LeGGPS2、AtCAO与AtHEMA1基因对烟草耐弱光性的影响
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摘要
弱光是目前制约设施生产的重要因素。研究表明弱光逆境不仅会引起植物叶绿素含量的变化,而且影响光合速率、光合产物的运输与分配,进而影响植物的生长发育。转基因技术具有其独特的优势,是国际上公认的改良作物农艺性状的可行技术,且在提高作物光合速率方面已取得一定进展,但弱光性状是多基因控制的数量性状,利用转化色素合成关键基因提高作物光合速率及耐弱光性的报道迄今少见。
本研究通过分析叶绿素和类胡萝卜素的合成途径,从拟南芥中克隆叶绿素合成初期的关键基因谷氨酰-tRNA还原酶基因(AtHEMA1)、催化叶绿素b合成的关键基因叶绿素酸酯a氧化酶基因(AtCAO);并构建了含番茄牻牛儿基牻牛儿基焦磷酸合成酶基因(LeGGPS2)的双元表达载体pVCT2295,含AtCAO、AtHEMA1基因的双元表达载体pVCT2299;通过农杆菌介导法将LeGGPS2基因、AtCAO与AtHEMA1基因转入烟草。对转基因烟草植株进行弱光处理后,分析导入基因对烟草光合色素含量、光合速率、光合产物及植株生长发育的影响。
主要结果如下：
1.构建了含LeGGPS2基因的双元表达载体pVCT2295,并成功转化烟草
将从番茄品种Ailsa Craig (Solanwn lycopersicon L. cv. Ailsa Craig)中克隆的LeGGPS2基因插入到载体pVCT2024中,构建了含有LeGGPS2等基因的双元载体pVCT2295,经农杆菌介导转化烟草。对Kan抗性烟草植株进行PCR鉴定、荧光检测,证明获得了整合有Pnos-nptⅡ、35S-LeGGPS2和35S-GFP外源基因的转基因烟草植株9株。
2. LeGGPS2基因增强了弱光下烟草的光合性能,且增加了生物量积累
弱光处理后,转基因烟草株系TG1、TG2的类胡萝卜素含量、叶绿素含量、光合速率均比野生烟草增加,达到了差异显著性水平,可见转入LeGGPS2基因可提高烟草弱光下的光合性能。另外,转基因烟草的叶面积增长量、单位叶面积重量、总干重、根冠干重比,均比野生烟草高,证明转入该基因可提高烟草的生物量积累并促进其向根部的分配。
3.克隆了AtCAO与AtHEMA1基因
参照GenBank中公布的拟南芥叶绿素酸酯a加氧酶(AtCAO)、谷氨酰tRNA还原酶(AtHEMAl)基因序列,设计特异PCR引物,采用PrimStar HS DNA聚合酶从拟南芥(Arabidopsis thaliana L. Ecotype Columbia) gDNA中扩增目的片段,分别TA克隆进T-载体pMD18-T和pMD19-T,构成载体pVCT1263、pVCT1298。测序结果显示与网上公布序列一致,表明成功克隆了AtCAO与AtHEMA1基因。
4.成功获得了分别含AtCAO基因、AtHEMA1基因的植物表达载体pVCT2296和pVCT2298
将克隆的AtCAO基因插入到载体pVCT2100中,构建含有nptⅡ AtCAO基因的表达载体pVCT2296;将克隆的AtHEMA1基因插入到载体pVCT2101中,构建含有npⅡ、AtHEMA1基因的表达载体pVCT2298;通过PCR和酶切鉴定,证明载体pVCT2296、pVCT2298构建成功。
5.获得了含AtCAO与AtHEMAl基因的植物表达载体pVCT2299并成功转化烟草
将克隆的AtCAO基因插入到载体pVCT2298中,构建了含有nptⅡ、AtCAO, AtHEMA1基因的表达载体pVCT2299,并经过农杆菌介导转化烟草,对Kan抗性烟草植株进行PCR鉴定,证明获得了整合有nptⅡ、AtCAO、AtHEMA1外源基因的转基因烟草植株24株。进一步对选取的转基因烟草株系TL1、TL2进行Southern杂交,证实获得了含单拷贝基因的烟草植株。
6.转AtCAO和AtHEMA1双价基因不仅促进烟草叶绿素b合成,且促进生长发育,显示出能增强烟草的耐弱光性
弱光下转基因烟草相比野生烟草,叶绿素总量增加不显著,但叶绿素b含量增加16%～17%,叶绿素a/b比值降低9%-12%,光合作用增强42%～65%,均达到差异显著水平,而且生长发育比野生烟草快。
Low light is an important factor of restricting the production facilities. The study shows that low light stress can cause changes in the content of chlorophyll, the intensity of photosynthesis, the transport and distribution of photosynthetic products, thus affecting the growth and development of plants. Transgenic technology has its unique advantages and is internationally recognized as viable technology for improving crop agronomic traits, and some progress has been made in improving the photosynthetic rate of plants, but the low light stress is a scalar property manipulated by many genes, so far it is reported rarely to improve plant photosynthetic rate and resistant to low light by transforming some key genes in pigment synthesis.
In this study, we cloned glutamyl-tRNA reductase gene (AtHEMAl) from the genome of Arabidopsis.thaliana, which is the early key gene for chlorophyll synthesis gene, also we cloned the chlorophyllide a oxygenase(AtCAO), which catalytic synthesis of chlorophyll b and then we constructed the expression vector pVCT2299, in which contain the AtCAO gene and AtHEMAl gene. At the same time, we constructed the expression vector contain LeGGPS2gene. All the vectors were successfully transformed into tobacco by Agrobacterium-mediated transformation. After low light treatment, we analysed the content of pigment, the intensity of photosynthesis and the growth and development of transgenic tobacco. The main results are as follows:
(1) Constructed the expression vector pVCT2295which contains LeGGPS2gene and successfully transformed it into tobacco by Agrobaclerium-mediated leaf disc transformation.
The LeGGPS2gene was amplfied from tomato cultivar Ailsa Craig. We constructed the binary vector pVCT2295which contain35S-LeGGPS2gene by inserting the LeGGPS2gene fragment into pVCT2024. Then the pVCT2295vector was transformed into tobacco through Agrobaclerium-mediated leaf disc transformation. PCR analysis and fluorescence detection proved that we have got nine tobaccoes which contain nptll, LeGGPS2and GFP genes.
(2) Transgenic tobacco with LeGGPS2gene had increased the photosynthetic performance and the biomass accumulation after low light treatment
After low light treatment, we found the transgenic tobacco have higher contents of carotenoid and chlorophyll,also increase intensity of photosynthesis comare with wild tobacco, and some transgenic lines are significant difference, which indicated that the LeGGPS2gene can improve the tobacco photosynthesis performance under low light. In addition, the transgenic tobacco leaf area, the unit weight of leaf area, total dry weight, root and shoot dry weight ratio were higher than wild tobacco, which proved the transgenic tobacco had increased the biomass accumulation and promoted its trsnsport into roots.
(3) Cloned the AtCAO and AtHEMAl genes
According to sequence published in the GenBank, we amplified the chlorophyllide a oxygenase (AtCAO) and glutamyl-tRNA reductase gene from Arabidopsis (Arabidopsis thaliana L. Ecotype Columbia) gDNA with specific PCR primers and PrimStar HS DNA polymerase. Then the fragments were TA cloned into vector pMD18-T, pMD19-T.respectively The sequence analysis display it is in accordance with the online sequence, so we obtained pVCT1263, pVCT1298vector successfully.
(4) Successfully constructed the expression vector pVCT2296and pVCT2298.
We constructed the expression vector pVCT2296which contain nptll and AtCAO genes by inserting AtCAO gene into pVCT2100and constructed the expression vector pVCT2298which contain nptll and AtHEMAl genes by inserting AtHEMAl gene into pVCT2101. The PCR identification and restriction enzyme digestion proved we obtained the correct vector pVCT2296and pVCT2298.
(5) Constructed the expression vector pVCT2299contain AtCAO and AtHEMAl gene and successfully transformed it into tobacco by Agrobaclerium-mediated leaf disc transformation.
We constructed the expression vector pVCT2299, in which contain nptll, AtCAO and AtHEMAl genes, by inserting AtCAO gene into pVCT2298and transformed it into tobacco using Agrobacterium-mediated method. By the rusults of PCR identification of kan resistant tobacco, we have obtained24strains transgenic tobacco plants with nptII、AtCAO and AtHEMA1exogenous genes. We have conducted Southern blot for furthern analysis the copy number of transgenic tobacco, proved transgenic tobcco TL1and TL2had single-copy.
(6) The tansgenic tobaco with AtCAO and AtHEMAl genes not only can accelerate the amount of chlorophyll b, but also can accelerate the growth and development, which display increased resistance to low light
Both the amount of chlorophyll b and the rate of photosynthesis remarkably increased in the transgenic plant lines TL1and TL2compared with the wild type tobacco, while the chlorophyll a/b decreased significantly(P<0.01). We also found the trsngenic tobacco had inreased23%-35%in height compared with wild tobacco, and the index was significant difference (P<0.05). The height was closely related to the time of buds appearing, so the buds of the transgenic tobacco10-20days earlier than wild tobacco, which showed its low light tolerance.
本研究通过分析叶绿素和类胡萝卜素的合成途径,从拟南芥中克隆叶绿素合成初期的关键基因谷氨酰-tRNA还原酶基因(AtHEMA1)、催化叶绿素b合成的关键基因叶绿素酸酯a氧化酶基因(AtCAO);并构建了含番茄牻牛儿基牻牛儿基焦磷酸合成酶基因(LeGGPS2)的双元表达载体pVCT2295,含AtCAO、AtHEMA1基因的双元表达载体pVCT2299;通过农杆菌介导法将LeGGPS2基因、AtCAO与AtHEMA1基因转入烟草。对转基因烟草植株进行弱光处理后,分析导入基因对烟草光合色素含量、光合速率、光合产物及植株生长发育的影响。
主要结果如下：
1.构建了含LeGGPS2基因的双元表达载体pVCT2295,并成功转化烟草
将从番茄品种Ailsa Craig (Solanwn lycopersicon L. cv. Ailsa Craig)中克隆的LeGGPS2基因插入到载体pVCT2024中,构建了含有LeGGPS2等基因的双元载体pVCT2295,经农杆菌介导转化烟草。对Kan抗性烟草植株进行PCR鉴定、荧光检测,证明获得了整合有Pnos-nptⅡ、35S-LeGGPS2和35S-GFP外源基因的转基因烟草植株9株。
2. LeGGPS2基因增强了弱光下烟草的光合性能,且增加了生物量积累
弱光处理后,转基因烟草株系TG1、TG2的类胡萝卜素含量、叶绿素含量、光合速率均比野生烟草增加,达到了差异显著性水平,可见转入LeGGPS2基因可提高烟草弱光下的光合性能。另外,转基因烟草的叶面积增长量、单位叶面积重量、总干重、根冠干重比,均比野生烟草高,证明转入该基因可提高烟草的生物量积累并促进其向根部的分配。
3.克隆了AtCAO与AtHEMA1基因
参照GenBank中公布的拟南芥叶绿素酸酯a加氧酶(AtCAO)、谷氨酰tRNA还原酶(AtHEMAl)基因序列,设计特异PCR引物,采用PrimStar HS DNA聚合酶从拟南芥(Arabidopsis thaliana L. Ecotype Columbia) gDNA中扩增目的片段,分别TA克隆进T-载体pMD18-T和pMD19-T,构成载体pVCT1263、pVCT1298。测序结果显示与网上公布序列一致,表明成功克隆了AtCAO与AtHEMA1基因。
4.成功获得了分别含AtCAO基因、AtHEMA1基因的植物表达载体pVCT2296和pVCT2298
将克隆的AtCAO基因插入到载体pVCT2100中,构建含有nptⅡ AtCAO基因的表达载体pVCT2296;将克隆的AtHEMA1基因插入到载体pVCT2101中,构建含有npⅡ、AtHEMA1基因的表达载体pVCT2298;通过PCR和酶切鉴定,证明载体pVCT2296、pVCT2298构建成功。
5.获得了含AtCAO与AtHEMAl基因的植物表达载体pVCT2299并成功转化烟草
将克隆的AtCAO基因插入到载体pVCT2298中,构建了含有nptⅡ、AtCAO, AtHEMA1基因的表达载体pVCT2299,并经过农杆菌介导转化烟草,对Kan抗性烟草植株进行PCR鉴定,证明获得了整合有nptⅡ、AtCAO、AtHEMA1外源基因的转基因烟草植株24株。进一步对选取的转基因烟草株系TL1、TL2进行Southern杂交,证实获得了含单拷贝基因的烟草植株。
6.转AtCAO和AtHEMA1双价基因不仅促进烟草叶绿素b合成,且促进生长发育,显示出能增强烟草的耐弱光性
弱光下转基因烟草相比野生烟草,叶绿素总量增加不显著,但叶绿素b含量增加16%～17%,叶绿素a/b比值降低9%-12%,光合作用增强42%～65%,均达到差异显著水平,而且生长发育比野生烟草快。
Low light is an important factor of restricting the production facilities. The study shows that low light stress can cause changes in the content of chlorophyll, the intensity of photosynthesis, the transport and distribution of photosynthetic products, thus affecting the growth and development of plants. Transgenic technology has its unique advantages and is internationally recognized as viable technology for improving crop agronomic traits, and some progress has been made in improving the photosynthetic rate of plants, but the low light stress is a scalar property manipulated by many genes, so far it is reported rarely to improve plant photosynthetic rate and resistant to low light by transforming some key genes in pigment synthesis.
In this study, we cloned glutamyl-tRNA reductase gene (AtHEMAl) from the genome of Arabidopsis.thaliana, which is the early key gene for chlorophyll synthesis gene, also we cloned the chlorophyllide a oxygenase(AtCAO), which catalytic synthesis of chlorophyll b and then we constructed the expression vector pVCT2299, in which contain the AtCAO gene and AtHEMAl gene. At the same time, we constructed the expression vector contain LeGGPS2gene. All the vectors were successfully transformed into tobacco by Agrobacterium-mediated transformation. After low light treatment, we analysed the content of pigment, the intensity of photosynthesis and the growth and development of transgenic tobacco. The main results are as follows:
(1) Constructed the expression vector pVCT2295which contains LeGGPS2gene and successfully transformed it into tobacco by Agrobaclerium-mediated leaf disc transformation.
The LeGGPS2gene was amplfied from tomato cultivar Ailsa Craig. We constructed the binary vector pVCT2295which contain35S-LeGGPS2gene by inserting the LeGGPS2gene fragment into pVCT2024. Then the pVCT2295vector was transformed into tobacco through Agrobaclerium-mediated leaf disc transformation. PCR analysis and fluorescence detection proved that we have got nine tobaccoes which contain nptll, LeGGPS2and GFP genes.
(2) Transgenic tobacco with LeGGPS2gene had increased the photosynthetic performance and the biomass accumulation after low light treatment
After low light treatment, we found the transgenic tobacco have higher contents of carotenoid and chlorophyll,also increase intensity of photosynthesis comare with wild tobacco, and some transgenic lines are significant difference, which indicated that the LeGGPS2gene can improve the tobacco photosynthesis performance under low light. In addition, the transgenic tobacco leaf area, the unit weight of leaf area, total dry weight, root and shoot dry weight ratio were higher than wild tobacco, which proved the transgenic tobacco had increased the biomass accumulation and promoted its trsnsport into roots.
(3) Cloned the AtCAO and AtHEMAl genes
According to sequence published in the GenBank, we amplified the chlorophyllide a oxygenase (AtCAO) and glutamyl-tRNA reductase gene from Arabidopsis (Arabidopsis thaliana L. Ecotype Columbia) gDNA with specific PCR primers and PrimStar HS DNA polymerase. Then the fragments were TA cloned into vector pMD18-T, pMD19-T.respectively The sequence analysis display it is in accordance with the online sequence, so we obtained pVCT1263, pVCT1298vector successfully.
(4) Successfully constructed the expression vector pVCT2296and pVCT2298.
We constructed the expression vector pVCT2296which contain nptll and AtCAO genes by inserting AtCAO gene into pVCT2100and constructed the expression vector pVCT2298which contain nptll and AtHEMAl genes by inserting AtHEMAl gene into pVCT2101. The PCR identification and restriction enzyme digestion proved we obtained the correct vector pVCT2296and pVCT2298.
(5) Constructed the expression vector pVCT2299contain AtCAO and AtHEMAl gene and successfully transformed it into tobacco by Agrobaclerium-mediated leaf disc transformation.
We constructed the expression vector pVCT2299, in which contain nptll, AtCAO and AtHEMAl genes, by inserting AtCAO gene into pVCT2298and transformed it into tobacco using Agrobacterium-mediated method. By the rusults of PCR identification of kan resistant tobacco, we have obtained24strains transgenic tobacco plants with nptII、AtCAO and AtHEMA1exogenous genes. We have conducted Southern blot for furthern analysis the copy number of transgenic tobacco, proved transgenic tobcco TL1and TL2had single-copy.
(6) The tansgenic tobaco with AtCAO and AtHEMAl genes not only can accelerate the amount of chlorophyll b, but also can accelerate the growth and development, which display increased resistance to low light
Both the amount of chlorophyll b and the rate of photosynthesis remarkably increased in the transgenic plant lines TL1and TL2compared with the wild type tobacco, while the chlorophyll a/b decreased significantly(P<0.01). We also found the trsngenic tobacco had inreased23%-35%in height compared with wild tobacco, and the index was significant difference (P<0.05). The height was closely related to the time of buds appearing, so the buds of the transgenic tobacco10-20days earlier than wild tobacco, which showed its low light tolerance.
引文
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