应用回交育种和QTL聚合改良粳稻抗旱性
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摘要
在世界上许多地区干旱胁迫和水资源短缺是威胁作物生产最重要的限制因素,在我国淡水资源中大约有70%被用于农业,而水稻生产的用水量占全部农业用水的70%。在我国北方,人们对高品质粳稻日益增长的强烈需求使得近几年水稻种植面积急剧增加,这使得竞争有限的水资源成为这里一个严重问题。选育和种植高产、抗旱或节水粳稻新品种成为解决这一问题的最佳途径。因而,育种家们在选育抗旱性强或水分利用效率高的高产粳稻品种时所面临的两大挑战是:对改良抗旱性(或水分利用效率)有利用价值的遗传变异在哪里;应该采取怎样的育种策略。
本研究采用同样的回交育种策略挖掘籼稻基因库资源中的抗旱隐蔽基因,并采取聚合来源于不同供体抗旱基因的方法,以我国北方粳稻优良恢复系C418为轮回亲本,与5个籼稻供体亲本进行杂交和2次回交,对BC_2F_2混合群体进行严格的干旱筛选,共得到5个抗旱选择群体(113个抗旱选择导入系),对其选择导入效应进行了综合评价,对其中3个群体进行了抗旱QTL定位和效应分析;另应用8个优良抗旱选择导入系配制4个聚合杂交组合,于F_2代进行严格的抗旱筛选,构建抗旱聚合系群体,并对其进行了抗旱QTL定位和抗旱遗传网络剖析。获得主要试验结果如下:
1.对5个抗旱导入系群体抗旱选择导入效应进行了综合评价。其主要试验结论:导入系后代经过两个不同环境的鉴定,筛选出46(40.7%)株优良导入系,这是应用回交方法开发籼稻基因资源中隐蔽有利变异改良粳稻抗旱性的育种策略的有力证据;在这些株系的变化中株高的增加和抽穗期的缩短是两个相关的变化,这说明抗旱可能与GA途径有关,也说明是逃旱性的抗旱;这些导入系的产量构成因子中存在着大量变异,可以说明不同抗旱机制的存在;本研究产生的C418导入系既可以为剖析抗旱性与产量的关系提供有用材料,又可以作为进一步聚合育种的中间材料。
2.抗旱选择导入群体QTL定位及其效应分析。其主要试验结论:卡方检验时发现较小规模的选择导入系群体,例如C418/manauthukha群体和C418/teqing群体检测到的超导入位点比例较大,而较大群体C418/yuexiangzhan检测到的超导入位点比例较小,因此,为提高选择效率适当的群体大小是我们进行选择时应该考虑的问题;定位到的许多抗旱QTL在两个群体中同时存在,这些位点相对更加可靠,但是这些位点的基因往往功能相近,在聚合育种时应该尽量避免选择重复功能的供体片段,对于提高改良效果更加有利;定位结果中发现一些对产量具有正向加性效应的QTL,这些位点将会被用于进一步改良抗旱性的育种程序中,同时发现了四种条件下对产量均表现正向显性效应的位点,这对研究QTL的显性效应十分有利,并且对于杂交水稻育种来说更有应用价值。
3.抗旱聚合系QTL定位和抗旱遗传网络剖析。其主要试验结论:每个聚合群体卡方检验得到的抗旱相关QTL与单向方差分析得到的产量相关QTL都有一些位点是重叠的,这是我们所期望的结果,同时也验证了导入系群体QTL效应分析的结论,有些抗旱位点是对产量具有正向加性效应的,有些抗旱位点对产量是存在负向加性效应的。抗旱遗传网络分析得出群体Ⅰ中AL1处于总调控位置,AL2、AL3、AL4、M1、M3、M4、M5和M6处于调控关系上游;群体Ⅱ中AL1处于总调控位置,AL2、M1、M2、M3处于调控关系上游;群体Ⅲ中AL1处于总调控位置,M1、M2、M3、AL2和AL3处于调控关系上游;群体Ⅳ中AL1处于总调控位置,M1、M2、M3、M4、M5、M6、M7、M8、M9、M10、M12、M14和M16处于调控关系上游,这些基因对于抗旱遗传网络的调控力度最大,对整个抗旱体系的作用举足轻重,在育种选择过程当中要注意对这些导入位点的跟踪选择。
Drought stress and water shortage are the most important factors threatening crop production in many areas of the world. In China, agriculture uses～70% of its total fresh water resources and rice production alone takes up～70% of the water resources for agriculture. In North China, rice acreage has nearly doubled recently as the dramatically increased demand for high quality japonica rice in China. This has created a serious problem in competing the limited water resources in the area. Thus, development and adoption of high yielding, drought tolerant (DT) or water-use-efficient (WUE) japonica varieties are the most attractive solution to this problem. Thus, two major challenges facing plant breeders for developing high yielding japonica rice cultivars with desirable levels of DT and WUE are: where is the useful genetic variation for DT/WUE and what breeding strategy should be used.
In this study, we used BC breeding to improve japonica hybrid restorer line, C418 by mining the diversity hidden in the indica gene pool. Five BC_2F_2 bulk populations were developed by crossing C418 (the recurrent parent) with 5 indicia donors followed by two rounds backcrossing. The BC_2F_2 populations were subjected to two rounds of selection under severe drought, resulting in the development of 113 DT introgression lines (ILs). Evaluation was made for the lines about introgression effects. Also, we mapped the QTLs for drought tolerance and analyzed the effects of DT QTLs. Furthermore, we conducted 4 pyramiding populations using 8 superior lines improved for DT followed by severe drought selection performed for F_2. QTLs mapping and analyzing of DT genetic network were made for pyramiding DT lines. The main results summarized as follows:
1. Evaluation was made for 113 lines about introgression effects. Progeny testing across 2 diverse environments allowed the identification of 46 (40.7%) superior C418-ILs, demonstrating the power of BC breeding in exploiting favorable variation hidden in the indica gene pool for improving DT of japonica rice. Increased-height and accelerated-heading were 2 correlated changes in these ILs, suggesting the possible involvement of GA pathway and drought escaping. Considerable variations in yield components among these ILs indicate the presence of different DT mechanisms. The C418-ILs developed in this study were supposed to provide useful materials for dissection of DT and yield, and for further pyramiding breeding in rice.
2. We mapped the QTLs for drought tolerance of 113 lines and analyzed the effects of DT QTLs.We found that the percent of super introgressin loci in small populations were much larger than that in large population. So we should consider the proper size of populations when we select the DT lines. Many QTLs mapped in the 113 lines were concurrence in two populations and these loci were more reliable for breeder. But, the gene functions of these loci are mostly close and we have to avoid selecting these loci to increase the improving effects when we perform pyraming breeding. We also found the DT QTLs with positive additive effect for yield and these QTLs will be used to improve DT further. At the same time, somes QTLs with positive dominant effect for yield were mapped under 4 conditions and these QTLs will be used to study dominant effect of QTLs further which are very useful for Hybrid rice breeding.
3. QTLs mapping and analyzing of DT genetic network were made for pyramiding DT lines. DT QTLs of X~2 test and yield QTLs of One-way ANOVA were found overlapping in every population which was desired by us. The results also verified the conclusion that some DT QTLs were positive additive effect for yield and another DT QTLs were negative additive effect for yield. Genetic networks of pyramiding populations on DT indicated that AL1s were all in the chief positions in 4 populations and AL2, AL3, AL4, M1, M3, M4, M5 and M6 of population I , AL2, M1, M2 and M3 of population II, M1, M2, M3, AL2 and AL3 of population III, M1,M2,M3,M4,M5,M6,M7, M8, M9,M10,M12,M14 and M16 of population IV were in the upstream of the relationship which had the highest regulating ability to the genetic network of DT. The functions of these locus or genes are very essential in the DT genetic network. We should pay attentions to track and select these locus in our breeding practice.
本研究采用同样的回交育种策略挖掘籼稻基因库资源中的抗旱隐蔽基因,并采取聚合来源于不同供体抗旱基因的方法,以我国北方粳稻优良恢复系C418为轮回亲本,与5个籼稻供体亲本进行杂交和2次回交,对BC_2F_2混合群体进行严格的干旱筛选,共得到5个抗旱选择群体(113个抗旱选择导入系),对其选择导入效应进行了综合评价,对其中3个群体进行了抗旱QTL定位和效应分析;另应用8个优良抗旱选择导入系配制4个聚合杂交组合,于F_2代进行严格的抗旱筛选,构建抗旱聚合系群体,并对其进行了抗旱QTL定位和抗旱遗传网络剖析。获得主要试验结果如下:
1.对5个抗旱导入系群体抗旱选择导入效应进行了综合评价。其主要试验结论:导入系后代经过两个不同环境的鉴定,筛选出46(40.7%)株优良导入系,这是应用回交方法开发籼稻基因资源中隐蔽有利变异改良粳稻抗旱性的育种策略的有力证据;在这些株系的变化中株高的增加和抽穗期的缩短是两个相关的变化,这说明抗旱可能与GA途径有关,也说明是逃旱性的抗旱;这些导入系的产量构成因子中存在着大量变异,可以说明不同抗旱机制的存在;本研究产生的C418导入系既可以为剖析抗旱性与产量的关系提供有用材料,又可以作为进一步聚合育种的中间材料。
2.抗旱选择导入群体QTL定位及其效应分析。其主要试验结论:卡方检验时发现较小规模的选择导入系群体,例如C418/manauthukha群体和C418/teqing群体检测到的超导入位点比例较大,而较大群体C418/yuexiangzhan检测到的超导入位点比例较小,因此,为提高选择效率适当的群体大小是我们进行选择时应该考虑的问题;定位到的许多抗旱QTL在两个群体中同时存在,这些位点相对更加可靠,但是这些位点的基因往往功能相近,在聚合育种时应该尽量避免选择重复功能的供体片段,对于提高改良效果更加有利;定位结果中发现一些对产量具有正向加性效应的QTL,这些位点将会被用于进一步改良抗旱性的育种程序中,同时发现了四种条件下对产量均表现正向显性效应的位点,这对研究QTL的显性效应十分有利,并且对于杂交水稻育种来说更有应用价值。
3.抗旱聚合系QTL定位和抗旱遗传网络剖析。其主要试验结论:每个聚合群体卡方检验得到的抗旱相关QTL与单向方差分析得到的产量相关QTL都有一些位点是重叠的,这是我们所期望的结果,同时也验证了导入系群体QTL效应分析的结论,有些抗旱位点是对产量具有正向加性效应的,有些抗旱位点对产量是存在负向加性效应的。抗旱遗传网络分析得出群体Ⅰ中AL1处于总调控位置,AL2、AL3、AL4、M1、M3、M4、M5和M6处于调控关系上游;群体Ⅱ中AL1处于总调控位置,AL2、M1、M2、M3处于调控关系上游;群体Ⅲ中AL1处于总调控位置,M1、M2、M3、AL2和AL3处于调控关系上游;群体Ⅳ中AL1处于总调控位置,M1、M2、M3、M4、M5、M6、M7、M8、M9、M10、M12、M14和M16处于调控关系上游,这些基因对于抗旱遗传网络的调控力度最大,对整个抗旱体系的作用举足轻重,在育种选择过程当中要注意对这些导入位点的跟踪选择。
Drought stress and water shortage are the most important factors threatening crop production in many areas of the world. In China, agriculture uses～70% of its total fresh water resources and rice production alone takes up～70% of the water resources for agriculture. In North China, rice acreage has nearly doubled recently as the dramatically increased demand for high quality japonica rice in China. This has created a serious problem in competing the limited water resources in the area. Thus, development and adoption of high yielding, drought tolerant (DT) or water-use-efficient (WUE) japonica varieties are the most attractive solution to this problem. Thus, two major challenges facing plant breeders for developing high yielding japonica rice cultivars with desirable levels of DT and WUE are: where is the useful genetic variation for DT/WUE and what breeding strategy should be used.
In this study, we used BC breeding to improve japonica hybrid restorer line, C418 by mining the diversity hidden in the indica gene pool. Five BC_2F_2 bulk populations were developed by crossing C418 (the recurrent parent) with 5 indicia donors followed by two rounds backcrossing. The BC_2F_2 populations were subjected to two rounds of selection under severe drought, resulting in the development of 113 DT introgression lines (ILs). Evaluation was made for the lines about introgression effects. Also, we mapped the QTLs for drought tolerance and analyzed the effects of DT QTLs. Furthermore, we conducted 4 pyramiding populations using 8 superior lines improved for DT followed by severe drought selection performed for F_2. QTLs mapping and analyzing of DT genetic network were made for pyramiding DT lines. The main results summarized as follows:
1. Evaluation was made for 113 lines about introgression effects. Progeny testing across 2 diverse environments allowed the identification of 46 (40.7%) superior C418-ILs, demonstrating the power of BC breeding in exploiting favorable variation hidden in the indica gene pool for improving DT of japonica rice. Increased-height and accelerated-heading were 2 correlated changes in these ILs, suggesting the possible involvement of GA pathway and drought escaping. Considerable variations in yield components among these ILs indicate the presence of different DT mechanisms. The C418-ILs developed in this study were supposed to provide useful materials for dissection of DT and yield, and for further pyramiding breeding in rice.
2. We mapped the QTLs for drought tolerance of 113 lines and analyzed the effects of DT QTLs.We found that the percent of super introgressin loci in small populations were much larger than that in large population. So we should consider the proper size of populations when we select the DT lines. Many QTLs mapped in the 113 lines were concurrence in two populations and these loci were more reliable for breeder. But, the gene functions of these loci are mostly close and we have to avoid selecting these loci to increase the improving effects when we perform pyraming breeding. We also found the DT QTLs with positive additive effect for yield and these QTLs will be used to improve DT further. At the same time, somes QTLs with positive dominant effect for yield were mapped under 4 conditions and these QTLs will be used to study dominant effect of QTLs further which are very useful for Hybrid rice breeding.
3. QTLs mapping and analyzing of DT genetic network were made for pyramiding DT lines. DT QTLs of X~2 test and yield QTLs of One-way ANOVA were found overlapping in every population which was desired by us. The results also verified the conclusion that some DT QTLs were positive additive effect for yield and another DT QTLs were negative additive effect for yield. Genetic networks of pyramiding populations on DT indicated that AL1s were all in the chief positions in 4 populations and AL2, AL3, AL4, M1, M3, M4, M5 and M6 of population I , AL2, M1, M2 and M3 of population II, M1, M2, M3, AL2 and AL3 of population III, M1,M2,M3,M4,M5,M6,M7, M8, M9,M10,M12,M14 and M16 of population IV were in the upstream of the relationship which had the highest regulating ability to the genetic network of DT. The functions of these locus or genes are very essential in the DT genetic network. We should pay attentions to track and select these locus in our breeding practice.
引文
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