水稻白条纹叶突变体wsl1的遗传分析及基因精细定位
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摘要
从粳稻日本晴和籼稻R1128杂交衍生的重组自交系群体中获得一个稳定遗传的白条纹叶突变体wsl1(white stripe leaf 1),世代为F10。与亲本R1128相比,突变体wsl1表现出白条纹叶,同时叶脉呈现白化,该性状在苗期就出现并持续整个生育期;突变体的株高、每穗总粒数、剑叶长、生育期显著增加,而结实率显著下降,其他农艺性状没有显著变化。分蘖期突变体wsl1的叶绿素a、叶绿素b和胡萝卜素含量较杂交亲本R1128显著下降;透射电镜观察表明,与野生型相比,突变体的叶绿体形状异常,不规则。遗传分析表明,该突变性状由1对隐性核基因控制。精细定位后发现,目标基因WSL1位于第1染色体短臂上标记M1-54与标记M1-70之间,两者相距89.7 kb。生物信息学分析表明候选区间内共有8个开放阅读框,暂未发现已报道的叶色相关基因;其中LOC_Os01g02080编码肽基脯氨酰顺反异构酶, GO (Gene Ontology)分类显示其可能与类囊体形成有关,后续将通过比较测序、qRT-PCR等分子实验来确定候选基因。
A white stripe leaf mutant wsl1 was obtained from the recombinant inbred lines derived from the cross of Oryza sativa var. japonica Nipponbare and Oryza sativa L. subsp. indica R1128. The mutant wsl1 showed white striped leaves and albino veins firstly at the seedling stage and then through the whole growth period. Agronomic traits such as plant height, number of spikelets per panicle, flag leaf length and heading date were significantly increased, while the seed setting rate decreased significantly in the mutant. Compared with wild type R1128, the chlorophyll a, chlorophyll b, and carotene contents of mutant leaves obviously decreased. Microscope observation indicated there were significantly decreased normal chloroplast and a large number of abnormal chloroplasts in mutant. Genetic analysis indicated that the mutant phenotype was controlled by a single recessive nuclear gene.WSL1 was mapped on the short arm of chromosome 1, between markers M1-54 and M1-70, with physical distance of about 89.7 kb. There were eight new open reading frames in the candidate region. Among them LOC_Os01 g02080 encodes a peptide-based prolyl cis-trans isomerase, GO(Gene Ontology) classification showed that it might be related to thylakoid formation.
A white stripe leaf mutant wsl1 was obtained from the recombinant inbred lines derived from the cross of Oryza sativa var. japonica Nipponbare and Oryza sativa L. subsp. indica R1128. The mutant wsl1 showed white striped leaves and albino veins firstly at the seedling stage and then through the whole growth period. Agronomic traits such as plant height, number of spikelets per panicle, flag leaf length and heading date were significantly increased, while the seed setting rate decreased significantly in the mutant. Compared with wild type R1128, the chlorophyll a, chlorophyll b, and carotene contents of mutant leaves obviously decreased. Microscope observation indicated there were significantly decreased normal chloroplast and a large number of abnormal chloroplasts in mutant. Genetic analysis indicated that the mutant phenotype was controlled by a single recessive nuclear gene.WSL1 was mapped on the short arm of chromosome 1, between markers M1-54 and M1-70, with physical distance of about 89.7 kb. There were eight new open reading frames in the candidate region. Among them LOC_Os01 g02080 encodes a peptide-based prolyl cis-trans isomerase, GO(Gene Ontology) classification showed that it might be related to thylakoid formation.
引文
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[29]Sugimoto H,Kusumi K,Noguchi K,Yano M,Yoshimura A,Iba K.The rice nuclear gene,VIRESCENT 2,is essential for chloroplast development and encodes a novel type of guanylate kinase targeted to plastids and mitochondria.Plant J,2007,52:512-527.
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[31]Wang Y,Zhang J,Shi X,Peng Y,Li P,Lin D,Dong Y,Teng S.Temperature-sensitive albino gene TCD5,encoding a monooxygenase,affects chloroplast development at low temperatures.JExp Bot,2016,67:5187-5202.
[32]Wang L,Wang C,Wang Y,Niu M,Ren Y,Zhou K,Zhang H,Lin Q,Wu F,Cheng Z,Wang J,Zhang X,Guo X,Jiang L,Lei C,Wang J,Zhu S,Zhao Z,Wan J.WSL3,a component of the plastid-encoded plastid RNA polymerase,is essential for early chloroplast development in rice.Plant Mol Biol,2016,92:581-595.
[33]Su N,Hu M L,Wu D X,Wu F Q,Fei G L,Lan Y,Chen X L,Shu X L,Zhang X,Guo X P,Cheng Z J,Lei C L,Qi C K,Jiang L,Wang H,Wan J M.Disruption of a rice pentatricopeptide repeat protein causes a seedling specific albino phenotype and its utilization to enhance seed purity in hybrid rice production.Plant Physiol,2012,159:227-238.
[34]成钦淑,叶邦全,袁灿,李伟滔,尹俊杰,王静,贺闽,汪吉春,王玉平,李仕贵,陈学伟.水稻白条纹叶突变体st11的遗传分析与基因定位.中国水稻科学,2015,29:14-21.Cheng Q S,Ye B Q,Yuan C,Li W T,Yin J J,Wang J,He M,Wang J C,Wang Y P,Li S G,Chen X W.Genetic analysis and gene mapping of white stripe leaf mutant stl1 in rice.Chin J Rice Sci,2015,29:14-21(in Chinese with English abstract).
[35]Cao P H,Ren Y,Kun L X,Zhang T Y,Zhang P,Xiao L J,Zhang F L,Liu S J,Jiang L,Wan J M.Purine nucleotide biosynthetic gene GARS controls early chloroplast development in rice(Oryza sativa L.).Plant Cell Rep,2019,38:183-194.
[2]Du P,Ling Y H,Sang X C,Zhao F M,Xie R,Yang Z L,He G H.Gene mapping related to yellow green leaf in a mutant line in rice(Oryza sativa).Genes Genom,2009,31:165-171.
[3]张洪征,程治军,万建民.水稻白化突变体研究进展.生物技术通报,2013,(11):1-7.Zhang H Z,Cheng Z J,Wan J M.Progresses on the studying of rice leaf albino.Biotechnol Bull,2013,(11):1-7(in Chinese with English abstract).
[4]Nagamatsu T,Omura T.Linkage study of the genes belonging to the first chromosome in rice.Jpn J Breed,1962,12:231-236.
[5]Birky C W Jr,Perlman P S,Byers T J.Genetics biogenesis mitochondria chloroplasts.Q Rev Biol,1976,51:527-528.
[6]Aluru M R,Rodermel S R.Control of chloroplast redox by the immutants terminal oxidase.Physiol Plant,2004,120:4-11.
[7]孙立亭,林添资,王云龙,牛梅,胡婷婷,刘世家,王益华,万建民.水稻白条纹突变体st13的表型分析及基因定位.中国水稻科学,2017,31:355-363.Sun L T,Lin T Z,Wang Y L,Niu M,Hu T T,Liu S J,Wang Y H,Wan J M.Phenotypic analysis and gene mapping of a white stripe mutant st13 in rice.Chin J Rice Sci,2017,31:355-363(in Chinese with English abstract).
[8]Li H C,Qian Q,Wang B,Li X B,Zhu L H,Xu J C.Identification and chromosomal localization of rice white panicle.Chin Sci Bull,2003,48:268-270.
[9]许凤华,程治军,王久林,吴自明,孙伟,张欣,雷财林,王洁,吴赴清,郭秀平,刘玲珑,万建民.水稻白条纹叶Gws基因的精细定位与遗传分析.作物学报,2010,36:71-720.Xu F H,Cheng Z J,Wang J L,Wu Z M,Sun W,Zhang X,Lei CL,Wang J,Wu F Q,Guo X P,Liu L L,Wan J M.Genetic analysis and fine-mapping of Gws gene using green-white-stripe rice mutant.Acta Agron Sin,2010,36:713-720(in Chinese with English abstract).
[10]Fang J,Chai C,Qian Q.Mutations of genes in synthesis of the carotenoid precursors of ABA lead to preharvest sprouting and photo-oxidation in rice.Plant J,2008,54:177-189.
[11]Li N,Chu H W,Wen T Q,Zhang D B.Genetic analysis and mapping of the rice white midrib mutant Oswm.Acta Agric Shanghai,2007,23(1):1-4.
[12]Ge C W,Wang L,Ye W J,Wu L W,Cui Y T,Chen P,Pan J J,Zhang D,Hu J,Zeng D L,Dong G J,Qian Q,Guo L B,Xue D W.Single-point mutation of an histidine-aspartic domain-containing gene involving in chloroplast ribosome biogenesis leads to white fine stripe leaf in rice.Sci Rep,2017,7:3298.
[13]Lee S,Kim J H,Yoo E S,Lee C H,Hirochika H.Differential regulation of chlorophyll a oxygenase genes in rice.Plant Mol Biol,2005,57:805-818.
[14]Zhang T,Feng P,Li Y F,Yu P,Yu G L,Sang X C,Ling Y H,Zeng X Q,Li Y D,Huang J Y,Zhang T Q,Zhao F M,Wang N,Zhang C W,Yang Z L,Wu R H,He G H.Virescent-albino leaf 1 regulates leaf colour development and cell division in rice.J Exp Bot,2018,doi:10.1093/jxb/ery250.
[15]Rogers S O,Bendich A J.Extraction of DNA from milligram amounts of fresh,herbarium and mummified plant tissues.Plant Mol Biol,1985,5:69-76.
[16]初志战,刘小林,陈远玲,刘耀光.一个水稻白化致死突变基因的精细定位和遗传研究.中国水稻科学,2016,30:136-142.Chu Z Z,Liu X L,Chen Y L,Liu Y G.Genetic analysis and gene mapping of an albino lethal mutant in rice.Chin J Rice Sci,2016,30:136-142(in Chinese with English abstract).
[17]Gothandam K M,Kim E S,Cho H,Chung Y Y.OsPPR1,a pentatricopeptide repeat protein of rice is essential for the chloroplast biogenesis.Plant Mol Biol,2005,58:421-433.
[18]Su N,Hu M L,Wu D X.Disruption of a rice pentatricopeptide repeat protein causes a seedling-specific albino phenotype and its utilization to enhance seed purity in hybrid rice production.Plant Physiol,2012,159:227-238.
[19]马娇,任德勇,吴国超,朱小燕,马玲,桑贤春,凌英华,何光华.水稻叶缘白化突变体mal的遗传分析与基因定位.作物学报,2014,40:591-599.Ma J,Ren D Y,Wu G C,Zhu X Y,Ma L,Sang X C,Ling Y H,He G H.Genetic analysis and gene mapping of a marginal albino leaf mutant mal in rice.Acta Agron Sin,2014,40:591-599(in Chinese with English abstract).
[20]Siddappa K,Vasudev K L,Ganiger B S,Rathod R,Devar K V.Report of albino seedlings in Pongamia pinnata.Karnataka JAgric Sci,2004,17:884-885.
[21]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,2015,41:831-837.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,2015,41:831-837(in Chinese with English abstract).
[22]朱小燕,徐芳芳,桑贤春,蒋钰东,代高猛,王楠,张长伟,何光华.水稻叶脉白化突变体wpsm的遗传分析与基因定位.作物学报,2013,39:1409-1415.Zhu X Y,Xu F F,Sang X C,Jiang Y D,Dai G M,Wang N,Zhang C W,He G H.Genetic analysis and gene mapping of a rice white midrib mutant wpsm.Acta Agron Sin,2013,39:1409-1415(in Chinese with English abstract).
[23]周坤能,夏加发,马廷臣,王元垒,李泽福.水稻条纹叶和白穗基因SLWP的定位及变异分析.中国水稻科学,2018,32:325-334.Zhou K N,Xia J F,Ma T C,Wang Y L,Li Z F.Mapping and mutation analysis of stripe leaf and white panicle of gene SLWP in rice.Chin J Rice Sci,2018,32:325-334(in Chinese with English abstract).
[24]王兴春,王敏,季芝娟,陈钊,刘文真,韩渊怀,杨长登.水稻糖苷水解酶基因OsBE1在叶绿体发育中的功能.作物学报,2014,40:2090-2097.Wang X C,Wang M,Ji Z J,Chen Z,Liu W Z,Han Y H,Yang CD.Functional characterization of the clycoside hydrolase encoding gene Os BE1 during chloroplast development in Oryza sativa.Acta Agron Sin,2014,40:2090-2097(in Chinese with English abstract).
[25]Gothandam K M,Kim E S,Cho H,Chung Y Y.OsPPR1,a pentatricopeptide repeat protein of rice is essential for the chloroplast biogenesis.Plant Mol Biol,2005,58:421-433.
[26]Lin D,Jiang Q,Zheng K,Chen S,Zhou H,Gong X,Xu J,Teng S,Dong Y.Mutation of the rice ASL2 gene encoding plastid ribosomal protein L21 causes chloroplast developmental defects and seedling death.Plant Biol(Stuttg),2015,17:599-607.
[27]Zhang Z,Tan J,Shi Z,Xie Q,Xing Y,Liu C,Chen Q,Zhu H,Wang J,Zhang J,Zhang G.Albino leaf1 that encodes the sole octotricopeptide repeat protein is responsible for chloroplast development.Plant Physiol,2016,171:1182-1191.
[28]Wang Y,Wang C,Zheng M,Liu J,Xu Y,Li X,Niu M,Long W,Wang D,Wang H Y,William T,Wang Y,Wan J.White Panicle1,a Val-tRNA synthetase regulating chloroplast ribosome biogenesis in rice,is essential for early chloroplast development.Plant Physiol,2016,170:2110-2123.
[29]Sugimoto H,Kusumi K,Noguchi K,Yano M,Yoshimura A,Iba K.The rice nuclear gene,VIRESCENT 2,is essential for chloroplast development and encodes a novel type of guanylate kinase targeted to plastids and mitochondria.Plant J,2007,52:512-527.
[30]Yoo S C,Cho S H,Sugimoto H,Li J,Kusumi K,Koh H J,Iba K,Paek N C.Rice virescent3 and stripe1 encoding the large and small subunits of ribonucleotide reductase are required for chloroplast biogenesis during early leaf development.Plant Physiol,2009,150:388-401.
[31]Wang Y,Zhang J,Shi X,Peng Y,Li P,Lin D,Dong Y,Teng S.Temperature-sensitive albino gene TCD5,encoding a monooxygenase,affects chloroplast development at low temperatures.JExp Bot,2016,67:5187-5202.
[32]Wang L,Wang C,Wang Y,Niu M,Ren Y,Zhou K,Zhang H,Lin Q,Wu F,Cheng Z,Wang J,Zhang X,Guo X,Jiang L,Lei C,Wang J,Zhu S,Zhao Z,Wan J.WSL3,a component of the plastid-encoded plastid RNA polymerase,is essential for early chloroplast development in rice.Plant Mol Biol,2016,92:581-595.
[33]Su N,Hu M L,Wu D X,Wu F Q,Fei G L,Lan Y,Chen X L,Shu X L,Zhang X,Guo X P,Cheng Z J,Lei C L,Qi C K,Jiang L,Wang H,Wan J M.Disruption of a rice pentatricopeptide repeat protein causes a seedling specific albino phenotype and its utilization to enhance seed purity in hybrid rice production.Plant Physiol,2012,159:227-238.
[34]成钦淑,叶邦全,袁灿,李伟滔,尹俊杰,王静,贺闽,汪吉春,王玉平,李仕贵,陈学伟.水稻白条纹叶突变体st11的遗传分析与基因定位.中国水稻科学,2015,29:14-21.Cheng Q S,Ye B Q,Yuan C,Li W T,Yin J J,Wang J,He M,Wang J C,Wang Y P,Li S G,Chen X W.Genetic analysis and gene mapping of white stripe leaf mutant stl1 in rice.Chin J Rice Sci,2015,29:14-21(in Chinese with English abstract).
[35]Cao P H,Ren Y,Kun L X,Zhang T Y,Zhang P,Xiao L J,Zhang F L,Liu S J,Jiang L,Wan J M.Purine nucleotide biosynthetic gene GARS controls early chloroplast development in rice(Oryza sativa L.).Plant Cell Rep,2019,38:183-194.