大豆对大豆花叶病毒（SMV）的数量抗性研究及QTL定位
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摘要
大豆花叶病毒(Soybean Mosaic Virus,SMV)病是主要的大豆病害之一,严重影响大豆的产量和品质。目前,培育抗病品种仍是控制SMV流行、减少危害的最有效方法。大豆对SMV存在质量抗性与数量抗性两类,开展数量抗性研究对拓宽抗源、培育非专化抗性品种具有重要意义。
本研究利用圣豆一号×汾豆72的P1、P2、Fl、F2和F2：3五个世代进行了主基因+多基因混合遗传分析。结果表明大豆对SMV的数量抗性由一对加性主基因+加性-显性多基因共同控制。遗传参数的估计结果表明,主基因遗传率大于多基因遗传率,因此在数量抗性育种进程中要重视对主基因的选择,同时兼顾对多基因的积累。
利用圣豆一号×汾豆72组合的F2群体,构建了包含24个SSR标记覆盖9个连锁群的遗传图谱。利用构建的遗传连锁图谱,采用复合区间作图法(CIM)和多区间作图法(MIM)对F2和F2:3两个世代的数量抗性QTL进行了定位。结果表明：
(1)在F2群体中,使用MIM以SDI-1138为指标检测到2个QTL,位于C2和F连锁群,共可解释表型变异的21.70%。两个QTL间存在加性×加性(AA)互作效应,效应值为-0.24,贡献率为37.90%；以SDI-SY指标,使用MIM,检测到2个QTL和3个QTL间互作,位于C2和F连锁群。其中qSRM22-C2和qSRM22-F对表型变异的解释比例分别为31.90%和19.30%,其显性×显性(DD)互作效应贡献率较高,为19.8%；以DI为指标,两种方法共检测到4个QTL和2个QTL间互作,位于C2和F连锁群。4个QTL对表型变异的解释比例均在11.00%之上,其中qSRC23-C2最高,为30.81%；以发病率为指标,仅检测到1个QTL,位于F连锁群,贡献率为7.29%；以平均病级为指标,两种方法共检测到3个QTL和3个QTL间互作,位于F连锁群。其中qSRM25-F1和qSRM25-F2的贡献率分别为6.10%和18.30%,其显性×显性(DD)互作的贡献率较高,为17.50%。
(2)在F2：3群体中,以SDI-1138为指标两种方法共检测到6个QTL,分布于B2、C2、Dlb、I和M连锁群。其中CIM检测到4个QTL,但贡献率均较低,最高的仅有4.16%,使用MIM检测到2个QTL其中qSRM31-B2的贡献率最高,为53.5%；以SDI-SY为指标,两种方法共检测到6个QTL及1个QTL间互作,其中CIM检测到4个QTL,但贡献率均较低,平均仅为1.92%；MIM检测到2个QTL,qSRM32-B2和qSRM32-D1b,可解释表型变异的比例分别为19.90%和3.50%,两者的加性×显性(AD)效应贡献率为44.00%；以DI为指标,两种方法共检测到3个QTL和1个QTL间互作,位于D1b和I连锁群。其中qSRM33-Ⅰ的贡献率最高,为33.20%；以发病率为指标,使用MIM方法检测到3个QTL和2个互作QTL,位于B2、C2和M连锁群；共可解释表型变异的43.40%。其中qSRM34-B2和qSRM34-C2对表型变异的解释比例均为14.10%,两者的显性×显性(DD)互作效应较高,效应值为0.67,贡献率为50.50%；以平均病级为指标,两种方法检测到3个QTL和1个QTL间互作,位于D1b和I连锁群。qSRM35-D1、qSRM35-Ⅰ和qSRC35-Ⅰ的贡献率分别为7.10%、33.20%和3.16%,其中前两者的显性×显性(DD)贡献率为6.20%。
(3)比较F2世代和F2:3世代的定位结果,以LOD≥2.0和贡献率R2≥10.00%为主效QTL的认定标准,互作效应≥10.00%的QTL亦认定为主效QTL,在F2中使用CIM和MIM两种方法共检测到了11个主效QTL,分布于C2和F两个连锁群；在F2:3中两种方法共检测到8个主效QTL,分布于B2、C2、Dlb、I和M等5个连锁群；比较使用5个指标检测出的主效QTL数目可知,以DI为指标时检测的主效QTL最多,且其调查方便,因此本研究认为以DI为指标描述数量抗性较为合适。此外,在F2和F2:3两个世代中都检测到一个位于C2连锁群控制大豆对SMV数量抗性的主效QTL。这印证了遗传分析时发现的大豆对SMV的数量抗性由一对加性主基因+加性-显性多基因共同控制的遗传方式。
Soybean mosaic virus (SMV) is one of the major diseases in soybean, which causes severe yield loss and seed quality deficiency. Developing resistant cultivars is still the most effective means to control SMV at present. There are two kinds of resistance to SMV in soybean, qualitative resistance and quantitative resistance. It's of great importance to study the quantitative resistance to widen resistance-donor and cultivate non-specialized cultivars in soybean.
Genetic segregation analysis for disease index(DI) were conducted by using major gene plus polygenes mixed inheritance models and joint analysis method of P1, P2, F1, F2 and F2:3 generations. The results showed that quantitative resistance to SMV was controlled by an additive major gene plus additive-dominant polygenes. Estimates of genetic parameter indicated that heritability of major genes were higher than polygenes, it's necessary to pay more attention to major genes in resistance breeding.
A genetic linkage map based on F2 population of Shengdoul and Fendou72 crossing was constructed and QTLs in F2 and F2:3 were mapped on the genetic map with methods of composite interval method (CIM) and multiple interval method (MIM). The results showed that:(1) In F2, two QTLs were detected on C2 and F for SDI-1138 using MIM, totally explaining 21.70% of phenotypic variation and giving high contribution rate of 37.9% with QTL ineractions (AA). For SDI-SY, two QTLs and three QTL interactions were detected on C2 and F using MIM. Among them qSRM22-C2 & qSRM22-F explained 31.90% and 19.30% of phenotypic variation respectively, with high contribution rate of 19.80% from QTL interaction (DD). Four QTLs and two QTL interactions were dectected on C2 and F for DI using CIM & MIM. The contribution rate of four QTLs was all beyond 11.00% with qSRC23-C2 giving the highest of 30.81%. There was one QTL for incidence locationg on F with constribution rate of 7.290%. For the mean disease scale, three QTLs and three QTL interactions locating on F were detected using CIM and MIM. Among them qSRM25-F1 & qSRM25-F2 explained 6.10% and 18.30% of phenotypic variation respectively, and 17.50% of phenotypic variation for their QTL interaction (DD).
(2) In F2:3, six QTLs were detected on B2、C2、D1b、I and M for SDI-1138 using CIM and MIM. Four QTLs were detected by CIM, giving minor contribution rate with the highest of 4.16%. The qSRM31-B2 detecting by MIM explained 53.50% of phenotypic variation. For SDI-SY, six QTLs and one QTL were detected on B2、C2、D1b. I and M for SDI-SY using CIM and MIM. Four QTLs were detected by CIM, giving minor constribution rate averaging 1.92%. The qSRM32-B2 & qSRM32-D1b detecting by MIM explained 19.90% and 3.50% of phenotypic variation respectively, with QTL interaction (AD) of 44.00%. Three QTLs and one QTL interaction were dectected on D1b and I for DI using CIM & MIM. Among them gave the highest constribution rate of 33.20%. Three QTLs and two QTL interactions were detected on B2, C2 and M for incidence using MIM, totally explaining 43.40% of phenotypic variation. QTL interaction (DD) between qSRM34-B2 and qSRM34-C2 gave high constribution rate of 50.50%. For mean disease scale, three QTLs and one QTL were detected on D1b and I using CIM and MIM. Among them constribution rate of qSRM35-D1, qSRM35-Ⅰand qSRC35-Ⅰwere 7.10%,33.20% and 3.16% repectively. QTL interaction (DD) between qSRM35-D1 & qSRM35-Ⅰexplained 6.20% of phenotypic variation.
(3) LOD value more than 2.0 and contribution rate more than 10.00% for QTLs and QTL interactions were used as the criterion for major QTLs. Mapping results indicated that eleven major QTLs in F2 located on C2 and F, and eight major QTLs on B2、C2、D1b、I and M in F2：3. More major QTLs were detected for DI plus easily using made DI the best trait for quantitative resistance reseach. In addition, a major QTL controlling quantitative resistance to SMV on C2 was detected both in F2 and F2:3. Genetic analysis and QTL mapping were essentially consistent.
本研究利用圣豆一号×汾豆72的P1、P2、Fl、F2和F2：3五个世代进行了主基因+多基因混合遗传分析。结果表明大豆对SMV的数量抗性由一对加性主基因+加性-显性多基因共同控制。遗传参数的估计结果表明,主基因遗传率大于多基因遗传率,因此在数量抗性育种进程中要重视对主基因的选择,同时兼顾对多基因的积累。
利用圣豆一号×汾豆72组合的F2群体,构建了包含24个SSR标记覆盖9个连锁群的遗传图谱。利用构建的遗传连锁图谱,采用复合区间作图法(CIM)和多区间作图法(MIM)对F2和F2:3两个世代的数量抗性QTL进行了定位。结果表明：
(1)在F2群体中,使用MIM以SDI-1138为指标检测到2个QTL,位于C2和F连锁群,共可解释表型变异的21.70%。两个QTL间存在加性×加性(AA)互作效应,效应值为-0.24,贡献率为37.90%；以SDI-SY指标,使用MIM,检测到2个QTL和3个QTL间互作,位于C2和F连锁群。其中qSRM22-C2和qSRM22-F对表型变异的解释比例分别为31.90%和19.30%,其显性×显性(DD)互作效应贡献率较高,为19.8%；以DI为指标,两种方法共检测到4个QTL和2个QTL间互作,位于C2和F连锁群。4个QTL对表型变异的解释比例均在11.00%之上,其中qSRC23-C2最高,为30.81%；以发病率为指标,仅检测到1个QTL,位于F连锁群,贡献率为7.29%；以平均病级为指标,两种方法共检测到3个QTL和3个QTL间互作,位于F连锁群。其中qSRM25-F1和qSRM25-F2的贡献率分别为6.10%和18.30%,其显性×显性(DD)互作的贡献率较高,为17.50%。
(2)在F2：3群体中,以SDI-1138为指标两种方法共检测到6个QTL,分布于B2、C2、Dlb、I和M连锁群。其中CIM检测到4个QTL,但贡献率均较低,最高的仅有4.16%,使用MIM检测到2个QTL其中qSRM31-B2的贡献率最高,为53.5%；以SDI-SY为指标,两种方法共检测到6个QTL及1个QTL间互作,其中CIM检测到4个QTL,但贡献率均较低,平均仅为1.92%；MIM检测到2个QTL,qSRM32-B2和qSRM32-D1b,可解释表型变异的比例分别为19.90%和3.50%,两者的加性×显性(AD)效应贡献率为44.00%；以DI为指标,两种方法共检测到3个QTL和1个QTL间互作,位于D1b和I连锁群。其中qSRM33-Ⅰ的贡献率最高,为33.20%；以发病率为指标,使用MIM方法检测到3个QTL和2个互作QTL,位于B2、C2和M连锁群；共可解释表型变异的43.40%。其中qSRM34-B2和qSRM34-C2对表型变异的解释比例均为14.10%,两者的显性×显性(DD)互作效应较高,效应值为0.67,贡献率为50.50%；以平均病级为指标,两种方法检测到3个QTL和1个QTL间互作,位于D1b和I连锁群。qSRM35-D1、qSRM35-Ⅰ和qSRC35-Ⅰ的贡献率分别为7.10%、33.20%和3.16%,其中前两者的显性×显性(DD)贡献率为6.20%。
(3)比较F2世代和F2:3世代的定位结果,以LOD≥2.0和贡献率R2≥10.00%为主效QTL的认定标准,互作效应≥10.00%的QTL亦认定为主效QTL,在F2中使用CIM和MIM两种方法共检测到了11个主效QTL,分布于C2和F两个连锁群；在F2:3中两种方法共检测到8个主效QTL,分布于B2、C2、Dlb、I和M等5个连锁群；比较使用5个指标检测出的主效QTL数目可知,以DI为指标时检测的主效QTL最多,且其调查方便,因此本研究认为以DI为指标描述数量抗性较为合适。此外,在F2和F2:3两个世代中都检测到一个位于C2连锁群控制大豆对SMV数量抗性的主效QTL。这印证了遗传分析时发现的大豆对SMV的数量抗性由一对加性主基因+加性-显性多基因共同控制的遗传方式。
Soybean mosaic virus (SMV) is one of the major diseases in soybean, which causes severe yield loss and seed quality deficiency. Developing resistant cultivars is still the most effective means to control SMV at present. There are two kinds of resistance to SMV in soybean, qualitative resistance and quantitative resistance. It's of great importance to study the quantitative resistance to widen resistance-donor and cultivate non-specialized cultivars in soybean.
Genetic segregation analysis for disease index(DI) were conducted by using major gene plus polygenes mixed inheritance models and joint analysis method of P1, P2, F1, F2 and F2:3 generations. The results showed that quantitative resistance to SMV was controlled by an additive major gene plus additive-dominant polygenes. Estimates of genetic parameter indicated that heritability of major genes were higher than polygenes, it's necessary to pay more attention to major genes in resistance breeding.
A genetic linkage map based on F2 population of Shengdoul and Fendou72 crossing was constructed and QTLs in F2 and F2:3 were mapped on the genetic map with methods of composite interval method (CIM) and multiple interval method (MIM). The results showed that:(1) In F2, two QTLs were detected on C2 and F for SDI-1138 using MIM, totally explaining 21.70% of phenotypic variation and giving high contribution rate of 37.9% with QTL ineractions (AA). For SDI-SY, two QTLs and three QTL interactions were detected on C2 and F using MIM. Among them qSRM22-C2 & qSRM22-F explained 31.90% and 19.30% of phenotypic variation respectively, with high contribution rate of 19.80% from QTL interaction (DD). Four QTLs and two QTL interactions were dectected on C2 and F for DI using CIM & MIM. The contribution rate of four QTLs was all beyond 11.00% with qSRC23-C2 giving the highest of 30.81%. There was one QTL for incidence locationg on F with constribution rate of 7.290%. For the mean disease scale, three QTLs and three QTL interactions locating on F were detected using CIM and MIM. Among them qSRM25-F1 & qSRM25-F2 explained 6.10% and 18.30% of phenotypic variation respectively, and 17.50% of phenotypic variation for their QTL interaction (DD).
(2) In F2:3, six QTLs were detected on B2、C2、D1b、I and M for SDI-1138 using CIM and MIM. Four QTLs were detected by CIM, giving minor contribution rate with the highest of 4.16%. The qSRM31-B2 detecting by MIM explained 53.50% of phenotypic variation. For SDI-SY, six QTLs and one QTL were detected on B2、C2、D1b. I and M for SDI-SY using CIM and MIM. Four QTLs were detected by CIM, giving minor constribution rate averaging 1.92%. The qSRM32-B2 & qSRM32-D1b detecting by MIM explained 19.90% and 3.50% of phenotypic variation respectively, with QTL interaction (AD) of 44.00%. Three QTLs and one QTL interaction were dectected on D1b and I for DI using CIM & MIM. Among them gave the highest constribution rate of 33.20%. Three QTLs and two QTL interactions were detected on B2, C2 and M for incidence using MIM, totally explaining 43.40% of phenotypic variation. QTL interaction (DD) between qSRM34-B2 and qSRM34-C2 gave high constribution rate of 50.50%. For mean disease scale, three QTLs and one QTL were detected on D1b and I using CIM and MIM. Among them constribution rate of qSRM35-D1, qSRM35-Ⅰand qSRC35-Ⅰwere 7.10%,33.20% and 3.16% repectively. QTL interaction (DD) between qSRM35-D1 & qSRM35-Ⅰexplained 6.20% of phenotypic variation.
(3) LOD value more than 2.0 and contribution rate more than 10.00% for QTLs and QTL interactions were used as the criterion for major QTLs. Mapping results indicated that eleven major QTLs in F2 located on C2 and F, and eight major QTLs on B2、C2、D1b、I and M in F2：3. More major QTLs were detected for DI plus easily using made DI the best trait for quantitative resistance reseach. In addition, a major QTL controlling quantitative resistance to SMV on C2 was detected both in F2 and F2:3. Genetic analysis and QTL mapping were essentially consistent.
引文
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