牙鲆繁殖内分泌分子机理研究
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摘要
几乎所有的养殖鱼类都存在繁殖机能障碍。雌性鱼类通常表现为卵母细胞不能最终成熟、排卵和产卵；雄鱼表现为精液产量减少或者品质较差。本论文主要以人工养殖的牙鲆(Paralichthys olivaceus)和野生的漠斑牙鲆(Paralichthys lethostigma)为研究对象,运用组织学、免疫组织化学、激素放射免疫测定方法及分子生物学方法,对牙鲆性腺、血浆性类固醇激素及6个与繁殖相关的基因进行研究,以期提供一些对于理解人工养殖的比目鱼繁殖机制和改善工厂化养殖鱼类的繁殖调控有价值的参考信息,主要研究内容和结果如下：
1.P450芳香化酶是细胞色素P450超家族的一员,由CYP19基因编码；催化C19雄激素为芳香化的C18雌激素。为了解性腺型P450芳香化酶(CYP19α)在牙鲆繁殖过程中生理功能,采用RT-PCR方法检测CYP19αmRNA在不同组织表达和精巢不同发育阶段的表达。CYP19αmRNA大量表达在雌鱼卵巢和脾脏,在雄鱼精巢和脾脏CYP19αmRNA表达相对较弱,在其它组织没有检测到该基因表达。在雄鱼繁殖周期,CYP19αmRNA表达水平变化与血浆中雌二醇的变化趋势不一致。免疫组化研究表明：随着精巢的发育,精原细胞、精母细胞对芳香化酶抗体的免疫阳性反应逐渐变弱,精子细胞显免疫阴性反应。总体看来,芳香化酶在精巢发育的早期有较强的表达。这些结果说明CYP19α参与调节精子发生。
2.细胞色素P450c17是单一的酶,在性腺和肾上腺类固醇激素合成过程中具有调解17α-羟化酶和17,20-裂解酶活性的作用。在硬骨鱼类有二种类型的P450c17,P450c17-Ⅰ具有二种酶活性,P450c17-Ⅱ只有17α-羟化酶活性。在牙鲆克隆得到P450c17-Ⅰ和P450c17-Ⅱ的cDNA全长。RT-PCR分析表明P450c17-ⅠmRNA在心脏、精巢和卵巢表达丰富,P450c17-ⅡmRNA在脑、头肾、肾、精巢和卵巢表达丰富。在精巢不同发育阶段均检测到P450c17-Ⅰ和P450c17-ⅡmRNA表达,并且这二种基因的表达水平变化与血浆中睾酮水平和性腺指数变化趋势基本相似。这些研究结果的生理学意义需要进一步实验阐明。
3.雌激素通过结合核受体蛋白,雌激素受体,在繁殖组织生长、分化和维持繁殖能力方面发挥重要作用。本研究通过PCR-SSCP方法检测采集的雌性牙鲆雌激素受体β基因编码区多态性,并对雌激素受体β基因的多态性与繁殖性能指标进行关联分析。在雌激素受体β基因发现二个单核苷酸多态性位点：SNP1 (c.577delC)和SNP2 [c.A891T (p.Glnl 14Leu)]。方差分析表明SNP1与雌性牙鲆性腺指数相关(P<0.05),SNP2与雌性牙鲆的雌二醇水平和性腺指数相关(P<0.05)。在SNP2位点,AB基因型个体比AA基因型个体具有较高的雌激素水平和性腺指数(P<0.05)。本研究构建4个二倍型；通过多重比较发现二倍型D2对肝重指数效应显著高于其它二倍型(P<0.05)。
4.FOXL2基因具有参与卵巢发育,滤泡细胞分化和维持卵巢生理功能的作用。本研究采用PCR-SSCP技术对采集的雌性牙鲆FOXL2基因进行多态性检测,并对检测到的遗传多态性与其繁殖性能指标进行关联分析。结果表明：引物3,引物4,和引物5的PCR产物具有多态性；共检测到5个突变：SNP1 [c.540A>C (p.Asn102His)和c.591A>G (p.Asn119Asp)], SNP2 [c.864G>A (p.Lys210Glu)和c.875G>A]和SNP3(c.1169C>A)。采用最小二乘法对牙鲆的3个FOXL2基因突变位点和繁殖性能指标进行关联分析：在Forkhead区域,SNP1对牙鲆性腺指数效应均达到显著水平(P<0.05)。在Forkhead区域下游,SNP2对牙鲆雌二醇效应达到显著水平(P<0.05)。在3’-非编码区,SNP3对牙鲆肝重指数效应达到显著水平(P<0.05)。基于检测到的5个突变,构建得到5种二倍型组合。通过多重比较,二倍型D3对睾酮水平效应和二倍型D5对性腺指数效应显著高于其它二倍型(P<0.05)。
5.近期在几种硬骨鱼类的研究表明膜孕酮受体alpha (mPRα)通过调节孕激素非基因组功能诱导卵母细胞成熟和精子运动。在当前研究通过分析mPRα在雄性牙鲆精巢不同发育阶段mRNA表达进一步评价mPRα在配子和性腺的生理学作用。首先从牙鲆克隆得到mPRα的cDNA全长；预测的氨基酸序列与其它鱼类：nPRα的氨基酸序列高度同源如漠斑牙鲆(99%)、斑点犬牙石首鱼(94%)和大西洋绒须石首鱼(93%)。跨膜区域预测结果表明牙鲆mPRα蛋白具有七次跨膜结构。牙鲆mPRαmRNA广泛表达在各检测的组织,在脑、卵巢、精巢和肾脏表达丰富。牙鲆mPRαmRNA在精巢不同发育阶段的表达变化趋势与血浆雌二醇和性腺指数变化趋势大体相似。所获得数据支持我们的假设：mPRα是牙鲆成熟诱导类固醇受体。
6.近来我们在牙鲆发现一个具有孕激素脂联素受体特征的新基因PAQR-like基因,本研究报道新基因克隆和此基因部分特征。系统发生分析表明PAQR-like基因位于PAQR7和PAQR8分支之间。预测的氨基酸序列与三刺鱼未命名基因的氨基酸序列相似性为77%,与青鳉未命名基因的氨基酸序列相似性为72%。PAQR-like基因跨膜区域预测结果表明该基因具有7个跨膜区域,具有孕激素脂联素受体特征。RT-PCR分析表明PAQR-like基因mRNA在卵巢、头肾、精巢和肾脏表达丰富,在脑、鳃、肠、胃和幽门盲囊表达相对较弱。PAQR-like基因mRNA表达水平变化与性腺指数变化趋势基本相似。这些特征表明PAQR-like基因参与牙鲆繁殖生理活动。
7.漠斑牙鲆是与牙鲆很相似的海洋硬骨鱼类,其成熟诱导类固醇激素至今未见报道。当前研究目的是确定漠斑牙鲆卵母细胞成熟过程中产生的主要孕酮激素并调查在此过程中是否产生的主要孕酮激素分泌增加。完整的卵巢滤泡细胞经氚标记的孕烯醇酮孵育,孵育后的产物经薄层色谱方法确定。我们发现氚标记的孕烯醇酮通过生物化学方法主要转变为20p-S、17α,20β-DHP和11-脱氧皮质醇。卵母细胞成熟过程中20β-S大量生成,而生成的11-脱氧皮质醇逐渐减少,17α,20β-DHP生成量基本不变。同时体外实验表明20β-S和17α,20β-DHP是诱导漠斑牙鲆卵母细胞成熟最有效的二种激素。这些结果表明20β-S和17α,20β-DHP可能是漠斑牙鲆的成熟诱导类固醇激素。
Almost all fish reared in captivity exhibit some form of reproductive dysfunction. In females, there is often failure to undergo final oocyte maturation, ovulation and spawning; while in males milt production may be reduced and of low quality. This thesis mainly studied on cultured Japanese flounder (Paralichthys olivaceus) and wild southern flounder(Paralichthys lethostigma), from which the gonad, sexual steroid hormone of blood plasma, several gene related to fish reproduction were taken and studied by the methods of histological staining, immunohistochemical techniques, radioimmunoassay and molecular methods. The aim is to provide some valuable information for understanding the reproductive mechanisms in flatfish cultured and for improving strategies of reproductive manipulations in the aquaculture industry. Main contents and results are as follows: 1. P450 aromatase (P450arom, CYP19), a CYP19 gene product, is a member of the cytochrome P450 superfamily that catalyzes the formation of aromatic C18 estrogen from C19 androgen. To characterize the involvement of the gonadal P450 aromatase gene (CYP19α) during the reproduction of Japanese flounder, the expression of CYP19αmRNA was detected in different tissue of adult fish and also determined during the various developmental stages of testes. The CYP19αmRNA was highly abundant in ovary and spleen of female fish, less abundant in testis and spleen of male fish, but not present in other tissues by RT-PCR. The changes in mRNA level of CYP19αdid not share the same changes trend with serum E2 during the reproductive cycle of male fish Immunohistochemically, aromatase immunopositive reaction of spermatogonia and spermatocytes weaken gradually and spermids were immunonegative to aromatase during testis development. Overall, immunoreactive P450arom has stronger expression at early stage of testis development. These results suggest that CYP19a gene implicated in the regulation of spermatogenesis.
2. Cytochrome P450c17 is the single enzyme that mediates the 17a-hydroxylase and 17,20 lyase activities during the biosynthesis of steroid hormones in the gonads and adrenal gland. In teleost, there are two types of P450c17, P450c17-Ⅰhas both these enzymatic activity and P450cl7-II possesses only 17a-hydroxylase activities. In this study, the full length cDNA of P450c17-Ⅰand P450c17-Ⅱwere cloned from the Japanese flounder, which is an annually spawning teleost. RT-PCR analysis of Japanese flounder tissue extracts showed significant expression of P450c17-ⅠmRNA in the heart, testis and ovary. And the expression of P450c17-ⅡmRNA was relatively high in brain, head kidney, kidney, testis and ovary. We further examined the pattern of expression of two genes throughout the reproductive cycle. RT-PCR revealed that P450c17-Ⅰand P450c17-ⅡmRNA were expressed in testes of different stage. And the variation trend of serumal T level and GSI were similar with the pattern of these two genes mRNA expression. The physiological relevance of these findings remains to be further elucidated.
3. Estrogens play important roles in the growth and differentiation of reproductive tissues and in the maintenance of fertility by binding to a nuclear receptor protein, the estrogen receptors. The objectives of this study were to characterize polymorphisms within the coding region of estrogen receptorβ(ERβ) gene in the collected female Japanese flounder and to analyze the association of ERβpolymorphisms with reproductive indices by polymerase chain reaction single-strand conformation polymorphism (PCR-SSCP). Two single nucleotide polymorphisms (SNPs), SNP1 (c.577de1C) and SNP2 [c.A891T (p.Gln1 14Leu)], were identified in the ERβgene. A one-way ANOVA revealed that SNP1 was significantly associated with the gonadosomatic index (GSI) in female Japanese flounder (P<0.05). And SNP2 was significantly associated with the serum 17β-estradiol (E2) level and GSI (P<0.05). Individuals with genotype AB of SNP2 had significantly higher serum E2 level and GSI than those of genotype AA (P< 0.05).Moreover, the hepatosomatic index (HSI), a marker for genetic effects, was significantly higher for diplotype D2 compared with the other three diplotypes (P<0.05).
4. FOXL2 gene plays an important role in ovarian development, granulosa cell differentiation, and thus the proper maintenance of ovarian function. The aims of this study were to characterize polymorphisms within the FOXL2 gene in the collected female Japanese flounder and analyze the association of FOXL2 polymorphisms with reproductive performance by polymerase chain reaction-single stranded conformational polymorphism (PCR-SSCP). Results indicated that five Single nucleotide polymorphisms (SNPs), which were SNP1 [c.540A>C (p.Asn102His) and c.591A>G (p.Asn119Asp)], SNP2 [c.864G>A (p.Lys210Glu) and c.875G>A] and SNP3 (c.1169C>A), were identified in the FOXL2 gene. General Linear Model (GLM) analysis showed that SNP1 in the forkhead domain was significantly associated with gonadosomatic index (GSI) (P<0.05). SNP2 in the downstream of forkhead domain was significantly associated with serum 17β-estradiol (E2) level (P<0.05). And SNP3 in the 3'-UTR was significantly associated with hepatosomatic index (HSI) (P<0.05). Moreover, the evaluation of the genetic effects for both Testosterone (T) level of diplotype D3 and GSI of diplotype D5 suggested they were significantly higher than those of other four diplotypes (P<0.05), respectively.
5. Recent results suggested that membrane progestin receptor alpha (mPRα) mediates nongenomic actions of progestin hormones to induce oocyte maturation and sperm hypermotility in several teleost species. The role of mPRa in gamete and gonadal physiology was further evaluated in the present study by examining gonadal expression of mPRa during the gonadal development in male Japanese flounder.We cloned the mPRa cDNA from the Japanese flounder ovary. The predicted protein showed high sequence identity with Southern flounder (99%),Seatrout(94%) and Atlantic croaker(93%) mPRα. Computer modeling predicted that the deduced protein has seven transmembrance domains.The mPRα mRNA was detected in all the tissues tested with relatively greater expression in brain, ovary, testis and kidney. The mPRa mRNA was also expressed in Japanese flounder testes throughout the gonadal cycle. And the serumal E2 and GSI level showed the similar variation trend with the pattern of mPRa mRNA expression. Taken together, these data supported the hypothesis that mPRa is the MIS receptor in Japanese flounder.
6. Recently we discovered a previously uncharacterized gene with the characteristics of a progesterone and adipoQ receptor (PAQR) in a flatfish, Japanese flounder. Here, we reported the cloning and partial characteristics of this hitherto unknown gene. Phylogenetic analysis clearly indicated the PAQR-like gene clade was between the PAQR7 clade and PAQR8 clade. The predicted protein showed high sequence identity with three-spined stickleback (77%) and medaka (72%) unnamed gene. Structural analyses of the translated cDNA suggested it encoded protein with seven transmembrance domains, typical of progesterone and adipoQ receptor. RT-PCR analysis of Japanese flounder tissue extracts showed significant expression of the PAQR-like gene in ovary, head kidney, testis and kidney, and lower expression in brain, gill, intestine, stomach and pyloric caecum. The PAQR-like gene mRNA was also expressed in Japanese flounder testes throughout the gonadal cycle. And the variation trend of GSI was similar with the pattern of the PAQR-like gene mRNA expression. These characteristics suggested that the PAQR-like gene is related to reproductive physiology in this species.
7. The identity of the maturation-inducing steroid (MIS) in Southern flounder, a marine teleost which is similar to Japanese flounder, is unknown. The purpose of the present study was to identify the major progestin produced during oocyte maturation (OM) in southern flounder and investigate whether its secretion increases during this process. Intact of ovarian follicles undergoing OM were incubated in vitro with tritiated [3H]pregnenolone precursor and the tritiated products were extracted, purified, and identified by TLC. We found the tritiated products were biosynthesized from [3H]pregnenolone including 20β-S, 17a,20p-DHP,11-deoxycortisol. During the oocyte maturation, there were a significant increase in the production of 20β-S, whereas the production of 11-deoxycortisol decreased. And the amount of 17α,20β-DHP remained almost the same. In vitro,20β-S and 17α,20β-DHP were most effective steroids to induce maturation of southern flounder oocytes. These results suggest that that 20β-S and 17α,20P-DHP maybe the naturally occurring MIS in the female southern flounder.
1.P450芳香化酶是细胞色素P450超家族的一员,由CYP19基因编码；催化C19雄激素为芳香化的C18雌激素。为了解性腺型P450芳香化酶(CYP19α)在牙鲆繁殖过程中生理功能,采用RT-PCR方法检测CYP19αmRNA在不同组织表达和精巢不同发育阶段的表达。CYP19αmRNA大量表达在雌鱼卵巢和脾脏,在雄鱼精巢和脾脏CYP19αmRNA表达相对较弱,在其它组织没有检测到该基因表达。在雄鱼繁殖周期,CYP19αmRNA表达水平变化与血浆中雌二醇的变化趋势不一致。免疫组化研究表明：随着精巢的发育,精原细胞、精母细胞对芳香化酶抗体的免疫阳性反应逐渐变弱,精子细胞显免疫阴性反应。总体看来,芳香化酶在精巢发育的早期有较强的表达。这些结果说明CYP19α参与调节精子发生。
2.细胞色素P450c17是单一的酶,在性腺和肾上腺类固醇激素合成过程中具有调解17α-羟化酶和17,20-裂解酶活性的作用。在硬骨鱼类有二种类型的P450c17,P450c17-Ⅰ具有二种酶活性,P450c17-Ⅱ只有17α-羟化酶活性。在牙鲆克隆得到P450c17-Ⅰ和P450c17-Ⅱ的cDNA全长。RT-PCR分析表明P450c17-ⅠmRNA在心脏、精巢和卵巢表达丰富,P450c17-ⅡmRNA在脑、头肾、肾、精巢和卵巢表达丰富。在精巢不同发育阶段均检测到P450c17-Ⅰ和P450c17-ⅡmRNA表达,并且这二种基因的表达水平变化与血浆中睾酮水平和性腺指数变化趋势基本相似。这些研究结果的生理学意义需要进一步实验阐明。
3.雌激素通过结合核受体蛋白,雌激素受体,在繁殖组织生长、分化和维持繁殖能力方面发挥重要作用。本研究通过PCR-SSCP方法检测采集的雌性牙鲆雌激素受体β基因编码区多态性,并对雌激素受体β基因的多态性与繁殖性能指标进行关联分析。在雌激素受体β基因发现二个单核苷酸多态性位点：SNP1 (c.577delC)和SNP2 [c.A891T (p.Glnl 14Leu)]。方差分析表明SNP1与雌性牙鲆性腺指数相关(P<0.05),SNP2与雌性牙鲆的雌二醇水平和性腺指数相关(P<0.05)。在SNP2位点,AB基因型个体比AA基因型个体具有较高的雌激素水平和性腺指数(P<0.05)。本研究构建4个二倍型；通过多重比较发现二倍型D2对肝重指数效应显著高于其它二倍型(P<0.05)。
4.FOXL2基因具有参与卵巢发育,滤泡细胞分化和维持卵巢生理功能的作用。本研究采用PCR-SSCP技术对采集的雌性牙鲆FOXL2基因进行多态性检测,并对检测到的遗传多态性与其繁殖性能指标进行关联分析。结果表明：引物3,引物4,和引物5的PCR产物具有多态性；共检测到5个突变：SNP1 [c.540A>C (p.Asn102His)和c.591A>G (p.Asn119Asp)], SNP2 [c.864G>A (p.Lys210Glu)和c.875G>A]和SNP3(c.1169C>A)。采用最小二乘法对牙鲆的3个FOXL2基因突变位点和繁殖性能指标进行关联分析：在Forkhead区域,SNP1对牙鲆性腺指数效应均达到显著水平(P<0.05)。在Forkhead区域下游,SNP2对牙鲆雌二醇效应达到显著水平(P<0.05)。在3’-非编码区,SNP3对牙鲆肝重指数效应达到显著水平(P<0.05)。基于检测到的5个突变,构建得到5种二倍型组合。通过多重比较,二倍型D3对睾酮水平效应和二倍型D5对性腺指数效应显著高于其它二倍型(P<0.05)。
5.近期在几种硬骨鱼类的研究表明膜孕酮受体alpha (mPRα)通过调节孕激素非基因组功能诱导卵母细胞成熟和精子运动。在当前研究通过分析mPRα在雄性牙鲆精巢不同发育阶段mRNA表达进一步评价mPRα在配子和性腺的生理学作用。首先从牙鲆克隆得到mPRα的cDNA全长；预测的氨基酸序列与其它鱼类：nPRα的氨基酸序列高度同源如漠斑牙鲆(99%)、斑点犬牙石首鱼(94%)和大西洋绒须石首鱼(93%)。跨膜区域预测结果表明牙鲆mPRα蛋白具有七次跨膜结构。牙鲆mPRαmRNA广泛表达在各检测的组织,在脑、卵巢、精巢和肾脏表达丰富。牙鲆mPRαmRNA在精巢不同发育阶段的表达变化趋势与血浆雌二醇和性腺指数变化趋势大体相似。所获得数据支持我们的假设：mPRα是牙鲆成熟诱导类固醇受体。
6.近来我们在牙鲆发现一个具有孕激素脂联素受体特征的新基因PAQR-like基因,本研究报道新基因克隆和此基因部分特征。系统发生分析表明PAQR-like基因位于PAQR7和PAQR8分支之间。预测的氨基酸序列与三刺鱼未命名基因的氨基酸序列相似性为77%,与青鳉未命名基因的氨基酸序列相似性为72%。PAQR-like基因跨膜区域预测结果表明该基因具有7个跨膜区域,具有孕激素脂联素受体特征。RT-PCR分析表明PAQR-like基因mRNA在卵巢、头肾、精巢和肾脏表达丰富,在脑、鳃、肠、胃和幽门盲囊表达相对较弱。PAQR-like基因mRNA表达水平变化与性腺指数变化趋势基本相似。这些特征表明PAQR-like基因参与牙鲆繁殖生理活动。
7.漠斑牙鲆是与牙鲆很相似的海洋硬骨鱼类,其成熟诱导类固醇激素至今未见报道。当前研究目的是确定漠斑牙鲆卵母细胞成熟过程中产生的主要孕酮激素并调查在此过程中是否产生的主要孕酮激素分泌增加。完整的卵巢滤泡细胞经氚标记的孕烯醇酮孵育,孵育后的产物经薄层色谱方法确定。我们发现氚标记的孕烯醇酮通过生物化学方法主要转变为20p-S、17α,20β-DHP和11-脱氧皮质醇。卵母细胞成熟过程中20β-S大量生成,而生成的11-脱氧皮质醇逐渐减少,17α,20β-DHP生成量基本不变。同时体外实验表明20β-S和17α,20β-DHP是诱导漠斑牙鲆卵母细胞成熟最有效的二种激素。这些结果表明20β-S和17α,20β-DHP可能是漠斑牙鲆的成熟诱导类固醇激素。
Almost all fish reared in captivity exhibit some form of reproductive dysfunction. In females, there is often failure to undergo final oocyte maturation, ovulation and spawning; while in males milt production may be reduced and of low quality. This thesis mainly studied on cultured Japanese flounder (Paralichthys olivaceus) and wild southern flounder(Paralichthys lethostigma), from which the gonad, sexual steroid hormone of blood plasma, several gene related to fish reproduction were taken and studied by the methods of histological staining, immunohistochemical techniques, radioimmunoassay and molecular methods. The aim is to provide some valuable information for understanding the reproductive mechanisms in flatfish cultured and for improving strategies of reproductive manipulations in the aquaculture industry. Main contents and results are as follows: 1. P450 aromatase (P450arom, CYP19), a CYP19 gene product, is a member of the cytochrome P450 superfamily that catalyzes the formation of aromatic C18 estrogen from C19 androgen. To characterize the involvement of the gonadal P450 aromatase gene (CYP19α) during the reproduction of Japanese flounder, the expression of CYP19αmRNA was detected in different tissue of adult fish and also determined during the various developmental stages of testes. The CYP19αmRNA was highly abundant in ovary and spleen of female fish, less abundant in testis and spleen of male fish, but not present in other tissues by RT-PCR. The changes in mRNA level of CYP19αdid not share the same changes trend with serum E2 during the reproductive cycle of male fish Immunohistochemically, aromatase immunopositive reaction of spermatogonia and spermatocytes weaken gradually and spermids were immunonegative to aromatase during testis development. Overall, immunoreactive P450arom has stronger expression at early stage of testis development. These results suggest that CYP19a gene implicated in the regulation of spermatogenesis.
2. Cytochrome P450c17 is the single enzyme that mediates the 17a-hydroxylase and 17,20 lyase activities during the biosynthesis of steroid hormones in the gonads and adrenal gland. In teleost, there are two types of P450c17, P450c17-Ⅰhas both these enzymatic activity and P450cl7-II possesses only 17a-hydroxylase activities. In this study, the full length cDNA of P450c17-Ⅰand P450c17-Ⅱwere cloned from the Japanese flounder, which is an annually spawning teleost. RT-PCR analysis of Japanese flounder tissue extracts showed significant expression of P450c17-ⅠmRNA in the heart, testis and ovary. And the expression of P450c17-ⅡmRNA was relatively high in brain, head kidney, kidney, testis and ovary. We further examined the pattern of expression of two genes throughout the reproductive cycle. RT-PCR revealed that P450c17-Ⅰand P450c17-ⅡmRNA were expressed in testes of different stage. And the variation trend of serumal T level and GSI were similar with the pattern of these two genes mRNA expression. The physiological relevance of these findings remains to be further elucidated.
3. Estrogens play important roles in the growth and differentiation of reproductive tissues and in the maintenance of fertility by binding to a nuclear receptor protein, the estrogen receptors. The objectives of this study were to characterize polymorphisms within the coding region of estrogen receptorβ(ERβ) gene in the collected female Japanese flounder and to analyze the association of ERβpolymorphisms with reproductive indices by polymerase chain reaction single-strand conformation polymorphism (PCR-SSCP). Two single nucleotide polymorphisms (SNPs), SNP1 (c.577de1C) and SNP2 [c.A891T (p.Gln1 14Leu)], were identified in the ERβgene. A one-way ANOVA revealed that SNP1 was significantly associated with the gonadosomatic index (GSI) in female Japanese flounder (P<0.05). And SNP2 was significantly associated with the serum 17β-estradiol (E2) level and GSI (P<0.05). Individuals with genotype AB of SNP2 had significantly higher serum E2 level and GSI than those of genotype AA (P< 0.05).Moreover, the hepatosomatic index (HSI), a marker for genetic effects, was significantly higher for diplotype D2 compared with the other three diplotypes (P<0.05).
4. FOXL2 gene plays an important role in ovarian development, granulosa cell differentiation, and thus the proper maintenance of ovarian function. The aims of this study were to characterize polymorphisms within the FOXL2 gene in the collected female Japanese flounder and analyze the association of FOXL2 polymorphisms with reproductive performance by polymerase chain reaction-single stranded conformational polymorphism (PCR-SSCP). Results indicated that five Single nucleotide polymorphisms (SNPs), which were SNP1 [c.540A>C (p.Asn102His) and c.591A>G (p.Asn119Asp)], SNP2 [c.864G>A (p.Lys210Glu) and c.875G>A] and SNP3 (c.1169C>A), were identified in the FOXL2 gene. General Linear Model (GLM) analysis showed that SNP1 in the forkhead domain was significantly associated with gonadosomatic index (GSI) (P<0.05). SNP2 in the downstream of forkhead domain was significantly associated with serum 17β-estradiol (E2) level (P<0.05). And SNP3 in the 3'-UTR was significantly associated with hepatosomatic index (HSI) (P<0.05). Moreover, the evaluation of the genetic effects for both Testosterone (T) level of diplotype D3 and GSI of diplotype D5 suggested they were significantly higher than those of other four diplotypes (P<0.05), respectively.
5. Recent results suggested that membrane progestin receptor alpha (mPRα) mediates nongenomic actions of progestin hormones to induce oocyte maturation and sperm hypermotility in several teleost species. The role of mPRa in gamete and gonadal physiology was further evaluated in the present study by examining gonadal expression of mPRa during the gonadal development in male Japanese flounder.We cloned the mPRa cDNA from the Japanese flounder ovary. The predicted protein showed high sequence identity with Southern flounder (99%),Seatrout(94%) and Atlantic croaker(93%) mPRα. Computer modeling predicted that the deduced protein has seven transmembrance domains.The mPRα mRNA was detected in all the tissues tested with relatively greater expression in brain, ovary, testis and kidney. The mPRa mRNA was also expressed in Japanese flounder testes throughout the gonadal cycle. And the serumal E2 and GSI level showed the similar variation trend with the pattern of mPRa mRNA expression. Taken together, these data supported the hypothesis that mPRa is the MIS receptor in Japanese flounder.
6. Recently we discovered a previously uncharacterized gene with the characteristics of a progesterone and adipoQ receptor (PAQR) in a flatfish, Japanese flounder. Here, we reported the cloning and partial characteristics of this hitherto unknown gene. Phylogenetic analysis clearly indicated the PAQR-like gene clade was between the PAQR7 clade and PAQR8 clade. The predicted protein showed high sequence identity with three-spined stickleback (77%) and medaka (72%) unnamed gene. Structural analyses of the translated cDNA suggested it encoded protein with seven transmembrance domains, typical of progesterone and adipoQ receptor. RT-PCR analysis of Japanese flounder tissue extracts showed significant expression of the PAQR-like gene in ovary, head kidney, testis and kidney, and lower expression in brain, gill, intestine, stomach and pyloric caecum. The PAQR-like gene mRNA was also expressed in Japanese flounder testes throughout the gonadal cycle. And the variation trend of GSI was similar with the pattern of the PAQR-like gene mRNA expression. These characteristics suggested that the PAQR-like gene is related to reproductive physiology in this species.
7. The identity of the maturation-inducing steroid (MIS) in Southern flounder, a marine teleost which is similar to Japanese flounder, is unknown. The purpose of the present study was to identify the major progestin produced during oocyte maturation (OM) in southern flounder and investigate whether its secretion increases during this process. Intact of ovarian follicles undergoing OM were incubated in vitro with tritiated [3H]pregnenolone precursor and the tritiated products were extracted, purified, and identified by TLC. We found the tritiated products were biosynthesized from [3H]pregnenolone including 20β-S, 17a,20p-DHP,11-deoxycortisol. During the oocyte maturation, there were a significant increase in the production of 20β-S, whereas the production of 11-deoxycortisol decreased. And the amount of 17α,20β-DHP remained almost the same. In vitro,20β-S and 17α,20β-DHP were most effective steroids to induce maturation of southern flounder oocytes. These results suggest that that 20β-S and 17α,20P-DHP maybe the naturally occurring MIS in the female southern flounder.
引文
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