粳稻回交导入群体耐逆性筛选及QTL剖析
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摘要
盐害、冷害和高温是水稻生产面临的主要非生物胁迫。本研究以粳型超优1号为轮回亲本与11个来自不同国家的供体品种杂交和回交培育的BC2F4群体为材料，进行苗期及分蘖至成熟期耐盐、开花期耐热、孕穗期耐冷的鉴定与筛选，对目标性状的选择导入系进行正常和胁迫条件下的产量相关性状评价，剖析抗逆性的生理机制，并利用耐冷选择导入系进行耐冷QTL定位，获得了以下主要结果。
1.对回交群体进行苗期(2-3叶)盐胁迫(140mmol·L-1NaCl)筛选，经过初筛共筛选到125个耐盐单株，入选率为2.58%。对52个耐盐株系进行形态和离子的测定，结果表明：盐胁迫导致了根长、根鲜重和根干重的增加；苗长、地上鲜重和地上干重的显著减少，且根部变化幅度小于地上部，说明幼苗地上部分的反应比根部对盐害的反应敏感。在离子效应上，盐胁迫导致了地上部和根部钠离子的显著增加，其中地上部分增加了20倍以上，导入系增加的量小于轮回亲本。钾离子的吸收与钠离子相反，胁迫下减少了对钾离子的吸收，但导入系比轮回亲本表现吸收较多的钾离子。所以导入系耐盐性的增强在离子效应上主要表现为保持较低的叶片钠离子浓度、较低的钠离子转运和增强对钾离子的吸收来降低盐害对水稻生长的抑制和毒害作用。
2.对回交群体进行分蘖至成熟期的盐池(140mmol·L-1NaCl)筛选，筛选到150个耐盐导入系。其中籼、粳稻供体回交后代的在入选率和结实率上分别为6.9%和2.6%，表明籼、粳供体的回交后代在耐盐性上存在差异。耐盐导入系间具有不同的生理形态表现：一部分株系表现为抽穗期提早，属于避盐机制；另一部分株系表现为具有较高的株高、有效穗数和生物产量的株系，属于稀盐机制；还有一些株系表现为千粒重、每穗实粒数、结实率的显著提高，属于忍盐机制。机理间并不是独立的，一些导入系同时具有两种或两种以上的耐盐机制，这些机制间相互协调，彼此促进，提高耐盐性。
3.对回交群体进行开花期高温胁迫，筛选到124个耐高温单株，粳稻比籼稻供体的回交群体出现耐热个体数多，表明粳稻资源中同样存在耐热有利基因。导入系在两种环境下均出现广幅分离，在胁迫和正常条件下分别有80个和47个株系的结实率显著高于轮回亲本，从中鉴定出的耐热性和产量性状均显著好于轮回亲本的优良导入系有8个。对3个F2聚合群体耐高温筛选分别获得7、31和68株极显著好于聚合亲本的耐热个体，平均结实率在80%以上，显著高于聚合亲本和轮回亲本，显示出较理想的耐热聚合效果。
4.对回交群体进行孕穗期的冷水胁迫处理，筛选到324个耐冷单株。经苗期和孕穗期耐冷鉴定，其耐冷性比轮回亲本有显著提高的株系数分别为40个和242个。对116个强耐冷株系进行重复鉴定，在胁迫条件下，结实率和单株产量显著高于轮回亲本的导入系分别有94个和77个；正常条件下，结实率和单株产量显著高于轮回亲本的导入系分别有39个和34个，并且鉴定出18个耐冷性和产量性状都显著好于轮回亲本的优良导入系。而且对目标性状的选择还导致了株高、生物产量、每穗实粒数等相关联性状的显著提高。
5.利用以超优1号为轮回亲本，Chhomromg、Doddi和丰矮占1号为供体筛选获得的回交选择耐冷导入系进行卡方检测的耐冷QTL定位，同时对随机群体进行耐冷评价和方差分析定位耐冷QTL。采用耐冷极端选择导入系定位到29个QTL，方差分析检测到16个耐冷QTL。发现耐冷QTL主要有两种类型：一是影响随机群体冷胁迫与正常条件下结实率差值的QTL，这些等位基因具有提高冷胁迫下性状表达的稳定性，对增强耐冷性有贡献，共检测到8个QTL，分布在第3、5、6、7和8染色体上；第二类是在不同的选择群体中经卡方检测到的相同耐冷QTL，其受假阳性的影响相对较小，共有3个QTL，分别位于第2、3和9染色体上。本研究检测到的耐冷QTL位点，大多与前人发现的位点吻合，说明本研究利用选择群体进行QTL定位的方法有效。试验材料的获得是在高的选择压力下进行的，这些耐冷选择导入系将是生产上的育种群体，本研究将育种群体和基因定位群体相联系，缩短了遗传理论研究与育种实践的距离，对育种技术路线提供了有益的补充。
研究表明，不同供体的回交后代中均出现耐盐、耐冷和耐热的超亲分离，表明这些供体均带有控制这些性状的有利“隐蔽”基因，但各个组合间的抗逆性筛选效率存在较大差异，抗逆性表现不仅取决于供体本身，还取决于受体品种与供体品种的不同组合。本研究筛选出的导入系将成为进行大规模有利基因发掘以及通过有利基因的高效聚合来定向改良抗逆性分子育种的材料平台。
Abiotic stresses such as salinity, cold and heat are major threat to rice production. ElevenBC2F4backcross introgression populations derived from crosses between a japonica varietyChaoyou1as the recurrent parent and11donor parents collected from different countries werescreened for salt tolerance (ST) at the seedling and from tillering to maturity stages, heattolerance (HT) at flowering stage and cold tolerance (CT) at booting stage. The targetintrogression lines (ILs) were evaluated for yield and its related traits under normal and stresscondition to dissect physiological mechanisms of stress tolerance and map QTL for CT. Themain results were described as follows.
1. Eleven BC2F4backcross introgression populations were screened for salt tolerance (ST)at seedling stage. One hundred and twenty-five (125) ILs were selected on the basis of singleplant SDS under salt water (140mmol·L-1NaCl).12,11,21and10ILs selected from four ofthe BC populations including donors Chouyou1/X21, Chouyou1/Q5, Chouyou1/Chhomrongand Chouyou1/Doddi were evaluated for morphological characteristics and ion concentrationsin replicated experiments under stress and normal conditions. The ILs had increased rootlength, root fresh weight, root dry weight but reduced shoot length, shoot fresh weight, shootdry weight under salt stress. The amplitudes of variations in the root traits were smaller thanthose of shoot traits, indicating that shoots were more sensitive to salt stress. With regard toionic effect, salt stress resulted in accumulation of Na+and reduction of K+in the whole plant.Compared to the recurrent parent, the increase of amplitude of Na+and the reduction of Kwere observably lower for the ILs. Coinstantaneous accumulation of20times more Na+in theshoot after salt stress was observed. Most of the salinity tolerant ILs improve their salinitytolerances through lower shoot Na+concentration and lower root to shoot Na+translocation.
2. Eleven BC2F4backcross introgression populations were screened for salt tolerance (ST)at different developmental stages from tillering to maturity. One hundred and fifty (150) ILswere selected on the basis of single plant grain yield (GY) and spikelet fertility (SF) under saltwater (140mmol·L-1NaCl). Our results clearly showed that there was a rich hidden geneticvariation for ST in the donor lines and phenotypic selection under stress condition was apowerful way to identify transgressive segregaantis from the segregating BC populations.Further evaluation of126ILs selected from four BC populations in replicated experimentsunder stress and normal conditions identified10promising ILs with greatly improved ST and higher yield than CY1. Selection under salt stress for yield and SF, supplemented by acombination of desirable secondary traits, can greatly improve the efficiency and accuracy ofbreeding for high salt-tolerance. It is important to point out that three mechanisms includingosmotic adjustment, salt exclusion and tissue tolerance appeared to function together andaffect the same suite of ST related traits in the ILs; although their contributions to specific STtraits were very different for different ILs depending on the specific scenarios of stress.
3. Eleven BC2F4backcross introgression populations were screened for heat tolerance(HT) at flowering stage. Total of124ILs were selected, based on single plant SF under hightemperature (38℃). The results showed that the ILs had better tolerance to heat (HT) than therecurrent parent, Chaoyou1, and the frequencies of plants with higher HT were higher in thepopulations derived from japonica donors were higher than those from indica ones,suggesting that japonica varieties were better sources of desirable genes for improving HT.Progeny testing of the124ILs under the HT stress and normal conditions revealed that64.5%(80) of the lines had better HT than the RP, as measured by SF. The124selected ILspresented wide segregations for other measured traits as well. Eight ILs with greatly improvedHT and higher yield than CY1were selected. HT plants selected from three pyramidingpopulations had an average SF of more than80%, which was significantly higher than the RPand the parents used in pyramiding. One hundred and six (106) plants with significant higherHT than the pyramiding parental lines were selected
4. ILs from the eleven diverse donors were screened for CT at the reproductive stageunder the low temperature (LT) treatment created by irrigating the plants with cold water(19℃) at the reproductive stage. Three hundred and twenty four (324) BC2F5introgressionlines (ILs) were selected based on single plant SF. Progeny testing of the324ILs under thesimilar LT stress revealed that the efficiency of selection for CT was0.794. Further evaluationof116ILs selected from five BC populations in replicated experiments under LT and normalconditions identified18promising ILs with greatly improved CT and yield than CY1.Detailed comparisons between the ILs and CY1were conducted for yield and related traitsunder the LT stress and non-stress conditions to gain useful information and betterunderstanding of important issues related to breeding for CT using introgression breedingsuch as donor selection, selection efficiency, screening method and genetic drag fornon-target traits in the BC breeding process.
5. ILs were assessed to identify cold-tolerance quantitative trait loci (QTL) from thedonors including Chhomromg, Doddi and Fengaizhan1in the genetic background of japonicacultivar Chaoyou1. X2test was used for QTL identification for the CT-selected ILs, meanwhile, random populations were assessed to identify CT-QTL by single marker analysismethod. Total of29QTL associated with CT were detected by the selected ILs and16identified by single marker analysis. The CT-QTL were classified into two types, one are eightQTL affecting trait difference for seed setting rate of stress to control, which distributedchromosomes3,5~8and the alleles at the QTL improved trait stability under stress andactually contributed to CT. The second type is the three common QTL detected by theselected ILs from different populations with less negative chance of QTL detection, whichdistributed chromosomes2,3and9. Furthermore, most of the QTLs identified in this studywere consistent with those documented. Therefore, QTL detection using selected ILs based onwell-known X2test is a simple and effective. previously by other studies.. The use ofintrogression populations in QTL identification and selection for outstanding lines providesan efficient way for integrating gene discovery and practical breeding and as a result speed upthe application modern molecular techniques in breeding for complex traits.
The present study indicated that transgressive segregations were observed for ST, HTand CT in most of the backcross populations, indicating that all the11donors possesfavorable ‘hidden’ alleles for tolerances to the studied stresses. There were significantdifferences among populations in selection efficiency, reflecting stress tolerance depends notonly on donor itself but also on different combinations of the recurrent parent and the donorparent. The stress tolerant ILs will be partially a platform and valuable materials for favorableallele mining and improvement of stress tolerance by pyramiding of favorable genes.
1.对回交群体进行苗期(2-3叶)盐胁迫(140mmol·L-1NaCl)筛选，经过初筛共筛选到125个耐盐单株，入选率为2.58%。对52个耐盐株系进行形态和离子的测定，结果表明：盐胁迫导致了根长、根鲜重和根干重的增加；苗长、地上鲜重和地上干重的显著减少，且根部变化幅度小于地上部，说明幼苗地上部分的反应比根部对盐害的反应敏感。在离子效应上，盐胁迫导致了地上部和根部钠离子的显著增加，其中地上部分增加了20倍以上，导入系增加的量小于轮回亲本。钾离子的吸收与钠离子相反，胁迫下减少了对钾离子的吸收，但导入系比轮回亲本表现吸收较多的钾离子。所以导入系耐盐性的增强在离子效应上主要表现为保持较低的叶片钠离子浓度、较低的钠离子转运和增强对钾离子的吸收来降低盐害对水稻生长的抑制和毒害作用。
2.对回交群体进行分蘖至成熟期的盐池(140mmol·L-1NaCl)筛选，筛选到150个耐盐导入系。其中籼、粳稻供体回交后代的在入选率和结实率上分别为6.9%和2.6%，表明籼、粳供体的回交后代在耐盐性上存在差异。耐盐导入系间具有不同的生理形态表现：一部分株系表现为抽穗期提早，属于避盐机制；另一部分株系表现为具有较高的株高、有效穗数和生物产量的株系，属于稀盐机制；还有一些株系表现为千粒重、每穗实粒数、结实率的显著提高，属于忍盐机制。机理间并不是独立的，一些导入系同时具有两种或两种以上的耐盐机制，这些机制间相互协调，彼此促进，提高耐盐性。
3.对回交群体进行开花期高温胁迫，筛选到124个耐高温单株，粳稻比籼稻供体的回交群体出现耐热个体数多，表明粳稻资源中同样存在耐热有利基因。导入系在两种环境下均出现广幅分离，在胁迫和正常条件下分别有80个和47个株系的结实率显著高于轮回亲本，从中鉴定出的耐热性和产量性状均显著好于轮回亲本的优良导入系有8个。对3个F2聚合群体耐高温筛选分别获得7、31和68株极显著好于聚合亲本的耐热个体，平均结实率在80%以上，显著高于聚合亲本和轮回亲本，显示出较理想的耐热聚合效果。
4.对回交群体进行孕穗期的冷水胁迫处理，筛选到324个耐冷单株。经苗期和孕穗期耐冷鉴定，其耐冷性比轮回亲本有显著提高的株系数分别为40个和242个。对116个强耐冷株系进行重复鉴定，在胁迫条件下，结实率和单株产量显著高于轮回亲本的导入系分别有94个和77个；正常条件下，结实率和单株产量显著高于轮回亲本的导入系分别有39个和34个，并且鉴定出18个耐冷性和产量性状都显著好于轮回亲本的优良导入系。而且对目标性状的选择还导致了株高、生物产量、每穗实粒数等相关联性状的显著提高。
5.利用以超优1号为轮回亲本，Chhomromg、Doddi和丰矮占1号为供体筛选获得的回交选择耐冷导入系进行卡方检测的耐冷QTL定位，同时对随机群体进行耐冷评价和方差分析定位耐冷QTL。采用耐冷极端选择导入系定位到29个QTL，方差分析检测到16个耐冷QTL。发现耐冷QTL主要有两种类型：一是影响随机群体冷胁迫与正常条件下结实率差值的QTL，这些等位基因具有提高冷胁迫下性状表达的稳定性，对增强耐冷性有贡献，共检测到8个QTL，分布在第3、5、6、7和8染色体上；第二类是在不同的选择群体中经卡方检测到的相同耐冷QTL，其受假阳性的影响相对较小，共有3个QTL，分别位于第2、3和9染色体上。本研究检测到的耐冷QTL位点，大多与前人发现的位点吻合，说明本研究利用选择群体进行QTL定位的方法有效。试验材料的获得是在高的选择压力下进行的，这些耐冷选择导入系将是生产上的育种群体，本研究将育种群体和基因定位群体相联系，缩短了遗传理论研究与育种实践的距离，对育种技术路线提供了有益的补充。
研究表明，不同供体的回交后代中均出现耐盐、耐冷和耐热的超亲分离，表明这些供体均带有控制这些性状的有利“隐蔽”基因，但各个组合间的抗逆性筛选效率存在较大差异，抗逆性表现不仅取决于供体本身，还取决于受体品种与供体品种的不同组合。本研究筛选出的导入系将成为进行大规模有利基因发掘以及通过有利基因的高效聚合来定向改良抗逆性分子育种的材料平台。
Abiotic stresses such as salinity, cold and heat are major threat to rice production. ElevenBC2F4backcross introgression populations derived from crosses between a japonica varietyChaoyou1as the recurrent parent and11donor parents collected from different countries werescreened for salt tolerance (ST) at the seedling and from tillering to maturity stages, heattolerance (HT) at flowering stage and cold tolerance (CT) at booting stage. The targetintrogression lines (ILs) were evaluated for yield and its related traits under normal and stresscondition to dissect physiological mechanisms of stress tolerance and map QTL for CT. Themain results were described as follows.
1. Eleven BC2F4backcross introgression populations were screened for salt tolerance (ST)at seedling stage. One hundred and twenty-five (125) ILs were selected on the basis of singleplant SDS under salt water (140mmol·L-1NaCl).12,11,21and10ILs selected from four ofthe BC populations including donors Chouyou1/X21, Chouyou1/Q5, Chouyou1/Chhomrongand Chouyou1/Doddi were evaluated for morphological characteristics and ion concentrationsin replicated experiments under stress and normal conditions. The ILs had increased rootlength, root fresh weight, root dry weight but reduced shoot length, shoot fresh weight, shootdry weight under salt stress. The amplitudes of variations in the root traits were smaller thanthose of shoot traits, indicating that shoots were more sensitive to salt stress. With regard toionic effect, salt stress resulted in accumulation of Na+and reduction of K+in the whole plant.Compared to the recurrent parent, the increase of amplitude of Na+and the reduction of Kwere observably lower for the ILs. Coinstantaneous accumulation of20times more Na+in theshoot after salt stress was observed. Most of the salinity tolerant ILs improve their salinitytolerances through lower shoot Na+concentration and lower root to shoot Na+translocation.
2. Eleven BC2F4backcross introgression populations were screened for salt tolerance (ST)at different developmental stages from tillering to maturity. One hundred and fifty (150) ILswere selected on the basis of single plant grain yield (GY) and spikelet fertility (SF) under saltwater (140mmol·L-1NaCl). Our results clearly showed that there was a rich hidden geneticvariation for ST in the donor lines and phenotypic selection under stress condition was apowerful way to identify transgressive segregaantis from the segregating BC populations.Further evaluation of126ILs selected from four BC populations in replicated experimentsunder stress and normal conditions identified10promising ILs with greatly improved ST and higher yield than CY1. Selection under salt stress for yield and SF, supplemented by acombination of desirable secondary traits, can greatly improve the efficiency and accuracy ofbreeding for high salt-tolerance. It is important to point out that three mechanisms includingosmotic adjustment, salt exclusion and tissue tolerance appeared to function together andaffect the same suite of ST related traits in the ILs; although their contributions to specific STtraits were very different for different ILs depending on the specific scenarios of stress.
3. Eleven BC2F4backcross introgression populations were screened for heat tolerance(HT) at flowering stage. Total of124ILs were selected, based on single plant SF under hightemperature (38℃). The results showed that the ILs had better tolerance to heat (HT) than therecurrent parent, Chaoyou1, and the frequencies of plants with higher HT were higher in thepopulations derived from japonica donors were higher than those from indica ones,suggesting that japonica varieties were better sources of desirable genes for improving HT.Progeny testing of the124ILs under the HT stress and normal conditions revealed that64.5%(80) of the lines had better HT than the RP, as measured by SF. The124selected ILspresented wide segregations for other measured traits as well. Eight ILs with greatly improvedHT and higher yield than CY1were selected. HT plants selected from three pyramidingpopulations had an average SF of more than80%, which was significantly higher than the RPand the parents used in pyramiding. One hundred and six (106) plants with significant higherHT than the pyramiding parental lines were selected
4. ILs from the eleven diverse donors were screened for CT at the reproductive stageunder the low temperature (LT) treatment created by irrigating the plants with cold water(19℃) at the reproductive stage. Three hundred and twenty four (324) BC2F5introgressionlines (ILs) were selected based on single plant SF. Progeny testing of the324ILs under thesimilar LT stress revealed that the efficiency of selection for CT was0.794. Further evaluationof116ILs selected from five BC populations in replicated experiments under LT and normalconditions identified18promising ILs with greatly improved CT and yield than CY1.Detailed comparisons between the ILs and CY1were conducted for yield and related traitsunder the LT stress and non-stress conditions to gain useful information and betterunderstanding of important issues related to breeding for CT using introgression breedingsuch as donor selection, selection efficiency, screening method and genetic drag fornon-target traits in the BC breeding process.
5. ILs were assessed to identify cold-tolerance quantitative trait loci (QTL) from thedonors including Chhomromg, Doddi and Fengaizhan1in the genetic background of japonicacultivar Chaoyou1. X2test was used for QTL identification for the CT-selected ILs, meanwhile, random populations were assessed to identify CT-QTL by single marker analysismethod. Total of29QTL associated with CT were detected by the selected ILs and16identified by single marker analysis. The CT-QTL were classified into two types, one are eightQTL affecting trait difference for seed setting rate of stress to control, which distributedchromosomes3,5~8and the alleles at the QTL improved trait stability under stress andactually contributed to CT. The second type is the three common QTL detected by theselected ILs from different populations with less negative chance of QTL detection, whichdistributed chromosomes2,3and9. Furthermore, most of the QTLs identified in this studywere consistent with those documented. Therefore, QTL detection using selected ILs based onwell-known X2test is a simple and effective. previously by other studies.. The use ofintrogression populations in QTL identification and selection for outstanding lines providesan efficient way for integrating gene discovery and practical breeding and as a result speed upthe application modern molecular techniques in breeding for complex traits.
The present study indicated that transgressive segregations were observed for ST, HTand CT in most of the backcross populations, indicating that all the11donors possesfavorable ‘hidden’ alleles for tolerances to the studied stresses. There were significantdifferences among populations in selection efficiency, reflecting stress tolerance depends notonly on donor itself but also on different combinations of the recurrent parent and the donorparent. The stress tolerant ILs will be partially a platform and valuable materials for favorableallele mining and improvement of stress tolerance by pyramiding of favorable genes.
引文
[1]邴雷,赵宝存,沈银柱,黄占景,葛荣朝.2008.植物耐盐性及耐盐相关基因的研究进展.河北师范大学学报(自然科学版).32(2):243-248.
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