大麦黄绿叶色突变体ygl的农艺性状及其调控基因初步定位
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摘要
叶色突变是植物界广泛存在的一种现象,突变株系是解析植物光合作用和光形态建成等过程的理想材料。大麦黄绿叶色突变体ygl是本课题组在鄂大麦934的EMS突变体库中筛选获得的,该突变体的叶色在整个生育期都较野生型浅,而且苗期呈现黄绿色,后期表现为浅绿色。与野生型相比,ygl株高和千粒重分别表现为极显著和显著降低,其它重要产量性状,如穗数、穗长和穗粒数等无显著变化。进一步分析发现,ygl苗期叶片中基粒类囊体严重线性化,叶绿体超微结构受损。以正常叶色大麦品种Harrington和黄绿叶色突变体ygl为亲本构建了F_2分离群体,对F_2群体及其衍生的F_(2:3)家系进行分析发现,该性状由一个隐性核基因控制,命名为HvYGL(yellowgreenleaf)。利用上述F_2群体,通过基于SSR的BSA法,将该基因初步定位在大麦3H染色体上12.7 cM的区间内,其中 Bmag0209、 EBmac0871和 Bmag603三个标记与黄绿叶色性状共分离。
Leaf color mutations are common in plants.This kind of mutant has played an important role in illustrating the pathway of photosynthesis and photomorphism.Herein we describe an EMS mutant of barley cultivar Edamai 934,ygl,whose leaves are in yellow-green under the field conditions from the very beginning of its life,and in light green at the late stage of its life.Compared to the wild type,plant height and thousand grains weight of ygl were decreased significantly,while the other agronomic traits showed no significant changes,including spikes per plant,length of spike and grain number per spike.Further analysis showed that grana can not be stacked normally,and grana and stroma thylakoid was linear in leaves of ygl at seedling stage.Analysis of the F_2 population and F_(2:3) lines derived from barley cv.Harrington and ygl showed that this trait was controlled by a single recessive nuclear gene,designated as HvYGL in this study.HvYGL was initially mapped in a 12.7 cM interval on chromosome 3 H using the strategy of BSA based on SSR,and the three linked markers Bmag0209, EBmac0871 and Bmag603 co-segregated with the yellow-green leaf color phenomenon.The current study lays foundation for fine mapping and map-based cloning of the HvYGL gene.
Leaf color mutations are common in plants.This kind of mutant has played an important role in illustrating the pathway of photosynthesis and photomorphism.Herein we describe an EMS mutant of barley cultivar Edamai 934,ygl,whose leaves are in yellow-green under the field conditions from the very beginning of its life,and in light green at the late stage of its life.Compared to the wild type,plant height and thousand grains weight of ygl were decreased significantly,while the other agronomic traits showed no significant changes,including spikes per plant,length of spike and grain number per spike.Further analysis showed that grana can not be stacked normally,and grana and stroma thylakoid was linear in leaves of ygl at seedling stage.Analysis of the F_2 population and F_(2:3) lines derived from barley cv.Harrington and ygl showed that this trait was controlled by a single recessive nuclear gene,designated as HvYGL in this study.HvYGL was initially mapped in a 12.7 cM interval on chromosome 3 H using the strategy of BSA based on SSR,and the three linked markers Bmag0209, EBmac0871 and Bmag603 co-segregated with the yellow-green leaf color phenomenon.The current study lays foundation for fine mapping and map-based cloning of the HvYGL gene.
引文
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[19]AGRAWAL G,BLAXTER M,YAMAZAKI M,et al.Screening of the rice viviparous mutants generated by endogenous retrotransposon Tos17 insertion.Tagging of a zeaxanthin epoxidase gene and a novel OsTATC gene [J].Plant Physiology,2001,125(3):125:4.
[20]LIU X G,XU H,ZHANG J Y,et al.Effect of low temperature on chlorophyll biosynthesis in albinism line of wheat(Triticum aestivum) FA85 [J].Physiologia Plantarum,2012,145(3):385.
[21]PRINA A,ARIAS M,LAINEZ V,et al.A cytoplasmically inherited mutant controlling early chloroplast development in barley seedlings [J].Theoretical and Applied Genetics,2003,107(8):1411.
[22]WANG R,YANG F,ZHANG X,et al.Characterization of a thermo-inducible chlorophyll-deficient mutant in barley [J].Frontier in Plant Science,2017,8:4.
[2]XIA J,WANG Y,MA B,et al.Ultrastructure and gene mapping of the albino mutant al12 in rice(Oryza satave L.) [J].Acta Genetica Sinica,2006,33(12):1114.
[3]CHEN T,ZHANG Y,ZHAO L,et al.Fine mapping and candidate gene analysis of a green-revertible albino gene gra(t) in rice [J].Journal of Genetics and Genomics,2009,36,118.
[4]张向前,李晓燕,朱海涛,等.水稻阶段性返白突变体的鉴定和候选基因分析[J].科学通报,2010,55(23):2298.ZHANG X Q,LI X Y,ZHU H T,et al.Identification and candidate gene analysis of stage green-revertible albino mutant in rice [J].Chinese Science Bulletin,2010,55(23):2298.
[5]郭涛,黄宣,黄永相,等.一个控制水稻叶色变化转绿及多分蘖矮杆性状基因 hw-1(t)的鉴定 [J].作物学报,2012,38(1):25.GUO T,HUANG Y,HUANG Y,et al.Characterizations of a mutant gene hw-1(t) for green-revertible albino,high tillering and dwarf in rice(Oryza sativa L.) [J].Acta Agronomica Sinica,2012,38(1):25.
[6]ZHANG T,FENG P,LI Y,et al.VIRESCENT-ALBINO LEAF 1 regulates leaf colour development and cell division in rice [J].Journal of Experimental Botany,2018,4794.
[7]MEI J,LI F,LIU X,et al.Newly identified CSP41b gene localized in chloroplasts affects leaf color in rice [J].Plant Science,2017,256:41.
[8]MA X,SUN X,LI C,et al.Map-based cloning and characterization of the novel yellow-green leaf gene ys83 in rice(Oryza sativa) [J].Plant Physiology and Biochemistry,2017,111:4.
[9]王婷婷,翟立红,苏旭,等.Mutator诱导的玉米白化突变体插入位点的遗传分析及代谢途径构建[J].中国农业科学,2010,43(22):4573 .WANG T T.ZHAI L H,SU X,et al.Genetic analysis and construction of metabolic pathway by mutator- mediated albino mutant insertion sites in Zea Mays L.[J].Scientia Agricultura Sinica,2010,43(22):4573.
[10]朱碧岩,蔡建秋,程方民,等.大麦叶色转换突变系的转换特性及其基因控制[J].华南师范大学学报,1998(1):73.ZHU B,CAI J,CHEN F,et al.Study on characteristic of leaf colour conversion mutant lines of barly and their heredity[J].Journal of South China Normal University,1998(1):73.
[11]陈升位,王楠,陈疏影,等.3种裸大麦叶色突变体的选育及其突变性状的比对分析[J].云南农业大学学报,2014,29(1):3.CHEN S,WANG N,CHEN S,et al.Development of three leaf color mutants of naked barley and the comparative analysis of their mutant traits [J].Journal of Yunnan Agricultural University,2014,29(1):3.
[12]GALOVA E,BOHMOVA B,SEVCOVICOVA A.Analysis of some barley chlorophyll mutants and their response to temperature stress [J].Photosynthetica,2000,38(1):29-35.
[13]YARONSKAYA E,ZIEMANN V,WALTER G,et al.Metabolic control of the tetrapyrrole biosynthetic pathway for porphyrin distribution in the barley mutant albostrians [J].The Plant Journal,2003,35:513.
[14]李志勇.西藏野生大麦产量与农艺性状的相关分析与大麦叶色黄化突变体的基因定位[D].杭州:浙江大学,2016:38.LI Z Y.The correlation analysis between yield and agronomic traits in Tibetan wild Barley and the gene mapping of yellow leaves mutants [D].Hangzhou:Zhejiang University,2016:38.
[15]QIN D D,DONG J,XU F C,et al.Characterization and fine mapping of a novel barley stage green-revertible albino gene(HvSGRA) by bulked segregant analysis based on SSR assay and specific length amplified fragment sequencing [J].BMC Genomics,2015,16:840.
[16]MICHELMORE R W,PARAN I,KESSELI R.Identification of markers linked to disease-resistance genes by bulked segregant analysis:A rapid method to detect markers in specific genomic regions by using segregating populations [J].Proceedings of the National Academy of Sciences,1991,88(21):9828.
[17]PARKS B M,QUAIL P H.Phytochrome-deficient hy1 and hy2 long hypocotyl mutants of Arabidopsis are defective in phytochrome chromophore biosynthesis [J].The Plant Cell,1991,3(11):1178.
[18]FAMBRINI M,CASTAGNA A,VECCHIA F D,et al.Characterization of a pigment-deficient mutant of sunflower(Helianthus annuus L.) with abnormal chloroplast biogenesis,reduced PS II activity and low endogenous level of abscisic acid [J].Plant Science,2004,167(1):81.
[19]AGRAWAL G,BLAXTER M,YAMAZAKI M,et al.Screening of the rice viviparous mutants generated by endogenous retrotransposon Tos17 insertion.Tagging of a zeaxanthin epoxidase gene and a novel OsTATC gene [J].Plant Physiology,2001,125(3):125:4.
[20]LIU X G,XU H,ZHANG J Y,et al.Effect of low temperature on chlorophyll biosynthesis in albinism line of wheat(Triticum aestivum) FA85 [J].Physiologia Plantarum,2012,145(3):385.
[21]PRINA A,ARIAS M,LAINEZ V,et al.A cytoplasmically inherited mutant controlling early chloroplast development in barley seedlings [J].Theoretical and Applied Genetics,2003,107(8):1411.
[22]WANG R,YANG F,ZHANG X,et al.Characterization of a thermo-inducible chlorophyll-deficient mutant in barley [J].Frontier in Plant Science,2017,8:4.