臺灣博碩士論文加值系統

本研究藉由分析LHRH-A、E2和HCG等激素對具有雙性生殖腺的ㄧ齡黃鰭鯛腦區ERs、環化酵素 (CYP19a1b)、GnRHs、LHβ與FSHβ基因表現量的影響，希望能瞭解在黃鰭鯛卵黃生成過程中，E2在腦區扮演之角色；並探討LHRH-A、HCG、E2和ATD等激素對處於成熟階段二齡黃鰭鯛雌魚的影響，以釐清在黃鰭鯛最後成熟階段，E2在腦區扮演之角色和激素對卵質及卵群分佈的影響。
根據本研究在黃鰭鯛腦區中的實驗結果與先前實驗室在生殖腺、肝臟中所發現的結果，我們推測，在黃鰭鯛卵黃蓄積階段，GTH可能透過與腦中GTH受體作用後，使ERα、ERβ1、CYP19a1b大量表現。而透過CYP19a1b生成E2後，E2與ERα作用促使卵黃蓄積。SbGnRH可能參與黃鰭鯛卵細胞最後成熟 (final oocyte maturation, FOM) 階段的調控，而此時E2在腦區中生成後，E2會與GTH共同作用而刺激腦區ERs的表現，其中ERα可能參與卵黃蓄積和最後成熟的調控；ERβ1與ERβ2可能參與最後成熟調控；腦區GPER的表現則可能參與調控完成卵黃蓄積之卵細胞不進入最後成熟有關。
HCG+ E2處理使可以接受催熟之卵細胞增加，使在最後成熟之催熟下有更多卵完成FOM。預先以HCG+E2處理再進行LHRH-A催熟並抑制內源性E2生成之二段式催熟處理，可有效提升卵質而獲致良好排卵。多次產卵型魚種每次產卵之數量並不是很多，且在人工繁殖下往往只產卵一次，故本實驗結果可供多次產卵型魚種人工繁殖改善產卵量及卵質之應用。

The aims of this study were to comprehend the roles of Estradiol-17β (E2) in the brain of yellowfin porgy (Acanthopagrus latus) during vitellogenesis and at the final oocyte maturation (FOM). We analyzed the expression levels of brain ERs, CYP19a1b, GnRHs, the expression levels of pituitary FSHβ and LHβ after treated with LHRH-A, E2 and HCG in one-year-old yellowfin porgy during non-spawning season; and examined the egg quality, the distribution of oocytes diameters, the expression levels of ERs, CYP19a1b, GnRHs, FSHβ and LHβ in the brain or pituitary of the matured two-year-old female fish after LHRH-A, HCG, E2 and ATD treatments.
According to the results of the study and our previous studies on yellowfin porgy, we speculated that GTHs could stimulate the expression levels of brain ERα, ERβ1, and CYP19a1b may due to binding with the GTH receptors which were presented in the brain, and then E2 which was syntheses in the brain, interacted with ERα and stimulated vitellogenesis. Our study showed that sbGnRH involved of the regulation of FOM of yellowfin porgy. During the FOM, the brain E2 and GTHs simultaneously stimulated the expression of brain ERs; the expression of brain ERα might be involved in the regulation of yolk formation and the FOM; the expressions of brain ERβ1and ERβ2 might also regulate the FOM; the expression of brain GPER might be relate to inhibited GVBD in full-grown stage oocytes.
The treatment of HCG combined with E2 in fish could increase the ration of full growth oocyte in ovary, and as the result in increased the amount of ovulated oocyte in following stimulation. Pretreatment of HCG combined with E2, then induced FOM with LHRH-A and decease endogenous E2 at the same time, could improve egg quality and get success in spawning. Our study could provide the application for reproduction of multiple spawning fish, which were low quantity of oocyte in each batch of spawn, and difficult in multiple spawn in artificial environment.

目錄
中文摘要 I
英文摘要 IV
目錄 VIII
表目錄 IX
圖目錄 X
縮寫表 XVIII
第一章 緒論 1
第二章 黃鰭鯛腦區雌性素受體與環化酵素在卵黃生成階段之變化 14
第三章 黃鰭鯛腦區環化酵素與雌性素受體在成熟階段之特性 48
第四章 結論 87
參考文獻 88

鄭絢如,2005. 日本鰻生殖內分泌之研究. 國立海洋大學水產養殖學系研究所博士論文.
顏秀芳,2007. 黃鰭鯛卵細胞成熟之研究. 國立中山大學海洋生物研究所碩士論文.
孫淑芳,2010. 雌二醇對黃鰭鯛卵成熟之影響. 國立高雄海洋科技大學水產養殖研究所碩士論文.
Adams, B.A., Vickers, E.D., Warby, C., Park, M., Fischer, W.H., Grey Craig, A., Rivier, J.E., Sherwood, N.M., 2002. Three forms of gonadotropin-releasing hormone, including a novel form, in a basal salmonid, Coregonus clupeaformis. Biol. Reprod, 67, 232-239.
An, K.W., Nelson, E.R., Habibi, H.R., Choi, C.Y., 2008. Molecular characterization and expression of three GnRH forms mRNA during gonad sex-change process, and effect of GnRHa on GTH subunits mRNA in the protandrous black porgy (Acanthopagrus schlegeli). Gen. Comp. Endocrinol, 159, 38-45.
Andersson, E., Borg, B., Lambert, J.G.D., 1988. Aromatase activity in brain and pituitary of immature and mature atlantic salmon (Salmo salar L.) parr. Gen. Comp. Endocrinol, 72, 394-401.
Andreassen, T.K., Skjoedt, K., Anglade, I., Kah, O., Korsgaard, B., 2003. Molecular cloning, characterisation, and tissue distribution of oestrogen receptor alpha in eelpout (Zoarces viviparus). Gen. Comp. Endocrinol, 132, 356-368.
Anglade, I., Mazurais, D., Douard, V., Le Jossic-Corcos, C., Mananos, E.L., Michel, D., Kah, O., 1999. Distribution of glutamic acid decarboxylase mRNA in the forebrain of the rainbow trout as studied by in situ hybridization. J. Comp. Neurol, 410, 277-289.
Bhandari, R.K., Komuro, H., Nakamura, S., Higa, M., Nakamura, M., 2003. Gonadal restructuring and correlative steroid hormone profiles during natural sex change in protogynous honeycomb grouper (Epinephelus merra). Zoolog Sci, 20, 1399-1404.
Borg, B., 1994. Androgens in teleost fishes. Comparative Biochemistry and Physiology Part C: Pharmacology, Toxicology and Endocrinology, 109, 219-245.
Bowman, C.J., Kroll, K.J., Gross, T.G., Denslow, N.D., 2002. Estradiol-induced gene expression in largemouth bass (Micropterus salmoides). Mol. Cell. Endocrinol, 196, 67-77.
Boyce-Derricott, J., Nagler, J.J., Cloud, J.G., 2009. Regulation of hepatic estrogen receptor isoform mRNA expression in rainbow trout (Oncorhynchus mykiss). Gen. Comp. Endocrinol, 161, 73-78.
Brenner, R.M., West, N.B., McClellan, M.C., 1990. Estrogen and progestin receptors in the reproductive tract of male and female primates. Biol. Reprod, 42, 11-19.
Breton, B., Sambroni, É., Govoroun, M., Weil, C., 1997. Effects of steroids on GTH I and GTH II secretion and pituitary concentration in the immature rainbow trout Oncorhynchus mykiss. Proceedings of the Academy of Sciences, 320, 783-789.
Bustin, S.A., 2000. Absolute quantification of mRNA using real-time reverse transcription polymerase chain reaction assays. J. Mol. Endocrinol, 25, 169-193.
Campbell, B., Dickey, J.T., Swanson, P., 2003. Endocrine changes during onset of puberty in male spring Chinook salmon, Oncorhynchus tshawytscha. Biol. Reprod, 69, 2109-2117.
Chiang, E.F., Yan, Y.L., Guiguen, Y., Postlethwait, J., Chung, B., 2001. Two Cyp19 (P450 aromatase) genes on duplicated zebrafish chromosomes are expressed in ovary or brain. Mol. Biol. Evol, 18, 542-550.
Choi, C.Y., Habibi, H.R., 2003. Molecular cloning of estrogen receptor α and expression pattern of estrogen receptor subtypes in male and female goldfish. Mol. Cell. Endocrinol, 204, 169-177.
Collins, P.M., O'Neill, D.F., Barron, B.R., Moore, R.K., Sherwood, N.M., 2001. 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, 173-179.
Devlin, R.H., Nagahama, Y., 2002. Sex determination and sex differentiation in fish: an overview of genetic, physiological, and environmental influences. Aquaculture, 208, 191-364.
Dickey, J.T., Swanson, P., 1998. Effects of sex steroids on gonadotropin (FSH and LH) regulation in coho salmon (Oncorhynchus kisutch). J. Mol. Endocrinol, 21, 291-306.
Diotel, N., Page, Y.L., Mouriec, K., Tong, S., Pellegrini, E., Vaillant, C., Anglade, I., Brion, F., Pakdel, F., Chung, B., Kah, O., 2010. Aromatase in the brain of teleost fish. Front. Neuroendocrinol, 31, 172-192.
DuSell, C.D., Nelson, E.R., Wittmann, B.M., Fretz, J.A., Kazmin, D., Thomas, R.S., Pike, J.W., McDonnell, D.P., 2010. Regulation of aryl hydrocarbon receptor function by selective estrogen receptor modulators. Mol. Endocrinol, 24, 33-46.
Edwards, D.P., 2005. Regulation of signal transduction pathways by estrogen and progesterone. Annu. Rev. Physiol, 67, 335-376.
Fernald, R.D., White, R.B., 1999. Gonadotropin-Releasing Hormone Genes: Phylogeny, Structure, and Functions. Front. Neuroendocrinol, 20, 224-240.
Filardo, E.J., Quinn, J.A., Bland, K.I., Frackelton, A.R.,Jr, 2000. Estrogen-induced activation of Erk-1 and Erk-2 requires the G protein-coupled receptor homolog, GPR30, and occurs via trans-activation of the epidermal growth factor receptor through release of HB-EGF. Mol. Endocrinol, 14, 1649-1660.
Forlano, P.M., Deitcher, D.L., Myers, D.A., Bass, A.H., 2001. Anatomical distribution and cellular basis for high levels of aromatase activity in the brain of teleost fish: aromatase enzyme and mRNA expression identify glia as source. J. Neurosci, 21, 8943-8955.
Gelinas, D., A. Pitoc, G., V. Callard, G., 1998. Isolation of a goldfish brain cytochrome P450 aromatase cDNA:: mRNA expression during the seasonal cycle and after steroid treatment1. Mol. Cell. Endocrinol, 138, 81-93.
González, A., Piferrer, F., 2003. Aromatase activity in the European sea bass (Dicentrarchus labrax L.) brain. Distribution and changes in relation to age, sex, and the annual reproductive cycle. Gen. Comp. Endocrinol, 132, 223-230.
Graddy, L.G., Kowalski, A.A., Simmen, F.A., Davis, S.L., Baumgartner, W.W., Simmen, R.C., 2000. Multiple isoforms of porcine aromatase are encoded by three distinct genes. J. Steroid Biochem. Mol. Biol, 73, 49-57.
Greytak, S.R., Champlin, D., Callard, G.V., 2005. Isolation and characterization of two cytochrome P450 aromatase forms in killifish (Fundulus heteroclitus): differential expression in fish from polluted and unpolluted environments. Aquat. Toxicol, 71, 371-389.
Guiguen, Y., Fostier, A., Piferrer, F., Chang, C., 2010. Ovarian aromatase and estrogens: A pivotal role for gonadal sex differentiation and sex change in fish. Gen. Comp. Endocrinol, 165, 352-366.
Guraya, S.S., 1986. The cell and molecular biology of fish oogenesis. Monogr. Dev. Biol, 18, 1-223.
Haddy, J.A., Pankhurst, N.W., 2000. The efficacy of exogenous hormones in stimulating changes in plasma steroids and ovulation in wild black bream Acanthopagrus butcheri is improved by treatment at capture. Aquaculture, 191, 351-366.
Hawkins, M.B., Thornton, J.W., Crews, D., Skipper, J.K., Dotte, A., Thomas, P., 2000. Identification of a third distinct estrogen receptor and reclassification of estrogen receptors in teleosts. Proc. Natl. Acad. Sci, 97, 10751-10756.
Hewitt, S.C., Korach, K.S., 2003. Oestrogen receptor knockout mice: roles for oestrogen receptors alpha and beta in reproductive tissues. Reproduction, 125, 143-149.
Holland, M.C., Hassin, S., Zohar, Y., 1998. Effects of long-term testosterone, gonadotropin-releasing hormone agonist, and pimozide treatments on gonadotropin II levels and ovarian development in juvenile female striped bass (Morone saxatilis). Biol. Reprod, 59, 1153-1162.
Huggard, D., Khakoo, Z., Kassam, G., Mahmoud, S.S., Habibi, H.R., 1996. Effect of testosterone on maturational gonadotropin subunit messenger ribonucleic acid levels in the goldfish pituitary. Biol. Reprod, 54, 1184-1191.
Jeng, S., Chen, G., Lai, J., Huang, Y., Dufour, S., Chang, C., 2002. Regulation of pituitary gonadotropin II and growth hormone content by sex steroids and pituitary extract in the aquacultured Japanese eel, Anguilla japonica. Aquaculture, 209, 319-332.
Jeng, S., Pasquier, J., Yueh, W., Chen, G., Lee, Y., Dufour, S., Chang, C., 2012. Differential regulation of the expression of cytochrome P450 aromatase, estrogen and androgen receptor subtypes in the brain–pituitary–ovarian axis of the Japanese eel (Anguilla japonica) reveals steroid dependent and independent mechanisms. Gen. Comp. Endocrinol, 175, 163-172.
Kah, O., Dufour, S. 2011. Chapter 2 - Conserved and Divergent Features of Reproductive Neuroendocrinology in Teleost Fishes. Hormones and Reproduction of Vertebrates. pp. 15-42.
Kajimura, S., Yoshiura, Y., Suzuki, M., Aida, K., 2001. 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, 117-129.
Kang, B.J., Jung, J.-., Lee, J.M., Lim, S.-., Saito, H., Kim, M.H., Kim, Y.-., Saigusa, M., Han, C.-., 2007. Structural and expression analyses of two vitellogenin genes in the carp, Cyprinus carpio. Biochemistry and Molecular Biology, 148, 445-453.
Kavanaugh, S.I., Nozaki, M., Sower, S.A., 2008. Origins of gonadotropin-releasing hormone (GnRH) in vertebrates: identification of a novel GnRH in a basal vertebrate, the sea lamprey. Endocrinology, 149, 3860-3869.
Knapp, R., Carlisle, S.L. 2011. Chapter 3 - Testicular Function and Hormonal Regulation in Fishes. Academic Press, London, pp. 43-63.
Kuiper, G.G., Carlsson, B., Grandien, K., Enmark, E., Haggblad, J., Nilsson, S., Gustafsson, J.A., 1997. Comparison of the ligand binding specificity and transcript tissue distribution of estrogen receptors alpha and beta. Endocrinology, 138, 863-870.
Kusakabe, M., Nakamura, I., Evans, J., Swanson, P., Young, G., 2006. Changes in mRNAs encoding steroidogenic acute regulatory protein, steroidogenic enzymes and receptors for gonadotropins during spermatogenesis in rainbow trout testes. J. Endocrinol, 189, 541-554.
Leaños-Castañeda, O., Van Der Kraak, G., 2007. Functional characterization of estrogen receptor subtypes, ERα and ERβ, mediating vitellogenin production in the liver of rainbow trout. Toxicol. Appl. Pharmacol, 224, 116-125.
Lee, Y.H., Du, J.L., Yueh, W.S., Lin, B.Y., Huang, J.D., Lee, C.Y., Lee, M.F., Lau, E.L., Lee, F.Y., Morrey, C., Nagahama, Y., Chang, C.F., 2001. Sex change in the protandrous black porgy, Acanthopagrus schlegeli: a review in gonadal development, estradiol, estrogen receptor, aromatase activity and gonadotropin. J. Exp. Zool, 290, 715-726.
Lee, Y., Lee, F., Yueh, W., Tacon, P., Du, J., Chang, C., Jeng, S., Tanaka, H., Chang, C., 2000. Profiles of Gonadal Development, Sex Steroids, Aromatase Activity, and Gonadotropin II in the Controlled Sex Change of Protandrous Black Porgy, Acanthopagrus schlegeli Bleeker. Gen. Comp. Endocrinol, 119, 111-120.
Lei, Z.M., Rao, C.V., Kornyei, J.L., Licht, P., Hiatt, E.S., 1993. Novel expression of human chorionic gonadotropin/luteinizing hormone receptor gene in brain. Endocrinology, 132, 2262-2270.
Linard, B., Anglade, I., Corio, M., Navas, J.M., Pakdel, F., Saligaut, C., Kah, O., 1996. Estrogen receptors are expressed in a subset of tyrosine hydroxylase-positive neurons of the anterior preoptic region in the rainbow trout. Neuroendocrinology, 63, 156-165.
Ma, C.H., Dong, K.W., Yu, K.L., 2000. cDNA cloning and expression of a novel estrogen receptor β-subtype in goldfish (Carassius auratus)1. Biochimica Et Biophysica Acta (BBA) - Gene Structure and Expression, 1490, 145-152.
MacKay, M.E., Raelson, J., Lazier, C.B., 1996. Up-regulation of estrogen receptor mRNA and estrogen receptor activity by estradiol in liver of rainbow trout and other teleostean fish. Pharmacology, Toxicology and Endocrinology, 115, 201-209.
Majumder, S., Das, S., Moulik, S.R., Mallick, B., Pal, P., Mukherjee, D., 2015. G-protein coupled estrogen receptor (GPER) inhibits final oocyte maturation in common carp, Cyprinus carpio. Gen. Comp. Endocrinol, 211, 28-38.
Mateos, J., Mañanos, E., Carrillo, M., Zanuy, S., 2002. Regulation of follicle-stimulating hormone (FSH) and luteinizing hormone (LH) gene expression by gonadotropin-releasing hormone (GnRH) and sexual steroids in the Mediterranean Sea bass. Biochemistry and Molecular Biology, 132, 75-86.
Matsuo, H., Baba, Y., Nair, R.M.G., Arimura, A., Schally, A.V., 1971. Structure of the porcine LH- and FSH-releasing hormone. I. The proposed amino acid sequence. Biochem. Biophys. Res. Commun, 43, 1334-1339.
Menuet, A., Anglade, I., Le Guevel, R., Pellegrini, E., Pakdel, F., Kah, O., 2003. Distribution of aromatase mRNA and protein in the brain and pituitary of female rainbow trout: Comparison with estrogen receptor alpha. J. Comp. Neurol, 462, 180-193.
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, 1881-1892.
Menuet, A., Pellegrini, E., Brion, F., Gueguen, M.M., Anglade, I., Pakdel, F., Kah, O., 2005. Expression and estrogen-dependent regulation of the zebrafish brain aromatase gene. J. Comp. Neurol, 485, 304-320.
Miura, T., Higuchi, M., Ozaki, Y., Ohta, T., Miura, C., 2006. Progestin is an essential factor for the initiation of the meiosis in spermatogenetic cells of the eel. Proc. Natl. Acad. Sci, 103, 7333-7338.
Miyamoto, K., Hasegawa, Y., Nomura, M., Igarashi, M., Kangawa, K., Matsuo, H., 1984. Identification of the second gonadotropin-releasing hormone in chicken hypothalamus: evidence that gonadotropin secretion is probably controlled by two distinct gonadotropin-releasing hormones in avian species. Proc. Natl. Acad. Sci, 81, 3874-3878.
Mok, T.K., 1985. Induced spawning and larval rearing of the white seabream, Mylio berda. Aquaculture, 44, 41-49.
Murphy, M.J.,Jr, 1998. Molecular Action and Clinical Relevance of Aromatase Inhibitors. Oncologist, 3, 129-130.
Nagahama, Y., Yamashita, M., 2008. Regulation of oocyte maturation in fish. Dev. Growth Differ, 50 Suppl 1, S195-219.
Nagler, J.J., Krisfalusi, M., Cyr, D.G., 2000. Quantification of rainbow trout (Oncorhynchus mykiss) estrogen receptor-alpha messenger RNA and its expression in the ovary during the reproductive cycle. J. Mol. Endocrinol, 25, 243-251.
Nakamura, M., Kobayashi, T., Chang, X., Nagahama, Y., 1998. Gonadal sex differentiation in teleost fish. J. Exp. Zool, 281, 362-372.
Nelson, E.R., Habibi, H.R., 2010. Functional significance of nuclear estrogen receptor subtypes in the liver of goldfish. Endocrinology, 151, 1668-1676.
Nelson, E.R., Habibi, H.R., 2013. Estrogen receptor function and regulation in fish and other vertebrates. Gen. Comp. Endocrinol, 192, 15-24.
Ogiwara, K., Fujimori, C., Rajapakse, S., Takahashi, T., 2013. Characterization of luteinizing hormone and luteinizing hormone receptor and their indispensable role in the ovulatory process of the medaka. PLOS One, 8, e54482.
Otto, C., Rohde-Schulz, B., Schwarz, G., Fuchs, I., Klewer, M., Brittain, D., Langer, G., Bader, B., Prelle, K., Nubbemeyer, R., Fritzemeier, K., 2008. G Protein-Coupled Receptor 30 Localizes to the Endoplasmic Reticulum and Is Not Activated by Estradiol. Endocrinology, 149, 4846-4856.
Pang, Y., Thomas, P., 2009. Involvement of estradiol-17beta and its membrane receptor, G protein coupled receptor 30 (GPR30) in regulation of oocyte maturation in zebrafish, (Danio rario). Endocrinol, 161, 58-61.
Pang, Y., Dong, J., Thomas, P., 2008. Estrogen Signaling Characteristics of Atlantic Croaker G Protein-Coupled Receptor 30 (GPR30) and Evidence It Is Involved in Maintenance of Oocyte Meiotic Arrest. Endocrinology, 149, 3410-3426.
Pang, Y., Thomas, P., 2010. Role of G protein-coupled estrogen receptor 1, GPER, in inhibition of oocyte maturation by endogenous estrogens in zebrafish. Dev. Biol, 342, 194-206.
Pankhurst, N.W., Carragher, J.F., 1992. Oocyte maturation and changes in plasma steroid levels in snapper Pagrus (Chrysophrys) auratus (Sparidae) following treatment with human chorionic gonadotropin. Aquaculture, 101, 337-347.
Pennie, W.D., Aldridge, T.C., Brooks, A.N., 1998. Differential activation by xenoestrogens of ER alpha and ER beta when linked to different response elements. J. Endocrinol, 158, R11-4.
Powell, J.F., Fischer, W.H., Park, M., Craig, A.G., Rivier, J.E., White, S.A., Francis, R.C., Fernald, R.D., Licht, P., Warby, C., 1995. Primary structure of solitary form of gonadotropin-releasing hormone (GnRH) in cichlid pituitary; three forms of GnRH in brain of cichlid and pumpkinseed fish. Regul. Pept, 57, 43-53.
Prossnitz, E.R., Arterburn, J.B., Sklar, L.A., 2007. GPR30: A G protein-coupled receptor for estrogen. Mol. Cell. Endocrinol, 265–266, 138-142.
Roch, G.J., Busby, E.R., Sherwood, N.M., 2011. Evolution of GnRH: Diving deeper. Gen. Comp. Endocrinol, 171, 1-16.
Sabo-Attwood, T., Kroll, K.J., Denslow, N.D., 2004. Differential expression of largemouth bass (Micropterus salmoides) estrogen receptor isotypes alpha, beta, and gamma by estradiol. Mol. Cell. Endocrinol, 218, 107-118.
Schulz, R.W., Vischer, H.F., Cavaco, J.E.B., Santos, E.M., Tyler, C.R., Goos, H.J.T., Bogerd, J., 2001. Gonadotropins, their receptors, and the regulation of testicular functions in fish. Comparative Biochemistry and Physiology Part B: Biochemistry and Molecular Biology, 129, 407-417.
Schulz, R.W., Goos, H.J.T., 1999. Puberty in male fish: concepts and recent developments with special reference to the African catfish (Clarias gariepinus). Aquaculture, 177, 5-12.
Scott, A.P., Sumpter, J.P., Stacey, N., 2010. The role of the maturation-inducing steroid, 17,20beta-dihydroxypregn-4-en-3-one, in male fishes: a review. J. Fish Biol, 76, 183-224.
Senthilkumaran, B., Yoshikuni, M., Nagahama, Y., 2004. A shift in steroidogenesis occurring in ovarian follicles prior to oocyte maturation. Mol. Cell. Endocrinol, 215, 11-18.
Sherwood, N., Eiden, L., Brownstein, M., Spiess, J., Rivier, J., Vale, W., 1983. Characterization of a teleost gonadotropin-releasing hormone. Proc. Natl. Acad. Sci, U. S. A. 80, 2794-2798.
Sherwood, N.M., Sower, S.A., Marshak, D.R., Fraser, B.A., Brownstein, M.J., 1986. Primary structure of gonadotropin-releasing hormone from lamprey brain. J. Biol. Chem, 261, 4812-4819.
Slayden, O.D., Brenner, R.M., 2004. Hormonal regulation and localization of estrogen, progestin and androgen receptors in the endometrium of nonhuman primates: effects of progesterone receptor antagonists. Arch. Histol, Cytol. 67, 393-409.
Socorro, S., Power, D.M., Olsson, P.E., Canario, A.V., 2000. Two estrogen receptors expressed in the teleost fish, Sparus aurata: cDNA cloning, characterization and tissue distribution. J. Endocrinol, 166, 293-306.
Swanson, P., Suzuki, K., Kawauchi, H., Dickhoff, W.W., 1991. Isolation and characterization of two coho salmon gonadotropins, GTH I and GTH II. Biol. Reprod, 44, 29-38.
Taranger, G.L., Carrillo, M., Schulz, R.W., Fontaine, P., Zanuy, S., Felip, A., Weltzien, F., Dufour, S., Karlsen, Norberg, B., Andersson, E., Hansen, T., 2010. Control of puberty in farmed fish. Gen. Comp. Endocrinol, 165, 483-515.
Thompson, E.A.,Jr, Siiteri, P.K., 1974. The involvement of human placental microsomal cytochrome P-450 in aromatization. J. Biol. Chem, 249, 5373-5378.
Trubiroha, A., Kroupova, H., Wuertz, S., Kloas, W., 2012. Up-regulation of gonadotropin mRNA-expression at the onset of gametogenesis in the roach (Rutilus rutilus): Evidence for an important role of brain-type aromatase (cyp19a1b) in the pituitary. Gen. Comp. Endocrinol, 178, 529-538.
Trudeau, V.L., Murthy, C.K., Habibi, H.R., Sloley, B.D., Peter, R.E., 1993. Effects of sex steroid treatments on gonadotropin-releasing hormone-stimulated gonadotropin secretion from the goldfish pituitary. Biol. Reprod, 48, 300-307.
Tyler, C.R., Pottinger, T.G., Coward, K., Prat, F., Beresford, N., Maddix, S., 1997. Salmonid follicle-stimulating hormone (GtH I) mediates vitellogenic development of oocytes in the rainbow trout, Oncorhynchus mykiss. Biol. Reprod, 57, 1238-1244.
van der Kraak, G., 2009. Chapter 3 The GnRH System and the Neuroendocrine Regulation of Reproduction. Fish Physiology, 28, 115-149.
White, S.A., Kasten, T.L., Bond, C.T., Adelman, J.P., Fernald, R.D., 1995. Three gonadotropin-releasing hormone genes in one organism suggest novel roles for an ancient peptide. Proceedings of the National Academy of Sciences, 92, 8363-8367.
Yang, E.J., Nasipak, B.T., Kelley, D.B., 2007. Direct action of gonadotropin in brain integrates behavioral and reproductive functions. Proc. Natl. Acad. Sci, 104, 2477-2482.
Yaron, Z., Gur, G., Melamed, P., Rosenfeld, H., Elizur, A., Levavi-Sivan, B., 2003. Regulation of fish gonadotropins. Int. Rev. Cytol, 225, 131-185.
You, S., Kim, H., Hsu, C.C., El Halawani, M.E., Foster, D.N., 2000. Three different turkey luteinizing hormone receptor (tLH-R) isoforms I: characterization of alternatively spliced tLH-R isoforms and their regulated expression in diverse tissues. Biol. Reprod, 62, 108-116.
Zempo, B., Kanda, S., Okubo, K., Akazome, Y., Oka, Y., 2013. Anatomical distribution of sex steroid hormone receptors in the brain of female medaka. J. Comp. Neurol, 521, 1760-1780.