棉花与黄萎病菌的分子互作机制研究及GbWRKY1基因的功能鉴定
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摘要
棉花黄萎病是由大丽轮枝菌引起的土传真菌维管束病害,发病时会引起棉花叶片失绿变黄,蕾铃脱落,严重时造成棉株死亡,是当前影响棉花生产的主要病害之一。了解棉花与黄萎病菌互作的分子机制,分离抗病相关基因并利用基因工程技术对现有陆地棉品种进行抗性改造,已成为当前棉花分子生物学研究的重点内容之一本研究分别构建了抗黄萎病的海岛棉‘海7124’接种黄萎病菌后的抑制差减杂交cDNA文库以及通过大规模测序的RNA-Seq技术研究棉花抗黄萎病机制。通过基因表达分析,获得了大批棉花抗黄萎病相关基因,并对其中的一个WRKY类转录因子进行了较为详细的功能研究,取得的主要结果如下：1、构建了‘海7124’受黄萎病菌接种后的抑制差减杂交cDNA文库。对得到的211个差异表达基因功能分类表明主要包括代谢类相关基因、生物及非生物胁迫反应相关基因、细胞结构以及信号传导相关基因等,此外有38%左右的基因未在数据库中得到功能注释或者功能注释为未知功能蛋白。一些已知的植物防卫反应相关的基因,如PR蛋白、氧爆发相关基因以及一些次生代谢相关的基因等在‘海7124’接种黄萎病菌后受诱导表达明显。此外一些参与基因表达调控类基因,如WRKY类转录因子、MAPK类蛋白激酶等信号传导相关的基因也在棉花抗黄萎病反应中差异表达明显。对这些参与信号转导和表达调控类基因的详细分析会将有助于深入了解棉花抗黄萎病病反应的分子机制。运用Nothern杂交、RT-PCR和qRT-PCR技术对分离得到的这些差异表达基因进行验证,并对它们在一些信号分子,如乙烯、茉莉酸、水杨酸及H202等处理后的表达模式进行分析,推测棉花抗黄萎病反应所参与的信号路径。研究发现,乙烯作为重要的一类抗病信号分子可能在棉花抗黄萎病反应中具有重要作用。
2、为了进一步了解棉花抗黄萎病分子机制,我们利用RNA-Seq大规模测序分析技术对‘海7124’接种黄萎病菌后4h、12h、24h和48h的不同时间点取样研究参与棉花抗黄萎病反应的基因表达变化。通过数据库筛选及比对分析,共获得3442个差异表达基因。为了验证RNA-Seq大规模测序表达谱分析结果的准确性,我们选取了不同功能分类、不同表达模式的361个基因进行qRT-PCR分析,结果表明两种基因表达模式结果基本吻合,可对表达谱数据进行深入分析。对这些差异表达基因进行GO功能富集分析发现,它们主要参与代谢类相关途径和应对外界的刺激相关反应。我们对其中参与植物抗病免疫反应信号传导途径的四类基因包括富含亮氨酸重复的类受体激酶、激素信号路径相关基因、转录调控基因及次生代谢苯丙环类路径相关基因进行了深入分析。研究表明大多数苯丙环类物质代谢路径基因都参与了棉花与黄萎病菌的分子互作,并在抗病的‘海7124’和感病的陆地棉品系‘YZ-1’中均受黄萎病菌诱导,但在抗病材料中这些基因被诱导的速度和表达水平都高于感病的陆地棉。酶活检测的结果与基因表达分析的结果相似。采用组织化学检测的方法对接菌前后‘海7124’和‘YZ-1’的木质素沉积和维管结构的变化进行了比较分析,结果显示抗病材料的木质素合成增强并在次生壁中的积累和沉积使得维管结构增强了对病原菌的抗性,在海岛棉对黄萎病的抗性中起着重要作用。
3、GbWRKY1是从海岛棉中克隆得到的一个受黄萎病菌诱导的WRKY类转录因子。通过序列比对发现,GbWRKY1与拟南芥AtWRKY75相似性较高,同属于第二类C亚组的WRKY蛋白。通过在拟南芥中超量表达该基因发现转基因拟南芥能有效缓解对低磷逆境胁迫的反应,包括减少花青素的积累和更多的生物学产量。研究还表明,在正常条件下转基因拟南芥的侧根数增多、磷酸酶活性增强以及植株体内无机磷和总磷含量增加。对一系列受磷饥饿诱导基因表达研究发现,无论是磷转运子基因还是磷饥饿诱导相关基因,在低磷处理后转基因拟南芥中受诱导程度都低于野生型。由此推论,GbWRKY1可能是缓解磷饥饿胁迫信号传导中的一个正调控因子,在拟南芥中该基因的超量表达能在正常条件下模拟磷饥饿的信号,激活植物的低磷反应,包括侧根增多,体内积累吸收更多的磷,从而能在低磷处理下能够缓解植物的低磷胁迫反应。由于生长素对低磷胁迫反应有重要的调控机制,我们对超量表达GbWRKY1转基因拟南芥对外源生长素处理的响应进行了分析,并对其内源生长素含量和生长素响应的相关基因表达进行了检测。但结果显示转基因与野生型拟南芥在内源生长素含量上没有显著差异,而转基因植株对生长素的敏感性增强。GbWRKY1是磷饥饿反应的正调控因子,可能通过增强生长素响应信号路径和促进拟南芥侧根发生来调控拟南芥的低磷胁迫反应。GbWRKY1在棉花中受黄萎病菌诱导表达,它在棉花抗黄萎病反应中的作用还有待进一步研究。
Cotton Verticillium wilt caused by Verticillium dahliae is a soil-borne vascular disease. The representative symptoms caused by V. dahliae in the susceptible cotton include leaf curl, necrosis and defoliation. The disease has become the most economically important disease of cotton and severe outbreak of this disease causes reduction in fibre quality and significant yield losses. Through research on the molecular mechanism during the cotton defense response to V. dahliae, and identification of defense related genes for the use of genetic engineering methods to cultivate resistant cotton varieties will be the main direction of future cotton breeding. In this study, a suppression subtractive hybridization (SSH) based cDNA library constructed from V. dahliae inoculated Gossypium barbadense variety'Hai7124'and deep-sequencing technology based RNA-Seq were developed to better understand the molecular mechanism in cotton defense response to V. dahliae. Through gene expression analysis, a number of defense related genes were identified. Detailed functional identification of a defense related WRKY-type transcription factor isolated from the SSH cDNA library was studied in Arabidopsis. The main results of this study were as follow:
A suppression subtractive hybridization (SSH) based cDNA library was constructed from V. dahliae inoculated G. barbadense variety'Hai7124'. A total of211unique genes were differentially identified and classified into11functional categories. The largest groups contain genes involved in metabolism, stress/defence response, cell structure and signal transduction. More than one-third of the genes (38%) were identified as unknown classification or function. This study identified a set of disease-related genes involved in the process of the response, including pathogenesis-related genes of various classes, oxidative burst-related genes and secondary metabolism-related genes, which were better understand in the disease resistance. The characterization of some transcription factors and kinases enabled us to better understand the defence mechanisms. Northern blot analysis and quantitative real-time PCR (qPCR) were performed to investigate the expression patterns of some representative genes and characterize the role of some signal molecules (H2O2, ethylene, jasmonic acid and salicylic acid) in the cotton defence response. Our results suggested that a complicated and concerted mechanism involving multiple pathways including salicylic acid, jasmonic acid and ethylene was responsible for the cotton defence response to V. dahliae. The results suggested that ethylene play a putative role in the resistant response as a signal molecule in the cotton defense response.
The incompatible pathosystem between resistant cotton (Gossypium barbadense cv.'Hai7124') and V. dahliae strain V991was used to study the cotton transcriptome changes after pathogen inoculation by RNA-Seq. Of32(?)774genes detected by mapping the tags to assembly cotton contigs,3442defence-responsive genes were identified. Gene cluster analyses and functional assignments of differentially expressed genes indicated a significant transcriptional complexity. Quantitative real-time PCR (qPCR) was performed on selected genes with different expression levels and functional assignments to demonstrate the utility of RNA-Seq for gene expression profiles during the cotton defence response. Detailed elucidation of responses of leucine-rich repeat receptor-like kinases (LRR-RLKs), phytohormone signalling-related genes, and transcription factors described the interplay of signals that allowed the plant to fine-tune defence responses. On the basis of global gene regulation of phenylpropanoid metabolism-related genes, phenylpropanoid metabolism was deduced to be involved in the cotton defence response. A closer look at the expression of these genes, enzyme activity, and lignin levels revealed differences between'Hai7124' and 'YZ-1'. Both types of plants showed an increased level of expression of lignin synthesis-related genes and increased phenylalanine-ammonia lyase (PAL) and (POD) enzyme activity after inoculation with V. dahliae, but the increase was greater and faster in the'Hai7124'. Histochemical analysis of lignin revealed that'Hai7124'not only retains its vascular structure, but also accumulates high levels of lignin. Furthermore, quantitative analysis demonstrated increased lignification and cross-linking of lignin in stem of'Hai7124'. Overall, a critical role for lignin was believed to contribute to the cotton disease resistance to V. dahliae.
GbWRKY1is a cloned WRKY transcription factor from Gossypium barbadense cv.'Hai7124', which was firstly identified as defense related gene and showed highest similarity with AtWRKY75in Arabidopsis thaliana which was assigned to class Ⅱc of WRKY family. Overexpression of GbWRKYl in Arabidopsis resulted of attenuated Pi starvation stress response, including reduced accumulation of anthocyanin and more biological yield. Moreover, overexpression plants exhibited relative high level of total P and Pi as well as enhanced accumulation of acid phosphatases ativity. Genes encoding highaffinity Pi transporters (PT1, PT2, and PHT1;8) and phosphatases (PS1, ACP5, and RNS1) were upregulated in overexpression plants compared to WT plants under Pi-sufficient condition. The result demonstrated that GbWRKY1may be act as a positive regulator involved in the global Pi starvation response. Together with the fact that AtWRKY75was only induced upon Pi starvation, the results suggested that the overexpression of GbWRKY1caused plants exhibited a Pi starvation response in the Pi-sufficient condition, including enhanced number of LR and accmulation more Pi, which may attenuate Pi starvation stress response of overexpression plants. To explore whether the alterations of root system architecture and enhanced Pi signaling response in the overexpression plants by modulating the auxin signaling response, the response to exogenous IAA treatment was investigated. Together with the result of internal IAA concentration and gene expression level of TIR1and ARF19, it seemed that the overexpression plants was more sensitivity to exogenous auxin under Pi-sufficient conditions compared with the WT. Changes in the auxin sensitivity of the root seemed to be involved in the developmental response of the Arabidopsis root system to Pi starvation. Overall, the involvement of GbWRKY1in the modification of the root system by changed auxin sensitivity to regulate Pi starvation responses. The expression of GbWRKY1was induced by V. dahliae inoculation in cotton. The putative role of GbWRKY1in the cotton defense response still need further study.
2、为了进一步了解棉花抗黄萎病分子机制,我们利用RNA-Seq大规模测序分析技术对‘海7124’接种黄萎病菌后4h、12h、24h和48h的不同时间点取样研究参与棉花抗黄萎病反应的基因表达变化。通过数据库筛选及比对分析,共获得3442个差异表达基因。为了验证RNA-Seq大规模测序表达谱分析结果的准确性,我们选取了不同功能分类、不同表达模式的361个基因进行qRT-PCR分析,结果表明两种基因表达模式结果基本吻合,可对表达谱数据进行深入分析。对这些差异表达基因进行GO功能富集分析发现,它们主要参与代谢类相关途径和应对外界的刺激相关反应。我们对其中参与植物抗病免疫反应信号传导途径的四类基因包括富含亮氨酸重复的类受体激酶、激素信号路径相关基因、转录调控基因及次生代谢苯丙环类路径相关基因进行了深入分析。研究表明大多数苯丙环类物质代谢路径基因都参与了棉花与黄萎病菌的分子互作,并在抗病的‘海7124’和感病的陆地棉品系‘YZ-1’中均受黄萎病菌诱导,但在抗病材料中这些基因被诱导的速度和表达水平都高于感病的陆地棉。酶活检测的结果与基因表达分析的结果相似。采用组织化学检测的方法对接菌前后‘海7124’和‘YZ-1’的木质素沉积和维管结构的变化进行了比较分析,结果显示抗病材料的木质素合成增强并在次生壁中的积累和沉积使得维管结构增强了对病原菌的抗性,在海岛棉对黄萎病的抗性中起着重要作用。
3、GbWRKY1是从海岛棉中克隆得到的一个受黄萎病菌诱导的WRKY类转录因子。通过序列比对发现,GbWRKY1与拟南芥AtWRKY75相似性较高,同属于第二类C亚组的WRKY蛋白。通过在拟南芥中超量表达该基因发现转基因拟南芥能有效缓解对低磷逆境胁迫的反应,包括减少花青素的积累和更多的生物学产量。研究还表明,在正常条件下转基因拟南芥的侧根数增多、磷酸酶活性增强以及植株体内无机磷和总磷含量增加。对一系列受磷饥饿诱导基因表达研究发现,无论是磷转运子基因还是磷饥饿诱导相关基因,在低磷处理后转基因拟南芥中受诱导程度都低于野生型。由此推论,GbWRKY1可能是缓解磷饥饿胁迫信号传导中的一个正调控因子,在拟南芥中该基因的超量表达能在正常条件下模拟磷饥饿的信号,激活植物的低磷反应,包括侧根增多,体内积累吸收更多的磷,从而能在低磷处理下能够缓解植物的低磷胁迫反应。由于生长素对低磷胁迫反应有重要的调控机制,我们对超量表达GbWRKY1转基因拟南芥对外源生长素处理的响应进行了分析,并对其内源生长素含量和生长素响应的相关基因表达进行了检测。但结果显示转基因与野生型拟南芥在内源生长素含量上没有显著差异,而转基因植株对生长素的敏感性增强。GbWRKY1是磷饥饿反应的正调控因子,可能通过增强生长素响应信号路径和促进拟南芥侧根发生来调控拟南芥的低磷胁迫反应。GbWRKY1在棉花中受黄萎病菌诱导表达,它在棉花抗黄萎病反应中的作用还有待进一步研究。
Cotton Verticillium wilt caused by Verticillium dahliae is a soil-borne vascular disease. The representative symptoms caused by V. dahliae in the susceptible cotton include leaf curl, necrosis and defoliation. The disease has become the most economically important disease of cotton and severe outbreak of this disease causes reduction in fibre quality and significant yield losses. Through research on the molecular mechanism during the cotton defense response to V. dahliae, and identification of defense related genes for the use of genetic engineering methods to cultivate resistant cotton varieties will be the main direction of future cotton breeding. In this study, a suppression subtractive hybridization (SSH) based cDNA library constructed from V. dahliae inoculated Gossypium barbadense variety'Hai7124'and deep-sequencing technology based RNA-Seq were developed to better understand the molecular mechanism in cotton defense response to V. dahliae. Through gene expression analysis, a number of defense related genes were identified. Detailed functional identification of a defense related WRKY-type transcription factor isolated from the SSH cDNA library was studied in Arabidopsis. The main results of this study were as follow:
A suppression subtractive hybridization (SSH) based cDNA library was constructed from V. dahliae inoculated G. barbadense variety'Hai7124'. A total of211unique genes were differentially identified and classified into11functional categories. The largest groups contain genes involved in metabolism, stress/defence response, cell structure and signal transduction. More than one-third of the genes (38%) were identified as unknown classification or function. This study identified a set of disease-related genes involved in the process of the response, including pathogenesis-related genes of various classes, oxidative burst-related genes and secondary metabolism-related genes, which were better understand in the disease resistance. The characterization of some transcription factors and kinases enabled us to better understand the defence mechanisms. Northern blot analysis and quantitative real-time PCR (qPCR) were performed to investigate the expression patterns of some representative genes and characterize the role of some signal molecules (H2O2, ethylene, jasmonic acid and salicylic acid) in the cotton defence response. Our results suggested that a complicated and concerted mechanism involving multiple pathways including salicylic acid, jasmonic acid and ethylene was responsible for the cotton defence response to V. dahliae. The results suggested that ethylene play a putative role in the resistant response as a signal molecule in the cotton defense response.
The incompatible pathosystem between resistant cotton (Gossypium barbadense cv.'Hai7124') and V. dahliae strain V991was used to study the cotton transcriptome changes after pathogen inoculation by RNA-Seq. Of32(?)774genes detected by mapping the tags to assembly cotton contigs,3442defence-responsive genes were identified. Gene cluster analyses and functional assignments of differentially expressed genes indicated a significant transcriptional complexity. Quantitative real-time PCR (qPCR) was performed on selected genes with different expression levels and functional assignments to demonstrate the utility of RNA-Seq for gene expression profiles during the cotton defence response. Detailed elucidation of responses of leucine-rich repeat receptor-like kinases (LRR-RLKs), phytohormone signalling-related genes, and transcription factors described the interplay of signals that allowed the plant to fine-tune defence responses. On the basis of global gene regulation of phenylpropanoid metabolism-related genes, phenylpropanoid metabolism was deduced to be involved in the cotton defence response. A closer look at the expression of these genes, enzyme activity, and lignin levels revealed differences between'Hai7124' and 'YZ-1'. Both types of plants showed an increased level of expression of lignin synthesis-related genes and increased phenylalanine-ammonia lyase (PAL) and (POD) enzyme activity after inoculation with V. dahliae, but the increase was greater and faster in the'Hai7124'. Histochemical analysis of lignin revealed that'Hai7124'not only retains its vascular structure, but also accumulates high levels of lignin. Furthermore, quantitative analysis demonstrated increased lignification and cross-linking of lignin in stem of'Hai7124'. Overall, a critical role for lignin was believed to contribute to the cotton disease resistance to V. dahliae.
GbWRKY1is a cloned WRKY transcription factor from Gossypium barbadense cv.'Hai7124', which was firstly identified as defense related gene and showed highest similarity with AtWRKY75in Arabidopsis thaliana which was assigned to class Ⅱc of WRKY family. Overexpression of GbWRKYl in Arabidopsis resulted of attenuated Pi starvation stress response, including reduced accumulation of anthocyanin and more biological yield. Moreover, overexpression plants exhibited relative high level of total P and Pi as well as enhanced accumulation of acid phosphatases ativity. Genes encoding highaffinity Pi transporters (PT1, PT2, and PHT1;8) and phosphatases (PS1, ACP5, and RNS1) were upregulated in overexpression plants compared to WT plants under Pi-sufficient condition. The result demonstrated that GbWRKY1may be act as a positive regulator involved in the global Pi starvation response. Together with the fact that AtWRKY75was only induced upon Pi starvation, the results suggested that the overexpression of GbWRKY1caused plants exhibited a Pi starvation response in the Pi-sufficient condition, including enhanced number of LR and accmulation more Pi, which may attenuate Pi starvation stress response of overexpression plants. To explore whether the alterations of root system architecture and enhanced Pi signaling response in the overexpression plants by modulating the auxin signaling response, the response to exogenous IAA treatment was investigated. Together with the result of internal IAA concentration and gene expression level of TIR1and ARF19, it seemed that the overexpression plants was more sensitivity to exogenous auxin under Pi-sufficient conditions compared with the WT. Changes in the auxin sensitivity of the root seemed to be involved in the developmental response of the Arabidopsis root system to Pi starvation. Overall, the involvement of GbWRKY1in the modification of the root system by changed auxin sensitivity to regulate Pi starvation responses. The expression of GbWRKY1was induced by V. dahliae inoculation in cotton. The putative role of GbWRKY1in the cotton defense response still need further study.
引文
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