猪ADAMTS1基因的克隆、定位、表达规律、遗传效应和调控机理的研究
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摘要
含凝血酶敏感蛋白模体的去解联金属蛋白酶(a disintegrin and metalloproteinasewith thrombospondin motifs-1,ADAMTS1)是一种存在于哺乳动物和无脊椎动物中的细胞外金属蛋白酶。研究发现ADAMTS1基因通过孕酮受体信号途径在卵泡中发挥着重要的作用。在ADAMTS1基因敲除的雌性小鼠中,发现雌性生殖器官畸形和不能完成排卵等现象。同时发现其在卵丘-卵母细胞复合体(cumulus-oocytecomplexes,COC)的扩张过程中发挥着重要的作用。以上现象说明ADAMTS1基因是卵泡破裂过程中的关键因素并以蛋白酶的角色控制着排卵。因此,本研究对猪ADAMTS1基因进行了分离和定位,并对其表达规律、遗传效应和调控机理进行了研究,取得了以下结果:
(1)根据与人ADAMTS1基因有较高同源性的猪EST以及人和小鼠ADAMTS1cDNA相似性较高的区域设计引物,以梅山猪DNA作为模板扩增、克隆并测序,拼接后获得9026 bp的整合DNA序列,包含全部CDS和内含子以及部分5'和3'非翻译区。ADAMTS1基因GenBank登录号为DQ177331。
(2)利用Clustal X、ORF Finder、Signal P2.0、ANTHEPORT、PSORTⅡ、TMoreo等软件对猪ADAMTS1基因结构、编码的蛋白质结构、功能等特征进行了预测和分析。通过生物信息学分析,预测猪ADAMTS1蛋白铆定在细胞膜上,没有跨膜区域,拥有一个解联蛋白区,一个包含锌指结构的金属蛋白酶区和三个血小板反应蛋白结构、两个潜在的糖基化位点并存在很多半胱氨酸残基。
(3)根据猪ADAMTS1基因DNA序列设计引物,利用猪辐射杂种克隆板(ImpRH)将猪ADAMTS1基因定位于13号染色体SSC13q49。两点连锁分析发现与猪ADAMTS1基因连锁最紧密的标记是S0215(LOD=14.18)。
(4)根据已获得的DNA序列设计引物,分别以大白和梅山猪DNA作为模板进行扩增、克隆并测序。经序列比对,发现了2个SNP,第7外显子72bp处出现碱基C/G颠换,导致622位氨基酸由精氨酸变为脯氨酸;第7内含子512bp处出现碱基G/A转换,据此建立适宜的PCR-RFLP SNP分型方法。
(5)利用所建立的PCR-RFLP检测方法,对猪ADAMTS1基因的2个SNP在7个中外猪群体中进行基因型分型,分析了它们的基因型频率和基因频率;并在116头新清平母猪中进行了与产仔数和产活仔数性状的关联性分析,结果表明这两个标记均与猪产仔数和产活仔数性状存在显著或极显著相关。
(6)在组织和细胞水平分析了猪ADAMTS1基因的表达模式,结果表明:猪ADAMTS1基因受促性腺激素的诱导,在排卵前卵泡颗粒细胞中瞬时表达。在细胞水平分析了猪孕酮受体(progesterone receptor,PR)基因的表达模式,结果表明:猪孕酮受体基因受促性腺激素的诱导,在ADAMTS1基因表达之前在卵泡颗粒细胞中瞬时表达。利用Western blot检测PR在卵巢颗粒细胞中的表达,证实在PMSG/hCG诱导后的卵巢颗粒细胞中存在孕酮受体两个亚型的表达。
(7)利用基因组PCR步移方法获得猪ADAMTS1基因转录起始位点上游1458bp的启动子序列。对猪ADAMTS1基因的启动子区序列进行了生物信息学分析,发现两个孕酮受体结合位点。通过以上预测结果,以1458bp的启动子序列为模板,以PCR方法获得4个5'侧翼缺失片断和3个定点突变孕酮受体结合位点的片断。将这些片断插入到载体pEGFP-1的报告基因EGFP的上游,构建相应的报告基因表达载体。将这些载体分别转染COS-7细胞,荧光显微镜观察和流式细胞仪分析启动子活性,发现在所构建的7个启动子片断之间没有显著的差异,这说明以上7个片断的启动子活性没有显著差异。
(8)根据与人孕酮受体基因有较高同源性的猪EST以及人和小鼠孕酮受体cDNA相似性较高的区域设计引物,PCR扩增孕酮受体的DNA结合区域(DNAbinding domain,DBD),构建表达载体pcDNA-PR DBD,并将其与以上构建的EGFP报告基因表达载体分别共转染COS-7细胞。荧光显微镜观察和流式细胞仪分析启动子活性,发现拥有孕酮受体结合位点的启动子片断的启动子活性显著高于没有或是突变此位点的启动子片断。利用EMSA检测ADAMTS1基因启动子上两个PR结合位点与PR的结合能力。EMSA试验在PMSG/hCG处理后卵巢颗粒细胞中的核蛋白中分别检测到了ADAMTS1基因启动子上两个PR结合位点与PR的特异性结合。以上结果说明:在ADAMTS1基因的启动子上存在孕酮受体结合位点,并且当孕酮受体通过其DBD区域结合在这些位点上时会增强ADAMTS1基因启动子的活性,启动ADAMTS1基因的表达。
A disintegrin-like and metalloprotease(reprolysin type) with thrombospondin type 1 motif(ADAMTS) is a novel family of extracellular proteases found in both mammals and invertebrates.Later research showed that ADAMTS1 plays an important role in follicles via progesterone receptor(PR)-dependent pathways.Female fertility is impaired in ADAMTS1 knocked out mice,and this is accompanied by obvious abnormalities of the uterus and ovaries.In addition,there is evidence that progesterone- and PR-dependent functions in cumulus cells are essential for cumulus-oocyte complexes(COC) expansion in pig,possibly through the action of ADAMTS1.All these data suggest that ADAMTS1 plays a critical role in follicular rupture and represents a major advance in the proteolytic events that control ovulation.In this research,the ADAMTS1 gene were cloned and identified and the promoter regulation and genetics effect were analyzed.The main results are as follows:
1.The sequences of cDNAs of human and mouse ADAMTS1 genes were used to search against the porcine EST databases.Six porcine ESTs sharing＞80%identity to the corresponding human and mouse cDNAs were obtained and assembled into a contig. Based on the contig sequence,a set of primers was designed to amplify the DNA sequence of porcine ADAMTS1 gene.These primers yielded twelve overlapping PCR products that produced a consensus sequence of 9026-bp DNA sequence containing the full coding region,all 8 introns and part of 5'and 3' untranslated region of the porcine ADAMTS1 gene was obtained(Genbank accession number DQ177331).
2.Using CLUSTAL W and some related software;we analyzed the gene structure, protein structure and conserved motifs.The hydropathy plot and the TMAP prediction derived from the World-Wide Web service showed that porcine ADAMTS1 protein includes no transmembrane region and there are many cysteine residues and four putative N-glycosylation sites in the protein which indicates that the ADAMTS1 gene product is a putative cysteine-rich secretory glycoprotein. Sequence analyses of porcine ADAMTS1 protein by ExPASy revealed that the porcine ADAMTS1 protein consists of one disintegrinlike domain,one zinc-dependent metalloproteinase and three thrombospondin typeⅠrepeat.
3.Using the ImpRH panel,we determined that pig ADAMTS1 is closely linked with microsatellite marker S0215 on SSC13q49.
4.Comparison of the sequences of ADAMTS1 in different pig breeds by using BLAST revealed two SNP.One of them was found within exon 7 of which a G-C substitution changes the codon for arginine into proline that spans a PvuⅡrestriction site.The other was found within intron 7 of which a G-A substitution that spans a StyⅠrestriction site.
5.The substitutions were situated within PvuⅡand StyⅠrecognition site and developed as PCR-RFLP markers for further use in population variation investigations and association analysis with litter size.Allele frequencies of this SNP were investigated in seven pig breeds/lines.An association analysis in new Qingping female line(116 new Qingping female line pigs;247 litter records) suggested that different ADAMTS1 genotypes have significant differences in litter size.
6.RT-PCR results indicated that porcine ADAMTS1 gene is selectively induced in granulosa cells of preovulatory follicles by the hCG surge.Maximal levels of these proteases are observed at 12-16 h after an hCG surge,the time of ovulation.RT-PCR results indicated that porcine PR gene is induced in granulosa cells of preovulatory follicles by the hCG surge.Maximal levels of these proteases are observed at 8-12 h after an hCG surge,just before the ADAMTS1.Western blot results indicated that porcine PR protein is induced in granulosa cells of preovulatory follicles by the hCG surge.Maximal levels of these proteases are observed at 7-9 h after hCG surge.
7.Genome walking technique was used to clone the proximal promoter region of porcine ADAMTS1.A fragment of 1458bp upstream start code was obtained from genomic DNA of porcine.Use computer analyze,tow PR site were found.Based on this fragment,together with different mutants of the proximal promoter cloned by PCR were cloned upstream of the EGFP reporter constructs and were analyzed using transfection and flow cytometry.The result showed that all 7 EGFP reporter constructs have no significant differences.
8.Based on the porcine EST databases,a pair of primer was designed to amplify the DNA binding region of porcine PR gene.The DNA binding region of porcine PR gene was cloned into pcDNA3.1 vector.Co-transfection analysis was used to examine interaction between DNA binding region of porcine PR gene and ADAMTS1 promoter.The recombinant pcDNA-PR was co-transfected into COS-7 cells with ADAMTS1 promoter-EGFP.To confirm the interaction,three mutant ADAMTS1 promoter-EGFP for PR-binding site(mut) were also co-transfected with pcDNA-PR. Expression level in the cells were examined using flow cytometry.Electrophoratic mobility shift assay was employed to evaluate the combining activity of PR and ADAMTS1 promoter of granulosa cells.EMSA results indicated that the PR has binding activity on ADAMTS1 promer.The result showed that the PR plays important role in starting ADAMTS1 expression.
(1)根据与人ADAMTS1基因有较高同源性的猪EST以及人和小鼠ADAMTS1cDNA相似性较高的区域设计引物,以梅山猪DNA作为模板扩增、克隆并测序,拼接后获得9026 bp的整合DNA序列,包含全部CDS和内含子以及部分5'和3'非翻译区。ADAMTS1基因GenBank登录号为DQ177331。
(2)利用Clustal X、ORF Finder、Signal P2.0、ANTHEPORT、PSORTⅡ、TMoreo等软件对猪ADAMTS1基因结构、编码的蛋白质结构、功能等特征进行了预测和分析。通过生物信息学分析,预测猪ADAMTS1蛋白铆定在细胞膜上,没有跨膜区域,拥有一个解联蛋白区,一个包含锌指结构的金属蛋白酶区和三个血小板反应蛋白结构、两个潜在的糖基化位点并存在很多半胱氨酸残基。
(3)根据猪ADAMTS1基因DNA序列设计引物,利用猪辐射杂种克隆板(ImpRH)将猪ADAMTS1基因定位于13号染色体SSC13q49。两点连锁分析发现与猪ADAMTS1基因连锁最紧密的标记是S0215(LOD=14.18)。
(4)根据已获得的DNA序列设计引物,分别以大白和梅山猪DNA作为模板进行扩增、克隆并测序。经序列比对,发现了2个SNP,第7外显子72bp处出现碱基C/G颠换,导致622位氨基酸由精氨酸变为脯氨酸;第7内含子512bp处出现碱基G/A转换,据此建立适宜的PCR-RFLP SNP分型方法。
(5)利用所建立的PCR-RFLP检测方法,对猪ADAMTS1基因的2个SNP在7个中外猪群体中进行基因型分型,分析了它们的基因型频率和基因频率;并在116头新清平母猪中进行了与产仔数和产活仔数性状的关联性分析,结果表明这两个标记均与猪产仔数和产活仔数性状存在显著或极显著相关。
(6)在组织和细胞水平分析了猪ADAMTS1基因的表达模式,结果表明:猪ADAMTS1基因受促性腺激素的诱导,在排卵前卵泡颗粒细胞中瞬时表达。在细胞水平分析了猪孕酮受体(progesterone receptor,PR)基因的表达模式,结果表明:猪孕酮受体基因受促性腺激素的诱导,在ADAMTS1基因表达之前在卵泡颗粒细胞中瞬时表达。利用Western blot检测PR在卵巢颗粒细胞中的表达,证实在PMSG/hCG诱导后的卵巢颗粒细胞中存在孕酮受体两个亚型的表达。
(7)利用基因组PCR步移方法获得猪ADAMTS1基因转录起始位点上游1458bp的启动子序列。对猪ADAMTS1基因的启动子区序列进行了生物信息学分析,发现两个孕酮受体结合位点。通过以上预测结果,以1458bp的启动子序列为模板,以PCR方法获得4个5'侧翼缺失片断和3个定点突变孕酮受体结合位点的片断。将这些片断插入到载体pEGFP-1的报告基因EGFP的上游,构建相应的报告基因表达载体。将这些载体分别转染COS-7细胞,荧光显微镜观察和流式细胞仪分析启动子活性,发现在所构建的7个启动子片断之间没有显著的差异,这说明以上7个片断的启动子活性没有显著差异。
(8)根据与人孕酮受体基因有较高同源性的猪EST以及人和小鼠孕酮受体cDNA相似性较高的区域设计引物,PCR扩增孕酮受体的DNA结合区域(DNAbinding domain,DBD),构建表达载体pcDNA-PR DBD,并将其与以上构建的EGFP报告基因表达载体分别共转染COS-7细胞。荧光显微镜观察和流式细胞仪分析启动子活性,发现拥有孕酮受体结合位点的启动子片断的启动子活性显著高于没有或是突变此位点的启动子片断。利用EMSA检测ADAMTS1基因启动子上两个PR结合位点与PR的结合能力。EMSA试验在PMSG/hCG处理后卵巢颗粒细胞中的核蛋白中分别检测到了ADAMTS1基因启动子上两个PR结合位点与PR的特异性结合。以上结果说明:在ADAMTS1基因的启动子上存在孕酮受体结合位点,并且当孕酮受体通过其DBD区域结合在这些位点上时会增强ADAMTS1基因启动子的活性,启动ADAMTS1基因的表达。
A disintegrin-like and metalloprotease(reprolysin type) with thrombospondin type 1 motif(ADAMTS) is a novel family of extracellular proteases found in both mammals and invertebrates.Later research showed that ADAMTS1 plays an important role in follicles via progesterone receptor(PR)-dependent pathways.Female fertility is impaired in ADAMTS1 knocked out mice,and this is accompanied by obvious abnormalities of the uterus and ovaries.In addition,there is evidence that progesterone- and PR-dependent functions in cumulus cells are essential for cumulus-oocyte complexes(COC) expansion in pig,possibly through the action of ADAMTS1.All these data suggest that ADAMTS1 plays a critical role in follicular rupture and represents a major advance in the proteolytic events that control ovulation.In this research,the ADAMTS1 gene were cloned and identified and the promoter regulation and genetics effect were analyzed.The main results are as follows:
1.The sequences of cDNAs of human and mouse ADAMTS1 genes were used to search against the porcine EST databases.Six porcine ESTs sharing＞80%identity to the corresponding human and mouse cDNAs were obtained and assembled into a contig. Based on the contig sequence,a set of primers was designed to amplify the DNA sequence of porcine ADAMTS1 gene.These primers yielded twelve overlapping PCR products that produced a consensus sequence of 9026-bp DNA sequence containing the full coding region,all 8 introns and part of 5'and 3' untranslated region of the porcine ADAMTS1 gene was obtained(Genbank accession number DQ177331).
2.Using CLUSTAL W and some related software;we analyzed the gene structure, protein structure and conserved motifs.The hydropathy plot and the TMAP prediction derived from the World-Wide Web service showed that porcine ADAMTS1 protein includes no transmembrane region and there are many cysteine residues and four putative N-glycosylation sites in the protein which indicates that the ADAMTS1 gene product is a putative cysteine-rich secretory glycoprotein. Sequence analyses of porcine ADAMTS1 protein by ExPASy revealed that the porcine ADAMTS1 protein consists of one disintegrinlike domain,one zinc-dependent metalloproteinase and three thrombospondin typeⅠrepeat.
3.Using the ImpRH panel,we determined that pig ADAMTS1 is closely linked with microsatellite marker S0215 on SSC13q49.
4.Comparison of the sequences of ADAMTS1 in different pig breeds by using BLAST revealed two SNP.One of them was found within exon 7 of which a G-C substitution changes the codon for arginine into proline that spans a PvuⅡrestriction site.The other was found within intron 7 of which a G-A substitution that spans a StyⅠrestriction site.
5.The substitutions were situated within PvuⅡand StyⅠrecognition site and developed as PCR-RFLP markers for further use in population variation investigations and association analysis with litter size.Allele frequencies of this SNP were investigated in seven pig breeds/lines.An association analysis in new Qingping female line(116 new Qingping female line pigs;247 litter records) suggested that different ADAMTS1 genotypes have significant differences in litter size.
6.RT-PCR results indicated that porcine ADAMTS1 gene is selectively induced in granulosa cells of preovulatory follicles by the hCG surge.Maximal levels of these proteases are observed at 12-16 h after an hCG surge,the time of ovulation.RT-PCR results indicated that porcine PR gene is induced in granulosa cells of preovulatory follicles by the hCG surge.Maximal levels of these proteases are observed at 8-12 h after an hCG surge,just before the ADAMTS1.Western blot results indicated that porcine PR protein is induced in granulosa cells of preovulatory follicles by the hCG surge.Maximal levels of these proteases are observed at 7-9 h after hCG surge.
7.Genome walking technique was used to clone the proximal promoter region of porcine ADAMTS1.A fragment of 1458bp upstream start code was obtained from genomic DNA of porcine.Use computer analyze,tow PR site were found.Based on this fragment,together with different mutants of the proximal promoter cloned by PCR were cloned upstream of the EGFP reporter constructs and were analyzed using transfection and flow cytometry.The result showed that all 7 EGFP reporter constructs have no significant differences.
8.Based on the porcine EST databases,a pair of primer was designed to amplify the DNA binding region of porcine PR gene.The DNA binding region of porcine PR gene was cloned into pcDNA3.1 vector.Co-transfection analysis was used to examine interaction between DNA binding region of porcine PR gene and ADAMTS1 promoter.The recombinant pcDNA-PR was co-transfected into COS-7 cells with ADAMTS1 promoter-EGFP.To confirm the interaction,three mutant ADAMTS1 promoter-EGFP for PR-binding site(mut) were also co-transfected with pcDNA-PR. Expression level in the cells were examined using flow cytometry.Electrophoratic mobility shift assay was employed to evaluate the combining activity of PR and ADAMTS1 promoter of granulosa cells.EMSA results indicated that the PR has binding activity on ADAMTS1 promer.The result showed that the PR plays important role in starting ADAMTS1 expression.
引文
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