利用野生番茄Solanum pennellii LA716渐渗系群体进行番茄耐盐相关QTLs定位
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摘要
番茄(Solanum Lycopersicum)原产中南美洲,属茄科番茄属,广泛栽培于世界各地,可鲜食,也可加工成不同类型的番茄制品。盐渍化是造成土壤退化、作物减产的重要因素。培育耐盐的品种是提高盐碱地上番茄产量的有效方法。本文利用对盐敏感的栽培种番茄M82与耐盐的野生潘那利番茄S. pennellii LA716构建的渐渗系群体(introgression lines, ILs),分别对番茄芽期以及苗期的耐盐QTLs进行定位,分析了QTLs的遗传效应和互作效应；并对苗期耐盐主效QTLStq7b进行了精细定位；结合田间试验进行耐盐性鉴定筛选。研究结果对于探讨番茄耐盐机理和利用分子标记辅助选育耐盐番茄品种具有重要的理论指导意义。主要研究结果如下：
1.番茄芽期耐盐QTLs的定位及其效应分析
通过芽期耐盐性鉴定,定位了分布在1、2、4、5、6、8、12共7条染色体上的10个QTLs (Stq1、Stq2a、 Stq2b、Stq2c、Stq4、 Stq5、Stq6、Stq8a、Stq8b和Stq12)影响番茄芽期的耐盐性,这些QTLs均来自S. pennellii LA716。在盐胁迫条件下,这些包含QTLs的渐渗系较M82的发芽指数减少21.52～36.22%。
这10个QTLs与来自类番茄S. lycopersicoides LA2951的3个芽期耐盐QTLs进行同位比较,只有第2条染色体上的Stq2c与来自S. lycopersicoides的St2b同位,其它QTLs均不同位。
QTLs遗传效应分析表明,有4个QTLs (Stq1、 Stq2c、Stq5和Stq8b)表现明显的显性效应,其余6个QTLs (Stq2a、 Stq2b、Stq4、Stq6、Stq8a和Stq12)均表现隐性效应。QTLs互作分析表明呈现典型的小于加性的效应。
2.番茄苗期耐盐QTLs的定位及其效应分析
通过苗期耐盐性鉴定,定位了分布在2、6、7、8、10共5条染色体上的7个QTLs (Stq2a、Stq2b、Stq6a、Stq7a、Stq7b、Stq8和Stq10)影响番茄的耐盐性,这些QTLs均来自S. pennellii LA716,在盐胁迫条件下,这些包含QTLs的渐渗系可较M82的成活率提高18.9～83.8%,而包含在IL6-4与IL6-3的1个QTL (Stq6b)有待于进一步确定。
这7个QTLs与来自类番茄S.lycopersicoides LA2951的7个苗期耐盐QTLs进行同位比较,在第2条染色体上Stq2a和St2同位,在第6条染色体上Stq6a和St6同位,其它QTLs均不同位。
QTLs遗传效应分析表明,除Stq6b外,其余QTLs均表现明显的显性效应。QTLs互作分析表明呈现典型的小于加性的效应,但来自第7条染色体上的2个QTLs互作却表现消减效应。
3.番茄苗期耐盐主效QTLStq7b的精细定位
根据已定位的耐盐主效QTL所在的亚渐渗系片段IL7-5-5与M82的耐盐性差异,利用CAPS、SSR和AFLP等分子标记进行亲本间的多态性标记筛选与创制,共得到6个分子标记(3个CAPS、1个SSR和2个AFLP)与苗期耐盐主效QTL紧密连锁、可用于精细定位。
利用M82与含苗期耐盐QTLStq7b的渐渗系IL7-5为亲本,建立1338株F2群体,进行苗期耐盐性鉴定；以IL7-5-5上的特异标记SSR285筛选1338株F2群体建立了一个255株的重组群体,再利用已获得的IL7-5-5上的另5个特异性标记,对255个重组单株进行基因型分析,建立IL7-5-5片段上更饱和的标记连锁图谱,这6个标记覆盖了整个IL7-5-5上0.9cM的染色体片段,标记间平均遗传距离0.15cM。并将苗期耐盐QTLStq7b定位在CAPS标记U231219和TG418之间,分别相距0.36和0.16cM,且与C2_At4g30580共分离,其贡献率为7.34%,加性效应5.45,LOD值2.24。
根据Stq7b精细定位的结果,利用生物信息学的方法,以已测序的S. lycopersicum H1706全基因组序列为参考,对该区域的候选基因进行预测分析。候选基因(Solyc07g005440)和(Solyc07g005520)的分别具有典型的NAF和BTB结构域,初步推断前者在应答非生物胁迫时起着正面效应,后者在应答植物的盐胁迫,维持细胞膨压,保护细胞免受非生物胁迫中起着关键的作用。
4.耐盐渐渗系及品系的田间筛选
筛选出耐盐性良好,产量、单果重等性状良好的渐渗系材料1个(IL7-5)；耐盐性良好,产量、单果重等性状较差的渐渗系材料3个(IL6-2、IL6-4和IL7-1)；耐盐性较好,产量、单果重等性状良好的渐渗系材料2个(IL2-1-1与IL2-6-5)。从26个杂交组合中筛选出4个组合,其耐盐性好,移栽成活率和产量均高于对照。
Tomato (Solanum lycopersicum) originated from middle-south America, it was grown around the world, and can be used as fresh and processing products marketing. Soil salinization is a key factor of land degradation, leading to falling crop yields. Breeding for salt-tolerance (ST) cultivars have been recognized as an effective way to improve tomato productivity in salt-affected soil. In this paper, the introgression line population (ILs), which was developed from a salt-sensitive parent S. lycopersicum M82and a ST wild species S. pennellii LA716, was employed to identify QTLs conferring ST during seed germination and vegetative stage respectively. Meanwhile the genetic effect and interaction of some QTLs were also analyzed. The fine mapping of one major ST QTL during vegetative stage and the trial of the ST germplasm and hybrids were conducted. The main results obtained in this thesis were concluded as follows:
1. Identification and Effect Analysis of QTLs Conferring ST during Seed Germination Through salt tolerance identification in seed germination, ten unambiguous QTLs (Stq1, Stq2a, Stq2b, Stq2c, Stq4, Stq5, Stq6, Stq8a, Stq8b and Stq12) located on Chrom.l,2,4,5,6,8and12were identified, which were responsible for controlling ST during seed germination and derived from wild relative S. pennellii LA716. The germination index decreased about21.52～36.22%as compared to the control M82under salt stress.
Comparing these10QTLs with other3QTLs derived from S. lycopersicoides LA2951ILs, the results showed that only QTL Stq2c and QTL St2b (from S. lycopersicoides) sitting at the same position on Chrom.2.
Genetic analysis of4QTLs (Stql, Stq2c, Stq5and Stq8b) showed a significant dominant effect, and6QTLs (Stq2a, Stq2b, Stq4, Stq6, Stq8a. and Stq12) showed a recessive effect. The interaction of QTLs presented a typically less-than-additive effect.
2. Identification and Effect Analysis of QTLs Conferring ST during Vegetative Stage
Through salt tolerance identification in vegetative stage, seven unambiguous QTLs (Stq2a, Stq2b, Stq6a, Stq1a, Stq7b, Stq8and Stq10) located respectively on Chrom.2,6,7,8and10were identified, which were responsible for controlling ST during vegetative stage and derived from wild relative S. pennellii LA716. The survival percentage increased by18.9-83.8%as compared to the control M82under salt stress. One QTL possibly involved in IL6-4and IL6-3needed to be confirmed in the further experiments.
Comparing these7QTLs with other7QTLs derived from S. lycopersicoides LA2951ILs, QTL Stq2a and QTL St2, as well as QTL Stq6a and QTL St6, sat at the same position on Chrom.2and Chrom.6respectively.
Genetic analysis of some QTLs showed a significant dominant effect with the exception of Stq6b. The interaction of QTLs presented a typically less-than-additive but two QTLs located on Chrom.7gave a reduced effect.
3. Fine Mapping of Major ST QTLStq7b during Vegetative Stage
Screening with PCR polymorphism molecular markers based on the ST differences between M82and IL7-5-5,6molecular markers (3CAPS,1SSR and2AFLP markers) were confirmed, which were closely linked with major ST QTLStq7b and can be used for fine mapping.
1338F2individuals were derived from a cross of M82×IL7-5. The recombinant population of255individuals was selected via screening the1338F2individuals by SSR285, then screening the recombinant population of255individuals by others5markers. Finally, the fine mapping of QTLStq7b on IL7-5-5segment was constructed. The length of the introgressed segment was measured at0.9cM which is approximately the same as the length of IL7-5-5, and with an average genetic distance of0.15cM. ST QTLStq1b was identified in the fine mapping, its genetic distance to the maker U231219and TG418was0.36and0.16cM respectively, and co-segregating with the maker C2_At4g30580. The contribution of individual QTL to phenotypic variation was7.34%, the additive effect was5.45, and the LOD was2.24.
Prediction of candidate genes involved in the genome sequence of Stq7b was carried out by bioinformatics based on the fine mapping and the published complete genome sequence of S. lycopersicum H1706. The candidate gene (Solyc07g005440) and (Solyc07g005520) contained a typical NAF domain and BTB domain in terminal region respectively, the former may have a positive effect on responds to abiotic signals, and the latter may have a positive effect on responds to salt stress, which play an important role in protecting cell against abiotic stress by maintaining cell turgor.
4. Trial of the ST ILs and Hybrids in Salt-affected Soil
Comprehensive evaluation the salt-tolerance of ILs, six ILs were selected, IL7-5showed good ST, high yield and fruit weight. IL6-2, IL6-4and IL7-1presented good ST but lower yield and fruit weight. IL2-2-2and IL2-6-5presented not best ST but high yield and fruit weight. Comparing to the M82and others3controls,4combinations performed good ST with high survival percentage and high yield were selected from26hybrids.
1.番茄芽期耐盐QTLs的定位及其效应分析
通过芽期耐盐性鉴定,定位了分布在1、2、4、5、6、8、12共7条染色体上的10个QTLs (Stq1、Stq2a、 Stq2b、Stq2c、Stq4、 Stq5、Stq6、Stq8a、Stq8b和Stq12)影响番茄芽期的耐盐性,这些QTLs均来自S. pennellii LA716。在盐胁迫条件下,这些包含QTLs的渐渗系较M82的发芽指数减少21.52～36.22%。
这10个QTLs与来自类番茄S. lycopersicoides LA2951的3个芽期耐盐QTLs进行同位比较,只有第2条染色体上的Stq2c与来自S. lycopersicoides的St2b同位,其它QTLs均不同位。
QTLs遗传效应分析表明,有4个QTLs (Stq1、 Stq2c、Stq5和Stq8b)表现明显的显性效应,其余6个QTLs (Stq2a、 Stq2b、Stq4、Stq6、Stq8a和Stq12)均表现隐性效应。QTLs互作分析表明呈现典型的小于加性的效应。
2.番茄苗期耐盐QTLs的定位及其效应分析
通过苗期耐盐性鉴定,定位了分布在2、6、7、8、10共5条染色体上的7个QTLs (Stq2a、Stq2b、Stq6a、Stq7a、Stq7b、Stq8和Stq10)影响番茄的耐盐性,这些QTLs均来自S. pennellii LA716,在盐胁迫条件下,这些包含QTLs的渐渗系可较M82的成活率提高18.9～83.8%,而包含在IL6-4与IL6-3的1个QTL (Stq6b)有待于进一步确定。
这7个QTLs与来自类番茄S.lycopersicoides LA2951的7个苗期耐盐QTLs进行同位比较,在第2条染色体上Stq2a和St2同位,在第6条染色体上Stq6a和St6同位,其它QTLs均不同位。
QTLs遗传效应分析表明,除Stq6b外,其余QTLs均表现明显的显性效应。QTLs互作分析表明呈现典型的小于加性的效应,但来自第7条染色体上的2个QTLs互作却表现消减效应。
3.番茄苗期耐盐主效QTLStq7b的精细定位
根据已定位的耐盐主效QTL所在的亚渐渗系片段IL7-5-5与M82的耐盐性差异,利用CAPS、SSR和AFLP等分子标记进行亲本间的多态性标记筛选与创制,共得到6个分子标记(3个CAPS、1个SSR和2个AFLP)与苗期耐盐主效QTL紧密连锁、可用于精细定位。
利用M82与含苗期耐盐QTLStq7b的渐渗系IL7-5为亲本,建立1338株F2群体,进行苗期耐盐性鉴定；以IL7-5-5上的特异标记SSR285筛选1338株F2群体建立了一个255株的重组群体,再利用已获得的IL7-5-5上的另5个特异性标记,对255个重组单株进行基因型分析,建立IL7-5-5片段上更饱和的标记连锁图谱,这6个标记覆盖了整个IL7-5-5上0.9cM的染色体片段,标记间平均遗传距离0.15cM。并将苗期耐盐QTLStq7b定位在CAPS标记U231219和TG418之间,分别相距0.36和0.16cM,且与C2_At4g30580共分离,其贡献率为7.34%,加性效应5.45,LOD值2.24。
根据Stq7b精细定位的结果,利用生物信息学的方法,以已测序的S. lycopersicum H1706全基因组序列为参考,对该区域的候选基因进行预测分析。候选基因(Solyc07g005440)和(Solyc07g005520)的分别具有典型的NAF和BTB结构域,初步推断前者在应答非生物胁迫时起着正面效应,后者在应答植物的盐胁迫,维持细胞膨压,保护细胞免受非生物胁迫中起着关键的作用。
4.耐盐渐渗系及品系的田间筛选
筛选出耐盐性良好,产量、单果重等性状良好的渐渗系材料1个(IL7-5)；耐盐性良好,产量、单果重等性状较差的渐渗系材料3个(IL6-2、IL6-4和IL7-1)；耐盐性较好,产量、单果重等性状良好的渐渗系材料2个(IL2-1-1与IL2-6-5)。从26个杂交组合中筛选出4个组合,其耐盐性好,移栽成活率和产量均高于对照。
Tomato (Solanum lycopersicum) originated from middle-south America, it was grown around the world, and can be used as fresh and processing products marketing. Soil salinization is a key factor of land degradation, leading to falling crop yields. Breeding for salt-tolerance (ST) cultivars have been recognized as an effective way to improve tomato productivity in salt-affected soil. In this paper, the introgression line population (ILs), which was developed from a salt-sensitive parent S. lycopersicum M82and a ST wild species S. pennellii LA716, was employed to identify QTLs conferring ST during seed germination and vegetative stage respectively. Meanwhile the genetic effect and interaction of some QTLs were also analyzed. The fine mapping of one major ST QTL during vegetative stage and the trial of the ST germplasm and hybrids were conducted. The main results obtained in this thesis were concluded as follows:
1. Identification and Effect Analysis of QTLs Conferring ST during Seed Germination Through salt tolerance identification in seed germination, ten unambiguous QTLs (Stq1, Stq2a, Stq2b, Stq2c, Stq4, Stq5, Stq6, Stq8a, Stq8b and Stq12) located on Chrom.l,2,4,5,6,8and12were identified, which were responsible for controlling ST during seed germination and derived from wild relative S. pennellii LA716. The germination index decreased about21.52～36.22%as compared to the control M82under salt stress.
Comparing these10QTLs with other3QTLs derived from S. lycopersicoides LA2951ILs, the results showed that only QTL Stq2c and QTL St2b (from S. lycopersicoides) sitting at the same position on Chrom.2.
Genetic analysis of4QTLs (Stql, Stq2c, Stq5and Stq8b) showed a significant dominant effect, and6QTLs (Stq2a, Stq2b, Stq4, Stq6, Stq8a. and Stq12) showed a recessive effect. The interaction of QTLs presented a typically less-than-additive effect.
2. Identification and Effect Analysis of QTLs Conferring ST during Vegetative Stage
Through salt tolerance identification in vegetative stage, seven unambiguous QTLs (Stq2a, Stq2b, Stq6a, Stq1a, Stq7b, Stq8and Stq10) located respectively on Chrom.2,6,7,8and10were identified, which were responsible for controlling ST during vegetative stage and derived from wild relative S. pennellii LA716. The survival percentage increased by18.9-83.8%as compared to the control M82under salt stress. One QTL possibly involved in IL6-4and IL6-3needed to be confirmed in the further experiments.
Comparing these7QTLs with other7QTLs derived from S. lycopersicoides LA2951ILs, QTL Stq2a and QTL St2, as well as QTL Stq6a and QTL St6, sat at the same position on Chrom.2and Chrom.6respectively.
Genetic analysis of some QTLs showed a significant dominant effect with the exception of Stq6b. The interaction of QTLs presented a typically less-than-additive but two QTLs located on Chrom.7gave a reduced effect.
3. Fine Mapping of Major ST QTLStq7b during Vegetative Stage
Screening with PCR polymorphism molecular markers based on the ST differences between M82and IL7-5-5,6molecular markers (3CAPS,1SSR and2AFLP markers) were confirmed, which were closely linked with major ST QTLStq7b and can be used for fine mapping.
1338F2individuals were derived from a cross of M82×IL7-5. The recombinant population of255individuals was selected via screening the1338F2individuals by SSR285, then screening the recombinant population of255individuals by others5markers. Finally, the fine mapping of QTLStq7b on IL7-5-5segment was constructed. The length of the introgressed segment was measured at0.9cM which is approximately the same as the length of IL7-5-5, and with an average genetic distance of0.15cM. ST QTLStq1b was identified in the fine mapping, its genetic distance to the maker U231219and TG418was0.36and0.16cM respectively, and co-segregating with the maker C2_At4g30580. The contribution of individual QTL to phenotypic variation was7.34%, the additive effect was5.45, and the LOD was2.24.
Prediction of candidate genes involved in the genome sequence of Stq7b was carried out by bioinformatics based on the fine mapping and the published complete genome sequence of S. lycopersicum H1706. The candidate gene (Solyc07g005440) and (Solyc07g005520) contained a typical NAF domain and BTB domain in terminal region respectively, the former may have a positive effect on responds to abiotic signals, and the latter may have a positive effect on responds to salt stress, which play an important role in protecting cell against abiotic stress by maintaining cell turgor.
4. Trial of the ST ILs and Hybrids in Salt-affected Soil
Comprehensive evaluation the salt-tolerance of ILs, six ILs were selected, IL7-5showed good ST, high yield and fruit weight. IL6-2, IL6-4and IL7-1presented good ST but lower yield and fruit weight. IL2-2-2and IL2-6-5presented not best ST but high yield and fruit weight. Comparing to the M82and others3controls,4combinations performed good ST with high survival percentage and high yield were selected from26hybrids.
引文
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