利用F_2和F_3群体定位水稻粒形和千粒重QTL
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摘要
水稻籽粒的粒长、粒宽和粒厚共同塑成了水稻粒形,并决定水稻籽粒的千粒重及外观品质,而千粒重是决定水稻产量三要素之一。本研究以籼型温敏核不育系广占63-4S为母本,以大粒籼稻TGMS29为父本杂交衍生的F_2和F_3群体,对影响水稻籽粒粒形和千粒重的数量性状位点(quantitative trait loci, QTL)进行定位。2014年和2015年检测到影响籽粒粒长的QTL 5个,粒宽QTL 8个,粒厚的QTL 4个,千粒重QTL 4个。其中,两年重复检测到2个粒长QTL,分别位于第1染色体RM1和RM490之间,可解释的表型变异为21.3%、22%,和第2染色体RM240和RM208之间,可解释的表型变异为12.8%、15.7%。两年均被检测到的粒宽QTL位于第3染色体RM251和RM571之间,可解释表型变异分别为53.60%、55.00%。两年均被检测到的千粒重QTL位于第1染色体RM220和RM1之间,可解释表型变异分别为15.1%、15.4%。这些QTLs的鉴定为稻米粒形和千粒重的遗传研究提供了帮助,也为稻米品质和产量的改良提供了宝贵的基因资源。
The grain shape of rice consists of three factors, namely grain length, width and thickness, which determines the 1 000-grain weight and appearance quality, and the 1 000-grain weight is one of the three factors determining rice yield. In this study, QTL of grain shape and 1 000-grain weight was mapped using F_2 and F_3 populations derived from indica temperature-sensitive genic male-sterile lines(TGMS) GZ63-4 S as female parent and large-grain indica TGMS29 as male parent. In 2014 and 2015, 5 QTL affecting grain length, 8 QTLs affecting grain width, 4 QTL affecting grain thickness, and 4 QTL affecting 1 000-grain weight were detected. Among that, two QTLs for grain length were repeatedly detected, one of which was located between RM1 and RM490 on chromosome 1, interpreting 21.3% and 22% of the phenotypic variation in two years respectively, and the other was located between RM240 and RM208 on chromosome 2, interpreting 12.8% and 15.7% of the variation respectively.The QTL for grain width detected in both years was located between RM251 and RM571 on chromosome 3, interpreting 53.6% and 55% of the phenotypic variance, respectively. The QTL for 1 000-grain weight was located between RM220 and RM1 on chromosome 1, interpreting 15.1% and 15.4% of the phenotypic variance in year 2014 and 2015, respectively. The identification of these QTLs would be helpful for the genetic studies on grain shape and 1 000-grain weight of rice, and provided valuable genetic resources for the improvement of grain quality and grain yield in rice.
The grain shape of rice consists of three factors, namely grain length, width and thickness, which determines the 1 000-grain weight and appearance quality, and the 1 000-grain weight is one of the three factors determining rice yield. In this study, QTL of grain shape and 1 000-grain weight was mapped using F_2 and F_3 populations derived from indica temperature-sensitive genic male-sterile lines(TGMS) GZ63-4 S as female parent and large-grain indica TGMS29 as male parent. In 2014 and 2015, 5 QTL affecting grain length, 8 QTLs affecting grain width, 4 QTL affecting grain thickness, and 4 QTL affecting 1 000-grain weight were detected. Among that, two QTLs for grain length were repeatedly detected, one of which was located between RM1 and RM490 on chromosome 1, interpreting 21.3% and 22% of the phenotypic variation in two years respectively, and the other was located between RM240 and RM208 on chromosome 2, interpreting 12.8% and 15.7% of the variation respectively.The QTL for grain width detected in both years was located between RM251 and RM571 on chromosome 3, interpreting 53.6% and 55% of the phenotypic variance, respectively. The QTL for 1 000-grain weight was located between RM220 and RM1 on chromosome 1, interpreting 15.1% and 15.4% of the phenotypic variance in year 2014 and 2015, respectively. The identification of these QTLs would be helpful for the genetic studies on grain shape and 1 000-grain weight of rice, and provided valuable genetic resources for the improvement of grain quality and grain yield in rice.
引文
Bai X.F.,Luo L.J.,Yan W.H.,Kovi M.R.,Zhan W.,and Xing Y.Z.,2010,Genetic dissection of rice grain shape using a recombinant inbred line population derived from two contrasting parents and fine mapping a pleiotropic quantitative trait locus qgl7,BMC Genetics,11(1):16
Fan C.C.,Xing Y.Z.,Mao H.L.,Lu T.T.,Han B.,Xu C.G.,Li X.H.,and Zhang Q.F.,2006,Gs3,a major QTL for grain length and weight and minor QTL for grain width and thickness in rice,encodes a putative transmembrane protein,Theor.Appl.Genet.,112(6):1164-1171
Fu F.H.,Wang F.,Huang W.J.,Peng H.P.,Wu Y.Y.,and Huang D.J.,1994,Genetic analysis on grain characters in hybrid rice,Zuowu Xuebao(Acta Agronomica Sinica),1(7):39-45(符福鸿,王丰,黄文剑,彭惠普,伍应运,黄德娟,1994,杂交水稻谷粒性状的遗传分析,作物学报,1(7):39-45)
Jiang Y.D.,Luo J.T.,Kuang H.C.,Yang Y.,Chen G.Z.,He X.C.,Fu J.,Zheng J.,and Zeng Z.M.,2015,Identification of QTLs for grain shape and weight using F2generation of two Indica rice varieties,Fenzi Zhiwu Yuzhong(Molecular Plant Breeding),13(8):1689-1694(蒋钰东,罗俊涛,况浩池,杨扬,陈光珍,何兴财,付均,郑军,曾正明,2015,利用两个籼稻杂交F2定位水稻粒型,粒重,分子植物育种,13(8):1689-1694)
Li J.X.,Yu S.B.,Xu C.G.,Tan Y.F.,Gao Y.J.,Li X.H.,and Zhang Q.F.,2000,Analyzing quantitative trait loci for yield using a vegetatively replicated F2population from a cross between the parents of an elite rice hybrid,Theor.Appl.Genet.,101(1-2):248-254
Li Q.,Li L.,Yang X.H.,Warburton M.L.,Bai G.H.,Dai J.R.,and Li J.S.,2010,Relationship,evolutionary fate and function of two maize co-orthologs of rice GW2 associated with kernel size and weight,BMC Plant Biology,10(1):143
Li Y.B.,Fan C.C.,Xing Y.Z.,Jiang Y.H.,Luo L.J.,Sun L.,Shao D.,Xu C.J.,Li X.H.,Xiao J.H.,He Y.Q.,and Zhang Q.F.,2011,Natural variation in GS5 plays an important role in regulating grain size and yield in rice,Nat.Genet.,43(12):1266-1269
Lin H.X.,Min S.K.,Xiong Z.M.,Qian H.R.,Zhuang J.Y.,Lu J.,Zheng K.L.,and Huang N.,1995,RFLP mapping of QTLs for grain shape traits in indica rice(Oryza sattve L.subsp.indica),Zhongguo Nongye Kexue(Scientia Agricultura Sinica),28(4):1-7(林鸿宣,闵绍楷,熊振民,钱惠荣,庄杰云,陆军,郑康乐,黄宁,1995,应用RFLP图谱定位分析籼稻粒型数量性状基因定位,中国农业科学,28(4):1-7)
Lin L.H.,and Wu W.R.,2003,Mapping of QTLs underlying grain shape and grain weight in rice,Fenzi Zhiwu Yuzhong(Molecular Plant Breeding),1(3):337-342(林荔辉,吴为人,2003,水稻粒型和粒重的QTL定位分析,分子植物育种,1(3):337-342)
Liu D.,Wang J.Y.,Chai Y.S.,Sun Y.Y.,Sun G.H.,Feng Z.L.,and Chen W.F.,2015,Mapping of the QTLs for rice grain shape using recombinant inbred lines of Indica-Japonica hybridization in rice,Shenyang Nongye Daxue Xuebao(Journal of Shenyang Agricultural University),46(4):385-390(刘丹,王嘉宇,柴永山,孙玉友,孙国宏,冯章丽,陈温福,2015,利用籼粳交RILs群体的水稻粒形QTL定位,沈阳农业大学学报,46(4):385-390)
Ma M.L.,Liu Y.H.,Jiang L.,Lei E.,Su Y.L.,and Lu B.Y.,2015,QTLs mapping of grain length-width ration by using F2population in rice,Jiangxi Nongye Daxue Xuebao(Acta Agriculturae Universitatis Jiangxiensis),37(2):260-263(马孟莉,刘艳红,江玲,雷恩,苏一兰,卢丙越,2015,利用F2群体定位水稻谷粒长宽比QTL,江西农业大学学报,37(2):260-263)
Qi P.,Lin Y.S.,Song X.J.,Shen J.B.,Huang W.,Shan J.X.,Zhu M.Z.,Jiang L.W.,Gao J.P.,and Lin H.X.,2012,The novel quantitative trait locus GL3.1 controls rice grain size and yield by regulating Cyclin-T13,Cell Research,22(12):1666-1680
Sun B.,Zhan X.D.,Lin Z.C.,Gao Z.Q.,Yu P.,Liu Q.E.,Shen X.H.,Zhang Y.X.,Chen D.B.,Cheng S.H.,and Cao L.Y.,2015,Correlation analysis and QTL mapping of grain shape and grain weight in rice,Fenzi Zhiwu Yuzhong(Molecular Plant Breeding),13(12):2663-2672(孙滨,占小登,林泽川,高志强,于萍,刘群恩,沈希宏,张迎信,陈代波,程式华,曹立勇,2015,水稻粒形和粒重性状的相关性分析及QTL定位,分子植物育种,13(12):2663-2672)
Wang S.K.,Li S.,Liu Q.,Wu K.,Zhang J.Q.,Wang S.S.,Wang Y.,Chen X.B.,Zhang Y.,Gao C.X.,Wang F.,Huang H.X.,and Fu X.D.,2015,The OsSPL16-GW7 regulatory module determines grain shape and simultaneously improves rice yield and grain quality,Nature Genetics,47(8):949-954
Weng J.F.,Gu S.H.,Wan X.Y.,Gao H.,Guo T.,Su N.,Lei C.L.,Zhang X.,Cheng Z.J.,Guo X.P.,Wang J.L.,Jiang L.,Zhai H.Q.,and Wan J.M.,2008,Isolation and initial characterization of GW5,a major QTL associated with rice grain width and weight,Cell Res.,18(12):1199-1209
Xia D.,Zhou H.,Liu R.J.,Dan W.H.,Li P.B.,Wu B.,Chen J.X.,Wang L.Q.,Gao G.J.,Zhang Q.L.,and He Y.Q.,2018,GL3.3,a Novel QTL encoding a GSK3/SHAGGY-like kinase,epistatically interacts with GS3 to produce extra-long grains in rice,Molecular Plant,11(5):754-756
Xiao J.H.,and Zhang Q.F.,2010,Functional genomics research and green super rice,Fenzi Zhiwu Yuzhong(Molecular Plant Breeding),8(6):1050-1053(肖景华,张启发,2010,功能基因组与绿色超级稻,分子植物育种,8(6):1050-1053)
Xu J.J.,Wang B.H.,Wu Y.H.,Du P.N.,Wang J.,Wang M.,Yi C.D.,Gu M.H.,and Liang G.H.,2011,Fine mapping and candidate gene analysis of ptgms2-1,the photoperiod-thermo-sensitive genic male sterile gene in rice(Oryza sativa L.),Theor.Appl.Genet.,122(2):365-372
Xu J.J.,Zhao Q.,Tang Z.X.,Zhao Y.F.,Zhu L.,Xu C.W.,Gu M.H.,Han B.,and Liang G.H.,2011,Mapping of QTLs for grain shape using whole-genome sequenced chromosome segment substitution lines in rice,Zhongguo Nongye Kexue(Scientia Agricultura Sinica),25(4):365-369(徐建军,赵强,汤在祥,赵元凤,朱磊,徐辰武,顾铭洪,韩斌,梁国华,2011,利用重测序的染色体片段代换系群体定位水稻粒型QTL,中国水稻科学,25(4):365-369)
Yang L.S.,Bai Y.S.,Zhang P.J.,Xu C.W.,Hu X.M.,Wang W.M.,She D.H.,and Chen G.Z.,2001,Studies for the correlations between grain shape and rice quality,Zajiao Shuidao(Hybrid Rice),16(4):48-54(杨联松,白一松,张培江,许传万,胡兴明,王伍梅,佘德红,陈桂芝,2001,谷粒形状与稻米品质相关性研究,杂交水稻,16(4):48-54)
Yang W.N.,2011,Studies on digital technology of automatic extraction of parameters for rice yield related traits,Dissertation for Ph.D.,Huazhong University of Science and Technology,Supervisor:Liu Q.,pp.100-105(杨万能,2011,水稻产量相关性状参数自动提取的数字化技术研究,博士学位论文,华中科技大学,导师:刘谦,pp.100-105)
Yu B.,Nassirou T.Y.,Gao G.J.,and Zhang Q.L.,2012,QTLs analysis on appearance quality traits and kernel weight of rice,Hubei Nongye Kexue(Hubei Agricultural Sciences),51(19):187-192(于波,Nassirou T.Y.,高冠军,张庆路,2012,水稻外观品质性状和千粒重的QTL分析,湖北农业科学,51(19):187-192)
Zeng R.Z.,Talukdar A.,Liu F.,and Zhang G.Q.,2006,Mapping of the QTLs for grain shape using single segment substitu tion lines in rice,Zhongguo Nongye Kexue(Scientia Agricultura Sinica),39(4):647-654(曾瑞珍,Talukdar A.,刘芳,张桂权,2006,利用单片段代换系定位水稻粒形QTL,中国农业科学,39(4):647-654)
Zhang Y.D.,Zhang Y.H.,Dong S.L.,Chen T.,Zhao Q.Y.,Zhu Z.,Zhou L.H.,Yao Z.,Zhao L.,Yu X.,and Wang C.L.,2013,Identification of QTL for rice grain traits based on an extralarge grain material,Zhongguo Shuidao Kexue(China Journal Rice Science),27(2):122-128(张亚东,张颖慧,董少玲,陈涛,赵庆勇,朱镇,周丽慧,姚姝,赵凌,于新,王才林,2013,特大粒水稻材料粒型性状的QTL检测,中国水稻科学,27(2):122-128)
Zhao J.G.,Jiang K.F.,Yang L.,Yang Q.H.,Wan X.Q.,Cao Y.J.,You S.M.,Luo J.,Zhang T.,and Zheng J.K.,2013,QTLmapping for yield related components in a RIL population of rice,Zhongguo Shuidao Kexue(China Journal Rice Science),27(4):344-352(赵建国,蒋开锋,杨莉,杨乾华,万先齐,曹应江,游书梅,罗婧,张涛,郑家奎,2013,水稻产量相关性状QTL定位,中国水稻科学,27(4):344-352)
Zou D.T.,Liu Z.L.,Zhao H.W.,Sun J.,Zheng H.L.,Chen B.B.,and Liu B.W.,2014,Mapping QTLs for grain shape and weight using F2:3and BC2F2population in rice,Dongbei Nongye Daxue Xuebao(Journal of Northeast Agricultural U-niversity),45(9):9-17(邹德堂,刘忠良,赵宏伟,孙健,郑洪亮,陈宾宾,刘博文,2014,利用F2:3和BC2F2群体定位水稻粒型和粒重QTL,东北农业大学学报,45(9):9-17)
Fan C.C.,Xing Y.Z.,Mao H.L.,Lu T.T.,Han B.,Xu C.G.,Li X.H.,and Zhang Q.F.,2006,Gs3,a major QTL for grain length and weight and minor QTL for grain width and thickness in rice,encodes a putative transmembrane protein,Theor.Appl.Genet.,112(6):1164-1171
Fu F.H.,Wang F.,Huang W.J.,Peng H.P.,Wu Y.Y.,and Huang D.J.,1994,Genetic analysis on grain characters in hybrid rice,Zuowu Xuebao(Acta Agronomica Sinica),1(7):39-45(符福鸿,王丰,黄文剑,彭惠普,伍应运,黄德娟,1994,杂交水稻谷粒性状的遗传分析,作物学报,1(7):39-45)
Jiang Y.D.,Luo J.T.,Kuang H.C.,Yang Y.,Chen G.Z.,He X.C.,Fu J.,Zheng J.,and Zeng Z.M.,2015,Identification of QTLs for grain shape and weight using F2generation of two Indica rice varieties,Fenzi Zhiwu Yuzhong(Molecular Plant Breeding),13(8):1689-1694(蒋钰东,罗俊涛,况浩池,杨扬,陈光珍,何兴财,付均,郑军,曾正明,2015,利用两个籼稻杂交F2定位水稻粒型,粒重,分子植物育种,13(8):1689-1694)
Li J.X.,Yu S.B.,Xu C.G.,Tan Y.F.,Gao Y.J.,Li X.H.,and Zhang Q.F.,2000,Analyzing quantitative trait loci for yield using a vegetatively replicated F2population from a cross between the parents of an elite rice hybrid,Theor.Appl.Genet.,101(1-2):248-254
Li Q.,Li L.,Yang X.H.,Warburton M.L.,Bai G.H.,Dai J.R.,and Li J.S.,2010,Relationship,evolutionary fate and function of two maize co-orthologs of rice GW2 associated with kernel size and weight,BMC Plant Biology,10(1):143
Li Y.B.,Fan C.C.,Xing Y.Z.,Jiang Y.H.,Luo L.J.,Sun L.,Shao D.,Xu C.J.,Li X.H.,Xiao J.H.,He Y.Q.,and Zhang Q.F.,2011,Natural variation in GS5 plays an important role in regulating grain size and yield in rice,Nat.Genet.,43(12):1266-1269
Lin H.X.,Min S.K.,Xiong Z.M.,Qian H.R.,Zhuang J.Y.,Lu J.,Zheng K.L.,and Huang N.,1995,RFLP mapping of QTLs for grain shape traits in indica rice(Oryza sattve L.subsp.indica),Zhongguo Nongye Kexue(Scientia Agricultura Sinica),28(4):1-7(林鸿宣,闵绍楷,熊振民,钱惠荣,庄杰云,陆军,郑康乐,黄宁,1995,应用RFLP图谱定位分析籼稻粒型数量性状基因定位,中国农业科学,28(4):1-7)
Lin L.H.,and Wu W.R.,2003,Mapping of QTLs underlying grain shape and grain weight in rice,Fenzi Zhiwu Yuzhong(Molecular Plant Breeding),1(3):337-342(林荔辉,吴为人,2003,水稻粒型和粒重的QTL定位分析,分子植物育种,1(3):337-342)
Liu D.,Wang J.Y.,Chai Y.S.,Sun Y.Y.,Sun G.H.,Feng Z.L.,and Chen W.F.,2015,Mapping of the QTLs for rice grain shape using recombinant inbred lines of Indica-Japonica hybridization in rice,Shenyang Nongye Daxue Xuebao(Journal of Shenyang Agricultural University),46(4):385-390(刘丹,王嘉宇,柴永山,孙玉友,孙国宏,冯章丽,陈温福,2015,利用籼粳交RILs群体的水稻粒形QTL定位,沈阳农业大学学报,46(4):385-390)
Ma M.L.,Liu Y.H.,Jiang L.,Lei E.,Su Y.L.,and Lu B.Y.,2015,QTLs mapping of grain length-width ration by using F2population in rice,Jiangxi Nongye Daxue Xuebao(Acta Agriculturae Universitatis Jiangxiensis),37(2):260-263(马孟莉,刘艳红,江玲,雷恩,苏一兰,卢丙越,2015,利用F2群体定位水稻谷粒长宽比QTL,江西农业大学学报,37(2):260-263)
Qi P.,Lin Y.S.,Song X.J.,Shen J.B.,Huang W.,Shan J.X.,Zhu M.Z.,Jiang L.W.,Gao J.P.,and Lin H.X.,2012,The novel quantitative trait locus GL3.1 controls rice grain size and yield by regulating Cyclin-T13,Cell Research,22(12):1666-1680
Sun B.,Zhan X.D.,Lin Z.C.,Gao Z.Q.,Yu P.,Liu Q.E.,Shen X.H.,Zhang Y.X.,Chen D.B.,Cheng S.H.,and Cao L.Y.,2015,Correlation analysis and QTL mapping of grain shape and grain weight in rice,Fenzi Zhiwu Yuzhong(Molecular Plant Breeding),13(12):2663-2672(孙滨,占小登,林泽川,高志强,于萍,刘群恩,沈希宏,张迎信,陈代波,程式华,曹立勇,2015,水稻粒形和粒重性状的相关性分析及QTL定位,分子植物育种,13(12):2663-2672)
Wang S.K.,Li S.,Liu Q.,Wu K.,Zhang J.Q.,Wang S.S.,Wang Y.,Chen X.B.,Zhang Y.,Gao C.X.,Wang F.,Huang H.X.,and Fu X.D.,2015,The OsSPL16-GW7 regulatory module determines grain shape and simultaneously improves rice yield and grain quality,Nature Genetics,47(8):949-954
Weng J.F.,Gu S.H.,Wan X.Y.,Gao H.,Guo T.,Su N.,Lei C.L.,Zhang X.,Cheng Z.J.,Guo X.P.,Wang J.L.,Jiang L.,Zhai H.Q.,and Wan J.M.,2008,Isolation and initial characterization of GW5,a major QTL associated with rice grain width and weight,Cell Res.,18(12):1199-1209
Xia D.,Zhou H.,Liu R.J.,Dan W.H.,Li P.B.,Wu B.,Chen J.X.,Wang L.Q.,Gao G.J.,Zhang Q.L.,and He Y.Q.,2018,GL3.3,a Novel QTL encoding a GSK3/SHAGGY-like kinase,epistatically interacts with GS3 to produce extra-long grains in rice,Molecular Plant,11(5):754-756
Xiao J.H.,and Zhang Q.F.,2010,Functional genomics research and green super rice,Fenzi Zhiwu Yuzhong(Molecular Plant Breeding),8(6):1050-1053(肖景华,张启发,2010,功能基因组与绿色超级稻,分子植物育种,8(6):1050-1053)
Xu J.J.,Wang B.H.,Wu Y.H.,Du P.N.,Wang J.,Wang M.,Yi C.D.,Gu M.H.,and Liang G.H.,2011,Fine mapping and candidate gene analysis of ptgms2-1,the photoperiod-thermo-sensitive genic male sterile gene in rice(Oryza sativa L.),Theor.Appl.Genet.,122(2):365-372
Xu J.J.,Zhao Q.,Tang Z.X.,Zhao Y.F.,Zhu L.,Xu C.W.,Gu M.H.,Han B.,and Liang G.H.,2011,Mapping of QTLs for grain shape using whole-genome sequenced chromosome segment substitution lines in rice,Zhongguo Nongye Kexue(Scientia Agricultura Sinica),25(4):365-369(徐建军,赵强,汤在祥,赵元凤,朱磊,徐辰武,顾铭洪,韩斌,梁国华,2011,利用重测序的染色体片段代换系群体定位水稻粒型QTL,中国水稻科学,25(4):365-369)
Yang L.S.,Bai Y.S.,Zhang P.J.,Xu C.W.,Hu X.M.,Wang W.M.,She D.H.,and Chen G.Z.,2001,Studies for the correlations between grain shape and rice quality,Zajiao Shuidao(Hybrid Rice),16(4):48-54(杨联松,白一松,张培江,许传万,胡兴明,王伍梅,佘德红,陈桂芝,2001,谷粒形状与稻米品质相关性研究,杂交水稻,16(4):48-54)
Yang W.N.,2011,Studies on digital technology of automatic extraction of parameters for rice yield related traits,Dissertation for Ph.D.,Huazhong University of Science and Technology,Supervisor:Liu Q.,pp.100-105(杨万能,2011,水稻产量相关性状参数自动提取的数字化技术研究,博士学位论文,华中科技大学,导师:刘谦,pp.100-105)
Yu B.,Nassirou T.Y.,Gao G.J.,and Zhang Q.L.,2012,QTLs analysis on appearance quality traits and kernel weight of rice,Hubei Nongye Kexue(Hubei Agricultural Sciences),51(19):187-192(于波,Nassirou T.Y.,高冠军,张庆路,2012,水稻外观品质性状和千粒重的QTL分析,湖北农业科学,51(19):187-192)
Zeng R.Z.,Talukdar A.,Liu F.,and Zhang G.Q.,2006,Mapping of the QTLs for grain shape using single segment substitu tion lines in rice,Zhongguo Nongye Kexue(Scientia Agricultura Sinica),39(4):647-654(曾瑞珍,Talukdar A.,刘芳,张桂权,2006,利用单片段代换系定位水稻粒形QTL,中国农业科学,39(4):647-654)
Zhang Y.D.,Zhang Y.H.,Dong S.L.,Chen T.,Zhao Q.Y.,Zhu Z.,Zhou L.H.,Yao Z.,Zhao L.,Yu X.,and Wang C.L.,2013,Identification of QTL for rice grain traits based on an extralarge grain material,Zhongguo Shuidao Kexue(China Journal Rice Science),27(2):122-128(张亚东,张颖慧,董少玲,陈涛,赵庆勇,朱镇,周丽慧,姚姝,赵凌,于新,王才林,2013,特大粒水稻材料粒型性状的QTL检测,中国水稻科学,27(2):122-128)
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