稻米品质性状的QTL定位及遗传分析
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摘要
稻米品质包括加工品质、外观品质、蒸煮与食味品质和营养品质等,一般消费者偏向于关注外观品质、蒸煮与食味品质以及营养品质。传统育种对由多基因控制的数量性状的改良存在许多弊端,如性状不稳定,因此达到对这些性状改良的目的比较困难。本研究就是为充分利用现有的科技手段改良稻米品质提供一些参考信息。
     本研究利用日本优质粳稻品种越光和国内著名高产籼稻品种南京11构建的重组自交系(RIL Recombinant inbred lines)群体,在2006和2007两年间连续考察了稻谷和糙米的粒形(粒长、粒宽和长宽比)、千粒重、垩白米率、粗脂肪含量、总蛋白含量以及直链淀粉含量等11项稻米品质性状,结合分子数据从表型到遗传基础进行分析,将控制稻米品质性状的遗传基础分解为加性QTL、上位性QTL以及QTL与环境的互作。为下一步分子标记辅助选择改良稻米品质提供相关信息。
     结果如下:
     1.利用QTLNetWork软件对11项稻米主要性状进行分析,共检测到27个加性QTL,分别位于除第7、9、10和11染色体以外的8条染色体上;还检测到16对非加性上位性互作效应QTL。本研究所检测到27个加性QTL中有8个为前人曾经检测到在同一区间;19个为新的加性QTL,包括:控制稻谷粒长的qGL-3、qGL-4、qGL-6和qGL-12;控制稻谷粒宽的qGW-1、qGW-2和qGW-5;控制稻谷长/宽的qLWR-2和qLWR-3;控制糙米粒长的qBRL-3-1、qBRL-3-9和qBRL-4;控制糙米粒宽的qBRW-1;控制糙米长/宽的qLWRBR-2-11和qLWRBR-4;控制千粒重的qTSW-4和qTSW-5;控制垩白米率的qPGWC-6;控制脂肪含量的qRFC-12。
     2.稻米品质性状的加性QTL、上位性QTL以及QTL与环境的互作分析表明:稻谷粒长、稻谷粒宽、稻谷长/宽、糙米粒长、糙米粒宽、糙米长/宽、垩白米率、千粒重和脂肪含量主要受加性QTL控制外,还受到上位性QTL影响;蛋白质含量主要受加性QTL控制;直链淀粉含量主要受控于Wx基因,同时受微效位点的修饰外,还受到上位性QTL影响。
     3.在本研究所的结果中,普遍存在着QTL多效性:稻谷粒宽和糙米粒宽位于第1染色体上的QTL位点相同;稻谷粒宽、糙米粒宽和糙米长/宽分位于第2染色体上的QTL位点相同:稻谷长/宽、糙米粒宽和糙米长/宽位于第2染色体上的QTL位点相同;稻谷粒长和糙米粒长位于第3染色体上的QTL位点相同,稻谷长/宽和糙米粒长位于第3染色体上的QTL位点相同;稻谷粒长、糙米粒长、糙米长/宽和千粒重位于第4染色体上的QTL位点相同;稻谷粒长、垩白米率和直链淀粉含量位于第6染色体上的QTL位点相同。
The demand for superior grain quality is increasingly becoming a priority for international export markets in all of the cultivated rice-producing areas worldwide.The primary components of rice grain quality include appearance,eating,cooking,and milling quality,and nutritional qualities,all values that are determined by their physical-chemical properties and other socio-cultural factors.Among these quality properties,consumers pay more attention to the fine appearance,high eating and nutritional quality.Therefore,these are major goals in rice quality improvement.In order to characterized the genetic components,including additive effects,epistasis,and QTL-by-environment interaction,that are involved in the control of grain quality,data were collected from replicated tests of a recombinant inbred line population derived from the cross between Koshihikari,a top elite japonica variety with superior grain quality,and Nanjing11,a high-yield indica variety with poor grain quality,in two successive years.The results are as follows:
     1.The RIL population showed bimodal segregations for amylase content,indicating the involvement of a major gene for it;whereas,normal distributions of the others,indicating qualititative-inherited.A total of 27 QTL were identified with significant additive effects on nine chromosomes except chromosomes 7、9、10 and 11.Although only qAC-6 for amylase content had the largest effects.Epistasis analysis detected 16 significant additive-by-additive interactions on the traits.For grain length(GL),grain width(GW),grain length-width ratio(GLWR),grain length of rice seed of brown rice(MGL),grain width of rice seed of brown rice(BRW),grain length-width ratio of brown rice(LWRBR), thousand-seed weight(TSW),percentage of grains with chalkiness(PGWC),rice fat content(RFC),amylose content(AC).Only one additive QTL,qRPC-4,interacted with the environments for rice protein content(RPC).Compared with QTL previously reported for these traits,eleven additive QTL,namely qGL-3、qGL-4、qGL-6、qGL-12 qGW-1、qGW-2、qGW-5、qLWR-2、qLWR-3、qBRL-3-1、qBRL-3-9、qBRL-4、qBRW-1、qLWRBR-2-11、qLWRBR-4、qTSW-4、qTSW-5、qPGWC-6 and qRFC-12,are novel loci.Interestingly,the QTL qRFC-6a for RFC with the highest LOD score was located in the interval RM190-RM588 on chromosome 6,which corresponded to the waxy gene region.
     2.Based on the dissection of genetic basis in controlling grain quality,it revealed that grain length of rice seed of milled rice,grain width of rice seed of brown rice and fat content were mainly determined by additive QTL;additive effect and epistatic effect were the main genetic components for length-width ratio and thousand-seed weight rice;and amylose content was controlled by Wx and multiple minor loci.
     3.QTL multipurpose is normal in this research.We found 6 pairs QTLs there are locate the same place as follows:grain length and grain width of rice seed of brown rice each has the QTL of chromosome 1;grain width,grain width of rice seed of brown rice and grain length-width ratio of brown rice all have the same QTL of chromosome 2;grain length-width ratio,grain width of rice seed of brown rice and grain length-width ratio of brown rice all have the same chromosome 2 QTL;there is the same QTL of chromosome 3 for grain length and grain length of rice seed of brown rice;the same QTL is found in chromosome 3 for grain length-width ratio and grain length of rice seed of brown rice;in those QTLs,the same QTL is in the chromosome 4,and they are grain length,grain length of rice seed of brown rice,grain length-width ratio of brown rice and thousand-seed weight; and the last same QTL is in chromosome 6,the owner are grain length,percentage of grains with chalkiness and amylose content.
引文
包劲松,夏英武.稻米淀粉RVA谱的基因型/环境互作效应分析.中国农业科学,2001,34(2):123-127
    陈葆棠,彭仲明.水稻糊化温度的遗传分析.华中农业大学学报,1992,11(2):115-119
    陈秉发,陈建民,黄荣裕,等.稻米垩白遗传及无垩白稻谷单粒筛选技术初探.福建农业学报,2000,15(4):1-5
    陈建国.籼粳亚种间杂交水稻品质性状的遗传研究.浙江农业大学博士学位论文,1996
    陈建国,宋国清,瞿绍洪,等.杂交早稻米质性状的直接和母体遗传效应分析.中国水稻科学,1998,12(2):79-84
    陈建国,朱军.籼粳杂交稻米外观品质性状的遗传及基因型×环境互作效应研究.中国农业科学,1998,31(4):1-7
    郭益全,谢顺景.籼稻烹调与食用品质及谷粒性状之遗传.中华农业研究,1985,34(30):243-257
    郭银燕,张云康,陈润兴,等.早籼稻加工品质品种、地点、品种×地点互作效应的研究.遗传,1997,19(4):12-16
    郭银燕,张云康,杨尧城,等.浙江省早籼稻外观品质品种、地点、品种×地点互作效应研究.浙江农业大学学报,1998,24(1):20-26
    郝伟,金健,孙世勇,等.覆盖野生稻基因组的染色体片段替换系的构建及其米质相关数量性状基因座位的鉴定.植物生理与分子生物学学报,2006,32(3):354-362
    何光华,袁祚廉,郑家奎,等.水稻籽粒蛋白质、游离氨基酸含量的配合力与杂种优势分析.作物学报,1996,22(2):192-196
    何予卿,邢永忠,葛小佳,等.水稻米饭延伸指数相关性状的基因定位研究.分子植物育种,2003,1(5/6):616-619
    黄英金,刘宜柏,饶治祥,等.几个名优水稻品种特异品质性状的配合力分析.江西农业大学学报,1995,17(4):361-367
    黄祖六,谭学林,徐辰武,等.稻米胶稠度基因位点的标记和分析.中国农业科学,2000,33(6):1-5
    金庆生,裘伯钦,严文潮,等.Tagging of a gene for aroma iIl rice by RAPD and RFL P.浙江农业学报,1995,7(6):437-442
    李晨,潘大建,孙传清,等.水稻糙米高蛋白基因的QTL定位.植物遗传资源学报,2006,7(2): 170-174
    李仕贵,黎汉云,周开达,等.杂交水稻稻米外观品质性状的遗传相关分析.西南农业大学学报,1995,17(3):197-201
    李贤勇,何永歆,李顺武,等.水稻千粒重与垩白粒率的相关性分析.西南农业学报,2003,16(4):20-23
    李贤勇,王天凯,王楚桃.稻米蒸煮品质与营养品质的相关性分析.西南农业学报,2001,14(3):21-24
    李欣,陈宗祥,顾兴友,等.水稻香味基因的染色体定位研究.江苏农学院学报,1995,16(2):31-38
    李欣,顾铭洪,潘学彪.常见水稻品种稻米品质的研究.扬州大学学报,1987,8(1):1-8
    李欣,莫惠栋,王安民.粳型杂交稻米品质性状的遗传表达.中国水稻科学,1999,16(4):197-204
    李欣,汤述翥,陈宗祥,等.粳稻米糊化温度的遗传研究.江苏农学院学报,1995,16(1):15-20
    李欣,汤述翥,印志同,等.粳型杂种稻米品质性状的表现及遗传控制.作物学报,2000,26(4):411-419
    李欣,顾铭洪,程祝宽,等.水稻米香基因的染色体定位研究.江苏农学院学报,1996,17(3):33-39
    李泽福,万建民,夏加发,等.水稻外观品质的数量性状基因位点分析.遗传学报,2003,30(3):251-259
    梁敬琨,梁承邺,张隆芬,等.籼型杂交稻品质性状与其亲本关系的初步研究.中国水稻科学,1989,2(3):111-116
    林鸿宣,闵绍楷,熊振民,等.应用RFLP图谱定位分析籼稻粒形数量性状基因座位.中国农业科学,1995,28(4):1-7
    林建荣,吴明国,石春海.粳型杂交稻稻米外观品质性状的遗传效应研究.中国水稻科学,2001,15(2):93-96
    林荔辉,吴为人.水稻粒形和粒重的QTL定位分析.分子植物育种,2003,1(3):337-342
    罗玉坤,朱智伟,陈能,等.中国主要稻米的粒形及其品质特性.中国水稻科学,2006,18(2):165-169
    汤圣祥.我国杂交水稻蒸煮与食用品质的研究.中国农业科学,1987,20(5):17-22
    凌兆凤,莫惠栋,顾铭洪,等.籼稻米糊化温度的遗传研究.谷类作物品质性状遗传研究进展,江苏科技出版社,1990,106-112
    刘宜柏,黄英金.稻米食味品质的相关性研究.江西农业大学学报,1989,4:55-59.
    倪玲娟,全立勇,袁勤,等.杂交粳稻米质性状遗传效应.上海农业学报,1996,12(2):16-17
    祁祖白,李宝健,杨广文,等.水稻籽粒外观品质及脂肪的遗传研究.遗传学报,1983,10(6):452-458
    芮重庆,赵安常.籼稻粒重与粒形性状F_1遗传特性的双列分析.中国农业科学,1983,5:14-20
    沈圣泉,庄杰云,王淑珍,等.水稻米粒延伸性QTLs定位和基因型与环境互作分析.中国水稻科学,2007,19(4):319-322
    石春海,何慈信,朱军.稻米加工品质性状遗传效应及其与环境互作的遗传分析.遗传学报,1998,25(1):46-53
    石春海,何慈信,朱军,等.籼稻稻米外观品质性状的遗传主效应和环境互作效应分析.中国水稻科学,1999,16(3):179-182
    石春海,吴建国,樊龙江,等.不同环境下粕稻糙米重的发育遗传研究.植物学报,2001,43(6):603-609
    石春海,朱军.籼稻稻米外观品质与其它品质性状的相关性分析.浙江农业大学学报,1994,20(6):606-610
    石春海,朱军.籼稻稻米蒸煮品质的稻谷和母体遗传效应分析.中国水稻科学,1994,8(3):129-164.
    石春海.朱军.稻米营养品质稻谷效应和母体效应的遗传分析.遗传学报,1995,22(5):372-379
    汤圣祥,Khush GS.籼稻胶稠度的遗传.作物学报,1993,19(2):119-124
    汤圣祥,张云康,余汉勇.籼粳杂交稻米胶稠度的遗传.中国农业科学,1996,29(5):51-55
    汤圣祥,张云康,余汉勇.籼粳杂交稻米胶稠度的遗传特性分析.中国水稻科学,1996,10(4):197-200
    万向元,胡培松,王海莲,等.水稻品种直链淀粉含量、糊化温度和蛋白质含量的稳定性分析.中国农业科学,2007,38(1):1-6
    吴长明,孙传清,陈亮,等.控制稻米脂肪含量的QTLs分析.农业生物技术学报,2000,8(4):382-384
    吴长明,孙传清,陈亮,等.水稻直链淀粉含量与籼粳分化度的QTL及其相互关系研究.中国农业大学学报,2000,5(5):6-11
    吴长明,孙传清,付秀林,等.稻米品质性状与产量性状及籼粳分化度的相互关系研究.作物学报,2003,29(6):822-828
    汪斌,兰涛,吴为人.应用微卫星图谱定位稻米性状的QTL.福建农业学报,2003,18(1):11-15
    王余龙,姚友礼,李昙云,等.水稻籽粒品质性状与粒重关系的初步探讨.作物学报,1995,21(5):573-578
    武小金.稻米蒸煮食用品质性状的遗传研究.湖南农学院学报,1989,15(4):6-10
    邢永忠,谈移芳,徐才国,等.利用水稻重组自交系群体定位谷粒外观性状的数量性状基因.植物学报,2001,43(8):840-845
    徐辰武,张爱红.籼粳杂交稻米品质性状的遗传分析.作物学报,1996,22(5):530-534
    徐辰武,张爱红.几个籼粳交组合稻米品质性状遗传表达的鉴定.中国水稻科学,1998,12(1):51-54
    徐建龙,薛庆中,罗利军,等.水稻粒重及其相关性状的遗传剖析.中国水稻科,2002,16(1):6-10
    严长杰,梁国华,陈峰,等.利用籼粳回交群体分析水稻粒形性状相关QTLs.遗传学报,2003,30(8):711-716
    严长杰,徐辰武,裔传灯,等.利用SSR标记定位水稻糊化温度的QTLs.遗传学报,2001,28(11):1006-1011
    杨连松,白一松,张培江,等.水稻谷粒形状分类及其与稻米品质相关性研究.杂交水稻,2001,5
    杨仁崔,梁康迳,陈青华.稻米垩白直感遗传和杂交稻垩白米遗传分析.福建农林大学学报,1986,15(1):51-54
    易小平,陈芳远.籼型杂交水稻蒸煮品质、加工品质及营养品质的细胞质遗传效应.中国水稻科学,1992,6(4):187-189
    于永红,朱智伟,樊叶杨,等.应用重组自交系群体检测控制水稻糙米粗蛋白和粗脂肪含量的QTL.作物学报,2006,32(11):1712-1716
    袁玲,祝莉莉,何光存.稻米品质性状基因的SSR标记定位.武汉大学学报(理学报),2002,48(4):507-510
    甄海,吴东辉,伍时照,等.广东粕稻品种品质性状的相关与通径分析.华南农业大学学报,1996,17(4):41-45
    张光恒,曾大力,郭龙彪,等.水稻米粒延伸性的遗传剖析.遗传,2006,26(6):887-892
    张建军,李天,王勇.引进国外水稻品种稻米品质性状的相关性研究.陕西农业科学,2006,(3):18-21
    张小明,石春海,鲍根良,等.稻米外观和碾磨性状的数量遗传及相关性研究进展.浙江农业学报,2002,14(3):182-186
    张云康,林榕浑,闵捷,等.浙江水稻品种资源的品质研究.中国种业,1992,4:23-25
    朱碧岩,贾志宽.水稻品质性状遗传参数的分析.西北农林科技大学学报,1990,18(3):69-7
    李泽福,万建民,夏加发,翟虎渠,等.水稻外观品质的数量性状基因位点分析.遗传学报,Acta Genetica Sinica,March 2003,30(3):251-259
    万向元,刘世家,等.利用CSSLs群体研究稻米粒型QTL的表达稳定性.遗传学报,Acta Genetica Sinica,November 2004,31(11):1275-1283
    何平,李仕贵,等.影响稻米品质几个性状的基因座位分析.科学通报,1998年8月第43卷,第16期
    曾大力,钱前,阮刘青,滕胜,等.稻米垩白三维切面的遗传分析.中国水稻科学(Ch inese J R ice S ci),2002,16(1):11-14
    邢永忠,谈移芳,徐才国,等.利用水稻重组自交系群体定位谷粒外观性状的数量性状基因.植物学报,2001,43(8):840-845
    李合松,等.不同栽插方式对米粉稻生长发育和产量及稻米品质的影响.湖南农业大学学报(自然科学版),第33卷第4期Vol.33,No.4 Journal of Hunan Agricultural University(Natural Sciences)
    王红梅,刘巧泉,等.稻米蛋白营养品质及其遗传改良.植物生理学通讯,第43卷第2期
    胡孔峰,杨泽敏,雷振山,等.中国稻米品质研究的现状与展望.Chinese Agricultural Science Bulletin Vol.22 No.1 2006
    翁建峰,万向元,吴秀菊,王海莲,翟虎渠,万建民,等.利用CSSL群体研究稻米AC和PC相关QTL表达稳定性.作物学报,ACTA AGRONOMICA SINICA,第32卷第1期,Vol132,No11ppl14-19,Janl,2006
    王海莲,万向元,胡培松,翟虎渠,万建民,等.稻米脂肪含量近红外光谱分析技术研究.中国农业科学,2005,38(8):1540-1546,Scientia Agricultura Sini
    Ahn SN,Bollich CN,McClung AM,et al.RFLP analysis of genomic regions associated with cooked-kernel elongation in rice.Theor Appl Genet,1993,87:27-32
    Ahn SN,Bollich CN,Tanksley SD.RFLP tagging of a gene for aroma in rice.Theor Appl Genet,1992,84:825-828
    Allard RW.Formulas and tables to facilitate the calculation of recombination values in heredity.Hilgardia,1956,24:235-278
    Aluko G,Martinez C,Tohme J,et al.QTL mapping of grain quality traits from the interspoeific cross Oryza sativa × O.glaberrima.Theor Appl Genet,2006,109:630-639
    Bao JS,Harold C,He P,et al.Analysis of quantitative trait loci for starch properties of rice based on an RIL population.Acta Botanica Sinica,2003,45(8):986-994
    Bao JS,Wu YR,Hu B,et al.QTL for rice grain quality based on a DH population derived from parents with similar apparent amylose content.Euphytica,2002,128:317-324
    Bether DK,Hoff BJ.Inheritance of scent in American long grain rice.Crop Sci,1986,26:876-878
    Bollich CN,Webb BD.Inheritance of amylose in two hybrid populations of rice.Cereal Chem,1973,50:631-636
    Bradbury,Louis MT,Fitzgerald,et al.The gene for fragrance in rice.Plant Biotechnolo J,2007,3(3):363-371
    Chang TT,Somrith B.Genetic studies on the grain quality of rice.In:Chemical aspects of rice quality.IRRI,Los Banos,Philippines,1979,49-58
    Chang WL,Li WY.Inheritance of amylose content and gel consistency in rice.Botanical Bulletin of Academia Sinica,1981,22(1):35-47
    Chen N,Zhu ZW,Zhang BP,et al.Relationship between the eating quality and the physico chemical properties of high grain quality rice in China.Chinese Rice Research Newsletter,1997,5(1):7-8
    Chen X,Temnykh S,McCouch SR,et al.Development of a mierosatellite framework mapproviding genome-wide coverage in rice(Oryza sativa L.).Theor Appl Genet,1997,95:553-567
    Cordeiro GM,Christopher MJ,Henry RJ,et al.Identification of microsatellite markers for fragrance in rice by analysis of the rice genome sequence.Mol Breed,2002,9(4):245-250
    Dellaporta SL,Wood J,Hicks JB.A plant DNA minipreparation:Version Ⅱ.Plant Mol Biol Rep,1983,1:19-21
    Dhulappanavar CV.Inheritance of scent in rice.Euphytica,1976,25:659-662
    Fan CC,Xing YZ,Mao HL,et al.GS3,a major QTL for grain length of rice seed and weight and minor QTL for grain width of rice seed and thickness in rice,encodes a putative transmembrane protein.Theor Appl Genet,2006,112:1164-1171
    Fan CC,Yu XQ,Xing YZ,et al.The main effects,epistatic effects and environmental interactions of QTLs on the cooking and eating quality of rice in a doubled-haploid line population.Theor Appl Genet,2007,110:1445-1452
    Garland S,Lewin L,Blakeney AR,et al.PCR - based molecular markers for the fragrance gene in rice (Oryza sativa L.).Theor Appl Genet,2000,101:364-371
    Gravois KA,Moldenhauer KAK,Rohman PC.Genetic and genotype × environment effects for rough rue and head rice yield.Crop Sci,1991,31:907-911
    He P,Li SG,Qian Q,et al.Genetic analysis of rice grain quality.Theor Appl Genet,1999,98:502-508
    Heda GD,Reddy GM.Studies on protein content and yield levels in rice.Theor Appl Genet,1984,69:93-95
    Holguin VJE,Martinez RCP,Cuevas PF,et al.Effect of cytoplasm on the expression of white center and gelatinization temperature in rice(Oryza sativa L).Acta Agronomica,Universidad Nacional de Colobia.1995,45(2-7):7-16
    Hu ZL,Li P,Zhou MQ,et al.Mapping of quantitative trait loci(QTLs) for rice protein and fat content using doubled haploid lines.Euphytica,2006,165:47-54
    Huang N,ParcoA,Mew T,et al.RFLP mapping of isozymes,RAPD and QTLs for grain shape,brown planthopper resistance in a doubled haploid rice population.Mol Breed,1997,3:105-116
    Hussaln AA,Maurya DM,Vaish CP.Study on quality status of indigenous upland rice.Indian J Genet,1987,47(2):145-152
    Jin QS,Waters D,Cordeiro GM,et al.A single nucleotide polymorphism(SNP) marker linked to the fragrance gene in rice(Oryz a sativa L.).Plant Sci,2003,165:359-364
    Juliano BO.Rice chemistry and technology,edn.American Association of Cereal Chemists,Inc,St.Paul, Minn,1985
    Kang HJ,Cho YG,Lee YT,Kim YD,Eun MY,Shim JU.QTL mapping of genes related with grain chemical properties based on molecular map of rice.Korean J Crop Sci,1998,43(4):199-204
    Kaw RN,De La Cruz NM.Interrelations amonn physicochemical main quality characters in rice.J Genet & Breed,1990,44:169-142
    Kongseree N,Juliano B.O.Physico-chemical properties of rice grain and starch from lines differing in amylose content and gelatinisation temperature.J Agr Food Chem,1972,20(3):714-716
    Khush GS.Grain quality of hybrid rice.In:Hybrid rice IRRI,Manila,Philippine,1988,201-205
    Kubo T,Takano-kai N,Yoshimura A.RFLP mapping of genes for long kernel and awn on chromosome 3in rice.Rice Genet Newsl,2001,18:26-28
    Kumar I,and Khush GS.Genetic analysis of different amylose levels in rice(Oryza Sativa L).Crop Sci,1987,1167-1172
    Kumar I,Khush GS.Inheritance of amyfose content in rice(Oryza sativa L.).Euphytica,1988,38:261-269
    Lanceras JC,Huang ZL,Naivikul O,et al.Mapping of genes for cooking and eating qualities in Thai jasmine rice(KDML105).DNA Res,2000,28:93-101
    Lander ES,Green P,Abrahamson J,et al.MAPMAKER:an interactive computer package for constructing primary genetic linkage map of experimental and natural populations.Genomics,1987,1:174-181
    Li JM,Xiao JH,Grandillo S,et al.QTL detection for rice grain quality traits using an interspecific backcross population derived from cultivated Asian(O.Sativa L.) and African(O.glaberrium S)rice.Genome,2006,47:697-704
    Li Z,Pinson SRM,Park WD,et al.Epistasis for three grain yield components in rice(Oryza sativa L.).Genetics,1997,145:453-465
    Li ZF,Wan JM,Xia JF,et al.Mapping of quantitative trait loci controlling physico-chemical properties of rice grains(Oryza sativa L.).Breed Sci,2003,53:209-215
    Li ZK.QTL mapping in rice:a few critical considerations.In:Khush GS,Brar DS,Hardy Beds.,Rice genetics Ⅳ Science Publishers,New Delhi,2001:153-171
    Lincoln SE,Daly MJ,Lander ES,et al.Constructing genetic linkage maps with MAPMAKER/EXP version 3.0:A tutorial and reference manual.A Whitehead Institute for biomedical research technical report 3hird edition.Cambridge,Mass,1993a.
    Lincoln SE,Daly MJ,Lander ES,et al.Mapping genes controlling quantitative traits using MAPMAKER/QTL version 1.1:A tutorial and reference manual.A Whitehead Institute for biomedical research technical report third edition.Cambridge,Mass,1993b
    Lorieux MM,Petrov N,Huang E,et al.Aroma in rice genetic analysis of a quantitative trait.Theor Appl Genet,1996,93:1145-1151
    McCouch SR,Kochert G,Tanksley SD,et al.Molecular mapping of rice chromosomes.Theor Appl Genet,1988,76:815-829
    Mckenzie,KS and Nutger J.Genetic analysis of amylose content,alkali spreading score,and grain dimensions in rice.Crop Sci,1983,23(2):306-316
    Michelmore RW,Paran I,Kesseli RV.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.Proc NatlAcadSci,1991,88:9828-9832
    Peng J,Ronin,Fahima T,et al.Domestication quantitative trait loci in Triticum dicoccoides,the progenitor of wheat.Proc Natl Acad Sci,2003,100:2489-2494
    Pinson SRM.Inheritance of aroma in six rice cultivars.Crop Sci,1994,34:1151-1157
    Pooni HS,Kumar ISH,Khush GS.Genetic control of amylose content in a diallel set of rice crosses.Heredity,1993,71:603-616
    Reddy GM,Sarala AK.Study on the amylose content and gelatinization temperature in certain local cultivars and induced grain shape mutants in rice.Euphytica,1979,28:665-674
    Reddy PR.Irheritance of aroma in rice.Indian J Genet Plant Breed,1994,40:327-327
    Redona ED,Mackill DJ.Quantitative trait locus analysis for rice panicle and grain characteristics.Theor Appl Genet,1998,96:957-963.
    Sanguinetti C J,Dias Neto E,Simpson AJG.Rapid silver staining and recover of PCR products separated on polyacrylamide gels.Biotechniques,1994,17:915-919.
    Septiningsih EM,Trijatmiko KR,Moeljopawiro S,et al.Identification of quantitative trait loci for grain quality in an advanced backcross population derived from the Oryza sativa variety IR64 and the wild relative O.rufipogon.Theor Appl Genet,2003,107:1433-1441
    Shenoy VV and Seshu DV.et aL Inheritance of protein per grain in rice.Indica J Genet,1991,51(2):214-220
    Shi CH,Xue JM,Yu YG,et al.Analysis of genetic effect on nutrient quality Waits in indica rice.Theor Appl Genet,1996,92:1099-1102
    Shi CH.Genetic and heterosis analysis for cooking quality traits of indica rice in different environments.Theor Appl Genet,1997,95:294-300
    Shubash KDG.and Mosina SB.Inheritance of the amylose content of the grain of rice hybrids.Rice Abstracts,1987,10(4):1471
    Singh NB and Sight HG.Gene action for quality components in rice.Indian J Agri Sci,1982 52(8):485-488
    Singh RK,Singh US,Khush GS.Genetics and biotechnology of quality traits in aromatic rices.In:Singh RK et al(ed) Aromatic Rice,Oxford&IBH publishing Co.Pvt.Ltd.,New Delhi,India,2000,p47-69
    Song XJ,Huang W,Shi M,et al.A QTL for rice grain width of rice seed and weight encodes a previously unknown RING-type E3 ubiquitin ligase.Nat Genet,2007,39(5):623-630
    Sood BC,Siddiq EA.Genetic analysis of crude protein content in rice.Indian J Agri Sci,1986,56(11):796-797
    Tan YF,Sun M,Xing YZ,et al.Mapping quantitative trait loci for milling quality,protein content and color characteristics of rice using a recombinant inbred line population derived from an elite rice hybrid.Theor Appl Genet,2001,103:1037-1045
    Tan YF,Li JX,Yu SB,et al.The three important traits for cooking and eating quality of rice grains are controlled by a single locus in an elite rice hybrid,Shanyou 63.Theor Appl Genet,1999,99:642-648
    Tan YF,Xing YZ,Zhang QF,et al.Quantitative genetic basis of gelatinization temperature of rice.Cereal Chem,2001,78(6):666-674
    Tan YF,Xing YZ,Li JX,et al.Genetic bases of appearance quality of rice grains in Shanyou 63,an elite rice hybrid.Theor Appl Genet,2000,101:823-829
    Tanksley SD,Grandillo S,Fulton MT,et al.Advanced back-cross QTL analysis in a cross between an elite processing line of tomato and its wild relative L.nimrrinellifolizan.Theor Appl Genet,1996,92:216-224
    Temnykh S,Park WD,MeCoueh SR,et al.Mapping and genome organization of mierosatellite sequence in rice(Oryza sativa L.).Theor Appl Genet,2000,100:697-712
    Thomson M,Tai T,McClung A,et al.Mapping quantitative trait loci for yield,yield components and morphological traits in an advanced backeross population between Oryza rufipogon and the Oryza sativa cultivar Jefferson.Theor Appl Genet,2003,107:479-493
    Tian R,Jiang GH,Shen LH,Wang LQ,He YQ.Mapping quantitative trait loci underlying the cooking and eating quality office using a DH population.Mol Breed,2007,15:117-124
    Tomar JB and Nanda JS.Genetic and Association studies of Kernel shape in rice.Indican J Genet,1985,45(2):278-283
    Tomar JB and Nanda JS.Genetics of geltinization temperature and its association with protein content in rice(Oryza sativa L.).Rice Abstracts,1984,7(7):1004
    Tan YF,Xing YZ,Li JX,et al.Genetic bases of appearance quality of rice grains in Shanyou 63,an elite rice hybrid.Theor Appl Genet,2000,101:823-829
    Tsuzuki E,shimokawa E.Inheritance of aroma in rice.Euphytica,1990,46:157-159
    Visscher,P.M.,R.Thomson,C.S.Haley.Confidence intervals in QTL mapping by bootstrapping.Genetics,1996,143:1016-1020
    Vivekanandan P,Girdharan S.Genetic variability and character association for kernel and cooking quality traits in rice.Oryza,1998,35(3):242-245
    Wan XY,Wan JM,Su CC,et al.QTL detection for eating quality of cooked rice in a population of chromosome segment substitution lines.Theor Appl Genet,2006,110(1):71-79
    Wan XY,Wan JM,Weng JF,et al.Stability of QTLs for rice grain dimension and endosperm chalkiness characteristics across eight environments.Theor Appl Genet,2007,110(7):1634-1646
    Wang,D,Zhu J,Li Z.Mapping QTLs with epistatic effects and QTL environment interaction by mixed linear model approaches.Thero Appl Genet,1999,99:1255-1264
    Webb BD.Rice quality and grades.In:Lub BS(ed).Rice Utilization.2nd edition.New York,1992,89-119
    Xian-Jun Song,Wei Huang,Min Shi,Mei-Zhen Zhu,et al.A QTL for rice grain width and weight encodes a previously unknown RiNG-type E3 ubiquitin ligase,nature genetics 2007
    Xing YZ,Tan YF,Xu CG;et al.Mapping quantitative trait loci for grain appearance traits of rice using a recombinant inbred line population.Acta Botanica Sinica,2001,43:840-845
    Yoshida S,Ikegami M,Kuze J,et al.QTL analysis for plant and grain characters of sake-brewing rice using a doubled haploid population.Breed Sci,2002,52:309-317
    Yu,SB,Li JX,Xu CG,et al.Importance of epistasis as the genetic basis of heterosis in an elite rice hybrid.Proc Natl Acad Sci,1997,94:9226-9231
    Zaman FU,Siddiq EA,Parsad AB.Genetical analysis of gel consistency in rice.(Oryza sativa L.) Indian J Genet Plant Breed,1985,45(1):111-118
    Naoko Fujita,Mayumi Yoshida et al.Characterization of SSⅢa-Deficient Mutants of Rice:The Function of SSⅢa and Pleiotropic Effects by SSⅢa Deficiency in the Rice Endosperm Plant Physiology,August 2007,Vol.144,pp.2009-2023,