一个水稻花发育突变体的基因克隆及功能分析
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摘要
水稻是单子叶植物中发育分子生物学研究的模式植物,突变体则是功能基因组学研究的重要材料,因此水稻发育突变体的获得和研究,在揭示水稻发育相关基因的作用及其相互关系,探明植物发育,特别是花发育的奥秘中起着十分重要的作用。近年来有关水稻发育基因调控的研究已取得了长足的进展,通过突变体获得了一系列与水稻发育相关的基因,对高等植物发育调控机制的研究具有重要的意义。
我们在本实验室发展的日本晴突变体库中发现的一个水稻花器官突变体,进一步观察整体植株发现从株高、叶以及叶鞘都有差异,其中以花器官的差异最为明显。遗传分析表明该突变体的表型受位于八号染色体上一不完全显性单基因控制,利用图位克隆法精细定位后,推测侯选基因编码--WD40repeat蛋白，并与拟南芥中的TOPLESS基因同源,因此将这个基因暂命名为Ostopless8(Ostp18)。进一步对基因序列测序发现位于BAC AP005504上14053处A>T,从面导致GAG>GTG(Glu>Val).为研究该基因功能构建了互补、GUS、GFP、过表达等载体,其中将突变位点引入野生型日本晴中成功产生了突变体表型,GUS转基因结果显示该基因在各组织中的表达位置,其他载体正在转化中。为进一步了解该基因与其他基因之间的调控关系,本研究还选取了营养生长期野生型、突变体和杂合体叶片组织提取RNA做基因芯片,发现一些可能有关联的基因,并用Realtime-PCR进行验证。Ostpl8基因的克隆与功能分析为进一步了解水稻发育的分子调控机制打下了良好基础。
Rice is the model for studying development of monocotyledon in Molecular Biology and mutants are important materials for research into functional genomics. The acquisition and research on the mutants related to development play an important role in understanding the function and interaction of genes in development process, especially in floral development. Recently, there have been many progresses in genes regulating the development of rice. Many genes about this process have been cloned by mutants. So that, the study on molecular mechanism of monocotyledon development using rice mutants becomes one of the focuses in plant molecular genetics. The research about development regulation and control mechanism has a major significance in higher plant
We obtained a floral organ mutant from the Nipponbare mutant library which was constructed by our lab. Further observation of the whole plant found that there were differences from height, leaf, and sheath, particularly in floral organ. Genetic analysis showed the phenotype of the mutant is controlled by a semi-dominant gene on Chromosome8. We fine mapped this gene and predicted it as a WD40repeat protein which is homologous with TOPLESS in Arabidopsis thaliana by map-based cloning and we provisionally named it as Ostopless8(Ostpl8). The sequencing result showed that the A change to T and cause GAG change to GTG (Glu> Val). We constructed several vectors such as complementation, GUS, GFP and over-expression to research the gene's function. The mutation in Nipponbare successfully caused mutant phenotype and GUS transgene plant showed the expression location iof Ostpl8in tissue while other vectors were under transgene. We extract RNA from leaves of wild type, mutant and heterozygote in vegetative period for gene chip and found some relationship between genes, some of these results had been proved by Realtime-PCR. The cloning and function analysis of Ostpl8lays a good foundation for further studying of molecular mechanisms in regulation of rice development.
我们在本实验室发展的日本晴突变体库中发现的一个水稻花器官突变体,进一步观察整体植株发现从株高、叶以及叶鞘都有差异,其中以花器官的差异最为明显。遗传分析表明该突变体的表型受位于八号染色体上一不完全显性单基因控制,利用图位克隆法精细定位后,推测侯选基因编码--WD40repeat蛋白，并与拟南芥中的TOPLESS基因同源,因此将这个基因暂命名为Ostopless8(Ostp18)。进一步对基因序列测序发现位于BAC AP005504上14053处A>T,从面导致GAG>GTG(Glu>Val).为研究该基因功能构建了互补、GUS、GFP、过表达等载体,其中将突变位点引入野生型日本晴中成功产生了突变体表型,GUS转基因结果显示该基因在各组织中的表达位置,其他载体正在转化中。为进一步了解该基因与其他基因之间的调控关系,本研究还选取了营养生长期野生型、突变体和杂合体叶片组织提取RNA做基因芯片,发现一些可能有关联的基因,并用Realtime-PCR进行验证。Ostpl8基因的克隆与功能分析为进一步了解水稻发育的分子调控机制打下了良好基础。
Rice is the model for studying development of monocotyledon in Molecular Biology and mutants are important materials for research into functional genomics. The acquisition and research on the mutants related to development play an important role in understanding the function and interaction of genes in development process, especially in floral development. Recently, there have been many progresses in genes regulating the development of rice. Many genes about this process have been cloned by mutants. So that, the study on molecular mechanism of monocotyledon development using rice mutants becomes one of the focuses in plant molecular genetics. The research about development regulation and control mechanism has a major significance in higher plant
We obtained a floral organ mutant from the Nipponbare mutant library which was constructed by our lab. Further observation of the whole plant found that there were differences from height, leaf, and sheath, particularly in floral organ. Genetic analysis showed the phenotype of the mutant is controlled by a semi-dominant gene on Chromosome8. We fine mapped this gene and predicted it as a WD40repeat protein which is homologous with TOPLESS in Arabidopsis thaliana by map-based cloning and we provisionally named it as Ostopless8(Ostpl8). The sequencing result showed that the A change to T and cause GAG change to GTG (Glu> Val). We constructed several vectors such as complementation, GUS, GFP and over-expression to research the gene's function. The mutation in Nipponbare successfully caused mutant phenotype and GUS transgene plant showed the expression location iof Ostpl8in tissue while other vectors were under transgene. We extract RNA from leaves of wild type, mutant and heterozygote in vegetative period for gene chip and found some relationship between genes, some of these results had been proved by Realtime-PCR. The cloning and function analysis of Ostpl8lays a good foundation for further studying of molecular mechanisms in regulation of rice development.
引文
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Hong Z, Ueguchi-Tanaka M, Umemura K, et al. A rice brassinosteroid-deficient mutant, Ebisu Dwarf (D2), is caused by loss of function of a new member of cytochrome P450. Plant Cell 2003,15:2900-2910.
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Inukai Y, Miwa M, Nagato Y, et al. Characterization of rice mutants deficient in the formation of crown roots. Breed Sci,2001,51:123-129.
Inukai Y, Sakamoto T, Ueguchi-Tanaka M, et al. Crown rootless], which is essential for crown root formation in rice, is a target of an AUXIN RESPONSE FACTOR in auxin signaling. Plant Cell,2005,17:1387-1396.
Itoh J I, Kitano H, Matsuoka M, et al. SHOOT ORGANIZATION genes regulate the organization of shoot apical meristem and the leaf initiation pattern. Plant Cell, 2000,12:2161-2174.
Itoh J I, Nonomura K, Ikeda K, et al. Rice plant development:from zygote to spikelet[J]. Plant Cell Physiol,2005,46(1):23-47.
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Jackson D, Veit B, Hake S. Expression of maize KNOTTEDl related homeobox genes in the shoot apical meristem predicts patterns of morphogenesis in the vegetative shoot. Development,1992,120:405-413.
Kamiya N, Itoh, J I, Morikami A, et al. The SCARECROW gene's role in asymmetric cell divisions in rice plants. Plant J,2003a,36(1):45-54.
Kamiya N, Nagasaki H, Morikami A, et al. Isolation and characterization of a rice WUSCHEL-type homeobox gene that is specifically expressed in the central cells of a quiescent center in the root apical meristem. Plant J,2003b,35(4):429-441.
Kaufman P B. Development of the shoot of Oryza sativa Ⅰ:the shoot apex. Phytomorphology,1959,9:228-242.
Kaufman P B. Development of the shoot of Oryza sativa Ⅱ:leaf histogenesis. Phytomorphology,1959,9:277-311.
Kawata Shin-ichiro, S.H.. on the lateral root primordia formation in the crown roots of rice plant. Proc Crop Sci Soc Jpn,1976,33,423-431.
Keishi Komatsu, Masahiko Maekawa, Shin Ujiie. LAX and SPA:Major regulators of shoot branching in rice. Proc Natl Acad Sci USA,2003,100(20):11765-11770.
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