不同倍性鲫鲤脑垂体细胞结构和HPG轴相关基因的表达及育性研究
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摘要
世界上首次培育成功的两性可育异源四倍体鲫鲤群体在生物进化和鱼类遗传育种方面都有重要意义。用雄性四倍体鲫鲤与雌性二倍体日本白鲫交配产生了不育三倍体湘云鲫。四倍体、三倍体鱼的形成为不同倍性鱼的生物学比较提供了重要的生物学平台。鱼类的性腺发育与配子成熟主要受到脑-垂体-性腺轴(HPG)为核心的神经和内分泌系统调控,脑分泌促性腺激素释放激素(GnRH),激发脑垂体分泌促性腺激素(GTH)与性腺表面特异的促性腺激素受体(GTHR)结合,促使性腺分泌性类固醇激素,从而使性腺发育成熟。而脑垂体作为重要的内分泌器官,在内分泌激素调控中具有非常重要的作用。本研究首次比较分析了二倍体红鲫、三倍体湘云鲫和四倍体鲫鲤脑垂体分泌细胞结构,并对其Gnrh、Gth和Gthr基因的结构及表达进行了比较研究。主要内容如下:
1.对不同倍性鲫鲤脑垂体组织化学结构及分泌细胞大小进行比较分析。就同一类分泌细胞而言,三种鱼脑垂体细胞的体积随倍性增加而增大。在繁殖季节,四倍体鱼中腺垂体的GTH细胞所占比例最大,三倍体鱼最少,这与四倍体性腺发育提前,三倍体不育有关;而三倍体鱼STH细胞所占比例最高,四倍体鱼最少,这与三倍体生长速度快,而四倍体生长速度相对缓慢有关。
2.对不同倍性鲫鲤GTH、STH细胞的超微结构进行观察,发现四倍体与二倍体鱼的分泌细胞能进行正常的分泌活动。而繁殖季节三倍体鱼GTH细胞中的分泌颗粒的数量远不及同时期的二倍体和四倍体鱼,并且在繁殖季节后也只能观察到少量空泡状结构。推测由于某种因素抑制了分泌颗粒的排出,从而导致三倍体鱼不育。
3.对不同倍性鲫鲤Gnrh2 cDNA全长进行克隆和序列分析。三种鱼Gnrh2氨基酸序列与其他鲤科鱼类有较高的同源性。除GnRH2十肽和蛋白水解位点在三种鱼中完全一致外,四倍体鱼与红鲫的GnRH2氨基酸差异要比三倍体鱼与红鲫的差异小,推测是由于红鲫与四倍体鱼具有更近的亲缘关系所致。
4.比较研究Gnrh2基因在不同倍性鲫鲤不同组织中的表达情况。Gnrh2基因在三种鱼的中脑、垂体和性腺中均有表达,其转录产物主要定位于中脑的后部,说明Gnrh2基因可能主要起神经递质或神经调节剂的作用,但也参与调节促性腺激素、生长激素等的释放。繁殖季节Gnrh2基因在四倍体鱼中脑的表达水平最高,三倍体鱼最低;而繁殖季节后的情况相反。推测可能与繁殖季节三倍体鱼性腺不能正常发育且繁殖季节后其Gnrh2表达量不能正常下调有关。
5.对不同倍性鲫鲤Fshβ和Lhβ基因cDNA全序列进行克隆和同源性分析。三种鱼的两种基因与其他鲤科鱼类同源性较高。此外,三种鱼Fshβ基因均含有13个半胱氨酸残基,推测其通过第一和第二个半胱氨酸残基间的裂解位点使之形成与其他硬骨鱼类似的12个半胱氨酸残基,从而形成6个二硫键。而三倍体和四倍体鱼Fshβ基因第2个糖基化位点的缺失可能影响其空间结构及生物活性。
6.比较研究Fshβ和Lhβ基因在不同倍性鲫鲤不同组织中的表达情况。Fshβ和Lhβ基因只在三种鱼的脑垂体中表达,且均定位于中腺垂体区域,说明这三种鱼的促性腺激素主要是由中腺垂体的GTH细胞产生。繁殖季节,三种鱼Lhβ基因的表达量及分布范围均比Fshβ大,说明Lhβ在促进配子及性腺最终成熟过程中起主要作用。此外,繁殖季节四倍体鱼Fshβ和Lhβ基因表达水平略高于二倍体,可能与其性成熟年龄提前有关。而繁殖季节后三倍体鱼两个基因的表达量要高于二倍体与四倍体鱼,可能是下游因子的反馈调控抑制其表达量下调。
7.对不同倍性鲫鲤Fshr和Lhr基因的表达进行比较研究。三种鱼Fshr和Lhr基因均只在精巢和卵巢中表达,并主要定位于卵巢的滤泡细胞、颗粒细胞、放射膜外层及精巢的支持细胞、间质细胞,说明它们都与促进性腺发育及配子成熟有关。但三倍体鱼性腺中两种基因的表达量相对较少,导致GTHR与GTH结合的能力下降,不能产生足够的类固醇激素促进性腺的正常发育,并最终造成三倍体不育。
总之,上述结果说明不同倍性鱼脑垂体结构及Gnrh、Gth和Gthr基因表达存在的差异与三倍体不育及四倍体可育有关联,这也为探讨四倍体可育性和三倍体不育性的内分泌学机制提供细胞生物学及分子生物学依据。
The formation of allotetraploid population derived from the red crucian carps(Carassius auratus red var.,♀,2n=100) and common carps (Cyprinus carpio L.,♂,2n=100) played an important role in biological evolution and fish breeding.The sterile triploid crucian carp was produced by mating the males of allotetraploids with the females of Japanese crucian carp(Carassius cuvieri).Both tetraploids and triploids provided a biological platform for the comparative studies of different ploidy level fishes.In fishes,reproductive activities are primarily controlled by an endocrine system consisting of the brain-pituitary-gonad axis.Gonadotropin-releasing hormone(GnRH) produced in the brain stimulates the production and release of gonadotropin(GTH) in the pituitary.Then GTH acts on the receptors of gonads to stimulate the production of sex steroids,which regulate gonadal development.The pituitary is an important endocrine organ,which plays an important role in regulating the secretion of hormones related to the growth and development.In this paper,we firstly observed the histological and ultra-structure of endocrine cells in different ploidy fishes,and comparatively analyzed the molecular and expression characterizations of Gnrh,Gth and Gthr genes in different ploidy fishes.The major results were presented as follows:
1.The histological structure and the endocrine cells' size of pituitary in different ploidy fishes were comparatively studied.For each type of endocrine cells,the cell size was increased gradually with the increasing ploidy level.In breeding season,the proportion of gonadotropin cells (GTH) in tetraploids was the highest,while that in triploids was the lowest.It might be related to the earlier sexual maturity of tetraploids and sterility of triploids.However,the proportion of somatotropin(STH) cells in triploids was the highest,whereas that in tetraploids was the lowest.It might be connected with the faster growth rate of triploids and slower growth rate of tetraploids.
2.The ultra-structures of GTH and STH cells in different ploidy fishes were observed.These two types of endocrine cells in diploids and tetraploids had normal biological activities.But the number of endocrine particles in GTH cells of triploids was fewer than that of diploids and tetraploids within breeding season,and only a few cavities could be observed after breeding season.It presumed that some factors restrained the releasing of endocrine particles,which then led to the sterility.
3.The full lengths of Gnrh2 cDNAs in different ploidy fishes were obtained and their sequences were comparatively analyzed.It was indicated that Gnrh2 gene had high homology among the cyprinids.The results of amino acid sequences among different ploidy fishes also revealed that the GnRH2 decapeptide and proteolytic cleavage site were absolutely consistent.The differences in signal peptide and GAP between the diploids and tetraploids were fewer than those between diploids and triploids.The possible reason was that the red crucian carp was the original maternal parent of tetraploids,so they had closer relationship.
4.The expression patterns of Gnrh2 in different ploidy fishes were comparatively studied.In these three kinds of fishes,Gnrh2 was not only expressed in midbrains,but also expressed in pituitaries and gonads.And Gnrh2 mRNA was mainly located in posterior end of midbrain.It was indicated that Gnrh2 might mainly act as the neurotransmitter and neuromodulator,and it also might participate in the regulating of other hormones releasing,such as GTH and STH.In addition,the expression level of Gnrh2 was the highest in tetraploids and the lowest in triploids within the breeding season,while that of triploids was the highest and that of tetraploids was the lowest after breeding season.It might be related to the abnormal gonadal development of triploids within breeding season and the Gnrh2 expression might be less down-regulated after breeding season.
5.The full length of Fshβand LhβcDNAs in different ploidy fishes were obtained and the sequence homology was then analyzed.Both of the two genes had high homology among the cyprinids.In these three fishes, Fshβhad 13 cysteine residues,12 of which formed 6 conserved disulfide bridges by utilizing the cleavage sites between first and second cysteine residue.In addition,the lack of second glycosylation sites in Fshβof these three fishes might influence the spacial structure and biological activities.
6.The expression patterns of Fshβand Lhβin different ploidy fishes were comparatively studied.Both of these two genes only expressed in the pituitaries of these three fishes,and they mainly located in the meso-adenohypophysis, which indicated that the gonadotropins were secreted by GTH cells in meso-adenohypophysis.Within breeding season,the Lhβgene in three fishes had higher expression level and wider distribution range than that of Fshβ,which suggested that Lhβwas important for inducing final gonadal maturity.In addition,Fshβand Lhβexpression levels in tetraploids were a little higher than that of diploids for its earlier sexual maturation.While Fshβand Lhβin triploids had higher expression levels than in other two fishes after breeding season for restraining by the feedbake regulation of downstream factors.
7.The expression patterns of Fshr and Lhr in different ploidy fishes were comparatively studied.Both of these two genes only expressed in the ovaries and testis of three fishes,and they mainly located in the follicle cells,granulose cells,outside layer of zona radiate,Sertoli cells and Leydig cells.It was indicated that Fshr and Lhr gene might stimulate the development and maturation of gonads.However,the hybridization signals of Fshr and Lhr gene in triploids were less than other two fishes. It was suggested that the triploids had fewer GTHR on target cell surfaces which couldn't interacting with the GTH sufficiently,so the gonad could not produce enough sex steroids and finally led to sterility.
In a word,all the results indicated that the differences of the structure of pituitary endocrine cells and the expression of Gnrh,Gth and Gthr genes in different ploidy fishes were related to the sterility of triploids and fertility of tetraploids.It also provided cytological and molecular evidences for the endocrine mechanism of the fertile tetraploids and sterile triploids.
1.对不同倍性鲫鲤脑垂体组织化学结构及分泌细胞大小进行比较分析。就同一类分泌细胞而言,三种鱼脑垂体细胞的体积随倍性增加而增大。在繁殖季节,四倍体鱼中腺垂体的GTH细胞所占比例最大,三倍体鱼最少,这与四倍体性腺发育提前,三倍体不育有关;而三倍体鱼STH细胞所占比例最高,四倍体鱼最少,这与三倍体生长速度快,而四倍体生长速度相对缓慢有关。
2.对不同倍性鲫鲤GTH、STH细胞的超微结构进行观察,发现四倍体与二倍体鱼的分泌细胞能进行正常的分泌活动。而繁殖季节三倍体鱼GTH细胞中的分泌颗粒的数量远不及同时期的二倍体和四倍体鱼,并且在繁殖季节后也只能观察到少量空泡状结构。推测由于某种因素抑制了分泌颗粒的排出,从而导致三倍体鱼不育。
3.对不同倍性鲫鲤Gnrh2 cDNA全长进行克隆和序列分析。三种鱼Gnrh2氨基酸序列与其他鲤科鱼类有较高的同源性。除GnRH2十肽和蛋白水解位点在三种鱼中完全一致外,四倍体鱼与红鲫的GnRH2氨基酸差异要比三倍体鱼与红鲫的差异小,推测是由于红鲫与四倍体鱼具有更近的亲缘关系所致。
4.比较研究Gnrh2基因在不同倍性鲫鲤不同组织中的表达情况。Gnrh2基因在三种鱼的中脑、垂体和性腺中均有表达,其转录产物主要定位于中脑的后部,说明Gnrh2基因可能主要起神经递质或神经调节剂的作用,但也参与调节促性腺激素、生长激素等的释放。繁殖季节Gnrh2基因在四倍体鱼中脑的表达水平最高,三倍体鱼最低;而繁殖季节后的情况相反。推测可能与繁殖季节三倍体鱼性腺不能正常发育且繁殖季节后其Gnrh2表达量不能正常下调有关。
5.对不同倍性鲫鲤Fshβ和Lhβ基因cDNA全序列进行克隆和同源性分析。三种鱼的两种基因与其他鲤科鱼类同源性较高。此外,三种鱼Fshβ基因均含有13个半胱氨酸残基,推测其通过第一和第二个半胱氨酸残基间的裂解位点使之形成与其他硬骨鱼类似的12个半胱氨酸残基,从而形成6个二硫键。而三倍体和四倍体鱼Fshβ基因第2个糖基化位点的缺失可能影响其空间结构及生物活性。
6.比较研究Fshβ和Lhβ基因在不同倍性鲫鲤不同组织中的表达情况。Fshβ和Lhβ基因只在三种鱼的脑垂体中表达,且均定位于中腺垂体区域,说明这三种鱼的促性腺激素主要是由中腺垂体的GTH细胞产生。繁殖季节,三种鱼Lhβ基因的表达量及分布范围均比Fshβ大,说明Lhβ在促进配子及性腺最终成熟过程中起主要作用。此外,繁殖季节四倍体鱼Fshβ和Lhβ基因表达水平略高于二倍体,可能与其性成熟年龄提前有关。而繁殖季节后三倍体鱼两个基因的表达量要高于二倍体与四倍体鱼,可能是下游因子的反馈调控抑制其表达量下调。
7.对不同倍性鲫鲤Fshr和Lhr基因的表达进行比较研究。三种鱼Fshr和Lhr基因均只在精巢和卵巢中表达,并主要定位于卵巢的滤泡细胞、颗粒细胞、放射膜外层及精巢的支持细胞、间质细胞,说明它们都与促进性腺发育及配子成熟有关。但三倍体鱼性腺中两种基因的表达量相对较少,导致GTHR与GTH结合的能力下降,不能产生足够的类固醇激素促进性腺的正常发育,并最终造成三倍体不育。
总之,上述结果说明不同倍性鱼脑垂体结构及Gnrh、Gth和Gthr基因表达存在的差异与三倍体不育及四倍体可育有关联,这也为探讨四倍体可育性和三倍体不育性的内分泌学机制提供细胞生物学及分子生物学依据。
The formation of allotetraploid population derived from the red crucian carps(Carassius auratus red var.,♀,2n=100) and common carps (Cyprinus carpio L.,♂,2n=100) played an important role in biological evolution and fish breeding.The sterile triploid crucian carp was produced by mating the males of allotetraploids with the females of Japanese crucian carp(Carassius cuvieri).Both tetraploids and triploids provided a biological platform for the comparative studies of different ploidy level fishes.In fishes,reproductive activities are primarily controlled by an endocrine system consisting of the brain-pituitary-gonad axis.Gonadotropin-releasing hormone(GnRH) produced in the brain stimulates the production and release of gonadotropin(GTH) in the pituitary.Then GTH acts on the receptors of gonads to stimulate the production of sex steroids,which regulate gonadal development.The pituitary is an important endocrine organ,which plays an important role in regulating the secretion of hormones related to the growth and development.In this paper,we firstly observed the histological and ultra-structure of endocrine cells in different ploidy fishes,and comparatively analyzed the molecular and expression characterizations of Gnrh,Gth and Gthr genes in different ploidy fishes.The major results were presented as follows:
1.The histological structure and the endocrine cells' size of pituitary in different ploidy fishes were comparatively studied.For each type of endocrine cells,the cell size was increased gradually with the increasing ploidy level.In breeding season,the proportion of gonadotropin cells (GTH) in tetraploids was the highest,while that in triploids was the lowest.It might be related to the earlier sexual maturity of tetraploids and sterility of triploids.However,the proportion of somatotropin(STH) cells in triploids was the highest,whereas that in tetraploids was the lowest.It might be connected with the faster growth rate of triploids and slower growth rate of tetraploids.
2.The ultra-structures of GTH and STH cells in different ploidy fishes were observed.These two types of endocrine cells in diploids and tetraploids had normal biological activities.But the number of endocrine particles in GTH cells of triploids was fewer than that of diploids and tetraploids within breeding season,and only a few cavities could be observed after breeding season.It presumed that some factors restrained the releasing of endocrine particles,which then led to the sterility.
3.The full lengths of Gnrh2 cDNAs in different ploidy fishes were obtained and their sequences were comparatively analyzed.It was indicated that Gnrh2 gene had high homology among the cyprinids.The results of amino acid sequences among different ploidy fishes also revealed that the GnRH2 decapeptide and proteolytic cleavage site were absolutely consistent.The differences in signal peptide and GAP between the diploids and tetraploids were fewer than those between diploids and triploids.The possible reason was that the red crucian carp was the original maternal parent of tetraploids,so they had closer relationship.
4.The expression patterns of Gnrh2 in different ploidy fishes were comparatively studied.In these three kinds of fishes,Gnrh2 was not only expressed in midbrains,but also expressed in pituitaries and gonads.And Gnrh2 mRNA was mainly located in posterior end of midbrain.It was indicated that Gnrh2 might mainly act as the neurotransmitter and neuromodulator,and it also might participate in the regulating of other hormones releasing,such as GTH and STH.In addition,the expression level of Gnrh2 was the highest in tetraploids and the lowest in triploids within the breeding season,while that of triploids was the highest and that of tetraploids was the lowest after breeding season.It might be related to the abnormal gonadal development of triploids within breeding season and the Gnrh2 expression might be less down-regulated after breeding season.
5.The full length of Fshβand LhβcDNAs in different ploidy fishes were obtained and the sequence homology was then analyzed.Both of the two genes had high homology among the cyprinids.In these three fishes, Fshβhad 13 cysteine residues,12 of which formed 6 conserved disulfide bridges by utilizing the cleavage sites between first and second cysteine residue.In addition,the lack of second glycosylation sites in Fshβof these three fishes might influence the spacial structure and biological activities.
6.The expression patterns of Fshβand Lhβin different ploidy fishes were comparatively studied.Both of these two genes only expressed in the pituitaries of these three fishes,and they mainly located in the meso-adenohypophysis, which indicated that the gonadotropins were secreted by GTH cells in meso-adenohypophysis.Within breeding season,the Lhβgene in three fishes had higher expression level and wider distribution range than that of Fshβ,which suggested that Lhβwas important for inducing final gonadal maturity.In addition,Fshβand Lhβexpression levels in tetraploids were a little higher than that of diploids for its earlier sexual maturation.While Fshβand Lhβin triploids had higher expression levels than in other two fishes after breeding season for restraining by the feedbake regulation of downstream factors.
7.The expression patterns of Fshr and Lhr in different ploidy fishes were comparatively studied.Both of these two genes only expressed in the ovaries and testis of three fishes,and they mainly located in the follicle cells,granulose cells,outside layer of zona radiate,Sertoli cells and Leydig cells.It was indicated that Fshr and Lhr gene might stimulate the development and maturation of gonads.However,the hybridization signals of Fshr and Lhr gene in triploids were less than other two fishes. It was suggested that the triploids had fewer GTHR on target cell surfaces which couldn't interacting with the GTH sufficiently,so the gonad could not produce enough sex steroids and finally led to sterility.
In a word,all the results indicated that the differences of the structure of pituitary endocrine cells and the expression of Gnrh,Gth and Gthr genes in different ploidy fishes were related to the sterility of triploids and fertility of tetraploids.It also provided cytological and molecular evidences for the endocrine mechanism of the fertile tetraploids and sterile triploids.
引文
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