大豆生育期结构性状的遗传分析及相关基因的分子标记
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摘要
大豆的生育期性状不仅包括全生育期的长短,而且包括各发育阶段的组成,即生育期结构。本文分别以全生育期(M)一致、营养生长期(V)不同的材料作为亲本,构建分离群体,研究大豆生育期结构性状的遗传规律,定位相关基因,分析主效基因在不同环境条件下的效应;以V一致、M不同的材料作为亲本,构建分离群体,研究大豆生殖生长期的遗传规律,定位开花后不同光周期条件下大豆主要性状及其光周期反应敏感性相关基因,以期为大豆生育期结构性状的改良提供理论。
主要内容如下:
1.全生育期相同、营养生长期不同大豆品种生育期结构性状的遗传分析。以小黑豆和GR8836作为亲本,构建包括亲本、F_1、F_2、F_(2:3)的多世代联合分析群体,研究生育期结构性状的遗传规律。结果表明,V、M及生殖生长期(R)与营养生长期的比值(R /V)均是由1对主基因和多基因共同控制的数量性状,相应主效基因的遗传贡献率分别为77.59%、16.77%和54.68%。对于R/V比值而言,主基因表现负向加、显性效应,且效应值相近;对于V、M而言,主基因效应以正向加性为主。鉴于V和R/V的主基因遗传率较高,在育种中,可在早代进行有效选择。
2.大豆生育期结构性状相关基因的QTL定位。应用小黑豆×GR8836组合的F_2、F_(2:3)群体定位生育期结构性状及其光温反应敏感性相关基因,检测到1个位于C2连锁群上Satt557附近的QTL,其在春、夏播条件下同时与R/V比值、V及M相关基因连锁,表现出一定的环境稳定性。但该位点可以解释同一性状的表型变异不同,加、显性效应大小也有差别,加之本研究定位到的7个微效QTL在不同播季条件下均未被重复检测到,说明QTL效应受制于环境条件。另外,在C2连锁群上Satt286附近区域定位到1个与R/V比值光温反应敏感性相关的QTL,该QTL可解释21.16%的表型变异。利用与生育期结构性状主效QTL紧密连锁的标记Satt557和Satt365分析大豆生育期近等基因系(NIL),发现该QTL与大豆生育期主基因E1/e1紧密连锁。
3.主效QTL在不同类型大豆品种中的分布。利用Satt557和Satt365分析本研究定位的生育期结构主效QTL在生育期长短和结构均不同的中、美大豆成熟期组标准品种中的分布,发现该主效QTL在美国品种中的分布频率显著高于中国大豆品种。同时,美国品种携带的主效QTL连锁基因隐性位点的材料数占携带该主效基因材料总数的比率显著高于中国品种,且隐性位点主要分布在相对早熟的品种中,推测该基因的隐性位点在美国品种中的利用率较高。
4.生育期结构性状主基因E1/e1的效应及与环境条件的关系。利用生育期性状不同的大豆近等基因系(NIL)材料,分析E1/e1在不同光周期、不同播季条件下的效应。结果表明,E1/e1显性位点延长V、M,缩短R,减小R/V比值。E1与其它生育期基因间存在互作;对营养生长期而言,背景基因型中其它生育期基因显性位点有增强E1基因效应的作用,且显性位点越多,增效作用越大;E1对R/V比值的效应与背景基因型无明显相关性。E1与环境间存在互作,在多数情况下,E1在长日照条件下的效应大于短日条件,在春播条件下的效应大于夏播。
5.营养生长期相同、全生育期不同大豆品种R和R/V比值的遗传分析。以鲁豆9号和浙江四月白作为亲本,构建分离群体,研究R及R/V比值的遗传规律。结果表明,在营养生长期相同的条件下,生殖生长期长度是由1对加性主基因和加性-显性多基因控制的数量性状,F2代的主基因遗传贡献率为44.11%;R/V比值是由两对主基因和多基因共同控制的数量性状,主基因总的遗传贡献率为82.37%,且两对主基因的加性效应作用方向的相同,说明在该组合中,可早代选择R/V性状。
6.生殖生长期处于不同光周期条件下大豆农艺性状和品质性状相关基因的QTL分析。以保褐选3×叙永红豆组合衍生的重组近交系(RIL)群体为材料,于生殖生长期进行不同光周期处理,观察农艺性状和品质性状的变化,定位大豆农艺性状、品质性状及其光周期反应敏感性的相关基因。结果表明,大豆不同性状对生殖生长期光周期反应的敏感性不同。顺序为:生殖生长期长度>株高>主茎节数>蛋白质含量、脂肪含量>百粒重>单株荚数>蛋白质和脂肪总量。利用复合区间作图法定位到21个与上述8个性状及其光周期反应敏感性相关的QTL,每个QTL可解释的表型变异范围在5.83%～27.85%之间。针对同一性状,未检测到在长、短日条件下均起作用的主效QTL,说明光周期不仅影响大豆的表型,而且调控相关基因的表达。作者提出,光周期对大豆的生长发育和产量形成有决定性影响,改良大豆品种的适应能力应从钝化其光周期反应敏感性入手。本研究的结果可为高产、稳产、广适应大豆品种的选育提供新的思路。
Growth period traits of soybean include not only the whole growth period (M), but also the growth period structure, i.e., the relative length of various developmental phases. In this study, the inheritance and QTL mapping of the growth period structure traits of soybean were analyzed based on segregation population derived from parents having similar M and different V, and the inheritance of the length of reproductive period (R) and the sensitivity of agronomic and quality traits to post-flowering photopeirod were also analyzed using the population derived from the parents with similar V and different M. The results were summarized as follows:
1. The inheritance of growth period structure traits in the cross with parents having different V and M.
Joint segregation analysis based on P_1, P_2, F_1, F_2, F_(2:3) generation populations derived from Xiaoheidou and GR8836 indicated that one major gene modified by poly-genes controlled the traits of V, M and R/V ratio. The major gene heritability was 77.59%, 16.77% and 54.68%, respectively. The major effect gene had positive addictive effect for V and M, and negative addictive effect for R/V. Since the major gene of R/V and V had higher heritability, then it will be effective to select the genotypes with desirable growth period structive traits.
2. QTL mapping for growth period structure traits in soybean.
F_2, F_(2:3) populations derived from Xiaoheidou and GR8836 were grown in spring and summer to map the relative genes of growth period structure traits and photothermal response sensitivity. One major effect QTL closed to Satt557 on C2 linkage group was detected, which associated with R/V ratio, V and M under different planting seasons. It was shown that the major effect QTL was stable in different environments. Meanwhile, the phenotypic variance explained by QTL, addictive and dominant effects were different for each trait, and seven minor effects QTL for growth period triats were could not be detected repeatedly under different planting seasons. Those indicated that genes of growth period traits were influenced by environmental factors to a certain extent. One major effect QTL for photothermal response sensitivity of R/V was mapped, which associated with the marker of Satt286 lied on C2 linkage group. To find the relationship of major effect QTL and E-series matutity genes controlling flowering and maturity time, Satt557 and Satt365 were used to assay the banding pattern of 41 NILs. The results indicated that the major effect QTL mapped in this study was tightly linked to E1/e1.
3. Distribution of major QTL in different maturity groups soybean varieties.
The banding patterns of 38 American varieties and 64 Chinese varieties belonging to different maturity groups were assessed using the SSR primers of Satt557 and Satt365 tightly linked to the major effect gene of growth period structure traits. The results suggested that the distribution frequency of major effect gene in American varieties was higher than that in Chinese varieties. Meanwhile, the percentage of varieties carring recessive allele to all varieties carring major QTL was higher in American maturity group varieties than that in Chinese, and the recessive allele mainly distributed in the early maturity groups, indicating that the recessive allele was more frequently used in American varieties than in Chinese varieties.
4. Evaluation of the effects of major maturity gene E1/e1 and its dependence on photothermal environment.
Effects of E1/e1 on growth period structure traits under different photoperiods and planting seasons were investigated by using maturity near-isolines of soybean. Dominant allele of E1 delayed V and M, hastened R and decreased R/V in most genotypes studied. Gene effect was dependent on the genetic background as well. There were interactions between E1 and other matrutiy genes since the dominant allele could strengthen the effects of other maturity genes generally. The interaction between E1 and environment was also detected, E1 effect under long day (LD) was higher than under short day (SD), and that in spring sowing was higher than in summer sowing.
5. The inheritance of R and R/V in the cross with parents having different (M) and similar (V).
Joint segregation analysis based on populations derived from Ludou 9 and Zhejiangsiyuebai showed that R was controlled by one major gene plus poly-genes. Heritability of the major effect gene was 44.11% in F2 generation; R/V was controlled by two major genes modified by poly-genes. Total heritability of two major genes was up to 82.37%. The addictive effects of two major genes were all positive. It was proposed that R/V ratio could be selected in early generations.
6. QTL mapping and effect analysis of yield and quality traits in soybean under different post -flowering photoperiods.
181 recombinant inbred lines (RIL) derived from Baohexuan 3×Xuyonghongdou were constructed to map QTL associated with the agronomic and quality traits under both LD (16 h) and SD (12 h) and their photoperiod response sensitivity. The order of the photopeiod sensitivity of the following eight traits was as below: reproductive period length (R)>plant height>node number on the main stem>protein content, oil content>100-seed weight>pod number per plant>total percentage of protein and oil. By using MCIM method, a total of 21 QTL were identified and mapped on 11 linkage groups including A1, A2, B1, B2, C1, D1a, etc. QTL detected could explain 5.83%～21.44% of observed phenotypic variation. For the given agronomic or quality traits, no major QTL were explored both under SD and LD. It could be drawn that photoperiod sensitivity is the key trait for adaptability improvement of soybean. To breed elite varieties with ideal yield potential, quality and stress tolerance, the photoperiod-insensitive genes related to various traits should be explored and pyramided with other needed genes.
主要内容如下:
1.全生育期相同、营养生长期不同大豆品种生育期结构性状的遗传分析。以小黑豆和GR8836作为亲本,构建包括亲本、F_1、F_2、F_(2:3)的多世代联合分析群体,研究生育期结构性状的遗传规律。结果表明,V、M及生殖生长期(R)与营养生长期的比值(R /V)均是由1对主基因和多基因共同控制的数量性状,相应主效基因的遗传贡献率分别为77.59%、16.77%和54.68%。对于R/V比值而言,主基因表现负向加、显性效应,且效应值相近;对于V、M而言,主基因效应以正向加性为主。鉴于V和R/V的主基因遗传率较高,在育种中,可在早代进行有效选择。
2.大豆生育期结构性状相关基因的QTL定位。应用小黑豆×GR8836组合的F_2、F_(2:3)群体定位生育期结构性状及其光温反应敏感性相关基因,检测到1个位于C2连锁群上Satt557附近的QTL,其在春、夏播条件下同时与R/V比值、V及M相关基因连锁,表现出一定的环境稳定性。但该位点可以解释同一性状的表型变异不同,加、显性效应大小也有差别,加之本研究定位到的7个微效QTL在不同播季条件下均未被重复检测到,说明QTL效应受制于环境条件。另外,在C2连锁群上Satt286附近区域定位到1个与R/V比值光温反应敏感性相关的QTL,该QTL可解释21.16%的表型变异。利用与生育期结构性状主效QTL紧密连锁的标记Satt557和Satt365分析大豆生育期近等基因系(NIL),发现该QTL与大豆生育期主基因E1/e1紧密连锁。
3.主效QTL在不同类型大豆品种中的分布。利用Satt557和Satt365分析本研究定位的生育期结构主效QTL在生育期长短和结构均不同的中、美大豆成熟期组标准品种中的分布,发现该主效QTL在美国品种中的分布频率显著高于中国大豆品种。同时,美国品种携带的主效QTL连锁基因隐性位点的材料数占携带该主效基因材料总数的比率显著高于中国品种,且隐性位点主要分布在相对早熟的品种中,推测该基因的隐性位点在美国品种中的利用率较高。
4.生育期结构性状主基因E1/e1的效应及与环境条件的关系。利用生育期性状不同的大豆近等基因系(NIL)材料,分析E1/e1在不同光周期、不同播季条件下的效应。结果表明,E1/e1显性位点延长V、M,缩短R,减小R/V比值。E1与其它生育期基因间存在互作;对营养生长期而言,背景基因型中其它生育期基因显性位点有增强E1基因效应的作用,且显性位点越多,增效作用越大;E1对R/V比值的效应与背景基因型无明显相关性。E1与环境间存在互作,在多数情况下,E1在长日照条件下的效应大于短日条件,在春播条件下的效应大于夏播。
5.营养生长期相同、全生育期不同大豆品种R和R/V比值的遗传分析。以鲁豆9号和浙江四月白作为亲本,构建分离群体,研究R及R/V比值的遗传规律。结果表明,在营养生长期相同的条件下,生殖生长期长度是由1对加性主基因和加性-显性多基因控制的数量性状,F2代的主基因遗传贡献率为44.11%;R/V比值是由两对主基因和多基因共同控制的数量性状,主基因总的遗传贡献率为82.37%,且两对主基因的加性效应作用方向的相同,说明在该组合中,可早代选择R/V性状。
6.生殖生长期处于不同光周期条件下大豆农艺性状和品质性状相关基因的QTL分析。以保褐选3×叙永红豆组合衍生的重组近交系(RIL)群体为材料,于生殖生长期进行不同光周期处理,观察农艺性状和品质性状的变化,定位大豆农艺性状、品质性状及其光周期反应敏感性的相关基因。结果表明,大豆不同性状对生殖生长期光周期反应的敏感性不同。顺序为:生殖生长期长度>株高>主茎节数>蛋白质含量、脂肪含量>百粒重>单株荚数>蛋白质和脂肪总量。利用复合区间作图法定位到21个与上述8个性状及其光周期反应敏感性相关的QTL,每个QTL可解释的表型变异范围在5.83%～27.85%之间。针对同一性状,未检测到在长、短日条件下均起作用的主效QTL,说明光周期不仅影响大豆的表型,而且调控相关基因的表达。作者提出,光周期对大豆的生长发育和产量形成有决定性影响,改良大豆品种的适应能力应从钝化其光周期反应敏感性入手。本研究的结果可为高产、稳产、广适应大豆品种的选育提供新的思路。
Growth period traits of soybean include not only the whole growth period (M), but also the growth period structure, i.e., the relative length of various developmental phases. In this study, the inheritance and QTL mapping of the growth period structure traits of soybean were analyzed based on segregation population derived from parents having similar M and different V, and the inheritance of the length of reproductive period (R) and the sensitivity of agronomic and quality traits to post-flowering photopeirod were also analyzed using the population derived from the parents with similar V and different M. The results were summarized as follows:
1. The inheritance of growth period structure traits in the cross with parents having different V and M.
Joint segregation analysis based on P_1, P_2, F_1, F_2, F_(2:3) generation populations derived from Xiaoheidou and GR8836 indicated that one major gene modified by poly-genes controlled the traits of V, M and R/V ratio. The major gene heritability was 77.59%, 16.77% and 54.68%, respectively. The major effect gene had positive addictive effect for V and M, and negative addictive effect for R/V. Since the major gene of R/V and V had higher heritability, then it will be effective to select the genotypes with desirable growth period structive traits.
2. QTL mapping for growth period structure traits in soybean.
F_2, F_(2:3) populations derived from Xiaoheidou and GR8836 were grown in spring and summer to map the relative genes of growth period structure traits and photothermal response sensitivity. One major effect QTL closed to Satt557 on C2 linkage group was detected, which associated with R/V ratio, V and M under different planting seasons. It was shown that the major effect QTL was stable in different environments. Meanwhile, the phenotypic variance explained by QTL, addictive and dominant effects were different for each trait, and seven minor effects QTL for growth period triats were could not be detected repeatedly under different planting seasons. Those indicated that genes of growth period traits were influenced by environmental factors to a certain extent. One major effect QTL for photothermal response sensitivity of R/V was mapped, which associated with the marker of Satt286 lied on C2 linkage group. To find the relationship of major effect QTL and E-series matutity genes controlling flowering and maturity time, Satt557 and Satt365 were used to assay the banding pattern of 41 NILs. The results indicated that the major effect QTL mapped in this study was tightly linked to E1/e1.
3. Distribution of major QTL in different maturity groups soybean varieties.
The banding patterns of 38 American varieties and 64 Chinese varieties belonging to different maturity groups were assessed using the SSR primers of Satt557 and Satt365 tightly linked to the major effect gene of growth period structure traits. The results suggested that the distribution frequency of major effect gene in American varieties was higher than that in Chinese varieties. Meanwhile, the percentage of varieties carring recessive allele to all varieties carring major QTL was higher in American maturity group varieties than that in Chinese, and the recessive allele mainly distributed in the early maturity groups, indicating that the recessive allele was more frequently used in American varieties than in Chinese varieties.
4. Evaluation of the effects of major maturity gene E1/e1 and its dependence on photothermal environment.
Effects of E1/e1 on growth period structure traits under different photoperiods and planting seasons were investigated by using maturity near-isolines of soybean. Dominant allele of E1 delayed V and M, hastened R and decreased R/V in most genotypes studied. Gene effect was dependent on the genetic background as well. There were interactions between E1 and other matrutiy genes since the dominant allele could strengthen the effects of other maturity genes generally. The interaction between E1 and environment was also detected, E1 effect under long day (LD) was higher than under short day (SD), and that in spring sowing was higher than in summer sowing.
5. The inheritance of R and R/V in the cross with parents having different (M) and similar (V).
Joint segregation analysis based on populations derived from Ludou 9 and Zhejiangsiyuebai showed that R was controlled by one major gene plus poly-genes. Heritability of the major effect gene was 44.11% in F2 generation; R/V was controlled by two major genes modified by poly-genes. Total heritability of two major genes was up to 82.37%. The addictive effects of two major genes were all positive. It was proposed that R/V ratio could be selected in early generations.
6. QTL mapping and effect analysis of yield and quality traits in soybean under different post -flowering photoperiods.
181 recombinant inbred lines (RIL) derived from Baohexuan 3×Xuyonghongdou were constructed to map QTL associated with the agronomic and quality traits under both LD (16 h) and SD (12 h) and their photoperiod response sensitivity. The order of the photopeiod sensitivity of the following eight traits was as below: reproductive period length (R)>plant height>node number on the main stem>protein content, oil content>100-seed weight>pod number per plant>total percentage of protein and oil. By using MCIM method, a total of 21 QTL were identified and mapped on 11 linkage groups including A1, A2, B1, B2, C1, D1a, etc. QTL detected could explain 5.83%～21.44% of observed phenotypic variation. For the given agronomic or quality traits, no major QTL were explored both under SD and LD. It could be drawn that photoperiod sensitivity is the key trait for adaptability improvement of soybean. To breed elite varieties with ideal yield potential, quality and stress tolerance, the photoperiod-insensitive genes related to various traits should be explored and pyramided with other needed genes.
引文
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