转hrf1基因水稻抗白叶枯病机理研究及水稻抗病相关基因Oscyp71Z2的功能分析
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摘要
水稻白叶枯病是由黄单胞病菌侵染引起的水稻三大病害之一,每年造成10%到80%的产量损失,威胁着世界粮食安全。目前,大量使用化学农药已经严重破坏了生态环境。同时,水稻品种抗性易于丧失对水稻抗病育种提出更高的要求。经济有效且环境友好的控制病害途径之一就是利用生物技术创制持久的、广谱抗性水稻栽培品种。而鉴定水稻抗病相关基因是培育抗病水稻新品种的基础。进一步发掘具有潜在利用价值的激发子蛋白编码基因为通过改善水稻自身防御体系来控制病害提供了可利用的资源。
来源于白叶枯菌株JXOⅢ的harpinxoo蛋白通过激活多重防卫反应,如系统获得抗病性(system acquired resistance, SAR)、过敏反应(hypersensitive response, HR),从而增强植物对多种病害的抗性。然而,仍旧不清楚植保素以及信号分子活性氧是否也在harpinxoo蛋白所激发的水稻抗病信号通路中起作用。在前期研究工作中,本实验室获得的部分转编码harpinXoo基因hrfl水稻T3代纯合体增强了对白叶枯病的抗性。在此基础之上,本研究选择转hrfl基因水稻T3代纯合体NJH12用来分析hrfl异源表达增强水稻白叶枯病抗性的机理。本研究表明,在水稻中异源表达harpinxoo蛋白编码基因hrf1激发了植保素生物合成通路,同时抑制了内源H202产生,并增强了对白叶枯病菌的抗性。在水稻中异源表达hrfl基因显著诱导植保素合成相关基因表达,促使快速持续地大量合成植保素。随着抗氧化酶活性的变化,转基因系的ROS-清除能力升高,H202量显著减少。通过扫描电镜和能谱分析,转基因水稻的硅含量显著升高,并且有部分的硅质体的分布也发生变化。在接菌的条件下,转基因系的防卫反应相关基因的表达量也显著升高。这些结果为更好地理解防卫反应在hrfl介导的白叶枯病抗性中所起到的作用提供了证据,并且也为获得非特异性病害抗性转基因植物提供了可利用的资源。
从转hrf1基因水稻NJH12芯片数据中筛选到一个表达倍数高达114.6倍的抗病相关基因。随后经生物信息学分析该基因是一个细胞色素P450基因,命名为Oscyp71Z2,是CYP71Z亚家族成员之一。从水稻品种日本晴中克隆Oscyp71Z2全长基因、Oscyp71Z2RNAi靶片段以及启动子上游序列分别构建超量表达载体、RNAi载体以及启动子/GUS融合载体,利用农杆菌介导法转化水稻日本晴。常规PCR和Southern blot结果表明T-DNA区已整合到水稻基因组中。Northern blot和Western blot结果表明超量表达株系Oscyp71Z2的转录本和蛋白量较野生型高,RNAi株系中Oscyp71Z2的蛋白量较野生型低,而转录本却都未检测到。GUS活性实验表明该基因主要在叶片、节、主根和颖片中表达。本研究表明,Oscyp71Z2超量表达株系在孕穗期减少了白叶枯病菌的病斑面积,并抑制了白叶枯病菌在水稻叶片中的定殖。同时,随着植保素合成相关基因的表达量升高,超量表达株系的植保素迅速、大量地积累。进一步分析活性氧清除能力的升高,导致超量表达株系的H202产生受到抑制。另一方面,Oscyp71Z2RNAi株系对白叶枯病菌抗性与野生型没有显著差异,但部分抑制了植保素积累。这些结果表明,Oscyp71Z2可能通过调控植保素和H202生物合成通路从而在水稻白叶枯病菌不亲合互作中起到重要作用。
Bacterial blight is one of the most destructive rice diseases, which caused by Xanthomonas oryzae pv. oryzae (Xoo), and results in10%to80%yield losses, endangering worldwide food security. Presently heavily application of chemical pesticides for controling disease has seriously damaged ecological environment. Meanwhile, easy lost of rice variety resistance to disease posed higher demond for rice breeding. An economically effective and environmentally sound approach to control disease is the utilization of cultivars that possessed durable, broad-spectrum resistance to disease by biotechnology. Identification of rice resistance-related genes is the base to cultivate the new rice disease-resistant varieties. The exploration of gene code elicitor protein with potential application value provides available resource for controlling disease by inducing multiple defense responses.
Harpinxoo proteins from the Xoo stain JXOIII increase plant disease resistance by activating multiple defense responses in plants, such as systemic acquired resistance and hypersensitive response. However, it is unclear whether phytoalexin production and ROS burst are involved in the disease resistance conferred by the expression of the harpinxoo protein in rice. In preliminary studies, our group showed some hrfl (code harpinxoo protein)-transformed rice T3homozygote increased resistance to Xoo. On the basis, in this article hrfl-transformed rice T3homozygote NJH12has been choiced for analyzing the resistance mechanism to Xoo. In this article, ectopic expression of hrfl in rice enhanced resistance to bacterial blight. Accompanying the activation of genes related to the phytoalexin biosynthesis pathway in hrfl-transformed rice, phytoalexins quickly and consistently accumulated concurrent with the limitation of bacterial growth rate. Moreover, the hrfl-transformed rice showed an increased ability for ROS scavenging and decreased hydrogen peroxide (H2O2) concentration. Furthermore, the localization and relative quantification of silicon deposition in rice leaves was detected by scanning electron microscopy (SEM) and energy-dispersive X-ray spectrometer (EDS). Finally, the transcript levels of defense response genes increased in transformed rice. These data are significant because they provide evidence for a better understanding the role of defense responses in the incompatible interaction between Xoo and hrf1-transformed plants. The data also supply an opportunity for generating nonspecific resistance to pathogens.
Based on the transcriptome profile of the transgenic rice line NJH12expressing hrfl from Xoo, a resistance-related gene showing114.6-fold increase of expression level was identified. The gene is the cytochrome P450gene, named as Oscyp71Z2analyzed by bioinformation, which belongs to the CYP71Z subfamily. Oscyp71Z2CDS, Oscyp71Z2RNAi target fragment and promoter sequence have cloned from rice cultivar Nipponbare transcript and genome, respectively make transformation vectors for overexpression, RNAi and Oscyp71Z2expression pattern, then transformed into rice Nipponbare genome mediated by Agrobacterium tumefaciens. The results from normal PCR and Southern blot showed T-DNA has inserted into rice genome. The data from Northern blot and Western blot showed the transcript and protein level in Oscyp71Z2-overexpressiong plant was higher than that in wild type Nipponbare. On the contrary, the protein level in Oscyp71Z-RNAi plant was lower than that in wild type, the tanscript hasa no detected in both. A strong GUS signal was detected mainly in the leaves, nodes, primary roots and glume. Overexpression of Oscyp71Z2enhanced resistance to Xoo at the booting stage, and inhibited multiplication of Xoo PXO99A. The accumulation of phytoalexins was rapidly and strongly induced in Oscyp71Z2-overexpressing plants, and the transcript levels of genes related to the phytoalexin biosynthesis pathway were elevated. The H2O2concentration in Oscyp71Z2-overexpressing plants was reduced in accordance with the increase in ROS-scavenging ability due to the induction of superoxide dismutase (SOD) as well as peroxidase (POD) and catalase (CAT) activation. We also showed that suppression of Oscyp71Z2had no significantly effect on disease resistance to Xoo in rice, although inhibited part phytoalexin production. These results demonstrated that Oscyp71Z2plays an important role for bacterial blight resistance by regulating the diterpenoid phytoalexin biosynthesis and H2O2generation.
来源于白叶枯菌株JXOⅢ的harpinxoo蛋白通过激活多重防卫反应,如系统获得抗病性(system acquired resistance, SAR)、过敏反应(hypersensitive response, HR),从而增强植物对多种病害的抗性。然而,仍旧不清楚植保素以及信号分子活性氧是否也在harpinxoo蛋白所激发的水稻抗病信号通路中起作用。在前期研究工作中,本实验室获得的部分转编码harpinXoo基因hrfl水稻T3代纯合体增强了对白叶枯病的抗性。在此基础之上,本研究选择转hrfl基因水稻T3代纯合体NJH12用来分析hrfl异源表达增强水稻白叶枯病抗性的机理。本研究表明,在水稻中异源表达harpinxoo蛋白编码基因hrf1激发了植保素生物合成通路,同时抑制了内源H202产生,并增强了对白叶枯病菌的抗性。在水稻中异源表达hrfl基因显著诱导植保素合成相关基因表达,促使快速持续地大量合成植保素。随着抗氧化酶活性的变化,转基因系的ROS-清除能力升高,H202量显著减少。通过扫描电镜和能谱分析,转基因水稻的硅含量显著升高,并且有部分的硅质体的分布也发生变化。在接菌的条件下,转基因系的防卫反应相关基因的表达量也显著升高。这些结果为更好地理解防卫反应在hrfl介导的白叶枯病抗性中所起到的作用提供了证据,并且也为获得非特异性病害抗性转基因植物提供了可利用的资源。
从转hrf1基因水稻NJH12芯片数据中筛选到一个表达倍数高达114.6倍的抗病相关基因。随后经生物信息学分析该基因是一个细胞色素P450基因,命名为Oscyp71Z2,是CYP71Z亚家族成员之一。从水稻品种日本晴中克隆Oscyp71Z2全长基因、Oscyp71Z2RNAi靶片段以及启动子上游序列分别构建超量表达载体、RNAi载体以及启动子/GUS融合载体,利用农杆菌介导法转化水稻日本晴。常规PCR和Southern blot结果表明T-DNA区已整合到水稻基因组中。Northern blot和Western blot结果表明超量表达株系Oscyp71Z2的转录本和蛋白量较野生型高,RNAi株系中Oscyp71Z2的蛋白量较野生型低,而转录本却都未检测到。GUS活性实验表明该基因主要在叶片、节、主根和颖片中表达。本研究表明,Oscyp71Z2超量表达株系在孕穗期减少了白叶枯病菌的病斑面积,并抑制了白叶枯病菌在水稻叶片中的定殖。同时,随着植保素合成相关基因的表达量升高,超量表达株系的植保素迅速、大量地积累。进一步分析活性氧清除能力的升高,导致超量表达株系的H202产生受到抑制。另一方面,Oscyp71Z2RNAi株系对白叶枯病菌抗性与野生型没有显著差异,但部分抑制了植保素积累。这些结果表明,Oscyp71Z2可能通过调控植保素和H202生物合成通路从而在水稻白叶枯病菌不亲合互作中起到重要作用。
Bacterial blight is one of the most destructive rice diseases, which caused by Xanthomonas oryzae pv. oryzae (Xoo), and results in10%to80%yield losses, endangering worldwide food security. Presently heavily application of chemical pesticides for controling disease has seriously damaged ecological environment. Meanwhile, easy lost of rice variety resistance to disease posed higher demond for rice breeding. An economically effective and environmentally sound approach to control disease is the utilization of cultivars that possessed durable, broad-spectrum resistance to disease by biotechnology. Identification of rice resistance-related genes is the base to cultivate the new rice disease-resistant varieties. The exploration of gene code elicitor protein with potential application value provides available resource for controlling disease by inducing multiple defense responses.
Harpinxoo proteins from the Xoo stain JXOIII increase plant disease resistance by activating multiple defense responses in plants, such as systemic acquired resistance and hypersensitive response. However, it is unclear whether phytoalexin production and ROS burst are involved in the disease resistance conferred by the expression of the harpinxoo protein in rice. In preliminary studies, our group showed some hrfl (code harpinxoo protein)-transformed rice T3homozygote increased resistance to Xoo. On the basis, in this article hrfl-transformed rice T3homozygote NJH12has been choiced for analyzing the resistance mechanism to Xoo. In this article, ectopic expression of hrfl in rice enhanced resistance to bacterial blight. Accompanying the activation of genes related to the phytoalexin biosynthesis pathway in hrfl-transformed rice, phytoalexins quickly and consistently accumulated concurrent with the limitation of bacterial growth rate. Moreover, the hrfl-transformed rice showed an increased ability for ROS scavenging and decreased hydrogen peroxide (H2O2) concentration. Furthermore, the localization and relative quantification of silicon deposition in rice leaves was detected by scanning electron microscopy (SEM) and energy-dispersive X-ray spectrometer (EDS). Finally, the transcript levels of defense response genes increased in transformed rice. These data are significant because they provide evidence for a better understanding the role of defense responses in the incompatible interaction between Xoo and hrf1-transformed plants. The data also supply an opportunity for generating nonspecific resistance to pathogens.
Based on the transcriptome profile of the transgenic rice line NJH12expressing hrfl from Xoo, a resistance-related gene showing114.6-fold increase of expression level was identified. The gene is the cytochrome P450gene, named as Oscyp71Z2analyzed by bioinformation, which belongs to the CYP71Z subfamily. Oscyp71Z2CDS, Oscyp71Z2RNAi target fragment and promoter sequence have cloned from rice cultivar Nipponbare transcript and genome, respectively make transformation vectors for overexpression, RNAi and Oscyp71Z2expression pattern, then transformed into rice Nipponbare genome mediated by Agrobacterium tumefaciens. The results from normal PCR and Southern blot showed T-DNA has inserted into rice genome. The data from Northern blot and Western blot showed the transcript and protein level in Oscyp71Z2-overexpressiong plant was higher than that in wild type Nipponbare. On the contrary, the protein level in Oscyp71Z-RNAi plant was lower than that in wild type, the tanscript hasa no detected in both. A strong GUS signal was detected mainly in the leaves, nodes, primary roots and glume. Overexpression of Oscyp71Z2enhanced resistance to Xoo at the booting stage, and inhibited multiplication of Xoo PXO99A. The accumulation of phytoalexins was rapidly and strongly induced in Oscyp71Z2-overexpressing plants, and the transcript levels of genes related to the phytoalexin biosynthesis pathway were elevated. The H2O2concentration in Oscyp71Z2-overexpressing plants was reduced in accordance with the increase in ROS-scavenging ability due to the induction of superoxide dismutase (SOD) as well as peroxidase (POD) and catalase (CAT) activation. We also showed that suppression of Oscyp71Z2had no significantly effect on disease resistance to Xoo in rice, although inhibited part phytoalexin production. These results demonstrated that Oscyp71Z2plays an important role for bacterial blight resistance by regulating the diterpenoid phytoalexin biosynthesis and H2O2generation.
引文
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