甘蓝型油菜隐性细胞核雄性不育上位抑制基因的精细定位
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摘要
隐性核不育由于具有不育性彻底且稳定、恢复源广和细胞质来源丰富等优点,在油菜杂种优势利用中具有很大的潜力。然而,由于不能获得高效而经济的核不育系保持方法,大部分的隐性核不育不能广泛应用于油菜杂交种生产。幸运的是,隐性上位互作核不育系9012AB的发现克服了传统隐性核不育在杂种优势应用上的困难。遗传分析表明,9012AB的育性受两对重叠隐性不育基因(ms3和ms4)和一对隐性上位抑制基因(esp)互作控制,两对隐性不育基因纯合时导致雄性不育,但任何一个不育基因为显性或隐性上位抑制基因纯合时育性恢复。根据这一遗传模型,将纯合不育系和临保系杂交就可以获得100%不育群体,避免了传统隐性核不育系在杂交种生产过程中拔去母本行50%可育株的麻烦。在中国,这种隐性核不育系的保持途径已经成功应用于甘蓝型油菜杂交种的生产。值得注意的是,通过传统的育种方法选育不育系及其临保系是一项繁琐、低效的工作,因此,为了加快育种进程,必须开发与ms3、ms4和esp紧密连锁的分子标记。本研究以甘蓝型油菜隐性上位互作核不育系9012AB、GosAB及其临保系T45为研究材料,构建了两个BC1定位群体,并开发与esp基因紧密连锁的分子标记,进而实现esp基因的精细定位,取得的主要结果如下：
1.esp基因连锁分子标记的开发。本研究采用以下3种策略开发与esp基因紧密连锁的分子标记。第一、BSA法结合AFLP技术。通过筛选2,816对AFLP引物组合,获得了17个与esp基因连锁的AFLP标记,其中10个AFLP标记在包含143个单株的群体Ⅰ中跟esp基因是共分离的。为了方便大群体分析,将其中7个共分离的AFLP标记转化成显性的SCAR标记。第二、遗传图谱整合。根据esp基因在甘蓝型油菜N7连锁群上的相对位置,从芸薹属A7连锁群相对应的区间挑选PCR标记对含有188个单株的群体Ⅱ进行分析,结果表明,总共有11个PCR标记,包括1个SCAR标记、7个SSR标记和3个IP标记,跟esp基因是连锁的。另外,3个基于白菜BAC序列的SSR标记在群体Ⅱ也是跟esp基因连锁的。第三、比较作图。通过对上述两个定位群体中部分标记序列的BLASTN分析并结合前人的研究结果,发现甘蓝型油菜N7连锁群esp基因及其邻近区间由G、H、F和B等4个保守区段组成,其中F保守区段将esp基因包含在内。随后,F保守区段中的拟南芥基因被用于IP标记的开发。IP引物设计在与拟南芥基因外显子高度同源的芸薹属EST和GSS序列上,总共设计122对引物。亲本多态性检测和群体分析表明,一共有14个IP标记在两个BC1定位群体中都是与esp基因紧密连锁的。
2.esp基因的精细定位。将与esp基因紧密连锁的PCR标记进一步分析3,878个单株的群体Ⅰ和3,484个单株的群体Ⅱ。为了提高大群体的分析效率,采用侧翼分子标记分析策略。在群体Ⅰ中,esp基因被定位在SCAR标记WSC6和IP标记IP5-3之间,其遗传距离分别为0.206cM和0.129cM。在群体Ⅱ中,esp基因被定位在IP标记IP36和IP2-3之间,其遗传距离分别为0.057cM和0.057cM。整合两个群体的定位结果,esp基因被IP标记IP36和IP5-3所限定在一个很小的遗传区间。虽然一共分析包含7,362个单株的两个BC1定位群体,仍然有6个SCAR标记和3个IP标记在两个群体中都是与esp基因共分离。
3.甘蓝型油菜和拟南芥在esp区间的微观共线性分析。利用来源于拟南芥的IP标记精细定位esp基因为分析甘蓝型油菜和拟南芥之间的微观共线性提供了便利。甘蓝型油菜和拟南芥在esp区间的完美共线性被染色体片段的倒位和基因的插入所打断。由于倒位的发生,导致了由esp两侧最近的标记IP36和IP5-3所限定的拟南芥候选区间从一个变成为两个。第一个拟南芥候选区间是由标记IP5-3和IP9-4界定,区间大小为54kb,包含12个预测基因。第二个拟南芥候选区间是由标记At3G24315和IP36界定,区间大小为40kb,包含8个预测基因。遗憾的是,在这20个拟南芥预测基因中,没有鉴定出esp的候选基因。
It is widely accepted that recessive genic male sterility (RGMS) systems have great potential in rapeseed heterosis utilization because of their stable and complete sterility, extensive distribution of restorers and diverse cytoplasmic sources. However, for most of them, their potential application for the commercial production of rapeseed hybrids is limited by the inability to maintain the male-sterile female parent line in an efficient and cost-effective way. About 50% male fertile plants must be removed during hybrid seed production, which increases the cost of hybrid seeds and limits its wide application. Fortunately, this difficulty can be overcome when a novel RGMS line,9012AB, is involved. Genetic analysis indicated that the sterility of 9012AB is controlled by two recessive genes (ms3 and ms4) interacting with one recessive epistatic suppressor gene (esp). Homozygous recessive sterile gene at both loci can result in male sterility, but the fertility can be restored when the esp gene is in recessive homozygosity or either of the sterile genes is not recessively homozygous. Based on this genetic model, a 100% male-sterile population could be produced by crossing a homozygous male-sterile line with a temporary maintainer line. This strategy is used successfully for the commercial production of canola hybrids (Brassica napus) in China. Care needs to be taken that the breeding of elite RGMS lines and their temporary maintainers by conventional methods is very laborious and time-consuming. It is therefore highly desirable to develop molecular marker tightly linked to the genes (ms3, ms4 and esp), which will greatly accelerate these breeding process. The purposes of this investigation are to develop molecular markers tightly linked to the esp gene, using two BC1 mapping populations derived from recessive epistatic genic male sterility line (9012A and GosA) and their temporary maintainer (T45), and then to construct a high-resolution map surrounding the esp gene. The main results were as follows:
1. Development of molecular markers linked to the esp gene. Several strategies were used to develop molecular markers associated with the target gene. Firstly, AFLP technology combined with bulked segregant analysis (BSA). From the survey of 2,816 AFLP primer combinations, seventeen tightly linked AFLP markers were obtained. Among them, ten AFLP makers co-segregated with the target gene in the populationⅠof 143 individuals. Given that AFLP has limitations in large-scale population analysis, seven co-segregated AFLP markers were converted to dominant SCAR makers. Secondly, genetic map integration. Considering the esp gene was mapped on linkage group N7 of Brassica napus, PCR markers surrounding corresponding region were selected from linkage group A7 to analyze the populationⅡof 188 plants. Results indicated that 14 PCR markers including 1 SCAR marker,7 SSR markers,2 IP markers and 3 BAC sequence-based markers were assicated with the esp gene. Thirdly, comparative mapping. Sequences of the esp-linked PCR markers were submitted to NCBI for BLASTN analysis. Combined with previous reports on comparative analysis, we found that the esp region in Brassica napus linkage group N7 is constituted of blocks G-H-F-B. The Arabidopsis genes of block F encompassing the esp gene were employed for the development of Intron Polymorphism (IP) markers. IP primers were designed from exon sequences which showed strong nucleotide conservation between Arabidopsis and the corresponding EST or GSS sequences described for any Brassica species, and 122 primer pairs were designed. After the polymorphism survey in parents and mapping population,14 IP markers were found to be tightly linked to the esp gene in both populationⅠand populationⅡ.
2. Fine mapping of the esp gene. The esp-linked PCR markers described above were used to tested on 3,878 plants from populationⅠand 3,484 plants from populationⅡ. We employed the flanking marker approach to improve the efficiency of large population analysis. In the populationⅠ, the esp gene was then genetically restricted to a region of 0.335 cM,0.206 cM from SCAR marker WSC6 and 0.129 cM from IP marker IP5-3. In the populationⅡ, the target gene was mapped between IP marker IP36 and IP marker IP02-3 with the distance of 0.057 cM and 0.057 cM, respectively. Integrating the mapping results from two larger populations, the esp was fine-mapped to the interval between IP markers IP36 and IP05-3. Despite of 7,362 plants from two mapping population were analyzed, six SCAR makers and three IP makers were still co-segregated with the esp gene.
3. Microcollinearity between Brassica napus and Arabidopsis at the esp locus region. The presence of many Arabidopsis-derived IP markers in the local high-resolution map allowed us to study the collinearity between Brassica napus and Arabidopsis at the esp locus region. Microcollinearity was found between the region on N7 carrying the esp gene and Arabidopsis chromosome 3, interrupted by chromosome segment inversion and gene inserts. Due to the inversion directly flanking the map position of esp, extending the search for candidate genes to an area delimited by the flanking markers, IP36 and IP5-3, identifies two regions on Arabidopsis chromosome 3. The first is a collinear region between IP markers IP5-3 and IP9-4 covering an interval of 54kb and containing 12 annotated genes. The second candidate region which locate between IP markers At3G24315 and IP36 cover an interval of 40kb and include 8 annotated genes. Unfortunately, no obvious candidate gene for esp was identified among 20 annotated genes in two Arabidopsis collinear region.
1.esp基因连锁分子标记的开发。本研究采用以下3种策略开发与esp基因紧密连锁的分子标记。第一、BSA法结合AFLP技术。通过筛选2,816对AFLP引物组合,获得了17个与esp基因连锁的AFLP标记,其中10个AFLP标记在包含143个单株的群体Ⅰ中跟esp基因是共分离的。为了方便大群体分析,将其中7个共分离的AFLP标记转化成显性的SCAR标记。第二、遗传图谱整合。根据esp基因在甘蓝型油菜N7连锁群上的相对位置,从芸薹属A7连锁群相对应的区间挑选PCR标记对含有188个单株的群体Ⅱ进行分析,结果表明,总共有11个PCR标记,包括1个SCAR标记、7个SSR标记和3个IP标记,跟esp基因是连锁的。另外,3个基于白菜BAC序列的SSR标记在群体Ⅱ也是跟esp基因连锁的。第三、比较作图。通过对上述两个定位群体中部分标记序列的BLASTN分析并结合前人的研究结果,发现甘蓝型油菜N7连锁群esp基因及其邻近区间由G、H、F和B等4个保守区段组成,其中F保守区段将esp基因包含在内。随后,F保守区段中的拟南芥基因被用于IP标记的开发。IP引物设计在与拟南芥基因外显子高度同源的芸薹属EST和GSS序列上,总共设计122对引物。亲本多态性检测和群体分析表明,一共有14个IP标记在两个BC1定位群体中都是与esp基因紧密连锁的。
2.esp基因的精细定位。将与esp基因紧密连锁的PCR标记进一步分析3,878个单株的群体Ⅰ和3,484个单株的群体Ⅱ。为了提高大群体的分析效率,采用侧翼分子标记分析策略。在群体Ⅰ中,esp基因被定位在SCAR标记WSC6和IP标记IP5-3之间,其遗传距离分别为0.206cM和0.129cM。在群体Ⅱ中,esp基因被定位在IP标记IP36和IP2-3之间,其遗传距离分别为0.057cM和0.057cM。整合两个群体的定位结果,esp基因被IP标记IP36和IP5-3所限定在一个很小的遗传区间。虽然一共分析包含7,362个单株的两个BC1定位群体,仍然有6个SCAR标记和3个IP标记在两个群体中都是与esp基因共分离。
3.甘蓝型油菜和拟南芥在esp区间的微观共线性分析。利用来源于拟南芥的IP标记精细定位esp基因为分析甘蓝型油菜和拟南芥之间的微观共线性提供了便利。甘蓝型油菜和拟南芥在esp区间的完美共线性被染色体片段的倒位和基因的插入所打断。由于倒位的发生,导致了由esp两侧最近的标记IP36和IP5-3所限定的拟南芥候选区间从一个变成为两个。第一个拟南芥候选区间是由标记IP5-3和IP9-4界定,区间大小为54kb,包含12个预测基因。第二个拟南芥候选区间是由标记At3G24315和IP36界定,区间大小为40kb,包含8个预测基因。遗憾的是,在这20个拟南芥预测基因中,没有鉴定出esp的候选基因。
It is widely accepted that recessive genic male sterility (RGMS) systems have great potential in rapeseed heterosis utilization because of their stable and complete sterility, extensive distribution of restorers and diverse cytoplasmic sources. However, for most of them, their potential application for the commercial production of rapeseed hybrids is limited by the inability to maintain the male-sterile female parent line in an efficient and cost-effective way. About 50% male fertile plants must be removed during hybrid seed production, which increases the cost of hybrid seeds and limits its wide application. Fortunately, this difficulty can be overcome when a novel RGMS line,9012AB, is involved. Genetic analysis indicated that the sterility of 9012AB is controlled by two recessive genes (ms3 and ms4) interacting with one recessive epistatic suppressor gene (esp). Homozygous recessive sterile gene at both loci can result in male sterility, but the fertility can be restored when the esp gene is in recessive homozygosity or either of the sterile genes is not recessively homozygous. Based on this genetic model, a 100% male-sterile population could be produced by crossing a homozygous male-sterile line with a temporary maintainer line. This strategy is used successfully for the commercial production of canola hybrids (Brassica napus) in China. Care needs to be taken that the breeding of elite RGMS lines and their temporary maintainers by conventional methods is very laborious and time-consuming. It is therefore highly desirable to develop molecular marker tightly linked to the genes (ms3, ms4 and esp), which will greatly accelerate these breeding process. The purposes of this investigation are to develop molecular markers tightly linked to the esp gene, using two BC1 mapping populations derived from recessive epistatic genic male sterility line (9012A and GosA) and their temporary maintainer (T45), and then to construct a high-resolution map surrounding the esp gene. The main results were as follows:
1. Development of molecular markers linked to the esp gene. Several strategies were used to develop molecular markers associated with the target gene. Firstly, AFLP technology combined with bulked segregant analysis (BSA). From the survey of 2,816 AFLP primer combinations, seventeen tightly linked AFLP markers were obtained. Among them, ten AFLP makers co-segregated with the target gene in the populationⅠof 143 individuals. Given that AFLP has limitations in large-scale population analysis, seven co-segregated AFLP markers were converted to dominant SCAR makers. Secondly, genetic map integration. Considering the esp gene was mapped on linkage group N7 of Brassica napus, PCR markers surrounding corresponding region were selected from linkage group A7 to analyze the populationⅡof 188 plants. Results indicated that 14 PCR markers including 1 SCAR marker,7 SSR markers,2 IP markers and 3 BAC sequence-based markers were assicated with the esp gene. Thirdly, comparative mapping. Sequences of the esp-linked PCR markers were submitted to NCBI for BLASTN analysis. Combined with previous reports on comparative analysis, we found that the esp region in Brassica napus linkage group N7 is constituted of blocks G-H-F-B. The Arabidopsis genes of block F encompassing the esp gene were employed for the development of Intron Polymorphism (IP) markers. IP primers were designed from exon sequences which showed strong nucleotide conservation between Arabidopsis and the corresponding EST or GSS sequences described for any Brassica species, and 122 primer pairs were designed. After the polymorphism survey in parents and mapping population,14 IP markers were found to be tightly linked to the esp gene in both populationⅠand populationⅡ.
2. Fine mapping of the esp gene. The esp-linked PCR markers described above were used to tested on 3,878 plants from populationⅠand 3,484 plants from populationⅡ. We employed the flanking marker approach to improve the efficiency of large population analysis. In the populationⅠ, the esp gene was then genetically restricted to a region of 0.335 cM,0.206 cM from SCAR marker WSC6 and 0.129 cM from IP marker IP5-3. In the populationⅡ, the target gene was mapped between IP marker IP36 and IP marker IP02-3 with the distance of 0.057 cM and 0.057 cM, respectively. Integrating the mapping results from two larger populations, the esp was fine-mapped to the interval between IP markers IP36 and IP05-3. Despite of 7,362 plants from two mapping population were analyzed, six SCAR makers and three IP makers were still co-segregated with the esp gene.
3. Microcollinearity between Brassica napus and Arabidopsis at the esp locus region. The presence of many Arabidopsis-derived IP markers in the local high-resolution map allowed us to study the collinearity between Brassica napus and Arabidopsis at the esp locus region. Microcollinearity was found between the region on N7 carrying the esp gene and Arabidopsis chromosome 3, interrupted by chromosome segment inversion and gene inserts. Due to the inversion directly flanking the map position of esp, extending the search for candidate genes to an area delimited by the flanking markers, IP36 and IP5-3, identifies two regions on Arabidopsis chromosome 3. The first is a collinear region between IP markers IP5-3 and IP9-4 covering an interval of 54kb and containing 12 annotated genes. The second candidate region which locate between IP markers At3G24315 and IP36 cover an interval of 40kb and include 8 annotated genes. Unfortunately, no obvious candidate gene for esp was identified among 20 annotated genes in two Arabidopsis collinear region.
引文
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