用DefH9-iaaM和TERF1基因转化糖橙的研究
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摘要
柑橘是世界第一大水果,2006年,世界柑橘年产量1.1亿吨,面积10730万亩,年贸易额达65亿美元,堪称世界第一大水果。我国是柑橘的重要原产地之一,柑橘资源丰富,优良品种繁多,栽培历史悠久。
2006年我国柑橘面积已超过170万公顷,占世界柑橘总面积的23%,居世界第一位,产量近1800万吨,居世界第二位。柑橘已经成为我国南方经济的支柱产业。但是我国柑橘产业虽有一定的基础,但与世界柑橘生产先进国家相比,差距还较大。特别是在品种选育方面还需进一步加强。柑橘是木本植物,存在童期长、珠心胚干扰、高度杂合、遗传背景复杂等缺点,导致常规育种的方法不仅存在很大的盲目性,而且育种效率低。随着生物技术的发展,与传统育种方法相比,利用基因工程培育新品种不受基因型限制,目的基因来源广泛,可以定向地改变目标性状等优点,采用成年态材料作为转化受体,还可以大大缩短育种周期,研究外源基因对植物开花和结果的影响。
糖橙由于纯甜似蜜,深受消费者的喜爱。但糖橙种籽较多,目前还没有无核的糖橙品种,同时细菌病害和真菌病害危害较多,增强抗病性也是主要育种目标之一。因此本项研究的目的是建立糖橙的高效再生转化体系和成年态再生转化体系,在此基础上把defH9-iaaM基因,即来自金鱼草的iaaM基因和来自根癌农杆菌的DefH9启动子导入柑橘中,以期望获得具有单性结实能力的新种质。其次,将通过酵母单杂交从番茄中克隆的转录因子TERF1导入糖橙中,期望能够获得对柑橘主要病害如溃疡病、炭疽病有广谱抗性的新种质。通过对转基因的种质进行转基因表达、形态学观测和抗病性等方面的研究,综合评价转基因育种对品种改良方面的作用。
本试验获得主要研究结果如下:
(1)建立糖橙实生态的再生培养体系。以30d苗龄的糖橙实生态节间茎段不定芽分化率较高,暗培养可以显著的提高不定芽的再生率。筛选出适宜实生态节间茎段再生的不定芽诱导的培养基:MS+5mg/L 6-BA;不定芽增殖培养基:MS+1.0 mg/L 6-BA+NAA 1.0 mg/L;生根培养基:MS+0.5 mg/L NAA。其中采用两步法诱导不定芽再生和生根效果较好。
(2)建立了成年态糖橙的不定芽再生体系。用改良的消毒方法G,即采枝梢前一周,每隔一天下午,于温室中将要采取的枝梢用10%的次氯酸钠溶液擦拭一遍,共擦拭4次,且供试植株须隔离放置。再采用70%的酒精处理30s,再用1%的升汞溶液加几滴吐温浸泡24min,无菌水冲洗4次进行消毒。可显著的提高成年态节间茎段的成活率。成年态节间茎段培养基以6-BA 5 mg/L+2,4-D 2 mg/L+IAA 2mg/L培养的效果最好,茎段愈伤诱导率和不定芽诱导率最高可达17.8%和2.87个。对成年态不定芽的嫁接进行了研究,认为经过改良的“⊥”压法成活率较高,可以达到78.94%。
(3)建立了将DefH9-iaaM基因和TERF1基因两个外源基因导入到实生态糖橙的转化体系。研究了不同根癌农杆菌对糖橙实生态节间茎段的转化率,结果显示EHA105的转化能力强于LBA4404。浸染时间以15min为宜。针对糖橙实生态节间茎段和成年态节间茎段再生能力的差异性,认为卡那霉素浓度为100mg/L时适合对糖橙实生态节间茎段再生不定芽进行筛选,卡那霉素浓度为30mg/L时适合对柑橘成年态节间茎段再生不定芽进行筛选,通过PCR鉴定,不定芽的转化率高达29.4%。
(4)通过PCR鉴定,结果显示以获得了6株转DefH9-iaaM基因植株,通过形态学的研究证实,这6株转基因植株在株高、节间长度、叶面积方面比对照有明显的增长、增宽现象。利用PCR、RT-PCR、Southern杂交技术对转TERF1基因的糖橙进行了分子检测,结果显示,共获得了7株转TERF1基因植株株系。通过对3,7,11号植株的Southern杂交,结果显示TERF1基因已经成功的整合到了糖橙的基因组中,且均为为单拷贝插入。从植株形态特征表现上看,转TERF1基因的植株与未转化植株无显著差异。
(5)对转TERF1基因的糖橙进行离体溃疡病和炭疽病的接种试验。通过对病斑大小和数量的分析,结果表明1,3,7,8,11号株系对柑橘溃疡病和炭疽病表现出广谱抗性。接种溃疡病病害时,3号株系对溃疡病的抗病性最强,发病级别为0级。其余转基因植株溃疡病的发病率比对照植株下降了70%左右。进行炭疽病的离体接种7d后发现,转基因植株的叶片基本上都表现出针孔处病斑减小。从照片上可以看到3号和7号转TERF1基因糖橙叶片不仅基本上观察不到炭疽病的典型病斑,而且叶片仍然可以保持翠绿。
Citrus is the most important fruit crop in the world.Now its annual production in the world is 110 million metric tons,with a total acreage of 7.15 million hectares (ha) and an annual volume of international trade about 650 million dollars.China is recognized as one of the key original centers of citrus,with a long history of cultivation,plenty of natural resources and selected varieties for commercial use.
In 2006,the planting area of citrus in China exceeded 1.7 million ha,which was 23%of the total acreage and ranked the first place in the world.The annual yield of citrus was nearly 18 million ton accepting the second biggest country for citrus production in the world.China has a long history of citrus breeding and has established the foundation of citrus industry,and the distance on technology application is widely accepted compared with those advanced countries of citrus production.In order to catch up with the international trade standard,the most important thing is to produce fruit of high quality and to satisfy the consumers.The criteria for evaluating the citrus cultivars are summarized as the following five main aspects:the appearance,taste,nutritive content,seeds number and easy-peeling by hand,et al.From the view of botany,the citrus is as a woody,low branching plant with a distinct juvenile stage,polyembryony,high genetic heterozygosity and arthenogenesis,and these characteristics have impeded the traditional breeding in citrus,such as the blindness in crossing and low efficiency for new cultivar development.With the development of biological technology,using transgenic technology to breed new variety is the best way to modify the genetic expressing or silencing.Transferring the foreign genes into mature explants will save us a lot of breeding time,and the gene expression in flower and fruit can be investigated much earlier than traditional did.
Succari is a very popular sweet orange,almost acidless,and is extending quickly in the south of China,but with plenty of seeds.Liking other sweet oranges, Succari trees can be attacked by many biotic(bacteria,fungi,etc.) and abiotic stresses. The objective of this project is to establish the regeneration and transformation system of Succari orange and to transfer a parthenocarpy gene 'defH9-iaaM' and a transcription factor 'TERFI' gene into the genome by using seedlings and adult inter-nodal segments as explants.Hopefully to gain a new germplasm or breeding material,which carrying parthenocarpy gene and got a broad-spectrum resistance to diseases.The DefH9-iaaM gene consists of an iaaM gene and a DefH9 promotor.The transcription factors TERF1 gene as cloned from tomato by yeast-one-hybrid system. After the disease inoculation and morphological trait observation,we evaluated the modification capability of the wild plants by foreign gene.
The main results are showed as follows:
1.Establishing the regeneration system from Succari orange seedling sterms intemodal segments.The results indicated that 30-day-old seedlings were the most efficient explants,and the co-culture in darkness for 6 days gave much improvement for good regeneration.The optimum medium for adventitious buds regeneration was MS + 6-BA(5.0mg/L).And MS medium supplied with 6-BA1.0mg/L and NAA 0.1mg/L showed the highest bud multiplication ratio.The shoot tissues got rooted successfully after 20 days in 0.5mg/L NAA.The better way to get more adventitious buds is using the "two-step" method,which is to put the explants from the medium plus the auxin first,and then remove to medium with no hormones after 6 days.
2.To set up an efficient regeneration system for mature sterms internodal segments.It shows that pretreatment disinfection method can raise significantly the survival rate to 80%.And the medium supplied with 6-BA 5 mg/L,2,4-D 2 mg/L and IAA 2mg/L could staminate the callus from cutting ends.And the callus induction rate reached up to 17.8%and 2.87 for the adventitious buds.The effect of three grafting methods on improving survival rate of adventitious buds was investigated,and the result showed that modified "⊥" way was the best one,which 78.94%of survival rate.
3.Development of the transformation system and the transfer of DefH9-iaaM gene and TERF1 gene into Succari.The effects of transformed by different Agrobacterium strains was observed,and it seemed that EHA105 was better than LBA4404.And 15 min was the best infection time.The selection concentration of km is different between the seedlings and mature explants,and 100 mg/L for seedling explants and 30 mg/L for mature explants were justified.
4.PCR results indicated that six regenerate plants of Succari were transgenic with the DefH9-iaaM gene,and the six transgenic plants showed evident increase in plant height,stem length,and leaf area.Through the PCR and RT-PCR analyses,it was showed that 7 regenerants were cooperated with the TERF1 gene.No.3,No.7 and No. 11 transgenic lines were verified by Southern blotting,and the results showed that one copy of the TERF1 gene were insert the plant genome.
5.After inoculation with the citrus canker and anthracnose pathogen the transgenic plants of TERF1 gene,the results showed that the TERF1 gene can improve the resistance to both diseases.The transgenic plants could reduce citrus bacterial canker incidence rate,by 70%.The No.3 showed 0%of the incidence rate.When the transgenic leaves and control ones were both infected with the citrus anthracnose,all the leaves showed blots symptom of disease.The blots on transgenic leaves were smaller than that on the controls.Leaves of No.3 and No.7 plants did not have the disease blots,and the color of leaves also kept green.
2006年我国柑橘面积已超过170万公顷,占世界柑橘总面积的23%,居世界第一位,产量近1800万吨,居世界第二位。柑橘已经成为我国南方经济的支柱产业。但是我国柑橘产业虽有一定的基础,但与世界柑橘生产先进国家相比,差距还较大。特别是在品种选育方面还需进一步加强。柑橘是木本植物,存在童期长、珠心胚干扰、高度杂合、遗传背景复杂等缺点,导致常规育种的方法不仅存在很大的盲目性,而且育种效率低。随着生物技术的发展,与传统育种方法相比,利用基因工程培育新品种不受基因型限制,目的基因来源广泛,可以定向地改变目标性状等优点,采用成年态材料作为转化受体,还可以大大缩短育种周期,研究外源基因对植物开花和结果的影响。
糖橙由于纯甜似蜜,深受消费者的喜爱。但糖橙种籽较多,目前还没有无核的糖橙品种,同时细菌病害和真菌病害危害较多,增强抗病性也是主要育种目标之一。因此本项研究的目的是建立糖橙的高效再生转化体系和成年态再生转化体系,在此基础上把defH9-iaaM基因,即来自金鱼草的iaaM基因和来自根癌农杆菌的DefH9启动子导入柑橘中,以期望获得具有单性结实能力的新种质。其次,将通过酵母单杂交从番茄中克隆的转录因子TERF1导入糖橙中,期望能够获得对柑橘主要病害如溃疡病、炭疽病有广谱抗性的新种质。通过对转基因的种质进行转基因表达、形态学观测和抗病性等方面的研究,综合评价转基因育种对品种改良方面的作用。
本试验获得主要研究结果如下:
(1)建立糖橙实生态的再生培养体系。以30d苗龄的糖橙实生态节间茎段不定芽分化率较高,暗培养可以显著的提高不定芽的再生率。筛选出适宜实生态节间茎段再生的不定芽诱导的培养基:MS+5mg/L 6-BA;不定芽增殖培养基:MS+1.0 mg/L 6-BA+NAA 1.0 mg/L;生根培养基:MS+0.5 mg/L NAA。其中采用两步法诱导不定芽再生和生根效果较好。
(2)建立了成年态糖橙的不定芽再生体系。用改良的消毒方法G,即采枝梢前一周,每隔一天下午,于温室中将要采取的枝梢用10%的次氯酸钠溶液擦拭一遍,共擦拭4次,且供试植株须隔离放置。再采用70%的酒精处理30s,再用1%的升汞溶液加几滴吐温浸泡24min,无菌水冲洗4次进行消毒。可显著的提高成年态节间茎段的成活率。成年态节间茎段培养基以6-BA 5 mg/L+2,4-D 2 mg/L+IAA 2mg/L培养的效果最好,茎段愈伤诱导率和不定芽诱导率最高可达17.8%和2.87个。对成年态不定芽的嫁接进行了研究,认为经过改良的“⊥”压法成活率较高,可以达到78.94%。
(3)建立了将DefH9-iaaM基因和TERF1基因两个外源基因导入到实生态糖橙的转化体系。研究了不同根癌农杆菌对糖橙实生态节间茎段的转化率,结果显示EHA105的转化能力强于LBA4404。浸染时间以15min为宜。针对糖橙实生态节间茎段和成年态节间茎段再生能力的差异性,认为卡那霉素浓度为100mg/L时适合对糖橙实生态节间茎段再生不定芽进行筛选,卡那霉素浓度为30mg/L时适合对柑橘成年态节间茎段再生不定芽进行筛选,通过PCR鉴定,不定芽的转化率高达29.4%。
(4)通过PCR鉴定,结果显示以获得了6株转DefH9-iaaM基因植株,通过形态学的研究证实,这6株转基因植株在株高、节间长度、叶面积方面比对照有明显的增长、增宽现象。利用PCR、RT-PCR、Southern杂交技术对转TERF1基因的糖橙进行了分子检测,结果显示,共获得了7株转TERF1基因植株株系。通过对3,7,11号植株的Southern杂交,结果显示TERF1基因已经成功的整合到了糖橙的基因组中,且均为为单拷贝插入。从植株形态特征表现上看,转TERF1基因的植株与未转化植株无显著差异。
(5)对转TERF1基因的糖橙进行离体溃疡病和炭疽病的接种试验。通过对病斑大小和数量的分析,结果表明1,3,7,8,11号株系对柑橘溃疡病和炭疽病表现出广谱抗性。接种溃疡病病害时,3号株系对溃疡病的抗病性最强,发病级别为0级。其余转基因植株溃疡病的发病率比对照植株下降了70%左右。进行炭疽病的离体接种7d后发现,转基因植株的叶片基本上都表现出针孔处病斑减小。从照片上可以看到3号和7号转TERF1基因糖橙叶片不仅基本上观察不到炭疽病的典型病斑,而且叶片仍然可以保持翠绿。
Citrus is the most important fruit crop in the world.Now its annual production in the world is 110 million metric tons,with a total acreage of 7.15 million hectares (ha) and an annual volume of international trade about 650 million dollars.China is recognized as one of the key original centers of citrus,with a long history of cultivation,plenty of natural resources and selected varieties for commercial use.
In 2006,the planting area of citrus in China exceeded 1.7 million ha,which was 23%of the total acreage and ranked the first place in the world.The annual yield of citrus was nearly 18 million ton accepting the second biggest country for citrus production in the world.China has a long history of citrus breeding and has established the foundation of citrus industry,and the distance on technology application is widely accepted compared with those advanced countries of citrus production.In order to catch up with the international trade standard,the most important thing is to produce fruit of high quality and to satisfy the consumers.The criteria for evaluating the citrus cultivars are summarized as the following five main aspects:the appearance,taste,nutritive content,seeds number and easy-peeling by hand,et al.From the view of botany,the citrus is as a woody,low branching plant with a distinct juvenile stage,polyembryony,high genetic heterozygosity and arthenogenesis,and these characteristics have impeded the traditional breeding in citrus,such as the blindness in crossing and low efficiency for new cultivar development.With the development of biological technology,using transgenic technology to breed new variety is the best way to modify the genetic expressing or silencing.Transferring the foreign genes into mature explants will save us a lot of breeding time,and the gene expression in flower and fruit can be investigated much earlier than traditional did.
Succari is a very popular sweet orange,almost acidless,and is extending quickly in the south of China,but with plenty of seeds.Liking other sweet oranges, Succari trees can be attacked by many biotic(bacteria,fungi,etc.) and abiotic stresses. The objective of this project is to establish the regeneration and transformation system of Succari orange and to transfer a parthenocarpy gene 'defH9-iaaM' and a transcription factor 'TERFI' gene into the genome by using seedlings and adult inter-nodal segments as explants.Hopefully to gain a new germplasm or breeding material,which carrying parthenocarpy gene and got a broad-spectrum resistance to diseases.The DefH9-iaaM gene consists of an iaaM gene and a DefH9 promotor.The transcription factors TERF1 gene as cloned from tomato by yeast-one-hybrid system. After the disease inoculation and morphological trait observation,we evaluated the modification capability of the wild plants by foreign gene.
The main results are showed as follows:
1.Establishing the regeneration system from Succari orange seedling sterms intemodal segments.The results indicated that 30-day-old seedlings were the most efficient explants,and the co-culture in darkness for 6 days gave much improvement for good regeneration.The optimum medium for adventitious buds regeneration was MS + 6-BA(5.0mg/L).And MS medium supplied with 6-BA1.0mg/L and NAA 0.1mg/L showed the highest bud multiplication ratio.The shoot tissues got rooted successfully after 20 days in 0.5mg/L NAA.The better way to get more adventitious buds is using the "two-step" method,which is to put the explants from the medium plus the auxin first,and then remove to medium with no hormones after 6 days.
2.To set up an efficient regeneration system for mature sterms internodal segments.It shows that pretreatment disinfection method can raise significantly the survival rate to 80%.And the medium supplied with 6-BA 5 mg/L,2,4-D 2 mg/L and IAA 2mg/L could staminate the callus from cutting ends.And the callus induction rate reached up to 17.8%and 2.87 for the adventitious buds.The effect of three grafting methods on improving survival rate of adventitious buds was investigated,and the result showed that modified "⊥" way was the best one,which 78.94%of survival rate.
3.Development of the transformation system and the transfer of DefH9-iaaM gene and TERF1 gene into Succari.The effects of transformed by different Agrobacterium strains was observed,and it seemed that EHA105 was better than LBA4404.And 15 min was the best infection time.The selection concentration of km is different between the seedlings and mature explants,and 100 mg/L for seedling explants and 30 mg/L for mature explants were justified.
4.PCR results indicated that six regenerate plants of Succari were transgenic with the DefH9-iaaM gene,and the six transgenic plants showed evident increase in plant height,stem length,and leaf area.Through the PCR and RT-PCR analyses,it was showed that 7 regenerants were cooperated with the TERF1 gene.No.3,No.7 and No. 11 transgenic lines were verified by Southern blotting,and the results showed that one copy of the TERF1 gene were insert the plant genome.
5.After inoculation with the citrus canker and anthracnose pathogen the transgenic plants of TERF1 gene,the results showed that the TERF1 gene can improve the resistance to both diseases.The transgenic plants could reduce citrus bacterial canker incidence rate,by 70%.The No.3 showed 0%of the incidence rate.When the transgenic leaves and control ones were both infected with the citrus anthracnose,all the leaves showed blots symptom of disease.The blots on transgenic leaves were smaller than that on the controls.Leaves of No.3 and No.7 plants did not have the disease blots,and the color of leaves also kept green.
引文
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