半滑舌鳎性别相关基因P450芳香化酶、FTZ-F1和DMRT1基因克隆及表达分析
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摘要
遗传基因作为性别分化的基础,提供了两性分化的可能性,而性分化则是在各种遗传基因和外部环境因子的相互作用下实现的。在本研究中通过外源激素甲基睾酮(MT)和温度处理半滑舌鳎获得性逆转个体,分析其中性别相关基因的表达变化情况有助于了解性别分化的作用机理。半滑舌鳎是近年来新开发的一种理想的增养殖对象,雌性个体生长速度比雄性快2-3倍。探索温度和MT对该鱼的性别决定和性别分化的影响,获得性逆转亲本,用于培育全雌后代,具有重要的应用前景和经济效益。
1温度和MT对半滑舌鳎性腺分化的影响
通过石蜡组织切片,对半滑舌鳎早期性腺分化进行组织学观察发现:在24℃饲养条件下,孵化后30d半滑舌鳎性腺开始分化,其中具有裂隙的性腺原基未来发育为卵巢,而不具裂隙的原基未来发育为精巢。在孵化后25-100d对半滑舌鳎进行不同温度(16℃、20℃、24℃、28℃、32℃)处理以及采用不同浓度梯度(20μg/L·H2O、30μg/L·H2O、50μg/L·H2O、80μg/L·H2O、100μg/L·H2O)的雄激素甲基睾酮(MT)浸浴处理,在9月龄时利用石蜡组织切片鉴定表型性别,观察温度和MT对性腺分化的影响,结果表明:28℃和32℃高温能显著提高群体中的雄性比例,分别达到69.2%(0.010.05)、57.1%(P>0.05);24℃处理群体中雌雄个体比例接近1∶1。在20μg/L·H2O、30μg/L·H2O、50μg/L·H2O、80μg/L·H2O、100μg/L·H2O的MT浸浴处理都能显著提高群体中的雄性比例,分别达到100%(P<0.01)、97.7%(P<0.01)、100%(P<0.01)、97.4%(P<0.01)、100%(P<0.01)。半滑舌鳎雌性特异的遗传性别鉴定技术也检测到高温处理组及MT处理出现了性逆转的雄性个体,这些结果表明在孵化后25-100d采用20-100μg/L·H2O浓度的甲基睾酮浸浴处理和28℃、32℃高温处理半滑舌鳎鱼苗,都能有效的诱导半滑舌鳎发生由雌性向雄性的性别逆转。
2半滑舌鳎性腺型芳香化酶(tsP450aromA)基因和脑型芳香化酶(tsP450aromB)基因的克隆及表达分析
采用同源克隆策略和RACE的方法,分别从半滑舌鳎卵巢和脑中分离了性腺型芳香化酶(tsP450aromA)和脑型芳香化酶(tsP450aromB)。tsP450aromA cDNA全长2266bp,该基因编码了526个氨基酸。氨基酸序列和系统发生分析表明,tsP450aromA属于卵巢型P450aromA,tsP450aromA的氨基酸序列与其它鱼类P450aromA的同源性较高(59%～77%),与脑型P450aromB的同源性较低(56%～60.7%)。Genome walking获得的启动子序列分析发现具TATA框和潜在的转录调节因子,包括半个雌激素应答元件(ERE half),两个Ad4-binding motifs。RT-PCR分析表明:tsP450aromA的表达具有明显组织特异性,tsP450aromA只在性腺中表达,且卵巢中表达量远高于精巢,而在雌雄鱼的其它组织中都不表达。在性腺发育过程中,精巢和卵巢中的芳香化酶的表达都逐渐增强,但卵巢中的表达量始终高于精巢。经过甲基睾酮浸浴处理和高温诱导半滑舌鳎由雌性性逆转为雄性后,性腺中芳香化酶的表达量降低。这些结果表明tsP450aromA参与了半滑舌鳎的性腺发育和性别决定过程。(tsP450aromB)cDNA全长2184 bp,该基因编码了498个氨基酸。氨基酸序列和系统发生分析表明,tsP450aromB属于脑型P450arom,tsP450aromB的氨基酸序列与其它硬骨鱼类的脑型P450aromB的同源性较高(48.3%～66.1%),与性腺型P450aromA的同源性较低(34.2%～49.9%),与自身的性腺型芳香化酶同源性为45.1%。RT-PCR分析表明:tsP450aromB mRNA的表达具有明显组织特异性,tsP450aromB只在性腺、脑、鳃、皮肤中表达,且脑中表达量远高于性腺,而在雌雄鱼的其它组织中都不表达。经过甲基睾酮浸浴处理和高温诱导半滑舌鳎由雌性性反转为雄性后,脑中tsP450aromB的表达量降低,这些结果表明tsP450aromB参与了半滑舌鳎的性腺分化和性别决定过程。
3半滑舌鳎FTZ-F1(tsFTZ-F1)基因的克隆及表达分析
采用同源克隆策略和RACE的方法,从半滑舌鳎精巢分离了3143 bp长的半滑舌鳎FTZ-F1(tsFTZ-F1)的全长cDNA,该序列包含1458bp开放阅读框(ORF),66 bp长的5’末端非编码区(UTR), 1619 bp长的3’末端UTR。mRNA的组织分布、氨基酸序列、功能分析和系统发生分析表明:tsFTZ-F1属于SF-1/Ad4BP类群。RT-PCR分析表明:tsFTZ-F1 mRNA的分布广泛,几乎在所有组织都有表达,但在性腺、肾脏、脑、头肾组织中表达最强,其他组织表达较弱,雌鱼脑和头肾中的表达量明显高于雄性。胚胎发育过程中表达量都高于孵化后仔鱼的表达量,表明tsFTZ-F1参与了半滑舌鳎的器官形成过程。经过甲基睾酮浸浴处理后,由雌性转变为雄性的性逆转雄性个体中,性腺中tsFTZ-F1和tsP450aromA的表达量降低。这些结果表明,tsFTZ-F1参与了半滑舌鳎的性别逆转过程,它的作用途径可能是作为转录调节因子,结合于芳香化酶的启动子实现对芳香化酶的表达调控。
4半滑舌鳎DMRT1(tsDMRT1)基因的克隆及表达分析
采用同源克隆策略和5’-RACE,从半滑舌鳎精巢分离了608 bp长的tsDMRT1基因cDNA的部分序列,含45 bp的5’-UTR和起始密码子,该片断包含了187个氨基酸残基,其中包括DMRT1基因保守的DM结构域。RT-PCR分析表明:tsDMRT1只在精巢中表达,其它组织中都不表达。经过甲基睾酮浸浴处理和高温诱导半滑舌鳎由雌性性逆转为雄性后,精巢中tsDMRT1也有表达。这些结果表明tsDMRT1参与了半滑舌鳎的精巢分化过程。
The sex related genes as the foundation of sex differentiation provide the possibility of sex differentiation, but the sex differentiation is the reciprocity of sex related genes and the environmental factors. With the sex reversal obtained by MT immersion and temperature treatment during sex differentiation, it will be helpful to understand the mechanism of sex determination to analyze the sex-related gene expression in sex reversal. Half-smooth tongue-sole (Cynoglossus semilaevis) is a newly exploited and commercially important cultured marine flatfish, in which females grow 2-3 times faster than males. Thus, the development of all-female tongue sole stock with temperatures and MT would be of significant benefit for aquaculture. 1 Gonadal differentiation and effects of temperature and MT on sex determination in C. semilaevis
Gonadal differentiation of C. semilaevis was observed by histological sectioning. The histological differentiation occurred firstly in 30 days post hatching(dph). There were two types of gonad. The one with cavity will develop into ovary, and the other one without cavity will develop into testis. By histological sectioning to distinguish the phenotypic sex and by female-specific marker to determination the genetic sex, effects of temperature and MT on gonad differentiation in tongue sole was addressed in a separate experiment. Juvenile tongue sole were grown at 16℃, 20℃, 24℃,28℃or 32℃from 25 to 100dph, respectively. High temperatures(28 or 32℃) induced phenotypic sex reversal in juvenile tongue sole producing a higher proportion of males ( 69.2% males at 28℃, 0.01
1温度和MT对半滑舌鳎性腺分化的影响
通过石蜡组织切片,对半滑舌鳎早期性腺分化进行组织学观察发现:在24℃饲养条件下,孵化后30d半滑舌鳎性腺开始分化,其中具有裂隙的性腺原基未来发育为卵巢,而不具裂隙的原基未来发育为精巢。在孵化后25-100d对半滑舌鳎进行不同温度(16℃、20℃、24℃、28℃、32℃)处理以及采用不同浓度梯度(20μg/L·H2O、30μg/L·H2O、50μg/L·H2O、80μg/L·H2O、100μg/L·H2O)的雄激素甲基睾酮(MT)浸浴处理,在9月龄时利用石蜡组织切片鉴定表型性别,观察温度和MT对性腺分化的影响,结果表明:28℃和32℃高温能显著提高群体中的雄性比例,分别达到69.2%(0.01
2半滑舌鳎性腺型芳香化酶(tsP450aromA)基因和脑型芳香化酶(tsP450aromB)基因的克隆及表达分析
采用同源克隆策略和RACE的方法,分别从半滑舌鳎卵巢和脑中分离了性腺型芳香化酶(tsP450aromA)和脑型芳香化酶(tsP450aromB)。tsP450aromA cDNA全长2266bp,该基因编码了526个氨基酸。氨基酸序列和系统发生分析表明,tsP450aromA属于卵巢型P450aromA,tsP450aromA的氨基酸序列与其它鱼类P450aromA的同源性较高(59%～77%),与脑型P450aromB的同源性较低(56%～60.7%)。Genome walking获得的启动子序列分析发现具TATA框和潜在的转录调节因子,包括半个雌激素应答元件(ERE half),两个Ad4-binding motifs。RT-PCR分析表明:tsP450aromA的表达具有明显组织特异性,tsP450aromA只在性腺中表达,且卵巢中表达量远高于精巢,而在雌雄鱼的其它组织中都不表达。在性腺发育过程中,精巢和卵巢中的芳香化酶的表达都逐渐增强,但卵巢中的表达量始终高于精巢。经过甲基睾酮浸浴处理和高温诱导半滑舌鳎由雌性性逆转为雄性后,性腺中芳香化酶的表达量降低。这些结果表明tsP450aromA参与了半滑舌鳎的性腺发育和性别决定过程。(tsP450aromB)cDNA全长2184 bp,该基因编码了498个氨基酸。氨基酸序列和系统发生分析表明,tsP450aromB属于脑型P450arom,tsP450aromB的氨基酸序列与其它硬骨鱼类的脑型P450aromB的同源性较高(48.3%～66.1%),与性腺型P450aromA的同源性较低(34.2%～49.9%),与自身的性腺型芳香化酶同源性为45.1%。RT-PCR分析表明:tsP450aromB mRNA的表达具有明显组织特异性,tsP450aromB只在性腺、脑、鳃、皮肤中表达,且脑中表达量远高于性腺,而在雌雄鱼的其它组织中都不表达。经过甲基睾酮浸浴处理和高温诱导半滑舌鳎由雌性性反转为雄性后,脑中tsP450aromB的表达量降低,这些结果表明tsP450aromB参与了半滑舌鳎的性腺分化和性别决定过程。
3半滑舌鳎FTZ-F1(tsFTZ-F1)基因的克隆及表达分析
采用同源克隆策略和RACE的方法,从半滑舌鳎精巢分离了3143 bp长的半滑舌鳎FTZ-F1(tsFTZ-F1)的全长cDNA,该序列包含1458bp开放阅读框(ORF),66 bp长的5’末端非编码区(UTR), 1619 bp长的3’末端UTR。mRNA的组织分布、氨基酸序列、功能分析和系统发生分析表明:tsFTZ-F1属于SF-1/Ad4BP类群。RT-PCR分析表明:tsFTZ-F1 mRNA的分布广泛,几乎在所有组织都有表达,但在性腺、肾脏、脑、头肾组织中表达最强,其他组织表达较弱,雌鱼脑和头肾中的表达量明显高于雄性。胚胎发育过程中表达量都高于孵化后仔鱼的表达量,表明tsFTZ-F1参与了半滑舌鳎的器官形成过程。经过甲基睾酮浸浴处理后,由雌性转变为雄性的性逆转雄性个体中,性腺中tsFTZ-F1和tsP450aromA的表达量降低。这些结果表明,tsFTZ-F1参与了半滑舌鳎的性别逆转过程,它的作用途径可能是作为转录调节因子,结合于芳香化酶的启动子实现对芳香化酶的表达调控。
4半滑舌鳎DMRT1(tsDMRT1)基因的克隆及表达分析
采用同源克隆策略和5’-RACE,从半滑舌鳎精巢分离了608 bp长的tsDMRT1基因cDNA的部分序列,含45 bp的5’-UTR和起始密码子,该片断包含了187个氨基酸残基,其中包括DMRT1基因保守的DM结构域。RT-PCR分析表明:tsDMRT1只在精巢中表达,其它组织中都不表达。经过甲基睾酮浸浴处理和高温诱导半滑舌鳎由雌性性逆转为雄性后,精巢中tsDMRT1也有表达。这些结果表明tsDMRT1参与了半滑舌鳎的精巢分化过程。
The sex related genes as the foundation of sex differentiation provide the possibility of sex differentiation, but the sex differentiation is the reciprocity of sex related genes and the environmental factors. With the sex reversal obtained by MT immersion and temperature treatment during sex differentiation, it will be helpful to understand the mechanism of sex determination to analyze the sex-related gene expression in sex reversal. Half-smooth tongue-sole (Cynoglossus semilaevis) is a newly exploited and commercially important cultured marine flatfish, in which females grow 2-3 times faster than males. Thus, the development of all-female tongue sole stock with temperatures and MT would be of significant benefit for aquaculture. 1 Gonadal differentiation and effects of temperature and MT on sex determination in C. semilaevis
Gonadal differentiation of C. semilaevis was observed by histological sectioning. The histological differentiation occurred firstly in 30 days post hatching(dph). There were two types of gonad. The one with cavity will develop into ovary, and the other one without cavity will develop into testis. By histological sectioning to distinguish the phenotypic sex and by female-specific marker to determination the genetic sex, effects of temperature and MT on gonad differentiation in tongue sole was addressed in a separate experiment. Juvenile tongue sole were grown at 16℃, 20℃, 24℃,28℃or 32℃from 25 to 100dph, respectively. High temperatures(28 or 32℃) induced phenotypic sex reversal in juvenile tongue sole producing a higher proportion of males ( 69.2% males at 28℃, 0.01
males at 32℃, 0.01
mperature(16℃, 20℃) induced phenotypic sex reversal and produced a slightly high proportion of males(56.5% males at 16℃,P>0.05; 57.1% males at 20℃, P>0.05). Raising tongue sole at the temperature(24℃) held sex ratios close to 1:1. Juvenile tongue sole were immersed with MT of 20μg/L·H2O、30μg/L·H2O、50μg/L·H2O、80μg/L·H2O、100μg/L·H2O from 25 to 100dph, respectively. High and low concentration of MT induced phenotypic sex reversal in juvenile tongue sole producing a significantly higher proportion of males (100%(P<0.01)、97.7%(P<0.01)、100%(P<0.01)、97.4%(P<0.01)、100%(P<0.01), respectively). The female-specific marker also suggested that high temperature and MT immersion induced the sex reversal from the female to male. These findings indicate that sex differentiation in tongue sole are affected by temperature and MT.
2 Molecular cloning, characterization and expression analysis of ovarian P450arom(tsP450aromA) and brain P450arom(tsP450aromB) in C. semilaevis The 2266 bp full-length ovarian P450 aromatase (termed tsP450aromA) cDNA encoding a deduced protein of 526 residues was isolated from the ovary of the C. semilaevis by using homologous cloning and RACE. Sequence and phylogenic analyses showed that the tongue sole P450aromA belonged to ovarian P450arom subfamilies. The putative tsP450aromA amino acid residues shared 59～77% sequence identify with other fish ovarian P450arom, and shared 56～60.7% sequence identify with brain P450arom of other fish. The promoter region (5'- flanking region) of tsP450aromA gene was cloned from tongue sole containing two Ad4 binding motifs and a half site of the palindrome sequences of estrogen-responsive element (ERE half). The RT-PCR analysis reveals that tsP450aromA mRNA was expressed highly in the ovary and weakly in the testis, but not other tissues. The P450aromA mRNA increased with the gonad development, but it is less abundant in testis in all phase. However, the tsP450aromA decreased in sex reversal male treatment by MT or high temperature. These results suggested that the tsP450aromA is involved in course of the gonad development and sex determination in tongue sole.
The 2184 bp full-length brain P450 aromatase (termed tsP450aromB) cDNA encoding a protein of 498 amino acid residues was isolated from the brain of the C. semilaevis by using homologous cloning and RACE. Sequence and phylogenic analyses showed that the tsP450aromB belonged to brain P450arom subfamilies. The putative tsP450aromB amino acid residues shared 48.3～66.1% sequence identify with other fish brain P450arom, and shared 34.2～49.9% sequence identify with ovarian P450arom of other fish. The sequence identify between the tsP450aromA and tsP450aromB is 45.1%. The RT-PCR analysis reveals that tsP450aromB mRNA was expressed highly in the brain and gill and weakly in the gonad, but not in other tissues. The tsP450aromB decreased in sex-reversed male treatment by MT or high temperature. These results suggested that the tsP450aromB is involved in course of the gonad development and sex determination in tongue sole.
3 Molecular cloning, characterization and expression analysis of FTZ-F1 homologue(tsFTZ-F1) in C. semilaevis
The full-length cDNA of the C. semilaevis FTZ-F1 homologue (tsFTZ-F1) was isolated from the testis by using homologous cloning, and the cDNA is 3143 bp in length, including the 5’terminal UTR of 66 bp, the encoding region of 1458 bp and the 3’terminal UTR of 1619 bp. Sequence, tissue distribution, function and phylogenic analyses of the FTZ-F1 showed that tsFTZ-F1belonged to SF-1/Ad4BP group. The RT-PCR analysis reveals that tsFTZ-F1 mRNA was highly expressed in the gonad, kidney, brain, head kidney and weakly in other tissues. However, the expression level in the brain and head kidney of female was much higher than those of male. It suggested that the tsFTZ-F1 involve in the organogenesis for the tsFTZ-F1 expression in embryo of different phase was higher aboundantly than that of larvae from hatching to 25 days post hatching. The expression of tsFTZ-F1 and tsP450aromA in the testis of sex-reversal decreased significantly in response to MT immersion, and it suggested the tsFTZ-F1 involve in the course of sex reversal. The tsFTZ-F1 probably acts as a transcriptional modulator by binding to the promoter of tsP450aromA to implement the expression of tsP450aromA.
4 Molecular cloning, characterization and expression analysis of DMRT1 (tsDMRT1) in C. semilaevis
The 608 bp cDNA of the partial C. semilaevis DMRT1 homologue (tsDMRT1) was isolated from the testis by using homologous cloning and 5’-RACE, and including the 5’terminal UTR of 45 bp and starting coden. It code a deduced protein of 187 residues, including DM dormain. The RT-PCR analysis reveals that tsDMRT1 was only expressed in the testis, but not in other tissues. The DMRT1 also expressed in sex-reversed male treatment by MT or high temperature. These results suggested that the DMRT1 is involved in course of the gonad development and testis differentiation in tongue sole.
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