不同诱变方法对油菜种子诱变效果及突变体的研究
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摘要
诱变可以产生丰富的突变,诱变技术已广泛的应用于作物育种和生物学研究中。射线辐射,化学诱变,空间搭载等诱变方法也较早的应用于油菜育种中,而不同诱变方法的诱变效果存在差异。
本研究旨在利用诱变技术处理油菜种子,比较不同剂量的γ射线辐射、EMS和空间搭载诱变对甘蓝型油菜种子品质性状的诱变效果,探索利用诱变技术改良甘蓝型油菜种子品质的有效方法,期望获得品质优良的突变体。并对诱变后代中发现的矮秆突变体和苗期黄化突变体进行了遗传分析和基因定位。主要研究结果如下：
1.不同诱变方法对甘蓝型油菜种子品质性状的诱变效果比较
分别用800 Gy、1000 Gy和1200 Gy的Y射线,0.5%,1.0%和1.5%(v/v)的EMS溶液以及太空辐射处理三个甘蓝型油菜品系06-425、06-3741和06-7317。分析各种不同处理的M3代种子的品质,结果表明γ射线和EMS处理都能显著增加种子不同品质性状的变异,但诱变的效果存在差异。1.5%的EMS在黑籽甘蓝型油菜中能诱发较多的变异,更适合用于改良油菜种子品质。而对于黄籽品系则适合较低剂量的诱变剂处理。
2.平阳霉素对甘蓝型油菜小孢子出胚率的影响
利用不同浓度的平阳霉素处理5个甘蓝型油菜品系的游离小孢子,结果表明,0.1μg/ml的平阳霉素处理甘蓝型油菜小孢子30分钟能显著提高甘蓝型油菜小孢子出胚率,随着平阳霉素浓度的增加甘蓝型油菜小孢子出胚率也随之降低。但平阳霉素处理却降低了品系T8再生植株的加倍率,而对品系B409和T10再生植株的加倍率没有显著影响。利用AFLP技术分析了平阳霉素对品系T8、T10和B409双单倍体植株的诱变频率。结果表明,在甘蓝型油菜小孢子培养过程中本身就会产生0.095%到0.114%的变异,而平阳霉素处理没有显著增加其变异。
3.矮秆突变体的研究
在EMS诱变的M3株系中发现一个株高极度矮化的矮秆突变株系。遗传分析表明,其矮秆性状受一对隐性核基因控制,不存在细胞质效应。通过外源赤霉素处理和内源赤霉素含量的测定,结果表明外源赤霉素不能恢复矮秆突变体(bnaC.dwf)的株高,而且其叶片和茎干中的内源赤霉素含量也显著高于原始亲本。因此推断赤霉素传导途径基因的突变导致了突变体的矮化。利用AFLP标记结合群体集团分离分组法(BSA)共筛选到9个与BnaC.DWF基因紧密连锁的AFLP标记,并成功的将其中的四个AFLP标记转化为SCAR标记。并利用与BnaC.DWF基因连锁的SSR标记将BnaC.DWF基因定位于甘蓝型油菜C8连锁群。
4.苗期黄化突变体的研究
在EMS诱变的M2群体中发现了一个苗期黄化的突变体。遗传分析表明,其黄化性状受一对隐性核基因控制,不存在细胞质效应。利用AFLP标记结合群体集团分离分组法(BSA)共筛选到8个与BnaC. YGL基因紧密连锁的AFLP标记,并成功的将其中的四个AFLP标记转化为SCAR标记。并利用与BnaC. YGL基因连锁的SSR标记将BnaC. YGL基因定位于甘蓝型油菜C7连锁群。
Induced mutagenesis has become one of most important tool in plant biology and breeding for changing plant traits, studying gene function, and locating genes on chromosomes. The use of ionizing radiation, chemical mutagens and space environment for inducing variation is well established. However, the ionizing radiations, chemical mutagens and space environment have different efficiency in inducing mutations as they bring about both chromosome aberrations and gene mutations.
In order to find the best induced mutagenesis and dosage to improve seeds quality of Brassica napus, Comparing effectiveness of different mutagens (gamma rays, Ethyl methane sulphonate and space environment) for seed quality in Brassica napus were studied in three B. napus lines. Then, the variation analysis of seeds compositions were described in the progeny to prove the feasibility of improving rapeseed seeds quality by induced mutagenesis. At the same time, some high oleic acid, high protein, low linolenic acid mutants were expected to be screened. Furthermore, we have obtained one dwarf mutant and the other one chlorophyll-deficient seedling with yellow-green leaves mutant from the B. napus inbred line T6 through chemical mutagen ethylmethane sulfonate (EMS). Genetic analysis and mapping of these two mutants were carried out. The main results were as follows:
1. Comparing effectiveness of different mutagens (gamma rays, Ethyl methane sulphonate and space environment) for seed quality in B. napus
Three different B. napus lines were treated with y ray (800 Gy,1000 Gy and 1200 Gy), EMS (0.5%,1.0% and 1.5%) and space radiation respectively. Analysis of the seeds compositions of M3 seeds suggested that both y ray EMS can increase the variation of seeds composition. However, different variation was detected between different treated methods. For rapeseed seeds compositions breeding,1.5% EMS was more effective than other ways in the black seeds lines. At the same time, lower dose of y ray (800 Gy) and EMS (0.5%) was more preferable for yellow seed lines.
2. The effects of bleomycin on microspore embryogenesis in B. napus
The effects of bleomycin on microspore embryogenesis and cell division were investigated using three concentrations of bleomycin in five semi-winter genotypes of B. napus viz. T8, T10, B409, P30, and DH1142. Inclusion of bleomycin in the culture medium at a concentration of 0.1μg ml-1 for 30 min significantly improved embryo production and cell division in all five genotypes. Embryo production was induced at rates two-and four-fold higher than controls after bleomycin treatment. Fifty plants regenerated by microspore embryogenesis treated with bleomycin in addition to non-treated controls of T8, T10, and B409 were selected for AFLP analysis. The results suggest that microspore culture is capable of producing 0.095% to 0.114% genetic variation, and there was no effect of bleomycin treatment on genetic stabilisation of doubled haploid populations versus the non-treated control.
3. The research of one dwarf mutant
In the present study, we have obtained one dwarf mutant (bnaC.dwf) from the B. napus inbred line T6 through chemical mutagen ethyl methanesulfonate (EMS). We have determined the phenotypic effects and genetic characteristics of dwarf mutant(bnaC.dwf). Genetic analysis revealed that one recessive gene is responsible for controlling the phenotypic expression of dwarf mutant. The dwarf mutant was insensitive to exogenous GA3 for plant height, suggesting that it is significantly playing a crucial role in the gibberellins response pathway. Amplified Fragment Length Polymorphism (AFLP) technique was applied for selecting markers linked to the BnaC.DWF gene which assisted in screening of dwarf and normal individuals in the BC4 population. We have identified nine AFLP markers linked to the BnaC.DWF gene, and four AFLP markers were successfully converted into Sequence Characterised Amplified Region (SCAR) markers. The BnaC.DWF gene was further mapped to the linkage region C8 by Simple Sequence Repeat (SSR) markers.
4. The research of one chlorophyll-deficient seedling with yellow-green leaves mutant
In the present study, we have obtained one chlorophyll-deficient seedling with yellow-green leaves mutant (bnaC.ygl) from the B. napus inbred line T6 through chemical mutagen ethyl methanesulfonate (EMS). We have determined the phenotypic effects and genetic characteristics of chlorophyll-deficient seedling mutant (bnaC.ygl). Genetic analysis revealed that one recessive gene is responsible for controlling the phenotypic expression of dwarf mutant. Amplified Fragment Length Polymorphism (AFLP) technique was applied for selecting markers linked to the BnaC.DWF gene which assisted in screening of dwarf and normal individuals in the BC4 population. We have identified nine AFLP markers linked to the BnaC.DWF gene, and four AFLP markers were successfully converted into Sequence Characterised Amplified Region (SCAR) markers. The BnaC.DWF gene was further mapped to the linkage region C8 by Simple Sequence Repeat (SSR) markers.
本研究旨在利用诱变技术处理油菜种子,比较不同剂量的γ射线辐射、EMS和空间搭载诱变对甘蓝型油菜种子品质性状的诱变效果,探索利用诱变技术改良甘蓝型油菜种子品质的有效方法,期望获得品质优良的突变体。并对诱变后代中发现的矮秆突变体和苗期黄化突变体进行了遗传分析和基因定位。主要研究结果如下：
1.不同诱变方法对甘蓝型油菜种子品质性状的诱变效果比较
分别用800 Gy、1000 Gy和1200 Gy的Y射线,0.5%,1.0%和1.5%(v/v)的EMS溶液以及太空辐射处理三个甘蓝型油菜品系06-425、06-3741和06-7317。分析各种不同处理的M3代种子的品质,结果表明γ射线和EMS处理都能显著增加种子不同品质性状的变异,但诱变的效果存在差异。1.5%的EMS在黑籽甘蓝型油菜中能诱发较多的变异,更适合用于改良油菜种子品质。而对于黄籽品系则适合较低剂量的诱变剂处理。
2.平阳霉素对甘蓝型油菜小孢子出胚率的影响
利用不同浓度的平阳霉素处理5个甘蓝型油菜品系的游离小孢子,结果表明,0.1μg/ml的平阳霉素处理甘蓝型油菜小孢子30分钟能显著提高甘蓝型油菜小孢子出胚率,随着平阳霉素浓度的增加甘蓝型油菜小孢子出胚率也随之降低。但平阳霉素处理却降低了品系T8再生植株的加倍率,而对品系B409和T10再生植株的加倍率没有显著影响。利用AFLP技术分析了平阳霉素对品系T8、T10和B409双单倍体植株的诱变频率。结果表明,在甘蓝型油菜小孢子培养过程中本身就会产生0.095%到0.114%的变异,而平阳霉素处理没有显著增加其变异。
3.矮秆突变体的研究
在EMS诱变的M3株系中发现一个株高极度矮化的矮秆突变株系。遗传分析表明,其矮秆性状受一对隐性核基因控制,不存在细胞质效应。通过外源赤霉素处理和内源赤霉素含量的测定,结果表明外源赤霉素不能恢复矮秆突变体(bnaC.dwf)的株高,而且其叶片和茎干中的内源赤霉素含量也显著高于原始亲本。因此推断赤霉素传导途径基因的突变导致了突变体的矮化。利用AFLP标记结合群体集团分离分组法(BSA)共筛选到9个与BnaC.DWF基因紧密连锁的AFLP标记,并成功的将其中的四个AFLP标记转化为SCAR标记。并利用与BnaC.DWF基因连锁的SSR标记将BnaC.DWF基因定位于甘蓝型油菜C8连锁群。
4.苗期黄化突变体的研究
在EMS诱变的M2群体中发现了一个苗期黄化的突变体。遗传分析表明,其黄化性状受一对隐性核基因控制,不存在细胞质效应。利用AFLP标记结合群体集团分离分组法(BSA)共筛选到8个与BnaC. YGL基因紧密连锁的AFLP标记,并成功的将其中的四个AFLP标记转化为SCAR标记。并利用与BnaC. YGL基因连锁的SSR标记将BnaC. YGL基因定位于甘蓝型油菜C7连锁群。
Induced mutagenesis has become one of most important tool in plant biology and breeding for changing plant traits, studying gene function, and locating genes on chromosomes. The use of ionizing radiation, chemical mutagens and space environment for inducing variation is well established. However, the ionizing radiations, chemical mutagens and space environment have different efficiency in inducing mutations as they bring about both chromosome aberrations and gene mutations.
In order to find the best induced mutagenesis and dosage to improve seeds quality of Brassica napus, Comparing effectiveness of different mutagens (gamma rays, Ethyl methane sulphonate and space environment) for seed quality in Brassica napus were studied in three B. napus lines. Then, the variation analysis of seeds compositions were described in the progeny to prove the feasibility of improving rapeseed seeds quality by induced mutagenesis. At the same time, some high oleic acid, high protein, low linolenic acid mutants were expected to be screened. Furthermore, we have obtained one dwarf mutant and the other one chlorophyll-deficient seedling with yellow-green leaves mutant from the B. napus inbred line T6 through chemical mutagen ethylmethane sulfonate (EMS). Genetic analysis and mapping of these two mutants were carried out. The main results were as follows:
1. Comparing effectiveness of different mutagens (gamma rays, Ethyl methane sulphonate and space environment) for seed quality in B. napus
Three different B. napus lines were treated with y ray (800 Gy,1000 Gy and 1200 Gy), EMS (0.5%,1.0% and 1.5%) and space radiation respectively. Analysis of the seeds compositions of M3 seeds suggested that both y ray EMS can increase the variation of seeds composition. However, different variation was detected between different treated methods. For rapeseed seeds compositions breeding,1.5% EMS was more effective than other ways in the black seeds lines. At the same time, lower dose of y ray (800 Gy) and EMS (0.5%) was more preferable for yellow seed lines.
2. The effects of bleomycin on microspore embryogenesis in B. napus
The effects of bleomycin on microspore embryogenesis and cell division were investigated using three concentrations of bleomycin in five semi-winter genotypes of B. napus viz. T8, T10, B409, P30, and DH1142. Inclusion of bleomycin in the culture medium at a concentration of 0.1μg ml-1 for 30 min significantly improved embryo production and cell division in all five genotypes. Embryo production was induced at rates two-and four-fold higher than controls after bleomycin treatment. Fifty plants regenerated by microspore embryogenesis treated with bleomycin in addition to non-treated controls of T8, T10, and B409 were selected for AFLP analysis. The results suggest that microspore culture is capable of producing 0.095% to 0.114% genetic variation, and there was no effect of bleomycin treatment on genetic stabilisation of doubled haploid populations versus the non-treated control.
3. The research of one dwarf mutant
In the present study, we have obtained one dwarf mutant (bnaC.dwf) from the B. napus inbred line T6 through chemical mutagen ethyl methanesulfonate (EMS). We have determined the phenotypic effects and genetic characteristics of dwarf mutant(bnaC.dwf). Genetic analysis revealed that one recessive gene is responsible for controlling the phenotypic expression of dwarf mutant. The dwarf mutant was insensitive to exogenous GA3 for plant height, suggesting that it is significantly playing a crucial role in the gibberellins response pathway. Amplified Fragment Length Polymorphism (AFLP) technique was applied for selecting markers linked to the BnaC.DWF gene which assisted in screening of dwarf and normal individuals in the BC4 population. We have identified nine AFLP markers linked to the BnaC.DWF gene, and four AFLP markers were successfully converted into Sequence Characterised Amplified Region (SCAR) markers. The BnaC.DWF gene was further mapped to the linkage region C8 by Simple Sequence Repeat (SSR) markers.
4. The research of one chlorophyll-deficient seedling with yellow-green leaves mutant
In the present study, we have obtained one chlorophyll-deficient seedling with yellow-green leaves mutant (bnaC.ygl) from the B. napus inbred line T6 through chemical mutagen ethyl methanesulfonate (EMS). We have determined the phenotypic effects and genetic characteristics of chlorophyll-deficient seedling mutant (bnaC.ygl). Genetic analysis revealed that one recessive gene is responsible for controlling the phenotypic expression of dwarf mutant. Amplified Fragment Length Polymorphism (AFLP) technique was applied for selecting markers linked to the BnaC.DWF gene which assisted in screening of dwarf and normal individuals in the BC4 population. We have identified nine AFLP markers linked to the BnaC.DWF gene, and four AFLP markers were successfully converted into Sequence Characterised Amplified Region (SCAR) markers. The BnaC.DWF gene was further mapped to the linkage region C8 by Simple Sequence Repeat (SSR) markers.
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