DAZ基因家族DAZL/BOULE与犏牛雄性不育关系的研究
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摘要
牦牛是牛属动物中唯一能适应高寒气候环境而延续至今的牛种,属珍稀畜种资源和生产性能低的原始牛种,也是世界屋脊的“景观牛种”。为了提高其生产性能,人们一直利用普通牛同牦牛进行种间杂交,获得具有明显杂种优势的一代杂种犏牛。但由于犏牛雄性不育,种间杂交不能达到改良牦牛遗传本质和充分利用牦牛遗传资源的目的,杂种优势的利用和新型牛种的培育均受到了极大的限制,成为牦牛杂交改良的“拦路虎”。近几十年来,为探讨犏牛雄性不育的问题,国内外许多学者对犏牛雄性不育这一复杂的生物现象坐了很多有意义的研究和探讨,但至今尚未找到犏牛雄性不育的主导原因。DAZ基因家族包括DAZ(Deleted in Azoospermia)、DAZL(DAZ-like)和BOULE基因,是精子生成的重要调控因子。DAZL起源于BOULE基因,位于人类2号染色体,BOULE位于人类的3号染色体上,是DAZ基因家族的祖先基因。
为了探讨犏牛雄性不育问题,本研究以成年健康雄性黄牛、牦牛和犏牛为研究对象,运用组织切片、RT-PCR克隆测序、Real-time PCR技术、原位杂交技术以及二级结构预测和系统发育分析对DAZL和BOULE基因的组织表达、在睾丸组织中的表达差异水平分析,和BOULE基因在黄牛精子发生中的组织定位,为研究犏牛雄性不育与DAZ基因家族的关系提供理论基础。
1.牦牛、犏牛DAZL基因的克隆、序列分析及表达的研究
本试验根据黄牛DAZL基因序列EF501823设计引物扩增出牦牛和犏牛全长分别为1772bp、1778bp的序列,两段序列包含完整的ORF编码区,全长为885bp,编码295个氨基酸。生物信息学研究发现,黄牛编码区序列与牦牛和犏牛分别为同源性分别为99.43%、98.31%,与其他动物也具有较高的同源性。含有DAZ家族所具有的典型的RRM结构域、DAZ重复基序,同时在N端还具有高度保守的PABP作用位点和Pumilio-2作用位点,说明脊椎动物的DAZL基因在进化上是高度保守的。系统发育树分析,聚类结果与经典分类学结果基本一致,实时荧光定量PCR显示,DAZL在三种牛睾丸组织中的表达量中,黄牛与牦牛差异显著(P<0.05),黄牛与犏牛,犏牛与牦牛差异不显著(P>0.05).推测DAZL基因与犏牛雄性不育无关.
2.牦牛、犏牛BOULE基因的克隆、序列分析及表达的研究
本试验根据黄牛BOULE基因序列(NM_001102115)设计引物扩增出牦牛和犏牛的全长分别为1647bp、1650bp,该序列包含完整的编码区,二者也都含有885bp的相同的完整开放阅读框(ORF),编码一个295个氨基酸蛋白,进行同源性比对,结果发现牦牛、犏牛和黄牛BOULE基因编码区核苷酸序列的同源性为99.85%,氨基酸序列的同源性为98.31%,在编码区内,三个物种共发现了4处碱基差异(0.45／100bp),与其他动物也具有较高的同源性,也含有DAZ基因家族所具有的典型的RRM结构域、DAZ重复基序。系统发育分析与经典分类结果也基本一致。实时荧光定量PCR显示,BOULE基因在三种牛睾丸组织中的表达量中,黄牛与牦牛差异不显著(P>0.05),黄牛、牦牛,与犏牛差异显著(P<0.05)。推测BOULE基因与犏牛雄性不育有一定的关系。
3.黄牛、牦牛、犏牛睾丸组织学观察和BOULE m RNA在睾丸组织的表达研究
制作黄牛、牦牛和犏牛睾丸组织的的石蜡切片并HE染色发现,黄牛睾丸组织生精小管发育正常,可见各级生精细胞,内有不同时期大量的生精细胞,间质组织和生精小管无间隙小或间隙小,而牦牛睾丸组织生精小管也发育正常,可见各级生精细胞,犏牛睾丸组织生精小管官腔大小及形态不一致,管壁皱缩,生精小管上皮多由1～2层精原细胞构成,有些精原细胞已经死亡,初级精母细胞很少,无次级精母细胞犏牛生精小管由1～2层精原细胞构成,精母细胞少,间质组织与基底膜间隙宽,由组织液填充。原位杂交结果显示,BOULE基因主要分布在初级精母细胞和次级精母细胞中,在其余各级细胞中未发现其表达,推测犏牛精子发生失败,生精细胞凋亡等表象可能与BOULE基因的表达有一定的关系。
The yak,known as the "landscapes species of the bos" in the roof of the world, which belong the unique topographic features and low reproduction the original bovine, and the only one bovine now is well adapted to high altitude enviroments. To improve its productivity, hybridization between cattle and yaks has been carried out since long long ago, Cattle yaks, the F1 hybrid between cattle and yaks, exhibit significant hybrid vigor. However, the males are sterile, which greatly restricts the utilization of this hybrid vigor, and is a "stumbling block" in the varietal improvement of the yak and use of genetic resources. Over the past decades, the cattle-yak males sterile were significant studied by a lot of scholars about the complex phenomenon at home and abroad, but the main reason about the males sterile was had not found. Genes of the DAZ(Deleted in Azoospermia) gene famiy, DAZ、DAZL(DAZ-like)and BOULE, DAZL was originated from BOULE on Chromosome 2q, BOULE is the ancestral gene of DAZ gene family on Chromosome 3q.
1) Cloning, characterization and expression of coding region of DAZL gene in yak and cattle-yak
Primers were designed from the sequence of the cattle DAZL m RNA (EF501823), the PCR amplification results s howed that the sequences were 1782 base pairs long and 1778 base pairs long of yak and cattle-yak,the encoding region contained 885bp encoding 285 amino acid residues, the encoding region contained 885bp encoding 285 amino acid residues. The homologies of the coding region amino acid residues between the yak and cattle-yak with the cattle were 99.43%and 98.31%respectively.The yak and the cattle-yak DAZL gene including one typical RNA recognitong motif (RRM)domains, multiple DAZ repeats, the PABP and the Pumilio-2 highly conserved action Site in N-terminal as in DAZ family, the DAZL gene was conserved in vertebrate evolution. The cluster result shows that was identical the classical classification. The results of Real-time PCR showed that the level of DAZL in cattle、yak and cattle-yak, the yak was significantly lower than that in cattle (P<0.05) and, and there was not significant difference between the yak and the cattle-yak, the cattle and the cattle-yak (P> 0.05),which proves that DAZL probably has no relationship with cattle-yak infertility.
2) Cloning、characterization and expression of coding region of BOULE gene in yak and cattle-yak
Primers were designed from the sequence of the cattle BOULE m RNA (NM_001102115), the PCR amplification results s howed that the sequences were 1647 base pairs long and 1650 base pairs long of yak and cattle-yak,the encoding region contained 885bp encoding 285amino acid residues. The homologies of the coding region amino acid residues between the yak and cattle-yak with the cattle were 99.85%and 98.31%respectively, There are 4 base difference in the three species (0.45/1 OObp),it has the high homology with other species. The yak and the cattle-yak BOULE gene also including one typical RNArecognitong motif (RRM)domains, multiple DAZ repeats. The cluster result shows that was identical the classical classification. The results of Real-time PCR showed that the level ofBOULE in cattle、yak and cattle-yak, the cattle-yak was significantly lower than that in cattle and yak (P<0.05), there was not significant difference between the yak and the cattle (P> 0.05),which proves that BOULE probably has relationship with cattle-yak infertility.
3) Histological observation in the testis tissue of cattle,yak and cattle-yak and the expression of BOULE m RNA in the testis tissue of cattle.
Manufaction of paraffin section on the testis of cattle,yak and cattle-yak and then stained with HE. The results shows that the seminiferous tubule of cattle-yak is much smaller than that of cattle and yak, the space between tubules is big, the size and shape of lumen is different, the tube wall is wizened, and there is no or only a thin flat of spermatogonia in epithelium of seminiferous tubule. Little primary spermatocyte can be seen, and no secondary spermatocyte, no sperm in epididymis; however, the seminiferous tubule tissue of cattle and yak testis develop normally, and there are all levels of spermatogenic. The sperm, which can be observed in epididymis, develop well. In situ hybridization demonstrated that the BOULE gene mainly distribution and expression on the primary spermatocyte and secondary spermatocyte.
The failure of spermatogenesis and spermatogenic cell with BOULE gene may leads to cattle-yak infertility.
为了探讨犏牛雄性不育问题,本研究以成年健康雄性黄牛、牦牛和犏牛为研究对象,运用组织切片、RT-PCR克隆测序、Real-time PCR技术、原位杂交技术以及二级结构预测和系统发育分析对DAZL和BOULE基因的组织表达、在睾丸组织中的表达差异水平分析,和BOULE基因在黄牛精子发生中的组织定位,为研究犏牛雄性不育与DAZ基因家族的关系提供理论基础。
1.牦牛、犏牛DAZL基因的克隆、序列分析及表达的研究
本试验根据黄牛DAZL基因序列EF501823设计引物扩增出牦牛和犏牛全长分别为1772bp、1778bp的序列,两段序列包含完整的ORF编码区,全长为885bp,编码295个氨基酸。生物信息学研究发现,黄牛编码区序列与牦牛和犏牛分别为同源性分别为99.43%、98.31%,与其他动物也具有较高的同源性。含有DAZ家族所具有的典型的RRM结构域、DAZ重复基序,同时在N端还具有高度保守的PABP作用位点和Pumilio-2作用位点,说明脊椎动物的DAZL基因在进化上是高度保守的。系统发育树分析,聚类结果与经典分类学结果基本一致,实时荧光定量PCR显示,DAZL在三种牛睾丸组织中的表达量中,黄牛与牦牛差异显著(P<0.05),黄牛与犏牛,犏牛与牦牛差异不显著(P>0.05).推测DAZL基因与犏牛雄性不育无关.
2.牦牛、犏牛BOULE基因的克隆、序列分析及表达的研究
本试验根据黄牛BOULE基因序列(NM_001102115)设计引物扩增出牦牛和犏牛的全长分别为1647bp、1650bp,该序列包含完整的编码区,二者也都含有885bp的相同的完整开放阅读框(ORF),编码一个295个氨基酸蛋白,进行同源性比对,结果发现牦牛、犏牛和黄牛BOULE基因编码区核苷酸序列的同源性为99.85%,氨基酸序列的同源性为98.31%,在编码区内,三个物种共发现了4处碱基差异(0.45／100bp),与其他动物也具有较高的同源性,也含有DAZ基因家族所具有的典型的RRM结构域、DAZ重复基序。系统发育分析与经典分类结果也基本一致。实时荧光定量PCR显示,BOULE基因在三种牛睾丸组织中的表达量中,黄牛与牦牛差异不显著(P>0.05),黄牛、牦牛,与犏牛差异显著(P<0.05)。推测BOULE基因与犏牛雄性不育有一定的关系。
3.黄牛、牦牛、犏牛睾丸组织学观察和BOULE m RNA在睾丸组织的表达研究
制作黄牛、牦牛和犏牛睾丸组织的的石蜡切片并HE染色发现,黄牛睾丸组织生精小管发育正常,可见各级生精细胞,内有不同时期大量的生精细胞,间质组织和生精小管无间隙小或间隙小,而牦牛睾丸组织生精小管也发育正常,可见各级生精细胞,犏牛睾丸组织生精小管官腔大小及形态不一致,管壁皱缩,生精小管上皮多由1～2层精原细胞构成,有些精原细胞已经死亡,初级精母细胞很少,无次级精母细胞犏牛生精小管由1～2层精原细胞构成,精母细胞少,间质组织与基底膜间隙宽,由组织液填充。原位杂交结果显示,BOULE基因主要分布在初级精母细胞和次级精母细胞中,在其余各级细胞中未发现其表达,推测犏牛精子发生失败,生精细胞凋亡等表象可能与BOULE基因的表达有一定的关系。
The yak,known as the "landscapes species of the bos" in the roof of the world, which belong the unique topographic features and low reproduction the original bovine, and the only one bovine now is well adapted to high altitude enviroments. To improve its productivity, hybridization between cattle and yaks has been carried out since long long ago, Cattle yaks, the F1 hybrid between cattle and yaks, exhibit significant hybrid vigor. However, the males are sterile, which greatly restricts the utilization of this hybrid vigor, and is a "stumbling block" in the varietal improvement of the yak and use of genetic resources. Over the past decades, the cattle-yak males sterile were significant studied by a lot of scholars about the complex phenomenon at home and abroad, but the main reason about the males sterile was had not found. Genes of the DAZ(Deleted in Azoospermia) gene famiy, DAZ、DAZL(DAZ-like)and BOULE, DAZL was originated from BOULE on Chromosome 2q, BOULE is the ancestral gene of DAZ gene family on Chromosome 3q.
1) Cloning, characterization and expression of coding region of DAZL gene in yak and cattle-yak
Primers were designed from the sequence of the cattle DAZL m RNA (EF501823), the PCR amplification results s howed that the sequences were 1782 base pairs long and 1778 base pairs long of yak and cattle-yak,the encoding region contained 885bp encoding 285 amino acid residues, the encoding region contained 885bp encoding 285 amino acid residues. The homologies of the coding region amino acid residues between the yak and cattle-yak with the cattle were 99.43%and 98.31%respectively.The yak and the cattle-yak DAZL gene including one typical RNA recognitong motif (RRM)domains, multiple DAZ repeats, the PABP and the Pumilio-2 highly conserved action Site in N-terminal as in DAZ family, the DAZL gene was conserved in vertebrate evolution. The cluster result shows that was identical the classical classification. The results of Real-time PCR showed that the level of DAZL in cattle、yak and cattle-yak, the yak was significantly lower than that in cattle (P<0.05) and, and there was not significant difference between the yak and the cattle-yak, the cattle and the cattle-yak (P> 0.05),which proves that DAZL probably has no relationship with cattle-yak infertility.
2) Cloning、characterization and expression of coding region of BOULE gene in yak and cattle-yak
Primers were designed from the sequence of the cattle BOULE m RNA (NM_001102115), the PCR amplification results s howed that the sequences were 1647 base pairs long and 1650 base pairs long of yak and cattle-yak,the encoding region contained 885bp encoding 285amino acid residues. The homologies of the coding region amino acid residues between the yak and cattle-yak with the cattle were 99.85%and 98.31%respectively, There are 4 base difference in the three species (0.45/1 OObp),it has the high homology with other species. The yak and the cattle-yak BOULE gene also including one typical RNArecognitong motif (RRM)domains, multiple DAZ repeats. The cluster result shows that was identical the classical classification. The results of Real-time PCR showed that the level ofBOULE in cattle、yak and cattle-yak, the cattle-yak was significantly lower than that in cattle and yak (P<0.05), there was not significant difference between the yak and the cattle (P> 0.05),which proves that BOULE probably has relationship with cattle-yak infertility.
3) Histological observation in the testis tissue of cattle,yak and cattle-yak and the expression of BOULE m RNA in the testis tissue of cattle.
Manufaction of paraffin section on the testis of cattle,yak and cattle-yak and then stained with HE. The results shows that the seminiferous tubule of cattle-yak is much smaller than that of cattle and yak, the space between tubules is big, the size and shape of lumen is different, the tube wall is wizened, and there is no or only a thin flat of spermatogonia in epithelium of seminiferous tubule. Little primary spermatocyte can be seen, and no secondary spermatocyte, no sperm in epididymis; however, the seminiferous tubule tissue of cattle and yak testis develop normally, and there are all levels of spermatogenic. The sperm, which can be observed in epididymis, develop well. In situ hybridization demonstrated that the BOULE gene mainly distribution and expression on the primary spermatocyte and secondary spermatocyte.
The failure of spermatogenesis and spermatogenic cell with BOULE gene may leads to cattle-yak infertility.
引文
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