大豆对大豆花叶病毒抗病基因的遗传分析、精细定位和标记辅助选择研究
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摘要
大豆花叶病毒(Soybean mosaic virus,SMV)病是大豆生产上的一种世界性病害,严重影响大豆产量和品质。培育抗病品种是防治大豆花叶病毒病,保证大豆优质、高产、稳产最经济、有效和安全的途径。国内外学者在抗源筛选、抗性遗传和抗性基因的分子标记定位等方面进行了广泛的研究。本研究在南京农业大学国家大豆改良中心完成对全国SMV株系划分命名及对各地区目前SMV株系的组成、分布和流行情况分析的基础上,有针对性地选择各大豆产区的主要流行株系(SC3、SC7、SC18、SC15),对最新育成品种及少量地方和引进品种进行抗性评价和抗源筛选；利用多个抗SMV材料配制杂交组合,分析大豆对SMV株系SC3的抗性遗传方式及抗性基因间等位关系；对大豆花叶病毒抗病基因RSC12进行初步定位,并对已初步定位的基因RSC12和RSC14Q进行标记辅助选择的可行性研究；在对RSC14Q初步定位基础上,构建次级F2群体,利用以基因组序列为基础开发的新的分子标记,对RSC14Q进行精细定位。这些研究为抗SMV育种提供了抗性种质和抗源信息,为明确抗性遗传方式,开展抗性基因的标记辅助选择提供理论和方法指导,为SMV抗性基因RSC14Q的图位克隆奠定基础。主要研究结果如下：
1.大豆对SMV株系SC3、SC7、SC15、SC18的抗性评价
除对华南热带区域的13个大豆品种接种当地主要流行广分布SMV株系SC18和SC15外,其余的238个大豆品种均接种SC3和SC7株系。共筛选到26份抗性较好的材料。科丰1号、齐黄1号、PI96983、PI486355、Kwanggyo这5个品种对SC3和SC7均表现抗侵染,齐黄22、大白麻、早熟18、徐豆1号、Davis、Buflla这6个品种只对SC3表现抗侵染,Q0807、南农307、Q0806、中作056082等15个品种接种SC3和SC7后发病症状较轻,病情指数均在20%以内,对SMV有较好的抗扩展能力。
251份品种无论接种优势流行弱毒株系(SC3和SC18)还是优势流行强毒株系(SC7和SC15),品种抗性分布都有如下规律：抗性表现为中间类型(中抗和中感)的品种所占比例很高,均在60%以上,表现高抗病和高感的品种比例很低,低于5%。并且抗弱毒株系的品种所占比例高于抗强毒株系的品种比例。
国内黄淮海大豆生态区的山东、北京、山西3省(市)的抗性资源丰富,品种平均病情轻,品种间抗性变异度大。
2.大豆对SMV株系SC3的抗性遗传与抗病基因的等位性分析
七个抗性材料分别与感病材料南农1138-2杂交获得杂交后代。对各组合亲本及后代接种SMV株系SC3鉴定其抗感表型,并对调查结果做卡方适合性测验。结果显示：齐黄1号、科丰1号、Davis、广吉、早熟18、徐豆1号、PI96983分别与感病品种南农1138-2杂交的F1表现抗病,F2呈3抗：1感的分离比例,F2：3家系符合1抗：2分离：1感的分离比例,初步表明它们各有一对基因控制对SMV株系SC3的抗性,且抗性基因为显性。
针对这七个抗性材料配制抗抗杂交组合,对各组合亲本及后代接种SMV株系SC3鉴定其抗感表型,并对调查结果做卡方适合性测验。结果显示：齐黄1号×广吉、齐黄1号×早熟18、Davis×广吉、Davis×早熟18、徐豆1号×广吉、PI96983×广吉六组抗抗杂交组合的F1,F2均未发现感病植株,初步表明齐黄1号、广吉、早熟18、Davis、徐豆1号、PI96983所携带的对SC3的抗性基因是等位的或紧密连锁的。科丰1号×广吉和科丰1号×早熟18两组抗抗杂交组合F1都表现为抗病,F2经卡方测验呈15抗：1感的分离比例,初步表明科丰1号所携带的对SC3的抗性基因与广吉、早熟18分别携带的对SC3的抗性基因在不同的位点上,且两个基因独立遗传。
3.大豆对SMV株系SC12的抗性基因的定位及对RSC12、 RSC14Q的标记辅助选择研究
杂交组合齐黄22(R)×南农1138-2(S)的亲本及后代接种SMV株系SC12,F1表现抗病,F2群体抗感分离比例符合3：1,F2：3家系呈1抗：2分离：1感的分离比例,表明齐黄22对SMV株系SC12的抗性是由一对显性基因控制,以Rsc12表示。
利用齐黄22×南农1138-2的包含219个单株的F2作图群体,采用常规摩擦接种法对群体各单株进行表型鉴定,依据分离群体分组分析法,应用Mapmaker3.0软件进行表型与基因型的连锁分析,对抗性基因Rsc12进行标记定位,将Rscl2定位在F连锁群上,7个SSR标记均与抗病基因RSC12连锁,标记与RSCl2顺序和距离为：Sat_2976.4cM Sat_2344.9cM Sat_1541.1cM Satt1140.7cM SOYHSP1761.6cM Satt3342.4cM RSC126.3cM Sct033。
利用抗病基因RSC12、Rsc14Q两侧的SSR分子标记Satt334和Sct033对齐黄22×南农1138-2的F2、F3和F4世代以及齐黄1号×南农1138-2的F5、F6世代进行抗病基因的标记辅助选择效率研究。结果表明：单独使用标记Satt334或Sct_033对Rscl2在齐黄22×南农138-2的三个世代以及对Rscl4Q在齐黄1号×南农1138的两个世代MAS准确率均在85%以上,同时使用两个标记准确率达95%。证明2个标记可以有效地代替人工接种鉴定用于大豆抗SMV育种中对抗病基因RSC12、RSC14Q的选择。
4.大豆对SMV抗病基因RSC14Q的精细定位研究
利用RSC14Q的初步定位结果,在重组自交系中筛选目标区间杂合、遗传背景纯合的单株,即剩余杂合体,共筛选到的四株剩余杂合体。构建了一个由剩余杂合体自交获得的包含680个单株的次级分离群体。经接种鉴定群体符合3抗：1感的分离比例,可以作为Rscl4Q基因的定位研究的定位群体。
根据大豆基因组序列信息,针对目标区段开发新的标记,成功开发出了2个InDel标记和1个SNP标记,所获得的InDel标记分别命名为MY750, MY26530,所获得的SNP标记命名为MY525。新标记的获得有助于提高目的基因区域遗传图谱的精度,精细定位RSC14Q。
本研究将RSC14Q定位在Satt334和MY750之间,分别与之相距0.6cM和0.5cM,根据已公布的大豆基因组序列,将包含RSC14Q的区段限定在1.18Mb区间内。根据单株及其后代的鉴定表型及标记数据,将RSC14Q锁定在标记MY525和MY750之间,依据大豆基因组序列信息,进一步将RSC14Q限定在616kb区间内,为图位克隆Rscl4Q基因奠定了基础。
Soybean mosaic virus (SMV) disease is the major virus disease in soybean (Glycine max (L.) Merr.) worldwide, resulting in substantial yield losses and significant seed-quality deterioration. Breeding resistance varieties is the most economical, effective and environmentally sound approach to control this disease and guarantee a high and stable yield and good quality in soybean production. At present, the identification and distribution of strain groups of SMV in China has been accomplished by Nanjing Agriculture University. In this study, the major prevalent strains (SC3、SC7、SC18、SC15) in soybean production regions domestic were selected to evaluate the resistance of the new cultivars to SMV. Diallel crosses among susceptible cultivar and several resistant cultivars were made to investigate the inheritance of SMV resistance to SC3and to determine the allelic relationship of resistance genes in different resistant cultivars. The resistance gene to SC12was primary mapped and the feasibility of marker-assisted selection (MAS) for RSC12was evaluated by related markers. RSC14Q to SC14was fine mapped via an advanced F2population and the new developed molecular markers. This research supplied some useful information for soybean breeding for disease resistance and provided a foundation for MAS and map-based cloning of resistant genes. The main results were as follows:
1. Evaluation of the soybean cultivars resistance to SMV strains SC3, SC7, SC15and SC18
13cultivars from the south China tropicalt ecoregion were all inoculated with SMV strain SC18and SC15. The other238cultivars were all inoculated with SC3and SC7.26cultivars were seleceted for their better resistance to SMV. Kefeng No.1, Qihuang No.1, PI96983, PI486355and Kwanggyo were the ones resistant in infection to SMV strains SC3and SC7. Qihuang22, Dabaima, Zaoshul8, Xudou No.1, Davis and Buflla were only resistant in infection to SMV strain SC3.15cultivars including Q0807, Nannong307, Q0806, Zhongzuo056082and so on all showed light symptoms after inoculation of SC3and SC7and their disease index were all within20%, which indicated that they had better resistance in development to SMV strains.
The resistance distribution of251cultivars evaluated by weak strains (SC3and SC18) or virulent strains (SC7and SC15) showed that the middle type (middle resistant and middle susceptible) cultivars with a high ratio were more than60%while cultivars in the two ends grades were less than5%. And the cultivars resistant to weak strains were more than to virulent.
Soybean resource resistant to SMV is abundant in the provinces of Jing, Lu, Jin in Huang-Huai-Hai ecoregion, where soybean average diseases index was low and variance degree of the resistance among the soybeans is high.
2. Studies on inheritance and allelism of resistance genes to SMV strain SC3in soybeans
Seven cultivars (Qihuang No.1, Kefeng No.1, Davis, Kwanggyo, Zaoshu18, Xudou No.1and P196983) resistant(R) to SMV strain SC3were crossed respectively with the susceptible(S) cultivar Nannong1138-2to determine the inheritance of their resistance reaction to SC3. As a result, F1from each cross were all resistant to SC3. F2from each cross were all fitted a ratio of3(R):1(S), and their separate F2:3lines segregated with a fitness to1(R):2(segregating):1(S). The results showed that a dominant gene controlled the resistance to SC3in each of Qihuang No.1, Kefeng No.1, Davis, Kwanggyo, Zaoshu18, Xudou No.1and PI96983.
The R parents were also crossed with each other to evaluate the allelic relationships between the genes in them. There were no susceptible plant in F1and F2from the correspond crosses of Qihuang No.1×Kwanggyo, Qihuang No.1×Zaoshu18, Davis×Kwanggyo, Davis×Zaoshu18, Xudou No.1×Kwanggyo, PI96983×Kwanggyo, which indicated that the single dominant gene with resistance to SC3in Qihuang No.1, Kwanggyo, Zaoshul8, Davis, Xudou No.1and PI96983were allelic at a common locus or very closely linked. F1from the two crosses of Kefeng No.1×Kwanggyo and Kefeng No.1×Zaoshu18were all resistant while their correspond F2population segregated in a ratio of15(R):1(S), which indicated that the single dominant gene resistant to SC3in Kefeng No.1and the gene resistant to SC3in each of Kwanggyo and Zaoshul8were not at the same locus and the two different genes inherited independently in their separate hybrid progenies.
3. Molecular mapping RSC12and marker assisted selection of resistance genes RSC12and RSC14Q to SMV in soybean
The P1, P2, F1plants, F2population and F2:3lines from the cross of Qihuang22and Nannong1138-2were all inoculated with the SMV strain SC12for identification of their resistance in the greenhouse. Qihuang22and F1individuals were resistant, and Nannong1138-2were susceptible. F2plants exhibited a good fit to3R:1S, and F2:3lines segregated with an acceptable fitness to1R:2segregating:1S. These results obviously indicated that a single dominant gene, designated as RSC12, controlled resistance to SC12in Qihuang22.
A F2population of Qihuang22(R) XNannong1138-2(S) with219individuals was constructed for molecular mapping of resistance gene RSC12to SMV in soybean. Linkage analysis between inoculation phenotype and genetic markers by using the software MAPMAKER/EXP3.0b demonstrated that the resistance gene Rsc12was located on the linkage group F and linked with seven SSR markers. The order and genetic distance linked RSC12were Sat_2976.4cM Sat_2344.9cM Sat_1541.1cM Satt1140.7cM SOYHSP1761.6cM Satt3342.4cM RSC126.3cM Sct_033.
The MAS efficiency of SSR markers Satt334and Sct_033for Rsc12and RSC14Q was evaluated in F2, F3and F4populations from Qihuang22XNannong1138-2and F5, F6populations from Qihuang No.1XNannong1138-2. The results indicated that the MAS efficiency of Satt334and Sct_033for RSC12or RsSC14Q was more than85%, and that the MAS efficiency reached as high as95%when these two markers were co-used. Therefore, the two SSR markers can be used effectively in selecting for resistance genes RSC12and RSC14Q to SMV instead of inoculation identification
4. Fine mapping of resistance gene RSC14Q to SMV
Based on the primary mapping of RSC14Q on the linkage group F (Li et al,2006),4residual heterozygous lines (RHLs) which have a heterozygous aim segment and a homozygous genetic background were screened from a recombinant inbred line (RIL) of soybean. The RILs were obtained from a cross between Qihuang No.1(R) and Nannong1138-2(S). The advanced F2population with680individuals constructed by selfing of4RHLs (F8) could be used for fine mapping of RSC147Q since its segregation ratio fitted to3R:1S after phenotype identification.
Based on the physical position of RSC14Q region, two InDel markers (MY750and MY26530) and one SNP marker (MY525) were newly developed by re-sequencing PCR products using primers designed from the soybean genome sequence. The new markers obtained to saturate the genetic map of target gene were helpful to fine map RSC14Q·
By linkage analysis between inoculation phenotype and genetic markers, RSC14Q was located at interval between Sart334and MY750, with genetic distance of0.6cM and0.5cM, respectively, which corresponded to a physical distance on the Williams82draft assembly (Glyma1.13) of1.18Mb. And then according to the genetic analysis and recombination information of the two individuals and their separate F2:3lines, RSC14Q was concluded to be located at a narrower interval between MY525and MY750. Thus the genomic region containing RSC14Q was further confined to616kb interval based on the soybean genome information. These results provided a foundation for MAS and map-based cloning of RSC14Q·
1.大豆对SMV株系SC3、SC7、SC15、SC18的抗性评价
除对华南热带区域的13个大豆品种接种当地主要流行广分布SMV株系SC18和SC15外,其余的238个大豆品种均接种SC3和SC7株系。共筛选到26份抗性较好的材料。科丰1号、齐黄1号、PI96983、PI486355、Kwanggyo这5个品种对SC3和SC7均表现抗侵染,齐黄22、大白麻、早熟18、徐豆1号、Davis、Buflla这6个品种只对SC3表现抗侵染,Q0807、南农307、Q0806、中作056082等15个品种接种SC3和SC7后发病症状较轻,病情指数均在20%以内,对SMV有较好的抗扩展能力。
251份品种无论接种优势流行弱毒株系(SC3和SC18)还是优势流行强毒株系(SC7和SC15),品种抗性分布都有如下规律：抗性表现为中间类型(中抗和中感)的品种所占比例很高,均在60%以上,表现高抗病和高感的品种比例很低,低于5%。并且抗弱毒株系的品种所占比例高于抗强毒株系的品种比例。
国内黄淮海大豆生态区的山东、北京、山西3省(市)的抗性资源丰富,品种平均病情轻,品种间抗性变异度大。
2.大豆对SMV株系SC3的抗性遗传与抗病基因的等位性分析
七个抗性材料分别与感病材料南农1138-2杂交获得杂交后代。对各组合亲本及后代接种SMV株系SC3鉴定其抗感表型,并对调查结果做卡方适合性测验。结果显示：齐黄1号、科丰1号、Davis、广吉、早熟18、徐豆1号、PI96983分别与感病品种南农1138-2杂交的F1表现抗病,F2呈3抗：1感的分离比例,F2：3家系符合1抗：2分离：1感的分离比例,初步表明它们各有一对基因控制对SMV株系SC3的抗性,且抗性基因为显性。
针对这七个抗性材料配制抗抗杂交组合,对各组合亲本及后代接种SMV株系SC3鉴定其抗感表型,并对调查结果做卡方适合性测验。结果显示：齐黄1号×广吉、齐黄1号×早熟18、Davis×广吉、Davis×早熟18、徐豆1号×广吉、PI96983×广吉六组抗抗杂交组合的F1,F2均未发现感病植株,初步表明齐黄1号、广吉、早熟18、Davis、徐豆1号、PI96983所携带的对SC3的抗性基因是等位的或紧密连锁的。科丰1号×广吉和科丰1号×早熟18两组抗抗杂交组合F1都表现为抗病,F2经卡方测验呈15抗：1感的分离比例,初步表明科丰1号所携带的对SC3的抗性基因与广吉、早熟18分别携带的对SC3的抗性基因在不同的位点上,且两个基因独立遗传。
3.大豆对SMV株系SC12的抗性基因的定位及对RSC12、 RSC14Q的标记辅助选择研究
杂交组合齐黄22(R)×南农1138-2(S)的亲本及后代接种SMV株系SC12,F1表现抗病,F2群体抗感分离比例符合3：1,F2：3家系呈1抗：2分离：1感的分离比例,表明齐黄22对SMV株系SC12的抗性是由一对显性基因控制,以Rsc12表示。
利用齐黄22×南农1138-2的包含219个单株的F2作图群体,采用常规摩擦接种法对群体各单株进行表型鉴定,依据分离群体分组分析法,应用Mapmaker3.0软件进行表型与基因型的连锁分析,对抗性基因Rsc12进行标记定位,将Rscl2定位在F连锁群上,7个SSR标记均与抗病基因RSC12连锁,标记与RSCl2顺序和距离为：Sat_2976.4cM Sat_2344.9cM Sat_1541.1cM Satt1140.7cM SOYHSP1761.6cM Satt3342.4cM RSC126.3cM Sct033。
利用抗病基因RSC12、Rsc14Q两侧的SSR分子标记Satt334和Sct033对齐黄22×南农1138-2的F2、F3和F4世代以及齐黄1号×南农1138-2的F5、F6世代进行抗病基因的标记辅助选择效率研究。结果表明：单独使用标记Satt334或Sct_033对Rscl2在齐黄22×南农138-2的三个世代以及对Rscl4Q在齐黄1号×南农1138的两个世代MAS准确率均在85%以上,同时使用两个标记准确率达95%。证明2个标记可以有效地代替人工接种鉴定用于大豆抗SMV育种中对抗病基因RSC12、RSC14Q的选择。
4.大豆对SMV抗病基因RSC14Q的精细定位研究
利用RSC14Q的初步定位结果,在重组自交系中筛选目标区间杂合、遗传背景纯合的单株,即剩余杂合体,共筛选到的四株剩余杂合体。构建了一个由剩余杂合体自交获得的包含680个单株的次级分离群体。经接种鉴定群体符合3抗：1感的分离比例,可以作为Rscl4Q基因的定位研究的定位群体。
根据大豆基因组序列信息,针对目标区段开发新的标记,成功开发出了2个InDel标记和1个SNP标记,所获得的InDel标记分别命名为MY750, MY26530,所获得的SNP标记命名为MY525。新标记的获得有助于提高目的基因区域遗传图谱的精度,精细定位RSC14Q。
本研究将RSC14Q定位在Satt334和MY750之间,分别与之相距0.6cM和0.5cM,根据已公布的大豆基因组序列,将包含RSC14Q的区段限定在1.18Mb区间内。根据单株及其后代的鉴定表型及标记数据,将RSC14Q锁定在标记MY525和MY750之间,依据大豆基因组序列信息,进一步将RSC14Q限定在616kb区间内,为图位克隆Rscl4Q基因奠定了基础。
Soybean mosaic virus (SMV) disease is the major virus disease in soybean (Glycine max (L.) Merr.) worldwide, resulting in substantial yield losses and significant seed-quality deterioration. Breeding resistance varieties is the most economical, effective and environmentally sound approach to control this disease and guarantee a high and stable yield and good quality in soybean production. At present, the identification and distribution of strain groups of SMV in China has been accomplished by Nanjing Agriculture University. In this study, the major prevalent strains (SC3、SC7、SC18、SC15) in soybean production regions domestic were selected to evaluate the resistance of the new cultivars to SMV. Diallel crosses among susceptible cultivar and several resistant cultivars were made to investigate the inheritance of SMV resistance to SC3and to determine the allelic relationship of resistance genes in different resistant cultivars. The resistance gene to SC12was primary mapped and the feasibility of marker-assisted selection (MAS) for RSC12was evaluated by related markers. RSC14Q to SC14was fine mapped via an advanced F2population and the new developed molecular markers. This research supplied some useful information for soybean breeding for disease resistance and provided a foundation for MAS and map-based cloning of resistant genes. The main results were as follows:
1. Evaluation of the soybean cultivars resistance to SMV strains SC3, SC7, SC15and SC18
13cultivars from the south China tropicalt ecoregion were all inoculated with SMV strain SC18and SC15. The other238cultivars were all inoculated with SC3and SC7.26cultivars were seleceted for their better resistance to SMV. Kefeng No.1, Qihuang No.1, PI96983, PI486355and Kwanggyo were the ones resistant in infection to SMV strains SC3and SC7. Qihuang22, Dabaima, Zaoshul8, Xudou No.1, Davis and Buflla were only resistant in infection to SMV strain SC3.15cultivars including Q0807, Nannong307, Q0806, Zhongzuo056082and so on all showed light symptoms after inoculation of SC3and SC7and their disease index were all within20%, which indicated that they had better resistance in development to SMV strains.
The resistance distribution of251cultivars evaluated by weak strains (SC3and SC18) or virulent strains (SC7and SC15) showed that the middle type (middle resistant and middle susceptible) cultivars with a high ratio were more than60%while cultivars in the two ends grades were less than5%. And the cultivars resistant to weak strains were more than to virulent.
Soybean resource resistant to SMV is abundant in the provinces of Jing, Lu, Jin in Huang-Huai-Hai ecoregion, where soybean average diseases index was low and variance degree of the resistance among the soybeans is high.
2. Studies on inheritance and allelism of resistance genes to SMV strain SC3in soybeans
Seven cultivars (Qihuang No.1, Kefeng No.1, Davis, Kwanggyo, Zaoshu18, Xudou No.1and P196983) resistant(R) to SMV strain SC3were crossed respectively with the susceptible(S) cultivar Nannong1138-2to determine the inheritance of their resistance reaction to SC3. As a result, F1from each cross were all resistant to SC3. F2from each cross were all fitted a ratio of3(R):1(S), and their separate F2:3lines segregated with a fitness to1(R):2(segregating):1(S). The results showed that a dominant gene controlled the resistance to SC3in each of Qihuang No.1, Kefeng No.1, Davis, Kwanggyo, Zaoshu18, Xudou No.1and PI96983.
The R parents were also crossed with each other to evaluate the allelic relationships between the genes in them. There were no susceptible plant in F1and F2from the correspond crosses of Qihuang No.1×Kwanggyo, Qihuang No.1×Zaoshu18, Davis×Kwanggyo, Davis×Zaoshu18, Xudou No.1×Kwanggyo, PI96983×Kwanggyo, which indicated that the single dominant gene with resistance to SC3in Qihuang No.1, Kwanggyo, Zaoshul8, Davis, Xudou No.1and PI96983were allelic at a common locus or very closely linked. F1from the two crosses of Kefeng No.1×Kwanggyo and Kefeng No.1×Zaoshu18were all resistant while their correspond F2population segregated in a ratio of15(R):1(S), which indicated that the single dominant gene resistant to SC3in Kefeng No.1and the gene resistant to SC3in each of Kwanggyo and Zaoshul8were not at the same locus and the two different genes inherited independently in their separate hybrid progenies.
3. Molecular mapping RSC12and marker assisted selection of resistance genes RSC12and RSC14Q to SMV in soybean
The P1, P2, F1plants, F2population and F2:3lines from the cross of Qihuang22and Nannong1138-2were all inoculated with the SMV strain SC12for identification of their resistance in the greenhouse. Qihuang22and F1individuals were resistant, and Nannong1138-2were susceptible. F2plants exhibited a good fit to3R:1S, and F2:3lines segregated with an acceptable fitness to1R:2segregating:1S. These results obviously indicated that a single dominant gene, designated as RSC12, controlled resistance to SC12in Qihuang22.
A F2population of Qihuang22(R) XNannong1138-2(S) with219individuals was constructed for molecular mapping of resistance gene RSC12to SMV in soybean. Linkage analysis between inoculation phenotype and genetic markers by using the software MAPMAKER/EXP3.0b demonstrated that the resistance gene Rsc12was located on the linkage group F and linked with seven SSR markers. The order and genetic distance linked RSC12were Sat_2976.4cM Sat_2344.9cM Sat_1541.1cM Satt1140.7cM SOYHSP1761.6cM Satt3342.4cM RSC126.3cM Sct_033.
The MAS efficiency of SSR markers Satt334and Sct_033for Rsc12and RSC14Q was evaluated in F2, F3and F4populations from Qihuang22XNannong1138-2and F5, F6populations from Qihuang No.1XNannong1138-2. The results indicated that the MAS efficiency of Satt334and Sct_033for RSC12or RsSC14Q was more than85%, and that the MAS efficiency reached as high as95%when these two markers were co-used. Therefore, the two SSR markers can be used effectively in selecting for resistance genes RSC12and RSC14Q to SMV instead of inoculation identification
4. Fine mapping of resistance gene RSC14Q to SMV
Based on the primary mapping of RSC14Q on the linkage group F (Li et al,2006),4residual heterozygous lines (RHLs) which have a heterozygous aim segment and a homozygous genetic background were screened from a recombinant inbred line (RIL) of soybean. The RILs were obtained from a cross between Qihuang No.1(R) and Nannong1138-2(S). The advanced F2population with680individuals constructed by selfing of4RHLs (F8) could be used for fine mapping of RSC147Q since its segregation ratio fitted to3R:1S after phenotype identification.
Based on the physical position of RSC14Q region, two InDel markers (MY750and MY26530) and one SNP marker (MY525) were newly developed by re-sequencing PCR products using primers designed from the soybean genome sequence. The new markers obtained to saturate the genetic map of target gene were helpful to fine map RSC14Q·
By linkage analysis between inoculation phenotype and genetic markers, RSC14Q was located at interval between Sart334and MY750, with genetic distance of0.6cM and0.5cM, respectively, which corresponded to a physical distance on the Williams82draft assembly (Glyma1.13) of1.18Mb. And then according to the genetic analysis and recombination information of the two individuals and their separate F2:3lines, RSC14Q was concluded to be located at a narrower interval between MY525and MY750. Thus the genomic region containing RSC14Q was further confined to616kb interval based on the soybean genome information. These results provided a foundation for MAS and map-based cloning of RSC14Q·
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