玉米C型胞质雄性不育恢复相关基因的克隆
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摘要
细胞质雄性不育的利用是实现杂种优势的一条重要途径。玉米(Zea mays L.)是世界上最早利用雄性不育进行杂交种子生产的作物,在降低生产成本、提高种子纯度等方面起到了积极的作用。目前在玉米种子生产上主要利用C型胞质雄性不育,然而玉米C型胞质雄性不育的恢复机理比较复杂,其恢复机理目前还不十分清楚,在一定程度上制约了其在生产上的利用。本研究利用转座子标签法对玉米C型胞质雄性不育的育性恢复相关基因进行克隆,并对候选基因的功能进行初步分析,取得了以下主要结论:
利用Mu转座子插入突变的方法,从29750株Cms-C87-1(Rf4Rf4)×Mu7的F1群体中筛选到5株雄性不育的插入突变体Rf4-m,并对其进行了不育细胞质鉴定,证明该突变是由核基因引起的。利用改良的AFLP方法对Cms-C87-1(Rf4Rf4)、Mu7和突变体Rf4-m进行多态性分析。在引物组合M1/BglⅡ-2的选择性扩增结果中,发现了一个750bp不同于两亲本的特异扩增片段。把特异的扩增片段连接转化测序,获得的序列中包含Mu转座子的部分序列和一段未知序列,推测可能是候选基因的部分序列。根据候选基因的侧翼序列设计引物,利用3’RACE快速扩增技术,以玉米雄穗的小花分化期的cDNA为模板,获得了一个209bp的cDNA序列。
利用生物信息学的方法对候选基因序列进行延伸,获得了一个全长为955bp的cDNA序列,基因组序列分析结果表明其DNA全长2625bp,包含3个内含子和4个外显子,其中有一个321bp的开放阅读框,编码106个氨基酸,功能结构域分析表明该基因是一个基本转录因子TFIIA。该基因与水稻、拟南芥等生物体的TFIIA基因有很高的相似性,同时与水稻的TFIIA基因有很近的同源关系。
TFIIA转录因子是一个核内蛋白,它参与了依赖于RNA聚合酶Ⅱ的DNA的转录。TFIIA是基本转录因子之一,这些基本转录因子是所有依赖于RNA聚合酶Ⅱ的转录所必需的。已有研究表明,动物中的TFIIA影响环磷酸腺苷应答元件调节器(CREM),而CREM基因编码的蛋白在雄性生殖细胞的发育过程中起着重要作用。TFIIA通过对关键基因CREM的调控,间接的调控精雄性生殖细胞的成熟发育。
The use of cytoplasmic male sterility (CMS) is an important approach in the utilization of heterosis. Maize (Zea mays L.) is the first crop that uses cytoplasmic male sterility for hybrid seed production. Since it can reduce the cost and improve the purity of hybrid seed, so the cytoplasmic male sterility (CMS) has become a main type in seed production. However, the genetic of fertility restoration in C-cytoplasmic male sterility is very complicated, and its restoration mechanism is not clear, then it becomes a main restrictive factor in using cytoplasmic male sterility in hybrid seed production. In this study, a related gene of fertility restoration for the CMS-C in maize was cloned by means of transposon tagging method, and its function of the candidate gene was preliminary analyzed. The main results showed in the hereinafter paragraphs.
Five mutants of the related fertility restoration were selected in a 29,750 F1 progeny plants between 87-1(Rf4Rf4) and Mu7, and the cytoplasmic type was confirmed by means of three special primers of different cytoplasmic types in maize. The special amplification band was selected using an improvement approach named Mu-AFLP method between two parents and the mutants, an about 750bp AFLP band was found when using a nested Mu7-specific primer BglⅡ-2 and a selective primer M1.A Mu7-flanking fragment was obtained after sequenced, which contained partial sequences of Mu transposon and an unknown sequence.
According to the sequence of the candidate gene, the PCR primers were designed and a 209bp cDNA was obtained from the cDNA coming from the floret differentiation stage of tassel, by mean of 3’rapid amplification of cDNA end (RACE) method.
Using the results of 3’RACE and electronic splicing in Genbank,a 955bp cDNA sequence was obtained; it contained a 321bp open reading frame which encoded 106 amino acid. The full length of the candidate gene was consisted of 2403 bp DNA sequence, contained three introns and four extrons. The results of functional domain analysis showed that the protein was a basic transcription factor TFIIA. It has homologous with the TFIIA gene of Oryza sativa and Arabidopsis thalian, and it has close relationships with the Oryza sativa. Transcription factor TFIIA was a nuclear protein involving in the RNA polymerase II-dependent DNAtranscription, and it was one of several basic transcription factors that were required for all transcription events using RNA polymerase II.
The previous reports have indicated that the CREM gene could be affected by the TFIIA in animals, which took an important place in the development of male germinal cells, and TFIIA could indirect regulate the development of maturity at different stages of spermiogenesis through regulating the key gene CREM. So we can suppose that the candidate gene, TFIIA, perhaps has some function on pollen development in maize.
利用Mu转座子插入突变的方法,从29750株Cms-C87-1(Rf4Rf4)×Mu7的F1群体中筛选到5株雄性不育的插入突变体Rf4-m,并对其进行了不育细胞质鉴定,证明该突变是由核基因引起的。利用改良的AFLP方法对Cms-C87-1(Rf4Rf4)、Mu7和突变体Rf4-m进行多态性分析。在引物组合M1/BglⅡ-2的选择性扩增结果中,发现了一个750bp不同于两亲本的特异扩增片段。把特异的扩增片段连接转化测序,获得的序列中包含Mu转座子的部分序列和一段未知序列,推测可能是候选基因的部分序列。根据候选基因的侧翼序列设计引物,利用3’RACE快速扩增技术,以玉米雄穗的小花分化期的cDNA为模板,获得了一个209bp的cDNA序列。
利用生物信息学的方法对候选基因序列进行延伸,获得了一个全长为955bp的cDNA序列,基因组序列分析结果表明其DNA全长2625bp,包含3个内含子和4个外显子,其中有一个321bp的开放阅读框,编码106个氨基酸,功能结构域分析表明该基因是一个基本转录因子TFIIA。该基因与水稻、拟南芥等生物体的TFIIA基因有很高的相似性,同时与水稻的TFIIA基因有很近的同源关系。
TFIIA转录因子是一个核内蛋白,它参与了依赖于RNA聚合酶Ⅱ的DNA的转录。TFIIA是基本转录因子之一,这些基本转录因子是所有依赖于RNA聚合酶Ⅱ的转录所必需的。已有研究表明,动物中的TFIIA影响环磷酸腺苷应答元件调节器(CREM),而CREM基因编码的蛋白在雄性生殖细胞的发育过程中起着重要作用。TFIIA通过对关键基因CREM的调控,间接的调控精雄性生殖细胞的成熟发育。
The use of cytoplasmic male sterility (CMS) is an important approach in the utilization of heterosis. Maize (Zea mays L.) is the first crop that uses cytoplasmic male sterility for hybrid seed production. Since it can reduce the cost and improve the purity of hybrid seed, so the cytoplasmic male sterility (CMS) has become a main type in seed production. However, the genetic of fertility restoration in C-cytoplasmic male sterility is very complicated, and its restoration mechanism is not clear, then it becomes a main restrictive factor in using cytoplasmic male sterility in hybrid seed production. In this study, a related gene of fertility restoration for the CMS-C in maize was cloned by means of transposon tagging method, and its function of the candidate gene was preliminary analyzed. The main results showed in the hereinafter paragraphs.
Five mutants of the related fertility restoration were selected in a 29,750 F1 progeny plants between 87-1(Rf4Rf4) and Mu7, and the cytoplasmic type was confirmed by means of three special primers of different cytoplasmic types in maize. The special amplification band was selected using an improvement approach named Mu-AFLP method between two parents and the mutants, an about 750bp AFLP band was found when using a nested Mu7-specific primer BglⅡ-2 and a selective primer M1.A Mu7-flanking fragment was obtained after sequenced, which contained partial sequences of Mu transposon and an unknown sequence.
According to the sequence of the candidate gene, the PCR primers were designed and a 209bp cDNA was obtained from the cDNA coming from the floret differentiation stage of tassel, by mean of 3’rapid amplification of cDNA end (RACE) method.
Using the results of 3’RACE and electronic splicing in Genbank,a 955bp cDNA sequence was obtained; it contained a 321bp open reading frame which encoded 106 amino acid. The full length of the candidate gene was consisted of 2403 bp DNA sequence, contained three introns and four extrons. The results of functional domain analysis showed that the protein was a basic transcription factor TFIIA. It has homologous with the TFIIA gene of Oryza sativa and Arabidopsis thalian, and it has close relationships with the Oryza sativa. Transcription factor TFIIA was a nuclear protein involving in the RNA polymerase II-dependent DNAtranscription, and it was one of several basic transcription factors that were required for all transcription events using RNA polymerase II.
The previous reports have indicated that the CREM gene could be affected by the TFIIA in animals, which took an important place in the development of male germinal cells, and TFIIA could indirect regulate the development of maturity at different stages of spermiogenesis through regulating the key gene CREM. So we can suppose that the candidate gene, TFIIA, perhaps has some function on pollen development in maize.
引文
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