. 19:1998;349-357.
37. Collins P.M., O'Neill D.F., Barron B.R., Moore R.K., Sherwood N.M. Gonadotropin-releasing hormone content in the brain and pituitary of male and female grass rockfish (Sebastes rastrelliger) in relation to seasonal changes in reproductive status. Biol. Reprod. 65:2001;173-179.
38. Conn P.M., Crowley W.F. Jr. Gonadotropin-releasing hormone and its analogues. Annu. Rev. Med. 45:1994;391-405.
39. Conte F.A., Grumbach M.M., Kaplan S.L. A diphasic pattern of gonadotropin secretion in patients with the syndrome of gonadal dysgenesis. J. Clin. Endocrinol. Metab. 40:1975;670-674.
40. Copeland P.A., Thomas P. Isolation of gonadotropin subunits and evidence for two distinct gonadotropins in Atlantic croaker (Micropogonias undulatus). Gen. Comp. Endocrinol. 91:1993;115-125.
41. Crim L.W., St.Arnaud R., Lavoie M., Labrie F. A study of LH-RH receptors in the pituitary gland of the winter flounder (Pseudopleuronectes americanus Walbaum). Gen. Comp. Endocrinol. 69:1988;372-377.
42. Davies B., Bromage N., Swanson P. The brain-pituitary-gonadal axis of female rainbow trout Oncorhynchus mykiss: effects of photoperiod manipulation. Gen. Comp. Endocrinol. 115:1999;155-166.
43. De Leeuw R., Conn P.N., van't Veer C., Goos H.J.Th., van Oordt P.G.W.J. Characterization of the receptor for gonadotropin-releasing hormone in the pituitary of African catfish, Clarias gariepinus. Fish Physiol. Biochem. 5:1988;99-107.
44. Dellovade T., Schwanzel-Fukuda M., Gordan J., Pfaff D. Aspects of GnRH neurobiology conserved across vertebrate forms. Gen. Comp. Endocrinol. 112:1998;276-282.
45. Dickey J.T., Swanson P. Effects of sex steroids on gonadotropin (FSH and LH) regulation in coho salmon (Oncorhynchus kisutch). J. Mol. Endocrinol. 21:1998;291-306.
46. Doerr-Schott J. Immunohistochemical detection by light and electron microscopy of pituitary hormones in cold-blooded vertebrates. I. Fish and amphibians. Gen. Comp. Endocrinol. 28:1976;487-512.
47. Dubois E.A., Zandbergen M.A., Peute J., Goos H.J.Th. Evolutionary development of three gonadotropin-releasing hormone (GnRH) systems in vertebrates. Brain Res. Bull. 57:2002;413-418.
48. Dufau M.L. The luteinizing hormone receptor. Annu. Rev. Physiol. 60:1998;461-496.
49. Dufour S., Delerue-Le Belle N., Fontaine Y.A. Effects of steroid hormones on pituitary immunoreactive gonadotropin in European freshwater eel, Anguilla anguilla L. Gen. Comp. Endocrinol. 52:1983;190-197.
50. Dufour S., Montero M., Le Belle N., Bassompierre M., King J.A., Millar R.P. Differential distribution and response to experimental sexual maturation of two forms of gonadotropin-releasing hormone (GnRH) in the European eel, Anguilla anguilla. Fish Physiol. Biochem. 11:1993;99-106.
51. Dufour, S., Huang, Y.S., Rousseau, K., et al., 2000. Puberty in teleosts: new insights into the role of peripheral signals in the stimulation of pituitary gonadotropins. In: Norberg, B., Kjesbu, O.S., Taranger, G.L., Andersson, E., Stefansson, S.O. (Eds.), Proceedings of the Sixth International Symposium on the Reproductive Physiology of Fish. John Grieg A/S, Bergen, Norway, pp. 455-461.
52. Dulka J.G., Stacey N.E., Sorensen P.W., Van Der Kraak G.J. A steroid sex pheromone synchronizes male-female spawning readiness in goldfish. Nature. 325:1987;251-253.
53. Elizur A., Zmora N., Rosenfeld H., et al. Gonadotropins beta-GtHI and beta-GtHII from the gilthead seabream, Sparus aurata. Gen. Comp. Endocrinol. 102:1996;39-46.
54. Felsenstein, J., 1995. PHYLIP (Phylogeny Inference Package) Version 3.5. Dept. Genet., Univ. Washington, Seattle.
55. Fernald R.D., White R.B. Gonadotropin-releasing hormone genes: Phylogeny, structure, and functions. Front. Neuroendocrin. 20:1999;224-240.
56. Fiddes J.C., Talmadge K. Structure, expression, and evolution of the genes for the human glycoprotein hormones. Recent Prog. Horm. Res. 40:1984;43-78.
57. Fontaine, Y.A., Dufour, S., 1987. Current status of LH-FSH-like gonadotropin in fish. In: Idler, D.R., Crim, L.W., Walsh, J.W. (Eds.), Proceedings of the Third International Symposium on the Reproductive Physiology of Fish. Memorial University of Newfoundland, St. John's, Canada, pp. 48-56.
58. Fox K.M., Dias J.A., Van Roey P. Three-dimensional structure of human follicle-stimulating hormone. Mol. Endocrinol. 15:2001;378-389.
59. García-Hernández M.P., Koide Y., Díaz M.V., Kawauchi H. Isolation and characterization of two distinct gonadotropins from the pituitary gland of Mediterranean yellowtail, Seriola dumerilii (Risso, 1810). Gen. Comp. Endocrinol. 106:1997;389-399.
60. Gen K., Okuzawa K., Senthilkumaran B., Tanaka H., Moriyama S., Kagawa H. Unique expression of gonadotropin-I and -II subunit genes in male and female red seabream (Pagrus major) during sexual maturation. Biol. Reprod. 63:2000;308-319.
61. Gilkey J.C. Mechanisms of fertilization in fishes. Am. Zool. 21:1981;359-375.
62. Gjerde B. Response to individual selection for age at sexual maturity in Atlantic salmon. Aquaculture. 38:1984;229-240.
63. Gomez J.M., Weil C., Ollitrault M., Le Bail P.-Y., Breton B., Le Gac F. Growth hormone (GH) and gonadotropin subunit gene expression and pituitary and plasma changes during spermatogenesis and oogenesis in rainbow trout (Oncorhynchus mykiss). Gen. Comp. Endocrinol. 113:1999;413-428.
64. Gonzáles-Martínez D., Madigou T., Zmora N., et al. Differential expression of three different prepro-GnRH (gonadotrophin-releasing hormone) messengers in the brain of the European sea bass (Dicentrarchus labrax). J. Comp. Neurol. 429:2001;144-155.
65. Goos H.J.Th., de Leeuw R., Cook H., van Oordt P.G.W.J. Gonadotropic hormone-releasing hormone (GnRH) bioactivity in the brain of the immature rainbow trout, Salmo gairdneri: the effect of testosterone. Gen. Comp. Endocrinol. 64:1986;80-84.
66. Gothilf Y., Muñoz-Cueto J.A., Sagrillo C.A., et al. Three forms of gonadotropin-releasing hormone in a perciform fish (Sparus aurata): complementary deoxyribonucleic acid characterization and brain localization. Biol. Reprod. 55:1996;636-645.
67. Green J.D. The comparative anatomy of the hypophysis with special reference to its blood supply and innervation. Am. J. Anat. 88:1951;225-311.
68. Grier H.J. Comparative organization of Sertoli cells including the Sertoli cell barrier. Russel L.D., Griswold M.D. The Sertoli Cell. 1993;704-739 Cache River Press.
69. Griswold M.D. Interactions between germ cells and Sertoli cells in the testis. Biol. Reprod. 52:1995;211-216.
70. Grumbach M.M., Roth J.C., Kaplan S.L., Kelch R.P. Hypothalamic-pituitary regulation of puberty in man: evidence and concepts derived from clinical research. Grumbach M.M., Grave G.D., Meyer F.E. Control of the Onset of Puberty. 1974;115-166 Wiley, New York.
71. Habibi H.R., Peter R.E., Sokolowska M., Rivier J.E., Vale W.W. Characterization of gonadotropin-releasing hormone (GnRH) binding to pituitary receptors in goldfish (Carassius auratus). Biol. Reprod. 36:1987;844-853.
72. Habibi H.R., Pati D. Extrapituitary GnRH receptors in goldfish. Fish Physiol. Biochem. 11:1993;43-49.
73. Habibi H.R., Pati D., Ouwens M., Goos H.J.Th. Presence of gonadotropin-releasing hormone (GnRH) binding sites and compounds with GnRH-like activity in the ovary of African catfish, Clarias gariepinus. Biol. Reprod. 50:1994;643-652.
74. Hall P.F. Testicular steroid synthesis: organization and regulation. Knobil E., Neill J.D. The Physiology of Reproduction. second ed. 1994;1335-1362 Raven Press, New York.
75. Harmin S.A., Crim L.W. Influence of gonadotropic hormone releasing hormone analog (GnRHa) on plasma sex steroid profiles and milt production in male winter flounder, Pseudopleuronectes americanus (Walbaum). Fish Physiol. Biochem. 10:1993;399-419.
76. Harmin S.A., Crim L.W., Wiegand M.D. Plasma sex steroid profiles and the seasonal reproductive cycle in male and female winter flounder, Pleuronectes americanus. Mar. Biol. 121:1995;601-610.
77. Hart N.F. Fertilization in teleost fishes: mechanisms of sperm-egg interactions. Int. Rev. Cytol. 121:1990;1-66.
78. Hassin S., Elizur A., Zohar Y. Molecular cloning and sequence analysis of striped bass (Morone saxatilis) gonadotropin-I and -II subunits. J. Mol. Endocrinol. 15:1995;23-35.
79. Hassin, S., 1998. Pubertal development in the striped bass (Morone saxatilis, Teleostei): ontogeny and regulation of gonadotropin-I and -II gene expression. Ph.D. Thesis, University of Maryland, USA, pp. 188.
80. Hassin S., Holland M.C.H., Zohar Y. Early maturity in the male striped bass, Morone saxatilis: follicle-stimulating hormone and luteinizing hormone gene expression and their regulation by gonadotropin-releasing hormone analogue and testosterone. Biol. Reprod. 63:2000;1691-1697.
81. Hillensjö T., LeMarie W.J. Gonadotropin releasing hormone agonists stimulate meiotic maturation of follicle-enclosed rat oocytes in vitro. Nature. 287:1980;145-146.
82. Hirai, T., Oba, Y., Yao, Z.X., et al., 2000. Putative gonadotropn receptors in tilapia (Oreochromis niloticus) gonads: cDNA cloning and expression during oogenesis. In: Norberg, B., Kjesbu, O.S., Taranger, G.L., Andersson, E., Stefansson, S.O. (Eds.), Proceedings of the Sixth International Symposium on the Reproductive Physiology of Fish. John Grieg A/S, Bergen, Norway, p. 201.
83. Holland M.C.H., Gothilf Y., Meiri I., et al. Levels of the native forms of GnRH in the pituitary of gilthead seabream, Sparus aurata, at several characteristic stages of the gonadal cycle. Gen. Comp. Endocrinol. 112:1998;394-405.
84. Holland M.C., Hassin S., Zohar Y. Seasonal fluctuations in pituitary levels of the three forms of gonadotropin-releasing hormone in striped bass, Morone saxatilis (Teleostei), during juvenile and pubertal development. J. Endocrinol. 169:2001;527-538.
85. Holmes R.L., Ball J.N. The Pituitary Gland: A Comparative Account. 1974;397 Cambridge University Press, London.
86. Holtzman S., Schreibman M.P., Swanson P., Dilernia A.D. Gonadotropes in maturing winter flounder, Pseudopleuronectes americanus. Am. Zool. 29:1989;A18.
87. Huelsenbeck J.P., Ronquist F. MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics. 17:2001;754-755.
88. Huth J.R., Mountjoy K., Perini F., Ruddon R.W. Intracellular folding pathway of human chorionic gonadotropin beta subunit. J. Biol. Chem. 267:1992;8870-8879.
89. Huth J.R., Mountjoy K., Perini F., Bedows E., Ruddon R.W. Domain-dependent protein folding is indicated by the intracellular kinetics of disulfide bond formation of human chorionic gonadotropin beta subunit. J. Biol. Chem. 267:1992;21396-21403.
90. Idler D.R., Schmidt P.J., Ronald A.P. Isolation and identification of 11-ketotestosterone in salmon plasma. Can. J. Biochem. Physiol. 38:1960;1053-1057.
91. Idler D.R., Fagerlund U.H.M., Ronald A.P. Isolation of pregn-4-ene-17α,20β-diol-3-one from the plasma of Pacific salmon (Oncorhynchus nerka). Biochem. Biophys. Res. Commun. 2:1960;133-137.
92. Idler D.R., MacNab H.C. The biosynthesis of 11-ketotestosterone and 11β-hydroxytestosterone by Atlantic salmon tissues in vitro. Can. J. Biochem. 45:1967;581-589.
93. Idler D.R., Ng T.B. Teleost gonadotropins: isolation, biochemistry and function. Hoar W.S., Randall D.J., Donaldson E.M. Fish Physiology, vol. IXA. Reproduction. 1983;187-221 Academic Press, New York.
94. Idler, D.R., Everard, B.A., 1987. Mammalian, salmon and chicken-like LHRHs from hypothalami of winter flounder (Pseudopleuronectes americanus) as evidenced by chromatographic mobility and immunoreactivity. In: Idler, D.R., Crim, L.W., Walsh, J.W. (Eds.), Proceedings of the Third Symposium on the Reproductive Physiology of Fish. Memorial University of Newfoundland, St. John's, Canada, p. 30.
95. f0.7 VG-GTH have excellent homology with the N-terminal sequence of proopiomelanocortin. In: Scott, A.P., Sumpter, J.P., Kime, D.E., Rolfe, M.S. (Eds.), Proceedings of the Fourth International Symposium on the Reproductive Physiology of Fish. FishSymp91, Sheffield, UK, pp. 8-10.
96. Ikeuchi T., Todo T., Kobayashi T., Nagahama Y. cDNA cloning of a novel androgen receptor subtype. J. Biol. Chem. 274:1999;25205-25209.
97. Illing N., Troskie B.E., Nahorniak C.S., Hapgoood J.P., Peter R.E., Millar R.P. Two gonadotropin-releasing hormone receptor subtypes with distinct ligand selectivity and differential distribution in brain and pituitary in the goldfish (Carassius auratus). Proc. Natl. Acad. Sci. USA. 96:1999;2526-2531.
98. Iwakoshi E., Takuwa-Kuroda K., Fujisawa Y., et al. Isolation and characterization of a GnRH-like peptide from Octopus vulgaris. Biochem. Biophys. Res. Commun. 291:2002;1187-1193.
99. Janssen, P., 1996. Reproduction of the flounder, Platichthys flesus (L.), in relation to environmental pollution - steroids and vitellogenesis. Ph.D. Thesis, University of Utrecht, The Netherlands, pp. 174.
100. Kagawa H., Kawazoe I., Tanaka H., Okuzawa K. Immunocytochemical identification of two distinct gonadotropic cells (GTH I and GTH II) in the pituitary of bluefin tuna, Thunnus thynnus. Gen. Comp. Endocrinol. 110:1998;11-18.
101. Kah, O., Madigou, T., Mazurais, D., Le Dréan, G., 2000. Aspects of the central regulation of reproduction in teleost fish. In: Norberg, B., Kjesbu, O.S., Taranger, G.L., Andersson, E., Stefansson, S.O. (Eds.), Proceedings of the Sixth International Symposium on the Reproductive Physiology of Fish. John Grieg A/S, Bergen, Norway, pp. 27-34.
102. Kajimura S., Yoshiura Y., Suzuki M., Aida K. cDNA cloning of two gonadotropin β subunits (GTH-Iβ and -IIβ) and their expression profiles during gametogenesis in the Japanese flounder (Paralichthys olivaceus). Gen. Comp. Endocrinol. 122:2001;117-129.
103. Kakizawa S., Kaneko T., Hirano T. Effects of hypothalamic factors on somatolactin secretion from the organ-cultured pituitary of rainbow trout. Gen. Comp. Endocrinol. 105:1997;71-78.
104. Kallman K.D., Schreibman M.P. A sex-linked gene controlling gonadotrop differentiation and its significance in determining the age of sexual maturation and size of the platyfish, Xiphophorus maculatus. Gen. Comp. Endocrinol. 21:1973;287-304.
105. Kessler M.J., Reddy M.S., Shah R.H., Bahl O.P. Structures of N-glycosidic carbohydrate units of human chorionic gonadotropin. J. Biol. Chem. 254:1979;7901-7908.
106. Kessler M.J., Mise T., Ghai R.D., Bahl O.P. Structure and location of the O-glycosidic carbohydrate units of human chorionic gonadotropin. J. Biol. Chem. 254:1979;7909-7914.
107. King J.A., Millar R.P. Heterogeneity of vertebrate luteinizing hormone-releasing hormone. Science. 206:1979;67-69.
108. King J.A., Millar R.P. Evolution of gonadotropin-releasing hormones. Trends Endocrinol. Metab. 3:1992;339-346.
109. King J.A., Millar R.P. Coordinated evolution of GnRHs and their receptors. Parhar I.S., Sakuma Y. GnRH Neurons: Gene to Behavior. 1997;51-77 Brain Shuppan, Tokyo.
110. Kobayashi, T., Andersen, Ø. Molecular cloning and characterization of the receptors for follicle stimulating hormone and luteinizing hormone in Atlantic halibut (Hippoglossus hippoglossus) In preparation.
111. Koide Y., Noso T., Schouten G., et al. Maturational gonadotropin from the African catfish, Clarias gariepinus: purification, characterization, localization, and biological activity. Gen. Comp. Endocrinol. 87:1992;327-341.
112. Koide Y., Itoh H., Kawauchi H. Isolation and characterization of two distinct gonadotropins, GTHI and GTHII, from bonito (Katsuwonus pelamis) pituitary glands. Int. J. Peptide Protein Res. 41:1993;52-65.
113. Konstantinou H., Shen D.C. The social and reproductive behavior of the eyed flounder, Bothus ocellatus, with notes on the spawning of Bothus lunatus and Bothus ellipticus. Environ. Biol. Fish. 44:1995;311-324.
114. Kumar T.R., Palapattu G., Wang P., et al. Transgenic models to study gonadotropin function: the role of follicle-stimulating hormone in gonadal growth and tumorigenesis. Mol. Endocrinol. 13:1999;851-865.
115. Kumar R.S., Ijiri S., Trant J.M. Molecular biology of the channel catfish gonadotropin receptors: 1. Cloning of a functional luteinizing hormone receptor and preovulatory induction of gene expression. Biol. Reprod. 64:2001;1010-1018.
116. Kumar R.S., Ijiri S., Trant J.M. Molecular biology of the channel catfish gonadotropin receptors: 2. Complementary DNA cloning, functional expression, and seasonal gene expression of the follicle-stimulating hormone receptor. Biol. Reprod. 65:2001;710-717.
117. Kyle A.L., Stacey N.E., Peter R.E. Ventral telencephalic lesions: effects on bisexual behavior, activity, and olfaction in the male goldfish. Behav. Neural Biol. 36:1982;229-241.
118. Laan M., Richmond H., He C., Campbell R.K. Zebrafish as a model for vertebrate reproduction: characterization of the first functional zebrafish (Danio rerio) gonadotropin receptor. Gen. Comp. Endocrinol. 125:2002;349-364.
119. Lapthorn A.J., Harris D.C., Littlejohn A., et al. Crystal structure of human chorionic gonadotropin. Nature. 369:1994;455-461.
120. Le Bail P.-Y. Growth-reproduction interaction in salmonids. Zohar Y., Breton B. Reproduction in Fish-Basic and Applied Aspects in Endocrinology and Genetics. 1988;91-108 INRA, Paris.
121. Le Gac F., Blaise O., Fostier A., et al. Growth hormone (GH) and reproduction: a review. Fish Physiol. Biochem. 11:1993;219-232.
122. Lejeune H., Chuzel F., Thomas T., et al. Paracrine regulation of Leydig cells. Ann. Endocrinol. 57:1996;55-63.
123. Lescheid D.W., Powell D.F.F., Fischer W.H., et al. Mammalian gonadotropin-releasing hormone (GnRH) identified by primary structure in Russian sturgeon, Acipenser gueldenstaedti. Regul. Peptides. 55:1995;299-309.
124. Li M.D., Ford J.J. A comprehensive evolutionary analysis based on nucleotide and amino acid sequences of the α- and β-subunits of glycoprotein hormone gene family. J. Endocrinol. 156:1998;529-542.
125. Licht P., Papkoff H. Separation of two distinct gonadotropins from the pituitary gland of the bullfrog Rana catesbeiana. Endocrinology. 94:1974;1587-1594.
126. Lin H.R., Peter R.E. Hormones and spawning in fish. Asian Fish. Sci. 9:1996;21-33.
127. Liu H.W., Stickney R.R., Dickhoff W.W. Changes in plasma concentrations of sex steroids in adult Pacific halibut, Hippoglossus stenolepsis. J. World Aquacult. Soc. 22:1991;30-35.
128. Liu Y.C., Kato Y., Inoue K., Tanaka S., Kurosumi K. Co-localization of LHβ and FSHβ mRNAs in the porcine anterior pituitary by in situ hybridization with biotinylated probes. Biochem. Biophys. Res. Commun. 154:1988;80-84.
129. Loir M., Le Gac F. Insulin-like growth factor-I and -II binding and action on DNA synthesis in rainbow trout spermatogonia and spermatocytes. Biol. Reprod. 51:1994;1154-1163.
130. Loir M. Spermatogonia of rainbow trout: II. In vitro study of the influence of pituitary hormones, growth factors and steroids on mitotic activity. Mol. Reprod. Dev. 53:1999;434-442.
131. Lovejoy D.A., Fischer W.H., Ngamvongchon S., et al. Distinct sequence of gonadotropin-releasing hormone (GnRH) in dogfish brain provides insight into GnRH evolution. Proc. Natl. Acad. Sci. USA. 89:1992;6373-6377.
132. Madigou T., Mañanos-Sanchez E., Hulshof S., et al. Cloning, tissue distribution, and central expression of the gonadotropin-releasing hormone receptor in the rainbow trout (Oncorhynchus mykiss). Biol. Reprod. 63:2000;1857-1866.
133. Manabe H., Ide M., Shinomiya A. Mating system of the lefteye flounder, Engyprosopon grandisquama. Ichthyol. Res. 47:2000;69-74.
134. Mañanós E.L., Swanson P., Stubblefield J., Zohar Y. Purification of gonadotropin II from a teleost fish, the hybrid striped bass, and development of a specific enzyme-linked immunosorbent assay. Gen. Comp. Endocrinol. 108:1997;209-222.
135. Manzella S.M., Dharmesh S.M., Beranek M.C., Swanson P., Baenziger J.U. Evolutionary conservation of the sulfated oligosaccharides on vertebrate glycoprotein hormones that control circulatory half-life. J. Biol. Chem. 270:1995;21665-21671.
136. Marchant T.A., Chang J.P., Nahorniak C.S., Peter R.P. Evidence that gonadotropin-releasing hormone also functions as a growth hormone releasing factor in the goldfish. Endocrinology. 124:1989;2509-2518.
137. Matsuo H., Baba Y., Nair R.M.G., Arimura A., Schally A.V. Structure of the porcine LH- and FSH-releasing hormone. I. The proposed amino acid sequence. Biochem. Biophys. Res. Commun. 43:1971;1334-1339.
138. Mayer I., Borg B., Schulz R. Conversion of 11-ketoandrostenedione to 11-ketotestosterone by blood cells of six fish species. Gen. Comp. Endocrinol. 77:1990;70-74.
139. Melamed P., Gur G., Rosenfeld H., Elizur A., Yaron Z. The mRNA levels of GtH Iβ and GtH IIβ and GH in relation to testicular development and testosterone treatment in pituitary cells of male tilapia. Fish Physiol. Biochem. 17:1997;93-98.
140. Melamed P., Gur G., Rosenfeld H., Elizur A., Schulz R.W., Yaron Z. Reproductive development of male and female tilapia hybrids (Oreochromis niloticus×O. aureus) and changes in mRNA levels of gonadotropin (GtH) Iβ and GtH IIβ subunits. J. Exp. Zool. 286:2000;64-75.
141. Methven D.A., Crim L.W., Norberg B., Brown B.A., Goff G.P., Huse I. Seasonal reproduction and plasma levels of sex steroids and vitellogenin in Atlantic halibut (Hippoglossus hippoglossus). Can. J. Fish. Aquat. Sci. 49:1992;754-759.
142. Millar R.P. GnRH II and type II GnRH receptors. Trends Endocrinol. Metab. 14:2003;35-43.
143. Minucci S., DiMatteo L., Pierantoni R., Varriale B., Rastogi R.K., Chieffi G. In vivo and in vitro stimulatory effect of a gonadotropin-releasing hormone analog (HOE 766) on spermatogonial multiplication in the frog, Rana esculenta. Endocrinology. 119:1986;731-736.
144. 90 of the β-subunit of human chorionic gonadotropin are crucial for heterodimer formation with the α-subunit: experimental evidence for the conclusions from the crystal structure of hCG. Biochim. Biophys. Acta. 1645:2003;49-55.
145. Miura T., Yamauchi K., Takahashi H., Nagahama Y. Human chorionic gonadotropin induces all stages of spermatogenesis in vitro in the male Japanese eel (Anguilla japonica). Dev. Biol. 146:1991;258-262.
146. Miura T., Yamauchi K., Takahashi H., Nagahama Y. Hormonal induction of all stages of spermatogenesis in vitro in the male Japanese eel (Anguilla japonica). Proc. Natl. Acad. Sci. USA. 88:1991;5774-5778.
147. Miura T., Yamauchi K., Takahashi H., Nagahama Y. The role of hormones in the aquisition of sperm motility in salmonid fish. J. Exp. Zool. 261:1992;359-363.
148. Miura T., Miura C., Yamauchi K., Nagahama Y. Human recombinant activin induces proliferation of spermatogonia in vitro in the Japanese eel Anguilla japonica. Fish. Sci. 61:1995;434-437.
149. Miura C., Miura T., Yamashita M., Yamauchi K., Nagahama Y. Hormonal induction of all stages of spermatogenesis in germ-somatic cell coculture from immature Japanese eel testis. Dev. Growth Differ. 38:1996;257-262.
150. Miura T., Kawamura S., Miura C., Yamauchi K. Impaired spermatogenesis in the Japanese eel, Anguilla japonica: possibility of the existence of factors that regulate entry of germ cells into meiosis. Dev. Growth Differ. 39:1997;685-691.
151. Miwa S., Yan L., Swanson P. Localization of two gonadotropin receptors in the salmon gonad by in vitro ligand autoradiography. Biol. Reprod. 50:1994;629-642.
152. Montaner A.D., Park M., Fischer W.H., et al. Primary structure of a novel gonadotropin-releasing hormone in the brain of a teleost, pejerrey. Endocrinology. 142:2001;1453-1460.
153. Montero M., Le Belle N., King J.A., Millar R.P., Dufour S. Differential regulation of the two forms of gonadotropin-releasing hormone (mGnRH and cGnRH-II) by sex steroids in the European female silver eel (Anguilla anguilla). Neuroendocrinology. 61:1995;525-535.
154. Montero M., Dufour S. Gonadotropin-releasing hormones (GnRH) in fishes: evolutionary data on their structure, localization, regulation, and function. Zool. Stud. 35:1996;149-160.
155. Moon S.H., Lim H.K., Kwon J.Y., Lee J.K., Chang Y.J. Increase plasma 17-hydroxyprogesterone and milt production in response to gonadotropin- releasing hormone agonist in captive male starry flounder, Platichthys stellatus. Aquaculture. 218:2003;703-716.
156. Moulton S.M., Burton M.P.M. Histological observations on spermatogenesis in winter flounder, Pleuronectes americanus Walbaum, from Conception Bay, Newfoundland, Canada. Can. J. Zool. 77:1999;1682-1689.
157. Mousa M.A., Mousa S.A. Immunohistochemical localization of gonadotropin releasing hormones in the brain and pituitary gland of the Nile perch, Lates niloticus (Teleostei, Centropomidae). Gen. Comp. Endocrinol. 130:2003;245-255.
158. Moyle W.R., Bahl O.P., März L. Role of the carbohydrate of human
159. Mugnier, C., Gaignon, J.L., Lebegue, E., Fauvel, C., Fostier, A., 1995. Maturation inducing steroid in turbot, Scophthalmus maximus. In: Goetz, F.W., Thomas, P. (Eds.), Proceedings of the Fifth International Symposium on the Reproductive Physiology of Fish. FishSymp95, Austin, TX, USA, p. 323.
160. Mugnier C., Gaignon J.L., Fostier A. In vitro synthesis of 17,20β,21-trihydroxy-4-pregnen-3-one by ovaries of turbot (Scophthalmus maximus L.) during oocyte maturation. Gen. Comp. Endocrinol. 107:1997;63-73.
161. Nabissi M., Pati D., Polzonetti-Magni A.M., Habibi H.R. Presence and activity of compounds with GnRH-like actvity in the ovary of seabream, Sparus aurata. Am. J. Physiol. 272:1997;R111-R117.
162. Nader M.R., Miura T., Ando N., Miura C., Yamauchi K. Recombinant human insulin-like growth factor I stimulates all stages of 11-ketotestosterone- induced spermatogenesis in the Japanese eel, Anguilla japonica, in vitro. Biol. Reprod. 61:1999;944-947.
163. Nævdal G. Genetic factors in connection with age at maturation. Aquaculture. 33:1983;97-106.
164. Nagahama Y., Hirose K., Young G., Adachi S., Suzuki K., Tamaoki B.-I. Relative in vitro effectiveness of 17α,20β-dihydroxy-4-pregnen-3-one and other pregnene derivaties on germinal vesicle breakdown in oocytes of ayu (Plecoglossus altivelis), amago salmon (Oncorhynchus rhodurus), rainbow trout (Salmo gairdneri), and goldfish (Carassius auratus). Gen. Comp. Endocrinol. 51:1983;15-23.
165. Nagahama Y. Endocrine regulation of gametogenesis in fish. Int. J. Dev. Biol. 38:1994;217-229.
166. Nagahama Y., Miura T., Kobayashi T., Ding J. The role of activins in spermatogenesis in fish. Aono T., Sugino H., Vale W.W. Inhibin, Activin and Follistatin. 1997;196-203 Springer, New York.
167. Nagahama, Y., 2000. Gonadal steroid hormones: major regulators of gonadal sex differentiation and gametogenesis in fish. In: Norberg, B., Kjesbu, O.S., Taranger, G.L., Andersson, E., Stefansson, S.O. (Eds.), Proceedings of the Sixth International Symposium on the Reproductive Physiology of Fish. John Grieg A/S, Bergen, Norway, pp. 211-222.
168. Naito N., Hyodo S., Okumoto N., Urano N., Nakai Y. Differential production and regulation of gonadotropins (GTH I and GTH II) in the pituitary gland of rainbow trout, Oncorhynchus mykiss, during ovarian development. Cell Tissue Res. 266:1991;457-467.
169. Naito N., De Jesus E.G., Nakai Y., Hirano T. Ontogeny of pituitary cell types and the hypothalamo-hypophysial relationship during early development of chum salmon, Oncorhynchus keta. Cell Tissue Res. 272:1993;429-437.
170. Neill J.D., Duck L.W., Selllers J.C., Musgrove I.C. A gonadotropin-releasing hormone (GnRH) receptor specific for GnRH II in primates. Biochem. Biophys. Res. Commun. 282:2001;1012-1018.
171. Nelson J.S. Fishes of the World. third ed. 1994;600 Wiley, New York.
172. Ng T.B., Idler D.R. 'Big' and 'little' forms of plaice vitellogenic and maturational hormones. Gen. Comp. Endocrinol. 34:1978;408-420.
173. Ng T.B., Idler D.R. A vitellogenic hormone with a large and small form from salmon pituitaries. Gen. Comp. Endocrinol. 35:1978;189-195.
174. Ngamvongchon S., Lovejoy D.A., Fischer W.H., et al. Primary structures of two forms of gonadotropin-releasing hormone, one distinct and one conserved, from catfish brain. Mol. Cell. Neurosci. 3:1992;17-22.
175. Nikolics K., Mason A.J., Szonyi E., Ramachandran J., Seeburg P.H. A prolactin-inhibiting factor within the precursor for human gonadotropin- releasing hormone. Nature. 316:1985;511-517.
176. Norberg B., Weltzien F.-A., Karlsen Ø., Holm J.C. Effects of photoperiod on sexual maturation and somatic growth in male Atlantic halibut (Hippoglossus hippoglossus L.). Comp. Biochem. Physiol. B. 129:2001;357-365.
177. Norris D.O. Vertebrate Endocrinology. third ed. 1997;Academic Press, San Diego.
178. Nozaki M., Naito N., Swanson P., et al. Salmonid pituitary gonadotrophs I. Distinct cellular distributions of two gonadotropins, GTH I and GTH II. Gen. Comp. Endocrinol. 77:1990;348-357.
179. Oba Y., Hirai T., Yoshiura Y., Yoshikuni M., Kawauchi H., Nagahama Y. Cloning, functional characterization, and expression of a gonadotropin receptor cDNA in theovary and testis of amago salmon (Oncorhynchus rhodurus). Biochem. Biophys. Res. Commun. 263:1999;584-590.
180. Oba Y., Hirai T., Yoshiura Y., Yoshikuni M., Kawauchi H., Nagahama Y. The duality of fish gonadotropin receptors: cloning and functional characterization of a second gonadotropin receptor cDNA expressed in the ovary and testis of amago salmon (Oncorhynchus rhodurus). Biochem. Biophys. Res. Commun. 265:1999;366-371.
181. Oba, Y., Hirai, T., Yoshiura, Y., Nagahama, Y., 2000. Fish pituitary glycoprotein receptors: cloning and characterization of two different gonadotropin receptors from gonads and two thyrotropin-like receptors from thyroid of amago salmon (Oncorhynchus rhodurus). In: Norberg, B., Kjesbu, O.S., Taranger, G.L., Andersson, E., Stefansson, S.O. (Eds.), Proceedings of the Sixth International Symposium on the Reproductive Physiology of Fish. John Grieg A/S, Bergen, Norway, pp. 164-166.
182. Oba Y., Hirai T., Yoshiura Y., Kobayashi T., Nagahama Y. Fish gonadotropin and thyrotropin receptors: the evolution of glycoprotein hormone receptors in vertebrates. Comp. Biochem. Physiol. B. 129:2001;441-448.
183. Oka Y. GnRH neuronal system of fish brain as a model system for the study of peptidergic neuromodulation. Parhar I.S., Sakuma Y. GnRH Neurons: Genes to Behavior. 1997;245-276 Brain Shuppan, Tokyo.
184. Okada T., Kawazoe I., Kimura S., Sasamoto Y., Aida K., Kawauchi H. Purification and characterization of gonadotropin I and II from pituitary glands of tuna (Thunnus obesus). Int. J. Peptide Protein Res. 43:1994;69-80.
185. Okubo K., Suetake H., Aida K. Expression of two gonadotropin-releasing hormone (GnRH) precursor genes in various tissues of the Japanese eel and evolution of GnRH. Zool. Sci. 16:1999;471-478.
186. Okubo K., Suetake H., Usami T., Aida K. Molecular cloning and tissue-specific expression of a gonadotropin-releasing hormone receptor in the Japanese eel. Gen. Comp. Endocrinol. 119:2000;181-192.
187. Okubo K., Nagata S., Ko R., et al. Identification and characterization of two distinct GnRH receptor subtypes in a teleost, the medaka Oryzias latipes. Endocrinology. 142:2001;4729-4739.
188. Okuzawa K., Amano M., Kobayashi M., et al. Differences in salmon GnRH and chicken GnRH-II contents in discrete brain areas of male and female rainbow trout according to age and state of maturity. Gen. Comp. Endocrinol. 80:1990;116-126.
189. Okuzawa K., Granneman J., Bogerd J., et al. Distinct expression of GnRH genes in the red seabream brain. Fish Physiol. Biochem. 17:1997;71-79.
190. Okuzawa K., Gen K., Bruysters M., et al. Seasonal variation of the three native gonadotropin-releasing hormone messenger ribonucleic acids levels in the brain of female red seabream. Gen. Comp. Endocrinol. 130:2003;324-332.
191. Olivereau M. Les cellules gonadotropes hypophysaires du Saumon de l'Atlantique: unicité ou dualité? Gen. Comp. Endocrinol. 28:1976;82-95.
192. Olivereau M. Les cellules gonadotropes chez les Salmonidés. Ann. Biol. Anim. Biochim. Biophys. 18:1978;793-798.
193. Olster D.H., Foster D.L. Control of gonadotropin secretion inthe male during puberty: a decrease in response to steroid inhibitory feedback in the absence of an increase in steroid-independent drive in the sheep. Endocrinology. 118:1986;2225-2234.
194. Oppen-Berntsen D.O., Olsen S.O., Rong C.J., Taranger G.L., Swanson P., Walther B.T. Plasma levels of eggshell zr-proteins, estradiol-17β, and gonadotropins during an annual reproductive cycle of Atlantic salmon (Salmo salar). J. Exp. Zool. 268:1994;59-70.
195. O'Shaughnessy P.J., Dudley K., Rajapaksha W.R.A.K. Expression of follicle stimulating hormone-receptor mRNA during gonadal development. Mol. Cell. Endocrinol. 125:1996;169-175.
196. Papkoff H., Gospodarowicz D., Li C.H. Purification and properties of ovine follicle-stimulating hormone. Arch. Biochem. Biophys. 120:1967;434-439.
197. Papkoff H., Farmer S.W., Licht P. Isolation and characterization of follicle-stimulating hormone and luteinizing hormone and its subunits from snapping turtle (Chelydra sepentina) pituitaries. Endocrinology. 98:1976;767-777.
198. Parhar I., Koibuchi N., Sakai M., Iwata M., Yamaoka S. Gonadotropin-releasing hormone (GnRH): expression during salmon smoltification. Neurosci. Lett. 172:1994;15-18.
199. Parhar I.S., Soga T., Ishikawa Y., Nagahama Y., Sakuma Y. Neurons synthesizing gonadotropin-releasing hormone mRNA subtypes have multiple developmental origins in the medaka. J. Comp. Neurol. 401:1998;217-226.
200. Parhar I.S., Soga T., Sakuma Y., Millar R.P. Spatio-temporal expression of gonadotropin-releasing hormone receptor subtypes in gonadotropes, somatotropes and lactotropes in the cichlid fish. J. Neuroendocrinol. 14:2002;657-665.
201. Pati D., Habibi H.R. Characterization of gonadotropin-releasing hormone (GnRH) receptors in the ovary of common carp (Cyprinus carpio). Can. J. Physiol. Pharmacol. 70:1992;268-274.
202. Pati D., Habibi H.R. Direct action of GnRH variants on goldfish oocyte meiosis and follicular steroidogenesis. Mol. Cell. Endocrinol. 160:2000;75-88.
203. Pescovitz O.H., Srivastava C.H., Breyer P.R., Monts B.A. Paracrine control of spermatogenesis. Trends Endocrinol. Metab. 5:1994;126-131.
204. Peter R.E. The brain and neurohormones in teleost reproduction. Hoar W.S., Randall D.J., Donaldson E.M. Fish Physiology, vol. IXA. Reproduction. 1983;97-135 Academic Press, New York.
205. Peter R.E., Yu K.L. Neuroendocrine regulation of ovulation in fishes: basic and applied aspects. Rev. Fish Biol. Fish. 7:1997;173-197.
206. Planas J., Bern H.A., Millar R.P. Effects of GnRH-associated peptide and its component peptides on prolactin secretion from the tilapia pituitary in vitro. Gen. Comp. Endocrinol. 77:1990;386-396.
207. Powell J.F.F., Zohar Y., Elizur A., et al. Three forms of gonadotropin-releasing hormone characterized from brains of one species. Proc. Natl. Acad. Sci. USA. 91:1994;12081-12085.
208. Powell J.F.F., Reska-Skinner S.M., Prakash O.M., et al. Two new forms of gonadotropin-releasing hormone in a protochordate and the evolutionary implications. Proc. Natl. Acad. Sci. USA. 91:1994;10461-10464.
209. Powell J.F.F., Fischer W.H., Park M., et al. Primary structure of a solitary form of gonadotropin-releasing hormone (GnRH) in cichlid pituitary; three forms of GnRH in brain of cichlid and pumpkinseed fish. Regul. Peptides. 57:1995;43-53.
210. Powell J.F.F., Stranden E.M., Carolsfeld J., et al. Primary forms of 3 forms of gonadotropin-releasing hormone (GnRH) from the pacu brain. Regul. Peptides. 68:1997;189-195.
211. Power D.M. Immunocytochemical identification of growth hormone, prolactin, and gonadotropin cells in the pituitary of male plaice (Pleuronectes platessa) during gonadal maturation. Gen. Comp. Endocrinol. 85:1992;358-366.
212. Prat F., Sumpter J.P., Tyler C.R. Validation of radioimmunoassays for two salmon gonadotropins (GtH I and GtH II) and their plasma concentrations throughout the reproductive cycle in male and female rainbow trout (Oncorhynchus mykiss). Biol. Reprod. 54:1996;1375-1382.
213. Pudney J. Spermatogenesis in nonmammalian vertebrates. Microsc. Res. Tech. 32:1995;459-497.
214. Quérat B. Molecular evolution of the glycoprotein hormones in vertebrates. Davey K.G., Peter R.E., Tobe S.S. Perspectives in Comparative Endocrinology. 1994;27-35 National Research Council of Canada, Ottawa, Canada.
215. Quérat B., Sellouk A., Salmon C. Phylogenetic analysis of the vertebrate glycoprotein hormone family including new sequences of sturgeon (Acipenser baerii) β subunits of the two gonadotropins and the thyroid-stimulating hormone. Biol. Reprod. 63:2000;222-228.
216. Ramirez V.D., McCann S.M. Inhibitory effect of testosterone on luteinizing hormone secretion in immature and adult rats. Endocrinology. 76:1965;412-417.
217. Robison R.R., White R.B., Illing N., et al. Gonadotropin-releasing hormone receptor in the teleost Haplochromis burtoni: structure, location and function. Endocrinology. 142:2001;1737-1743.
218. Rodrígez L., Carrillo M., Sorbera L.A., et al. Pituitary levels of three forms of GnRH in the male European sea bass (Dicentrarchus labrax, L.) during sex differentiation and first spawning season. Gen. Comp. Endocrinol. 120:2000;67-74.
219. Rodríguez-Gómez F.J., Rendon-Unceta M.C., Sarasquete C., Munoz-Cueto J.A. Localization of tyrosine hydroxylase-immunoreactivity in the brain of the Senegalese sole, Solea senegalensis. J. Chem. Neuroanat. 19:2000;17-32.
220. Roos P. Human follicle-stimulating hormone. Its isolation from the pituitary gland and from postmenopausal urine and a study of some chemical, physical, immunological, and biological properties of the hormone from these two sources. Acta Endocrinol. (Copenh.). 59:(Suppl. 131):1968;1-93.
221. Roy P., Datta M., Dasgupta S., Bhattacharya S. Gonadotropin-releasing hormone stimulates thyroid activity in a freshwater muriel, Channa gachau (Ham.) and carps Catla catla (Ham.) and Cirrhinus mrigala (Ham.). Gen. Comp. Endocrinol. 117:2000;456-463.
222. Ryan K.D., Robinson S.L., Tritt S.H., Zeleznik A.J. Sexual maturation in the female ferret: circumventing the gonadostat. Endocrinology. 122:1988;1201-1207.
223. Sairam M.R., Krishnamurthy H. The role of follicle-stimulating hormone in spermatogenesis: lessons from knockout animal models. Arch. Med. Res. 32:2001;601-608.
224. Schlatt S., Meinhardt A., Nieschlag E. Paracrine regulation of cellular interactions in the testis: factors in search of a function. Eur. J. Endocrinol. 137:1997;107-117.
225. Schreibman M.P., Leatherland J.F., McKeown B.A. Functional morphology of the teleost pituitary gland. Am. Zool. 13:1973;719-742.
226. Schreibman M.P., Margolis-Nunno H., Halpern-Sebold L.R., Goos H.J.Th., Perlman P.W. The influence of androgen administration on the structure and function of the brain-pituitary-gonad axis of sexually immature platyfish, Xiphophorus maculatus. Cell Tissue Res. 245:1986;519-524.
227. Schulz R.W., Bosma P.T., Zandbergen M.A., et al. Two gonadotropin- releasing hormones in the African catfish, Clarias gariepinus: localization, pituitary receptor binding, and gonadotropin release actvity. Endocrinology. 133:1993;1569-1577.
228. Schulz R.W., Goos H.J.Th. Puberty in male fish: concepts and recent developments with special reference to the African catfish (Clarias gariepinus). Aquaculture. 177:1999;5-12.
229. Schulz, R.W., Bogerd, J., Goos, H.J.Th., 2000. Spermatogenesis and its endocrine regulation. In: Norberg, B., Kjesbu, O.S., Taranger, G.L., Andersson, E., Stefansson, S.O. (Eds.), Proceedings of the Sixth International Symposium on the Reproductive Physiology of Fish. John Grieg A/S, Bergen, Norway, pp. 225-232.
230. Schulz R.W., Vischer H.F., Cavaco J.E.B., et al. Gonadotropins, their receptors, and the regulation of testicular functions in fish. Comp. Biochem. Physiol. B. 129:2001;407-417.
231. Scott A.P., Bye V.J., Baynes S.M., Springate J.R.C. Seasonal variations in plasma concentrations of 11-ketotestosterone and testosterone in male rainbow trout, Salmo gairdneri Richardson. J. Fish Biol. 17:1980;495-505.
232. Scott A.P., Canario A.V. 17α,20β-dihydroxy-4-pregnen-3-one 20-sulphate: a major new metabolite of the teleost oocyte maturation-inducing steroid. Gen. Comp. Endocrinol. 85:1992;91-100.
233. Selman K., Wallace R.A. Gametogenesis in Fundulus heteroclitus. Am. Zool. 26:1986;173-192.
234. Senthilkumaran B., Okuzawa K., Gen K., Ookura T., Kagawa H. Distribution and seasonal variations in levels of three native GnRHs in the brain and pituitary of perciform fish. J. Neuroendocrinol. 11:1999;181-186.
235. Sherwood N.M., Eiden L., Brownstein M., Spiess J., Rivier J., Vale W. Characterization of a teleost gonadotropin-releasing hormone. Proc. Natl. Acad. Sci. USA. 80:1983;2794-2798.
236. Sherwood N.M., Sower S.A., Marshak D.R., Fraser B.A., Brownstein M.J. Primary structure of gonadotropin-releasing hormone from lamprey brain. J. Biol.Chem. 261:1986;4812-4819.
237. Simoni M., Gromoll J., Nieschlag E. The follicle-stimulating hormone receptor: biochemistry, molecular biology, physiology, and pathophysiology. Endocr. Rev. 18:1997;739-773.
238. Sohn Y.C., Yoshiura Y., Kobayashi M., Aida K. Seasonal changes in mRNA levels of gonadotropin and thyrotropin subunits in the goldfish, Carassius auratus. Gen. Comp. Endocrinol. 113:1999;436-444.
239. Sol S.Y., Olson O.P., Lomax D.P., Johnson L.L. Gonadal development and associated changes in plasma reproductive steroids in English sole, Pleuronectes vetulus, from Puget Sound, Washington. Fish. Bull. 96:1998;859-870.
240. Sower S.A., Chiang Y.C., Lovas S., Conlon M. Primary structure and biological activity of a third gonadotropin-releasing hormone from lamprey brain. Endocrinology. 132:1993;1125-1131.
241. Squire P.G., Li C.H. Purification and properties of interstitial cell-stimulating hormone from sheep pituitary glands. J. Biol. Chem. 234:1959;520-525.
242. Stacey N.E. Roles of hormones and pheromones in fish reproductive behavior. Crews D. Psychobiology of Reproductive Behavior. 1987;28-65 Prentice-Hall, Englewood Cliffs, NY.
243. Stearns S.C. The Evolution of Life Histories. 1992;249 Oxford University Press, Oxford.
244. Stocco D.M., Clark B.J. Regulation of the acute production of steroids in steroidogenic cells. Endocr. Rev. 17:1996;221-244.
245. Stockell-Hartree A., Cunningham F.J. Purification of chicken pituitary follicle-stimulating hormone and luteinizing hormone. J. Endocrinol. 43:1969;609-616.
246. Stoner A.W., Bejda A.J., Manderson J.P., Phelan B.A., Stehlik L.L., Pessutti J.P. Behavior of winter flounder, Pseudopleuronectes americanus, during the reproductive season: laboratory and field observations on spawning, feeding, and locomotion. Fish. Bull. 97:1999;999-1016.
247. Styne D.M. Physiology of puberty. Horm. Res. 41:(Suppl. 2):1994;3-6.
248. Sumpter J.P., Scott A.P. Seasonal variations in plasma and pituitary levels of gonadotrophin in males and females of two strains of rainbow trout (Salmo gairdneri). Gen. Comp. Endocrinol. 75:1989;376-388.
249. Suter K.J., Pohl C.R., Plant T.M. Indirect assessment of pulsatile gonadotropin-releasing hormone release in agonadal prepubertal rhesus monkey (Macaca mulatta). J. Endocrinol. 160:1999;35-41.
250. Suzuki K., Kawauchi H., Nagahama Y. Isolation and characterization of two distinct gonadotropins from chum salmon pituitary glands. Gen. Comp. Endocrinol. 71:1988;292-301.
251. Suzuki K., Kawauchi H., Nagahama Y. Isolation and characterization of subunits from two distinct salmon gonadotropins. Gen. Comp. Endocrinol. 71:1988;302-306.
252. Suzuki K., Nagahama Y., Kawauchi H. Steroidogenic activities of two distinct salmon gonadotropins. Gen. Comp. Endocrinol. 71:1988;452-458.
253. Suzuki K., Kanamori A., Nagahama Y., Kawauchi H. Development of salmon GtH I and GtH II radioimmunoassays. Gen. Comp. Endocrinol. 71:1988;459-467.
254. Swanson P., Bernard M., Nozaki M., Suzuki K., Kawauchi H., Dickhoff W.W. Gonadotropins I and II in juvenile coho salmon. Fish Physiol. Biochem. 7:1989;169-176.
255. Swanson, P., 1991. Salmon gonadotropins: reconciling old and new ideas. In: Scott, A.P., Sumpter, J.P., Kime, D.E., Rolfe, M.S. (Eds.), Proceedings of the Fourth International Symposium on the Reproductive Physiology of Fish. FishSymp91, Sheffield, UK, pp. 2-7.
256. Swanson P., Suzuki K., Kawauchi H., Dickhoff W.W. Isolation and characterization of two coho salmon gonadotropins, GtH I and GtH II. Biol. Reprod. 44:1991;29-38.
257. Tanaka H., Kagawa H., Okuzawa K., Hirose K. Purification of gonadotropins (Pm GTH I and II) from red seabream (Pagrus major) and development of a homologous radioimmunoassay for PmGTH II. Fish Physiol. Biochem. 10:1993;409-418.
258. Temple J.L., Millar R.P., Rissman E.F. An evolutionary conserved form of gonadotropin-releasing hormone coordinates energy and reproductive behavior. Endocrinology. 144:2003;13-19.
259. Tensen C., Okuzawa K., Blomenröhr M., et al. Distinct efficacies for two endogenous ligands on a single cognate gonadoliberin receptor. Eur. J. Biochem. 243:1997;134-140.
260. Thompson J.D., Gibson T.J., Plewniak F., Jeanmougin F., Higgins D.G. The CLUSTALX windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucl. Acids Res. 25:1997;4876-4882.
261. Thorpe, J.E., 1986. Age at first maturity in Atlantic salmon, Salmo salar: freshwater period influences and conflicts with smolting. In: Meerburg, D.J. (Ed.), Salmonid age at maturity. Can. Spec. Publ. Fish. Aquat. Sci. 89, 7-14.
262. Thorpe J.E. Developmental variation in salmonid populations. J. Fish Biol. 35:(Suppl. A):1989;295-303.
263. Tilly J.L., Aihara T., Nishimori K., et al. Expression of recombinant human follicle-stimulating hormone receptor: species-specific ligand binding, signal transduction, and identification of multiple ovarian messenger ribonucleic acid transcripts. Endocrinology. 131:1992;799-806.
264. Todo T., Ikeuchi T., Kobayashi T., Nagahama Y. Fish androgen receptor: cDNA cloning, steroid activation of transcription in transfected mammalian cells, and tissue mRNA levels. Biochem. Biophys. Res. Commun. 254:1999;378-383.
265. Todo T., Ikeuchi T., Kobayashi T., et al. Characterization of a testicular 17α,20β-dihydroxy-4-pregnen-3-one (a spermiation-inducing steroid in fish) receptor from a teleost, Japanese eel (Anguilla japonica). FEBS Lett. 465:2000;12-17.
266. Trant J.M., Thomas P., Shackleton C.H.L. Identification of 17α,20β,21-trihydroxy-4-pregnen-3-one as the major ovarian steroid produced by the teleost Micropogonias undulatus during final oocyte maturation. Steroids. 47:1986;89-99.
267. Troskie B., Illing N., Rumbak E., et al. Identification of three putative GnRH receptor subtypes in vertebrates. Gen. Comp. Endocrinol. 112:1998;296-302.
268. Truscott B., So Y.P., Nagler J.J., Idler D.R. Steroids involved with final oocyte maturation in the winter flounder. J. Steroid Biochem. Mol. Biol. 42:1992;351-356.
269. Tsutsumi M., Zhou W., Millar R.P., et al. Cloning and functional expression of a mouse gonadotropin-releasing hormone receptor. Mol. Endocrinol. 6:1992;1163-1169.
270. Tvedt H.B., Benfey T.J., Martin-Robichaud D.J., Power J. The relationship between sperm density, spermatocrit, sperm motility and fertilization success in Atlantic halibut, Hippoglossus hippoglossus. Aquaculture. 194:2001;191-200.
271. Ueda H., Young G., Crim L.W., Kambegawa A., Nagahama Y. 17α,20β-Dihydroxy-4-pregnen-3-one: plasma levels during sexual maturation and in vitro production by the testes of amago salmon (Oncorhynchus rhodurus) and rainbow trout (Salmo gairdneri). Gen. Comp. Endocrinol. 51:1983;106-112.
272. Ueda H., Kambegawa A., Nagahama Y. Involvement of gonadotrophin and steroid hormones in spermiation in the amago salmon, Oncorhynchus rhodurus, and goldfish, Carassius auratus. Gen. Comp. Endocrinol. 59:1985;24-30.
273. Ulloa-Aguirre A., Timossi C., Damian-Matsumura P., Dias J.A. Role of glycosylation in function of follicle-stimulating hormone. Endocrine. 11:1999;205-215.
274. 2 receptors and secretion of follicle-stimulating hormone and luteinizing hormone during the annual reproductive cycle of female rainbow trout. J. Neuroendocrinol. 12:2000;1219-1226.
275. 2 receptors in the pituitary of the African catfish, Clarias gariepinus. Gen. Comp. Endocrinol. 80:1988;107-115.
276. Van Der Kraak G., Suzuki K., Peter R.E., Itoh H., Kawauchi H. Properties of common carp gonadotropin I and gonadotropin II. Gen. Comp. Endocrinol. 85:1992;217-229.
277. Vermeirssen E.L.M., Scott A.P., Mylonas P.P., Zohar Y. Gonadotropin releasing hormone agonist stimulates milt fluidity and plasma concentrations of 17,20β-dihydroxylated and 5β-reduced, 3α-hydroxylated C21 steroids in male plaice (Pleuronectes platessa). Gen. Comp. Endocrinol. 112:1998;163-177.
278. Vermeirssen E.L.M., Shields R.J., de Quero C.M., Scott A.P. Gonadotrophin-releasing hormone agonist raises plasma concentrations of progestogens and enhances milt fluidity in male Atlantic halibut (Hippoglossus hippoglossus). Fish Physiol. Biochem. 22:2000;77-87.
279. Vischer H.F., Bogerd J. Cloning and functional characterization of a gonadal luteinizing hormone receptor complementary DNA from the African catfish (Clarias gariepinus). Biol. Reprod. 68:2003;262-271.
280. Volkoff, H., Peter, R.E., 2000. Effects of two forms of gonadotropin-releasing hormone and a GnRH antagonist on the spawning behavior of the female goldfish, Carassius auratus. In: Norberg, B., Kjesbu, O.S., Taranger, G.L., Andersson, E., Stefansson, S.O. (Eds.), Proceedings of the Sixth International Symposium on the Reproductive Physiology of Fish. John Grieg A/S, Bergen, Norway, pp. 129-131.
281. Voss J.W., Rosenfeld M.G. Anterior pituitary development: short tales from dwarf mice. Cell. 70:1992;527-530.
282. Wang L., Bogerd J., Choi H.S., et al. Three distinct types of GnRH receptors characterized in the bullfrog. Proc. Natl. Acad. Sci. USA. 98:2001;361-366.
283. Weber G.M., Powell J.F.F., Park M., et al. Evidence that gonadotropin-releasing hormone (GnRH) functions as a prolactin-releasing factor in a teleost fish (Oreochromis mossambicus) and primary structures for three native GnRH molecules. J. Endocrinol. 155:1997;121-132.
284. Weil C., Crim L.W., Wilson C.E., Cauty C. Evidence of GnRH receptors in cultured pituitary cells of the winter flounder (Pseudopleuronectes americanus W.). Gen. Comp. Endocrinol. 85:1992;156-164.
285. Weltzien, F.-A., Andersen, Ø., Kobayashi, T., Swanson, P., Norberg, B., 2000. FSH and LH from pituitary glands of Atlantic halibut (Hippoglossus hippoglossus L.). In: Norberg, B., Kjesbu, O.S., Taranger, G.L., Andersson, E., Stefansson, S.O. (Eds.), Proceedings of the Sixth International Symposium on the Reproductive Physiology of Fish. John Grieg A/S, Bergen, Norway, pp. 475-477.
286. Weltzien F.-A., Taranger G.L., Karlsen Ø., Norberg B. Testicular development and plasma androgen levels in Atlantic halibut (Hippoglossus hippoglossus L.). Comp. Biochem. Physiol. A. 132:2002;567-575.
287. Weltzien F.-A., Norberg B., Helvik J.V., Andersen Ø., Swanson P., Andersson E. Identification and localization of eight distinct hormone-producing cell types in the pituitary of male Atlantic halibut (Hippoglossus hippoglossus L.). Comp. Biochem. Physiol. A. 134:2003;315-327.
288. Weltzien F.-A., Norberg B., Swanson P. Isolation and characterization of FSH and LH from pituitary glands of Atlantic halibut (Hippoglossus hippoglossus L.). Gen. Comp. Endocrinol. 131:2003;97-105.
289. Weltzien F.-A., Kobayashi T., Andersson E., Norberg B., Andersen Ø. Molecular characterization and expression of FSHβ, LHβ, and common-α subunit in male Atlantic halibut (Hippoglossus hippoglossus). Gen. Comp. Endocrinol. 131:2003;87-96.
290. White S.A., Kasten T.L., Bond C.T., Adelman J.T., Fernald R.D. Genomic structure and expression sites of three gonadotropin-releasing hormone genes in one organism suggest novel roles for an ancient peptide. Neurobiology. 92:1995;8363-8367.
291. White S.A., Fernald R.D. Genomic structure and expression sites of three gonadotropin-releasing hormone genes in one species. Gen. Comp. Endocrinol. 112:1998;17-25.
292. Wingstrand K.G. Comparative anatomy and evolution of the hypophysis. Harris G.W., Donovan B.T. The Pituitary Gland, vol. 1. 1966;58-126 Butterworths, London.
293. Wu H., Lustbader J.W., Liu Y., Canfield R.E., Hendrickson W.A. Structure of human chorionic gonadotropin at 2.6 Å resolution from MAD analysis of the selenomethionyl protein. Structure. 2:1994;545-558.
294. Xiong F., Chin R., Gong Z.-Y., et al. Control of salmon pituitary hormone gene expression. Fish Physiol. Biochem. 11:1993;63-70.
295. Yamamoto N., Oka Y., Kawashima S. Lesions of gonadotropin-releasing hormone-immunoreactive terminal nerve cells: effects on the reproductive behavior of male dwarf gouramis. Neuroendocrinology. 65:1997;403-412.
296. Yan L., Swanson P., Dickhoff W.W. A two-receptor model for salmon gonadotropins (GTH I and GTH II). Biol. Reprod. 47:1992;418-427.
297. Yen S.S.C., Jaffe R.B., Barbieri R.L. Reproductive Endocrinology: Physiology, Pathophysiology and Clinical Management. fourth ed. 1999;W.B. Saunders, Philadelphia.
298. 8] GnRH) in frog brain. Mol. Cell. Endocrinol. 164:2000;197-204.
299. Yoshiura Y., Suetake H., Aida K. Duality of gonadotropin in a primitive teleost, Japanese eel (Anguilla japonica). Gen. Comp. Endocrinol. 114:1999;121-131.
300. Yu K., Sherwood N., Peter R. Differential distribution of two molecular forms of gonadotropin-releasing hormone in discrete brain areas of the goldfish (Carassius auratus). Peptides. 9:1988;625-630.
301. Yu K.L., He M.L., Chik C.C., Lin X.W., Chang J.P., Peter R.E. mRNA expression of gonadotropin-releasing hormones (GnRHs) and GnRH receptor in goldfish. Gen. Comp. Endocrinol. 112:1998;303-311.
302. Yu K.L., Rosenblum P.M., Peter R.E. In vitro release of gonadotropin-releasing hormone from the brain preoptic-anterior hypothalamic region and pituitary of female goldfish. Gen. Comp. Endocrinol. 81:1991;256-267.
303. Zenkevics H., Vose V., Vosekalne I., Bucena A. Partial deglycosylation of alpha subunit modifies sturgeon gonadotropin function. Acta Biochim. Pol. 47:2000;815-819.
304. Zhang L., Wayne N.L., Sherwood N.M., Postigo H.R., Tsai P.S. Biological and immunological characterization of multiple GnRH in an opisthobranch mollusk, Aplysia californica. Gen. Comp. Endocrinol. 117:2000;77-89.
305. Zohar Y., Elizur A., Sherwood N.M., Powell J.F.F., Rivier J.E., Zmora N. Gonadotropin-releasing activities of the three native forms of gonadotropin-releasing hormone present in the brain of gilthead seabream, Sparus aurata. Gen. Comp. Endocrinol. 97:1995;289-299.