辣椒WRKY转录因子CaWRKY6和CaWRKY30基因的克隆、表达及功能分析
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摘要
辣椒(Capsicum annuum L.)是我国最主要的蔬菜作物之一,具有重要的经济价值。但辣椒也容易受到各种病原物的危害,近年来由根结线虫(Meloidogyne spp.)侵染引起的根结线虫病已成为辣椒生产中的主要病害之一,由于高抗根结线虫品种的缺乏,生产上主要采用一些高毒、高残留的化学药剂进行防治,从而导致产品的农药残留超标、生态环境的污染和土壤质量的退化,成为蔬菜清洁生产中一个亟待解决的突出问题。
本研究主要从分子水平上来探索辣椒与南方根结线虫早期亲和与不亲和互作的机制,克隆此过程中的关键调控基因并进行功能分析,以期为辣椒的抗性育种提供新思路。
本研究以含有单显性抗根结线虫基因Me3的辣椒HDA149与南方根结线虫亲和与不亲和互作为基础,采用新一代Solexa高通量测序技术,构建了辣椒与南方根结线虫亲和与不亲和互作的转录组cDNA序列数据库。在此基础上,利用RACE方法从辣椒与根结线虫不亲和互作的辣椒根组织中分离出两个推定的WRKY转录因子,并对两个WRKY基因的分子特征和相关功能进行了研究,实验获得的主要结果如下：
1.鉴定了针对Me3基因不同毒力的南方根结线虫(M.incognita)接种辣椒后的表型差异；在此基础上,建立了辣椒与根结线虫亲和互作转录组(SR)cDNA序列数据库和不亲和互作转录组(SR)的cDNA序列数据库,总共获得了214,909条EST序列,其中HR含190,213条,SR含177,407条；筛选了SR和HR转录组中显著上调表达的EST序列,分别为3,615和1,982条,并对其进行了功能GO分类；发现了仅在SR或HR转录组中表达的特异性EST序列,分别为24,696和37,502条,其中具有生物学分析价值的序列分别为382条和187条,同时对其功能进行了GO分类；筛选了部分SR中的EST序列如生长素相关蛋白、乙烯应答转录因子等,HR中如病程相关蛋白、过氧化物酶基因等进行了实时荧光定量RT-PCR验证；
2.克隆得到了两个辣椒WRKY转录因子CaWRKY30和CaWRKY6, CaWRKY30的全长cDNA1533bp, NCBI GenBank登录号FJ360844,编码区全长1095bp,编码推定的364个氨基酸,分子量大小为41.2kD,等电点6.57; CaWRKY6的全长CDNA1944bp,编码区全长1662bp, NCBI GenBank登录号GQ253367,编码推定的553个氨基酸,分子量大小60.18kD,等电点6.96；
3.克隆得到了CaWRKY30和CaWRKY6基因编码区的基因组DNA序列,CaWRKY30基因编码区DNA全长1743bp,含有3个外显子和2个内含子；CaWRKY6基因编码区DNA全长2530bp,含有6个外显子和5个内含子；Southern杂交证实CaWRKY30和CaWRKY6基因在基因组中都以单拷贝的形式存在,是单拷贝基因；
4. CaWRKY30基因受水杨酸(SA),辣椒疫霉菌、烟草花叶病毒、辣椒青枯菌、非毒性南方根结线虫的诱导而上调表达,但表达受到茉莉酸甲酯(MeJA)的抑制；CaWRKY6基因受辣椒疫霉菌、烟草花叶病毒、辣椒青枯菌、非毒性南方根结线虫的诱导而上调表达,但受茉莉酸甲酯(MeJA)的抑制；建立并优化了CaWRKY6与CaWRKY30基因的原核表达系统；
5.基因的亚细胞定位分析表明CaWRKY30和CaWRKY6蛋白都定位于细胞核,是在细胞核中起转录调控作用的转录因子；
6.构建了含有WRKY基因特异性片段的TRV表达载体,病毒侵染辣椒后RT-PCR验证WRKY基因的沉默效应。CaWRKY30基因沉默的辣椒H1)A149植株接种非毒性南方根结线虫后,辣椒植株的抗性没有改变,根结和卵块数量与对照相比差异不显著,推测这可能与WRKY基因的功能互补、或者沉默不彻底而只需少量转录本就可正常发挥功能的特性有关；CaWRKY6基因沉默的辣椒植株则在根结线虫侵染后,根结和卵块显著增加,CaWRKY6基因的沉默降低了辣椒对根结线虫的抗性。
7.构建了CaWRKY30和CaWRKY6基因分别与pCAMBIA2300融合的植物表达载体,通过农杆菌介导的转化方法分别导入到了番茄MoneyMaker和烟草植株(Nicotiana tabacum L)中,并分别获得了CaWRKY30和CaWRKY6基因的转基因T0代植株；转基因超表达CaWRKY30的番茄和超表达CaWRKY6基因的烟草对表型没有任何影响；转基因超表达CaWRKY30基因的番茄植株对南方根结线虫入侵的敏感性增大,植株的抗性降低,CaWRKY30基因在介导植物的根结线虫应答反应中起负调控作用。
辣椒与根结线虫互作转录组cDNA数据库的构建为深入分析这一互作机制,挖掘抗性相关基因提供了丰富的cDNA资源；同时抗性相关WRKY基因的克隆与功能分析为将来利用WRKY基因的转录调控来创新辣椒的抗线虫育种提供了新的证据,更为利用WRKY基因的来构建高效的抗线虫分子育种途径奠定了基础。
Pepper(Capsicum annuum L.) is one of the most economically important vegetables in China, which is susceptible to the adverse phytopathogens during production, and then leads to yields losses and quality decline. In recent years, the root-knot disease (RKN), one of the plant parasitic nematodes has become a major and seriously disease in pepper production. Due to lack of cultivars with the high-level resistance to RKN, Many methods were used to control this disease, however series of problems on pesticides residues, ecosystem environment pollution and soil deteriorated were brought from highly toxic and high residue chemicals in the field. Thus emergency problems must face up in the vegetable production.
The mechanism on compatible and incompatible interaction between pepper planta and RKN were performed on molecular biology in this study. Moreover relative key resistant genes were isolated and functioned from host plant pepper, the aims to provide novel strategy for pepper resistance-breeding program in the future.
In this paper, to understand the key processes governing resistance mechanisms in pepper HDA149(Capsicum annuum L.), which contains Me3nematode-resistance gene that confers the pepper resistance to the nematodes, upon infection with RKN(Meloidogyne incognita), two trancriptomes, from the root tips infested with virulent and avirulent M. incognita, were sequenced by llumina/Solexa high-throughput deep-sequencing technology. On this basis, two WRKY transcription factors were isolated from the pepper root tissue during the incompatible interaction between pepper HDA149and M. incognita by RACE method. Then, the characterization、expression and function were analysed. The main results are as following:
1. The phenotypic differences were identified after the pepper HDA149were innoculated with virulent and avirulent M. incognita, respectively. Based on this, after trancriptomes were sequenced by llumina/Solexa high-throughput deep-sequencing technology, two transcriptome cDNA databases were constructed, named HR transcriptome and SR transcriptome, respectively. A total of214909ESTs (190213in HR and177407in SR) were clustered by a computational pipeline.24696and37502unigenes were unique expressing in compatible and incompatible, respectively, among of them382in SR and187in HR were selected to future analysis. Among of the commonly transcripts,3,615and1,982unigenes were differentially up-regulation expression in SR and HR transcriptome, respectively. Several genes from the HR sample, i,e., fatty acid hydroperoxide lyase, peroxidase, pathogen-related proteins, and from SR samples,i,e., auxin-induced protein, ethylene-response transcription factor, phenylalanine ammonium-lyase, were selected and their differential expression in the SR and HR samples collected from different time points after nematode inoculation was confirmed by quantity RT-PCR process.
2. Two WRKY transcription factor, named CaWRKY30and CaWRKY6, were isolated from the pepper HDA149root tissue. The nucleotide sequence of CaWRKY30cDNA is1,533bp long and contains an open reading frame (ORF) encoding a polypeptide of364amino acids. The molecular mass of the predicted protein is41.2kDa, and the isoelectric point was calculated to be6.57. The nucleotide sequence of CaWRKY6cDNA is1,944bp long, and contains an ORF encoding a polypeptide of553amino acids. The molecular mass of the predicted protein is60.2kDa and the isoelectric point was calculated to be6.96.
3. The CaWRKY30and CaWRKY6genomic sequences were obtained. The genome DNA length of CaWRKY30was1,743bp, consisted of2introns and3exons, and the genome DNA length of CaWRKY6was2,530bp, contained5introns and6exons. Southern blot demonstrated that CaWRKY30and CaWRKY6only have single copy in pepper genome.
4. The CaWRKY30and CaWRKY6expression patterns were analysied after the pepper plants treatment with varous pathogens and signaling molecules, i,e,. salicylic acid and methyl jasmonate. The CaWRKY30transcripts were up-regulate after the pepper plants innoculated with signaling molecules SA, different pathogens, such as Ralstonia solanacerum, Tobacco mosaic virus (TMV), Phytophthora capsici and avirulent M. incognita, but the transcripts were suppressed by the MeJA. Similarly, the transcripts of CaWRKY6were up-regulate by the SA, R.solanacerum, TMV, P.capsici, avirulent M. incognita, and suppressed by the MeJA and virulent M. incognita. The phylogenetic relationship of CaWRKY proteins were analysis after the consensus maximum parsimony (MP) tree were constructed.
5. we demonstrated that the transiently expressed both the CaWRKY30::GFP and CaWRKY6::GFP fusion proteins were localized exclusively to the nuclei of onion epidermal cells. With the control vector alone, the GFP signal was distributed in both the nucleus and cytoplasm. The nuclear localization of CaWRKY30and CaWRKY6protein supports its role as a transcriptional regulator.
6. TRV vectors which contain specific regions of CaWRKY30and CaWRKY6gene were constructed, and then innoculated to the pepper HDA149, respectively. RT-PCR showed that the virus were successfully infected pepper plants and induced WRKY gene silence. Pepper HDA149which CaWRKY30gene was silenced didn't shown any changes against o the avirulent M.incogntia infection compared to the control, these results may be interpretated by the functional redundancy of WRKY genes or silence is not thorough and lower transcripts of CaWRKY30gene can maintain its roles. In contrast, silenceing of CaWRKY6in pepper HDA149increases the number of root knot and egg masses, suggested that CaWRKY6have an important role in RKN resistance.
7. Plant expresssion vector of pCAMBIA2300-CaWRKY30and pCAMBIA2300-CaWRKY6were constructed, and kanamycin resistant tomato (Moneymaker) plant of CaWRKY30, tobacco plants of CaWRKY6were obtained by Agrobacterium mediated transformation approach. CaWRKY30overexpressing transgenic plants showed increased the growth of aivrulent M. incognita, and more root-knot and egg masses compared to the control were observed after infected with M. incognita in CaWRKY30overexpressing transgenic tomato plants, CaWRKY30protien is a negative regulator of RKN defense.
In a word, comprehensive of the nematode-infested characterization by transcriptome sequencing would provides a rich cDNA resource for further analyzing the function of key regulatory genes involved in nemaotdes resistance in pepper. CaWRKY30and CaWRKY6have an important role in pepper plants responses to pathogens infection, it is a potential way to improve the plant resistance by the use of specific WRKY transcription factor, i,e., CaWRKY30and CaWRKY6.
本研究主要从分子水平上来探索辣椒与南方根结线虫早期亲和与不亲和互作的机制,克隆此过程中的关键调控基因并进行功能分析,以期为辣椒的抗性育种提供新思路。
本研究以含有单显性抗根结线虫基因Me3的辣椒HDA149与南方根结线虫亲和与不亲和互作为基础,采用新一代Solexa高通量测序技术,构建了辣椒与南方根结线虫亲和与不亲和互作的转录组cDNA序列数据库。在此基础上,利用RACE方法从辣椒与根结线虫不亲和互作的辣椒根组织中分离出两个推定的WRKY转录因子,并对两个WRKY基因的分子特征和相关功能进行了研究,实验获得的主要结果如下：
1.鉴定了针对Me3基因不同毒力的南方根结线虫(M.incognita)接种辣椒后的表型差异；在此基础上,建立了辣椒与根结线虫亲和互作转录组(SR)cDNA序列数据库和不亲和互作转录组(SR)的cDNA序列数据库,总共获得了214,909条EST序列,其中HR含190,213条,SR含177,407条；筛选了SR和HR转录组中显著上调表达的EST序列,分别为3,615和1,982条,并对其进行了功能GO分类；发现了仅在SR或HR转录组中表达的特异性EST序列,分别为24,696和37,502条,其中具有生物学分析价值的序列分别为382条和187条,同时对其功能进行了GO分类；筛选了部分SR中的EST序列如生长素相关蛋白、乙烯应答转录因子等,HR中如病程相关蛋白、过氧化物酶基因等进行了实时荧光定量RT-PCR验证；
2.克隆得到了两个辣椒WRKY转录因子CaWRKY30和CaWRKY6, CaWRKY30的全长cDNA1533bp, NCBI GenBank登录号FJ360844,编码区全长1095bp,编码推定的364个氨基酸,分子量大小为41.2kD,等电点6.57; CaWRKY6的全长CDNA1944bp,编码区全长1662bp, NCBI GenBank登录号GQ253367,编码推定的553个氨基酸,分子量大小60.18kD,等电点6.96；
3.克隆得到了CaWRKY30和CaWRKY6基因编码区的基因组DNA序列,CaWRKY30基因编码区DNA全长1743bp,含有3个外显子和2个内含子；CaWRKY6基因编码区DNA全长2530bp,含有6个外显子和5个内含子；Southern杂交证实CaWRKY30和CaWRKY6基因在基因组中都以单拷贝的形式存在,是单拷贝基因；
4. CaWRKY30基因受水杨酸(SA),辣椒疫霉菌、烟草花叶病毒、辣椒青枯菌、非毒性南方根结线虫的诱导而上调表达,但表达受到茉莉酸甲酯(MeJA)的抑制；CaWRKY6基因受辣椒疫霉菌、烟草花叶病毒、辣椒青枯菌、非毒性南方根结线虫的诱导而上调表达,但受茉莉酸甲酯(MeJA)的抑制；建立并优化了CaWRKY6与CaWRKY30基因的原核表达系统；
5.基因的亚细胞定位分析表明CaWRKY30和CaWRKY6蛋白都定位于细胞核,是在细胞核中起转录调控作用的转录因子；
6.构建了含有WRKY基因特异性片段的TRV表达载体,病毒侵染辣椒后RT-PCR验证WRKY基因的沉默效应。CaWRKY30基因沉默的辣椒H1)A149植株接种非毒性南方根结线虫后,辣椒植株的抗性没有改变,根结和卵块数量与对照相比差异不显著,推测这可能与WRKY基因的功能互补、或者沉默不彻底而只需少量转录本就可正常发挥功能的特性有关；CaWRKY6基因沉默的辣椒植株则在根结线虫侵染后,根结和卵块显著增加,CaWRKY6基因的沉默降低了辣椒对根结线虫的抗性。
7.构建了CaWRKY30和CaWRKY6基因分别与pCAMBIA2300融合的植物表达载体,通过农杆菌介导的转化方法分别导入到了番茄MoneyMaker和烟草植株(Nicotiana tabacum L)中,并分别获得了CaWRKY30和CaWRKY6基因的转基因T0代植株；转基因超表达CaWRKY30的番茄和超表达CaWRKY6基因的烟草对表型没有任何影响；转基因超表达CaWRKY30基因的番茄植株对南方根结线虫入侵的敏感性增大,植株的抗性降低,CaWRKY30基因在介导植物的根结线虫应答反应中起负调控作用。
辣椒与根结线虫互作转录组cDNA数据库的构建为深入分析这一互作机制,挖掘抗性相关基因提供了丰富的cDNA资源；同时抗性相关WRKY基因的克隆与功能分析为将来利用WRKY基因的转录调控来创新辣椒的抗线虫育种提供了新的证据,更为利用WRKY基因的来构建高效的抗线虫分子育种途径奠定了基础。
Pepper(Capsicum annuum L.) is one of the most economically important vegetables in China, which is susceptible to the adverse phytopathogens during production, and then leads to yields losses and quality decline. In recent years, the root-knot disease (RKN), one of the plant parasitic nematodes has become a major and seriously disease in pepper production. Due to lack of cultivars with the high-level resistance to RKN, Many methods were used to control this disease, however series of problems on pesticides residues, ecosystem environment pollution and soil deteriorated were brought from highly toxic and high residue chemicals in the field. Thus emergency problems must face up in the vegetable production.
The mechanism on compatible and incompatible interaction between pepper planta and RKN were performed on molecular biology in this study. Moreover relative key resistant genes were isolated and functioned from host plant pepper, the aims to provide novel strategy for pepper resistance-breeding program in the future.
In this paper, to understand the key processes governing resistance mechanisms in pepper HDA149(Capsicum annuum L.), which contains Me3nematode-resistance gene that confers the pepper resistance to the nematodes, upon infection with RKN(Meloidogyne incognita), two trancriptomes, from the root tips infested with virulent and avirulent M. incognita, were sequenced by llumina/Solexa high-throughput deep-sequencing technology. On this basis, two WRKY transcription factors were isolated from the pepper root tissue during the incompatible interaction between pepper HDA149and M. incognita by RACE method. Then, the characterization、expression and function were analysed. The main results are as following:
1. The phenotypic differences were identified after the pepper HDA149were innoculated with virulent and avirulent M. incognita, respectively. Based on this, after trancriptomes were sequenced by llumina/Solexa high-throughput deep-sequencing technology, two transcriptome cDNA databases were constructed, named HR transcriptome and SR transcriptome, respectively. A total of214909ESTs (190213in HR and177407in SR) were clustered by a computational pipeline.24696and37502unigenes were unique expressing in compatible and incompatible, respectively, among of them382in SR and187in HR were selected to future analysis. Among of the commonly transcripts,3,615and1,982unigenes were differentially up-regulation expression in SR and HR transcriptome, respectively. Several genes from the HR sample, i,e., fatty acid hydroperoxide lyase, peroxidase, pathogen-related proteins, and from SR samples,i,e., auxin-induced protein, ethylene-response transcription factor, phenylalanine ammonium-lyase, were selected and their differential expression in the SR and HR samples collected from different time points after nematode inoculation was confirmed by quantity RT-PCR process.
2. Two WRKY transcription factor, named CaWRKY30and CaWRKY6, were isolated from the pepper HDA149root tissue. The nucleotide sequence of CaWRKY30cDNA is1,533bp long and contains an open reading frame (ORF) encoding a polypeptide of364amino acids. The molecular mass of the predicted protein is41.2kDa, and the isoelectric point was calculated to be6.57. The nucleotide sequence of CaWRKY6cDNA is1,944bp long, and contains an ORF encoding a polypeptide of553amino acids. The molecular mass of the predicted protein is60.2kDa and the isoelectric point was calculated to be6.96.
3. The CaWRKY30and CaWRKY6genomic sequences were obtained. The genome DNA length of CaWRKY30was1,743bp, consisted of2introns and3exons, and the genome DNA length of CaWRKY6was2,530bp, contained5introns and6exons. Southern blot demonstrated that CaWRKY30and CaWRKY6only have single copy in pepper genome.
4. The CaWRKY30and CaWRKY6expression patterns were analysied after the pepper plants treatment with varous pathogens and signaling molecules, i,e,. salicylic acid and methyl jasmonate. The CaWRKY30transcripts were up-regulate after the pepper plants innoculated with signaling molecules SA, different pathogens, such as Ralstonia solanacerum, Tobacco mosaic virus (TMV), Phytophthora capsici and avirulent M. incognita, but the transcripts were suppressed by the MeJA. Similarly, the transcripts of CaWRKY6were up-regulate by the SA, R.solanacerum, TMV, P.capsici, avirulent M. incognita, and suppressed by the MeJA and virulent M. incognita. The phylogenetic relationship of CaWRKY proteins were analysis after the consensus maximum parsimony (MP) tree were constructed.
5. we demonstrated that the transiently expressed both the CaWRKY30::GFP and CaWRKY6::GFP fusion proteins were localized exclusively to the nuclei of onion epidermal cells. With the control vector alone, the GFP signal was distributed in both the nucleus and cytoplasm. The nuclear localization of CaWRKY30and CaWRKY6protein supports its role as a transcriptional regulator.
6. TRV vectors which contain specific regions of CaWRKY30and CaWRKY6gene were constructed, and then innoculated to the pepper HDA149, respectively. RT-PCR showed that the virus were successfully infected pepper plants and induced WRKY gene silence. Pepper HDA149which CaWRKY30gene was silenced didn't shown any changes against o the avirulent M.incogntia infection compared to the control, these results may be interpretated by the functional redundancy of WRKY genes or silence is not thorough and lower transcripts of CaWRKY30gene can maintain its roles. In contrast, silenceing of CaWRKY6in pepper HDA149increases the number of root knot and egg masses, suggested that CaWRKY6have an important role in RKN resistance.
7. Plant expresssion vector of pCAMBIA2300-CaWRKY30and pCAMBIA2300-CaWRKY6were constructed, and kanamycin resistant tomato (Moneymaker) plant of CaWRKY30, tobacco plants of CaWRKY6were obtained by Agrobacterium mediated transformation approach. CaWRKY30overexpressing transgenic plants showed increased the growth of aivrulent M. incognita, and more root-knot and egg masses compared to the control were observed after infected with M. incognita in CaWRKY30overexpressing transgenic tomato plants, CaWRKY30protien is a negative regulator of RKN defense.
In a word, comprehensive of the nematode-infested characterization by transcriptome sequencing would provides a rich cDNA resource for further analyzing the function of key regulatory genes involved in nemaotdes resistance in pepper. CaWRKY30and CaWRKY6have an important role in pepper plants responses to pathogens infection, it is a potential way to improve the plant resistance by the use of specific WRKY transcription factor, i,e., CaWRKY30and CaWRKY6.
引文
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