栉孔扇贝（Chlamys farreri）Wnt4，β-catenin及Dax1基因的克隆及其与性腺发育相关性的研究
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摘要
性腺发育是发育生物学的热点研究领域之一。有关性腺发育相关基因的研究在脊椎动物特别是哺乳动物中已有了相当的认识。然而，在无脊椎动物特别是海洋贝类中相关的研究还是非常有限的。本研究以我国重要的水产经济贝类——栉孔扇贝（Chlamys farreri）为研究对象，采用cDNA末端快速扩增（rapid-amplification of cDNAends，RACE）技术获得了Wnt4，β-catenin及Dax1基因cDNA全长，进一步采用半定量RT-PCR技术检测了3个目的基因的组织表达特点；采用相对定量RT-PCR（quantitative real-time PCR，qRT-PCR）技术比较了其在扇贝精、卵巢以及不同发育阶段性腺中的表达差异；采用免疫组织化学技术对其在扇贝性腺中的细胞学定位和配子发生过程中的作用进行了分析；采用抑制剂DKK-1（经典Wnt信号抑制剂）和槲皮素（β-catenin基因的抑制剂）处理体外原代培养的扇贝生长期精巢细胞，探讨了3个目的基因在扇贝性腺发育和配子发生过程中的作用关系。此外，本文还采用荧光原位杂交技术（Fluorescencein situ hybridization，FISH）对Dax1在染色体上的定位进行了鉴定。
Wnt4（Wingless-type MMTV integration site family，member4）属于Wnt家族一员，是一种生长因子。从栉孔扇贝（C. farreri）卵巢中克隆得到的Wnt4基因的cDNA全长为1239bp，其开放阅读框（open reading frame，ORF）为1068bp，可以编码355个氨基酸，推导的氨基酸序列含有Wnt家族特有序列，且与多个物种的Wnt4同源性都在60%以上。半定量RT-PCR结果显示，除肾脏外，Wnt4基因在栉孔扇贝（C. farreri）的精巢，卵巢，闭壳肌，肝胰腺，鳃，外套膜组织中均有表达，暗示其参与了多样的生物学过程；然而其表达量普遍较低，提示其可能作为信号分子发挥作用。qRT-PCR结果表明：Wnt4基因在成熟期的精、卵巢中表达量最高，且精巢表达量显著高于卵巢，推测其可能参与两性性腺的发育和成熟过程的调控，并在精巢中的作用明显于卵巢。
Armadillo（ARM）家族是一类进化上高度保守的蛋白，含有大约由50个氨基酸组成的ARM序列，并在细胞信号的传导和细胞骨架的调节等多个生命过程中发挥作用。β-catenin是ARM家族中的一员，其作为一种重要的信号传导蛋白广泛地参与动物的发育过程。栉孔扇贝（C. farreri）β-catenin基因cDNA序列全长为3353bp，其中ORF为2511bp，该基因编码836个氨基酸。推导的β-catenin氨基酸序列高度保守，并含有ARM家族特有的重复序列及典型的N-末端及C-末端区。半定量RT-PCR结果表明：β-catenin亦在扇贝的多种组织中表达；qRT-PCR结果显示：随着性腺的发育和成熟，β-catenin在性腺中的表达量逐渐升高，并在成熟期达到最高，随后的休止期性腺中表达量急剧下降到最低水平；与此同时，β-catenin的表达还呈现出性别二态性模式，即在卵巢中的表达量显著高于同时期的精巢（P<0.05）。免疫组织化学结果显示，栉孔扇贝（C. farreri）β-catenin蛋白主要定位在生殖细胞中，即精巢中的精原细胞和精母细胞、卵巢中的卵原细胞和卵母细胞，这一表达特征暗示了该基因可能参与栉孔扇贝（C.farreri）两性配子的发生过程。
DAX1[dosage-sensitive sex reversal-adrenal hypoplasia congenital (AHC)critical region on the X chromosome gene1]是核受体超家族的一员，在几个脊椎动物物种中被认为参与性别决定及性腺发育。栉孔扇贝（C. farreri）Dax1基因cDNA全长为2093bp，其中包括1404bp的ORF，编码467个氨基酸。栉孔扇贝（C.farreri）DAX1蛋白在其假定的DNA结合区中没有哺乳动物类的LXXLL保守序列。荧光原位杂交技术标定栉孔扇贝（C. farreri）Dax1基因位于一对亚端部着丝粒染色体的短臂上。半定量RT-PCR结果显示，栉孔扇贝（C. farreri）Dax1基因在所有检测的组织中均有表达；qRT-PCR检测栉孔扇贝（C. farreri）不同发育时期的性腺（增殖期，生长期，成熟期）中均表达Dax1基因，且在性腺中也呈现性别二态性表达特点，但其在精巢中的表达量显著性高于同时期的卵巢（p<0.05）；与β-catenin蛋白的细胞学定位一致，栉孔扇贝（C. farreri）DAX1蛋白也定位在精巢中的精原细胞、精母细胞以及卵巢的卵原细胞、卵母细胞中，表明DAX1可能参与了栉孔扇贝的配子发生过程。
抑制剂实验结果显示，当向栉孔扇贝（C. farreri）精巢细胞体外原代培养体系中添加0.1μg/ml和0.2μg/ml的DKK-148h时，qRT-PCR检测发现，原代培养细胞中的β-catenin表达量显著下调（p<0.05），表明在栉孔扇贝（C. farreri）精巢中存在经典的Wnt信号通路。当向栉孔扇贝（C. farreri）精巢细胞体外原代培养体系中添加50μmol/L和100μmol/L的槲皮素时，处理48h，Dax1基因的表达量显著性降低（p<0.05），暗示Dax1作为β-catenin的下游基因参与性腺的发育和配子发生过程。
Gonadal development is one of the major research fields in the developmentalbiology. It has been cognized to some extent on genes of gonadal development invertebrates especially in mammals. While, limited information about the field isavailable in invertebrates especially in marine bivalves. In the present study, wecloned full-length cDNA sequences of the genes Wnt4, β-catenin and Dax1from theimportant commercial bivalves in china-scallop Chlamys farreri by the method ofrapid-amplification of cDNA ends (RACE). Furthermore, tissue distributions of thethree target genes were detected by semi-quantitative RT-PCR technique (sqRT-PCR),and expressions of the three genes in gonads during different developmental stageswere analysized by quantitative real-time PCR (qRT-PCR) technique.Immunohistochemistry technique was employed to determine the cytologicallocations of the three genes and their functions were analysized during thegametogenesis. Two repressors, DKK-1(an inhibitor of canonical Wnt signalmolecule) and quercetin (an inhibitor of β-catenin) were used to treat C. farreritestis cells cultivated in vitro to probe into their roles of the three genes during thegonadal development and the gametogenesis. In addition, we identified the location ofthe gene Dax1in chromosomes by fluorescence in situ hybridization (FISH)technique.
Wnt4(Wingless-type MMTV integration site family, member4), a growth factor, ismember of Wnt family. The full-length cDNA sequence of Wnt4from C. farreri ovarywas obtained which was1,239bp including an open reading frame (ORF) of1,068bpwhich encoded355amino acids. The deduced amino acid sequence of Wnt4had theconserved sequence of Wnt family, and identity of over60%with most species Wnt4s. Semi-quantative RT-PCR result showed that C. farreri Wnt4expressed with lowexpression level in most detected tissues including testes, ovaries, adductor muscle,hepatopancreas, gill and mantle, except kidney, implying that Wnt4could involve in avarious of biological processes and played roles as a signal molecule in C. farreri.qRT-PCR results indicated that C. farreri Wnt4mRNA expression was the highestlevel in testis and ovary at mature stage, and higher in testis than in ovay at the samestage during the whole reproductive cycle. We inferred that Wnt4contributed to thedevelopmental processes of gonad and germ cell maturation.
Armadillo (ARM) family is composed of highly conserved proteins, containingthe ARM motif consisted of about50amino aids and playing roles in variousbiological processes, such as the transduction of cell signal and the regulation ofcytoskeleton. β-catenin is a member of ARM family and as a key signal transductionprotein wildly involved in the development process of animals. The whole cDNAsequence of C. farreri β-catenin was3,353bp in length including an ORF of2,511bpwhich encoded836amino acids. The deduced amino acid sequence of β-catenin washighly conserved and had the repeated sequences of ARM family, and the typicalN-terminal and C-terminal region. Semi-quantative RT-PCR result revealed that C.farreri β-catenin was also expressed in multiple tissues. qRT-PCR result showed thatthe expression of β-catenin in gonad increased gradually as the gonad developmentand mature, and reached the highest level at mature stage, and then it decreasedsharply to the lowest level at the resting stage. Meanwhile, the expression of β-cateninin gonad presented a sexually dimorphic pattern, that was with the significant higherexpression in ovaries than in testis at the same stage (P<0.05). Immunohistochemistryresult showed that β-catenin protein was mainly located in germ cells, such asspermatogonia and spermatocytes in testis, oogonia and oocytes in ovary, indicatingthat this gene maybe participate in the gametogenesis in C. farreri.
DAX1[dosage-sensitive sex reversal-adrenal hypoplasia congenital (AHC)critical region on the X chromosome gene1], a member of nuclear receptorsuperfamily, is known to involve in sex determination and gonadal development inseveral vertebrate species. The cDNA sequence of C. farreri Dax1is2,093bp in full-length including1,404bp ORF encoding a predicted467-amino acid. Unlike invertebrates, no conserved LXXLL-related motif existed in putative DNA bindingregion of C. farreri DAX1. FISH result showed that C. farreri Dax1was located onthe short arm of a pair of subtelocentric chromosome. Semi-quantitative RT-PCRrevealed that C. farreri Dax1mRNA expressed wildly in adult scallop tissues. Duringthe reproductive cycle (proliferative stage, growing stage and mature stage) of C.farreri, the Dax1also assumed a sex-dimorphic expression pattern, whereas with asignificant higher expression in testis than in ovay at the same stage (p<0.05). Inaccordance with the cytolocalization of β-catenin, C. farreri DAX1was also mainlylocated in spermatogonia and spermatocytes of testis, in oogonia and oocytes of ovary,implying that DAX1may involved in gametogenesis in C. farreri.
When DKK-1with concentrations of0.1μg/ml and0.2μg/ml were added to themedium for48h, the expression of β-catenin in the C. farreri testis cells in vitrowas significantly down-regulated (p<0.05) in treated groups compared with thecontrol group detected by qRT-PCR, which indicated the canonical Wnt pathwayexisted in C. farreri testis. Furthermore，when, quercetin with concentrations of50μmol/L and100μmol/L were added to the medium for48h respectively, also theexpression level of Dax1significantly decreased compared with the control, implyingDax1acts as the downstream gene of β-catenin to regulate the gonadal developmentand gametogenesis.
Wnt4（Wingless-type MMTV integration site family，member4）属于Wnt家族一员，是一种生长因子。从栉孔扇贝（C. farreri）卵巢中克隆得到的Wnt4基因的cDNA全长为1239bp，其开放阅读框（open reading frame，ORF）为1068bp，可以编码355个氨基酸，推导的氨基酸序列含有Wnt家族特有序列，且与多个物种的Wnt4同源性都在60%以上。半定量RT-PCR结果显示，除肾脏外，Wnt4基因在栉孔扇贝（C. farreri）的精巢，卵巢，闭壳肌，肝胰腺，鳃，外套膜组织中均有表达，暗示其参与了多样的生物学过程；然而其表达量普遍较低，提示其可能作为信号分子发挥作用。qRT-PCR结果表明：Wnt4基因在成熟期的精、卵巢中表达量最高，且精巢表达量显著高于卵巢，推测其可能参与两性性腺的发育和成熟过程的调控，并在精巢中的作用明显于卵巢。
Armadillo（ARM）家族是一类进化上高度保守的蛋白，含有大约由50个氨基酸组成的ARM序列，并在细胞信号的传导和细胞骨架的调节等多个生命过程中发挥作用。β-catenin是ARM家族中的一员，其作为一种重要的信号传导蛋白广泛地参与动物的发育过程。栉孔扇贝（C. farreri）β-catenin基因cDNA序列全长为3353bp，其中ORF为2511bp，该基因编码836个氨基酸。推导的β-catenin氨基酸序列高度保守，并含有ARM家族特有的重复序列及典型的N-末端及C-末端区。半定量RT-PCR结果表明：β-catenin亦在扇贝的多种组织中表达；qRT-PCR结果显示：随着性腺的发育和成熟，β-catenin在性腺中的表达量逐渐升高，并在成熟期达到最高，随后的休止期性腺中表达量急剧下降到最低水平；与此同时，β-catenin的表达还呈现出性别二态性模式，即在卵巢中的表达量显著高于同时期的精巢（P<0.05）。免疫组织化学结果显示，栉孔扇贝（C. farreri）β-catenin蛋白主要定位在生殖细胞中，即精巢中的精原细胞和精母细胞、卵巢中的卵原细胞和卵母细胞，这一表达特征暗示了该基因可能参与栉孔扇贝（C.farreri）两性配子的发生过程。
DAX1[dosage-sensitive sex reversal-adrenal hypoplasia congenital (AHC)critical region on the X chromosome gene1]是核受体超家族的一员，在几个脊椎动物物种中被认为参与性别决定及性腺发育。栉孔扇贝（C. farreri）Dax1基因cDNA全长为2093bp，其中包括1404bp的ORF，编码467个氨基酸。栉孔扇贝（C.farreri）DAX1蛋白在其假定的DNA结合区中没有哺乳动物类的LXXLL保守序列。荧光原位杂交技术标定栉孔扇贝（C. farreri）Dax1基因位于一对亚端部着丝粒染色体的短臂上。半定量RT-PCR结果显示，栉孔扇贝（C. farreri）Dax1基因在所有检测的组织中均有表达；qRT-PCR检测栉孔扇贝（C. farreri）不同发育时期的性腺（增殖期，生长期，成熟期）中均表达Dax1基因，且在性腺中也呈现性别二态性表达特点，但其在精巢中的表达量显著性高于同时期的卵巢（p<0.05）；与β-catenin蛋白的细胞学定位一致，栉孔扇贝（C. farreri）DAX1蛋白也定位在精巢中的精原细胞、精母细胞以及卵巢的卵原细胞、卵母细胞中，表明DAX1可能参与了栉孔扇贝的配子发生过程。
抑制剂实验结果显示，当向栉孔扇贝（C. farreri）精巢细胞体外原代培养体系中添加0.1μg/ml和0.2μg/ml的DKK-148h时，qRT-PCR检测发现，原代培养细胞中的β-catenin表达量显著下调（p<0.05），表明在栉孔扇贝（C. farreri）精巢中存在经典的Wnt信号通路。当向栉孔扇贝（C. farreri）精巢细胞体外原代培养体系中添加50μmol/L和100μmol/L的槲皮素时，处理48h，Dax1基因的表达量显著性降低（p<0.05），暗示Dax1作为β-catenin的下游基因参与性腺的发育和配子发生过程。
Gonadal development is one of the major research fields in the developmentalbiology. It has been cognized to some extent on genes of gonadal development invertebrates especially in mammals. While, limited information about the field isavailable in invertebrates especially in marine bivalves. In the present study, wecloned full-length cDNA sequences of the genes Wnt4, β-catenin and Dax1from theimportant commercial bivalves in china-scallop Chlamys farreri by the method ofrapid-amplification of cDNA ends (RACE). Furthermore, tissue distributions of thethree target genes were detected by semi-quantitative RT-PCR technique (sqRT-PCR),and expressions of the three genes in gonads during different developmental stageswere analysized by quantitative real-time PCR (qRT-PCR) technique.Immunohistochemistry technique was employed to determine the cytologicallocations of the three genes and their functions were analysized during thegametogenesis. Two repressors, DKK-1(an inhibitor of canonical Wnt signalmolecule) and quercetin (an inhibitor of β-catenin) were used to treat C. farreritestis cells cultivated in vitro to probe into their roles of the three genes during thegonadal development and the gametogenesis. In addition, we identified the location ofthe gene Dax1in chromosomes by fluorescence in situ hybridization (FISH)technique.
Wnt4(Wingless-type MMTV integration site family, member4), a growth factor, ismember of Wnt family. The full-length cDNA sequence of Wnt4from C. farreri ovarywas obtained which was1,239bp including an open reading frame (ORF) of1,068bpwhich encoded355amino acids. The deduced amino acid sequence of Wnt4had theconserved sequence of Wnt family, and identity of over60%with most species Wnt4s. Semi-quantative RT-PCR result showed that C. farreri Wnt4expressed with lowexpression level in most detected tissues including testes, ovaries, adductor muscle,hepatopancreas, gill and mantle, except kidney, implying that Wnt4could involve in avarious of biological processes and played roles as a signal molecule in C. farreri.qRT-PCR results indicated that C. farreri Wnt4mRNA expression was the highestlevel in testis and ovary at mature stage, and higher in testis than in ovay at the samestage during the whole reproductive cycle. We inferred that Wnt4contributed to thedevelopmental processes of gonad and germ cell maturation.
Armadillo (ARM) family is composed of highly conserved proteins, containingthe ARM motif consisted of about50amino aids and playing roles in variousbiological processes, such as the transduction of cell signal and the regulation ofcytoskeleton. β-catenin is a member of ARM family and as a key signal transductionprotein wildly involved in the development process of animals. The whole cDNAsequence of C. farreri β-catenin was3,353bp in length including an ORF of2,511bpwhich encoded836amino acids. The deduced amino acid sequence of β-catenin washighly conserved and had the repeated sequences of ARM family, and the typicalN-terminal and C-terminal region. Semi-quantative RT-PCR result revealed that C.farreri β-catenin was also expressed in multiple tissues. qRT-PCR result showed thatthe expression of β-catenin in gonad increased gradually as the gonad developmentand mature, and reached the highest level at mature stage, and then it decreasedsharply to the lowest level at the resting stage. Meanwhile, the expression of β-cateninin gonad presented a sexually dimorphic pattern, that was with the significant higherexpression in ovaries than in testis at the same stage (P<0.05). Immunohistochemistryresult showed that β-catenin protein was mainly located in germ cells, such asspermatogonia and spermatocytes in testis, oogonia and oocytes in ovary, indicatingthat this gene maybe participate in the gametogenesis in C. farreri.
DAX1[dosage-sensitive sex reversal-adrenal hypoplasia congenital (AHC)critical region on the X chromosome gene1], a member of nuclear receptorsuperfamily, is known to involve in sex determination and gonadal development inseveral vertebrate species. The cDNA sequence of C. farreri Dax1is2,093bp in full-length including1,404bp ORF encoding a predicted467-amino acid. Unlike invertebrates, no conserved LXXLL-related motif existed in putative DNA bindingregion of C. farreri DAX1. FISH result showed that C. farreri Dax1was located onthe short arm of a pair of subtelocentric chromosome. Semi-quantitative RT-PCRrevealed that C. farreri Dax1mRNA expressed wildly in adult scallop tissues. Duringthe reproductive cycle (proliferative stage, growing stage and mature stage) of C.farreri, the Dax1also assumed a sex-dimorphic expression pattern, whereas with asignificant higher expression in testis than in ovay at the same stage (p<0.05). Inaccordance with the cytolocalization of β-catenin, C. farreri DAX1was also mainlylocated in spermatogonia and spermatocytes of testis, in oogonia and oocytes of ovary,implying that DAX1may involved in gametogenesis in C. farreri.
When DKK-1with concentrations of0.1μg/ml and0.2μg/ml were added to themedium for48h, the expression of β-catenin in the C. farreri testis cells in vitrowas significantly down-regulated (p<0.05) in treated groups compared with thecontrol group detected by qRT-PCR, which indicated the canonical Wnt pathwayexisted in C. farreri testis. Furthermore，when, quercetin with concentrations of50μmol/L and100μmol/L were added to the medium for48h respectively, also theexpression level of Dax1significantly decreased compared with the control, implyingDax1acts as the downstream gene of β-catenin to regulate the gonadal developmentand gametogenesis.
引文
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