不结球白菜冷诱导差异表达基因的筛选及功能分析
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摘要
随着全球极端灾害性天气频发,冷害成为蔬菜生产上的一个严重问题,造成蔬菜品质下降以及严重减产,直接影响冬、春季蔬菜周年供应。冷害影响植物的生长和发育,包括：渗透胁迫和次级胁迫如过氧化物胁迫等,常常导致植物细胞骨架结构破坏、膜脂相变、细胞水分亏缺、体内酶的活性降低和光合速率下降。但植物对冷害胁迫的响应并非是完全被动的,为了适应抵抗冷害,植物在长期的进化过程中,形成了相应的保护机制。对模式植物拟南芥的研究表明,冷驯化是一个由众多基因协同调控,有着复杂调控网络的过程,其中少数关键性调控基因控制着大量基因的表达。不结球白菜(Brassica campestris ssp. chinensis Makino)经过一段零度以上的冷诱导,即冷驯化过程,可以获得一定的抗寒能力,但对其机理的研究较少。采用cDNA-AFLP技术,以不结球白菜抗寒自交系043为材料,从转录组水平上分离冷诱导差异表达的相关基因,为探明不结球白菜冷驯化的分子机理提供理论依据。主要研究结果如下：
1.不结球白菜冷诱导相关基因差异表达的cDNA-AFLP分析
采用cDNA-AFLP技术,对不结球白菜抗寒自交系043在冷诱导(4℃)下差异表达的基因进行了分析,256对引物组合共筛选了9960个cDNA片段,获得差异片段77个。其中上调表达的基因片段为57个,下调表达的基因片段为20个。对其功能比对发现,这些TDFs分别参与了基础代谢、光合作用和能量、转运、信号转导、次生代谢、防御响应、转录、细胞骨架结构,另外还有一些假想和功能未知蛋白。同时选取TDF16#、TDF21#和TDF57#进行实时定量PCR分析,结果表明这三个TDFs受冷胁迫的诱导表达,在不结球白菜冷驯化过程中可能具有重要作用。构建了不结球白菜在冷诱导下的基因表达谱,从转录组水平上鉴定了一批冷诱导相关基因,这些新基因可用于冷驯化的分子机理研究。
2.不结球白菜冷诱导相关基因BcWRKY46的克隆及功能分析
WRKY转录因子能广泛地参与植物的生物与非生物胁迫反应。以cDNA-AFLP技术,从不结球白菜抗寒自交系043冷诱导(4℃)的叶片中得到的一个差异表达片段TDF21#,进一步利用RACE技术获得了该基因cDNA全序列1175bp,包含有855bp的开放阅读框,编码284个氨基酸,命名为BcWRKY46。qRT-PCR结果表明,该基因表达不仅受冷胁迫诱导,还响应其它胁迫,包括盐、干旱和ABA。过量表达BcWRKY46的转基因烟草提高了抗寒、盐、干旱和ABA胁迫的能力。结果表明,BcWRKY46在提高ABA和非生物胁迫的抗性过程中有着重要的作用。
3.不结球白菜冷诱导相关基因BcMCSU的克隆及功能分析
采用cDNA-AFLP技术,从不结球白菜抗寒自交系043冷诱导(4℃)的叶片中得到的一个差异表达片段TDF16#,进一步利用RACE技术获得了该基因全长序列1194bp,包含有926bp的开放阅读框,编码307个氨基酸,命名为BcMCSU。其氨基酸序列与拟南芥AtMCSU编码的氨基酸序列具有较高同源性。蛋白结构域分析表明：所推导的氨基酸序列具有完整的结构,在其N-端和C-端分别含有一个MOSC结构域。采用Gateway技术构建植物过量表达载体,通过农杆菌介导的真空渗透法将其转入野生型拟南芥中,经除草剂筛选,获得了抗性转基因拟南芥。
4.不结球白菜冷诱导相关基因BcVIN3的克隆及功能分析
采用cDNA-AFLP技术,从不结球白菜抗寒自交系043冷诱导(4℃)的叶片中得到的一个差异表达片段TDF57#,进一步利用RACE技术获得了该基因全长序列1814bp,包含有1749bp的开放阅读框,编码582个氨基酸,命名为BcVIN3。其氨基酸序列与拟南芥AtVIN3编码的氨基酸序列具有较高同源性。蛋白结构域分析表明：所推导的氨基酸序列具有完整的结构,包含一个PHD-finger结构域。采用Gateway技术构建植物过量表达载体,通过花粉管通道法将其转入不结球白菜‘苏州青’中,经除草剂筛选,获得了抗性转基因不结球白菜。
Temperature fluctuations and extreme freezing temperature at crown level, occurring during winter and early spring, cause recurrent losses of vegetables production. Cold stress severely affect the growth and development of plants through the imposition of osmotic stress, and secondary stress such as oxidative stress leading to membrane disorganization, metabolic toxicity, and inhibition of photosynthesis. Many plants increase in freezing tolerance in response to low nonfreezing temperatures, a phenomenon known as cold acclimation. Freezing tolerance of such plants increases substantially after a period of exposure to low but non-freezing temperature. Cold acclimation of plant is a highly active process resulting from gene expression involved in physiological and metabolic adaptations to low temperature. Cold acclimation involves the remodeling of cell and tissue structures, the reprogramming of metabolism and gene expression, which encoding molecular chaperones, lipid, and sugar metabolism are suggested to be involved in cold adaptation. Studies on acquired freezing tolerance in Arabidopsis have contributed substantially towards the understanding of cold acclimation mechanisms. But the molecular basis of this acquired chilling tolerance or chilling acclimation is poorly understood in non-heading Chinese cabbage (Brassica campestris ssp. chinensis Makino). To date, non-heading Chinese cabbage has not been exploited to understand the molecular basis of its unusually high level of cold tolerance. Here cDNA-AFLP was used to identify transcripts that are strongly accumulated and induced in response to cold stress.
1. Differential expression analysis of candidate genes involved in cold response in non-heading Chinese cabbage using cDNA-AFLP
The objective of this study was to determine temporal expression profiles of transcripts during cold acclimation of non-heading Chinese cabbage cold-tolerant inbred line043by using cDNA-AFLP.256primer combinations were used to investigate9960cDNA fragments. A total of77differentially expressed cDNA fragments were detected,57of which were up-regulated and20were down-regulated under cold treatment (4℃). These TDFs were classified into several functional groups with Blast in GenBank. Except for18function-unknown TDFs, the remained up-or down-regulated TDFs conferred functions of metabolism, photosynthesis and energy, transport, signal transduction, secondary metabolism, defense response, transcription, cytoskeleton, function unknown and hypothetical protein. Meanwhile, the expression patterns of three TDFs, TDF16#, TDF21#, and TDF57#showed a significant increase of differential expression after cold stress was analysed by qRT-PCR. These three TDFs might play important roles in promoting adaptation to cold stress. The result showed that all of the three TDFs were induced by cold stress, which suggest that play important roles in promoting adaptation to cold stress. This study provides important clues to understanding cold acclimation and low-temperature regulation mechanisms in non-heading Chinese cabbage and the three TDFs involved in cold responses need further research to determine their usefulness in breeding new resistance cultivars.
2. Molecular cloning, characterization and function analysis of BcWRKY46from non-heading Chinese cabbage
WRKY transcription factors belong to one of the largest families of transcriptional regulators in plants and form integral parts of signaling webs that modulate many vital processes during plant growth and development. BcWRKY46, a cDNA clone encoding a polypeptide of284amino acids and exhibited the structural features of group III of WRKY protein family, was isolated from the cold-treated leaves of non-heading Chinese cabbage using the cDNA-AFLP technique. Expression of this gene was induced quickly and strongly in response to various environmental stresses, including low temperatures, ABA, salt and dehydration. Constitutive expression of BcWRKY46in tobacco under the control of the CaMV35S promoter reduced the susceptibility of transgenic tobacco to freezing, ABA, salt and dehydration stresses. Our studies suggest that BcWRKY46plays an important role in responding to ABA and abiotic stress.
3. Molecular cloning, characterization and function analysis of BcMCSU from non-heading Chinese cabbage
Using cDNA-AFLP techniques, a cold induced TDF16#homologous to AtMCSU was isolated from the leaves of non-heading Chinese cabbage which was named as BcMCSU. The full-length BcMCSU cDNA which consisted of a single open reading frame encoded a putative polypeptide of307amino acids. According to the functional domain analysis of the predicted amino acid sequence, the BcMCSU protein contains a Nifs domain at its N-terminus and a MOSC domain at the C-terminus. An efficiently overexpression vector of pEG103-BcMCSU was constructed and transformed into Arabidopsis thaliana by Agrobacterium mediated depressor permeating method. Transgenic Arabidopsis thaliana plants were obtained by resistance screening and PCR identification.
4. Molecular cloning, characterization and function analysis of BcVIN3from non-heading Chinese cabbage
Using cDNA-AFLP techniques, a cold induced TDF57#homologous to AtVIN3was isolated from the leaves of non-heading Chinese cabbage which was named as BcVIN3. The full-length cDNA sequence of BcVIN3was1814bp, and encoding a putative polypeptide of582amino acids. According to the functional domain analysis of the predicted amino acid sequence, the BcVIN3protein contains a PHD-finger domain. An efficiently overexpression vector of pEG103-BcVIN3was constructed and transformed into non-heading Chinese cabbage'Suzhouqing' by the pollen tube pathway method. Transgenic plants were obtained by resistance screening and PCR identification.
1.不结球白菜冷诱导相关基因差异表达的cDNA-AFLP分析
采用cDNA-AFLP技术,对不结球白菜抗寒自交系043在冷诱导(4℃)下差异表达的基因进行了分析,256对引物组合共筛选了9960个cDNA片段,获得差异片段77个。其中上调表达的基因片段为57个,下调表达的基因片段为20个。对其功能比对发现,这些TDFs分别参与了基础代谢、光合作用和能量、转运、信号转导、次生代谢、防御响应、转录、细胞骨架结构,另外还有一些假想和功能未知蛋白。同时选取TDF16#、TDF21#和TDF57#进行实时定量PCR分析,结果表明这三个TDFs受冷胁迫的诱导表达,在不结球白菜冷驯化过程中可能具有重要作用。构建了不结球白菜在冷诱导下的基因表达谱,从转录组水平上鉴定了一批冷诱导相关基因,这些新基因可用于冷驯化的分子机理研究。
2.不结球白菜冷诱导相关基因BcWRKY46的克隆及功能分析
WRKY转录因子能广泛地参与植物的生物与非生物胁迫反应。以cDNA-AFLP技术,从不结球白菜抗寒自交系043冷诱导(4℃)的叶片中得到的一个差异表达片段TDF21#,进一步利用RACE技术获得了该基因cDNA全序列1175bp,包含有855bp的开放阅读框,编码284个氨基酸,命名为BcWRKY46。qRT-PCR结果表明,该基因表达不仅受冷胁迫诱导,还响应其它胁迫,包括盐、干旱和ABA。过量表达BcWRKY46的转基因烟草提高了抗寒、盐、干旱和ABA胁迫的能力。结果表明,BcWRKY46在提高ABA和非生物胁迫的抗性过程中有着重要的作用。
3.不结球白菜冷诱导相关基因BcMCSU的克隆及功能分析
采用cDNA-AFLP技术,从不结球白菜抗寒自交系043冷诱导(4℃)的叶片中得到的一个差异表达片段TDF16#,进一步利用RACE技术获得了该基因全长序列1194bp,包含有926bp的开放阅读框,编码307个氨基酸,命名为BcMCSU。其氨基酸序列与拟南芥AtMCSU编码的氨基酸序列具有较高同源性。蛋白结构域分析表明：所推导的氨基酸序列具有完整的结构,在其N-端和C-端分别含有一个MOSC结构域。采用Gateway技术构建植物过量表达载体,通过农杆菌介导的真空渗透法将其转入野生型拟南芥中,经除草剂筛选,获得了抗性转基因拟南芥。
4.不结球白菜冷诱导相关基因BcVIN3的克隆及功能分析
采用cDNA-AFLP技术,从不结球白菜抗寒自交系043冷诱导(4℃)的叶片中得到的一个差异表达片段TDF57#,进一步利用RACE技术获得了该基因全长序列1814bp,包含有1749bp的开放阅读框,编码582个氨基酸,命名为BcVIN3。其氨基酸序列与拟南芥AtVIN3编码的氨基酸序列具有较高同源性。蛋白结构域分析表明：所推导的氨基酸序列具有完整的结构,包含一个PHD-finger结构域。采用Gateway技术构建植物过量表达载体,通过花粉管通道法将其转入不结球白菜‘苏州青’中,经除草剂筛选,获得了抗性转基因不结球白菜。
Temperature fluctuations and extreme freezing temperature at crown level, occurring during winter and early spring, cause recurrent losses of vegetables production. Cold stress severely affect the growth and development of plants through the imposition of osmotic stress, and secondary stress such as oxidative stress leading to membrane disorganization, metabolic toxicity, and inhibition of photosynthesis. Many plants increase in freezing tolerance in response to low nonfreezing temperatures, a phenomenon known as cold acclimation. Freezing tolerance of such plants increases substantially after a period of exposure to low but non-freezing temperature. Cold acclimation of plant is a highly active process resulting from gene expression involved in physiological and metabolic adaptations to low temperature. Cold acclimation involves the remodeling of cell and tissue structures, the reprogramming of metabolism and gene expression, which encoding molecular chaperones, lipid, and sugar metabolism are suggested to be involved in cold adaptation. Studies on acquired freezing tolerance in Arabidopsis have contributed substantially towards the understanding of cold acclimation mechanisms. But the molecular basis of this acquired chilling tolerance or chilling acclimation is poorly understood in non-heading Chinese cabbage (Brassica campestris ssp. chinensis Makino). To date, non-heading Chinese cabbage has not been exploited to understand the molecular basis of its unusually high level of cold tolerance. Here cDNA-AFLP was used to identify transcripts that are strongly accumulated and induced in response to cold stress.
1. Differential expression analysis of candidate genes involved in cold response in non-heading Chinese cabbage using cDNA-AFLP
The objective of this study was to determine temporal expression profiles of transcripts during cold acclimation of non-heading Chinese cabbage cold-tolerant inbred line043by using cDNA-AFLP.256primer combinations were used to investigate9960cDNA fragments. A total of77differentially expressed cDNA fragments were detected,57of which were up-regulated and20were down-regulated under cold treatment (4℃). These TDFs were classified into several functional groups with Blast in GenBank. Except for18function-unknown TDFs, the remained up-or down-regulated TDFs conferred functions of metabolism, photosynthesis and energy, transport, signal transduction, secondary metabolism, defense response, transcription, cytoskeleton, function unknown and hypothetical protein. Meanwhile, the expression patterns of three TDFs, TDF16#, TDF21#, and TDF57#showed a significant increase of differential expression after cold stress was analysed by qRT-PCR. These three TDFs might play important roles in promoting adaptation to cold stress. The result showed that all of the three TDFs were induced by cold stress, which suggest that play important roles in promoting adaptation to cold stress. This study provides important clues to understanding cold acclimation and low-temperature regulation mechanisms in non-heading Chinese cabbage and the three TDFs involved in cold responses need further research to determine their usefulness in breeding new resistance cultivars.
2. Molecular cloning, characterization and function analysis of BcWRKY46from non-heading Chinese cabbage
WRKY transcription factors belong to one of the largest families of transcriptional regulators in plants and form integral parts of signaling webs that modulate many vital processes during plant growth and development. BcWRKY46, a cDNA clone encoding a polypeptide of284amino acids and exhibited the structural features of group III of WRKY protein family, was isolated from the cold-treated leaves of non-heading Chinese cabbage using the cDNA-AFLP technique. Expression of this gene was induced quickly and strongly in response to various environmental stresses, including low temperatures, ABA, salt and dehydration. Constitutive expression of BcWRKY46in tobacco under the control of the CaMV35S promoter reduced the susceptibility of transgenic tobacco to freezing, ABA, salt and dehydration stresses. Our studies suggest that BcWRKY46plays an important role in responding to ABA and abiotic stress.
3. Molecular cloning, characterization and function analysis of BcMCSU from non-heading Chinese cabbage
Using cDNA-AFLP techniques, a cold induced TDF16#homologous to AtMCSU was isolated from the leaves of non-heading Chinese cabbage which was named as BcMCSU. The full-length BcMCSU cDNA which consisted of a single open reading frame encoded a putative polypeptide of307amino acids. According to the functional domain analysis of the predicted amino acid sequence, the BcMCSU protein contains a Nifs domain at its N-terminus and a MOSC domain at the C-terminus. An efficiently overexpression vector of pEG103-BcMCSU was constructed and transformed into Arabidopsis thaliana by Agrobacterium mediated depressor permeating method. Transgenic Arabidopsis thaliana plants were obtained by resistance screening and PCR identification.
4. Molecular cloning, characterization and function analysis of BcVIN3from non-heading Chinese cabbage
Using cDNA-AFLP techniques, a cold induced TDF57#homologous to AtVIN3was isolated from the leaves of non-heading Chinese cabbage which was named as BcVIN3. The full-length cDNA sequence of BcVIN3was1814bp, and encoding a putative polypeptide of582amino acids. According to the functional domain analysis of the predicted amino acid sequence, the BcVIN3protein contains a PHD-finger domain. An efficiently overexpression vector of pEG103-BcVIN3was constructed and transformed into non-heading Chinese cabbage'Suzhouqing' by the pollen tube pathway method. Transgenic plants were obtained by resistance screening and PCR identification.
引文
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