利用RILs群体对水稻正季和再生季重要农艺性状的QTL比较分析
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摘要
水稻多数性状是由多基因控制的数量性状,对数量性状基因位点进行鉴定在作物遗传育种上具有重要意义。本研究以粳糯稻品种糯稻89-1为母本,籼稻品种蜀恢527为父本进行杂交,从F_2开始通过单粒传法获得重组自交系群体(RILs),2007年正季种植F_(2:6)群体,在当年正季考察了株高、有效穗、穗长等9个农艺性状,再生季考察了有效穗、穗长、再生力等8个农艺性状,此外,利用该群体还考察了水稻发芽率及芽期和再生稻桩耐寒性状,同时利用SSR标记构建遗传图谱和对这些性状进行QTL定位分析,得到的主要研究结果如下:
1.在RILs群体中,所考察的性状均表现为连续变异,且都接近正态分布,表明这些性状均为数量性状。方差分析表明,正季和再生季所有农艺性状株系间基因型差异显著或极显著。表明该重组自交系群体内株系间存在广泛的遗传变异。
2.对所考察的性状进行相关分析,发现多数性状存在相关性。在正季,株高与齐穗期、结实率与千粒重等分别显著正相关,株高、穗长、每穗总粒数、每穗实粒数四者彼此极显著正相关,着粒密度、每穗总粒数、每穗实粒数三者彼此极显著正相关,有效穗与穗长、每穗总粒数极显著负相关,千粒重与着粒密度极显著负相关。在再生季,再生力与每穗总粒数、着粒密度显著负相关,与有效穗极显著正相关,与芽期耐寒性显著正相关,其他农艺性状间的相关性与正季的表现规律相似。在两季均考察的7个性状中,同一性状两季间表现出极显著正相关,其中千粒重的相关系数最大,接近1;正季有效穗、每穗总粒数、着粒密度、每穗实粒数与再生季多数性状相关性达显著或极显著水平,而正季千粒重除与再生季千粒重和着粒密度相关性达极显著水平外,与其他再生季性状相关性不显著。
3.选用SSR引物对双亲进行多态性检测,有118对引物在双亲间表现多态性,用这些引物对169个株系进行检测,构建了一张包含105个分子标记、12个连锁群的连锁遗传图,覆盖了水稻基因组1144.1 cM,标记间的平均距离为10.90 cM,采用基于最小二乘的复合区间定位法对上述性状进行了QTL定位。
4.对各性状分别进行QTL定位分析,所考察的性状均检测到了相应的QTL。正季共检测到QTL 27个,分布在1、2、3、5、6、7、8、10染色体上,再生季共检测到QTL 27个,分布在1、2、3、4、5、6、7、10染色体上。对正季和再生季分别分析并比较两季检测到的QTL发现,在两季共同定位的7个农艺性状中能同时检测到的QTL共有11个,分布于第1、2、6、7、10染色体上。分别是穗长1个(qP16),每穗总粒数3个(qSnp1-1,qSnp1-2,qSnp6),着粒密度2个(qDp1-1,qDp1-2),千粒重5个(qKw2-1,qKw2-2,qKw2-3,qKw7-1,qKw10)。另外3个性状,即有效穗、每穗实粒数、结实率在两季中未检测到相同的QTL。两季中,千粒重检测到的QTL重复率最高,共检测到5个相同的QTL,且同一QTL在两季间贡献率大小表现稳定,表明千粒重性状表现受环境影响小。
本研究首次检测到2个新的控制再生力的QTL位点,分别是qRa4和qRa5,分别解释表型变异的8.17%和7.09%。
本研究首次检测到2个耐寒主效QTL。其中控制水稻芽期耐寒性的QTL 2个,即qCtb3和qCtb5,分别解释表型变异的75.57%和79.04%;检测到控制再生稻桩越冬耐寒性的主效QTL 1个(qCtr5),与qCtb5位于同一标记区间。此外,还检测到1个控制水稻发芽率的QTL(qGr10)。
5.发现有控制不同性状QTL定位在同一位点或区域。如在第1号染色体上的标记区间RM259～RM220内同一位点,同时定位到控制每穗总粒数和着粒密度的QTL(qSnp1-1(R)、qDp1-1(M)),在同一区间相邻位点定位到1个控制穗长的QTL(qPl1-1(R));在第6号染色体上的标记区间RM340～RM528内同一位点,同时定位到了控制株高、穗长和每穗总粒数的QTL(qPh6-2(M)、qP16(R)、qSnp6(R));在第7号染色体上的标记区间RM346～RM351内同一位点,同时定位到了控制每穗实粒数和结实率的QTL(qGpp7(M)、qSsr7(M))。这种控制不同性状的QTL定位在同一位置或区域,可能是QTL紧密连锁或一因多效。研究结果对指导水稻育种实践具有重要意义。
The most traits of crop are quantitative character controlled by more than one gene.It is very important for crop breeding to identify the quantitativetrait locus (QTL).This research used glutinous rice 89-1 as female parent,used Shuhui 527 as male parent,and got the F2 recombination inbred lines(RILs) population through the method of one seed reproduction,then plant them in 2007 suitable season,and inspected the plant height,the number of effective panicle,spike length and other six economical characters,then inspected the number of effective panicle,spike length, regeneration capacity and other five character in retooning rice.In additional,we also surveyed the germination percentage of rice and the cold-hardiness in the regeneration season.At the same time,we constructed the genetic map using SSR marker,and map the QTL character.The main results of the research as follow:
1.All the inspected trait showed continuous variation,and approach to normal distribution in RILs population,which explained the characters all are quantitative trait.The analysis of variance showed all the gene difference between the economical characters was significance or extremely significance.This suggested the wide range genetic variation between the RILs population
2.The correlation analysis of all inspected characters showed many characters are correlative.In the positive season,the plant height,economic panicle and the spike density were significant positive association,the plant height,spike length,the total number of every panicle,and the economical number of very spike were extremely significant positive association.The spike density,the total number of every panicle,and the economical number of every panicle,were extremely significant positive association.The number of productive spike,spike length and the total number of every panicle were extremely significant positive association.The economical number of every number setting percentage was extremely significant positive association.The kilo-grain weight and spike density were extremely significant negative association.The full panicle stage and the spike length were significant negative association.The full panicle stage,total number of every panicle, spike density and the economical number of every panicle,were extremely significant negative association.In the regeneration season,the capacity of regeneration,the total number of every panicle,spike density was extremely significant positive association. The setting percentage and the kilo-grains weight were significant positive association. Panicle lengths,the total number of every panicle and the spike density were extremely significant positive association.The total number of every panicle,the economical number and the spike density were extremely significant positive association.
3.We analyzed the polymorphism of parents using SSR primers.There are 118 pair primers presenting polymorphism between parents.We detected 169 strains using these primers,and then we constructed a genetic linkage map that containing 105 molecule markers and 12 linkage groups.This map covers 1144.1 cM of rice genome and the average length between markers is 10.90cM.All above results come from the least squared method of composite interval mapping(CIM).
4.Analyzing QTLs of corresponding traits,all detected traits can find QTLs. There are 27 QTLs distributing 1,2,3,5,6,7,8 and 10 chromosomes that detected in the positive season.In retooning season,27 QTL were still detected that distributing 1,2,3,4,5,6,7 and 10 chromosomes.Comparing two seasons' results,there are 11 QTLs distributing 1,2,6,7 and 10 chromosomes corresponding 7 traits such as Panicle length(qP16),total number of every panicle(qSnp1-1,qSnp1-2,qSnp6),spike density(qDp1-1,qDp1-2),kilo-grains weight(qKw2-1,qKw2-2,qKw2-3,qKw7-1, qKw10).Three other traits containing number of effective panicle,effective number of every panicle and maturing rate were not detected.In two seasons,because the recurrence rate of kilo-grains weight is highest,we think hereditary capacity of kilo-grains weight is very strong and is almost not affected by environment.
At the same time,we firstly' detected two QTLs of regenerative capacity(qRa4, qRa5) respectively offering 8.17%and 7.09%of phenotypic variation.
We also firstly detected two QTLs of cold resistance such as qCtb3 and qCtb5 regulating the cold resistance of rice shoot stage these respectively offer 75.57%and 79.04%of phenotypic variation,qCtr5 located the same marker with qCtb5 controlling cold resistance of overwintering rice stub.In addition,we detected a QTL of controlling percentage of germination(qGr10).
5.This research indicated some QTLs of different traits can be located in the same locus or area.For example,a locus between RM259-RM220 in 1 chromosome can control total number of every panicle(qSnp1-1(R)) and spike density (qDp1-1(M)).In this area,a QTL of Panicle length(qP11-1(R)) was also detected.A locus between RM340-RM528 in 6 chromosome can control total number of every panicle,panicle length and height(qPh6-2(M),qP16(R),qSnp6(R)).A locus between RM346-RM351 in 7 chromosome can control effective number of every panicle and maturing rate(qGpp7(M),qSsr7(M)).These results these being from pleiotropism or close linkage of QTLs could induce a huge improvement of rice breeding.
1.在RILs群体中,所考察的性状均表现为连续变异,且都接近正态分布,表明这些性状均为数量性状。方差分析表明,正季和再生季所有农艺性状株系间基因型差异显著或极显著。表明该重组自交系群体内株系间存在广泛的遗传变异。
2.对所考察的性状进行相关分析,发现多数性状存在相关性。在正季,株高与齐穗期、结实率与千粒重等分别显著正相关,株高、穗长、每穗总粒数、每穗实粒数四者彼此极显著正相关,着粒密度、每穗总粒数、每穗实粒数三者彼此极显著正相关,有效穗与穗长、每穗总粒数极显著负相关,千粒重与着粒密度极显著负相关。在再生季,再生力与每穗总粒数、着粒密度显著负相关,与有效穗极显著正相关,与芽期耐寒性显著正相关,其他农艺性状间的相关性与正季的表现规律相似。在两季均考察的7个性状中,同一性状两季间表现出极显著正相关,其中千粒重的相关系数最大,接近1;正季有效穗、每穗总粒数、着粒密度、每穗实粒数与再生季多数性状相关性达显著或极显著水平,而正季千粒重除与再生季千粒重和着粒密度相关性达极显著水平外,与其他再生季性状相关性不显著。
3.选用SSR引物对双亲进行多态性检测,有118对引物在双亲间表现多态性,用这些引物对169个株系进行检测,构建了一张包含105个分子标记、12个连锁群的连锁遗传图,覆盖了水稻基因组1144.1 cM,标记间的平均距离为10.90 cM,采用基于最小二乘的复合区间定位法对上述性状进行了QTL定位。
4.对各性状分别进行QTL定位分析,所考察的性状均检测到了相应的QTL。正季共检测到QTL 27个,分布在1、2、3、5、6、7、8、10染色体上,再生季共检测到QTL 27个,分布在1、2、3、4、5、6、7、10染色体上。对正季和再生季分别分析并比较两季检测到的QTL发现,在两季共同定位的7个农艺性状中能同时检测到的QTL共有11个,分布于第1、2、6、7、10染色体上。分别是穗长1个(qP16),每穗总粒数3个(qSnp1-1,qSnp1-2,qSnp6),着粒密度2个(qDp1-1,qDp1-2),千粒重5个(qKw2-1,qKw2-2,qKw2-3,qKw7-1,qKw10)。另外3个性状,即有效穗、每穗实粒数、结实率在两季中未检测到相同的QTL。两季中,千粒重检测到的QTL重复率最高,共检测到5个相同的QTL,且同一QTL在两季间贡献率大小表现稳定,表明千粒重性状表现受环境影响小。
本研究首次检测到2个新的控制再生力的QTL位点,分别是qRa4和qRa5,分别解释表型变异的8.17%和7.09%。
本研究首次检测到2个耐寒主效QTL。其中控制水稻芽期耐寒性的QTL 2个,即qCtb3和qCtb5,分别解释表型变异的75.57%和79.04%;检测到控制再生稻桩越冬耐寒性的主效QTL 1个(qCtr5),与qCtb5位于同一标记区间。此外,还检测到1个控制水稻发芽率的QTL(qGr10)。
5.发现有控制不同性状QTL定位在同一位点或区域。如在第1号染色体上的标记区间RM259～RM220内同一位点,同时定位到控制每穗总粒数和着粒密度的QTL(qSnp1-1(R)、qDp1-1(M)),在同一区间相邻位点定位到1个控制穗长的QTL(qPl1-1(R));在第6号染色体上的标记区间RM340～RM528内同一位点,同时定位到了控制株高、穗长和每穗总粒数的QTL(qPh6-2(M)、qP16(R)、qSnp6(R));在第7号染色体上的标记区间RM346～RM351内同一位点,同时定位到了控制每穗实粒数和结实率的QTL(qGpp7(M)、qSsr7(M))。这种控制不同性状的QTL定位在同一位置或区域,可能是QTL紧密连锁或一因多效。研究结果对指导水稻育种实践具有重要意义。
The most traits of crop are quantitative character controlled by more than one gene.It is very important for crop breeding to identify the quantitativetrait locus (QTL).This research used glutinous rice 89-1 as female parent,used Shuhui 527 as male parent,and got the F2 recombination inbred lines(RILs) population through the method of one seed reproduction,then plant them in 2007 suitable season,and inspected the plant height,the number of effective panicle,spike length and other six economical characters,then inspected the number of effective panicle,spike length, regeneration capacity and other five character in retooning rice.In additional,we also surveyed the germination percentage of rice and the cold-hardiness in the regeneration season.At the same time,we constructed the genetic map using SSR marker,and map the QTL character.The main results of the research as follow:
1.All the inspected trait showed continuous variation,and approach to normal distribution in RILs population,which explained the characters all are quantitative trait.The analysis of variance showed all the gene difference between the economical characters was significance or extremely significance.This suggested the wide range genetic variation between the RILs population
2.The correlation analysis of all inspected characters showed many characters are correlative.In the positive season,the plant height,economic panicle and the spike density were significant positive association,the plant height,spike length,the total number of every panicle,and the economical number of very spike were extremely significant positive association.The spike density,the total number of every panicle,and the economical number of every panicle,were extremely significant positive association.The number of productive spike,spike length and the total number of every panicle were extremely significant positive association.The economical number of every number setting percentage was extremely significant positive association.The kilo-grain weight and spike density were extremely significant negative association.The full panicle stage and the spike length were significant negative association.The full panicle stage,total number of every panicle, spike density and the economical number of every panicle,were extremely significant negative association.In the regeneration season,the capacity of regeneration,the total number of every panicle,spike density was extremely significant positive association. The setting percentage and the kilo-grains weight were significant positive association. Panicle lengths,the total number of every panicle and the spike density were extremely significant positive association.The total number of every panicle,the economical number and the spike density were extremely significant positive association.
3.We analyzed the polymorphism of parents using SSR primers.There are 118 pair primers presenting polymorphism between parents.We detected 169 strains using these primers,and then we constructed a genetic linkage map that containing 105 molecule markers and 12 linkage groups.This map covers 1144.1 cM of rice genome and the average length between markers is 10.90cM.All above results come from the least squared method of composite interval mapping(CIM).
4.Analyzing QTLs of corresponding traits,all detected traits can find QTLs. There are 27 QTLs distributing 1,2,3,5,6,7,8 and 10 chromosomes that detected in the positive season.In retooning season,27 QTL were still detected that distributing 1,2,3,4,5,6,7 and 10 chromosomes.Comparing two seasons' results,there are 11 QTLs distributing 1,2,6,7 and 10 chromosomes corresponding 7 traits such as Panicle length(qP16),total number of every panicle(qSnp1-1,qSnp1-2,qSnp6),spike density(qDp1-1,qDp1-2),kilo-grains weight(qKw2-1,qKw2-2,qKw2-3,qKw7-1, qKw10).Three other traits containing number of effective panicle,effective number of every panicle and maturing rate were not detected.In two seasons,because the recurrence rate of kilo-grains weight is highest,we think hereditary capacity of kilo-grains weight is very strong and is almost not affected by environment.
At the same time,we firstly' detected two QTLs of regenerative capacity(qRa4, qRa5) respectively offering 8.17%and 7.09%of phenotypic variation.
We also firstly detected two QTLs of cold resistance such as qCtb3 and qCtb5 regulating the cold resistance of rice shoot stage these respectively offer 75.57%and 79.04%of phenotypic variation,qCtr5 located the same marker with qCtb5 controlling cold resistance of overwintering rice stub.In addition,we detected a QTL of controlling percentage of germination(qGr10).
5.This research indicated some QTLs of different traits can be located in the same locus or area.For example,a locus between RM259-RM220 in 1 chromosome can control total number of every panicle(qSnp1-1(R)) and spike density (qDp1-1(M)).In this area,a QTL of Panicle length(qP11-1(R)) was also detected.A locus between RM340-RM528 in 6 chromosome can control total number of every panicle,panicle length and height(qPh6-2(M),qP16(R),qSnp6(R)).A locus between RM346-RM351 in 7 chromosome can control effective number of every panicle and maturing rate(qGpp7(M),qSsr7(M)).These results these being from pleiotropism or close linkage of QTLs could induce a huge improvement of rice breeding.
引文
[1]焦险峰,杨邦杰,裴志远.基于分层抽样的中国水稻种植面积遥感调查方法研究[J].农业工程学报,2006,(5)
[2]李仕贵.水稻几个重要性状的遗传分析和分子标记定位[D].四川农业大学博士论文,1998
[3]钱前,程式华.水稻遗传学和功能基因组学[M].北京:科学出版社,2006
[4]陈瑞,程在全,黄兴奇等.水稻优良性状控制基因的定位进展及其在染色体上的分布[J].HEREDITAS(Beijing)2007,29(4):399-412
[5]Burr.B.Burr F.A.Recombinant inbreds for molecular mapping in maize:theoretical and practical considerations,TTG,1991,7:55-60
[6]高用明,朱军,宋佑胜等.水稻永久F2群体抽穗期QTL的上位性及其与环境互作效应的分析[J].作物学报,2004,30(9):849-854
[7]吴为人,唐定中,李维明.数量性状的遗传剖析和分子剖析[J].作物学报,2000,26(4):501-507
[8]Lander,.A..Barlow,.S..E.Lincoln etal.MAPMARKER:an interactive computer package for constructing.primary.genetic.linkagemaps.ofexperimental.and.natural.populations.Genomics,1987,1:174-181
[9]Suiter KA,Wendel JF,Case JS.LINKAGE-Ⅰ:a Pascal computer program for the detection and analysis of genetic linkage.J Hered,1987,74:203-204
[10]Morton NE,Andrews V.MAP,an expert system for,multiple pair wise linkage analysis.AnnHum Genet,1989:53:263-269
[11]Shoemaker J,Zaitlin D A comparison of three Agrigeoietics maize RFLP linkage map.Maize Genet Coop Newslett,1992,66:65-68
[12]Manly KF,Elliott RW.RI Manager,a microcomputer program for analysis of data from recombinant inbred strains.Mamm Genome,1991,1(2):123-126
[13]Stam P.Construction of integrated genetic linkage maps by means of a new computer-package Join Map.Plant J,1993,3:739-744
[14]McCouch S R,Kochert G,Yu Z H etal.Molecular mapping of rice chromosomes.Theor Appl Genet,1988,76:815-829
[15]杨俊品,曹永国,.玉米SSR遗传图谱的构建及产量性状基因定位[J].遗传学报,2001,28(8):778-784
[16]Paterson A H,Lander E S,Hewitt J D etal.Resolution of quantitative traits into Mendelian factors using a complete likage map of restriction fragment length polymorphisms.Nature,1988,335:721-726
[17]Tanksley S D,Paterson A,Paterson A.RFLP mapping in plant breeding[J].生物技术,1989,7:257-264
[18]Apuya N R,Frazier B L,Keim P et al.Restriction fragment length polymorphisms asgenetic markers in soybean.Theor Appl Genet,1988,75:889-901
[19]袁爱萍,侯爱斌,毛雪等.高粱基因组学研究的新进展[J].生物技术通报,2003,01:8-12
[20]Heun M.Mapping quantitative powdery mildew resistance of barly using a restriction fragment polymorphism map.Genome,1992,35:1019-1025
[21]Gill K S,Lubbers E L,Gill B S etal.A genetic linkage mapof Triticum Tauschii(DD) and its relationship to the D genome of bread wheat(AABDD).Genome,1991,34:362 -374
[22][14]McCouch S R,Kochert G,Yu Z H etal.Molecular mapping of rice chromosomes.Theor Appl Genet,1988,76:815-829
[23]McMouch S.R,S.D.Tanksley.Development and use of RFLP in rice breeding and genetics.Rice Biotechnology,1992,109-133
[24]徐吉臣,朱立煌,陈英等.用双单倍体群体构建水稻的分子连锁图谱[J].遗传学报,1994,21(3):205-214
[25]李平.水稻分子图谱的构建与基因分析,四川雅安,四川农业大学,1994年
[26]谭震波.水稻分子图谱的构建及数量性状基因的研究,四川雅交,四川农业大学,1996年
[27]Causse MA,Fulton TM.Saturated molecular map of the rice genome based on an interspecifi backcross population.Genetics,1994,138:1251-1274
[28]Kurata N,Nagamura Y,Harushima Y etal.A 300 Kilobase interval genetic map of rice including 883 expressed sequence.Nature Genetics,1994,8:365-372
[29]Harushima Y,Yano M,Shomura A et al.A high-desity rice genetic linkage map with 2275markers using a single F2 population.Genetics,1998,148:479-494
[30]Saito A,Yano M,Kishimoto N etal.Linkage map of restriction fragment length polymorphism loci in rice.Japan J Breed;1991,41:665-670
[31]Xiao J,Li J,Yuan L etal.Identification of QTLs affecting traits of agronomic importance in a recombinant inbred population derived from a subspecifiC rice cross.Theor Appl Genet,1996,92:230-244
[32]唐定中.水稻分子标记连锁图的构建与应用.福建神州,福建农业大学,1998年
[33]翁清妹,吴为人,李维明等.用2个不同来源的DNA探针构建水稻RFLP连锁图谱[J].福律农林大学学报,2000,29(2),129-133
[34]李维明,卢浩然,唐定中等.用籼交重组自交系群体构建的分子遗传图谱及其与籼/粳交群体的分子图谱的比较[J].中国水稻科学,2000,14(2):71-78
[35]Tanksley S.D.Mapping polygenes.Annu Rev Genet,1993,27:205-233
[36]Stuber C.W,Moll R.H,Goodman M.M.Allozyme frequency changes associated with selection for increased grain yield in maize(Zea mat's L.).Genetics,1980,95:225-236
[37]Sax K.The association of size difference with seed-coat pattern and pigmentation in phaseolus valgaris[J].Genetics,1923,8:522-560
[38]Lander E.S,Botstein D.Mapping mendelian factors underlying guantitative traits using RFLP linkage maps.Genetics,1989,121:185-199
[39]Zeng.Z.B.Precision mapping of quantitative trait loci.loci.Genetics,1994,136:1457-1468
[40]Soller M,Brody T,Genizi A.On the power of experimental designs for the detection of linkage between marker loci and quantitative loci in crosses between inbred lines.Theor Appi.Genet,1976,47:35-39
[41]Simpson S P.detection of Linkage between quantitative traits loci and restriction fragment length polymorphism using inbred lines.Theor Appl Genet,1989,77:815-819
[42]李维明,吴为人,卢浩然.检测作物数量性状基因与遗传标记连锁关系的方差分析法及其应用[J].作物学报,1993,19(2):97-102
[43]Kearsey M J,Hyne V.QTL analysis:a simple marker regression approach.Theor Appl Genet,1994,89:698-702
[44]Jiang C,Zeng Z B.Mapping quantitative trait loci with dominant and missing markers in various crosses from two inbred lines.Genetica,1997,101:47-58
[45]朱军.复杂数量性状基因定位的混合模型方法,全国作物育种学术讨论会论文集(干连铮戴景瑞主编),中国农业出版社,1998 19-20
[46]Zhu J,Weir BS:Mixed model approaches for genetic analysis of quantitative traits,In Chen LS,Ruan S q Zhu J,ed,Advanced Topics in Biomathematics,Proceedings of International Conference on Mathematical Biology.World Scientific Publishing Co,Singapore 1998 321-330
[47]高用明,朱军.植物QTL定位方法的研究进展[J].遗传,2000,22(3):175-179
[48]PIEPHO,H.P,G.KOCH.Codominant analysis of banding data from a dominant marker system by normal mixtures.Genetics,2000,155:1459-1468.
[49]Zhu J.Analysis of conditional genetic effects and variance components in developmental genetics.Genetics,1995,141:1633-1639(条件 QTL)
[50]杨绍华.水稻若干重要农艺性状的QTL定位[D].扬州大学硕士论文.2001
[51]李余生,汤国辉,管荣展等.不同环境下水稻产量相关性状的QTL分析[J].江苏农业学报,2007,23(6):509-515
[52]陈庆全,余四斌,李春海等.不同环境条件下水稻结实率的QTL定位分析[J].农业生物技术学报,2007,15(5):834-840
[53]严长杰,李欣.利用分子标记定位水稻芽期耐冷性基因[J].中国水稻科学,1999,13(3):134-138
[54]乔永利,韩龙直,安永平等.水稻芽期耐冷性QTL定位的分子定位[J].中国农业科学,2005,38(2):217-221
[55]陈玮,李炜.水稻RIL群体芽期耐冷性基因的分子标记定位[J].武汉植物学研究,2005,23(2):116-120
[56]ChungG S.The rice cold tolerance program in Korea[C].In:Re.port of a Rice Cold Tolerance Workshop.Manila:IRRI,1979.7-19
[57]Li C C,Rutger J N.Inheritance of cool temperature seedling vigorin rice and its relationship with other agronomic charactcrs[J].CropSci,1980,(20):295-298
[58]陈大洲,钟平安,肖叶青等.利用SSR标记定位东乡野生稻苗期耐冷性基因[J].江西农业大学学报,2002,24(6):735-756
[59]钱前,曾大力,何平等.水稻籼粳交DH群体苗期耐冷性QTLs分析[J].科学通报,1999,44(22):2402-2407
[60]詹庆才,曾曙珍,熊伏星等.水稻苗划耐冷性QTLs的分子定位[J].湖南农业大学学报,2003,29(1):7-11
[61]胡莹,王奕众.水稻RIL群体苗期耐冷性QTL分析[J].武汉植物学研究,2005,23(3):211-215
[62]Tsunoda K,Fujimure K,Nakahori T,etc.Studies on the testing methods for cool-tolerance in rice plants.Ⅰ.An improved method by means of short term treatment with cool and deep water.Japan J Breeding,1968,18(1):33-40
[63]曾亚文,申时全,文国松.小白谷×大理早籼重组自交系耐冷性状遗传研究[J].西南农业大学学报,2001,(6):494-497
[64]叶昌荣.云南稻种冲腿的孕穗期耐冷性QTL分析[J].中国水稻科学,2001,15(1):13-16
[65]刘凤霞,孙传清.江西东乡野生稻孕穗开花期耐冷基因定位[J].科学通报,2003,48(17):1864-1867
[66]王连敏,王立志,张国民.花期低温对水稻结实的影响[J].中国农业气象,1997,18(5):9-11
[67]Jeong E G.Molecular gentic analysis of quantitative trait loci related to cold tolerance in rice.Ph.D thesis.Kangwon National University.2001
[68]赵正武,王述明,雷树凡等.糯89-1腋芽休眠越冬及种子耐寒性研究[J].植物遗传资源科学,2000,1(3):25-28
[69]赵正武,水稻越冬耐冷性遗传变异及QTL定位研究[D].四川农业大学博士论文,2006
[70]张成良.东乡野生稻耐寒性研究现状与展望[J].种子(SEED),2006,25(10):44-47
[71]郭远明.中国培育“傻瓜水稻”挑战千年稻种耕作制.[2003-6-27].http://www.sina.com.cn.
[72]陈大洲,肖叶青.东乡野生稻耐冷性的遗传改良初步研究[J].江西农业大学学报,2003,25(1):8-11
[73]丁彦,周清明.水稻再生力利用研究及展望[J].湖南农业科学,2005,(2):11-1
[74]唐浩,陈立云,杨益善等.水稻的再生率及其与产量性状的关系[J].杂交水稻,2003,18(3):55-58
[75]谭震波,沈利爽,袁祚廉等.水稻再生能力和头季稻产量性状的QTL定位及其遗传效应分析[J].作物学报,1997,23(3):289-295
[76]郑景生,李义珍,林文雄.应用SSR标记定位水稻再生力和再生产量及其构成的QTL[J].分子植物育种,2004,2(3):342-347
[77]李太贵.在低温下筛选水稻不同生长期耐寒品种的室内方法[J].国外农业科技,1981(4):18-21
[78]易镇邪,屠乃美,陈平平.杂交稻新组合再生稻头季及再生季源库特征分析[J].中国水稻科学,2005,19(3):243-248
[79]曹立勇,占小登,庄杰云等.水稻产量性状的QTL定位与上位性分析[J].中国农业科学,2003,36(11):1241-1247
[80]郭龙彪,罗利军,邢永忠等.水稻重要农艺性状的两年QTL剖析[J].中国水稻科学,2003,17(3):211-218
[81]许凌,张亚东,朱镇等.不同年份水稻产量性状的QTL分析[J].中国水稻科学,2008,22(4):370-376
[82]张启军.利用SSR和四个测序水稻品种构建分子图谱及QTLs分析[D],博士论文,2004
[83]封功能.水稻籼粳交产量相关性状的QTL定位[D].扬州大学硕士论文,2004
[84]陈庆全,余四斌,李春海等.不同环境条件下水稻结实率的QTL定位分析[J].农业生物技术学报,2007,15(5):834-840
[85]鄂志国,吴殿星,庄杰云等.水稻千粒重QTL图谱整合研究[J].科技通报,2008,(24)5:617-622
[86]Lin W.,Li Y.Z.,Jiang Z.W.,and Zheng J.S.,2001,Morphologic and functional difference of root systems among ratooning rice varieties and its correlation with yield,Fujian Nongye Xuebao(Fujian Joural of Agricultural Sciences),16(1):1-4
[87]Liu Y.S.,Zhou K.D.,Zeng R.Y.,and Luo W.Z.,1992,Evaluation for rice ratooning ability of intersubspecific hybrid and its relation to agronomic characters of mother plant,Zhongguo Shuidao Kexie(Chinese Journal of rice Science),6(4):151-154
[88]张露霞,王松凤,江玲等.利用重组自交系群体检测水稻芽期耐寒性QTL[J].南京农业大学学报,2007,30(4):1-5
[89]Wang GL,Mackill DJ,et al,1994,RFLP mapping of gengs conferring complete and partial resistance to blast in a durably resistant rice cultivar.Genetics 136:1421-1434
[90]Yano M,Harushima,et al,1994,Strategy for genetic dissection of quantitative traits into single mendelian factors using DNA markers.Rice genome,3(2):5
[91]Maeda,H.,T.Ishii,l,et al,1995,Molecular mapping of semidwarfing gene,sd-1,using RAPD and RFLP markers Breeding Science,45:93
[2]李仕贵.水稻几个重要性状的遗传分析和分子标记定位[D].四川农业大学博士论文,1998
[3]钱前,程式华.水稻遗传学和功能基因组学[M].北京:科学出版社,2006
[4]陈瑞,程在全,黄兴奇等.水稻优良性状控制基因的定位进展及其在染色体上的分布[J].HEREDITAS(Beijing)2007,29(4):399-412
[5]Burr.B.Burr F.A.Recombinant inbreds for molecular mapping in maize:theoretical and practical considerations,TTG,1991,7:55-60
[6]高用明,朱军,宋佑胜等.水稻永久F2群体抽穗期QTL的上位性及其与环境互作效应的分析[J].作物学报,2004,30(9):849-854
[7]吴为人,唐定中,李维明.数量性状的遗传剖析和分子剖析[J].作物学报,2000,26(4):501-507
[8]Lander,.A..Barlow,.S..E.Lincoln etal.MAPMARKER:an interactive computer package for constructing.primary.genetic.linkagemaps.ofexperimental.and.natural.populations.Genomics,1987,1:174-181
[9]Suiter KA,Wendel JF,Case JS.LINKAGE-Ⅰ:a Pascal computer program for the detection and analysis of genetic linkage.J Hered,1987,74:203-204
[10]Morton NE,Andrews V.MAP,an expert system for,multiple pair wise linkage analysis.AnnHum Genet,1989:53:263-269
[11]Shoemaker J,Zaitlin D A comparison of three Agrigeoietics maize RFLP linkage map.Maize Genet Coop Newslett,1992,66:65-68
[12]Manly KF,Elliott RW.RI Manager,a microcomputer program for analysis of data from recombinant inbred strains.Mamm Genome,1991,1(2):123-126
[13]Stam P.Construction of integrated genetic linkage maps by means of a new computer-package Join Map.Plant J,1993,3:739-744
[14]McCouch S R,Kochert G,Yu Z H etal.Molecular mapping of rice chromosomes.Theor Appl Genet,1988,76:815-829
[15]杨俊品,曹永国,.玉米SSR遗传图谱的构建及产量性状基因定位[J].遗传学报,2001,28(8):778-784
[16]Paterson A H,Lander E S,Hewitt J D etal.Resolution of quantitative traits into Mendelian factors using a complete likage map of restriction fragment length polymorphisms.Nature,1988,335:721-726
[17]Tanksley S D,Paterson A,Paterson A.RFLP mapping in plant breeding[J].生物技术,1989,7:257-264
[18]Apuya N R,Frazier B L,Keim P et al.Restriction fragment length polymorphisms asgenetic markers in soybean.Theor Appl Genet,1988,75:889-901
[19]袁爱萍,侯爱斌,毛雪等.高粱基因组学研究的新进展[J].生物技术通报,2003,01:8-12
[20]Heun M.Mapping quantitative powdery mildew resistance of barly using a restriction fragment polymorphism map.Genome,1992,35:1019-1025
[21]Gill K S,Lubbers E L,Gill B S etal.A genetic linkage mapof Triticum Tauschii(DD) and its relationship to the D genome of bread wheat(AABDD).Genome,1991,34:362 -374
[22][14]McCouch S R,Kochert G,Yu Z H etal.Molecular mapping of rice chromosomes.Theor Appl Genet,1988,76:815-829
[23]McMouch S.R,S.D.Tanksley.Development and use of RFLP in rice breeding and genetics.Rice Biotechnology,1992,109-133
[24]徐吉臣,朱立煌,陈英等.用双单倍体群体构建水稻的分子连锁图谱[J].遗传学报,1994,21(3):205-214
[25]李平.水稻分子图谱的构建与基因分析,四川雅安,四川农业大学,1994年
[26]谭震波.水稻分子图谱的构建及数量性状基因的研究,四川雅交,四川农业大学,1996年
[27]Causse MA,Fulton TM.Saturated molecular map of the rice genome based on an interspecifi backcross population.Genetics,1994,138:1251-1274
[28]Kurata N,Nagamura Y,Harushima Y etal.A 300 Kilobase interval genetic map of rice including 883 expressed sequence.Nature Genetics,1994,8:365-372
[29]Harushima Y,Yano M,Shomura A et al.A high-desity rice genetic linkage map with 2275markers using a single F2 population.Genetics,1998,148:479-494
[30]Saito A,Yano M,Kishimoto N etal.Linkage map of restriction fragment length polymorphism loci in rice.Japan J Breed;1991,41:665-670
[31]Xiao J,Li J,Yuan L etal.Identification of QTLs affecting traits of agronomic importance in a recombinant inbred population derived from a subspecifiC rice cross.Theor Appl Genet,1996,92:230-244
[32]唐定中.水稻分子标记连锁图的构建与应用.福建神州,福建农业大学,1998年
[33]翁清妹,吴为人,李维明等.用2个不同来源的DNA探针构建水稻RFLP连锁图谱[J].福律农林大学学报,2000,29(2),129-133
[34]李维明,卢浩然,唐定中等.用籼交重组自交系群体构建的分子遗传图谱及其与籼/粳交群体的分子图谱的比较[J].中国水稻科学,2000,14(2):71-78
[35]Tanksley S.D.Mapping polygenes.Annu Rev Genet,1993,27:205-233
[36]Stuber C.W,Moll R.H,Goodman M.M.Allozyme frequency changes associated with selection for increased grain yield in maize(Zea mat's L.).Genetics,1980,95:225-236
[37]Sax K.The association of size difference with seed-coat pattern and pigmentation in phaseolus valgaris[J].Genetics,1923,8:522-560
[38]Lander E.S,Botstein D.Mapping mendelian factors underlying guantitative traits using RFLP linkage maps.Genetics,1989,121:185-199
[39]Zeng.Z.B.Precision mapping of quantitative trait loci.loci.Genetics,1994,136:1457-1468
[40]Soller M,Brody T,Genizi A.On the power of experimental designs for the detection of linkage between marker loci and quantitative loci in crosses between inbred lines.Theor Appi.Genet,1976,47:35-39
[41]Simpson S P.detection of Linkage between quantitative traits loci and restriction fragment length polymorphism using inbred lines.Theor Appl Genet,1989,77:815-819
[42]李维明,吴为人,卢浩然.检测作物数量性状基因与遗传标记连锁关系的方差分析法及其应用[J].作物学报,1993,19(2):97-102
[43]Kearsey M J,Hyne V.QTL analysis:a simple marker regression approach.Theor Appl Genet,1994,89:698-702
[44]Jiang C,Zeng Z B.Mapping quantitative trait loci with dominant and missing markers in various crosses from two inbred lines.Genetica,1997,101:47-58
[45]朱军.复杂数量性状基因定位的混合模型方法,全国作物育种学术讨论会论文集(干连铮戴景瑞主编),中国农业出版社,1998 19-20
[46]Zhu J,Weir BS:Mixed model approaches for genetic analysis of quantitative traits,In Chen LS,Ruan S q Zhu J,ed,Advanced Topics in Biomathematics,Proceedings of International Conference on Mathematical Biology.World Scientific Publishing Co,Singapore 1998 321-330
[47]高用明,朱军.植物QTL定位方法的研究进展[J].遗传,2000,22(3):175-179
[48]PIEPHO,H.P,G.KOCH.Codominant analysis of banding data from a dominant marker system by normal mixtures.Genetics,2000,155:1459-1468.
[49]Zhu J.Analysis of conditional genetic effects and variance components in developmental genetics.Genetics,1995,141:1633-1639(条件 QTL)
[50]杨绍华.水稻若干重要农艺性状的QTL定位[D].扬州大学硕士论文.2001
[51]李余生,汤国辉,管荣展等.不同环境下水稻产量相关性状的QTL分析[J].江苏农业学报,2007,23(6):509-515
[52]陈庆全,余四斌,李春海等.不同环境条件下水稻结实率的QTL定位分析[J].农业生物技术学报,2007,15(5):834-840
[53]严长杰,李欣.利用分子标记定位水稻芽期耐冷性基因[J].中国水稻科学,1999,13(3):134-138
[54]乔永利,韩龙直,安永平等.水稻芽期耐冷性QTL定位的分子定位[J].中国农业科学,2005,38(2):217-221
[55]陈玮,李炜.水稻RIL群体芽期耐冷性基因的分子标记定位[J].武汉植物学研究,2005,23(2):116-120
[56]ChungG S.The rice cold tolerance program in Korea[C].In:Re.port of a Rice Cold Tolerance Workshop.Manila:IRRI,1979.7-19
[57]Li C C,Rutger J N.Inheritance of cool temperature seedling vigorin rice and its relationship with other agronomic charactcrs[J].CropSci,1980,(20):295-298
[58]陈大洲,钟平安,肖叶青等.利用SSR标记定位东乡野生稻苗期耐冷性基因[J].江西农业大学学报,2002,24(6):735-756
[59]钱前,曾大力,何平等.水稻籼粳交DH群体苗期耐冷性QTLs分析[J].科学通报,1999,44(22):2402-2407
[60]詹庆才,曾曙珍,熊伏星等.水稻苗划耐冷性QTLs的分子定位[J].湖南农业大学学报,2003,29(1):7-11
[61]胡莹,王奕众.水稻RIL群体苗期耐冷性QTL分析[J].武汉植物学研究,2005,23(3):211-215
[62]Tsunoda K,Fujimure K,Nakahori T,etc.Studies on the testing methods for cool-tolerance in rice plants.Ⅰ.An improved method by means of short term treatment with cool and deep water.Japan J Breeding,1968,18(1):33-40
[63]曾亚文,申时全,文国松.小白谷×大理早籼重组自交系耐冷性状遗传研究[J].西南农业大学学报,2001,(6):494-497
[64]叶昌荣.云南稻种冲腿的孕穗期耐冷性QTL分析[J].中国水稻科学,2001,15(1):13-16
[65]刘凤霞,孙传清.江西东乡野生稻孕穗开花期耐冷基因定位[J].科学通报,2003,48(17):1864-1867
[66]王连敏,王立志,张国民.花期低温对水稻结实的影响[J].中国农业气象,1997,18(5):9-11
[67]Jeong E G.Molecular gentic analysis of quantitative trait loci related to cold tolerance in rice.Ph.D thesis.Kangwon National University.2001
[68]赵正武,王述明,雷树凡等.糯89-1腋芽休眠越冬及种子耐寒性研究[J].植物遗传资源科学,2000,1(3):25-28
[69]赵正武,水稻越冬耐冷性遗传变异及QTL定位研究[D].四川农业大学博士论文,2006
[70]张成良.东乡野生稻耐寒性研究现状与展望[J].种子(SEED),2006,25(10):44-47
[71]郭远明.中国培育“傻瓜水稻”挑战千年稻种耕作制.[2003-6-27].http://www.sina.com.cn.
[72]陈大洲,肖叶青.东乡野生稻耐冷性的遗传改良初步研究[J].江西农业大学学报,2003,25(1):8-11
[73]丁彦,周清明.水稻再生力利用研究及展望[J].湖南农业科学,2005,(2):11-1
[74]唐浩,陈立云,杨益善等.水稻的再生率及其与产量性状的关系[J].杂交水稻,2003,18(3):55-58
[75]谭震波,沈利爽,袁祚廉等.水稻再生能力和头季稻产量性状的QTL定位及其遗传效应分析[J].作物学报,1997,23(3):289-295
[76]郑景生,李义珍,林文雄.应用SSR标记定位水稻再生力和再生产量及其构成的QTL[J].分子植物育种,2004,2(3):342-347
[77]李太贵.在低温下筛选水稻不同生长期耐寒品种的室内方法[J].国外农业科技,1981(4):18-21
[78]易镇邪,屠乃美,陈平平.杂交稻新组合再生稻头季及再生季源库特征分析[J].中国水稻科学,2005,19(3):243-248
[79]曹立勇,占小登,庄杰云等.水稻产量性状的QTL定位与上位性分析[J].中国农业科学,2003,36(11):1241-1247
[80]郭龙彪,罗利军,邢永忠等.水稻重要农艺性状的两年QTL剖析[J].中国水稻科学,2003,17(3):211-218
[81]许凌,张亚东,朱镇等.不同年份水稻产量性状的QTL分析[J].中国水稻科学,2008,22(4):370-376
[82]张启军.利用SSR和四个测序水稻品种构建分子图谱及QTLs分析[D],博士论文,2004
[83]封功能.水稻籼粳交产量相关性状的QTL定位[D].扬州大学硕士论文,2004
[84]陈庆全,余四斌,李春海等.不同环境条件下水稻结实率的QTL定位分析[J].农业生物技术学报,2007,15(5):834-840
[85]鄂志国,吴殿星,庄杰云等.水稻千粒重QTL图谱整合研究[J].科技通报,2008,(24)5:617-622
[86]Lin W.,Li Y.Z.,Jiang Z.W.,and Zheng J.S.,2001,Morphologic and functional difference of root systems among ratooning rice varieties and its correlation with yield,Fujian Nongye Xuebao(Fujian Joural of Agricultural Sciences),16(1):1-4
[87]Liu Y.S.,Zhou K.D.,Zeng R.Y.,and Luo W.Z.,1992,Evaluation for rice ratooning ability of intersubspecific hybrid and its relation to agronomic characters of mother plant,Zhongguo Shuidao Kexie(Chinese Journal of rice Science),6(4):151-154
[88]张露霞,王松凤,江玲等.利用重组自交系群体检测水稻芽期耐寒性QTL[J].南京农业大学学报,2007,30(4):1-5
[89]Wang GL,Mackill DJ,et al,1994,RFLP mapping of gengs conferring complete and partial resistance to blast in a durably resistant rice cultivar.Genetics 136:1421-1434
[90]Yano M,Harushima,et al,1994,Strategy for genetic dissection of quantitative traits into single mendelian factors using DNA markers.Rice genome,3(2):5
[91]Maeda,H.,T.Ishii,l,et al,1995,Molecular mapping of semidwarfing gene,sd-1,using RAPD and RFLP markers Breeding Science,45:93
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