水稻苗期与耐冷性有关的几个性状的动态基因定位
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摘要
本研究选用以水稻品种IR64和Azucena为亲本的105个DH群体为遗传材料，利用包括175个标记的水稻连锁图谱，采用基于混合线性模型的QTL定位软件QTLMapper1.6(Wang等，1999；朱军，1999)，结合Zhu(1995)提出的条件遗传分析方法，将苗期分为5个连续的阶段，研究水稻3叶期进行低温处理后，控制水稻苗期株高、根长、出叶数和苗重的QTL数目、位置和基因效应(包括加性、加性×处理互作、加×加上位性以及加×加上位性与处理互作效应)。主要研究结果如下：
1．本文通过分析苗期5个阶段4个性状在适温和冷处理条件下的数据，表明在个体发育过程中，一些具有较大显著主效应的QTLs可以使得个体能够在不同环境下保持相同的生长发育模式的同时，另外一些QTLs(具有显著QE互作效应的QTLs)决定了个体在不同环境下所发生的变异。因而在分子标记辅助选择育种中，对加性主效应的选择适用于各种不同的环境条件，而对QE效应的选择则适用于特定的环境条件。并且控制这些性状的增或减效等位基因往往分布在不同的亲本中。本研究中，水稻苗期株高、根长、出叶数和苗重，4个性状检测到的具有条件QE互作效应的QTLs的数目、效应大小及方向都不相同，决定了不同的性状在冷处理过后具有不同的恢复生长的速率。并且，由于控制不同性状的QTLs位于同一个基因位点(一因多效)或者基因间存在紧密连锁的现象，不同性状之间存在相关性。因而，对一个性状的选择会影响到其它的性状。
2．采用动态基因定位方法，条件和非条件定位方法检测到控制苗高、根长、叶片数和苗重的具有加性和加性×处理互作效应的QTLs分别为23、22、14和23个。但是所有这些与株高、根长、出叶数和苗重4个性状有关的QTLs中，分别只有3个、1个、0个、2个QTLs可以在5个观测时期都检测到显著的加性效应值，并且在不同的时期其效应值大小、作用方式也不相同。其它的QTLs只在1个或几个时期检测到显著的效应值，说明在不同的发育阶段基因是选择性表达的。即使是在几个阶段都表达的QTL，其表达方式也是有所不同的。本研究中采用的条件分析方法可以对特定发育阶段内的净基因表达效应进行有效的估算和检测，从而可以有效的探明基因在不同发育阶段的作用方式，为更清楚地了解各个性状在不同的时期对冷害的反应作用机理提供了一个有力的手段。
3．上位性是数量性状的重要遗传基础。本研究中条件和非条件定位方法共检测到30对影响苗高、36对影响根长、33对影响叶片数和23对影响苗重的上位性QTLs，但是分
浙江大学硕十学位论文
摘要
别有50%、47%、62%和42%的参与上位性的QTLs不具有显著的加性和(或)加性X处理
互作效应。少1且，一个QTL与多个QTLs同时存在互作是非常普遍的现象，而且存在多位
点互作时，其效应值方向也多不相同。另外，还发现，许多上位性互作只能用条件方法检
测到，说明大量的上位性互作只存在很短的一段时间，但是对各个性状的最终建成却起着
不容忽视的作用，尤其是对叶片数性状来说。
A population of 105 DH lines derived from a cross between irrigated indica variety IR64 and upland japonica variety Azucena was used in the experiments. The genetic linkage map of this population, containing 175 markers distributed among 12 chromosomes. In the present research, the dynamic behavior of four seedling traits including seedling height, root length, leaf number, and seedling weight after 3-leaf stage cold treatment were analyzed for detecting QTLs associated with additive and epistatic effects as well as their QE interaction effects by combining the statistical procedures for analyzing conditional genetic effects (Zhu 1995) and the QTL mapping method based on mixed model approaches (Wang et al. 1999; Zhu 1999). The QTL mapping analysis revealed the following results:
1. Through analyzing the data of four seedling traits at five measuring stages in two treatments, an important result could be obtained, that is, while a set of QTLs with relatively high magnitude of effects (those QTLs with significant genetic main effects) held the key to growth under different environments (treatments), few QTLs (those detected with significant QE effects) may, however, determine the differential response of the crop to environmental variation which could inversely guide the breeders to formulate an appropriate breeding strategy. In other words, selection on QTLs with additive main effects applied to different environments (treatments), while selection on QTLs with QE interaction could adapt to a special environment. And the alleles contributed to the seedling traits dispersed between two parents. In the study, the numbers ^ magnitude and direction of QTLs with conditional QE effects were different for different stages and traits determined the dissimilar growth resuming rates
at different stages and for different seedling traits. Furthmore, the phenomenon of pleiotropic effects or close linkage of genes were the main causes for correlations among traits which resulted in selection for one will result in a correlated response with another.
2. QTL mapping studies using age-specific measures clearly demonstrated distinct age-specific gene action. In our study, using unconditional and conditioanl mapping methods, there were 23 QTLs with additive and additive x treatment effects for seedling height, 22 QTLs for root length, 14 QTLs for leaf number, and 23 QTLs for seedling weight, respectively. But
only 3, 1,0, and 2 QTLs with significant additive effects at all measuring stages for seedling height, root length, leaf number, and seedling weight, respectively. And other QTLs could be detected only at one or several stages. This suggested that genes controlling these seedling traits expressed selectively at different stages. Conditional mapping method could estimate and detect the net effects of gene expression that were independent of the causal genetic effects in earlier stages. And could reveal the gene expression pattern efficiently at different stage. This technique could serve as an important tool for studying the inheritance of response to stress.
3. The present study showed the importance of epistasis as a genetic basis of the quantitative traits and revealed several important features of this phenomenon. In the study, with two mapping methods, there were 30 pairs of QTLs involved in epistasis for seedling height, 36 pairs for root length, 33 pairs for leaf number, and 25 pairs for seedling weight with proportion of 50%, 47%, 62%, and 42% for each seedling traits, respectively, without additive and/or additive x treatment interaction effects. Moreover, it was common for several loci to interact with more than one non-allelic locus at the same stage and/or across different stages, and, the interactions often showed differences in the direction of effects. It was indicated that the possibility of multilocus associations existed for trait development. Beside this, a lot of interactions could detected only by conditional mapping method indicated that many epistasis interactions
1．本文通过分析苗期5个阶段4个性状在适温和冷处理条件下的数据，表明在个体发育过程中，一些具有较大显著主效应的QTLs可以使得个体能够在不同环境下保持相同的生长发育模式的同时，另外一些QTLs(具有显著QE互作效应的QTLs)决定了个体在不同环境下所发生的变异。因而在分子标记辅助选择育种中，对加性主效应的选择适用于各种不同的环境条件，而对QE效应的选择则适用于特定的环境条件。并且控制这些性状的增或减效等位基因往往分布在不同的亲本中。本研究中，水稻苗期株高、根长、出叶数和苗重，4个性状检测到的具有条件QE互作效应的QTLs的数目、效应大小及方向都不相同，决定了不同的性状在冷处理过后具有不同的恢复生长的速率。并且，由于控制不同性状的QTLs位于同一个基因位点(一因多效)或者基因间存在紧密连锁的现象，不同性状之间存在相关性。因而，对一个性状的选择会影响到其它的性状。
2．采用动态基因定位方法，条件和非条件定位方法检测到控制苗高、根长、叶片数和苗重的具有加性和加性×处理互作效应的QTLs分别为23、22、14和23个。但是所有这些与株高、根长、出叶数和苗重4个性状有关的QTLs中，分别只有3个、1个、0个、2个QTLs可以在5个观测时期都检测到显著的加性效应值，并且在不同的时期其效应值大小、作用方式也不相同。其它的QTLs只在1个或几个时期检测到显著的效应值，说明在不同的发育阶段基因是选择性表达的。即使是在几个阶段都表达的QTL，其表达方式也是有所不同的。本研究中采用的条件分析方法可以对特定发育阶段内的净基因表达效应进行有效的估算和检测，从而可以有效的探明基因在不同发育阶段的作用方式，为更清楚地了解各个性状在不同的时期对冷害的反应作用机理提供了一个有力的手段。
3．上位性是数量性状的重要遗传基础。本研究中条件和非条件定位方法共检测到30对影响苗高、36对影响根长、33对影响叶片数和23对影响苗重的上位性QTLs，但是分
浙江大学硕十学位论文
摘要
别有50%、47%、62%和42%的参与上位性的QTLs不具有显著的加性和(或)加性X处理
互作效应。少1且，一个QTL与多个QTLs同时存在互作是非常普遍的现象，而且存在多位
点互作时，其效应值方向也多不相同。另外，还发现，许多上位性互作只能用条件方法检
测到，说明大量的上位性互作只存在很短的一段时间，但是对各个性状的最终建成却起着
不容忽视的作用，尤其是对叶片数性状来说。
A population of 105 DH lines derived from a cross between irrigated indica variety IR64 and upland japonica variety Azucena was used in the experiments. The genetic linkage map of this population, containing 175 markers distributed among 12 chromosomes. In the present research, the dynamic behavior of four seedling traits including seedling height, root length, leaf number, and seedling weight after 3-leaf stage cold treatment were analyzed for detecting QTLs associated with additive and epistatic effects as well as their QE interaction effects by combining the statistical procedures for analyzing conditional genetic effects (Zhu 1995) and the QTL mapping method based on mixed model approaches (Wang et al. 1999; Zhu 1999). The QTL mapping analysis revealed the following results:
1. Through analyzing the data of four seedling traits at five measuring stages in two treatments, an important result could be obtained, that is, while a set of QTLs with relatively high magnitude of effects (those QTLs with significant genetic main effects) held the key to growth under different environments (treatments), few QTLs (those detected with significant QE effects) may, however, determine the differential response of the crop to environmental variation which could inversely guide the breeders to formulate an appropriate breeding strategy. In other words, selection on QTLs with additive main effects applied to different environments (treatments), while selection on QTLs with QE interaction could adapt to a special environment. And the alleles contributed to the seedling traits dispersed between two parents. In the study, the numbers ^ magnitude and direction of QTLs with conditional QE effects were different for different stages and traits determined the dissimilar growth resuming rates
at different stages and for different seedling traits. Furthmore, the phenomenon of pleiotropic effects or close linkage of genes were the main causes for correlations among traits which resulted in selection for one will result in a correlated response with another.
2. QTL mapping studies using age-specific measures clearly demonstrated distinct age-specific gene action. In our study, using unconditional and conditioanl mapping methods, there were 23 QTLs with additive and additive x treatment effects for seedling height, 22 QTLs for root length, 14 QTLs for leaf number, and 23 QTLs for seedling weight, respectively. But
only 3, 1,0, and 2 QTLs with significant additive effects at all measuring stages for seedling height, root length, leaf number, and seedling weight, respectively. And other QTLs could be detected only at one or several stages. This suggested that genes controlling these seedling traits expressed selectively at different stages. Conditional mapping method could estimate and detect the net effects of gene expression that were independent of the causal genetic effects in earlier stages. And could reveal the gene expression pattern efficiently at different stage. This technique could serve as an important tool for studying the inheritance of response to stress.
3. The present study showed the importance of epistasis as a genetic basis of the quantitative traits and revealed several important features of this phenomenon. In the study, with two mapping methods, there were 30 pairs of QTLs involved in epistasis for seedling height, 36 pairs for root length, 33 pairs for leaf number, and 25 pairs for seedling weight with proportion of 50%, 47%, 62%, and 42% for each seedling traits, respectively, without additive and/or additive x treatment interaction effects. Moreover, it was common for several loci to interact with more than one non-allelic locus at the same stage and/or across different stages, and, the interactions often showed differences in the direction of effects. It was indicated that the possibility of multilocus associations existed for trait development. Beside this, a lot of interactions could detected only by conditional mapping method indicated that many epistasis interactions
引文
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