大黄鱼sox9a/b基因的克隆与表达分析
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摘要
为探究sox9基因在大黄鱼性别决定与分化过程中的作用,利用RACE方法克隆得到大黄鱼sox9a和sox9b 2个基因,采用实时荧光定量PCR (qRT-PCR)分析其在雌雄个体不同组织和不同发育阶段的表达规律,并检测了其在17β-雌二醇处理后的遗传雄性和17α-甲基睾酮诱导处理后遗传雌性幼鱼中的表达变化。结果显示,大黄鱼sox9a c DNA全长2 442 bp(NCBI登录号:MH996431),包含476 bp 5′UTR、466 bp 3′UTR、1 500 bp ORF,编码499个氨基酸。大黄鱼sox9b cDNA全长为2 199 bp (NCBI登录号:MH996432),包含335 bp5′UTR、415 bp 3′UTR、1 449 bp ORF,编码482个氨基酸。qRT-PCR分析显示,sox9a基因主要在性腺、眼、脑、肝脏中表达,精巢中的表达量显著高于卵巢;sox9b在大黄鱼各组织中广泛表达,但以精巢中表达量最高,而卵巢中只有痕量表达。在鱼苗性腺发育初期,sox9a/b的表达量都维持在较低水平;随着鱼苗长大,sox9a/b的表达量在84 dph、123 dph 2次达到高峰,随后开始下降,10 mph后又逐渐上升。17β-雌二醇处理能够显著下调遗传雄性大黄鱼sox9a和sox9b基因在性腺中的表达,17α-甲基睾酮处理则显著上调遗传雌性大黄鱼sox9a和sox9b基因在性腺中的表达。研究表明,sox9a/b基因与大黄鱼性别发育和分化过程密切相关,对大黄鱼精巢的发育与维持起重要的调控作用,而且两个基因的功能可能具有一定程度的分化。
To elucidate the role of sox9in sex determination and differentiation of the large yellow croaker(Larimichthys crocea), the full length of sox9a and sox9b were cloned using reverse transcription-polymerase chain reaction(qRT-PCR) and rapid amplification of cDNA ends(RACE). The difference of the gene expression in various tissues and development stages was analyzed through quantitative real-time PCR. Expression profiles of sox9a/b after 17β-estradiol or 17α-methyl testosterone treatments were also examined. The full-length cDNA of sox9a gene is 2 442 bp(NCBI: MH996431), including a 476 bp 5′ UTR, a 466 bp 3′ UTR and a 1 500 bp ORF coding a polypeptide of 499 amino acids. The full-length cDNA of sox9b gene is 2 199 bp(NCBI: MH996432),including a 335 bp 5′ UTR, a 415 bp 3′ UTR and a 1 449 bp ORF coding a polypeptide of 482 amino acids.Quantitative Real-time PCR results showed that sox9a was primarily expressed in gonad, eye, brain, liver, and the expression level in testis was significantly higher than that in ovaries. sox9b was widely expressed in multiple tissues in large yellow croaker; the expression level was the highest in testes, but can be barely detected in ovaries.At early developmental stages of fry, sox9a/b was expressed at a lower level. Sox9 a/b peaked at 84 dph(day post hatch) and 123 dph, then their expression declined and gradually rose again at 10 mph. In addition, 17β-estradiol can significantly down-regulate the expression of sox9a and sox9b in testes. 17α-methyl testosterone can significantly elevate the expression of sox9a and sox9b in gonads. The study demonstrated that sox9a/b may play important roles in sex determination and differentiation in the large yellow croaker. However, the functions of the two genes may be different.
To elucidate the role of sox9in sex determination and differentiation of the large yellow croaker(Larimichthys crocea), the full length of sox9a and sox9b were cloned using reverse transcription-polymerase chain reaction(qRT-PCR) and rapid amplification of cDNA ends(RACE). The difference of the gene expression in various tissues and development stages was analyzed through quantitative real-time PCR. Expression profiles of sox9a/b after 17β-estradiol or 17α-methyl testosterone treatments were also examined. The full-length cDNA of sox9a gene is 2 442 bp(NCBI: MH996431), including a 476 bp 5′ UTR, a 466 bp 3′ UTR and a 1 500 bp ORF coding a polypeptide of 499 amino acids. The full-length cDNA of sox9b gene is 2 199 bp(NCBI: MH996432),including a 335 bp 5′ UTR, a 415 bp 3′ UTR and a 1 449 bp ORF coding a polypeptide of 482 amino acids.Quantitative Real-time PCR results showed that sox9a was primarily expressed in gonad, eye, brain, liver, and the expression level in testis was significantly higher than that in ovaries. sox9b was widely expressed in multiple tissues in large yellow croaker; the expression level was the highest in testes, but can be barely detected in ovaries.At early developmental stages of fry, sox9a/b was expressed at a lower level. Sox9 a/b peaked at 84 dph(day post hatch) and 123 dph, then their expression declined and gradually rose again at 10 mph. In addition, 17β-estradiol can significantly down-regulate the expression of sox9a and sox9b in testes. 17α-methyl testosterone can significantly elevate the expression of sox9a and sox9b in gonads. The study demonstrated that sox9a/b may play important roles in sex determination and differentiation in the large yellow croaker. However, the functions of the two genes may be different.
引文
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[3]Wagner T,Wirth J,Meyer J,et al.Autosomal sex reversal and campomelic dysplasia are caused by mutations in and around the SRY-related gene SOX9[J].Cell,1994,79(6):1111-1120.
[4]Barrionuevo F,Bagheri-Fam S,Klattig J,et al.Homozygous inactivation of Sox9 causes complete XYsex reversal in mice[J].Biology of Reproduction,2006,74(1):195-201.
[5]Qin Y J,Kong L K,Poirier C,et al.Long-range activation of Sox9 in Odd sex(Ods)mice[J].Human Molecular Genetics,2004,13(12):1213-1218.
[6]Qin Y J,Bishop C E.Sox9 is sufficient for functional testis development producing fertile male mice in the absence of Sry[J].Human Molecular Genetics,2005,14(9):1221-1229.
[7]Kim G J,Sock E,Buchberger A,et al.Copy number variation of two separate regulatory regions upstream of SOX9 causes isolated 46,XY or 46,XX disorder of sex development[J].Journal of Medical Genetics,2015,52(4):240-247.
[8]Cox J J,Willatt L,Homfray T,et al.A SOX9 duplication and familial 46,XX developmental testicular disorder[J].The New England Journal of Medicine,2011,364(1):91-93.
[9]Vidal V P I,Chaboissier M C,De Rooij D G,et al.Sox9induces testis development in XX transgenic mice[J].Nature Genetics,2001,28(3):216-217.
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[12]文爱韵,尤锋,孙鹏,等.牙鲆sox9基因的克隆与表达研究[J].海洋科学进展,2011,29(1):97-104.Wen A Y,You F,Sun P,et al.Study on cloning and expression of gene sox9 in olive flounder(Paralichthys olivaceus)[J].Advances in Marine Science,2011,29(1):97 -104(in Chinese).
[13]Takamatsu N,Kanda H,Ito M,et al.Rainbow trout SOX9:cDNA cloning,gene structure and expression[J].Gene,1997,202(1-2):167-170.
[14]杜启艳,常重杰,王凤羽,等.鲤鱼中Sox9b基因的克隆和表达[J].实验生物学报,2005,38(5):397-403.Du Q Y,Chang Z J,Wang F Y,et al.The cloning and expression analysis of Sox9b in Cyprinus carpio[J].Acta Biologiae Experimentalis Sinica,2005,38(5):397-403(in Chinese).
[15]俞菊华,李建林,曹丽萍,等.黄颡鱼sox9基因的分离及分析[J].农业生物技术学报,2005,13(5):620-623.Yu J H,Li J L,Cao L P,et al.Isolation and analysis of sox9 derived from Yellow Catfish(Pelteobagrus fulvidraco)[J].Journal of Agricultural Biotechnology,2005,13(5):620-623(in Chinese).
[16]陈建华,何毛贤,牟幸江,等.金钱鱼sox9 cDNA克隆及其表达分析[J].动物学杂志,2015,50(1):93-102.Chen J H,He M X,Mu X J,et al.cDNA cloning and mRNA expression analysis of sox9 in Scatophagus argus[J].Chinese Journal of Zoology,2015,50(1):93-102(in Chinese).
[17]贺超,洪广,吴静娴,等.泰国斗鱼sox9基因的克隆及组织表达研究[J].安徽农业科学,2017,45(8):151-155,179.He C,Hong G,Wu J X,et al.Study on cloning and tissue expression of sox9 in Betta splendens[J].Journal of Anhui Agricultural Sciences,2017,45(8):151-155,179(in Chinese).
[18]Luo Y S,Hu W,Liu X C,et al.Molecular cloning and mRNA expression pattern of sox9 during sex reversal in orange-spotted grouper(Epinephelus coioides)[J].Aquaculture,2010,306(1/2/3/4):322-328.
[19]Kobayashi T,Kajiura-Kobayashi H,Guan G J,et al.Sexual dimorphic expression of DMRT1 and sox9a during gonadal differentiation and hormone-induced sex reversal in the teleost fish Nile tilapia(Oreochromis niloticus)[J].Developmental Dynamics,2008,237(1):297-306.
[20]常重杰,周荣家,余其兴.大鳞副泥鳅中Sox9基因保守区的序列分析[J].遗传学报,2000,27(2):121-126.Chang Z J,Zhou R J,Yu Q X.The conservative region sequence analysis of PdSox9 in Paramisgurnus dabryanus[J].Acta Genetica Sinica,2000,27(2):121-126 (in Chinese).
[21]Zhou R J,Liu L,Guo Y Q,et al.Similar gene structure of two Sox9a genes and their expression patterns during gonadal differentiation in a teleost fish,rice field eel(Monopterus albus)[J].Molecular Reproduction and Development,2003,66(3):211-217.
[22]Johnsen H,Tveiten H,Torgersen J S,et al.Divergent and sex-dimorphic expression of the paralogs of the Sox9-Amh-Cyp19a1 regulatory cascade in developing and adult atlantic cod(Gadus morhua L.)[J].Molecular Reproduction and Development,2013,80(5):358-370.
[23]Chiang E F L,Pai C I,Wyatt M,et al.Two Sox9 genes on duplicated zebrafish chromosomes:expression of similar transcription activators in distinct sites[J].Developmental Biology,2001,231(1):149-163.
[24]Yokoi H,Kobayashi T,Tanaka M,et al.sox9 in a teleost fish,medaka(Oryzias latipes):evidence for diversified function of Sox9 in gonad differentiation[J].Molecular Reproduction and Development,2002,63(1):5-16.
[25]林爱强,谢仰杰,徐双斌,等.大黄鱼gsdf和amh基因的克隆及表达分析[J].南方水产科学,2017,13(6):1-13.Lin A Q,Xie Y J,Xu S B,et al.Cloning and expression profiling of gsdf and amh genes in large yellow croaker(Larimichthys crocea)[J].South China Fisheries Science,2017,13(6):1-13(in Chinese).
[26]陈芸,周鹏,张子平,等.大黄鱼cyp19a/b基因的克隆与表达分析[J].集美大学学报(自然科学版),2015,20(2):81-89.Chen Y,Zhou P,Zhang Z P,et al.Cloning and expression of cyp19a/b gene in large yellow croaker Larimichthys crocea[J].Journal of Jimei University(Natural Science),2015,20(2):81-89(in Chinese).
[27]周鹏.若干大黄鱼性腺发育相关基因的克隆与表达[D].厦门:集美大学,2009.Zhou P.Cloning and expression patterns of gonad related genes from large yellow croaker[D].Xiamen:Jimei University,2009(in Chinese).
[28]林晓煜,肖世俊,李完波,等.大黄鱼性别特异SNP标记的开发与验证[J].水产学报,2018,42(9):1329-1337.Lin X Y,Xiao S J,Li W B,et al.Development and validation of sex-specific SNP markers in Larimichthys crocea[J].Journal of Fisheries of China,2018,42(9):1329-1337(in Chinese).
[29]Margarit E,Coll M D,Oliva R,et al.SRY gene transferred to the long arm of the X chromosome in a Y-positive XX true hermaphrodite[J].American Journal of Medical Genetics,2000,90(1):25-28.
[30]Sekido R,Lovell-Badge R.Sex determination involves synergistic action of SRY and SF1 on a specific Sox9enhancer[J].Nature,2008,453(7197):930-934.
[31]Kim Y,Kobayashi A,Sekido R,et al.Fgf9 and Wnt4 act as antagonistic signals to regulate mammalian sex determination[J].PLoS Biology,2006,4(6):e187.
[32]梅洁,桂建芳.鱼类性别异形和性别决定的遗传基础及其生物技术操控[J].中国科学:生命科学,2015,58(2):124-136.Mei J,Gui J F.Genetic basis and biotechnological manipulation of sexual dimorphism and sex determination in fish[J].Science China Life Sciences,2015,58(2):124-136.
[33]Ospina-álvarez N,Piferrer F.Temperature-dependent sex determination in fish revisited:prevalence,a single sex ratio response pattern,and possible effects of climate change[J].PLoS ONE,2008,3(7):e2837.
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