水稻小穗发育相关基因MFS1的图位克隆与功能分析
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摘要
水稻(Oryza sativa L.)是世界上重要的粮食作物,也是单子叶植物的模式植物。水稻是研究花发育的理想材料,与双子叶模式植物相比,水稻花序具有禾本科植物独有的结构单元一小穗,而且具有丰富的遗传与基因组资源。深入研究水稻生殖发育过程,不仅有助于深入理解水稻小穗或小花的形成机制,而且对丰富植物发育生物学知识和提高水稻产量都具有重要意义。
在本研究中,克隆了一个水稻小穗发育相关基因MULTI-FLIRET SPIKELET1(MFS1),我们对其进行了表型和细胞学观察,表达模式分析以及功能验证。主要结果如下：
1.mfs1突变体表型观察
从EMS诱变库中发现mfS1-1和mfs1-2两个等位小穗突变体,相比而言mfs1-1突变表型更严重和丰富,以下主要描述mfs1-1突变体。开花期的mfs1-1小穗护颖退化类似于副护颖,小穗轴伸长,形成额外的外稃,内稃主体部分退化,内三轮的花器官没有发现同源转化现象但花器官数目不确定。扫描电镜显示,早期护颖原基发育迟缓和退化,部分小花形成扩大的花分生组织和两个退化的内稃原基,雄蕊发育不同步。
2.MFS1遗传分析与分子定位
利用三个杂交组合进行遗传分析和分子定位,杂交F,代植株表型与野生型相似,F2代植株发生分离,正常株和突变株分离比均符合3：1,表明MFS1基因受单隐性基因控制。运用SSR标记和InDel标记进行基因定位,MFS1基因被最终定位在第5染色体InDel17和InDel24之间。
3.MFS1基因的克隆与蛋白质分析
DNA和cDNA测序分析发现一个编码APETALA2/EREBP结构域蛋白的基因Os05g41760分别在mfS1-1和mfs1-2突变体中存在一个C-T的替换,引起了氨基酸的改变,导致蛋白质的功能变异,初步确定其为候选基因。MFS1属于AP2/ERF家族中一个功能未知的分支,蛋白质定位在细胞核内。
4.MFS1基因的表达模式分析
利用半定量RT-PCR,qPCR和原位杂交技术进行了表达模式分析。半定量RT-PCR和qPCR结果显示MFS1基因在根,茎,叶和花中都表达,尤其在花序早期表达强烈。原位杂交显示MFS1在分生组织和护颖原基中表达强烈,随着花器官的发育,表达量逐渐减弱。
5.MFS1基因的功能验证
构建MFS1基因的基因组互补载体,转化mfs1-1突变体的愈伤组织,获得转基因植株,通过GUS染色检测转基因植株。结果表明,mfs1-1突变体表型完全恢复,确定了Os05g41760基因是MFS1基因。
构建MFS1基因的RNAi转基因表达载体,转化籼稻品种缙恢10号和粳稻品种中花11的愈伤组织,获得转基因植株,GUS染色、PCR检测转基因植株。RNAi转基因植株小穗表现为护颖退化,小穗轴伸长,额外的外稃状器官和内稃退化,与mfs1突变体的表型高度相似,进一步证明了OS05g41760基因是MFS1基因和MFS1基因的功能。
6.MFS1调控其它花发育相关基因的表达
qPCR分析和原位杂交分析结果表明,在mfs1-1突变体中,A功能基因OSMADs15表达量增加,AP2/ERF结构域基因SNB、OsIDS1和FZP表达量减少及护颖特征基因G1表达量减少。
Rice (Oryza sativa L.) is not only an important food crops in the world and a model plant in monocots but also an ideal object for studying flower development. Compaired with the model plants in dicots, rice bears a unique inflorescence consisting of spikelets to grass species, and has abundant genetic and genomic resources. So, reseach on rice reproductive development process would facilitate understanding of the molecular mechanism involved in spikelet development in rice and enrich knowledge of plant developmental biology for us. Meantime, it is very important to improve rice yields.
In the study, we cloned a MULTI-FLORET SPIKELET1(MFS1) gene associated with spikelet development in rice. We did investgation of mfsl mutants, and analysed the expression patterns and functions of MFS1gene. The results as follows:
1. Phenotypic observation of mfsl mutants
Two mutants of rice, mfsl-1and mfsl-2, were identified from ethylmethane sulfonate (EMS)-treated Jinhui10(Oryza sativa L). Because mfsl-1showed more severe defects than mfsl-2, the main focus of our study was mfsl-1.The mfsl-1spikelet displayed degenerated the sterile lemma, the elongated rachilla, the extra lemma-like organ and the degraded palea. Additionally, mfsl-1flowers produced varied numbers of inner floral organs. Scanning Elctron Microscropy (SEM) showed that the sterile lemma and palea was degraded in mfsl-1, floral meristem was enlarged and the stamen development was not synchronous in a few cases.
2. Genetic analysis and molecule mapping of MFSl
We employed3crosses to conduct the genetic analysis. The F1population showed wild-type phenotype and the segregation rates of wild-type and mfsl plants fit the ratio of3:1, suggesting that the MFS1gene was controlled by a single recessive gene. Next, we used the SSR markers and InDel markers to map the MFS1gene, the MFS1gene was finally mapped between Indell7and Indel24on chromosome5.
3. Cloning and protein analysis of the MFS1gene
Sequencing of DNA and cDNA analysis found a single nucleotide substitution from C to T within a predicted AP2/ERF transcription factor (Os05g041760) in different positions of the two mfsl alleles, causing a Ala66to Val66substitution in mfs1-1and a Thr51to Ile51substitution in mfs1-2. So, we infer that the Os05g041760is the candidate gene. MFS1was shown to belong to a clade of unknown function in the AP2/ERF family, and localized in the nucleus.
4. Expression patterns analysis of the MFS1gene
Semi-quantitative RT-PCR and quantitative RT-PCR (qPCR) analysis exhibited that MFS1was universally expressed in various tissues including roots, stems, leaves and panicles, especially high in young panicle. In situ hybridization analysis showed MFS1was strong expressed in the meristems and sterile lemma primordium. The MFS1expression was reduced gradually as the floral organ started to develop.
5. Function analysis of the MFS1gene
The MFS1complementation vector constructed and transformed into mfsl-1. we accuquired the transgenic positive plants and detected them GUS expression identification. In the transgenic plants, the mutant phenotypes were completely rescued in transgenic plants, suggesting that the Os05g41760is the MFS1gene.
The MFS1interference vectors were constructed and transformed into Jinhui10(Oryza sativa L. ssp. indica) and Zhonghuall (Oryza sativa L. ssp. japonica). We accuquired the transgenic positive plants and detected them by PCR and GUS expression identification. In the transgenic plants, pleiotropic spikelet defects were similar to those of mfs1-1. The result further confirmed that the Os05g41760is the MFS1gene and the functions of the MFS1gene.
6. MFS1regulates the expression of related floral development genes
The expression patterns showed the expression of A function gene OsMADS15was increased, the expressions of the AP2/ERF domian genes (FZP, SNB and OsIDSl) and the sterile lemma identity gene G1were decreased.
在本研究中,克隆了一个水稻小穗发育相关基因MULTI-FLIRET SPIKELET1(MFS1),我们对其进行了表型和细胞学观察,表达模式分析以及功能验证。主要结果如下：
1.mfs1突变体表型观察
从EMS诱变库中发现mfS1-1和mfs1-2两个等位小穗突变体,相比而言mfs1-1突变表型更严重和丰富,以下主要描述mfs1-1突变体。开花期的mfs1-1小穗护颖退化类似于副护颖,小穗轴伸长,形成额外的外稃,内稃主体部分退化,内三轮的花器官没有发现同源转化现象但花器官数目不确定。扫描电镜显示,早期护颖原基发育迟缓和退化,部分小花形成扩大的花分生组织和两个退化的内稃原基,雄蕊发育不同步。
2.MFS1遗传分析与分子定位
利用三个杂交组合进行遗传分析和分子定位,杂交F,代植株表型与野生型相似,F2代植株发生分离,正常株和突变株分离比均符合3：1,表明MFS1基因受单隐性基因控制。运用SSR标记和InDel标记进行基因定位,MFS1基因被最终定位在第5染色体InDel17和InDel24之间。
3.MFS1基因的克隆与蛋白质分析
DNA和cDNA测序分析发现一个编码APETALA2/EREBP结构域蛋白的基因Os05g41760分别在mfS1-1和mfs1-2突变体中存在一个C-T的替换,引起了氨基酸的改变,导致蛋白质的功能变异,初步确定其为候选基因。MFS1属于AP2/ERF家族中一个功能未知的分支,蛋白质定位在细胞核内。
4.MFS1基因的表达模式分析
利用半定量RT-PCR,qPCR和原位杂交技术进行了表达模式分析。半定量RT-PCR和qPCR结果显示MFS1基因在根,茎,叶和花中都表达,尤其在花序早期表达强烈。原位杂交显示MFS1在分生组织和护颖原基中表达强烈,随着花器官的发育,表达量逐渐减弱。
5.MFS1基因的功能验证
构建MFS1基因的基因组互补载体,转化mfs1-1突变体的愈伤组织,获得转基因植株,通过GUS染色检测转基因植株。结果表明,mfs1-1突变体表型完全恢复,确定了Os05g41760基因是MFS1基因。
构建MFS1基因的RNAi转基因表达载体,转化籼稻品种缙恢10号和粳稻品种中花11的愈伤组织,获得转基因植株,GUS染色、PCR检测转基因植株。RNAi转基因植株小穗表现为护颖退化,小穗轴伸长,额外的外稃状器官和内稃退化,与mfs1突变体的表型高度相似,进一步证明了OS05g41760基因是MFS1基因和MFS1基因的功能。
6.MFS1调控其它花发育相关基因的表达
qPCR分析和原位杂交分析结果表明,在mfs1-1突变体中,A功能基因OSMADs15表达量增加,AP2/ERF结构域基因SNB、OsIDS1和FZP表达量减少及护颖特征基因G1表达量减少。
Rice (Oryza sativa L.) is not only an important food crops in the world and a model plant in monocots but also an ideal object for studying flower development. Compaired with the model plants in dicots, rice bears a unique inflorescence consisting of spikelets to grass species, and has abundant genetic and genomic resources. So, reseach on rice reproductive development process would facilitate understanding of the molecular mechanism involved in spikelet development in rice and enrich knowledge of plant developmental biology for us. Meantime, it is very important to improve rice yields.
In the study, we cloned a MULTI-FLORET SPIKELET1(MFS1) gene associated with spikelet development in rice. We did investgation of mfsl mutants, and analysed the expression patterns and functions of MFS1gene. The results as follows:
1. Phenotypic observation of mfsl mutants
Two mutants of rice, mfsl-1and mfsl-2, were identified from ethylmethane sulfonate (EMS)-treated Jinhui10(Oryza sativa L). Because mfsl-1showed more severe defects than mfsl-2, the main focus of our study was mfsl-1.The mfsl-1spikelet displayed degenerated the sterile lemma, the elongated rachilla, the extra lemma-like organ and the degraded palea. Additionally, mfsl-1flowers produced varied numbers of inner floral organs. Scanning Elctron Microscropy (SEM) showed that the sterile lemma and palea was degraded in mfsl-1, floral meristem was enlarged and the stamen development was not synchronous in a few cases.
2. Genetic analysis and molecule mapping of MFSl
We employed3crosses to conduct the genetic analysis. The F1population showed wild-type phenotype and the segregation rates of wild-type and mfsl plants fit the ratio of3:1, suggesting that the MFS1gene was controlled by a single recessive gene. Next, we used the SSR markers and InDel markers to map the MFS1gene, the MFS1gene was finally mapped between Indell7and Indel24on chromosome5.
3. Cloning and protein analysis of the MFS1gene
Sequencing of DNA and cDNA analysis found a single nucleotide substitution from C to T within a predicted AP2/ERF transcription factor (Os05g041760) in different positions of the two mfsl alleles, causing a Ala66to Val66substitution in mfs1-1and a Thr51to Ile51substitution in mfs1-2. So, we infer that the Os05g041760is the candidate gene. MFS1was shown to belong to a clade of unknown function in the AP2/ERF family, and localized in the nucleus.
4. Expression patterns analysis of the MFS1gene
Semi-quantitative RT-PCR and quantitative RT-PCR (qPCR) analysis exhibited that MFS1was universally expressed in various tissues including roots, stems, leaves and panicles, especially high in young panicle. In situ hybridization analysis showed MFS1was strong expressed in the meristems and sterile lemma primordium. The MFS1expression was reduced gradually as the floral organ started to develop.
5. Function analysis of the MFS1gene
The MFS1complementation vector constructed and transformed into mfsl-1. we accuquired the transgenic positive plants and detected them GUS expression identification. In the transgenic plants, the mutant phenotypes were completely rescued in transgenic plants, suggesting that the Os05g41760is the MFS1gene.
The MFS1interference vectors were constructed and transformed into Jinhui10(Oryza sativa L. ssp. indica) and Zhonghuall (Oryza sativa L. ssp. japonica). We accuquired the transgenic positive plants and detected them by PCR and GUS expression identification. In the transgenic plants, pleiotropic spikelet defects were similar to those of mfs1-1. The result further confirmed that the Os05g41760is the MFS1gene and the functions of the MFS1gene.
6. MFS1regulates the expression of related floral development genes
The expression patterns showed the expression of A function gene OsMADS15was increased, the expressions of the AP2/ERF domian genes (FZP, SNB and OsIDSl) and the sterile lemma identity gene G1were decreased.
引文
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