四种细胞色素P450酶在半滑舌鳎繁殖生理机能中的作用研究
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摘要
本文首次克隆得到半滑舌鳎(Cynoglossus semilaevis) P450c17-I、P450c17-Ⅱ全长cDNA,系统研究了P450c17 (-Ⅰ、Ⅱ)、P450c19 (A, B) mRNA时空表达,并首次结合性腺发育、血清T及E2含量,探讨这几种酶类在雌雄半滑舌鳎繁殖周期中的分子生理机制。
1.人工养殖半滑舌鳎卵巢发育及产卵类型研究
人工养殖半滑舌鳎成熟鱼卵巢经历Ⅱ期、Ⅲ期、Ⅳ期、Ⅴ期、Ⅵ期,再到重复发育Ⅱ期；人工养殖条件下,3-5月份大部分鱼类处于Ⅱ期卵巢,6月份开始,卵母细胞内开始沉积卵黄颗粒,9-11月份,卵黄颗粒充盈整个胞质,基本达到生长成熟,排卵后至翌年2月份,卵巢进入退化吸收期或重复发育Ⅱ期；繁殖期GSI达到最高峰；产卵后或重复发育Ⅱ期,GSI显著下降,此时HSI显著升高至全年最高峰；半滑舌鳎在一年中只有1个繁殖期,进行分批产卵。
2.半滑舌鳎血浆性类固醇激素水平的周年变化
采用125I放射免疫法测定雌鱼血浆E2、T含量,发现其与卵巢发育过程(GSI)基本相一致。繁殖季节,E2、T为全年最高值(P<0.05)；进入越冬期后,两者含量均下降；到饲育期继续大幅下降(P<0.05)；至促熟期,E2显著升高(P<0.05),而T降至本法检测不到。结果表明,雌鱼血浆E2含量的规律性变动,与卵巢发育、卵黄形成是密切相关的；T作为E2前体,其变化趋势与卵巢发育具有相关性,是卵巢发育到一定阶段所必需。
雄鱼血浆T、E2含量与精巢发育(GSI)基本一致。最高值出现在繁殖季节,之后进入11～翌年1月份,两者含量均显著下降(P<0.05)；2～4月份T含量略微上升,E2含量则继续下降,之后到5～7月份,T、E2含量均显著升高(P<0.05)。结果表明,睾酮对精细胞的生长发育有直接作用,能够促进精子的发生和成熟。
3.半滑舌鳎两种P450c19时空表达分析
半滑舌鳎两种芳香化酶(P450arom A、P450aromB)空间表达发现,P450aromA只在性腺表达,P450aromB只在脑、鳃表达。P450aroms在各个发育期雌雄鱼的性腺和脑内均有表达。卵巢P450aromA表达模式与血清E2、T变化趋势一致,暗示P450aromA在卵巢发育和卵母细胞生长中起到了重要作用。而精巢内P450aromA时间表达却与E2、T变化趋势呈无相关性。P450aromB时间表达发现,退化吸收期雌鱼脑内表达量显著升高至全年最大值；排精后雄鱼脑内也出现类似结果。这些结果表明雌性繁殖周期很大程度上受CYP19A调控外,还受CYP19B调控。
4.新型程序化设计简并引物克隆半滑舌鳎CYP17基因片段
采用一种新的简并引物设计方法—CODEHOP法来克隆半滑舌鳎CYP17基因。该法结合NCBI、Blastp、BIockMaker、CODEHOP等生物信息学资源和DNAMAN、Oligo6.0等生物软件,程序性设计半滑舌鳎CYP17基因的CODEHOP引物,从半滑舌鳎卵巢组织中成功克隆了CYP17基因,该基因片段与其它物种CYP17家族具有较高同源性,经鉴定,该基因属于鱼类CYP17基因群。实验结果表明,程序性设计简并引物较传统简并引物特异性更高,假阳性率少,更有利于实验成功。
5.半滑舌鳎P450c17-Ⅰ克隆及时空表达分析
本研究克隆了P450c17-Ⅰ全长cDNA,为2469 bp,编码509个氨基酸。同时也得到一条较短cDNA,长1742bp,推测是经多聚腺苷酸化而来。P450c17-Ⅰ酶与胖头鱼岁、斑马鱼、日本鳗鲡、罗非鱼、三刺鱼、青鳉、白斑角鲨、鸡、鼠和人的P450c17-Ⅰ酶具有高同源性。空间表达发现：P450c17-Ⅰ主要在性腺内表达,其次是脑,在胃、肠、鳃、脾脏、肾、头肾、心内检测到弱带。性腺和脑内P450c17-Ⅰ时间表达模式表明其表达具有卵巢发育正相关性。另结合雌鱼血清T和E2水平,表明P450c17-Ⅰ在类固醇激素途径发生转变中起到了重要作用。
6.半滑舌鳎P450c17-Ⅱ克隆及时空表达分析
本研究克隆了P450c17-Ⅱ全长cDNA,为2102bp,编码524个氨基酸。半滑舌鳎P450c17-Ⅱ酶,与其他物种P450c17-Ⅱ酶同源性较其他物种P450c17-Ⅰ酶同源性高。时空表达发现其只在性腺、头肾、脑内表达。卵巢和脑内P450c17-Ⅱ时间表达模式都具有发育正相关性,而P450c17-Ⅱ在精巢内的时间表达却与性腺发育呈负相关。本研究通过性腺、脑、头肾内P450c17-Ⅱ表达,结合性腺发育期、血清中E2、T,探讨P450c17-Ⅱ与性腺发育关系,为国内鲆鲽鱼类繁殖生理学提供理论基础。
The full length of P450c17-Ⅰand P450c17-ⅡcDNA is cloned from half-smooth tongue sole Cynoglossus semilaevis, and the spatial and temporal expression of P450c17(-Ⅰ,-Ⅱ) are well studied, as well as P450c19(A,B).The results of the spatial and temporal expression of the four enzymes are firstly linked to ovarian development, serum sexual steroid hormones testosterone(T) and estradiol-17β(E2) level, so as to work over the enzymes' molecular physiological mechanism in female and male half-smooth tongue sole.
1. Studies on ovarian development and spawn type of cultured female half-smooth tongue sole Cynoglossus semilaevis
Seasonal variations of ovarian development and the relationship with Gonadosomatic index (GSI)、hepatosomatic index (HSI) was studied in the cultured half-smooth tongue sole Cynoglossue semilaevis. The results show that ovarian development consists of phaseⅡ,Ⅲ,Ⅳ,Ⅴ,Ⅵ, then cycles to recrudescent phaseⅡ; under artificial culture, most fish has phaseⅡovarium from March to May, and the oocytes begin to accumulate vitellin granule from June, then the whole cytoplasm is stuffed with vitellin granule ranging from September to November, suggesting the oocytes growth to mature, after spawning, the ovarium come to assimilative phaseⅥor recrudescent phaseⅡ;GSI reaches to an peak during reproduction, then dramatically drop after spawning or recrudescent phaseⅡwhile HSI significantly increases to maximum; there is only one breeding season in the half-smooth tongue sole, and it belongs to batch-spawning fish.
2. Study on the seasonal changes of serum sexual steroid hormones in half-smooth tongue sole Cynoglossus semilaevis
Serum sexual steroid hormones testosterone(T) and estradiol-17β(E2) levels are measured by 125I radioimmunoassay(RIA). Serum E2 and T levels are shown to be accord with ovarian developmental phase (GSI) in female hale-smooth tongue sole. During the reproductive period, serum E2 and T levels significantly increase, and reach to the highest level throughout the whole year(P<0.05). After spawn, the females live through winter, and both of serum E2 and T levels decline. In the period of artificially enriching and culturing, serum E2 level remarkably increase(P<0.05), but the content of serum T drops markedly to the value that can't be detected. The results show that seasonal changes of female half-smooth tongue sole serum E2 level is closely correlated with ovarian development and vitellus' formation; and as the precursor of E2, the changes of T level has some relationship with ovarian development, furthermore it is essential for certain ovarian developmental phase.
Serum T and E2 levels are also shown to be accord with testis development(GSI) in male half-smooth tongue sole. The peak of serum T and E2 levels both occur at reproductive season; from November to January of next year, both of the contents drop rapidly(P<0.05); during February to April, serum T level slightly increase, but the content of serum E2 still decrease; in the period of May to July, serum T and E2 levels both significantly increase(P<0.05). The results show that T directly effects on the growth of sperm cell, and can boost sperm' formation and maturation.
3. Spatial and temporal expression analysis of half-smooth tongue sole Cynoglossus semilaevis P450c19s
The spatial expression of P450aromA and P450aromB show that P450aromA only expresses in gonad, and P450aromB only expresses in brain and gill. Further, P450aroms express in the gonad and brain throughout the whole gonad development in female and male C. semilaevis, respectively. The temporal expression pattern of P450aromA is consistent with the changes of serum E2 and T level in ovaries, indicating that it plays an important role in ovarian development and oocyte growth. However, the pattern of P450aromA in testis is not related with the changes of serum E2 and T level. The results of P450aromB'temporal expression shows that, in female tongue sole of atretic follicles stage, P450aromB expression in brain increases significantly and reaches to the peak; the similar result occurs in male tongue sole brain after spermiation. These results suggested the reproductive cycle of female tongue sole is not only regulated by CYP19A, but also regulated by CYP19B.
4. Programming Design of Degenerate Primers and Cloning Half-smooth Tongue Sole Cynoglossus semilaevis CYP17 partial cDNA
A new method of designing degenerate primers-CODEHOP is introduced to clone half-smooth tongue sole Cynoglossus semilaevis CYP17 gene. With Bioinformatics Resources such as NCBI、Blastp、BlockMaker、CODEHOP and biological software such as DNAMAN、Oligo6.0, the CODEHOP primer is programming designed, and finally we succeeded to clone the gene from the ovaries of half-smooth tongue sole Cynoglossus semilaevis. The fragment of the gene has high identities with the CYP17 family of other species, and it is clustered into the CYP17 gene group of all the other vertebrates, especially into the fish subgroup. The experiment results showed that compared with traditional degenerate primers, programming designed primers had more Specific ,lower positive rate, and more benefit for accomplishing the experiment.
5. Cloning, spatial and temporal expression analysis of half-smooth tongue sole Cynoglossus semilaevis P450c17-Ⅰ
A full length cDNA of P450c17-Ⅰhas been isolated from half-smooth tongue sole (Cynoglossus semilaevis) ovary. It is 2469 bp in length, encoding 509 amino acids. Also a relative shorter cDNA (1742bp) was found, likely due to a polyadenylation. The putative P450c17-Ⅰenzyme shares high sequence identity with that of fathead minnow, zebrafish, the Japanese eel, tilapia, three spined stickleback, medaka dogfish, chicken, rat, and human. Semi-quantitative RT-PCR analysis of spatial expression shows that it was predominantly expressed in the ovary, followed by brain, and faint signals were also detected in stomach, intestine, gill, spleen, kidney, head kidney. Further examination of temporal expression pattern of P450c17-Ⅰin ovary and brain revealed developmental stage-dependency. In addition to this our data on T and E2 levels further endorse the critical role of P450c17-Ⅰduring shift in steroidogenesis.
6. Cloning, spatial and temporal expression analysis of half-smooth tongue sole Cynoglossus semilaevis P450c17-Ⅱ
A full length cDNA of P450c17-Ⅱhas been isolated from half-smooth tongue sole (Cynoglossus semilaevis) ovary. It is 2102bp in length, encoding 524 amino acids. The putative tongue sole P450c17-Ⅱenzyme shares higher sequence identity with that of other animals, than the identity of P450c17-Ⅰenzyme of the same species. Semi-quantitative RT-PCR analysis of spatial expression shows that it was specially expressed in the ovary, brain and head kidney. Further examination of temporal expression pattern of P450c17-Ⅱin ovary and brain revealed developmental stage-dependency. However, the expression pattern of P450c17-Ⅱin testis shows negative relationship with testis development. P450c17-Ⅱexpression in gonad, brain and head kidney is studied, combined with serum E2 and T levels, so as to find out the relationship between P450cl7-Ⅱand gonad development, and provide theory foundation of flatfish reproductive physiology.
1.人工养殖半滑舌鳎卵巢发育及产卵类型研究
人工养殖半滑舌鳎成熟鱼卵巢经历Ⅱ期、Ⅲ期、Ⅳ期、Ⅴ期、Ⅵ期,再到重复发育Ⅱ期；人工养殖条件下,3-5月份大部分鱼类处于Ⅱ期卵巢,6月份开始,卵母细胞内开始沉积卵黄颗粒,9-11月份,卵黄颗粒充盈整个胞质,基本达到生长成熟,排卵后至翌年2月份,卵巢进入退化吸收期或重复发育Ⅱ期；繁殖期GSI达到最高峰；产卵后或重复发育Ⅱ期,GSI显著下降,此时HSI显著升高至全年最高峰；半滑舌鳎在一年中只有1个繁殖期,进行分批产卵。
2.半滑舌鳎血浆性类固醇激素水平的周年变化
采用125I放射免疫法测定雌鱼血浆E2、T含量,发现其与卵巢发育过程(GSI)基本相一致。繁殖季节,E2、T为全年最高值(P<0.05)；进入越冬期后,两者含量均下降；到饲育期继续大幅下降(P<0.05)；至促熟期,E2显著升高(P<0.05),而T降至本法检测不到。结果表明,雌鱼血浆E2含量的规律性变动,与卵巢发育、卵黄形成是密切相关的；T作为E2前体,其变化趋势与卵巢发育具有相关性,是卵巢发育到一定阶段所必需。
雄鱼血浆T、E2含量与精巢发育(GSI)基本一致。最高值出现在繁殖季节,之后进入11～翌年1月份,两者含量均显著下降(P<0.05)；2～4月份T含量略微上升,E2含量则继续下降,之后到5～7月份,T、E2含量均显著升高(P<0.05)。结果表明,睾酮对精细胞的生长发育有直接作用,能够促进精子的发生和成熟。
3.半滑舌鳎两种P450c19时空表达分析
半滑舌鳎两种芳香化酶(P450arom A、P450aromB)空间表达发现,P450aromA只在性腺表达,P450aromB只在脑、鳃表达。P450aroms在各个发育期雌雄鱼的性腺和脑内均有表达。卵巢P450aromA表达模式与血清E2、T变化趋势一致,暗示P450aromA在卵巢发育和卵母细胞生长中起到了重要作用。而精巢内P450aromA时间表达却与E2、T变化趋势呈无相关性。P450aromB时间表达发现,退化吸收期雌鱼脑内表达量显著升高至全年最大值；排精后雄鱼脑内也出现类似结果。这些结果表明雌性繁殖周期很大程度上受CYP19A调控外,还受CYP19B调控。
4.新型程序化设计简并引物克隆半滑舌鳎CYP17基因片段
采用一种新的简并引物设计方法—CODEHOP法来克隆半滑舌鳎CYP17基因。该法结合NCBI、Blastp、BIockMaker、CODEHOP等生物信息学资源和DNAMAN、Oligo6.0等生物软件,程序性设计半滑舌鳎CYP17基因的CODEHOP引物,从半滑舌鳎卵巢组织中成功克隆了CYP17基因,该基因片段与其它物种CYP17家族具有较高同源性,经鉴定,该基因属于鱼类CYP17基因群。实验结果表明,程序性设计简并引物较传统简并引物特异性更高,假阳性率少,更有利于实验成功。
5.半滑舌鳎P450c17-Ⅰ克隆及时空表达分析
本研究克隆了P450c17-Ⅰ全长cDNA,为2469 bp,编码509个氨基酸。同时也得到一条较短cDNA,长1742bp,推测是经多聚腺苷酸化而来。P450c17-Ⅰ酶与胖头鱼岁、斑马鱼、日本鳗鲡、罗非鱼、三刺鱼、青鳉、白斑角鲨、鸡、鼠和人的P450c17-Ⅰ酶具有高同源性。空间表达发现：P450c17-Ⅰ主要在性腺内表达,其次是脑,在胃、肠、鳃、脾脏、肾、头肾、心内检测到弱带。性腺和脑内P450c17-Ⅰ时间表达模式表明其表达具有卵巢发育正相关性。另结合雌鱼血清T和E2水平,表明P450c17-Ⅰ在类固醇激素途径发生转变中起到了重要作用。
6.半滑舌鳎P450c17-Ⅱ克隆及时空表达分析
本研究克隆了P450c17-Ⅱ全长cDNA,为2102bp,编码524个氨基酸。半滑舌鳎P450c17-Ⅱ酶,与其他物种P450c17-Ⅱ酶同源性较其他物种P450c17-Ⅰ酶同源性高。时空表达发现其只在性腺、头肾、脑内表达。卵巢和脑内P450c17-Ⅱ时间表达模式都具有发育正相关性,而P450c17-Ⅱ在精巢内的时间表达却与性腺发育呈负相关。本研究通过性腺、脑、头肾内P450c17-Ⅱ表达,结合性腺发育期、血清中E2、T,探讨P450c17-Ⅱ与性腺发育关系,为国内鲆鲽鱼类繁殖生理学提供理论基础。
The full length of P450c17-Ⅰand P450c17-ⅡcDNA is cloned from half-smooth tongue sole Cynoglossus semilaevis, and the spatial and temporal expression of P450c17(-Ⅰ,-Ⅱ) are well studied, as well as P450c19(A,B).The results of the spatial and temporal expression of the four enzymes are firstly linked to ovarian development, serum sexual steroid hormones testosterone(T) and estradiol-17β(E2) level, so as to work over the enzymes' molecular physiological mechanism in female and male half-smooth tongue sole.
1. Studies on ovarian development and spawn type of cultured female half-smooth tongue sole Cynoglossus semilaevis
Seasonal variations of ovarian development and the relationship with Gonadosomatic index (GSI)、hepatosomatic index (HSI) was studied in the cultured half-smooth tongue sole Cynoglossue semilaevis. The results show that ovarian development consists of phaseⅡ,Ⅲ,Ⅳ,Ⅴ,Ⅵ, then cycles to recrudescent phaseⅡ; under artificial culture, most fish has phaseⅡovarium from March to May, and the oocytes begin to accumulate vitellin granule from June, then the whole cytoplasm is stuffed with vitellin granule ranging from September to November, suggesting the oocytes growth to mature, after spawning, the ovarium come to assimilative phaseⅥor recrudescent phaseⅡ;GSI reaches to an peak during reproduction, then dramatically drop after spawning or recrudescent phaseⅡwhile HSI significantly increases to maximum; there is only one breeding season in the half-smooth tongue sole, and it belongs to batch-spawning fish.
2. Study on the seasonal changes of serum sexual steroid hormones in half-smooth tongue sole Cynoglossus semilaevis
Serum sexual steroid hormones testosterone(T) and estradiol-17β(E2) levels are measured by 125I radioimmunoassay(RIA). Serum E2 and T levels are shown to be accord with ovarian developmental phase (GSI) in female hale-smooth tongue sole. During the reproductive period, serum E2 and T levels significantly increase, and reach to the highest level throughout the whole year(P<0.05). After spawn, the females live through winter, and both of serum E2 and T levels decline. In the period of artificially enriching and culturing, serum E2 level remarkably increase(P<0.05), but the content of serum T drops markedly to the value that can't be detected. The results show that seasonal changes of female half-smooth tongue sole serum E2 level is closely correlated with ovarian development and vitellus' formation; and as the precursor of E2, the changes of T level has some relationship with ovarian development, furthermore it is essential for certain ovarian developmental phase.
Serum T and E2 levels are also shown to be accord with testis development(GSI) in male half-smooth tongue sole. The peak of serum T and E2 levels both occur at reproductive season; from November to January of next year, both of the contents drop rapidly(P<0.05); during February to April, serum T level slightly increase, but the content of serum E2 still decrease; in the period of May to July, serum T and E2 levels both significantly increase(P<0.05). The results show that T directly effects on the growth of sperm cell, and can boost sperm' formation and maturation.
3. Spatial and temporal expression analysis of half-smooth tongue sole Cynoglossus semilaevis P450c19s
The spatial expression of P450aromA and P450aromB show that P450aromA only expresses in gonad, and P450aromB only expresses in brain and gill. Further, P450aroms express in the gonad and brain throughout the whole gonad development in female and male C. semilaevis, respectively. The temporal expression pattern of P450aromA is consistent with the changes of serum E2 and T level in ovaries, indicating that it plays an important role in ovarian development and oocyte growth. However, the pattern of P450aromA in testis is not related with the changes of serum E2 and T level. The results of P450aromB'temporal expression shows that, in female tongue sole of atretic follicles stage, P450aromB expression in brain increases significantly and reaches to the peak; the similar result occurs in male tongue sole brain after spermiation. These results suggested the reproductive cycle of female tongue sole is not only regulated by CYP19A, but also regulated by CYP19B.
4. Programming Design of Degenerate Primers and Cloning Half-smooth Tongue Sole Cynoglossus semilaevis CYP17 partial cDNA
A new method of designing degenerate primers-CODEHOP is introduced to clone half-smooth tongue sole Cynoglossus semilaevis CYP17 gene. With Bioinformatics Resources such as NCBI、Blastp、BlockMaker、CODEHOP and biological software such as DNAMAN、Oligo6.0, the CODEHOP primer is programming designed, and finally we succeeded to clone the gene from the ovaries of half-smooth tongue sole Cynoglossus semilaevis. The fragment of the gene has high identities with the CYP17 family of other species, and it is clustered into the CYP17 gene group of all the other vertebrates, especially into the fish subgroup. The experiment results showed that compared with traditional degenerate primers, programming designed primers had more Specific ,lower positive rate, and more benefit for accomplishing the experiment.
5. Cloning, spatial and temporal expression analysis of half-smooth tongue sole Cynoglossus semilaevis P450c17-Ⅰ
A full length cDNA of P450c17-Ⅰhas been isolated from half-smooth tongue sole (Cynoglossus semilaevis) ovary. It is 2469 bp in length, encoding 509 amino acids. Also a relative shorter cDNA (1742bp) was found, likely due to a polyadenylation. The putative P450c17-Ⅰenzyme shares high sequence identity with that of fathead minnow, zebrafish, the Japanese eel, tilapia, three spined stickleback, medaka dogfish, chicken, rat, and human. Semi-quantitative RT-PCR analysis of spatial expression shows that it was predominantly expressed in the ovary, followed by brain, and faint signals were also detected in stomach, intestine, gill, spleen, kidney, head kidney. Further examination of temporal expression pattern of P450c17-Ⅰin ovary and brain revealed developmental stage-dependency. In addition to this our data on T and E2 levels further endorse the critical role of P450c17-Ⅰduring shift in steroidogenesis.
6. Cloning, spatial and temporal expression analysis of half-smooth tongue sole Cynoglossus semilaevis P450c17-Ⅱ
A full length cDNA of P450c17-Ⅱhas been isolated from half-smooth tongue sole (Cynoglossus semilaevis) ovary. It is 2102bp in length, encoding 524 amino acids. The putative tongue sole P450c17-Ⅱenzyme shares higher sequence identity with that of other animals, than the identity of P450c17-Ⅰenzyme of the same species. Semi-quantitative RT-PCR analysis of spatial expression shows that it was specially expressed in the ovary, brain and head kidney. Further examination of temporal expression pattern of P450c17-Ⅱin ovary and brain revealed developmental stage-dependency. However, the expression pattern of P450c17-Ⅱin testis shows negative relationship with testis development. P450c17-Ⅱexpression in gonad, brain and head kidney is studied, combined with serum E2 and T levels, so as to find out the relationship between P450cl7-Ⅱand gonad development, and provide theory foundation of flatfish reproductive physiology.
引文
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