水稻几个重要性状的QTL定位及抗白叶枯病基因分子标记辅助选择
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摘要
产量相关性状、株高、抽穗期、耐热性及幼苗活力相关性状为水稻的重要性状，本论文构建了水稻DH、RIL遗传群体，并进行了分子标记遗传作图，在此基础上考察了水稻的上述几个重要性状，定位了这几个性状的QTL，最后应用分子标记辅助选择，进行了水稻白叶枯病抗性基因的转育。主要的研究内容和结果如下：
1、以与水稻耐贮藏材料(Daw Dam)有丰富分子标记多态性的常规材料2070、明恢63、Dulur、协青早B为亲本，分别与之杂交，采用改良一步成苗花药培养法对其F_1代进行花药培养，以一步成苗法为对照，评价新方法在构建DH作图群体中的应用潜力。研究结果表明，改良一步成苗法显著地提高了愈伤组织分化率、植株再生率、绿苗率、绿苗素质，且得到的DH系有良好田间表现，是构建DH作图群体的理想方法，但不同基因型间的改进效果表现出一定的差异性，三个参试组合中Daw Dam／2070、Daw Dam／明恢63的培养效率有较大增幅。利用改良一步成苗法已获得一定数量的DH群体，为开展进一步研究奠定了基础。
2、重组自交系群体是构建遗传连锁图谱的最常用群体之一；以中国水稻研究所育成的高产但感穗瘟的品种中156与半矮杆抗稻瘟病品种谷梅2号杂交，建立由304个RIL(重组自交系)组成的品种间重组自交系群体，应用RFLP, RAPD, SSLP, RGA和CG共168个DNA标记，构建了全长为1447.9cM、覆盖水稻基因组12条染色体的连锁图。
以中156／谷梅2号304个F_9重组自交系及双亲，于2001年分单季和连晚两季在杭州中国水稻研究所试验场以完全随机区组种植实验材料，记载抽穗期、考种株高、穗长、单株有效穗数、每穗颖花数、每穗实粒数、结实率及千粒重等7个性状，将不同季别当作环境因子处理，采用QTLMapper统计软件进行水稻株高、抽穗期和产量性状的QTL定位、上位性分析及其与环境的互作效应分析，同时对株高和单株有效穗数进行了条件QTL定位(剔除抽穗期对株高和单株有效穗数的影响)。
上述8个性状共检测到42个主效应QTLs，分别位于除第9染色体以外的11条染色体上，其中只有6个主效应QTLs表现出与环境(季别)之间存在显著互作(GE)。共检测到39对影响株高、抽穗期及产量构成性状的加性×加性上位性显著互作效应，其中株高6对，抽穗期2对，穗长2对，单株有效穗数2对，每穗颖花数9对，每穗实粒数9对，结实率6对，千粒重3对，它们分别解释这些性状总变异的12.12％、1.38％、
3．76％、5．00％、10．24％。20．50％、16．31％和 4．48％。在所有的上位性效应中，多数加性
＼加性上位性互作效应的贡献率及效应均较小，没有检测到上位性效应与环境的显著
互作。
在对株高和单株有效穗数条件QTL定位表明，共检测到7个影响株高的主效应QTLS
＼ 和3个影响单株有效穗数的主效应QTLS，以及6对影响株高、3对影响有效穗数的上位
性QTLS。在影响株高的 QTLS中，qPH7－1和 qPH10仅表现出与环境的互作；qPH7－2既
表现加性效应也表现出与环境的互作效应，另外4个QTLS未检测出与环境的互作；3
个影响单株有效穗数的叮LS都仅表现为加性效应，而未检测出与环境的互作：检测到
6对影响株高和 3对影响单株有效穗数的上位性互作QTLS，贡献率分别在 1．9％到4．2％
之间和3．26％到3．％％之间；没有检测到上位性叮L与环境的互作效应。同时也显示抽
穗期这一性状对株高和单株有效穗数QTLS的表达既有抑制作用、也具有较大的贡献率。
3、耐热性是水稻（OI’Ma sa ti va L．）抗逆研究中最重要的性状之一。本研究应
应。
5、选择抗白叶枯病的具Xa－21基因的供体亲本IRBB60与感病的多系l号与R2070
的杂种后代儿）目交。利用与h刁1基因紧密连锁的分子标记p仪248进行Xa上1基因
标记辅助选择抗白叶枯病恢复系的选育，经过多次日交筛选出抗白叶枯病杂交水稻恢
】复系，所组配的新组合（中 gA八8006）据初步试种，表现出穗大粒多，株型挺、耐肥
抗倒的特点，2001年参加浙江省“8812”联品，表现抗白叶枯病和稻瘟病，比对照汕
优63增产6．67巩2002年参加南方稻区区试和浙江、江西、湖南等省区试，可望在生
产上应用。
Most agronomic characteristics are important in rice such as PH (plant height), HD (heading date), HR (heat-resistance) and yield components or seedling vigor .components. Both DH (doubled haploid) and RIL (recombinant inbred lines) populations were derived. A genetic map was constructed by molecular marker analysis. QTLs associated with these traits above were mapped according to the data for phenotype. At the same time, we transferred gene resistant to bacteria blight into rice on the basis of marker-assisted selection. Following are major contents and results studied.
Crosses were produced by using a storable rice variety (Daw Dam) and 2070, Ming Hui 63, Dulur as well as Xie Qing Zao B as parents, among these materials there are abundant polymorphisms of DNA molecular markers. Their Ffs were applied to generate DH lines by an improved method of direct induction of pollen plant. The method of the direct induction of pollen plant was used as check treatment to evaluate the improved method. The results showed that the improved method increased the re-differentiation percentage of pollen callus, the percentage of regeneration plantlets, and the percentage of green regeneration plantlets and produced more vigorous seedlings. The improved method was a competent method of establishing DH population for mapping lox-3 gene. It was also showed that the efficiency of the new method was different among genotypes and out of three experimental crosses Daw Dam/2070 and Daw Dam/Minghui 63 showed more efficient. More than 60 DH lines have been obtained in the present experiment.
A RIL population consists of 304 individuals was derived from a cross between Zhongl56, a species sensitive to rice blight but has high yield, and Gumei2, a species resistant to rice blight, which is semi-dwarf. Both parents are selected in China National Rice Research Institute (CNRRI). Applied to RFLP, RAPD, SSLP, RGA and CG, a linkage map consisting of 168 markers was constructed, which covers 1447.9cM of the rice genome and covers all of chromosomes in rice.
The parents and a Fg including 304 individuals were grown in the paddy field in China National Rice Research Institute (CNRRI), Hangzhou, China in 2001. The experiments were
IV
carried out in two enviroments followed a randomized complete block design. Recorded the data of HD (heading data), PH (plant height), PL (panicle length), PNPP (panicle number per plant), SNPP (spikelets number per panicle), FGN (filled grain number), GF (grain fertility) and KW (kilo-grain weight). The identification of QTL for PH and HD and yield components and analysis of epistasis and QTL * environment interaction were conducted by using the statistic software of MAPMARKER/QTL. In addition, conditional QTL analysis for PH and PNPP was made.
42 QTLs with additive effects covering all chromosomes except 9 chromosome were detected. 6 QTLs have significant QE interaction. 39 QTLs with additive x additive epistatic effects for PH, HD and yield components were detected, including 6 QTLs controlling PH, 2 QTLs controlling HD, 2 QTLs controlling PL, 2 QTLs controlling PNPP, 9 QTLs controlling SNPP, 9 QTLs controlling FGN, 6 QTLs controlling GF, 3 QTLs controlling KW. The phenotypic variances these QTLs explained separately were 12.12%, 1.38%, 3.76%, 5.00%, 10.24%, 20.50%, 16.31% and 4.48%. For all epistatic effects, most effects and variances were lesser. No significant interaction between epistatic effects and environment was detected.
In conditional QTL analysis for PH and PNPP, 7 and 3 additive QTLs and 6 and 3 epistatic QTLs for PH and PNPP were detected. For all PH QTLs, qPH7-l and qPHIO showed QE interaction only; qPH7-2 showed both additive effect and QE interaction. No QE interaction for other 4 QTLs was detected. All 3 QTLs controlling PNPP showed additive effects only and no QE interaction was detected. The variances of 6 and 3 epistatic QTLs for PH and PNPP were from 1.9% to 4.2% and from 3.26% to 3.96%. No epistatic QE interaction was detected. Meanwhile conditional QTL analysis showed that HD
1、以与水稻耐贮藏材料(Daw Dam)有丰富分子标记多态性的常规材料2070、明恢63、Dulur、协青早B为亲本，分别与之杂交，采用改良一步成苗花药培养法对其F_1代进行花药培养，以一步成苗法为对照，评价新方法在构建DH作图群体中的应用潜力。研究结果表明，改良一步成苗法显著地提高了愈伤组织分化率、植株再生率、绿苗率、绿苗素质，且得到的DH系有良好田间表现，是构建DH作图群体的理想方法，但不同基因型间的改进效果表现出一定的差异性，三个参试组合中Daw Dam／2070、Daw Dam／明恢63的培养效率有较大增幅。利用改良一步成苗法已获得一定数量的DH群体，为开展进一步研究奠定了基础。
2、重组自交系群体是构建遗传连锁图谱的最常用群体之一；以中国水稻研究所育成的高产但感穗瘟的品种中156与半矮杆抗稻瘟病品种谷梅2号杂交，建立由304个RIL(重组自交系)组成的品种间重组自交系群体，应用RFLP, RAPD, SSLP, RGA和CG共168个DNA标记，构建了全长为1447.9cM、覆盖水稻基因组12条染色体的连锁图。
以中156／谷梅2号304个F_9重组自交系及双亲，于2001年分单季和连晚两季在杭州中国水稻研究所试验场以完全随机区组种植实验材料，记载抽穗期、考种株高、穗长、单株有效穗数、每穗颖花数、每穗实粒数、结实率及千粒重等7个性状，将不同季别当作环境因子处理，采用QTLMapper统计软件进行水稻株高、抽穗期和产量性状的QTL定位、上位性分析及其与环境的互作效应分析，同时对株高和单株有效穗数进行了条件QTL定位(剔除抽穗期对株高和单株有效穗数的影响)。
上述8个性状共检测到42个主效应QTLs，分别位于除第9染色体以外的11条染色体上，其中只有6个主效应QTLs表现出与环境(季别)之间存在显著互作(GE)。共检测到39对影响株高、抽穗期及产量构成性状的加性×加性上位性显著互作效应，其中株高6对，抽穗期2对，穗长2对，单株有效穗数2对，每穗颖花数9对，每穗实粒数9对，结实率6对，千粒重3对，它们分别解释这些性状总变异的12.12％、1.38％、
3．76％、5．00％、10．24％。20．50％、16．31％和 4．48％。在所有的上位性效应中，多数加性
＼加性上位性互作效应的贡献率及效应均较小，没有检测到上位性效应与环境的显著
互作。
在对株高和单株有效穗数条件QTL定位表明，共检测到7个影响株高的主效应QTLS
＼ 和3个影响单株有效穗数的主效应QTLS，以及6对影响株高、3对影响有效穗数的上位
性QTLS。在影响株高的 QTLS中，qPH7－1和 qPH10仅表现出与环境的互作；qPH7－2既
表现加性效应也表现出与环境的互作效应，另外4个QTLS未检测出与环境的互作；3
个影响单株有效穗数的叮LS都仅表现为加性效应，而未检测出与环境的互作：检测到
6对影响株高和 3对影响单株有效穗数的上位性互作QTLS，贡献率分别在 1．9％到4．2％
之间和3．26％到3．％％之间；没有检测到上位性叮L与环境的互作效应。同时也显示抽
穗期这一性状对株高和单株有效穗数QTLS的表达既有抑制作用、也具有较大的贡献率。
3、耐热性是水稻（OI’Ma sa ti va L．）抗逆研究中最重要的性状之一。本研究应
应。
5、选择抗白叶枯病的具Xa－21基因的供体亲本IRBB60与感病的多系l号与R2070
的杂种后代儿）目交。利用与h刁1基因紧密连锁的分子标记p仪248进行Xa上1基因
标记辅助选择抗白叶枯病恢复系的选育，经过多次日交筛选出抗白叶枯病杂交水稻恢
】复系，所组配的新组合（中 gA八8006）据初步试种，表现出穗大粒多，株型挺、耐肥
抗倒的特点，2001年参加浙江省“8812”联品，表现抗白叶枯病和稻瘟病，比对照汕
优63增产6．67巩2002年参加南方稻区区试和浙江、江西、湖南等省区试，可望在生
产上应用。
Most agronomic characteristics are important in rice such as PH (plant height), HD (heading date), HR (heat-resistance) and yield components or seedling vigor .components. Both DH (doubled haploid) and RIL (recombinant inbred lines) populations were derived. A genetic map was constructed by molecular marker analysis. QTLs associated with these traits above were mapped according to the data for phenotype. At the same time, we transferred gene resistant to bacteria blight into rice on the basis of marker-assisted selection. Following are major contents and results studied.
Crosses were produced by using a storable rice variety (Daw Dam) and 2070, Ming Hui 63, Dulur as well as Xie Qing Zao B as parents, among these materials there are abundant polymorphisms of DNA molecular markers. Their Ffs were applied to generate DH lines by an improved method of direct induction of pollen plant. The method of the direct induction of pollen plant was used as check treatment to evaluate the improved method. The results showed that the improved method increased the re-differentiation percentage of pollen callus, the percentage of regeneration plantlets, and the percentage of green regeneration plantlets and produced more vigorous seedlings. The improved method was a competent method of establishing DH population for mapping lox-3 gene. It was also showed that the efficiency of the new method was different among genotypes and out of three experimental crosses Daw Dam/2070 and Daw Dam/Minghui 63 showed more efficient. More than 60 DH lines have been obtained in the present experiment.
A RIL population consists of 304 individuals was derived from a cross between Zhongl56, a species sensitive to rice blight but has high yield, and Gumei2, a species resistant to rice blight, which is semi-dwarf. Both parents are selected in China National Rice Research Institute (CNRRI). Applied to RFLP, RAPD, SSLP, RGA and CG, a linkage map consisting of 168 markers was constructed, which covers 1447.9cM of the rice genome and covers all of chromosomes in rice.
The parents and a Fg including 304 individuals were grown in the paddy field in China National Rice Research Institute (CNRRI), Hangzhou, China in 2001. The experiments were
IV
carried out in two enviroments followed a randomized complete block design. Recorded the data of HD (heading data), PH (plant height), PL (panicle length), PNPP (panicle number per plant), SNPP (spikelets number per panicle), FGN (filled grain number), GF (grain fertility) and KW (kilo-grain weight). The identification of QTL for PH and HD and yield components and analysis of epistasis and QTL * environment interaction were conducted by using the statistic software of MAPMARKER/QTL. In addition, conditional QTL analysis for PH and PNPP was made.
42 QTLs with additive effects covering all chromosomes except 9 chromosome were detected. 6 QTLs have significant QE interaction. 39 QTLs with additive x additive epistatic effects for PH, HD and yield components were detected, including 6 QTLs controlling PH, 2 QTLs controlling HD, 2 QTLs controlling PL, 2 QTLs controlling PNPP, 9 QTLs controlling SNPP, 9 QTLs controlling FGN, 6 QTLs controlling GF, 3 QTLs controlling KW. The phenotypic variances these QTLs explained separately were 12.12%, 1.38%, 3.76%, 5.00%, 10.24%, 20.50%, 16.31% and 4.48%. For all epistatic effects, most effects and variances were lesser. No significant interaction between epistatic effects and environment was detected.
In conditional QTL analysis for PH and PNPP, 7 and 3 additive QTLs and 6 and 3 epistatic QTLs for PH and PNPP were detected. For all PH QTLs, qPH7-l and qPHIO showed QE interaction only; qPH7-2 showed both additive effect and QE interaction. No QE interaction for other 4 QTLs was detected. All 3 QTLs controlling PNPP showed additive effects only and no QE interaction was detected. The variances of 6 and 3 epistatic QTLs for PH and PNPP were from 1.9% to 4.2% and from 3.26% to 3.96%. No epistatic QE interaction was detected. Meanwhile conditional QTL analysis showed that HD
引文
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