向日葵染色体核型及其遗传转化的研究
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摘要
向日葵是世界上重要的油料作物和生物能源材料,在中国人民的饮食结构中是食用油的主要来源之一。由于向日葵是公认的难转化作物,其转化频率不高,并且重复性差,其主要原因是从转化的细胞或组织分化再生植株困难。因此,本研究主要内容之一是探讨向日葵离体培养再生难的问题,以促进向日葵植株再生技术日臻完善,不断提高植株再生频率。研究结果表明,向日葵不同基因型外植体在含适宜IAA、6-BA等激素的培养基中均较易形成愈伤组织;但愈伤组织分化不定芽的能力则较困难;不同基因型向日葵外植体的不定芽诱导率依次为子叶节>下胚轴>子叶>真叶;诱导子叶节不定芽分化的适宜6-BA浓度为1.2mg/L,下胚轴为1.8mg/L,子叶为0.6mg/L,真叶却未见到诱导出不定芽;幼胚大小为≤2mm时,体细胞胚发生频率较高(45.5%);体细胞胚胎发生的适宜培养基为MS+0.4mg/L 2,4-D+120g/L蔗糖。
核型分析是细胞遗传学、染色体工程、基因定位、细胞分类学以及现代进化理论等学科的基本研究方法,对向日葵染色体核型进行分析,这将为向日葵的细胞学分类、选择杂交亲本组合、培育新品种及转基因向日葵的细胞学检测等方面提供细胞学依据。结果表明,8个基因型向日葵的染色体数均为2n =2x = 34,除了RHA266之外,都为中部着丝粒或近中部着丝粒染色体;8个基因型向日葵具有2种核型:1B和2B,这表明不同基因型间染色体核型确实存在差异。
随着生物技术的发展,转基因植物产业将成为21世纪新的经济增长点。我们的设想是在21世纪里,向日葵的基因工程将从单基因性状向多基因发展。也就是说把抗虫、抗病、高产等多种基因都组合在一起,得到更好的向日葵品种。同时还想利用转基因工程,把有益健康的基因转移到向日葵中去,以改良向日葵的品质,使向日葵生产的食品成为健康食品、功能食品、营养食品。另外,通过向日葵转基因工程,人们还想尽快培育出抗旱、抗碱、抗土地贫瘠、抗重金属污染的向日葵。由此看来,对转基因向日葵的研究和开发其前景是非常远大的。关于对类胡萝卜素合成的有关酶基因转化向日葵的研究,无论是国外,还是国内至今有关这方面的研究报道较少。本实验通过农杆菌介导法和花粉管通道法,用类胡萝卜素生物合成代谢途径中关键酶PSY和LycB基因转化向日葵,以期获得类胡萝卜素含量较高的优质油用向日葵,达到改善向日葵品质的目的。研究结果表明:转化受体筛选剂Hyg使用浓度为8mg/L较适宜;共培养时间3d为宜;侵染时间8min为宜;重悬液浓度OD600以0.6为佳;以干燥处理4h的子叶节为外植体进行遗传转化,子叶节抗性芽率较不处理有明显的提高;柱头滴加法和子房注射法导入目的基因较适宜的时间为授粉后4-6h;柱头滴加法的抗性植株率低于子房注射法;PCR、Dot blot及Southern的分子检测证明,目的基因PSY和LycB已整合进向日葵基因组中。
Carotenoids,a kind of nutrition elements have various health benefits such as improving human's immunity, retarding the ageing process of organs, preventing cancer, reducing the risks of nyctalopia and infant amblyopia for lack of vitamin A.Human blood contains lycopene, xanthophyl,β-cryptoxanthin,α-carotene and other carotenoids although human body can not synthesize those, and take them from food. Fruit and vegetable, is the main source of carotenoids. Carotenoids are lipophilic soluble, which can’t be absorbed lack of lipid.β- carotene in albumen of“the golden rice”have gained a higher content, whereas, how much can be taken need a further research. Additionally, it is hard and poisonous to synthesize by chemical method, by which only a few kinds of carotenoids obtained and commercialized, such asβ- carotene, astaxanthin and canthaxanthin etc. In another way, we gotβ- carotene and astaxanthin by fermentation.
Karyotype analysis is a basic method for investigating the cell genetics, chromosome engineering, gene location, cytotaxonomy and modern evolution theory. It also provides cytology proof for cytotaxonomy breeding, selecting parents of hybrid Combination and cytology test of trans-genes sunflower. Carotenoids synthase genes were applied to transformation some plants such as tobacco, cole, tomato and rice. Although it is usually called as golden rice and golden cole for higher carotenoid content by genetic transformation, no related research and report about sunflower was found.
Sunflower, a kind of annual herbaceous plant belongs to composite family, which bears character of leanness, salt and drought resistance. Seed of Sunflower, the main source of protein and plant lipid, is one of the important biomass energy. Sunflower plays an important role in supplying lipid and protein and improving food structure as well as promoting development of animal husbandry and processing industry. Scientists aim to breed new breeds with high yield and good quality by modern biotechnology, whereas, sunflower is a generally acknowledged recalcitrant crop for transformation. So far no efficient and stable transformation system was set up because of frequency of plant regeneration.
Researchers have attached importance to utilizing tissue culture and gene engineering to meliorate quality of sunflower since the 1980s, but most research focus on improving the method of plant regeneration and transformation as well as on promoting transformation efficiency. No satisfying result was reported, except single or multi-genes transformed sunflower and very few characters were altered.
There will be significant economy and society benefit for health promotion and promoting the quality of sunflower through transformation of carotenoids synthase gene to enhance the carotenoids content.
Karyotype analysis of eight genotypes of sunflower showed that chromosome numbers in the eight sunflowers were 2n = 2x = 34. All the chromosomes were median and submedian centromeric. Their karyotype formulas were F53: 2n=2x=34=24m+10sm、2603: 2n=2x=34=24m+10sm、2736: 2n=2x=34=26m+8sm、89B1: 2n=2x=34=22m+12sm、HA300: 2n=2x=34=28m+6sm、B7: 2n=2x=34=22m+ 12sm、RHA266: 2n=2x=34=30m+2sm+2st、PR29: 2n=2x=34=30m+4sm;The eight genotypes of sunflowers shared two karyotypes (1B and 2B), which indicated there were some karyotypes diversity within different genotypes of sunflowers.
The regeneration frequency of induced bud with different explants in different culture medium was further researched by taking different genotypes sunflower as material. The result showed that, it is easy to form the callus that explants of different genotype sunflower in the culture medium which contain suitable endocrine such as IAA、6-BA and so on, but the possibility that callus forms bud is slim. The frequencies of induced bud which explants of different genotype sunflower are in turn as followed: cotyledenary node > hypocotyls> cotyledon> euphylla; the suitable concentration of 6-BA to induce the cotyledenary node to generate the bud is 1.2mg/l, hypocotyls is 1.8mg/l, cotyledon is 0.6mg/l, euphylla hasn’t been induced to form the bud. The capability of induced adventitious bud is very different with different genotypes of sunflower. The regenerated frequency of PR29 showed to be the highest.
The young embryos of different genotype sunflower were also used as material. The rule of somatic embryogenesis was studied preliminarily, including the size of young embryo, the concentration of sucrose, carbohydrate, endocrine (2,4-D and 6-BA). The results showed that somatic embryogenesis was induced on callus or between or marginal of two cotyledons, but the most be on one side of the cotyledon in line. The somatic embryogenesis frequency showed the highest as sucrose (120 g/L) added into the medium. When the size of young embryo is less than 2mm, the frequency of the embryo to embryogenesis was suitable (45.5%). The optimum culture medium for somatic embryogenesis is Ms+0.4mg/l+2,4-D+120 mg/l sucrose. Except that, the somatic embryogenesis frequency varied with genotypes, and RHA266 gained the highest (38.4%), on the contrary 2736 got the lowest(10.1%).
Cotyledonary nodes of three different genotypes of sunflowers (RHA266, PR29 and 2603) were transformed with Agrobracterium tumefaciens EHA101 carrying the PSY gene. Meanwhile the factors affecting transformation were studied. Transgenic sunflower’s plants were examined by PCR and Southern hybridiazation analysis. The results as follows: The Optimum concentration of Hyg is 8mg/L. The time of co-culture in 3 days might be suitable. Eight minutes is suitable for infection. The optimum range of OD600 is 0.6. PCR and PCR-Southern blot showed that the target LycB gene integrated into the genome of sunflower.
Carotenoid biosynthesis enzyme gene (LycB) was transformed to 3 genotypes of sunflowers via pollen tube pathway technique. The results showed that stigma method affecting seed operated in 2~4 hrs pollinated, and 4hr later, no effect was observed. Meanwhile, times of operation had slim effect on seed. Ovary injection method gave more effect on seed, which brought a very low seed ratio in 2hrs, after that gained height in another 2hrs, whereas, seed ratio got no strong promotion in 8hrs. 40 mg/L kanamycin is the best density to select the positive via pollen tube pathway technique. Additionally, different induction methods gained distinguished positive ratio, that is to say, ovary injection method gave higher setting rate than stigma. The LycB gene has integrated into genome of sunflower testified by the analysis of Southern blot.
核型分析是细胞遗传学、染色体工程、基因定位、细胞分类学以及现代进化理论等学科的基本研究方法,对向日葵染色体核型进行分析,这将为向日葵的细胞学分类、选择杂交亲本组合、培育新品种及转基因向日葵的细胞学检测等方面提供细胞学依据。结果表明,8个基因型向日葵的染色体数均为2n =2x = 34,除了RHA266之外,都为中部着丝粒或近中部着丝粒染色体;8个基因型向日葵具有2种核型:1B和2B,这表明不同基因型间染色体核型确实存在差异。
随着生物技术的发展,转基因植物产业将成为21世纪新的经济增长点。我们的设想是在21世纪里,向日葵的基因工程将从单基因性状向多基因发展。也就是说把抗虫、抗病、高产等多种基因都组合在一起,得到更好的向日葵品种。同时还想利用转基因工程,把有益健康的基因转移到向日葵中去,以改良向日葵的品质,使向日葵生产的食品成为健康食品、功能食品、营养食品。另外,通过向日葵转基因工程,人们还想尽快培育出抗旱、抗碱、抗土地贫瘠、抗重金属污染的向日葵。由此看来,对转基因向日葵的研究和开发其前景是非常远大的。关于对类胡萝卜素合成的有关酶基因转化向日葵的研究,无论是国外,还是国内至今有关这方面的研究报道较少。本实验通过农杆菌介导法和花粉管通道法,用类胡萝卜素生物合成代谢途径中关键酶PSY和LycB基因转化向日葵,以期获得类胡萝卜素含量较高的优质油用向日葵,达到改善向日葵品质的目的。研究结果表明:转化受体筛选剂Hyg使用浓度为8mg/L较适宜;共培养时间3d为宜;侵染时间8min为宜;重悬液浓度OD600以0.6为佳;以干燥处理4h的子叶节为外植体进行遗传转化,子叶节抗性芽率较不处理有明显的提高;柱头滴加法和子房注射法导入目的基因较适宜的时间为授粉后4-6h;柱头滴加法的抗性植株率低于子房注射法;PCR、Dot blot及Southern的分子检测证明,目的基因PSY和LycB已整合进向日葵基因组中。
Carotenoids,a kind of nutrition elements have various health benefits such as improving human's immunity, retarding the ageing process of organs, preventing cancer, reducing the risks of nyctalopia and infant amblyopia for lack of vitamin A.Human blood contains lycopene, xanthophyl,β-cryptoxanthin,α-carotene and other carotenoids although human body can not synthesize those, and take them from food. Fruit and vegetable, is the main source of carotenoids. Carotenoids are lipophilic soluble, which can’t be absorbed lack of lipid.β- carotene in albumen of“the golden rice”have gained a higher content, whereas, how much can be taken need a further research. Additionally, it is hard and poisonous to synthesize by chemical method, by which only a few kinds of carotenoids obtained and commercialized, such asβ- carotene, astaxanthin and canthaxanthin etc. In another way, we gotβ- carotene and astaxanthin by fermentation.
Karyotype analysis is a basic method for investigating the cell genetics, chromosome engineering, gene location, cytotaxonomy and modern evolution theory. It also provides cytology proof for cytotaxonomy breeding, selecting parents of hybrid Combination and cytology test of trans-genes sunflower. Carotenoids synthase genes were applied to transformation some plants such as tobacco, cole, tomato and rice. Although it is usually called as golden rice and golden cole for higher carotenoid content by genetic transformation, no related research and report about sunflower was found.
Sunflower, a kind of annual herbaceous plant belongs to composite family, which bears character of leanness, salt and drought resistance. Seed of Sunflower, the main source of protein and plant lipid, is one of the important biomass energy. Sunflower plays an important role in supplying lipid and protein and improving food structure as well as promoting development of animal husbandry and processing industry. Scientists aim to breed new breeds with high yield and good quality by modern biotechnology, whereas, sunflower is a generally acknowledged recalcitrant crop for transformation. So far no efficient and stable transformation system was set up because of frequency of plant regeneration.
Researchers have attached importance to utilizing tissue culture and gene engineering to meliorate quality of sunflower since the 1980s, but most research focus on improving the method of plant regeneration and transformation as well as on promoting transformation efficiency. No satisfying result was reported, except single or multi-genes transformed sunflower and very few characters were altered.
There will be significant economy and society benefit for health promotion and promoting the quality of sunflower through transformation of carotenoids synthase gene to enhance the carotenoids content.
Karyotype analysis of eight genotypes of sunflower showed that chromosome numbers in the eight sunflowers were 2n = 2x = 34. All the chromosomes were median and submedian centromeric. Their karyotype formulas were F53: 2n=2x=34=24m+10sm、2603: 2n=2x=34=24m+10sm、2736: 2n=2x=34=26m+8sm、89B1: 2n=2x=34=22m+12sm、HA300: 2n=2x=34=28m+6sm、B7: 2n=2x=34=22m+ 12sm、RHA266: 2n=2x=34=30m+2sm+2st、PR29: 2n=2x=34=30m+4sm;The eight genotypes of sunflowers shared two karyotypes (1B and 2B), which indicated there were some karyotypes diversity within different genotypes of sunflowers.
The regeneration frequency of induced bud with different explants in different culture medium was further researched by taking different genotypes sunflower as material. The result showed that, it is easy to form the callus that explants of different genotype sunflower in the culture medium which contain suitable endocrine such as IAA、6-BA and so on, but the possibility that callus forms bud is slim. The frequencies of induced bud which explants of different genotype sunflower are in turn as followed: cotyledenary node > hypocotyls> cotyledon> euphylla; the suitable concentration of 6-BA to induce the cotyledenary node to generate the bud is 1.2mg/l, hypocotyls is 1.8mg/l, cotyledon is 0.6mg/l, euphylla hasn’t been induced to form the bud. The capability of induced adventitious bud is very different with different genotypes of sunflower. The regenerated frequency of PR29 showed to be the highest.
The young embryos of different genotype sunflower were also used as material. The rule of somatic embryogenesis was studied preliminarily, including the size of young embryo, the concentration of sucrose, carbohydrate, endocrine (2,4-D and 6-BA). The results showed that somatic embryogenesis was induced on callus or between or marginal of two cotyledons, but the most be on one side of the cotyledon in line. The somatic embryogenesis frequency showed the highest as sucrose (120 g/L) added into the medium. When the size of young embryo is less than 2mm, the frequency of the embryo to embryogenesis was suitable (45.5%). The optimum culture medium for somatic embryogenesis is Ms+0.4mg/l+2,4-D+120 mg/l sucrose. Except that, the somatic embryogenesis frequency varied with genotypes, and RHA266 gained the highest (38.4%), on the contrary 2736 got the lowest(10.1%).
Cotyledonary nodes of three different genotypes of sunflowers (RHA266, PR29 and 2603) were transformed with Agrobracterium tumefaciens EHA101 carrying the PSY gene. Meanwhile the factors affecting transformation were studied. Transgenic sunflower’s plants were examined by PCR and Southern hybridiazation analysis. The results as follows: The Optimum concentration of Hyg is 8mg/L. The time of co-culture in 3 days might be suitable. Eight minutes is suitable for infection. The optimum range of OD600 is 0.6. PCR and PCR-Southern blot showed that the target LycB gene integrated into the genome of sunflower.
Carotenoid biosynthesis enzyme gene (LycB) was transformed to 3 genotypes of sunflowers via pollen tube pathway technique. The results showed that stigma method affecting seed operated in 2~4 hrs pollinated, and 4hr later, no effect was observed. Meanwhile, times of operation had slim effect on seed. Ovary injection method gave more effect on seed, which brought a very low seed ratio in 2hrs, after that gained height in another 2hrs, whereas, seed ratio got no strong promotion in 8hrs. 40 mg/L kanamycin is the best density to select the positive via pollen tube pathway technique. Additionally, different induction methods gained distinguished positive ratio, that is to say, ovary injection method gave higher setting rate than stigma. The LycB gene has integrated into genome of sunflower testified by the analysis of Southern blot.
引文
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