一个水稻低温移栽白条纹突变体wltt的鉴定和基因定位
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摘要
【目的】叶色突变相关基因的鉴定与克隆为研究叶绿体发育、叶绿素合成和光合作用等分子机制提供理论基础。【方法】从常规粳稻镇糯19杂交后代中分离出一个低温移栽后叶色转成白条纹的自然变异突变体,命名为wltt (white stripe leaf after transplanting at low temperature)。成熟期测定野生型和wltt的主要农艺性状,分别在苗期、移栽后15 d和同时期直播条件下测定新生叶片的色素含量并观察叶绿体的超微结构;将wltt和野生型正反交进行遗传分析;用wltt与籼稻9311杂交产生的F_2作为定位群体进行基因定位;采用RT-qPCR分析叶绿体发育、叶绿素合成和光合作用相关基因在野生型和wltt中的表达水平。【结果】wltt突变体在苗期表现正常绿色,移栽15 d后心叶出现白条纹叶表型,至分蘖末期心叶叶色恢复;而不经移栽,突变体不会出现白条纹叶。人工模拟实验表明该表型是由低温条件下根损伤引起的。与野生型相比,wltt突变体移栽后的新生叶色素含量显著降低,光合速率下降;同时株高变矮,穗长、剑叶长和每穗粒数均显著降低。叶绿体的超微结构显示,突变体的叶肉细胞中,仅少数细胞含有正常的叶绿体,其余大部分叶肉细胞不含叶绿体。进一步研究发现,突变体中部分光合系统相关基因和叶绿体发育相关基因表达下调,叶绿素生物合成相关的14个基因表达也下调。遗传分析表明,该突变性状受一对隐性核基因控制。利用wltt突变体/9311的F_2群体,将该基因定位于水稻第2染色体着丝粒附近853kb区间内。目前,该区间内没有叶色相关基因的报道。【结论】WLTT是低温条件下移栽调控叶片转色的关键基因,在叶绿体发育过程中发挥重要作用。
【Objective】 Isolation and characterization of leaf-color mutation related genes lays a firm theoretical foundation for dissecting the molecular mechanism underlying chloroplast development, chlorophyll biosynthesis, and photosynthesis in rice. 【Method】A spontaneous leaf-color mutant, termed as white stripe leaf after transplanting at low-temperature(wltt), was obtained from the progeny of japonica cultivar Zhennuo 19. The main agronomic traits of the wild type and wltt were determined at maturity. The pigment contents and ultrastructure of chloroplast of newly emerged leaves were analyzed at the seedling stage, fifteen days after transplanting, at the tillering stage under direct seeding. Genetic analysis was carried out by reciprocal cross of the wild type and wltt. An F_2 population derived from the cross wltt×9311 was used for gene mapping. Quantitative RT-PCR was carried out to analyze the relative expression of genes associated with chloroplast development and chlorophyll biogenesis in the wild type and the wltt mutant. 【Result】The white-striped leaves in the wltt mutant only emerged at 15 days after transplanting at low temperature such as 20℃. No white-striped leaf was observed under direct seeding treatment. However, leaves of the mutant developed normally at the late tillering stage. Simulation experiments showed that the mutant phenotype was caused by root injury at low temperature. Compared with the wild type, the pigment contents in white-stripe leaves of the wltt mutant were significantly decreased, accompanying by reduced photosynthetic rate. Simultaneously, most of the mesophyll cells had no chloroplasts. The expression levels of genes associated with chloroplast development, chlorophyll biosynthesis, and photosynthetic system were all down-regulated in the mutant. At maturity, the mutant was featured with reduced plant height, panicle length, flag leaf length and number of spikelets per panicle relative to its wild type. Genetic analysis revealed that the mutant phenotype was controlled by a single recessive nuclear gene. Moreover, the WLTT gene was mapped within an 853 kb region near the centromere on chromosome 2, between InDel markers L22 and L26, in which no gene related to leaf color was reported. 【Conclusion】WLTT is a key gene regulating leaf color after transplanting at low temperature, which plays an important role in chloroplast development.
【Objective】 Isolation and characterization of leaf-color mutation related genes lays a firm theoretical foundation for dissecting the molecular mechanism underlying chloroplast development, chlorophyll biosynthesis, and photosynthesis in rice. 【Method】A spontaneous leaf-color mutant, termed as white stripe leaf after transplanting at low-temperature(wltt), was obtained from the progeny of japonica cultivar Zhennuo 19. The main agronomic traits of the wild type and wltt were determined at maturity. The pigment contents and ultrastructure of chloroplast of newly emerged leaves were analyzed at the seedling stage, fifteen days after transplanting, at the tillering stage under direct seeding. Genetic analysis was carried out by reciprocal cross of the wild type and wltt. An F_2 population derived from the cross wltt×9311 was used for gene mapping. Quantitative RT-PCR was carried out to analyze the relative expression of genes associated with chloroplast development and chlorophyll biogenesis in the wild type and the wltt mutant. 【Result】The white-striped leaves in the wltt mutant only emerged at 15 days after transplanting at low temperature such as 20℃. No white-striped leaf was observed under direct seeding treatment. However, leaves of the mutant developed normally at the late tillering stage. Simulation experiments showed that the mutant phenotype was caused by root injury at low temperature. Compared with the wild type, the pigment contents in white-stripe leaves of the wltt mutant were significantly decreased, accompanying by reduced photosynthetic rate. Simultaneously, most of the mesophyll cells had no chloroplasts. The expression levels of genes associated with chloroplast development, chlorophyll biosynthesis, and photosynthetic system were all down-regulated in the mutant. At maturity, the mutant was featured with reduced plant height, panicle length, flag leaf length and number of spikelets per panicle relative to its wild type. Genetic analysis revealed that the mutant phenotype was controlled by a single recessive nuclear gene. Moreover, the WLTT gene was mapped within an 853 kb region near the centromere on chromosome 2, between InDel markers L22 and L26, in which no gene related to leaf color was reported. 【Conclusion】WLTT is a key gene regulating leaf color after transplanting at low temperature, which plays an important role in chloroplast development.
引文
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[9]谭炎宁,孙学武,袁定阳,孙志忠,余东,何强,段美娟,邓华凤,袁隆平.水稻单叶独立转绿型黄化突变体grc2的鉴定与基因精细定位.作物学报,2015,41(6):831-837.Tan Y N,Sun X W,Yuan D Y,Sun Z Z,Yu D,He Q,Duan M J,Deng H F,Yuan L P.Identification and fine mapping of green-revertible chlorina gene grc2 in rice(Oryza sativa L.).Acta Agron Sin,2015,41(6):831-837.(in Chinese with English abstract)
[10]钱前,朱旭东,曾大力,张小惠,严学强,熊振民.细胞质基因控制的新特异材料白绿苗的研究.作物品种资源,1996(4):11-12.Qian Q,Zhu X D,Zeng D L,Zhang X H,Yan X Q,Xiong Z M.The study on a new special material,white-green rice which controlled by plasma gene.J Crop Resour,1996(4):11-12.(in Chinese).
[11]李贤勇,王楚桃,李顺武,何永歆,陈世全.一个水稻高叶绿素含量基因的发现.西南农业学报,2002,15(4):122-123.Li X Y,Wang C T,Li S W,He Y X,Chen S Q.The discovery of a high chlorophyll content gene in rice.Southwest China J Agric Sci,2002,15(4):122-123.(in Chinese with English abstract)
[12]Lee S,Kim J H,Yoo E S,Lee C H,Hirochika H,An G.Differential regulation of chlorophyll a oxygenase genes in rice.Plant Mol Biol,2005,57(6):805-818.
[13]Yang Y L,Xu J,Huang L C,Leng Y J,Dai L P,Rao Y C,Chen L,Wang Y Q,Tu Z J,Hu J,Ren D Y,Zhang G H,Zhu L,Guo L B,Qian Q,Zeng D L.PGL,encoding chlorophyllide a oxygenase 1,impacts leaf senescence and indirectly affects grain yield and quality in rice.J Exp Bot,2016,67(5):1297-1310.
[14]Kusumi K,Yara A,Mitsui N,Tozawa Y,Iba K.Characterization of a rice nuclear-encoded plastid RNApolymerase gene Os RpoTp.Plant Cell Physiol,2004,45(9):1194-1201.
[15]Sugimoto H,Kusumi K,Tozawa Y,Yazaki J,Kishimoto N,Kikuchi S,Iba K.The virescent-2 mutation inhibits translation of plastid transcripts for the plastid genetic system at an early stage of chloroplast differentiation.Plant Cell Physiol,2004,45(8):985-996.
[16]Beale S I.Green genes gleaned.Trends Plant Sci,2005,10(7):309-312.
[17]Nagata N,Tanaka R,Satoh S,Tanaka A.Identification of a vinyl reductase gene for chlorophyll synthesis in Arabidopsis thaliana and implications for the evolution of Prochlorococcus species.Plant Cell,2005,17(1):233-240.
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