水稻RIL群体SSR标记遗传图谱构建和耐冷相关性状QTL的定位
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摘要
水稻是我国最重要的粮食作物，培育高产、优质和多抗的水稻品种是育种家所期望的，也是我国水稻生产的实际需要。由于水稻产量、品质和抗逆性大多数是复杂的数量性状，挖掘重要农艺性状相关的优异基因资源，并通过分子水平的定位，可为基因克隆和利用分子标记辅助选择技术进行水稻育种提供重要的基础。
本研究利用一个耐冷的粳稻品种大关稻与一个冷敏感的籼稻品种IR_(28)，通过杂交和F_2代单粒传法获得了295个重组自交家系群体(F_6)，随机选取227个家系(F_7)进行分子标记连锁图谱的构建，并进一步对苗期耐冷性和低温发芽力两个性状进行QTL的定位。主要研究过程和结果描述如下：
1、选择均匀覆盖水稻整个基因组的SSR引物1095对，筛选到在两亲本间呈多态性的引物269对，选带型清晰且在基因组中分布较均匀的标记构建了一个含167个SSR分子标记的连锁图谱。连锁群总长度为1837.2cM，标记间平均距离为11.0cM。群体标记均匀地分布在12条染色体上。根据对单标记位点分离的卡方检验结果，在构建的遗传图谱上发现有多个偏分离热点区域。利用一种新的基于多位点的统计方法对标记遗传图谱进行调整，根据调整的结果对偏分离座位(SDL)进行了定位，在12条染色体上发现有29个偏分离座位(SDL)。
2、利用建立的分子标记遗传图谱对水稻苗期耐冷性和低温发芽力性状进行QTL定位。表型鉴定在人工气候箱中进行，对二叶一心期的幼苗(播种后13天)在9℃的低温下冷处理5天，然后逐步恢复到25℃，5天后，根据死苗率进行耐冷分级，鉴定亲本和RIL群体的耐冷性。两亲本耐冷性差异显著(大关稻2级，IR_(28)为9级)，RILs间呈连续变异。在第8、11染色体上共检测到3个苗期耐冷QTLs。位于11号染色体上的qSCT-11-1能解析23.0％的表型变异，是主效基因，位于标记RM26281和RM3701之间；在qSCT-11-1旁侧检测到另一个QTL—qSCT-11-2，能解析10.1％的表型变异，位于标记RM7248和RM5731之间，与qSCT-11-1紧密连锁；在第8染色体的标记RM22491和RM22694之间检测到的qSCT8能解析8.1％的表型变异。在三个苗期耐冷QTL位置，来自粳稻种质大关稻的等位基因都提高苗期耐冷性。
3、利用建立的分子标记遗传图谱对水稻低温发芽性进行QTL定位，在14℃下鉴定亲本和RIL群体各家系在浸种后第8、10、12天的发芽率．在发芽的第8、10、12天，两亲本表现明显差异，RILs间呈连续变异。在第1、11染色体上共检测到2个低温发芽相关QTL。浸种后8天时检测到位于11号染色体上的qLTG-11能解析24.5％的表型变异。浸种后10天和12天时在第1染色体上都检测到一个以前未检测到的QTL，即qLTG-1；分别能解析18.4％和16.8％的表型变异。来自IR_(28)的qLTG-1位置的等位基因和来自大关稻qLTG-11位置的等位基因均能提高低温发芽能力。
Rice is one of the most critical cereals in China. It has been expected that breeding of varieties with high yield, good quality and multiple resistance by breeders and farmers. Because of the quantitatively-genetic nature of the traits related to yield, quality and resistancees to various stresses, it is fundamental to explore special resourcees with elite agronomic traits, mapping corresponding position by molecular markers, cloning of corresponding genes and breeding of rice by MAS (Molecular marker-assisted selection) .
In this study 227 lines (F7) were randomly selected among a RILs (Recombination Inbreeding Lines) population extracted from a cross between IR_(28), a cold-susceptible indica variety, and Da-guan-Dao, a cold-tolerance japanica, and used to construct a molecular linkage map. With this map the cold tolerance of rice seedling and seed germination ability under low temperature were genetically analyzed. The main results were described as follows:
1. Total 1095 SSR (Simple Sequence Repeat) markers were used to screen the polymorphism between two parents (IR_(28) and Da-Guan-Dao). Among 269 SSR markers with polymorphism, 167 were finally used to construct a genetic linkage map in a population of 227 lines. It spans 1837.2 cM in length covering all 12 chromosomes, with an average density of 11.0cM. Several distorted segregation regions was detected according to Chi-value of all 167 marker loci. Then the genetic map was tentatively adjusted with a multiple-point method. Based on the adjusted map 29 SDLs (Segregation-distorted loci) were located on 12 chromosomes.
2. The seedlings of 227 lines with 2-3 leaves were subjected to 9℃for 5 days and subsequently slow recovery of 5 days in growth chambers. The scoring was done to characterize the cold tolerance of two parents and RILs (Fs) according to the percentage of dead seedling after recovery. The result show that DaGuanDao was tolerant with a score of 2 while IR_(28) susceptible, with a score of 9. The RIL population exhibited all range of scores from 1-9, presenting continuous distribution. It was detected that three QTLs accounting for 38.1% of total phenotypic variation located on 8 and 11 chromosomes respectively. The qSCT-11-1, a major QTL, located between RM26281 and RM3701 on 11 chromosome accounted for 23.0% of total phenotypic variation. The qSCT-11-2 located between RM7248 and RM5731 closely flanking qSCT-11-1 explained 10.1% of total phenotypic variation. The qSCT-11-2 located on 11 chromosome flanked by RM22491 and RM22694 explained 8.1% of total phenotypic variation. All three QTL alleles from japonica parent decreased the percentage of seedling death.
3. In order to map QTLs conferring low temperature germinability (LTG) it was observed germination rate percentage at 14°C for 8、10、12 days of two parents and 227 RILs (Fg) after soaking. Germination rate percentage at 14°C for 8% 10% 12 days of two parents were obviously different and that of the RILs distributed continuously. Two Putative QTLs on chromosome 1 and 11, associated with low-temperature germination were detected. The total phynotypic variation explained by qLTG-1, a newly-detected QTL at 10 day and 12 day on chromosome 1 were 18.4% and 16.8% respectively. The additional QTL, qLTG-11, detected at 8 day on chromosome 11 accounted for 24.5% of the total phenotypic variation for low-temperature germinability. Both alleles in qLTG-1 From IR 28 and that at the region of qLTG-11 from DaGuanDao increased the low-temperature germination rate.
本研究利用一个耐冷的粳稻品种大关稻与一个冷敏感的籼稻品种IR_(28)，通过杂交和F_2代单粒传法获得了295个重组自交家系群体(F_6)，随机选取227个家系(F_7)进行分子标记连锁图谱的构建，并进一步对苗期耐冷性和低温发芽力两个性状进行QTL的定位。主要研究过程和结果描述如下：
1、选择均匀覆盖水稻整个基因组的SSR引物1095对，筛选到在两亲本间呈多态性的引物269对，选带型清晰且在基因组中分布较均匀的标记构建了一个含167个SSR分子标记的连锁图谱。连锁群总长度为1837.2cM，标记间平均距离为11.0cM。群体标记均匀地分布在12条染色体上。根据对单标记位点分离的卡方检验结果，在构建的遗传图谱上发现有多个偏分离热点区域。利用一种新的基于多位点的统计方法对标记遗传图谱进行调整，根据调整的结果对偏分离座位(SDL)进行了定位，在12条染色体上发现有29个偏分离座位(SDL)。
2、利用建立的分子标记遗传图谱对水稻苗期耐冷性和低温发芽力性状进行QTL定位。表型鉴定在人工气候箱中进行，对二叶一心期的幼苗(播种后13天)在9℃的低温下冷处理5天，然后逐步恢复到25℃，5天后，根据死苗率进行耐冷分级，鉴定亲本和RIL群体的耐冷性。两亲本耐冷性差异显著(大关稻2级，IR_(28)为9级)，RILs间呈连续变异。在第8、11染色体上共检测到3个苗期耐冷QTLs。位于11号染色体上的qSCT-11-1能解析23.0％的表型变异，是主效基因，位于标记RM26281和RM3701之间；在qSCT-11-1旁侧检测到另一个QTL—qSCT-11-2，能解析10.1％的表型变异，位于标记RM7248和RM5731之间，与qSCT-11-1紧密连锁；在第8染色体的标记RM22491和RM22694之间检测到的qSCT8能解析8.1％的表型变异。在三个苗期耐冷QTL位置，来自粳稻种质大关稻的等位基因都提高苗期耐冷性。
3、利用建立的分子标记遗传图谱对水稻低温发芽性进行QTL定位，在14℃下鉴定亲本和RIL群体各家系在浸种后第8、10、12天的发芽率．在发芽的第8、10、12天，两亲本表现明显差异，RILs间呈连续变异。在第1、11染色体上共检测到2个低温发芽相关QTL。浸种后8天时检测到位于11号染色体上的qLTG-11能解析24.5％的表型变异。浸种后10天和12天时在第1染色体上都检测到一个以前未检测到的QTL，即qLTG-1；分别能解析18.4％和16.8％的表型变异。来自IR_(28)的qLTG-1位置的等位基因和来自大关稻qLTG-11位置的等位基因均能提高低温发芽能力。
Rice is one of the most critical cereals in China. It has been expected that breeding of varieties with high yield, good quality and multiple resistance by breeders and farmers. Because of the quantitatively-genetic nature of the traits related to yield, quality and resistancees to various stresses, it is fundamental to explore special resourcees with elite agronomic traits, mapping corresponding position by molecular markers, cloning of corresponding genes and breeding of rice by MAS (Molecular marker-assisted selection) .
In this study 227 lines (F7) were randomly selected among a RILs (Recombination Inbreeding Lines) population extracted from a cross between IR_(28), a cold-susceptible indica variety, and Da-guan-Dao, a cold-tolerance japanica, and used to construct a molecular linkage map. With this map the cold tolerance of rice seedling and seed germination ability under low temperature were genetically analyzed. The main results were described as follows:
1. Total 1095 SSR (Simple Sequence Repeat) markers were used to screen the polymorphism between two parents (IR_(28) and Da-Guan-Dao). Among 269 SSR markers with polymorphism, 167 were finally used to construct a genetic linkage map in a population of 227 lines. It spans 1837.2 cM in length covering all 12 chromosomes, with an average density of 11.0cM. Several distorted segregation regions was detected according to Chi-value of all 167 marker loci. Then the genetic map was tentatively adjusted with a multiple-point method. Based on the adjusted map 29 SDLs (Segregation-distorted loci) were located on 12 chromosomes.
2. The seedlings of 227 lines with 2-3 leaves were subjected to 9℃for 5 days and subsequently slow recovery of 5 days in growth chambers. The scoring was done to characterize the cold tolerance of two parents and RILs (Fs) according to the percentage of dead seedling after recovery. The result show that DaGuanDao was tolerant with a score of 2 while IR_(28) susceptible, with a score of 9. The RIL population exhibited all range of scores from 1-9, presenting continuous distribution. It was detected that three QTLs accounting for 38.1% of total phenotypic variation located on 8 and 11 chromosomes respectively. The qSCT-11-1, a major QTL, located between RM26281 and RM3701 on 11 chromosome accounted for 23.0% of total phenotypic variation. The qSCT-11-2 located between RM7248 and RM5731 closely flanking qSCT-11-1 explained 10.1% of total phenotypic variation. The qSCT-11-2 located on 11 chromosome flanked by RM22491 and RM22694 explained 8.1% of total phenotypic variation. All three QTL alleles from japonica parent decreased the percentage of seedling death.
3. In order to map QTLs conferring low temperature germinability (LTG) it was observed germination rate percentage at 14°C for 8、10、12 days of two parents and 227 RILs (Fg) after soaking. Germination rate percentage at 14°C for 8% 10% 12 days of two parents were obviously different and that of the RILs distributed continuously. Two Putative QTLs on chromosome 1 and 11, associated with low-temperature germination were detected. The total phynotypic variation explained by qLTG-1, a newly-detected QTL at 10 day and 12 day on chromosome 1 were 18.4% and 16.8% respectively. The additional QTL, qLTG-11, detected at 8 day on chromosome 11 accounted for 24.5% of the total phenotypic variation for low-temperature germinability. Both alleles in qLTG-1 From IR 28 and that at the region of qLTG-11 from DaGuanDao increased the low-temperature germination rate.
引文
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