大豆种间杂交（Glycine max × G. soja）后代籽粒性状QTL定位
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摘要
野生大豆(Glycine soja Sieb.&Zucc.)是拓宽栽培大豆育成品种遗传基础、改善籽粒品质的有益基因源。百粒重、蛋白含量和泥膜是区别栽培大豆和野生大豆籽粒的3个显著特征。对野生大豆特别是对其重要性状的遗传研究将为认识和更好的利用这些珍贵资源提供坚实的科学基础。本研究以中国野生大豆ZYD2738为材料,与不同栽培大豆构建遗传群体,对其主要籽粒性状进行遗传分析,主要结果如下
1、构建遗传图谱2个。以冀豆12×ZYD2738组合85个单株F2群体和冀豆9号×ZYD2738组合236个单株F2群体为作图群体构建的遗传图谱,对大豆基因组覆盖率达69.2%。其中,冀豆12×ZYD2738组合遗传图谱包括了121个SSR标记,标记间平均距离10.7cM；冀豆9号×ZYD2738遗传图谱包括了143个SSR标记,标记间平均距离12.9cM。在冀豆9号×ZYD2738遗传图谱中,发现了18号染色体上的偏分离热点区域。
2、精细定位野生大豆泥膜基因B1,并利用精细定位区间两侧标记分析栽培大豆和野生大豆间的演化关系。以6个栽培大豆×野生大豆组合F2代进行遗传分析,结果表明,不同群体泥膜有无分别受1-2对基因控制遗传。以冀豆9号×ZYD2738组合204个F2纯合隐性单株和3072个F3次级群体单株为材料,将泥膜基因精细定位于13号染色体Indel标记B118和B1_9之间约43Kb范围内。利用B1精细定位区间两侧标记,以154份栽培大豆微核心种质和79份一年生野生大豆为材料,分析栽培大豆和野生大豆之间的演化关系。结果表明,不同地理来源群体间单倍型多样性水平由高到低依次为南方野生大豆≥北方野生大豆≥南方栽培大豆≥黄淮野生大豆≥北方栽培大豆≥黄淮栽培大豆,单倍型发生频率在各群体间存在差异。聚类分析可将栽培大豆和野生大豆区分开,揭示出北方和黄淮材料间较近的遗传关系,但定位区间单倍型地理来源特异不明显。
3、精细定位百粒重位点qSWT_13_1。冀豆12×ZYD2738组合F2:3群体在石家庄种植定位到5号、7号、10号和13号染色体上的4个QTL,F2：4群体在三亚种植定位到2号和10号染色体上的2个QTL；冀豆9号×ZYD2738组合F2：3群体在石家庄种植定位到9号、13号和14号染色体上的3个QTL, F2:4群体在三亚种植定位到6号和9号染色体上的2个QTL。利用F6：7次级分离群体,将qSWT_13_1精细定位在13号染色体Satt663和Satt114之间物理距离3.27Mb范围内。以341份一年生野生大豆和栽培大豆种质资源和17份全基因组重测序种质资源为材料,通过关联分析,进一步将精细定位区间缩小至QSSNP-Gm13-0095635和QSSNP-Gm13-0101267之间物理距离1.49Mb范围内。
4、定位野生大豆ZYD2738高蛋白位点qPRO_20_1并明确互作效应。以冀豆12×ZYD2738和冀豆9号×ZYD2738两个组合F2：3家系为材料,均定位到位于20号染色体的野生大豆高蛋白位点qPRO_20_1。在冀豆12×ZYD2738组合中,高蛋白位点与标记Satt239和Satt354相关,ZYD2738等位变异蛋白含量较冀豆12等位变异高出2.18%,在冀豆9号×ZYD2738组合中,高蛋白位点与标记Satt419和Satt270相关,ZYD2738等位变异蛋白含量较冀豆9号等位变异高出1.29%。在冀豆9号×ZYD2738组合中检测到了qPRO_20_1对另一个蛋白含量位点qPR0_19_1的屏蔽作用。分子标记辅助选择时,同时以qPRO_20_1和qPRO_19_1作为选择位点较仅以qPRO_20_1作选择位点,后代蛋白含量由40.43%提高到41.15%。
本研究获得了具有育种利用价值的QTL位点和种质资源,为利用野生大豆优异基因拓宽栽培大豆遗传基础奠定了理论和材料基础。
Wild soybean (Glycine soja) is an important genetic resource for cultivated soybean (Glycine max) improvement. Genetic study of important traits in the G.soja will provide a solid scientific basis for better understanding of the genetic mechanisms and practically using of those for the trait improvement. Three seed characters that distinguish cultivated with wild soybean are100-seed weight, protein content, and seed bloom. In the present research, we studied the genetic basis of those traits using the wild soybean accession ZYD2738that originated in China. The followings are the major results obtained from the research:
1.Two genetic linkage maps were developed by using the F2populations of Jidou12×ZYD2738and Jidou9hao X ZYD2738which were derived from crossing of wild soybean ZYD2738with two cultivars Jidou12and Jidou9hao. The genome coverage ratio in the genetic linkage map reached69.2%. A total of121SSR makers were anchored with average distance of10.7cM in85individuals of Jidou12×ZYD2738population whereas143SSR makers were anchored with average distance of12.9cM in236individuals of Jidou9hao×ZYD2738population by the linkage map. A segregation distortion region on chromosome18was found in the map of Jidou9hao×ZYD2738, but the distortion loci had little effects on linkage map.
2.The seed bloom locus (B1) was fine mapped, and the flanking makers were used to analysis the relationship between wild and cultivated germplasm. The results showed that the seed bloom may be controlled by one or two genes among six F2populations. By fine mapping in3072F3individuals, B1was further defined within43Kb region between Indel marker B1_18and B1_9on chromosome13, containing5candidate genes which needed to be confirmed. A total of161G.max and100G.soja were compared to estimate the relationship between wild and cultivated soybean based on B1flanking markers. The results showed that the diversity of G.soja was higher than G.max. The diversity of accessions from geological distribution ranked as South region> North region> Huanghuai region but no obviously geographic distinction was found among the distribution of haplotype.
3. The100-seed weight QTL qSWT_13_1was fine mapped on chromosome13. F2:3and F2:4populations of Jidou12X ZYD2738and Jidou9hao X ZYD2738were used to map the QTL of100-seed weight in2environments (Shijiazhuang and Sanya). Four QTL were mapped on chromosome5,7,10and13in Jidou12×ZYD2738population under Shijiazhuang environment, while2QTL were mapped on chromosome2and10in the same population under Sanya environment. In Jidou9hao×ZYD2738population, there were4QTL were mapped on chromosome9,13and14in Shijiazhuang, while2QTL were mapped on chromosome6and9in Sanya. The stable QTL qSWT_13_1in2populations was fine mapped on chromosome13flanked by SSR marker Satt663and Sattl14that spans3.27Mb physical distance in advanced analysis using F6:7sub-population, then the region was reduced to1.49Mb based on association mapping using the data of17re-sequenced accessions.
4. The F2:3progenies of Jidou12×ZYD2738and Jidou9hao×ZYD2738were used to discover the QTL for protein and oil content in Shijiazhuang environment. Six protein QTL and5oil QTL were identified in2populations. The qPRO_20_1on chromosome20was a stable QTL under2populations. The allele from ZYD2738contributed2.18%more protein than that from Jidou12and1.29%more protein than that from Jidou9hao. In the population of Jidou9hao×ZYD2738, qPRO_20_1showed epistatic role over the qPRO_19_l on chromosome19. For maker assistant breeding, the protein content was40.43%in progenies selected by qPRO_20_1compared to the41.15%in the progenies selected by both qPRO_19_1and qPRO_20_1.
The QTL and germplasm lines with breeding value were detected and summarized in the present study. The results will be helpful to expand the genetic base in soybean improvement program by introducing favorite genes from Gsoja.
1、构建遗传图谱2个。以冀豆12×ZYD2738组合85个单株F2群体和冀豆9号×ZYD2738组合236个单株F2群体为作图群体构建的遗传图谱,对大豆基因组覆盖率达69.2%。其中,冀豆12×ZYD2738组合遗传图谱包括了121个SSR标记,标记间平均距离10.7cM；冀豆9号×ZYD2738遗传图谱包括了143个SSR标记,标记间平均距离12.9cM。在冀豆9号×ZYD2738遗传图谱中,发现了18号染色体上的偏分离热点区域。
2、精细定位野生大豆泥膜基因B1,并利用精细定位区间两侧标记分析栽培大豆和野生大豆间的演化关系。以6个栽培大豆×野生大豆组合F2代进行遗传分析,结果表明,不同群体泥膜有无分别受1-2对基因控制遗传。以冀豆9号×ZYD2738组合204个F2纯合隐性单株和3072个F3次级群体单株为材料,将泥膜基因精细定位于13号染色体Indel标记B118和B1_9之间约43Kb范围内。利用B1精细定位区间两侧标记,以154份栽培大豆微核心种质和79份一年生野生大豆为材料,分析栽培大豆和野生大豆之间的演化关系。结果表明,不同地理来源群体间单倍型多样性水平由高到低依次为南方野生大豆≥北方野生大豆≥南方栽培大豆≥黄淮野生大豆≥北方栽培大豆≥黄淮栽培大豆,单倍型发生频率在各群体间存在差异。聚类分析可将栽培大豆和野生大豆区分开,揭示出北方和黄淮材料间较近的遗传关系,但定位区间单倍型地理来源特异不明显。
3、精细定位百粒重位点qSWT_13_1。冀豆12×ZYD2738组合F2:3群体在石家庄种植定位到5号、7号、10号和13号染色体上的4个QTL,F2：4群体在三亚种植定位到2号和10号染色体上的2个QTL；冀豆9号×ZYD2738组合F2：3群体在石家庄种植定位到9号、13号和14号染色体上的3个QTL, F2:4群体在三亚种植定位到6号和9号染色体上的2个QTL。利用F6：7次级分离群体,将qSWT_13_1精细定位在13号染色体Satt663和Satt114之间物理距离3.27Mb范围内。以341份一年生野生大豆和栽培大豆种质资源和17份全基因组重测序种质资源为材料,通过关联分析,进一步将精细定位区间缩小至QSSNP-Gm13-0095635和QSSNP-Gm13-0101267之间物理距离1.49Mb范围内。
4、定位野生大豆ZYD2738高蛋白位点qPRO_20_1并明确互作效应。以冀豆12×ZYD2738和冀豆9号×ZYD2738两个组合F2：3家系为材料,均定位到位于20号染色体的野生大豆高蛋白位点qPRO_20_1。在冀豆12×ZYD2738组合中,高蛋白位点与标记Satt239和Satt354相关,ZYD2738等位变异蛋白含量较冀豆12等位变异高出2.18%,在冀豆9号×ZYD2738组合中,高蛋白位点与标记Satt419和Satt270相关,ZYD2738等位变异蛋白含量较冀豆9号等位变异高出1.29%。在冀豆9号×ZYD2738组合中检测到了qPRO_20_1对另一个蛋白含量位点qPR0_19_1的屏蔽作用。分子标记辅助选择时,同时以qPRO_20_1和qPRO_19_1作为选择位点较仅以qPRO_20_1作选择位点,后代蛋白含量由40.43%提高到41.15%。
本研究获得了具有育种利用价值的QTL位点和种质资源,为利用野生大豆优异基因拓宽栽培大豆遗传基础奠定了理论和材料基础。
Wild soybean (Glycine soja) is an important genetic resource for cultivated soybean (Glycine max) improvement. Genetic study of important traits in the G.soja will provide a solid scientific basis for better understanding of the genetic mechanisms and practically using of those for the trait improvement. Three seed characters that distinguish cultivated with wild soybean are100-seed weight, protein content, and seed bloom. In the present research, we studied the genetic basis of those traits using the wild soybean accession ZYD2738that originated in China. The followings are the major results obtained from the research:
1.Two genetic linkage maps were developed by using the F2populations of Jidou12×ZYD2738and Jidou9hao X ZYD2738which were derived from crossing of wild soybean ZYD2738with two cultivars Jidou12and Jidou9hao. The genome coverage ratio in the genetic linkage map reached69.2%. A total of121SSR makers were anchored with average distance of10.7cM in85individuals of Jidou12×ZYD2738population whereas143SSR makers were anchored with average distance of12.9cM in236individuals of Jidou9hao×ZYD2738population by the linkage map. A segregation distortion region on chromosome18was found in the map of Jidou9hao×ZYD2738, but the distortion loci had little effects on linkage map.
2.The seed bloom locus (B1) was fine mapped, and the flanking makers were used to analysis the relationship between wild and cultivated germplasm. The results showed that the seed bloom may be controlled by one or two genes among six F2populations. By fine mapping in3072F3individuals, B1was further defined within43Kb region between Indel marker B1_18and B1_9on chromosome13, containing5candidate genes which needed to be confirmed. A total of161G.max and100G.soja were compared to estimate the relationship between wild and cultivated soybean based on B1flanking markers. The results showed that the diversity of G.soja was higher than G.max. The diversity of accessions from geological distribution ranked as South region> North region> Huanghuai region but no obviously geographic distinction was found among the distribution of haplotype.
3. The100-seed weight QTL qSWT_13_1was fine mapped on chromosome13. F2:3and F2:4populations of Jidou12X ZYD2738and Jidou9hao X ZYD2738were used to map the QTL of100-seed weight in2environments (Shijiazhuang and Sanya). Four QTL were mapped on chromosome5,7,10and13in Jidou12×ZYD2738population under Shijiazhuang environment, while2QTL were mapped on chromosome2and10in the same population under Sanya environment. In Jidou9hao×ZYD2738population, there were4QTL were mapped on chromosome9,13and14in Shijiazhuang, while2QTL were mapped on chromosome6and9in Sanya. The stable QTL qSWT_13_1in2populations was fine mapped on chromosome13flanked by SSR marker Satt663and Sattl14that spans3.27Mb physical distance in advanced analysis using F6:7sub-population, then the region was reduced to1.49Mb based on association mapping using the data of17re-sequenced accessions.
4. The F2:3progenies of Jidou12×ZYD2738and Jidou9hao×ZYD2738were used to discover the QTL for protein and oil content in Shijiazhuang environment. Six protein QTL and5oil QTL were identified in2populations. The qPRO_20_1on chromosome20was a stable QTL under2populations. The allele from ZYD2738contributed2.18%more protein than that from Jidou12and1.29%more protein than that from Jidou9hao. In the population of Jidou9hao×ZYD2738, qPRO_20_1showed epistatic role over the qPRO_19_l on chromosome19. For maker assistant breeding, the protein content was40.43%in progenies selected by qPRO_20_1compared to the41.15%in the progenies selected by both qPRO_19_1and qPRO_20_1.
The QTL and germplasm lines with breeding value were detected and summarized in the present study. The results will be helpful to expand the genetic base in soybean improvement program by introducing favorite genes from Gsoja.
引文
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