大豆对大豆花叶病毒抗性遗传、抗性基因精细定位及表达分析
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摘要
大豆花叶病毒(Soybean mosaic virus, SMV)病是世界大豆产区分布最广的病毒病害之一,在我国各大豆产区均有发生,严重影响大豆的产量与品质。培育和种植抗病品种是防治该病害最经济、安全、有效的途径,而抗病育种的关键是具备优异抗源,并阐明大豆对SMV的抗性遗传规律。进一步对抗性基因标记定位、对分子标记辅助选择以及抗性基因克隆、明确抗性基因作用也有重要意义。
因此,本研究针对新鉴定的部分SMV株系,筛选抗源、研究抗性遗传并精细定位抗性基因,为抗SMV育种提供抗性种质和技术支撑。利用实时荧光定量PCR(Quantitative real-time polymerase chain reaction, QRT-PCR)技术对抗性基因目标区段内的候选基因进行定量表达分析研究,进一步确定抗性品种抗病的关键基因,以阐明其抗病的分子机制,为大豆抗性基因的图位克隆和利用转基因技术提高大豆的抗病性奠定基础。同时采用分子标记辅助选择(Marker assisted selection, MAS)的方法,验证利用抗性相关的分子标记对SMV抗性基因聚合的可行性。主要研究结果如下：
1.抗源筛选
为抗SMV育种筛选优异抗源,本研究对新育成的参加2004-2007年国家及江苏、北京和山东等省市大豆区试的品种在接种我国大豆产区主要流行SMV株系SC3和SC7的条件下分别进行抗性评价,结果表明：在抗SMV鉴定的334个品种中,对SC3抗性较好(高抗和抗病)的品种数有148个,占参试品种数的44.31%；对SC7抗性较好的有71个,占参试品种数的21.26%；同时对两个株系抗性表现较好的有55个,占参试品种数的16.47%。这些抗性较好的品种既可用于大豆生产,也可作为抗源用于抗病品种选育和与抗性相关的研究。研究还显示,来自于西北和黄淮海大豆产区的参试品种一般抗性较好。
2.抗性遗传和抗性基因的等位性分析
利用在我国长江流域和北方大豆产区广分布的SMV株系SC8对部分抗病品种与感病品种的杂交后代进行鉴定,发现F1均表现抗病,F2分离结果经卡方测验符合3抗：1感分离比例,F2：3家系符合1抗：2分离：1感的分离比例,结果表明：科丰1号、PI96983、齐黄1号、大白麻、晋大74、汾豆56、中作229和NY58对SC8株系的抗性受1对显性基因控制。齐黄22x南农1138-2和其反交南农1138-2×齐黄22的F1均表现抗病、F2表现为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,结果表明齐黄22对SC8的抗性是由1对显性基因控制,且抗性基因是由细胞核遗传,无细胞质效应。抗抗组合的等位性测验显示抗病品种晋大74x汾豆56的F1表现抗病,F2和F2：3家系未发现感病株,表明晋大74与汾豆56携带对SC8株系的抗性基因是等位的或紧密连锁的。研究还显示齐黄1号与科丰1号、大白麻与汾豆56携带抗SC8的基因是不等位的,而且独立遗传。
大白麻×南农1138-2和南农1138-2x大白麻的F1、F2和F2：3接种SC4株系的鉴定结果表明,F1植株均表现抗病；经卡方测验,F2群体分离比例符合3抗：1感的比率；F2：3纯合抗病家系、抗感分离家系和纯合感病家系的比率符合1：2：1,表明大白麻对SC4的抗性是受细胞核遗传的1对显性基因控制,无细胞质效应。研究结果还显示：冀LD42、科丰1号、徐豆1号、晋大74、PI96983、齐黄22、跃进4号和汾豆56与感病品种杂交的F1均表现抗病,F2均符合3抗：1感分离比例,F2：3家系符合1抗：2分离：1感的分离比例,证明它们对SC4株系的抗性均各有1对基因控制,且抗病表现为显性。抗抗组合的等位性测验结果表明大白麻与汾豆56、科丰1号、齐黄1号早熟18四个品种携带抗SC4的基因是不等位的,冀LD42与汾豆56是不等位的,晋大74与中作229是不等位的,而且独立遗传。
3.抗性基因的精细定位
以抗病品种科丰1号和感病品种南农1138-2为亲本构建的F2群体(156个F2单株),重组自交家系群体(184个家系)和次级F2群体(181个单株)为材料,利用SNP和基因组SSR标记将科丰1号对SC8株系的抗性基因Rsc8精细定位在大豆的2号染色体上,Rsc8两侧最近的基因组SSR标记(BARCSOYSSR_02_0610和BARCSOYSSR_02_0616)之间的遗传距离为0.32cM,在大豆基因组上的物理距离不足200kb。
利用1047株大白麻×南农1138-2的F2作图群体,采用分离群体分组分析法,将对SC4株系表现抗病的大豆品种大白麻携带的抗性基因Rsc4定位在大豆的14号染色体上,连锁最紧密的基因组SSR标记是BARCSOYSSR_14_1413和BARCSOYSSR_14_1416,与RSC4的遗传距离分别为0.17cM和0.27cM,在大豆基因组上两个标记之间的物理距离不到110kb。
4.抗性候选基因的QRT-PCR (?)析
利用生物信息学的方法和QRT-PCR技术在抗性基因候选区段内搜索候选基因并进行分析：在Rsc8抗性基因区段内,14个预测基因中有11个基因在SMV诱导后的表达量在抗感品种间存在程度差异,进一步分析发现8个抗病候选基因的表达量在抗感品种间的响应模式或时间上有很大的差异,推测这些基因可能参与了科丰1号对SC8株系的抗性反应。在RSC4抗性基因所在的区段,初选的11个候选基因在抗感亲本间的QRT-PCR分析表明,9个基因在接种SC4株系后的表达量在不同时间点抗感品种间都有响应,深入比较发现7个候选基因在抗感品种间不同时间点的表达量有一定的差异,初步认为这些基因可能参与了大白麻对SC4株系的抗性反应。通过上面的分析,推测候选基因Glyma02g13310、13320、13400、13460、13470和Glyma14g38510、38560、38580、39300可能是抗病关键基因。
5.SMV抗性基因的聚合选择
针对SMV株系SC14、SC8和SC4,用齐黄1号(Rsc140)、科丰1号(RSC8)和大白麻(RSC4)做抗性基因的供体,利用获得的与各个抗性基因连锁的9个分子标记,对4个亲本复交(齐黄1号×科丰1号)×(大白麻×南农1138-2)后代进行了标记辅助选择,结合网室接种SC14、SC8和SC4进行表型鉴定,结果证实分子标记选择的携带3个抗性基因的20个F4代单株对三个株系均表现抗病,其中有2个单株携带的3个SMV抗性基因位点已经纯合。研究结果证实紧密连锁的分子标记可以有效地用于对SMV抗性基因的聚合。
Soybean mosaic disease, caused by soybean mosaic virus (SMV), is one of the most broadly distributed viral diseases worldwide in soybean [Glycine max (L.) Merr.]. It causes yield loss and seed quality deficiency seriously. Utilization of resistant varieties is the most economical and environmentally safe approach to controlling this disease. The success and failure of breeding resistant varieties depend on resistant resources, the realization of inheritance mechanisms of resistance and molecular mapping of resistance genes to SMV strains.
In China, SMV has been classified into21strains based on the reactions of SMV isolates on a set of soybean differentials. The objectives of this study were to identify resistant resources, to study the inheritance of resistance to the newly identified strains, to fine mapping the resistance genes, and to study feasibility of pyramiding of resistance genes for SMV using tightly linked molecular markers. Meanwhile, the mRNA expression profiles were analyzed by quantitative real-time polymerase chain reaction (QRT-PCR) in order to insight the resistance mechanism, to determine the soybean functional resistance genes. These will provide basis to cloning of resistance genes and improvement of the resistance of soybean through transgenic technique. The main results were as follows:
1. Identification of resistant resources
In order to provide resistant sources for breeding resistant varieties to SMV strains, the resistance to SMV of334cultivars from the soybean regional test in2004-2007was evaluated after inoculation with two SMV prevalent strains, i. e. SC3and SC7. The results showed that148(44.31%) and71(21.26%) cultivars were resistance to strains SC3and SC7respectively,55(16.47%) cultivars were resistance to both SC3and SC7. These resistance cultivars can not only be used directly in soybean production, but also be used as resistance resources in breeding programs. The study also showed that the cultivars from Northwest China and Huang-Huai-Hai valleys were more resistant to SMV than those from the other regions.
2. Analysis of inheritance and allelism of resistance genes
Crosses of resistant (R) and susceptible (S) parents were made and selfed to construct F1, F2and F2:3populations. They were used to determine the inheritance for resistance to SMV strain SC8. All F1showed resistant, F2generation segregated in3resistant:1susceptible ratio, and1R:2Seg.:1S in F2:3families. The results indicated that Kefeng No.1, PI96983, Qihuang No.1, Dabaima, Jinda74, Fendou56, Zhongzuo229, and NY58carried single resistant gene for SC8strain. Reciprocal crosses were made with Qihuang22and Nannong1138-2to investigate the inheritance of resistance to SC8strain. No cytoplasmic effect was found in the two kinds crosses. It indicated that Qihuang22carried a dominant resistant gene. The test of the allelism of the resistance genes showed that the single dominant gene in Jinda74and Fendou56were alleles or very closely linked; the resistance genes between Qihuang No.1and Kefeng No.1, between Kefeng No.1and Fendou56were not at same locus.
Ten soybean cultivars with resistance to SMV strain SC4, was crossed respectively with a susceptible cultivar (Nannong1138-2or8101) to determine the inheritance of their resistance reaction to SC4strain. Each R parents were also crossed with each other to test the allelism of the resistance genes. The results showed that a dominant gene controlled the resistance to SC4strain in each of Ji LD42, Kefeng No.1,Xudou No.1, Jinda74, P196983, Qihuang22, Yuejin No.4, and Fendou56, respectively. Reciprocal crosses were made with Dabaima and Nannong1138-2to investigate the inheritance of resistance to SC4. The result indicated that Dabaima carried single dominant resistant gene, no cytoplasmic effect was found in the two kind's crosses. The test of the allelism of the resistance genes showed the resistance genes between Dabaima and Fendou56, Kefeng No.1, Qihuang No.1, Zaoshul8; between Ji LD42and Fendou56; between Jinda74and Zhongzuo229were not at same locus.
3. Fine mapping for SMV resistance genes
In order to fine mapping the resistance gene to SMV strain SC8in Kefeng No.1, F2population (156individuals), recombined inbred lines (RIL) population (184families), F2 of secondary population (SP-F2,181individuals) of Kefeng No.1×Nannongl138-2were evaluated following inoculation with SC8and analyzed by simple sequence repeats (SSR), single nucleotide polymorphisms (SNP) and genomic-SSR markers. Results indicated that the resistance gene (designated as RSC8) in Kefeng No.l was located on soybean chromosome2(MLG D1b), closest markers locating two sides of RSC8respectively were BARCSOYSSR_02_0610and BARCSOYSSR_02_0616. The genetic distance of the two markers was0.32cM. Sequence analysis of soybean genome indicted that interval between the two genomic-SSR markers was less than200kb.
By applying bulked segregate analysis (BSA), two genomic-SSR markers (BARCSOYSSR_14_1413and BARCSOYSSR_14_1416) on soybean chromosome14(MLG B2) closely linked to SMV strain SC4resistant gene (designated as Rsc4) were identified in1047plants F2population of Dabaima×Nannongl138-2. The Rsc4was located at a distance of0.17cM from BARCSOYSSR141413and0.27cM from BARCSOYSSR_14_1416(on the other side). Sequence analysis of soybean genome indicted that interval between the two genomic-SSR markers was less than110kb.
4. QRT-PCR analysis of the resistance candidate genes
The analysis of bioinformatics and QRT-PCR of the candidate genes in the RSC8region indicated that the expression level of the eleven genes were different at some time points in the post inoculated with SC8. The eight genes maybe involved in resistance to SMV strain SC8in soybean. Seven candidate genes for Rsc4were identified by QRT-PCR and their expression levels were different between the two parents (Dabaima and Nannong1138-2). Based on the analysis of above, we inferred that candidate genes Glyma02g13310,13320,13400,13460,13470and Glymal4g38510,38560,38580, and39300could be key genes to activating plant defense pathways in response to SMV infection.
5. Pyramiding for SMV resistance genes
The purpose of the research was to pyramid resistance genes RSC14Q, RSC8and RSC4to SMV using marker assisted selection (MAS) and traditional breeding methods. Qihuang No.1, Kefeng No.1and Dabaima were used as the donor of resistance genes RSC14Q, RSC8and RSC4, respectively. A set of F2, F3and F4populations derived from multi-cross and self-cross of (Qihuang No.1×Kefeng No.1)x(DabaimaxNannong1138-2) were developed for selecting individuals carrying three resistant genes. Nine molecular markers linked to the three SMV resistance genes were used for MAS. The results indicated that twenty F4plants carry all three SMV resistance genes were confirmed are resistant to three SMV strains (SC14, SC8and SC4) by inoculation. The results confirmed that the feasibility of pyramiding of resistance genes for SMV in soybean by MAS.
因此,本研究针对新鉴定的部分SMV株系,筛选抗源、研究抗性遗传并精细定位抗性基因,为抗SMV育种提供抗性种质和技术支撑。利用实时荧光定量PCR(Quantitative real-time polymerase chain reaction, QRT-PCR)技术对抗性基因目标区段内的候选基因进行定量表达分析研究,进一步确定抗性品种抗病的关键基因,以阐明其抗病的分子机制,为大豆抗性基因的图位克隆和利用转基因技术提高大豆的抗病性奠定基础。同时采用分子标记辅助选择(Marker assisted selection, MAS)的方法,验证利用抗性相关的分子标记对SMV抗性基因聚合的可行性。主要研究结果如下：
1.抗源筛选
为抗SMV育种筛选优异抗源,本研究对新育成的参加2004-2007年国家及江苏、北京和山东等省市大豆区试的品种在接种我国大豆产区主要流行SMV株系SC3和SC7的条件下分别进行抗性评价,结果表明：在抗SMV鉴定的334个品种中,对SC3抗性较好(高抗和抗病)的品种数有148个,占参试品种数的44.31%；对SC7抗性较好的有71个,占参试品种数的21.26%；同时对两个株系抗性表现较好的有55个,占参试品种数的16.47%。这些抗性较好的品种既可用于大豆生产,也可作为抗源用于抗病品种选育和与抗性相关的研究。研究还显示,来自于西北和黄淮海大豆产区的参试品种一般抗性较好。
2.抗性遗传和抗性基因的等位性分析
利用在我国长江流域和北方大豆产区广分布的SMV株系SC8对部分抗病品种与感病品种的杂交后代进行鉴定,发现F1均表现抗病,F2分离结果经卡方测验符合3抗：1感分离比例,F2：3家系符合1抗：2分离：1感的分离比例,结果表明：科丰1号、PI96983、齐黄1号、大白麻、晋大74、汾豆56、中作229和NY58对SC8株系的抗性受1对显性基因控制。齐黄22x南农1138-2和其反交南农1138-2×齐黄22的F1均表现抗病、F2表现为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,结果表明齐黄22对SC8的抗性是由1对显性基因控制,且抗性基因是由细胞核遗传,无细胞质效应。抗抗组合的等位性测验显示抗病品种晋大74x汾豆56的F1表现抗病,F2和F2：3家系未发现感病株,表明晋大74与汾豆56携带对SC8株系的抗性基因是等位的或紧密连锁的。研究还显示齐黄1号与科丰1号、大白麻与汾豆56携带抗SC8的基因是不等位的,而且独立遗传。
大白麻×南农1138-2和南农1138-2x大白麻的F1、F2和F2：3接种SC4株系的鉴定结果表明,F1植株均表现抗病；经卡方测验,F2群体分离比例符合3抗：1感的比率；F2：3纯合抗病家系、抗感分离家系和纯合感病家系的比率符合1：2：1,表明大白麻对SC4的抗性是受细胞核遗传的1对显性基因控制,无细胞质效应。研究结果还显示：冀LD42、科丰1号、徐豆1号、晋大74、PI96983、齐黄22、跃进4号和汾豆56与感病品种杂交的F1均表现抗病,F2均符合3抗：1感分离比例,F2：3家系符合1抗：2分离：1感的分离比例,证明它们对SC4株系的抗性均各有1对基因控制,且抗病表现为显性。抗抗组合的等位性测验结果表明大白麻与汾豆56、科丰1号、齐黄1号早熟18四个品种携带抗SC4的基因是不等位的,冀LD42与汾豆56是不等位的,晋大74与中作229是不等位的,而且独立遗传。
3.抗性基因的精细定位
以抗病品种科丰1号和感病品种南农1138-2为亲本构建的F2群体(156个F2单株),重组自交家系群体(184个家系)和次级F2群体(181个单株)为材料,利用SNP和基因组SSR标记将科丰1号对SC8株系的抗性基因Rsc8精细定位在大豆的2号染色体上,Rsc8两侧最近的基因组SSR标记(BARCSOYSSR_02_0610和BARCSOYSSR_02_0616)之间的遗传距离为0.32cM,在大豆基因组上的物理距离不足200kb。
利用1047株大白麻×南农1138-2的F2作图群体,采用分离群体分组分析法,将对SC4株系表现抗病的大豆品种大白麻携带的抗性基因Rsc4定位在大豆的14号染色体上,连锁最紧密的基因组SSR标记是BARCSOYSSR_14_1413和BARCSOYSSR_14_1416,与RSC4的遗传距离分别为0.17cM和0.27cM,在大豆基因组上两个标记之间的物理距离不到110kb。
4.抗性候选基因的QRT-PCR (?)析
利用生物信息学的方法和QRT-PCR技术在抗性基因候选区段内搜索候选基因并进行分析：在Rsc8抗性基因区段内,14个预测基因中有11个基因在SMV诱导后的表达量在抗感品种间存在程度差异,进一步分析发现8个抗病候选基因的表达量在抗感品种间的响应模式或时间上有很大的差异,推测这些基因可能参与了科丰1号对SC8株系的抗性反应。在RSC4抗性基因所在的区段,初选的11个候选基因在抗感亲本间的QRT-PCR分析表明,9个基因在接种SC4株系后的表达量在不同时间点抗感品种间都有响应,深入比较发现7个候选基因在抗感品种间不同时间点的表达量有一定的差异,初步认为这些基因可能参与了大白麻对SC4株系的抗性反应。通过上面的分析,推测候选基因Glyma02g13310、13320、13400、13460、13470和Glyma14g38510、38560、38580、39300可能是抗病关键基因。
5.SMV抗性基因的聚合选择
针对SMV株系SC14、SC8和SC4,用齐黄1号(Rsc140)、科丰1号(RSC8)和大白麻(RSC4)做抗性基因的供体,利用获得的与各个抗性基因连锁的9个分子标记,对4个亲本复交(齐黄1号×科丰1号)×(大白麻×南农1138-2)后代进行了标记辅助选择,结合网室接种SC14、SC8和SC4进行表型鉴定,结果证实分子标记选择的携带3个抗性基因的20个F4代单株对三个株系均表现抗病,其中有2个单株携带的3个SMV抗性基因位点已经纯合。研究结果证实紧密连锁的分子标记可以有效地用于对SMV抗性基因的聚合。
Soybean mosaic disease, caused by soybean mosaic virus (SMV), is one of the most broadly distributed viral diseases worldwide in soybean [Glycine max (L.) Merr.]. It causes yield loss and seed quality deficiency seriously. Utilization of resistant varieties is the most economical and environmentally safe approach to controlling this disease. The success and failure of breeding resistant varieties depend on resistant resources, the realization of inheritance mechanisms of resistance and molecular mapping of resistance genes to SMV strains.
In China, SMV has been classified into21strains based on the reactions of SMV isolates on a set of soybean differentials. The objectives of this study were to identify resistant resources, to study the inheritance of resistance to the newly identified strains, to fine mapping the resistance genes, and to study feasibility of pyramiding of resistance genes for SMV using tightly linked molecular markers. Meanwhile, the mRNA expression profiles were analyzed by quantitative real-time polymerase chain reaction (QRT-PCR) in order to insight the resistance mechanism, to determine the soybean functional resistance genes. These will provide basis to cloning of resistance genes and improvement of the resistance of soybean through transgenic technique. The main results were as follows:
1. Identification of resistant resources
In order to provide resistant sources for breeding resistant varieties to SMV strains, the resistance to SMV of334cultivars from the soybean regional test in2004-2007was evaluated after inoculation with two SMV prevalent strains, i. e. SC3and SC7. The results showed that148(44.31%) and71(21.26%) cultivars were resistance to strains SC3and SC7respectively,55(16.47%) cultivars were resistance to both SC3and SC7. These resistance cultivars can not only be used directly in soybean production, but also be used as resistance resources in breeding programs. The study also showed that the cultivars from Northwest China and Huang-Huai-Hai valleys were more resistant to SMV than those from the other regions.
2. Analysis of inheritance and allelism of resistance genes
Crosses of resistant (R) and susceptible (S) parents were made and selfed to construct F1, F2and F2:3populations. They were used to determine the inheritance for resistance to SMV strain SC8. All F1showed resistant, F2generation segregated in3resistant:1susceptible ratio, and1R:2Seg.:1S in F2:3families. The results indicated that Kefeng No.1, PI96983, Qihuang No.1, Dabaima, Jinda74, Fendou56, Zhongzuo229, and NY58carried single resistant gene for SC8strain. Reciprocal crosses were made with Qihuang22and Nannong1138-2to investigate the inheritance of resistance to SC8strain. No cytoplasmic effect was found in the two kinds crosses. It indicated that Qihuang22carried a dominant resistant gene. The test of the allelism of the resistance genes showed that the single dominant gene in Jinda74and Fendou56were alleles or very closely linked; the resistance genes between Qihuang No.1and Kefeng No.1, between Kefeng No.1and Fendou56were not at same locus.
Ten soybean cultivars with resistance to SMV strain SC4, was crossed respectively with a susceptible cultivar (Nannong1138-2or8101) to determine the inheritance of their resistance reaction to SC4strain. Each R parents were also crossed with each other to test the allelism of the resistance genes. The results showed that a dominant gene controlled the resistance to SC4strain in each of Ji LD42, Kefeng No.1,Xudou No.1, Jinda74, P196983, Qihuang22, Yuejin No.4, and Fendou56, respectively. Reciprocal crosses were made with Dabaima and Nannong1138-2to investigate the inheritance of resistance to SC4. The result indicated that Dabaima carried single dominant resistant gene, no cytoplasmic effect was found in the two kind's crosses. The test of the allelism of the resistance genes showed the resistance genes between Dabaima and Fendou56, Kefeng No.1, Qihuang No.1, Zaoshul8; between Ji LD42and Fendou56; between Jinda74and Zhongzuo229were not at same locus.
3. Fine mapping for SMV resistance genes
In order to fine mapping the resistance gene to SMV strain SC8in Kefeng No.1, F2population (156individuals), recombined inbred lines (RIL) population (184families), F2 of secondary population (SP-F2,181individuals) of Kefeng No.1×Nannongl138-2were evaluated following inoculation with SC8and analyzed by simple sequence repeats (SSR), single nucleotide polymorphisms (SNP) and genomic-SSR markers. Results indicated that the resistance gene (designated as RSC8) in Kefeng No.l was located on soybean chromosome2(MLG D1b), closest markers locating two sides of RSC8respectively were BARCSOYSSR_02_0610and BARCSOYSSR_02_0616. The genetic distance of the two markers was0.32cM. Sequence analysis of soybean genome indicted that interval between the two genomic-SSR markers was less than200kb.
By applying bulked segregate analysis (BSA), two genomic-SSR markers (BARCSOYSSR_14_1413and BARCSOYSSR_14_1416) on soybean chromosome14(MLG B2) closely linked to SMV strain SC4resistant gene (designated as Rsc4) were identified in1047plants F2population of Dabaima×Nannongl138-2. The Rsc4was located at a distance of0.17cM from BARCSOYSSR141413and0.27cM from BARCSOYSSR_14_1416(on the other side). Sequence analysis of soybean genome indicted that interval between the two genomic-SSR markers was less than110kb.
4. QRT-PCR analysis of the resistance candidate genes
The analysis of bioinformatics and QRT-PCR of the candidate genes in the RSC8region indicated that the expression level of the eleven genes were different at some time points in the post inoculated with SC8. The eight genes maybe involved in resistance to SMV strain SC8in soybean. Seven candidate genes for Rsc4were identified by QRT-PCR and their expression levels were different between the two parents (Dabaima and Nannong1138-2). Based on the analysis of above, we inferred that candidate genes Glyma02g13310,13320,13400,13460,13470and Glymal4g38510,38560,38580, and39300could be key genes to activating plant defense pathways in response to SMV infection.
5. Pyramiding for SMV resistance genes
The purpose of the research was to pyramid resistance genes RSC14Q, RSC8and RSC4to SMV using marker assisted selection (MAS) and traditional breeding methods. Qihuang No.1, Kefeng No.1and Dabaima were used as the donor of resistance genes RSC14Q, RSC8and RSC4, respectively. A set of F2, F3and F4populations derived from multi-cross and self-cross of (Qihuang No.1×Kefeng No.1)x(DabaimaxNannong1138-2) were developed for selecting individuals carrying three resistant genes. Nine molecular markers linked to the three SMV resistance genes were used for MAS. The results indicated that twenty F4plants carry all three SMV resistance genes were confirmed are resistant to three SMV strains (SC14, SC8and SC4) by inoculation. The results confirmed that the feasibility of pyramiding of resistance genes for SMV in soybean by MAS.
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