玉米C型细胞质雄性不育的分子生物学研究
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摘要
细胞质雄性不育(CMS,cytoplasmic male sterility)是雄性不育中最重要的一种类型,在高等植物中普遍存在。细胞质雄性不育容易实现不育系、保持系和恢复系的配套,是当前玉米育种中利用的主要类型。玉米细胞质雄性不育的研究和利用已成为玉米育种学的重要内容。依据核基因对其育性恢复专效性的不同,玉米CMS可分为3大类型,即T-CMS,C-CMS,S-CMS。T型不育系育性高度稳定,但是对玉米小斑病T小种表现高度的专化感染;C型和S型不育系对玉米小斑病T小种不具有专化感染,但是S型不育系属于配子体不育类型,对环境较敏感,育性稳定性较差。所以,C型不育系的利用对于玉米育种学具有重要的意义。然而,由于C型不育系发现较迟,研究利用较晚,与T型和S型相比,其不育机制研究的还不够深入。
对2套同核异质不育系及保持系,包括48-2核背景(N48-2、C48-2、T48-2和S48-2)和黄早四核背景(N黄早四、C黄早四、T黄早四和S黄早四)的线粒体DNA进行了AFLP分析比较。从选择性扩增产物中筛选到了一条C48-2所特有的差异条带C48-223。对C48-223进行克隆、测序分析表明,它是玉米C型细胞质雄性不育线粒体atp6基因编码序列的一部分。Southern杂交分析证实了C48-223在C48-2线粒体基因组上的特异性。Northern结果显示,该片段在玉米苗期、四分体时期和单核期的表达量呈递增趋势。这说明该差异片段和玉米C型细胞质雄性不育有很高的相关性。从选择性扩增产物中筛选到了一条C黄早四所特有的差异条带C黄早四-1。对C黄早四-1进行克隆、测序分析表明,它与玉米C型细胞质雄性不育线粒体NADH基因编码序列的一部分高度同源。本文的研究结果表明,在单一不育系中出现的特异条带,有可能是核基因改变线粒体基因转录起始位点、转录物的加工位点或编辑位点的结果,而这种改变随核背景的不同而变化。这些新的特异条带的出现是否与细胞质雄性不育机制相关还有待于进一步的研究。
经过优化,确定适合玉米线粒体蛋白质使用的双向电泳体系为:采用含有7.0 mol L~(-1)尿素、4%CHAPS、2.0 mol L~(-1)硫脲、9.0 mmolL~(-1)DTT、0.8%两性电解质的裂解液提取线粒体蛋白质、12%的分离胶、90000 Vh的总聚焦时间、500μg的上样量和pH3-10的非线性胶条。采用固相pH梯度-SDS PAGE双向电泳对玉米C型细胞质雄性不育系(C48-2)及其保持系(N48-2)苗期叶片线粒体蛋白质、单核期和双核期花药线粒体蛋白质进行了分离,经过考马斯亮蓝染色,得到了重复性较好的双向电泳图谱。PDQUEST软件分别在苗期、单核期和双核期的双向电泳图谱上可识别约280、260、220个蛋白质点,将其中23个重复性比较好且差异表达在两倍以上的蛋白质点采用基质辅助激光解析分离飞行时间质谱进行了肽指纹图谱分析,然后用Mascot软件对NCBI数据库搜索,其中7个蛋白质点得到了鉴定。这7个蛋白质分别是F_1-ATP合成酶、ATP合成酶、磷脂酰肌醇-3激酶、蛋白质二硫键异构酶、谷氨酸脱氢酶和电压依赖阴离子通道蛋白(2个点)。它们分别参与ATP的合成、信号转导、新生肽段的折叠、氮代谢和线粒体外膜的通透性等。
总之,本文利用AFLP和双向电泳技术分别对玉米线粒体DNA和线粒体蛋白质进行了分析,为阐明玉米C型细胞质雄性不育的分子机制奠定了基础。
Cytoplasmic male sterility (CMS) is the most important type of male sterility in maize (Zea mays L.), which occurs in many plant species. The investigation and utilization of CMS have been the mainly content in maize breeding. Cytoplasmic. male sterility in maize (Zea mays L.) is categorized into T-CMS, C-CMS, and S-CMS types based on the specific restoration of nuclear gene, in which C-CMS has a great application potential in maize hybrid breeding due to its stable sterility and resistance to Helminthosporium maydis Nisik. The sterile mechanism of C-CMS is not fully understood for its later discovery and utilization than these of T-CMS and S-CMS.
Mitochondrial DNA extracted from two set of isonuclear alloplasm materials were analysed and compared each other with AFLP technique. One differential DNA segment of C48-2—named C48-223—was found when the mtDNA fragments were amplified with selective primers. The band was isolated , purified and sequenced. Based on the blasting result, the C48-223 is a part of atp6-C, which is a chimeric gene that uniquely associated with the C male-sterile cytoplasm of maize. The results of Southern blot indicate C48-223 is a real band. Via Northern blot, the signals can only be detected at the tetrad stage and uninucleate stage of anther RNA extracted from C48-2. Another differential DNA segment of C-huangzaosi—named C-huangzaosi1—was found when the mtDNA fragments were amplified with selective primers. Based on the blasting result, the C-huangzaosi1 is a part of NADH-C dehydrogenase subunit 1, which is uniquely associated with the C male-sterile cytoplasm of maize. All these indicate that the two fragments probably have strong relationship with the C male-sterile cytoplasm of maize.
After optimized the 2-D system, we choosed the 2-D system for maize mitochondrial proteins: extract reagent contain 7.0 mol L~(-1) Urea, 4%CHAPS, 2.0 mol L~(-1) thiourea, 9.0mmolL~(-1)DTT, 0.8% ampholyte; 12% polyacrylamide gel; total 90000Vh isoelectric focusing; 500μg of protein extract per isoelectric focusing unit; 17cm immobilized non-linear pH gtadients(pH3-10). The mitochondrial proteins of maize yellow seedling, uninucleate stage anther , and binucleate phase anther of C48-2 and N48-2 (maintenance line) were separated by two-dimensional electrophoresis with immobilized pH (3-10, non-linear) gradients as the first dimension and SDS-PAGE as the second. The Coomassie brilliant blue-stained stained proteins spots were analyzed using PDQUEST software, there were about 280, 260, 220detectable spots on 2D-gel of respective phase. With direct MALDI-TOF mass spectrometry analysis and protein database searching, 7 protein spots out of 23 were identified with significative data. They were F1-ATPsynthase, ATP synthase , Phosphatidylinositol 3-kinase , Glutamate dehydrogenase , voltage-dependent anion channel protein(two spots) and protein disulfide isomerase. F1-ATPsynthase and ATP synthase are relate with provide energy to cell. Glutamate dehydrogenase is present mainly in mitochondria in hight plants and play a important role in the course of nitrogen metabolism. VDAC plays an essential role not only in the permeability of the mitochondrial membrane, but also in apoptosis mediated by the mitochondrial pathway . Phosphatidylinositol-3 kinase (PI3K) is one of the most important regulatory proteins that is involved in different signaling pathways and controlling of key functions of the cell. The major role for protein disulfide isomerase is thought to be the formation and rearrangement of disulfide bonds.
In summary, AFLP and 2-D dimention were employed to compare the mtDNA and mitochondrial proteins between the C type cytoplasmic male sterility and Normal cytoplasm of maize. This study will facilitate the understanding of the molecular aspects of the cytoplasmic male sterility of maize.
对2套同核异质不育系及保持系,包括48-2核背景(N48-2、C48-2、T48-2和S48-2)和黄早四核背景(N黄早四、C黄早四、T黄早四和S黄早四)的线粒体DNA进行了AFLP分析比较。从选择性扩增产物中筛选到了一条C48-2所特有的差异条带C48-223。对C48-223进行克隆、测序分析表明,它是玉米C型细胞质雄性不育线粒体atp6基因编码序列的一部分。Southern杂交分析证实了C48-223在C48-2线粒体基因组上的特异性。Northern结果显示,该片段在玉米苗期、四分体时期和单核期的表达量呈递增趋势。这说明该差异片段和玉米C型细胞质雄性不育有很高的相关性。从选择性扩增产物中筛选到了一条C黄早四所特有的差异条带C黄早四-1。对C黄早四-1进行克隆、测序分析表明,它与玉米C型细胞质雄性不育线粒体NADH基因编码序列的一部分高度同源。本文的研究结果表明,在单一不育系中出现的特异条带,有可能是核基因改变线粒体基因转录起始位点、转录物的加工位点或编辑位点的结果,而这种改变随核背景的不同而变化。这些新的特异条带的出现是否与细胞质雄性不育机制相关还有待于进一步的研究。
经过优化,确定适合玉米线粒体蛋白质使用的双向电泳体系为:采用含有7.0 mol L~(-1)尿素、4%CHAPS、2.0 mol L~(-1)硫脲、9.0 mmolL~(-1)DTT、0.8%两性电解质的裂解液提取线粒体蛋白质、12%的分离胶、90000 Vh的总聚焦时间、500μg的上样量和pH3-10的非线性胶条。采用固相pH梯度-SDS PAGE双向电泳对玉米C型细胞质雄性不育系(C48-2)及其保持系(N48-2)苗期叶片线粒体蛋白质、单核期和双核期花药线粒体蛋白质进行了分离,经过考马斯亮蓝染色,得到了重复性较好的双向电泳图谱。PDQUEST软件分别在苗期、单核期和双核期的双向电泳图谱上可识别约280、260、220个蛋白质点,将其中23个重复性比较好且差异表达在两倍以上的蛋白质点采用基质辅助激光解析分离飞行时间质谱进行了肽指纹图谱分析,然后用Mascot软件对NCBI数据库搜索,其中7个蛋白质点得到了鉴定。这7个蛋白质分别是F_1-ATP合成酶、ATP合成酶、磷脂酰肌醇-3激酶、蛋白质二硫键异构酶、谷氨酸脱氢酶和电压依赖阴离子通道蛋白(2个点)。它们分别参与ATP的合成、信号转导、新生肽段的折叠、氮代谢和线粒体外膜的通透性等。
总之,本文利用AFLP和双向电泳技术分别对玉米线粒体DNA和线粒体蛋白质进行了分析,为阐明玉米C型细胞质雄性不育的分子机制奠定了基础。
Cytoplasmic male sterility (CMS) is the most important type of male sterility in maize (Zea mays L.), which occurs in many plant species. The investigation and utilization of CMS have been the mainly content in maize breeding. Cytoplasmic. male sterility in maize (Zea mays L.) is categorized into T-CMS, C-CMS, and S-CMS types based on the specific restoration of nuclear gene, in which C-CMS has a great application potential in maize hybrid breeding due to its stable sterility and resistance to Helminthosporium maydis Nisik. The sterile mechanism of C-CMS is not fully understood for its later discovery and utilization than these of T-CMS and S-CMS.
Mitochondrial DNA extracted from two set of isonuclear alloplasm materials were analysed and compared each other with AFLP technique. One differential DNA segment of C48-2—named C48-223—was found when the mtDNA fragments were amplified with selective primers. The band was isolated , purified and sequenced. Based on the blasting result, the C48-223 is a part of atp6-C, which is a chimeric gene that uniquely associated with the C male-sterile cytoplasm of maize. The results of Southern blot indicate C48-223 is a real band. Via Northern blot, the signals can only be detected at the tetrad stage and uninucleate stage of anther RNA extracted from C48-2. Another differential DNA segment of C-huangzaosi—named C-huangzaosi1—was found when the mtDNA fragments were amplified with selective primers. Based on the blasting result, the C-huangzaosi1 is a part of NADH-C dehydrogenase subunit 1, which is uniquely associated with the C male-sterile cytoplasm of maize. All these indicate that the two fragments probably have strong relationship with the C male-sterile cytoplasm of maize.
After optimized the 2-D system, we choosed the 2-D system for maize mitochondrial proteins: extract reagent contain 7.0 mol L~(-1) Urea, 4%CHAPS, 2.0 mol L~(-1) thiourea, 9.0mmolL~(-1)DTT, 0.8% ampholyte; 12% polyacrylamide gel; total 90000Vh isoelectric focusing; 500μg of protein extract per isoelectric focusing unit; 17cm immobilized non-linear pH gtadients(pH3-10). The mitochondrial proteins of maize yellow seedling, uninucleate stage anther , and binucleate phase anther of C48-2 and N48-2 (maintenance line) were separated by two-dimensional electrophoresis with immobilized pH (3-10, non-linear) gradients as the first dimension and SDS-PAGE as the second. The Coomassie brilliant blue-stained stained proteins spots were analyzed using PDQUEST software, there were about 280, 260, 220detectable spots on 2D-gel of respective phase. With direct MALDI-TOF mass spectrometry analysis and protein database searching, 7 protein spots out of 23 were identified with significative data. They were F1-ATPsynthase, ATP synthase , Phosphatidylinositol 3-kinase , Glutamate dehydrogenase , voltage-dependent anion channel protein(two spots) and protein disulfide isomerase. F1-ATPsynthase and ATP synthase are relate with provide energy to cell. Glutamate dehydrogenase is present mainly in mitochondria in hight plants and play a important role in the course of nitrogen metabolism. VDAC plays an essential role not only in the permeability of the mitochondrial membrane, but also in apoptosis mediated by the mitochondrial pathway . Phosphatidylinositol-3 kinase (PI3K) is one of the most important regulatory proteins that is involved in different signaling pathways and controlling of key functions of the cell. The major role for protein disulfide isomerase is thought to be the formation and rearrangement of disulfide bonds.
In summary, AFLP and 2-D dimention were employed to compare the mtDNA and mitochondrial proteins between the C type cytoplasmic male sterility and Normal cytoplasm of maize. This study will facilitate the understanding of the molecular aspects of the cytoplasmic male sterility of maize.
引文
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