日本结缕草IPT基因遗传转化及野牛草实生群体内遗传多样性研究
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摘要
日本结缕草和野牛草都是适应我国国情的优质暖季性草坪草。在进一步发展其优异的抗逆性同时,创造出具草坪优良性状特别是绿色期延长的新遗传种质是重要的育种目标。本研究建立了愈伤组织长期继代并具有高频再生能力的培养体系,分析了长期继代再生植株群体内存在的变异；并进一步利用该再生体系,以农杆菌介导的方法将IPT基因转入日本结缕草。此外,本研究针对草坪草容易变异的特征,对野牛草实生群体进行了形态与遗传物质多样性研究,结果如下：
1、日本结缕草IPT基因的遗传转化：
(1)以成熟胚为外植体诱导的愈伤组织长期继代高频再生体系的建立。诱导的胚性愈伤组织继代间隔时间为30d,在增加0.5×MS基本培养基Fe盐的继代培养基上,继代1-2次可以使愈伤组织的颗粒性增强。长期继代的愈伤组织分化率降低到0%,但在继代培养基中添加14mg/L CuS04预培养后,分化率可以提高到97%。继代30个月的愈伤组织切片观察,与新鲜愈伤组织相似,体胚发生和器官发生再生途径同时存在。
(2)长期培养愈伤组织再生植株的体细胞变异。以一粒种子诱导的愈伤组织增殖并长期继代再生的日本结缕草植株为材料研究发现,其在长期继代培养中容易发生变异。形态指标中匍匐茎数量和长度以及绿色期变异较大,有利于优异种质的筛选；利用ISSR和RAPD分子标记检测继代培养再生植株的遗传多样性,多态性位点数分别为83.6%和72.4%,样品特异性位点比例分别为15.1%、3.4%,显示组织培养中体细胞变异比率较高。两种分子标记遗传距离矩阵相关性分析,发现两者没有显著相关性r=-0.2397(P≦0.05),表明两种不同标记都能检测出再生群体的遗传多样性,但ISSR能扩增出更多的特异性带,在本实验中对日本结缕草的体细胞变异分析,ISSR比RAPD更合适。
(3)农杆菌介导的IPT基因的遗传转化。通过农杆菌介导法,成功地将特异抑制衰老的自我调控嵌合基因P SAGl2::IPT导入到日本结缕草的胚性愈伤组织,经筛选得到45株抗性植株,经田间鉴定转基因株系的枯黄期较野生型推后了15-71d,绿色期有了极大的改善,证明通过P SAGl2::IPT表达的调控增加结缕草体内的细胞分裂素含量以延长其绿色期是确实可行。PCR检测及Southern杂交,证实IPT已整合到结缕草的基因组中且得到表达,PCR检测阳性率达到51.1%。
2、野牛草实生群体多样性研究：
从102株实生群体中通过目测形态筛选出20株有明显差异的单株,通过表型性状调查和ISSR分子标记两种方法对野牛草实生群体的多样性进行分析,筛选出7条适合野牛草的ISSR引物,共扩增出89条带,多态性条带74条(83.14%)。单株间相似系数在0.58～0.82之间,遗传距离分布于0.17—0.42之间。表明野牛草实生群体内的遗传多样性较高,说明实生苗群体内可以采用单株选择法选育优良无性系。ISSR标记聚类结果与形态特征没有显著的相关关系,显示野牛草杂交亲本选择时不仅要考虑表型的差异,也要参考分子标记的差异。
Zoysia japonica and Buchloe dactyloides, as warm-season turfgrasses, were cultivated extensively since their good stress resistance. It is an important breeding goal to create new germplasms with excellent characters, especially with a longer blue-green period. In our present study, the variation of regenerated plants from embryogenic callus with long-term subculture history was studied based on the establishment of the long-term subculture and high efficient regeneration system, and the IPT gene was transformed into Z japonica. In addition, considering the easy mutation of turfgrass, the genetic diversity within the seedlings of B. dactyloides was also investigated. The main research results showed as follows:
1 Genetic transformation of IPT into Z. japonica
1) High efficient plant regeneration system for long-term subcultured calli derived from the mature embryos was established. The appropriate subculture interval was 30 d, and the subculture medium was supplemented with 0.5×Fe2+ of MS medium, which was used to improve the granule state of embryogenic callus after cultured for 30-60 d. The differentiation of long-term cultured callus was decreased to 0%. But when pre-cultured with 14 mg/L CuSO4, the frequency of differentiation was increased to 97%. Somatic embryogenesis and organogenesis were both observed on callus cultured for 30 month by histological analysis, which was the same as the fresh callus.
2) Genetic variation assessment of long-term subcultured regenerants. Variation was detected in the plants regenerated form the calli after long-term subculture. The number and length of stolons and blue-green period of individuals were varied significantly in the same grow season, which suggested selection of elite clones from those variations was possible. Based on ISSR and RAPD data,82.19% or 72.41% polymorphic loci was produced, respectively. The results suggested that the somatic variation during tissue culture was high in Z. japonica. The related coefficient of two markers genetic distance matrix was r=0.2397 (P≤0.05) which suggested there was not significant relation between the two markers. ISSR markers could amplify more specific loci, so ISSR was more preferable marker for detection of somatic variation in Z. japonica in this study.
3) Genetic transformation of Agrobacterium-mediated IPT gene. Psac 12::IPT was transferred into embryogenic callus of Z. japonica.45 resistant plants were obtained. PCR detection and Southern blotting confirmed the IPT gene was integrated into the genome. The positive rate of PCR was 51.1%. The observed phenotypic data showed the transgenic individuals turned yellow 15-71 d later than untransformed control plants in the field, which demonstrated that prolonging the green period was feasible via the transformation of PSAG12::IPT.
2 Genetic diversity within the seedlings of B. dactyloides
Analysis of genetic diversity within the population of buffalograss was carried out by morphological traits and ISSR markers.20 accessions within buffalograss population were selected and 74 (83.1%) polymorphic loci were detected out of 89 loci in total, which was derived from the ISSR-PCR analysis by seven ISSR primers. It showed that the genetic polymorphism in the population was high, the coefficient among each accessions ranged from 0.68 to 0.82, and genetic distance ranged from 0.17 to 0.42. The results indicated that individual selection was suitable in buffalograss breeding even within seedling population. Dendrogram obtained based on ISSR markers was not in accordance with that on morphologic characteristics, which suggested that not only morphological traits but also molecular genetic basis should be taken into account in selecting parents for bufflograss crossing.
1、日本结缕草IPT基因的遗传转化：
(1)以成熟胚为外植体诱导的愈伤组织长期继代高频再生体系的建立。诱导的胚性愈伤组织继代间隔时间为30d,在增加0.5×MS基本培养基Fe盐的继代培养基上,继代1-2次可以使愈伤组织的颗粒性增强。长期继代的愈伤组织分化率降低到0%,但在继代培养基中添加14mg/L CuS04预培养后,分化率可以提高到97%。继代30个月的愈伤组织切片观察,与新鲜愈伤组织相似,体胚发生和器官发生再生途径同时存在。
(2)长期培养愈伤组织再生植株的体细胞变异。以一粒种子诱导的愈伤组织增殖并长期继代再生的日本结缕草植株为材料研究发现,其在长期继代培养中容易发生变异。形态指标中匍匐茎数量和长度以及绿色期变异较大,有利于优异种质的筛选；利用ISSR和RAPD分子标记检测继代培养再生植株的遗传多样性,多态性位点数分别为83.6%和72.4%,样品特异性位点比例分别为15.1%、3.4%,显示组织培养中体细胞变异比率较高。两种分子标记遗传距离矩阵相关性分析,发现两者没有显著相关性r=-0.2397(P≦0.05),表明两种不同标记都能检测出再生群体的遗传多样性,但ISSR能扩增出更多的特异性带,在本实验中对日本结缕草的体细胞变异分析,ISSR比RAPD更合适。
(3)农杆菌介导的IPT基因的遗传转化。通过农杆菌介导法,成功地将特异抑制衰老的自我调控嵌合基因P SAGl2::IPT导入到日本结缕草的胚性愈伤组织,经筛选得到45株抗性植株,经田间鉴定转基因株系的枯黄期较野生型推后了15-71d,绿色期有了极大的改善,证明通过P SAGl2::IPT表达的调控增加结缕草体内的细胞分裂素含量以延长其绿色期是确实可行。PCR检测及Southern杂交,证实IPT已整合到结缕草的基因组中且得到表达,PCR检测阳性率达到51.1%。
2、野牛草实生群体多样性研究：
从102株实生群体中通过目测形态筛选出20株有明显差异的单株,通过表型性状调查和ISSR分子标记两种方法对野牛草实生群体的多样性进行分析,筛选出7条适合野牛草的ISSR引物,共扩增出89条带,多态性条带74条(83.14%)。单株间相似系数在0.58～0.82之间,遗传距离分布于0.17—0.42之间。表明野牛草实生群体内的遗传多样性较高,说明实生苗群体内可以采用单株选择法选育优良无性系。ISSR标记聚类结果与形态特征没有显著的相关关系,显示野牛草杂交亲本选择时不仅要考虑表型的差异,也要参考分子标记的差异。
Zoysia japonica and Buchloe dactyloides, as warm-season turfgrasses, were cultivated extensively since their good stress resistance. It is an important breeding goal to create new germplasms with excellent characters, especially with a longer blue-green period. In our present study, the variation of regenerated plants from embryogenic callus with long-term subculture history was studied based on the establishment of the long-term subculture and high efficient regeneration system, and the IPT gene was transformed into Z japonica. In addition, considering the easy mutation of turfgrass, the genetic diversity within the seedlings of B. dactyloides was also investigated. The main research results showed as follows:
1 Genetic transformation of IPT into Z. japonica
1) High efficient plant regeneration system for long-term subcultured calli derived from the mature embryos was established. The appropriate subculture interval was 30 d, and the subculture medium was supplemented with 0.5×Fe2+ of MS medium, which was used to improve the granule state of embryogenic callus after cultured for 30-60 d. The differentiation of long-term cultured callus was decreased to 0%. But when pre-cultured with 14 mg/L CuSO4, the frequency of differentiation was increased to 97%. Somatic embryogenesis and organogenesis were both observed on callus cultured for 30 month by histological analysis, which was the same as the fresh callus.
2) Genetic variation assessment of long-term subcultured regenerants. Variation was detected in the plants regenerated form the calli after long-term subculture. The number and length of stolons and blue-green period of individuals were varied significantly in the same grow season, which suggested selection of elite clones from those variations was possible. Based on ISSR and RAPD data,82.19% or 72.41% polymorphic loci was produced, respectively. The results suggested that the somatic variation during tissue culture was high in Z. japonica. The related coefficient of two markers genetic distance matrix was r=0.2397 (P≤0.05) which suggested there was not significant relation between the two markers. ISSR markers could amplify more specific loci, so ISSR was more preferable marker for detection of somatic variation in Z. japonica in this study.
3) Genetic transformation of Agrobacterium-mediated IPT gene. Psac 12::IPT was transferred into embryogenic callus of Z. japonica.45 resistant plants were obtained. PCR detection and Southern blotting confirmed the IPT gene was integrated into the genome. The positive rate of PCR was 51.1%. The observed phenotypic data showed the transgenic individuals turned yellow 15-71 d later than untransformed control plants in the field, which demonstrated that prolonging the green period was feasible via the transformation of PSAG12::IPT.
2 Genetic diversity within the seedlings of B. dactyloides
Analysis of genetic diversity within the population of buffalograss was carried out by morphological traits and ISSR markers.20 accessions within buffalograss population were selected and 74 (83.1%) polymorphic loci were detected out of 89 loci in total, which was derived from the ISSR-PCR analysis by seven ISSR primers. It showed that the genetic polymorphism in the population was high, the coefficient among each accessions ranged from 0.68 to 0.82, and genetic distance ranged from 0.17 to 0.42. The results indicated that individual selection was suitable in buffalograss breeding even within seedling population. Dendrogram obtained based on ISSR markers was not in accordance with that on morphologic characteristics, which suggested that not only morphological traits but also molecular genetic basis should be taken into account in selecting parents for bufflograss crossing.
引文
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