Isolation of CYP17 Ⅰ, 3β-HSD and AR Genes from Spotted Sea Bass (Lateolabrax maculatus) Testis and Their Responses to Hormones and Salinity Stimulations
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摘要
In this study, three cDNA sequences corresponding to cytochrome P450 C17(CYP17 Ⅰ), 3β-hydroxysteroid dehydrogenase(3β-HSD) and androgen receptor(AR) were isolated from spotted sea bass(Lateolabrax maculatus). The mRNA abundances of CYP17 Ⅰ and 3β-HSD increased from stage Ⅱ to stage V with a significant increase at stage Ⅴ, and the highest abundance of AR mRNA was detected at stage Ⅲ in testicular development cycle. CYP17 I, 3β-HSD and AR transcripts were obviously abundant in steroidogenesis tissues such as testis, brain, head kidney among others. Strong and positive signals were observed mainly in interstitial cell regions of L. maculatus testis as were measured with in situ hybridization method. Significant increases of CYP17 Ⅰ and3β-HSD transcripts were detected after 12-48 h hCG(human chorionic gonadotropin) and GnRHa(gonadotropin-releasing hormone analogue) treatments. However, an opposite relationship was found for AR in testis at the same time. In addition, decreasing trends of CYP17 Ⅰ and 3β-HSD mRNA were observed in testis of L. maculatus in freshwater group(FW) from day 2 to day 6, and mRNA abundance of AR increased in brackish water(BW) group from day 4 to day 8. These findings revealed that these three steroid synthesis genes are import for testicular development, hormone and salinity treatment, and provided also an insight into the mechanism of reproductive endocrine of L. maculatus.
In this study, three cDNA sequences corresponding to cytochrome P450 C17(CYP17 Ⅰ), 3β-hydroxysteroid dehydrogenase(3β-HSD) and androgen receptor(AR) were isolated from spotted sea bass(Lateolabrax maculatus). The mRNA abundances of CYP17 Ⅰ and 3β-HSD increased from stage Ⅱ to stage V with a significant increase at stage Ⅴ, and the highest abundance of AR mRNA was detected at stage Ⅲ in testicular development cycle. CYP17 Ⅰ, 3β-HSD and AR transcripts were obviously abundant in steroidogenesis tissues such as testis, brain, head kidney among others. Strong and positive signals were observed mainly in interstitial cell regions of L. maculatus testis as were measured with in situ hybridization method. Significant increases of CYP17 Ⅰ and3β-HSD transcripts were detected after 12-48 h hCG(human chorionic gonadotropin) and GnRHa(gonadotropin-releasing hormone analogue) treatments. However, an opposite relationship was found for AR in testis at the same time. In addition, decreasing trends of CYP17 Ⅰ and 3β-HSD mRNA were observed in testis of L. maculatus in freshwater group(FW) from day 2 to day 6, and mRNA abundance of AR increased in brackish water(BW) group from day 4 to day 8. These findings revealed that these three steroid synthesis genes are import for testicular development, hormone and salinity treatment, and provided also an insight into the mechanism of reproductive endocrine of L. maculatus.
In this study, three cDNA sequences corresponding to cytochrome P450 C17(CYP17 Ⅰ), 3β-hydroxysteroid dehydrogenase(3β-HSD) and androgen receptor(AR) were isolated from spotted sea bass(Lateolabrax maculatus). The mRNA abundances of CYP17 Ⅰ and 3β-HSD increased from stage Ⅱ to stage V with a significant increase at stage Ⅴ, and the highest abundance of AR mRNA was detected at stage Ⅲ in testicular development cycle. CYP17 Ⅰ, 3β-HSD and AR transcripts were obviously abundant in steroidogenesis tissues such as testis, brain, head kidney among others. Strong and positive signals were observed mainly in interstitial cell regions of L. maculatus testis as were measured with in situ hybridization method. Significant increases of CYP17 Ⅰ and3β-HSD transcripts were detected after 12-48 h hCG(human chorionic gonadotropin) and GnRHa(gonadotropin-releasing hormone analogue) treatments. However, an opposite relationship was found for AR in testis at the same time. In addition, decreasing trends of CYP17 Ⅰ and 3β-HSD mRNA were observed in testis of L. maculatus in freshwater group(FW) from day 2 to day 6, and mRNA abundance of AR increased in brackish water(BW) group from day 4 to day 8. These findings revealed that these three steroid synthesis genes are import for testicular development, hormone and salinity treatment, and provided also an insight into the mechanism of reproductive endocrine of L. maculatus.
引文
Alderdice,D.F.,1998.Osmotic and ionic regulation in teleost eggs and larvae.In:Fish Physiology Vol.11A.Hoar,W.S.,and Randall,D.J.,eds.,Academic Press,London,163-251.
Blázquez,M.,and Piferrer,F.,2005.Sea bass(Dicentrarchus labrax)androgen receptor:cDNA cloning,tissue-specific expression,and mRNA levels during early development and sex differentiation.Molecular and Cellular Endocrinology,237(1-2):37-48.
Chen,C.,F.,Wen,H.S.,Wang,Z.P.,and He,F.,2010.Cloning and expression of P450c17-I(17a-hydroxylase/17,20-lyase)in brain and ovary during gonad development in Cynoglossus semilaevis.Fish Physiology and Biochemistry,36(4):1001-1012.
Chen,G.,Bai,Y.,Ren,L.,Zhu,D.,Li,Y.H.,Fang,M.Y.,AlKateb,H.,and Doran,O.,2015.Metabolism of androstenone,17β-Estradiol and dihydrotestosterone in primary cultured pig hepatocytes and the role of 3β-hydroxysteroid dehydrogenase in this process.PLo S One,10(1):e113194.
Chi,M.L.,Wen,H.S.,Ni,M.,He,F.,Li,J.F.,Qian,K.,Zhang,P.,Chai,S.H.,Ding,Y.X.,and Yin,X.H.,2014.Molecular identification of genes involved in testicular steroid synthesis and characterization of the responses to hormones stimulation in testis of spotted sea bass(Lateolabrax japonicas).Steroids,84(6):92-102.
Chi,M.L.,Wen,H.S.,Ni,M.,Qian,K.,Zhang,P.,and Chai,S.H.,2015.The characteristics of vasa gene from Japanese sea bass(Lateolabrax japonicas)and its response to the external hormones.Journal of Ocean University of China,14(4):717-723.
Choi,C,Y.,An,K.W.,Jo,P.G.,Kang,D.Y.,and Chang,Y.J.,2007.Effects of gonadotropin-releasing hormone analog(GnRHa)on steroidogenic factor-1(SF-1)and estrogen receptorβ(ERβ)gene expression in the black porgy(Acanthopagrus schlegeli).Comparative Biochemistry and Physiology,Part B,147(1):82-86.
Davey,R.A.,and Grossmann,M.,2016.Androgen receptor structure,function and biology:From bench to bedside.The Clinical Biochemist Reviews,37(1):3-15.
Halm,S.,Kwon,J.Y.,Rand-Weaver,M.,Sumpter,J.P.,Pounds,N.,Hutchinson,T.H.,and Tyler,C.R.,2003.Cloning and gene expression of P450 17α-hydroxylase,17,20-lyase cDNAin the gonads and brain of the fathead minnow Pimephales promelas.General and Comparative Endocrinology,130(3):256-266.
Hoover,Z.,Weisgerber,J.N.,Pollock,M.S.,Chivers,D.P.,and Ferrari,M.C.O.,2013.Sub-lethal increases in salinity affect reproduction in fathead minnows.Science of the Total Environment,s463-464(5):334-339.
Huang,G.Y.,Ying,G.G.,Liu,S.,and Fang,Y.X.,2012.Regulation of reproduction-and biomarker-related gene expression by sex steroids in the livers and ovaries of adult female western mosquito fish(Gambusia affinis).Comparative Biochemistry and Physiology,Part A,162(1):36-43.
Ikeuchi,T.,Todo,T.,Kobayashi,T.,and Nagahama,Y.,2001.Two subtypes of androgen and progestogen receptors in fish testes.Comparative Biochemistry and Physiology,Part B,129(2-3):449-455.
Iwade,R.,Maruo,K.,Okada,G.,and Nakamura,M.,2008.Elevated expression of P450c17(CYP17)during testicular formation in the frog.General and Comparative Endocrinology,155(1):79-87.
Kazeto,Y.,Ijiri,S.,Matsubara,H.,Adachi,S.,and Yamauchi,K.,2003.Molecular cloning and characterization of 3 betahydroxysteroid dehydrogenase/Deltas-Delta4 isomerase c DNAs from Japanese eel ovary.The Journal of Steroid Biochemistry and Molecular Biology,85(1):49-56.
Kazeto,Y.,Ijiri,S.,Todo,T.,Adachi,S.,and Yamauchi,K.,2000.Molecular cloning and characterization of Japanese eel ovarian P450c17(CYP17)cDNA.General and Comparative Endocrinology,118(1):123-133.
Kemppainen,J.A.,Lane,M.V.,Sar,M.,and Wilson,E.M.,1992.Androgen receptor phosphorylation,turnover,nuclear transport and transcriptional activation.The Journal of Biological Chemistry,267(2):968-974.
Kim,S.J.,Ogasawara,K.,Park,J.G.,Takemura,A.,and Nakamura,M.,2002.Sequence and expression of androgen receptor and estrogen receptor gene in the sex types of protogynous wrasse,Halichoeres trimaculatus.General and Comparative Endocrinology,127(2):165-173.
Kobayashi,T.,Nakamura,M.,Kajiura-Kobayashi,H.,Young,G.,and Nagahama,Y.,1998.Immunolocalization of steroidogenic enzymes(P450scc,P450c17,P450arom,and 3β-HSD)in immature and mature testes of rainbow trout(Oncorhynchus mykiss).Cell and Tissue Research,292:573-577.
Kumar,S.,Ijiri,S.,and Trant,J.M.,2000.Changes in the expression of genes encoding steroidogenic enzymes in the channel catfish(Ictalurus punctatus)ovary throughout a reproductive cycle.Biology of Reproduction,63(6):1676-1682.
Leet,J.K.,Gall,H.E.,and Sepúlveda,M.S.,2011.A review of studies on androgen and estrogen exposure in fish early life stages:Effects on gene and hormonal control of sexual differentiation.Journal of Applied Toxicology,31(5):379.
Liang,Y.Q.,Huang,G.Y.,Liu,S.S.,Zhao,J.L.,Yang,Y,Y.,Chen,X,W.,Tian,F.,Jiang,Y.X.,and Ying,G.G.,2015.Longterm exposure to environmentally relevant concentrations of progesterone and norgestrel affects sex differentiation in zebrafish(Danio rerio).Aquatic Toxicology,160:172-179.
Liu,S.Z.,Qin,F.,Wang,H.P.,Wu,T.T.,Zhang,Y.Y.,Zheng,Y.,Li,M.,and Wang,Z.Z.,2012.Effects of 17α-ethinylestradiol and bisphenol A on steroidogenic messenger ribonucleic acid levels in the rare minnow gonads.Aquatic Toxicology,s122-123:19-27.
Nakamura,I.,Evans,J.C.,Kusakabe,M.,Nagahama,Y.,and Young,G.,2005.Changes in steroidogenic enzyme and steroidogenic acute regulatory protein messenger RNAs in ovarian follicles during ovarian development of rainbow trout(Oncorhyncus mykiss).General and Comparative Endocrinology,144(3):224-231.
Ogino,Y.,Katoh,H.,and Yamada,G.,2004.Androgen dependent development of a modified anal fin,gonopodium,as a model to understand the mechanism of secondary sexual character expression in vertebrates.FEBS Letters,575(1-3):119-126.
Raghuveer,K.,Garhwal,R.,Wang,D.S.,Bogerd,J.,Kirubagaran,R.,Rasheeda,M.K.,Sreenivasulu,G.,Bhattachrya,N.,Tarangini,S.,Nagahama,Y.,and Senthilkumaran,B.,2005.Effect of methyl testosterone-and ethynyl estradiol-induced sex differentiation on catfish,Clarias gariepinus:Expression profiles of DMRT1,Cytochrome P450aromatases and 3 betahydroxysteroid dehydrogenase.Fish Physiology and Biochemistry,31(2-3):143-147.
Schofield,P.J.,Peterson,M.S.,Lowe,M.R.,Brown-Peterson,N.J.,and Slack,W.T.,2011.Survival,growth and reproduction of non-indigenous Nile tilapia,Oreochromis niloticus(Linnaeus 1758).I.Physiological capabilities in various temperatures and salinities.Marine and Freshwater Research,62(5):439-449.
Senthilkumaran,B.,Sudhakumari,C.C.,Wang,D.S.,Sreenivasulu,G.,Kobayashi,T.,Kobayashi,H.K.,Yoshikuni,M.,and Nagahama,Y.,2009.Novel 3-hydroxysteroid dehydrogenases from gonads of the Nile tilapia:Phylogenetic significance and expression during reproductive cycle.Molecular and Cellular Endocrinology,299(2):146-152.
Shi,Y.,Liu,X.C.,Zhang,H.F.,Zhang,Y.,Lu,D.Q.,and Lin,H.R.,2012.Molecular identification of an androgen receptor and its changes in mRNA levels during 17α-methyltestosterone-induced sex reversal in the orange-spotted grouper Epinephelus coioides.Comparative Biochemistry and Physiology,Part B,163(1):43-50.
Sreenivasulu,G.,Swapna,I.,Rasheed,M.K.,Ijiri,S.,Adachi,S.,Thangaraj,K.,and Senthilkumaran,B.,2005.Expression of 20b-hydroxysteroid dehydrogenase and P450 17a-hydroxylase/c17-20 lyase during h CG-induced in vitro oocyte maturation in snake head murrel Channa striatus.Fish Physiology and Biochemistry,31(2-3):227-230.
Thomas,J.L.,and Bose,H.S.,2015.Regulation of human 3-beta-hydroxysteroid dehydrogenase type-2(3βHSD2)by molecular chaperones and the mitochondrial environment affects steroidogenesis.The Journal of Steroid Biochemistry and Molecular Biology,151:74-84.
Todo,T.,Ikeuchi,T.,Kobayashi,T.,and Nagahama,Y.,1999.Fish androgen receptor:cDNA cloning,steroid activation of transcription in transfected mammalian cells,and tissue mRNAlevels.Biochemical and Biophysical Research Communications,254:378-383.
Tompsett,A.R.,Park,J.W.,Zhang,X.,Jones,P.D.,Newsted,J.L.,Au,D.W.T.,Chen,E.X.H.,Yu,R.,Wu,R.S.S.,Kong,R.Y.C,Giesy,J.P.,and Hecker,M.,2009.In situ hybridization to detect spatial gene expression in medaka.Ecotoxicology and Environmental Safety,72(4):1257-1264.
Wang,Y.,and Ge,W.,2004.Cloning of zebrafish ovarian P450c17(CYP17,17a-hydroxylase/17,20-lyase)and characterization of its expression in gonadal and extra-gonadal tissues.General and Comparative Endocrinology,135(2):241-249.
Yu,H.,Cheng,H.,Guo,Y.,Xia,L.,and Zhou,R.,2003.Alternative splicing and differential expression of P450c17(CYP17)in gonads during sex transformation in the rice field eel.Biochemical and Biophysical Research Communications,307(1):165-171.
Yu,X.G.,2013.Study on key genes 3β-HSD and StAR of the steroidgenesis in Nile tilapia.Master thesis.Southwest University,Chongqing(in Chinese with English abstract).
Zhang,M.Z.,Gao,T.X.,Ruan,S.H.,Zhuang,Q.Z.,and Qu,Y.Q.,2001.Study on artificial cultivation and spawning inducement technique of Lateolabrax japonicus.Journal of Ocean University of Qingdao,31(3):195-200(in Chinese with English abstract).
Zhou,L.Y.,Wang,D.S.,Shibata,Y.,Paul-Prasanth,B.,Suzuki,A.,and Nagahama,Y.,2007.Characterization,expression and transcriptional regulation of P450c17-I and-II in the medaka,Oryzias latipes.Biochemical and Biophysical Research Communications,362(3):619-625.
Blázquez,M.,and Piferrer,F.,2005.Sea bass(Dicentrarchus labrax)androgen receptor:cDNA cloning,tissue-specific expression,and mRNA levels during early development and sex differentiation.Molecular and Cellular Endocrinology,237(1-2):37-48.
Chen,C.,F.,Wen,H.S.,Wang,Z.P.,and He,F.,2010.Cloning and expression of P450c17-I(17a-hydroxylase/17,20-lyase)in brain and ovary during gonad development in Cynoglossus semilaevis.Fish Physiology and Biochemistry,36(4):1001-1012.
Chen,G.,Bai,Y.,Ren,L.,Zhu,D.,Li,Y.H.,Fang,M.Y.,AlKateb,H.,and Doran,O.,2015.Metabolism of androstenone,17β-Estradiol and dihydrotestosterone in primary cultured pig hepatocytes and the role of 3β-hydroxysteroid dehydrogenase in this process.PLo S One,10(1):e113194.
Chi,M.L.,Wen,H.S.,Ni,M.,He,F.,Li,J.F.,Qian,K.,Zhang,P.,Chai,S.H.,Ding,Y.X.,and Yin,X.H.,2014.Molecular identification of genes involved in testicular steroid synthesis and characterization of the responses to hormones stimulation in testis of spotted sea bass(Lateolabrax japonicas).Steroids,84(6):92-102.
Chi,M.L.,Wen,H.S.,Ni,M.,Qian,K.,Zhang,P.,and Chai,S.H.,2015.The characteristics of vasa gene from Japanese sea bass(Lateolabrax japonicas)and its response to the external hormones.Journal of Ocean University of China,14(4):717-723.
Choi,C,Y.,An,K.W.,Jo,P.G.,Kang,D.Y.,and Chang,Y.J.,2007.Effects of gonadotropin-releasing hormone analog(GnRHa)on steroidogenic factor-1(SF-1)and estrogen receptorβ(ERβ)gene expression in the black porgy(Acanthopagrus schlegeli).Comparative Biochemistry and Physiology,Part B,147(1):82-86.
Davey,R.A.,and Grossmann,M.,2016.Androgen receptor structure,function and biology:From bench to bedside.The Clinical Biochemist Reviews,37(1):3-15.
Halm,S.,Kwon,J.Y.,Rand-Weaver,M.,Sumpter,J.P.,Pounds,N.,Hutchinson,T.H.,and Tyler,C.R.,2003.Cloning and gene expression of P450 17α-hydroxylase,17,20-lyase cDNAin the gonads and brain of the fathead minnow Pimephales promelas.General and Comparative Endocrinology,130(3):256-266.
Hoover,Z.,Weisgerber,J.N.,Pollock,M.S.,Chivers,D.P.,and Ferrari,M.C.O.,2013.Sub-lethal increases in salinity affect reproduction in fathead minnows.Science of the Total Environment,s463-464(5):334-339.
Huang,G.Y.,Ying,G.G.,Liu,S.,and Fang,Y.X.,2012.Regulation of reproduction-and biomarker-related gene expression by sex steroids in the livers and ovaries of adult female western mosquito fish(Gambusia affinis).Comparative Biochemistry and Physiology,Part A,162(1):36-43.
Ikeuchi,T.,Todo,T.,Kobayashi,T.,and Nagahama,Y.,2001.Two subtypes of androgen and progestogen receptors in fish testes.Comparative Biochemistry and Physiology,Part B,129(2-3):449-455.
Iwade,R.,Maruo,K.,Okada,G.,and Nakamura,M.,2008.Elevated expression of P450c17(CYP17)during testicular formation in the frog.General and Comparative Endocrinology,155(1):79-87.
Kazeto,Y.,Ijiri,S.,Matsubara,H.,Adachi,S.,and Yamauchi,K.,2003.Molecular cloning and characterization of 3 betahydroxysteroid dehydrogenase/Deltas-Delta4 isomerase c DNAs from Japanese eel ovary.The Journal of Steroid Biochemistry and Molecular Biology,85(1):49-56.
Kazeto,Y.,Ijiri,S.,Todo,T.,Adachi,S.,and Yamauchi,K.,2000.Molecular cloning and characterization of Japanese eel ovarian P450c17(CYP17)cDNA.General and Comparative Endocrinology,118(1):123-133.
Kemppainen,J.A.,Lane,M.V.,Sar,M.,and Wilson,E.M.,1992.Androgen receptor phosphorylation,turnover,nuclear transport and transcriptional activation.The Journal of Biological Chemistry,267(2):968-974.
Kim,S.J.,Ogasawara,K.,Park,J.G.,Takemura,A.,and Nakamura,M.,2002.Sequence and expression of androgen receptor and estrogen receptor gene in the sex types of protogynous wrasse,Halichoeres trimaculatus.General and Comparative Endocrinology,127(2):165-173.
Kobayashi,T.,Nakamura,M.,Kajiura-Kobayashi,H.,Young,G.,and Nagahama,Y.,1998.Immunolocalization of steroidogenic enzymes(P450scc,P450c17,P450arom,and 3β-HSD)in immature and mature testes of rainbow trout(Oncorhynchus mykiss).Cell and Tissue Research,292:573-577.
Kumar,S.,Ijiri,S.,and Trant,J.M.,2000.Changes in the expression of genes encoding steroidogenic enzymes in the channel catfish(Ictalurus punctatus)ovary throughout a reproductive cycle.Biology of Reproduction,63(6):1676-1682.
Leet,J.K.,Gall,H.E.,and Sepúlveda,M.S.,2011.A review of studies on androgen and estrogen exposure in fish early life stages:Effects on gene and hormonal control of sexual differentiation.Journal of Applied Toxicology,31(5):379.
Liang,Y.Q.,Huang,G.Y.,Liu,S.S.,Zhao,J.L.,Yang,Y,Y.,Chen,X,W.,Tian,F.,Jiang,Y.X.,and Ying,G.G.,2015.Longterm exposure to environmentally relevant concentrations of progesterone and norgestrel affects sex differentiation in zebrafish(Danio rerio).Aquatic Toxicology,160:172-179.
Liu,S.Z.,Qin,F.,Wang,H.P.,Wu,T.T.,Zhang,Y.Y.,Zheng,Y.,Li,M.,and Wang,Z.Z.,2012.Effects of 17α-ethinylestradiol and bisphenol A on steroidogenic messenger ribonucleic acid levels in the rare minnow gonads.Aquatic Toxicology,s122-123:19-27.
Nakamura,I.,Evans,J.C.,Kusakabe,M.,Nagahama,Y.,and Young,G.,2005.Changes in steroidogenic enzyme and steroidogenic acute regulatory protein messenger RNAs in ovarian follicles during ovarian development of rainbow trout(Oncorhyncus mykiss).General and Comparative Endocrinology,144(3):224-231.
Ogino,Y.,Katoh,H.,and Yamada,G.,2004.Androgen dependent development of a modified anal fin,gonopodium,as a model to understand the mechanism of secondary sexual character expression in vertebrates.FEBS Letters,575(1-3):119-126.
Raghuveer,K.,Garhwal,R.,Wang,D.S.,Bogerd,J.,Kirubagaran,R.,Rasheeda,M.K.,Sreenivasulu,G.,Bhattachrya,N.,Tarangini,S.,Nagahama,Y.,and Senthilkumaran,B.,2005.Effect of methyl testosterone-and ethynyl estradiol-induced sex differentiation on catfish,Clarias gariepinus:Expression profiles of DMRT1,Cytochrome P450aromatases and 3 betahydroxysteroid dehydrogenase.Fish Physiology and Biochemistry,31(2-3):143-147.
Schofield,P.J.,Peterson,M.S.,Lowe,M.R.,Brown-Peterson,N.J.,and Slack,W.T.,2011.Survival,growth and reproduction of non-indigenous Nile tilapia,Oreochromis niloticus(Linnaeus 1758).I.Physiological capabilities in various temperatures and salinities.Marine and Freshwater Research,62(5):439-449.
Senthilkumaran,B.,Sudhakumari,C.C.,Wang,D.S.,Sreenivasulu,G.,Kobayashi,T.,Kobayashi,H.K.,Yoshikuni,M.,and Nagahama,Y.,2009.Novel 3-hydroxysteroid dehydrogenases from gonads of the Nile tilapia:Phylogenetic significance and expression during reproductive cycle.Molecular and Cellular Endocrinology,299(2):146-152.
Shi,Y.,Liu,X.C.,Zhang,H.F.,Zhang,Y.,Lu,D.Q.,and Lin,H.R.,2012.Molecular identification of an androgen receptor and its changes in mRNA levels during 17α-methyltestosterone-induced sex reversal in the orange-spotted grouper Epinephelus coioides.Comparative Biochemistry and Physiology,Part B,163(1):43-50.
Sreenivasulu,G.,Swapna,I.,Rasheed,M.K.,Ijiri,S.,Adachi,S.,Thangaraj,K.,and Senthilkumaran,B.,2005.Expression of 20b-hydroxysteroid dehydrogenase and P450 17a-hydroxylase/c17-20 lyase during h CG-induced in vitro oocyte maturation in snake head murrel Channa striatus.Fish Physiology and Biochemistry,31(2-3):227-230.
Thomas,J.L.,and Bose,H.S.,2015.Regulation of human 3-beta-hydroxysteroid dehydrogenase type-2(3βHSD2)by molecular chaperones and the mitochondrial environment affects steroidogenesis.The Journal of Steroid Biochemistry and Molecular Biology,151:74-84.
Todo,T.,Ikeuchi,T.,Kobayashi,T.,and Nagahama,Y.,1999.Fish androgen receptor:cDNA cloning,steroid activation of transcription in transfected mammalian cells,and tissue mRNAlevels.Biochemical and Biophysical Research Communications,254:378-383.
Tompsett,A.R.,Park,J.W.,Zhang,X.,Jones,P.D.,Newsted,J.L.,Au,D.W.T.,Chen,E.X.H.,Yu,R.,Wu,R.S.S.,Kong,R.Y.C,Giesy,J.P.,and Hecker,M.,2009.In situ hybridization to detect spatial gene expression in medaka.Ecotoxicology and Environmental Safety,72(4):1257-1264.
Wang,Y.,and Ge,W.,2004.Cloning of zebrafish ovarian P450c17(CYP17,17a-hydroxylase/17,20-lyase)and characterization of its expression in gonadal and extra-gonadal tissues.General and Comparative Endocrinology,135(2):241-249.
Yu,H.,Cheng,H.,Guo,Y.,Xia,L.,and Zhou,R.,2003.Alternative splicing and differential expression of P450c17(CYP17)in gonads during sex transformation in the rice field eel.Biochemical and Biophysical Research Communications,307(1):165-171.
Yu,X.G.,2013.Study on key genes 3β-HSD and StAR of the steroidgenesis in Nile tilapia.Master thesis.Southwest University,Chongqing(in Chinese with English abstract).
Zhang,M.Z.,Gao,T.X.,Ruan,S.H.,Zhuang,Q.Z.,and Qu,Y.Q.,2001.Study on artificial cultivation and spawning inducement technique of Lateolabrax japonicus.Journal of Ocean University of Qingdao,31(3):195-200(in Chinese with English abstract).
Zhou,L.Y.,Wang,D.S.,Shibata,Y.,Paul-Prasanth,B.,Suzuki,A.,and Nagahama,Y.,2007.Characterization,expression and transcriptional regulation of P450c17-I and-II in the medaka,Oryzias latipes.Biochemical and Biophysical Research Communications,362(3):619-625.