大白菜细胞核雄性不育基因的分子标记及定位
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摘要
大白菜是起源于中国的十字花科芸薹属作物,也是中国及很多其他国家和地区十分重要的蔬菜作物之一。大白菜为典型的异花授粉作物,存在明显的杂种优势。利用自交不亲和系和雄性不育系制种是目前最常用的杂交制种技术。自交不亲和系在利用上存在亲本繁殖困难成本提高、多代自交生活力下降;易受环境影响使杂种纯度不高。利用雄性不育系制种是一种理想的杂种生产模式。雄性不育系又被分为细胞质雄性不育系和细胞核雄性不育系。细胞核雄性不育系不育性完全,不会影响植株生活力。目前,对大白菜细胞核雄性不育的遗传解释存在多种遗传模式,不同遗传模式将直接影响雄性不育的转育。同时,利用分子标记方法对不育基因和恢复基因的研究较为初步。
本研究对两个隐性细胞核雄性不育系(452AB和454AB)和一个显性细胞核雄性不育系(451AB)进行了遗传学研究,判断不育系的遗传模式,不育基因在自交系中的分布,并对三个不育系的育性恢复基因和不育基因进行了分子标记、基因定位。主要结果如下:
1.隐性雄性不育系452AB和454AB的可育株和不育株分别与显性雄性不育系451AB中的可育株杂交,并进一步回交。通过数据分析推断451AB系不育基因与454AB系恢复基因为等位基因,这两个系属于复等位遗传模式。452AB系与另外两个系不同,表现为独立遗传。三个不育系与17个自交系进行测交,判断自交系基因型。根据测交结果,4个自交系的可育基因对452AB系不育基因有保持作用。进一步研究判断452AB系也为一复等位雄性不育系。两复等位雄性不育系在基因组中的位置不同,并不存在关联。
2.隐性细胞核雄性不育系452AB包含164个单株。利用混合分组分离法(BSA)对不育系进行标记分析,并优化SRAP反应体系。在构建的基因池中筛选SRAP引物1 256对,SSR引物187对。获得与大白菜隐性细胞核雄性不育系恢复基因(BrMsf2)连锁的标记两个,PM8K4和Me2M49,与恢复基因的遗传距离分别为2.98 cM和10.92 cM。通过调查PM8K4标记在大白菜DH作图群体中的多态性,将该恢复基因定位在大白菜染色体A08。根据大白菜基因组测序信息,筛选来自染色体A08候选区域的Indel标记23个,其中scaffold20上的标记S20-1与恢复基因的遗传距离为4.22 cM。通过S20-1和PM8K4标记在染色体上的位置,将BrMsf2定位在scaffold20的附近。
3.显性细胞核雄性不育系451AB包含480个单株。利用混合分组分离法、SRAP和SSR标记技术筛选与不育基因BrMs1连锁的标记。在1 378对SRAP标记和238对SSR标记中,筛选到与不育基因连锁的SRAP标记5个(CuMe7BEm12, BMe9E32, PM8E38, E35BEm10, and M51K2),SSR标记1个(ENA6)。其中PM8E38标记与不育基因的距离最近,为0.37 cM。将该标记多态性片段进行克隆测序,成功的转化为较为简易的SCAR标记PM8E38S。通过SSR标记ENA6在DH作图群体中的位置,不育基因BrMs1被定位在大白菜染色体A07上。
4.利用2357个单株对451AB系不育基因进行精细定位。初步定位标记PM8E38S在精细定位群体中与不育基因的遗传距离为0.59cM。将初步定位标记的序列与基因组序列进行比对,没有比对在染色体A07上。筛选根据白菜基因组序列设计的内含子多态性(Intron Polymorphism,IP)标记13对,与芥菜中设计的IP标记28对,筛选到一个与不育基因距离较远的标记A0703-2。设计开发基于测序技术的酶切标签标记,并首次应用于雄性不育基因的精细定位中。筛选酶切标签标记76个,获得4个基因组位置已知的标记S17-52、S1727-5、S1727-6、S1729-5和6个基因组位置未知的标记SNONE-2、SNONE-7、SNONE-9,SNONE-16,SNONE-27和SNONE-44,其中SNONE-9与不育基因遗传距离最小(0.51 cM)。S1727-6是与不育基因距离最短(0.76 cM)的位置已知的标记。进一步对该区域Indel标记进行筛选,从26对引物中,筛选到4对存在多态性的Indel标记(S17-1、S17-2、S69-1和S69-2)。通过Indel标记以及酶切标签标记在基因组中的位置,将标记与基因之间的遗传距离与物理距离想对应。通过基因所在区域序列分析结果,预测该基因位于重复序列较多的中心粒区。
5. 454AB系为隐性细胞核雄性不育系,它包含320个单株。利用混合分组分离法根据育性构建育性池。筛选SRAP标记1 128个,SSR标记187个。筛选出与恢复基因BrMsf3连锁的SRAP标记两个,BMe10SA4和M52K2,与恢复基因的连锁距离为4.38 cM和7.46 cM,这两个标记位于基因的同侧; SSR标记ENA6与恢复基因的遗传连锁距离为11.57cM。通过ENA6标记在DH作图群体中的位置,将该恢复基因定位在大白菜染色体A07上。进一步筛选位于A07上的Indel标记26个,酶切标签标记65个,未发现与恢复基因(BrMsf3)连锁的标记。基因定位结果证实了451AB系和454AB系的复等位遗传关系。
6.通过遗传学分析和基因定位结果,得到451AB系和454AB系属于复等位遗传,不育基因位于染色体A07上,452AB系属于复等位遗传,不育基因位于染色体A08上,证明大白菜中存在一个以上复等位雄性不育系统。
Chinese cabbage (Brassica rapa L. ssp. pekinensis) is an important vegetable crop in China. It originated in China and has many advantageous economic characteristics. It has quite strong heterosis. Self-incompatible lines and male sterile lines are widely used in F1 seed production. The application of self-incompatible lines is limited by the reduction of fitness and vigor, the inconvenience of selfing, and the susceptibility of plants to environmental conditions. Male sterile lines are ideal systems for hybrid production. Plant male sterility can be generally classified into cytoplasmic male sterility (CMS) and genic male sterility (GMS). In GMS lines, sterility is stable and complete, and GMS does not result in reduced vigor. The study of genic male sterility of Brassica rapa showed different genetic models. The methods of transferring male sterility genes depend on certain genetic models. The genetic models of the genic male sterility lines had to be identified before they can be used for breeding. Molecular markers analysis of those male sterility genes just started in recent years.
This thesis deals with two recessive genic male sterility lines (452AB and 454AB) and one domiant genic male sterility line (451AB) to confirm their genetic models, to investigate distribution of different male sterile genes in male fertile inbred lines, and to find the molecular markers linked to the restoring genes and male sterility gene, and to map these genes. The main conclusions are as follows:
1. The genetic analysis in the recessive genic male sterility lines (452AB and 454AB) and dominant genic male sterility line (451AB) was conducted. The sterile plants and fertile plants in 452AB line and 454AB line were crossed with the fertile plants in 451AB line. According to the results, BrMs1 gene in 451AB and BrMsf3 gene in 454AB line were at the same allelic locus. So the genetic model of 451AB line and 454AB line was conformed to be a multiple-allele model. 452AB line had no relationship with 451AB line and 454AB line. For transferring these genes to other lines, male sterile plants were backcrossed with 17 inbred lines to confirm their genotypes. Four inbred lines can maintain sterility gene in 452AB line and resulted in 100% male sterile populations. The further experiments confirmed the genetic model of 452AB line also to be multiple-allele, but the genespositions were different from those in line 451AB and line 454AB.
2. Line 452AB was recessive genic male sterility (RGMS), which contained 164 individuals. Bulked segregant analysis (BSA) was used to screen markers linked to restoring gene BrMsf2. After SRAP (Sequence-related amplified polymorphism) markers system was optimized, 1 256 SRAP primer pairs and 187 simple sequence repeat (SSR) primer pairs were analyzed between fertile bulks. Two SRAP markers PM8K4 and Me2M49 were found, and the distance of PM8K4 and Me2M49 was 2.98 cM and 10.92 cM, respectively. PM8K4 was subsequently mapped on chromosome A08 using a doubled-haploid mapping population. Screened 23 Indel markers on A08 from Brassica rapa genome sequence, one marker (S20-1) in scaffold 20 showed polymorphism, and the genetic distance was 4.22 cM. According the positions of S20-1 and PM8K4 on chromosome A08, BrMsf2 was mapped around scaffold 20 in A08.
3. Line 451AB was dominant genic male sterility (DGMS), which produced 480 individuals. SRAP, SSR and sequence-characterized amplified region (SCAR) analyses were performed to screen markers linked to BrMs1. Five of the 1 378 SRAP primer pair combinations (CuMe7BEm12, BMe9E32, PM8E38, E35BEm10, and M51K2) and one of 238 SSRs (ENA6) revealed polymorphisms between bulks from male sterile and fertile progeny. PM8E38, the nearest marker linked to BrMs1 (0.37 cM), was converted to the SCAR marker PM8E38S. ENA6 was subsequently mapped on chromosome A07 using a doubled haploid mapping population.
4. A fine mapping population which contained 2 357 individuals was constructed in 451AB line. In this larger population, the distance between PM8E38S and BrMs1 was changed to 0.59 cM. None of them was mapped on A07 according to the alignment between the sequences of markers found in Chapter four and the whole genome of Brassica rapa. Firstly, Screened 13 Intron Polymorphism (IP) markers designed from Brassica rapa genome and 28 IP markers from Brassica juncea genome, one marker (A0703-2) showed polymorphism, but the genetic distance was larger than other markers. Secondly, a new marker technique based on genome sequencing named Specific Length Amplified Fragment (SLAF) was constructed, and first used on gene fine mapping. After screened 76 SLAF markers, 10 showed polymorphism. Four of them were position-known (S17-52、S1727-5、S1727-6、S1729-5), and six of them were position-unknown (SNONE-2、SNONE-7、SNONE-9,SNONE-16,SNONE-27和SNONE-44). The nearest position-known marker was S1727-6 (0.76 cM), and the nearest position-unknown marker was SNONE-9 (0.51 cM). Finally, 26 Indel markers on A07 were screened, four markers (S17-1、S17-2、S69-1 and S69-2) linked to BrMs1 gene. According the position of SLAF markers and Indel markers, the genetic distances of markers and gene corresponded to physical distances, and the predicted area in physical map was confirmed. By the sequence analysis in the predicted area, the position of BrMs1 was predicted in centriolar region.
5. Line 454AB was a RGMS line, which gave 320 individuals. SRAP and SSR techniques and bulked segregant analysis (BSA) were used to screen markers linked to the RGMS restoring gene (BrMsf3). Among the 1 128 SRAP primers, only BMe10/SA4 and M52/K2 showed polymorphism between bulks. The polymorphic bands were named as BMe10SA4 and M52K2. The distance of BMe10SA4 and M52K2 was 4.38 cM and 7.46 cM, respectively, and these two markers were on the same side of BrMsf3. SSR marker ENA6 was linked to BrMsf3, and the genetic distance was 11.57 cM. According to the position of ENA6 in genetic map, BrMsf3 was mapped on A07. Subsequently, 26 Indel markers and 65 SLAF markers from A07 were screened, and no one showed polymorphism. This result confirmed the multiple-allele genetic model for 451AB line and 454AB line.
6. According to the genetic analysis and results of gene mapping, 451AB line and 454AB line were multiple-allele male sterility lines, and the male sterility genes were on A07. 452AB line was also multiple-allele male sterility line, and the male sterility genes were on A08. That verified there were at least two multiple-allele male sterility systems in Chinese cabbage.
本研究对两个隐性细胞核雄性不育系(452AB和454AB)和一个显性细胞核雄性不育系(451AB)进行了遗传学研究,判断不育系的遗传模式,不育基因在自交系中的分布,并对三个不育系的育性恢复基因和不育基因进行了分子标记、基因定位。主要结果如下:
1.隐性雄性不育系452AB和454AB的可育株和不育株分别与显性雄性不育系451AB中的可育株杂交,并进一步回交。通过数据分析推断451AB系不育基因与454AB系恢复基因为等位基因,这两个系属于复等位遗传模式。452AB系与另外两个系不同,表现为独立遗传。三个不育系与17个自交系进行测交,判断自交系基因型。根据测交结果,4个自交系的可育基因对452AB系不育基因有保持作用。进一步研究判断452AB系也为一复等位雄性不育系。两复等位雄性不育系在基因组中的位置不同,并不存在关联。
2.隐性细胞核雄性不育系452AB包含164个单株。利用混合分组分离法(BSA)对不育系进行标记分析,并优化SRAP反应体系。在构建的基因池中筛选SRAP引物1 256对,SSR引物187对。获得与大白菜隐性细胞核雄性不育系恢复基因(BrMsf2)连锁的标记两个,PM8K4和Me2M49,与恢复基因的遗传距离分别为2.98 cM和10.92 cM。通过调查PM8K4标记在大白菜DH作图群体中的多态性,将该恢复基因定位在大白菜染色体A08。根据大白菜基因组测序信息,筛选来自染色体A08候选区域的Indel标记23个,其中scaffold20上的标记S20-1与恢复基因的遗传距离为4.22 cM。通过S20-1和PM8K4标记在染色体上的位置,将BrMsf2定位在scaffold20的附近。
3.显性细胞核雄性不育系451AB包含480个单株。利用混合分组分离法、SRAP和SSR标记技术筛选与不育基因BrMs1连锁的标记。在1 378对SRAP标记和238对SSR标记中,筛选到与不育基因连锁的SRAP标记5个(CuMe7BEm12, BMe9E32, PM8E38, E35BEm10, and M51K2),SSR标记1个(ENA6)。其中PM8E38标记与不育基因的距离最近,为0.37 cM。将该标记多态性片段进行克隆测序,成功的转化为较为简易的SCAR标记PM8E38S。通过SSR标记ENA6在DH作图群体中的位置,不育基因BrMs1被定位在大白菜染色体A07上。
4.利用2357个单株对451AB系不育基因进行精细定位。初步定位标记PM8E38S在精细定位群体中与不育基因的遗传距离为0.59cM。将初步定位标记的序列与基因组序列进行比对,没有比对在染色体A07上。筛选根据白菜基因组序列设计的内含子多态性(Intron Polymorphism,IP)标记13对,与芥菜中设计的IP标记28对,筛选到一个与不育基因距离较远的标记A0703-2。设计开发基于测序技术的酶切标签标记,并首次应用于雄性不育基因的精细定位中。筛选酶切标签标记76个,获得4个基因组位置已知的标记S17-52、S1727-5、S1727-6、S1729-5和6个基因组位置未知的标记SNONE-2、SNONE-7、SNONE-9,SNONE-16,SNONE-27和SNONE-44,其中SNONE-9与不育基因遗传距离最小(0.51 cM)。S1727-6是与不育基因距离最短(0.76 cM)的位置已知的标记。进一步对该区域Indel标记进行筛选,从26对引物中,筛选到4对存在多态性的Indel标记(S17-1、S17-2、S69-1和S69-2)。通过Indel标记以及酶切标签标记在基因组中的位置,将标记与基因之间的遗传距离与物理距离想对应。通过基因所在区域序列分析结果,预测该基因位于重复序列较多的中心粒区。
5. 454AB系为隐性细胞核雄性不育系,它包含320个单株。利用混合分组分离法根据育性构建育性池。筛选SRAP标记1 128个,SSR标记187个。筛选出与恢复基因BrMsf3连锁的SRAP标记两个,BMe10SA4和M52K2,与恢复基因的连锁距离为4.38 cM和7.46 cM,这两个标记位于基因的同侧; SSR标记ENA6与恢复基因的遗传连锁距离为11.57cM。通过ENA6标记在DH作图群体中的位置,将该恢复基因定位在大白菜染色体A07上。进一步筛选位于A07上的Indel标记26个,酶切标签标记65个,未发现与恢复基因(BrMsf3)连锁的标记。基因定位结果证实了451AB系和454AB系的复等位遗传关系。
6.通过遗传学分析和基因定位结果,得到451AB系和454AB系属于复等位遗传,不育基因位于染色体A07上,452AB系属于复等位遗传,不育基因位于染色体A08上,证明大白菜中存在一个以上复等位雄性不育系统。
Chinese cabbage (Brassica rapa L. ssp. pekinensis) is an important vegetable crop in China. It originated in China and has many advantageous economic characteristics. It has quite strong heterosis. Self-incompatible lines and male sterile lines are widely used in F1 seed production. The application of self-incompatible lines is limited by the reduction of fitness and vigor, the inconvenience of selfing, and the susceptibility of plants to environmental conditions. Male sterile lines are ideal systems for hybrid production. Plant male sterility can be generally classified into cytoplasmic male sterility (CMS) and genic male sterility (GMS). In GMS lines, sterility is stable and complete, and GMS does not result in reduced vigor. The study of genic male sterility of Brassica rapa showed different genetic models. The methods of transferring male sterility genes depend on certain genetic models. The genetic models of the genic male sterility lines had to be identified before they can be used for breeding. Molecular markers analysis of those male sterility genes just started in recent years.
This thesis deals with two recessive genic male sterility lines (452AB and 454AB) and one domiant genic male sterility line (451AB) to confirm their genetic models, to investigate distribution of different male sterile genes in male fertile inbred lines, and to find the molecular markers linked to the restoring genes and male sterility gene, and to map these genes. The main conclusions are as follows:
1. The genetic analysis in the recessive genic male sterility lines (452AB and 454AB) and dominant genic male sterility line (451AB) was conducted. The sterile plants and fertile plants in 452AB line and 454AB line were crossed with the fertile plants in 451AB line. According to the results, BrMs1 gene in 451AB and BrMsf3 gene in 454AB line were at the same allelic locus. So the genetic model of 451AB line and 454AB line was conformed to be a multiple-allele model. 452AB line had no relationship with 451AB line and 454AB line. For transferring these genes to other lines, male sterile plants were backcrossed with 17 inbred lines to confirm their genotypes. Four inbred lines can maintain sterility gene in 452AB line and resulted in 100% male sterile populations. The further experiments confirmed the genetic model of 452AB line also to be multiple-allele, but the genespositions were different from those in line 451AB and line 454AB.
2. Line 452AB was recessive genic male sterility (RGMS), which contained 164 individuals. Bulked segregant analysis (BSA) was used to screen markers linked to restoring gene BrMsf2. After SRAP (Sequence-related amplified polymorphism) markers system was optimized, 1 256 SRAP primer pairs and 187 simple sequence repeat (SSR) primer pairs were analyzed between fertile bulks. Two SRAP markers PM8K4 and Me2M49 were found, and the distance of PM8K4 and Me2M49 was 2.98 cM and 10.92 cM, respectively. PM8K4 was subsequently mapped on chromosome A08 using a doubled-haploid mapping population. Screened 23 Indel markers on A08 from Brassica rapa genome sequence, one marker (S20-1) in scaffold 20 showed polymorphism, and the genetic distance was 4.22 cM. According the positions of S20-1 and PM8K4 on chromosome A08, BrMsf2 was mapped around scaffold 20 in A08.
3. Line 451AB was dominant genic male sterility (DGMS), which produced 480 individuals. SRAP, SSR and sequence-characterized amplified region (SCAR) analyses were performed to screen markers linked to BrMs1. Five of the 1 378 SRAP primer pair combinations (CuMe7BEm12, BMe9E32, PM8E38, E35BEm10, and M51K2) and one of 238 SSRs (ENA6) revealed polymorphisms between bulks from male sterile and fertile progeny. PM8E38, the nearest marker linked to BrMs1 (0.37 cM), was converted to the SCAR marker PM8E38S. ENA6 was subsequently mapped on chromosome A07 using a doubled haploid mapping population.
4. A fine mapping population which contained 2 357 individuals was constructed in 451AB line. In this larger population, the distance between PM8E38S and BrMs1 was changed to 0.59 cM. None of them was mapped on A07 according to the alignment between the sequences of markers found in Chapter four and the whole genome of Brassica rapa. Firstly, Screened 13 Intron Polymorphism (IP) markers designed from Brassica rapa genome and 28 IP markers from Brassica juncea genome, one marker (A0703-2) showed polymorphism, but the genetic distance was larger than other markers. Secondly, a new marker technique based on genome sequencing named Specific Length Amplified Fragment (SLAF) was constructed, and first used on gene fine mapping. After screened 76 SLAF markers, 10 showed polymorphism. Four of them were position-known (S17-52、S1727-5、S1727-6、S1729-5), and six of them were position-unknown (SNONE-2、SNONE-7、SNONE-9,SNONE-16,SNONE-27和SNONE-44). The nearest position-known marker was S1727-6 (0.76 cM), and the nearest position-unknown marker was SNONE-9 (0.51 cM). Finally, 26 Indel markers on A07 were screened, four markers (S17-1、S17-2、S69-1 and S69-2) linked to BrMs1 gene. According the position of SLAF markers and Indel markers, the genetic distances of markers and gene corresponded to physical distances, and the predicted area in physical map was confirmed. By the sequence analysis in the predicted area, the position of BrMs1 was predicted in centriolar region.
5. Line 454AB was a RGMS line, which gave 320 individuals. SRAP and SSR techniques and bulked segregant analysis (BSA) were used to screen markers linked to the RGMS restoring gene (BrMsf3). Among the 1 128 SRAP primers, only BMe10/SA4 and M52/K2 showed polymorphism between bulks. The polymorphic bands were named as BMe10SA4 and M52K2. The distance of BMe10SA4 and M52K2 was 4.38 cM and 7.46 cM, respectively, and these two markers were on the same side of BrMsf3. SSR marker ENA6 was linked to BrMsf3, and the genetic distance was 11.57 cM. According to the position of ENA6 in genetic map, BrMsf3 was mapped on A07. Subsequently, 26 Indel markers and 65 SLAF markers from A07 were screened, and no one showed polymorphism. This result confirmed the multiple-allele genetic model for 451AB line and 454AB line.
6. According to the genetic analysis and results of gene mapping, 451AB line and 454AB line were multiple-allele male sterility lines, and the male sterility genes were on A07. 452AB line was also multiple-allele male sterility line, and the male sterility genes were on A08. That verified there were at least two multiple-allele male sterility systems in Chinese cabbage.
引文
1.鲍东兵,王舒华,佟景春,王立杰.大白菜细胞核雄性不育基因的等位性检测.辽宁师专学报, 1999, 1 (3): 91-95
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