甘蓝型油菜细胞质雄性不育系301A的遗传与分子生物学研究
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摘要
作物杂种优势利用，大大提高了油菜的产量，带来了相当可观的经济效益。细胞质雄性不育的利用，既可以减少人工去雄工作量，又有利于杂交种纯度的提高。目前生产中常用CMS的有：ogu CMS，Nap CMS，Polima CMS以及陕2A CMS。而且我国油菜细胞质雄性不育杂交种主要是以polima、陕2A CMS配制为主，且这两个不育系属于低温敏感型，持续低温易形成微粉，导致不育系自交产生不育的假杂种，这是影响杂交种纯度的主要原因。其次，人工去杂不彻底，隔离区不严而影响杂交种纯度，给生产造成很大的损失，因此，杂交种纯度鉴定越来越受重视。同时，长期利用同一胞质易造成细胞质单一化，因此，油菜杂种优势利用的当务之急是发掘和鉴定新的不育胞质类型。本实验通过形态学和细胞学观察、恢保关系鉴定、分子生物学等方法对新发现的细胞质雄性不育系301A进行研究，以期明确其细胞质雄性不育类型；同时对细胞质三系杂交种（合油杂2号）的纯度进行SSR鉴定体系的研究，为后续利用新细胞质进行三系品种选育及纯度鉴定提供研究基础。主要研究结果如下：
1.301A的花瓣皱缩，较宽大，雄蕊退化彻底，雌蕊及蜜腺发育正常；301A花器形态与Pol CMS、ogu CMS、Nap CMS有明显的差异，主要表现在：301A的长花丝和二弱雄蕊花药均为均最长。
2.以212A为对照，用30份甘蓝型油菜作测验种，分别与301A和212A杂交，测交结果表明：301A与212A的恢保关系完全相同。以301A保持系、Pol CMS、ogu CMS、NapCMS为对照，细胞学观察显示301A与Pol CMS、Nap CMS一样败育于孢原细胞时期，属于无花粉囊型，与ogu CMS（单核败育型）不同。
3.依据已报道的甘蓝型油菜细胞质雄性不育相关基因orf138、orf222、orf224设计引物，对301A、212A、ogu CMS、Nap CMS以及Cam的线粒体DNA做特异引物的PCR扩增，结果表明301A的线粒体基因组可能除了含有orf224基因的同源序列，还可能含有orf222基因的同源序列。
4.通过对细胞质三系杂交种（合油杂2号）进行SSR引物（http://ukcrop.net）筛选，一对引物（CB10524）分别在杂交种和其双亲之间存在扩增条带的多态性，表现为：杂交种条带均为父母本的互补型，很适合做杂交种纯度鉴定，为后续利用301A进行三系品种选育及纯度鉴定提供研究基础。
Crop heterosis utilization greatly improves the yield of rape, and brights considerablebenefits. The utilization of cytoplasmic male sterility not only economizes on man-power, butalso improves the hybrid purity. At present, Ogu CMS, Nap CMS, Polima CMS and Shan2ACMS are usually used in hybrid production. And in China Polima CMS and Shan2A CMSare mainly used to produce oilseed rape hybrids. However, the two sterile lines are bothsensitive to low temperature and easy to produce micro-podwer, which is the main reasonresulting in low hybrid purity. The incomplete remove of miscellaneous plants and the openisolation region also influence the purity and cause enormous losses. Therefore theidentification of hybrids purity is becoming more and more popular. Meanwhile the use of asingle cytoplasm for a long time hinders the development of rape hybrids. So exploiting andestablishing new CMS lines are of importance to oilseed rape hybrids production.301A is anew cytoplasmic male sterile line. In this study morphological observation, restoring andmaintaining relationship, cytological observation and molecular identification wereconducted to identify its cytoplasmic male sterility type. In addition, the purity of Brassicanapus cultivar Heyouza No.2was identified by SSR marker, which laid the foundation fordevelopment and purity identification of new cytoplasm male sterility three-line hybrid seeds.The main results were described as follows:
1. Petals of301A were shrunken and large surface.The stamen degeneratescompletely,the pistil and nectary growed well.In Seedling morphology,301A was differentfrom Pol CMS、ogu CMS、Nap CMS obviously，Length of the long filament and the twoweaker of301A are longest.
2.30species of Brassica napus L.were crossed with301A and212A,respectively.Thetest-cross results showed that301A was the same as212A in restoring and maintainingrelationship,both belonging to Pol CMS. With301A maintainer line、Pol CMS、ogu CMS andNap CMS as control samples, the result of cytological observation on anther showed that301A,like as Pol CMS and Nap CMS,was inhibited at the stage of archesporial celldifferentiation and different from the anther abortion period of ogu CMS started from thestage of tetrad to mononuclear.
3. Primers were designed according to the reported fertility-related geneorf138,orf222,orf224,.The mtDNA of4different CMS lines (including301A,212A, theimproved ogu CMS,nap CMS of Brassica napus L. and Cam were proceed PCRamplification with the distinctive primers.The result showed that mtDNA of301A maycontained both the homologous sequence of orf224and orf222.
4.The parental inbred lines of Brassica napus cultivar Heyouza No.2were used to studythe polymorphorism by SSR method.One pair of primer(CB10524) was found to bepolymorphic in the hybrid seeds and their parents and the polymorphism presented in theform that all the bands amplified from the hybrid seeds were complementary to those fromtheir parents, which was suitable for identification of seed purity and provided researchfoundation for using301A to breed new hybrid cultivars.
1.301A的花瓣皱缩，较宽大，雄蕊退化彻底，雌蕊及蜜腺发育正常；301A花器形态与Pol CMS、ogu CMS、Nap CMS有明显的差异，主要表现在：301A的长花丝和二弱雄蕊花药均为均最长。
2.以212A为对照，用30份甘蓝型油菜作测验种，分别与301A和212A杂交，测交结果表明：301A与212A的恢保关系完全相同。以301A保持系、Pol CMS、ogu CMS、NapCMS为对照，细胞学观察显示301A与Pol CMS、Nap CMS一样败育于孢原细胞时期，属于无花粉囊型，与ogu CMS（单核败育型）不同。
3.依据已报道的甘蓝型油菜细胞质雄性不育相关基因orf138、orf222、orf224设计引物，对301A、212A、ogu CMS、Nap CMS以及Cam的线粒体DNA做特异引物的PCR扩增，结果表明301A的线粒体基因组可能除了含有orf224基因的同源序列，还可能含有orf222基因的同源序列。
4.通过对细胞质三系杂交种（合油杂2号）进行SSR引物（http://ukcrop.net）筛选，一对引物（CB10524）分别在杂交种和其双亲之间存在扩增条带的多态性，表现为：杂交种条带均为父母本的互补型，很适合做杂交种纯度鉴定，为后续利用301A进行三系品种选育及纯度鉴定提供研究基础。
Crop heterosis utilization greatly improves the yield of rape, and brights considerablebenefits. The utilization of cytoplasmic male sterility not only economizes on man-power, butalso improves the hybrid purity. At present, Ogu CMS, Nap CMS, Polima CMS and Shan2ACMS are usually used in hybrid production. And in China Polima CMS and Shan2A CMSare mainly used to produce oilseed rape hybrids. However, the two sterile lines are bothsensitive to low temperature and easy to produce micro-podwer, which is the main reasonresulting in low hybrid purity. The incomplete remove of miscellaneous plants and the openisolation region also influence the purity and cause enormous losses. Therefore theidentification of hybrids purity is becoming more and more popular. Meanwhile the use of asingle cytoplasm for a long time hinders the development of rape hybrids. So exploiting andestablishing new CMS lines are of importance to oilseed rape hybrids production.301A is anew cytoplasmic male sterile line. In this study morphological observation, restoring andmaintaining relationship, cytological observation and molecular identification wereconducted to identify its cytoplasmic male sterility type. In addition, the purity of Brassicanapus cultivar Heyouza No.2was identified by SSR marker, which laid the foundation fordevelopment and purity identification of new cytoplasm male sterility three-line hybrid seeds.The main results were described as follows:
1. Petals of301A were shrunken and large surface.The stamen degeneratescompletely,the pistil and nectary growed well.In Seedling morphology,301A was differentfrom Pol CMS、ogu CMS、Nap CMS obviously，Length of the long filament and the twoweaker of301A are longest.
2.30species of Brassica napus L.were crossed with301A and212A,respectively.Thetest-cross results showed that301A was the same as212A in restoring and maintainingrelationship,both belonging to Pol CMS. With301A maintainer line、Pol CMS、ogu CMS andNap CMS as control samples, the result of cytological observation on anther showed that301A,like as Pol CMS and Nap CMS,was inhibited at the stage of archesporial celldifferentiation and different from the anther abortion period of ogu CMS started from thestage of tetrad to mononuclear.
3. Primers were designed according to the reported fertility-related geneorf138,orf222,orf224,.The mtDNA of4different CMS lines (including301A,212A, theimproved ogu CMS,nap CMS of Brassica napus L. and Cam were proceed PCRamplification with the distinctive primers.The result showed that mtDNA of301A maycontained both the homologous sequence of orf224and orf222.
4.The parental inbred lines of Brassica napus cultivar Heyouza No.2were used to studythe polymorphorism by SSR method.One pair of primer(CB10524) was found to bepolymorphic in the hybrid seeds and their parents and the polymorphism presented in theform that all the bands amplified from the hybrid seeds were complementary to those fromtheir parents, which was suitable for identification of seed purity and provided researchfoundation for using301A to breed new hybrid cultivars.
引文
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