芝麻隐性细胞核雄性不育基因的遗传定位和差异表达分析
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摘要
芝麻的杂种优势广泛存在,利用雄性不育生产杂交种是杂种优势利用的重要途径。我国已培育出多个隐性细胞核芝麻雄性不育系(RGMS),其育性稳定、无不良胞质效应、恢复源广,在生产F1种子中广泛应用,并已育成多个杂交品种。然而,直到目前为止,对芝麻细胞核雄性不育的遗传特性和败育机理了解较少,有必要对其进行深入的研究。本研究以芝麻隐性细胞核雄性不育两用系95ms-5AB为材料,利用形态学、细胞学及分子生物学等方法,对其开展组织形态、遗传定位和差异表达研究,主要结果和结论如下：
1.形态学研究发现,雄性不育系95ms-5可育株和不育株的植株形态无明显差异,但不育花药绿色、瘦瘪,花粉少而小,形态异常。可育花药长度和花柱长度都明显比不育花的长,而花冠长度和花丝长度无明显差异。与86ms-1转育得到的不育系相比,95ms-5A每朵花的平均花粉数量少,花粉偏小。95ms-5A和86ms-1转育不育系花粉离体培养均不萌发。
2.组织学观察结果说明95ms-5A雄配子的发育异常出现在单核小孢子期,其败育特征为四分体后小孢子形态异常,与降解不完全的胼胝质发生粘连及绒毡层降解推迟等。电镜观察可见不育花粉外壁凹陷,未见萌发沟和颗粒状突起。
3.95ms-5的测交和姊妹交后代遗传分析表明,95ms-5A的细胞核雄性不育受单一隐性基因Sims1(Sesamum indicum male sterility1)控制。与已报道的隐性细胞核雄性不育ms86-1的等位性分析表明,95ms-5A不育基因Sims1与ms86-1的不育基因不等位。
4.利用分离群体分组分析法(BSA)研究分析了SiMs1基因的扩增片断长度多态性(AFLP)连锁标记,并通过对近等基因系95ms-5A和95ms-5B姊妹交后代获得的237株定位群体扩增分析,构建了SiMs1基因的遗传连锁图,共有9个标记,分布于SiMs1基因的两侧。将SiMs1基因定位于8.0cM的遗传区间内,其两侧最近的标记P06MG04和P12EA14,距离目标基因分别为1.2cM和6.8cM。且标记P01MC08与SiMs1基因共分离。相关研究结果为芝麻SiMsl基因的分子标记辅助选择、图位克隆及杂优育种奠定了坚实基础。
5.利用cDNA-AFLP技术开展了95ms-5A不育花蕾和95ms-5B可育花蕾的转录谱分析,以阐明该隐性细胞核雄性不育的败育机理。共获得30条可重复的差异表达转录衍生片段(TDFs),其中在不育花蕾中呈下调表达的TDFs27条,上调表达的3条。成功克隆了27个TDFs,对其功能分析结果表明：11个TDFs的对应基因参与了能量代谢、信号传导和植物细胞壁的形成,其余16个TDFs与GenBank中未知功能蛋白同源或未匹配到同源序列。
6.对21个基因进行了花蕾不同发育时期的实时荧光定量PCR分析,它们的表达模式与cDNA-AFLP分析结果完全一致。上述差异基因不同发育阶段的表达模式和基因功能注释分析表明,95ms-5A中一些胼胝质壁或绒毡层降解相关基因,如胼胝质合成酶催化类亚基蛋白、类受体蛋白激酶等,在小孢子母细胞或四分体之前某个阶段表达受到抑制,这可能是导致绒毡层降解延迟和雄配子发育受阻的直接原因。一些功能未知的基因在可育与不育花蕾的某个时间段也表现出极显著的表达水平差异,也很有可能是导致雄性不育的重要原因。通过半定量Real-time PCR对其中8个差异表达基因的组织特异性表达分析,鉴定筛选到一个雄蕊特异表达新基因,该基因在不育花蕾和可育花蕾间表达差异极显著,且功能未知。
The hybrid vigor is widely existed in sesame (Sesamum indicum L.). One of the important ways for heterosis utilization is to use the genic male sterile lines for producing hybrid seeds. In China, several sesame recessive genic male sterile lines (RGMS) have been bred, which are characterized by male sterile stability, no negative cytoplasmic effects and broad restoring resources. Many hybrid varieties have been bred by producing F1seeds using RGMS lines. However, we have little knowledge about the genetic characterization and mechanism of sesame male sterility. Therefore, it is necessary to conduct further studies on these aspects. In this study,95ms-5AB, a two-type line, was used for studies on histology, mapping and differential expression by phenotypic, cytological and molecular biology methods. The main results and conclusions are as follows:
1. The phenotypic observations showed that95ms-5A plants did not show any other obvious differences from the male fertile95ms-5B plants, except for having greenish, shriveled and slim anthers with few, small and degenerative pollens. The lengths of anther and style in fertile flowers are larger than those of sterile flowers, but no difference in corolla and filament length between fertile and sterile flowers. Compared with the RGMS lines transferring from86ms-1,95ms-5A had less mean pollens and smaller pollen size. The pollen grains of95ms-5A and86ms-1lines could not germinate in vitro.
2. Histological observations revealed that abnormal inhibited male gametogenesis of95ms-5A appeared during the mononuclear microspore stage. It was characterized by collapsed microspores, adhesion with callose which did not fully degraded and the delayed degeneration of tapetum after the tetrad stage. The morphology of pollens was observed by scanning electron microscope. The sterile pollen wall was cupped and no germinal furrows and particles were observed on pollen exine.
3. Genetic analysis of test cross and sib-mating of95ms-5indicated that the male sterility of95ms-5A was controlled by a single recessive genic male sterile gene Simsl {Sesamum indicum male sterility1). Allelic test with ms86-1confirmed that Simsl in95ms-5A is different from ms86-1.
4. Amplified fragment length polymorphism (AFLP) markers linked to SiMsl were screened using bulked segregant analysis (BSA). A genetic linkage map of the SiMs1 gene was constructed using237plants derived from the sib-mating between the near-isogenic lines95ms-5A and95ms-5B. The SiMsl gene was found to be located in a region of8.0cM, with a distance of1.2cM from P06MG04and6.8cM from P12EA14, the closest markers on two sides. And marker P01MC08was identified to be co-segregated with SiMsl. The linkage map constructed in this study will be very useful for marker-assisted selection, and map-based cloning of SiMsl as well as for hybrid breeding in sesame.
5. cDNA-AFLP technique was used to investigate the transcriptional profiles of the sterile flower buds in95ms-5A and the fertile flower buds in95ms-5B in order to elucidate the mechanism of this RGMS. A total of30reproducible differentially expressed transcript-derived fragments (TDFs) were detected, including27down-regulated and three up-regulated TDFs in sterile buds. The functional annotation analysis of27successfully cloned TDFs indicated that the corresponding genes of11TDFs may participate in the processes of energy metabolism, signal transduction and plant cell development. The other16TDFs were homologous to the proteins with unknown functions or with no homo logy in GenBank.
6. Real-Time PCR analysis of21genes in buds at different stages was performed and their expression patterns were entirely the same with those identified by cDNA-AFLP. The expression patterns and functional annotation of above genes in different stages suggested that some genes, such as callose synthase catalytic subunit-like protein (TDF04), receptor-like protein kinase (TDF03) etc., related to the callose wall or tapetum degeneration, were suppressed in95ms-5A in a certain phase before the microspore mother cell or tetrad stage, which may lead to the delayed tapetum degeneration and blocked male gametogenesis. Some function unknown genes showed significant difference of expression level in certain stages between fertile and sterile buds, which may also be the reasons for male sterility. Semi-quantification Real-Time PCR was used to carry on tissue-special expressed analysis of8differential expression genes. One novel stamen-special expressed gene was identified, which showed significantly different expression level between fertile and sterile buds, and its function is unknown.
1.形态学研究发现,雄性不育系95ms-5可育株和不育株的植株形态无明显差异,但不育花药绿色、瘦瘪,花粉少而小,形态异常。可育花药长度和花柱长度都明显比不育花的长,而花冠长度和花丝长度无明显差异。与86ms-1转育得到的不育系相比,95ms-5A每朵花的平均花粉数量少,花粉偏小。95ms-5A和86ms-1转育不育系花粉离体培养均不萌发。
2.组织学观察结果说明95ms-5A雄配子的发育异常出现在单核小孢子期,其败育特征为四分体后小孢子形态异常,与降解不完全的胼胝质发生粘连及绒毡层降解推迟等。电镜观察可见不育花粉外壁凹陷,未见萌发沟和颗粒状突起。
3.95ms-5的测交和姊妹交后代遗传分析表明,95ms-5A的细胞核雄性不育受单一隐性基因Sims1(Sesamum indicum male sterility1)控制。与已报道的隐性细胞核雄性不育ms86-1的等位性分析表明,95ms-5A不育基因Sims1与ms86-1的不育基因不等位。
4.利用分离群体分组分析法(BSA)研究分析了SiMs1基因的扩增片断长度多态性(AFLP)连锁标记,并通过对近等基因系95ms-5A和95ms-5B姊妹交后代获得的237株定位群体扩增分析,构建了SiMs1基因的遗传连锁图,共有9个标记,分布于SiMs1基因的两侧。将SiMs1基因定位于8.0cM的遗传区间内,其两侧最近的标记P06MG04和P12EA14,距离目标基因分别为1.2cM和6.8cM。且标记P01MC08与SiMs1基因共分离。相关研究结果为芝麻SiMsl基因的分子标记辅助选择、图位克隆及杂优育种奠定了坚实基础。
5.利用cDNA-AFLP技术开展了95ms-5A不育花蕾和95ms-5B可育花蕾的转录谱分析,以阐明该隐性细胞核雄性不育的败育机理。共获得30条可重复的差异表达转录衍生片段(TDFs),其中在不育花蕾中呈下调表达的TDFs27条,上调表达的3条。成功克隆了27个TDFs,对其功能分析结果表明：11个TDFs的对应基因参与了能量代谢、信号传导和植物细胞壁的形成,其余16个TDFs与GenBank中未知功能蛋白同源或未匹配到同源序列。
6.对21个基因进行了花蕾不同发育时期的实时荧光定量PCR分析,它们的表达模式与cDNA-AFLP分析结果完全一致。上述差异基因不同发育阶段的表达模式和基因功能注释分析表明,95ms-5A中一些胼胝质壁或绒毡层降解相关基因,如胼胝质合成酶催化类亚基蛋白、类受体蛋白激酶等,在小孢子母细胞或四分体之前某个阶段表达受到抑制,这可能是导致绒毡层降解延迟和雄配子发育受阻的直接原因。一些功能未知的基因在可育与不育花蕾的某个时间段也表现出极显著的表达水平差异,也很有可能是导致雄性不育的重要原因。通过半定量Real-time PCR对其中8个差异表达基因的组织特异性表达分析,鉴定筛选到一个雄蕊特异表达新基因,该基因在不育花蕾和可育花蕾间表达差异极显著,且功能未知。
The hybrid vigor is widely existed in sesame (Sesamum indicum L.). One of the important ways for heterosis utilization is to use the genic male sterile lines for producing hybrid seeds. In China, several sesame recessive genic male sterile lines (RGMS) have been bred, which are characterized by male sterile stability, no negative cytoplasmic effects and broad restoring resources. Many hybrid varieties have been bred by producing F1seeds using RGMS lines. However, we have little knowledge about the genetic characterization and mechanism of sesame male sterility. Therefore, it is necessary to conduct further studies on these aspects. In this study,95ms-5AB, a two-type line, was used for studies on histology, mapping and differential expression by phenotypic, cytological and molecular biology methods. The main results and conclusions are as follows:
1. The phenotypic observations showed that95ms-5A plants did not show any other obvious differences from the male fertile95ms-5B plants, except for having greenish, shriveled and slim anthers with few, small and degenerative pollens. The lengths of anther and style in fertile flowers are larger than those of sterile flowers, but no difference in corolla and filament length between fertile and sterile flowers. Compared with the RGMS lines transferring from86ms-1,95ms-5A had less mean pollens and smaller pollen size. The pollen grains of95ms-5A and86ms-1lines could not germinate in vitro.
2. Histological observations revealed that abnormal inhibited male gametogenesis of95ms-5A appeared during the mononuclear microspore stage. It was characterized by collapsed microspores, adhesion with callose which did not fully degraded and the delayed degeneration of tapetum after the tetrad stage. The morphology of pollens was observed by scanning electron microscope. The sterile pollen wall was cupped and no germinal furrows and particles were observed on pollen exine.
3. Genetic analysis of test cross and sib-mating of95ms-5indicated that the male sterility of95ms-5A was controlled by a single recessive genic male sterile gene Simsl {Sesamum indicum male sterility1). Allelic test with ms86-1confirmed that Simsl in95ms-5A is different from ms86-1.
4. Amplified fragment length polymorphism (AFLP) markers linked to SiMsl were screened using bulked segregant analysis (BSA). A genetic linkage map of the SiMs1 gene was constructed using237plants derived from the sib-mating between the near-isogenic lines95ms-5A and95ms-5B. The SiMsl gene was found to be located in a region of8.0cM, with a distance of1.2cM from P06MG04and6.8cM from P12EA14, the closest markers on two sides. And marker P01MC08was identified to be co-segregated with SiMsl. The linkage map constructed in this study will be very useful for marker-assisted selection, and map-based cloning of SiMsl as well as for hybrid breeding in sesame.
5. cDNA-AFLP technique was used to investigate the transcriptional profiles of the sterile flower buds in95ms-5A and the fertile flower buds in95ms-5B in order to elucidate the mechanism of this RGMS. A total of30reproducible differentially expressed transcript-derived fragments (TDFs) were detected, including27down-regulated and three up-regulated TDFs in sterile buds. The functional annotation analysis of27successfully cloned TDFs indicated that the corresponding genes of11TDFs may participate in the processes of energy metabolism, signal transduction and plant cell development. The other16TDFs were homologous to the proteins with unknown functions or with no homo logy in GenBank.
6. Real-Time PCR analysis of21genes in buds at different stages was performed and their expression patterns were entirely the same with those identified by cDNA-AFLP. The expression patterns and functional annotation of above genes in different stages suggested that some genes, such as callose synthase catalytic subunit-like protein (TDF04), receptor-like protein kinase (TDF03) etc., related to the callose wall or tapetum degeneration, were suppressed in95ms-5A in a certain phase before the microspore mother cell or tetrad stage, which may lead to the delayed tapetum degeneration and blocked male gametogenesis. Some function unknown genes showed significant difference of expression level in certain stages between fertile and sterile buds, which may also be the reasons for male sterility. Semi-quantification Real-Time PCR was used to carry on tissue-special expressed analysis of8differential expression genes. One novel stamen-special expressed gene was identified, which showed significantly different expression level between fertile and sterile buds, and its function is unknown.
引文
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