十字花科黑腐病菌colRS双组分调控系统基因的研究
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摘要
十字花科黑腐病菌,学名为野油菜黄单胞菌野油菜致病变种(Xanthomonas campestris pv.campestris,简称Xcc),是一类γ-变形菌纲的革兰氏阴性细菌,能在世界范围侵染十字花科植物,给农业生产造成重大损失。该菌易于培养、保存,具有较好的遗传操作性,是研究微生物与寄主相互作用分子机理的重要模式细菌之一。Xcc属于典型的维管束病害病原菌,病菌通过水孔或伤口侵入植物体内,并在维管束木质部导管中扩展,导致系统病害。致病过程包括接触、识别、侵入、繁殖、扩展、显症等阶段,是一个复杂的、动态的多因子的相互作用过程。在整个侵染循环中,Xcc要历经腐生、附生和寄生阶段,经历着很大的环境变化,不断地与寄主以及环境因素发生互作。研究表明,在侵入寄主之前,在叶表成功定殖有利于病原菌侵入寄主体内。
根据研究报道,colR、colS(合称colRS)双组分调控系统在荧光假单胞菌WCS365菌株(Pseudomonas fluorescens WCS365)、恶臭假单胞菌KT2440(Pseudomonas putida KT2440)菌株中参与调节细菌在植物根际定殖过程。colS的编码蛋白ColS感受外界刺激,传递磷酸基团至colR的编码蛋白ColR,ColR调节其它基因表达。除了影响定殖外,目前了解到colR基因调节细菌抗逆性,colR与恶臭假单胞菌CD2菌株对重金属的抗逆性和恶臭假单胞菌PaW85菌株(Pseudomonas putida PaW85)对酚的抗逆性相关;colRS参与调节恶臭假单胞菌PaW85菌株Tn4652转座子转座等。双组分调控系统是细菌对环境变化做出应对的重要的调节基因,参与调节定殖、生长发育、抗逆性、致病性等多种细胞活动和功能。因此,双组分调控系统相关的基因及产物被认为是未来发展控制细菌病害的理想药物靶标。然而,迄今尚未有与定殖相关植物病原细菌双组分调控系统基因的报道。
在完成了Xcc 8004菌株全基因组序列测定的基础上,为了弄清该菌基因组中是否存在colRS基因的同源基因,我们用已报道的假单胞菌colRS基因编码的蛋白质序列对Xcc 8004全基因组数据库进行Blast搜索。结果表明,在Xcc 8004菌株基因组中,搜索到与已报道的colR、colS相似性很高的三对基因,XC1049和XC1050、XC3125和XC3126、XC3451和XC3452。为了了解这些基因是否与定殖相关,通过定点整合突变的方式,构建了以上六个基因的突变体NK1049、NK1050、NK3125和NK3126、NK3451和NK3452。植株实验检测结果表明,突变体NK1049定殖能力显著下降,将带有XC1049基因的pLAFR3导入NK1049突变体,能成功互补NK1049。表明XC1049基因与Xcc在寄主叶表定殖有关,故将XC1049命名为colR_(Xcc)。植株实验还显示,NK1049致病性基本丧失,在寄主植物满身红萝卜中生长迟缓,在辣椒品种ECW-10R诱导过敏反应延迟、强度显著下降。其它基因的突变体表型与野生型Xcc 8004菌株表型基本一致。
生理生化检测发现,在重金属、有机溶剂、渗透压、酸碱胁迫下,突变体NK1049可以忍受的胁迫水平明显低于野生型菌株Xcc 8004,colR_(Xcc)对Xcc 8004抗逆性具有显著影响;与Xcc 8004、互补株CNK1049相比,突变体NK1049在NYGB培养基、MMX培养基中生长较慢。其它检测如营养缺陷型检测、泳动观察、生物膜形成检测、胞外蛋白酶,胞外淀粉酶、胞外纤维素酶和胞外多糖等Xcc的致病因子检测没有发现突变体与Xcc8004有明显差异。抗生素抗性分析结果表明,突变体NK1049比Xcc 8004对抗生素敏感,细菌细胞外膜渗透性可能改变;多粘菌素与细胞外膜脂多糖结合影响膜渗透性,NK1049对多粘菌素(polymyxin)敏感,colR_(Xcc)可能改变脂多糖功能而影响膜的渗透性。RT-PCR分析发现,colR_(Xcc)调节hrp,基因表达,colR_(Xcc)突变后hrpC、hrpE转录单元表达量显著降低。
总之,本研究鉴定了十字花科黑腐病菌的一个与叶表定殖有关的基因colR_(Xcc)。实验表明,colR_(Xcc)与定殖、致病性、抗逆性及细胞生长相关,且调控该菌的hrp致病系统基因的表达。
Xanthomonas campestris pathovar campestris(Xcc)is the causal agent of black-rot disease,one of the most destructive diseases of cruciferous plants worldwide.To prevent the disease intensive studies have been carried out to investigate the mechanism of this model phytopathogen.Colonization is the initial phase of infection that Plant pathogens need to success colonize on host surface before entering into plant.Two-component regulatory system is widely used by bacteria for regulation of various cellular processes such as colonization,adaptation,growth,virulence,etc.It had been reported that colRS two-component regulatory system plays an important role in regulation of colonization.In addition,the research on colRS showed colR has function in adaptation in Pseudomonas putida CD2 and Pseudomonas putida PaW85 and colRS participates regulation of Tn4652 transposition in Pseudomonas putida PaW85,respectively.
There are 3 pairs of predicted colRS homologous genes in X.campestris pv.campestris 8004.To investigate whether colRS homologous genes contribute to colonization of Xcc,the integration mutants of colR,colS genes NK1049,NK1050,NK3125,NK3126,NK3451 and NK3452 were constructed and applied to colonization test.It was found that the mutant NK1049 has reduced colonizing-ability on plant and the phenotypcs of NK1049 could be restored by XC1049 in trans.XC1049 was then designated as colR_(Xcc).Plant-infection test also showed that NK1049 was impaired in virulence and could only cause weak and delayed HR on non-host plant.
General stress assay revealed that colR_(Xcc)mutant was defective in adaptation to osmotic,heavy-metal,phenol,pH stress.Monitoring growth rate showed the mutant NK1049 persisting slow growth in medium and plant leaves.Antibiotic sensitivity determination indicates that membrane function of colR_(Xcc)is likely altered.RT-PCR showed that mutation in colR_(Xcc)reduced the transcriptional level of hrpC and hrpE operons.
This work identifies the colR_(Xcc)as a key gene which is involved in colonization on host leaves,stress adaptation,growth,infection of plant, and regulation of hrp genes in X.campestris pv.campestris.
根据研究报道,colR、colS(合称colRS)双组分调控系统在荧光假单胞菌WCS365菌株(Pseudomonas fluorescens WCS365)、恶臭假单胞菌KT2440(Pseudomonas putida KT2440)菌株中参与调节细菌在植物根际定殖过程。colS的编码蛋白ColS感受外界刺激,传递磷酸基团至colR的编码蛋白ColR,ColR调节其它基因表达。除了影响定殖外,目前了解到colR基因调节细菌抗逆性,colR与恶臭假单胞菌CD2菌株对重金属的抗逆性和恶臭假单胞菌PaW85菌株(Pseudomonas putida PaW85)对酚的抗逆性相关;colRS参与调节恶臭假单胞菌PaW85菌株Tn4652转座子转座等。双组分调控系统是细菌对环境变化做出应对的重要的调节基因,参与调节定殖、生长发育、抗逆性、致病性等多种细胞活动和功能。因此,双组分调控系统相关的基因及产物被认为是未来发展控制细菌病害的理想药物靶标。然而,迄今尚未有与定殖相关植物病原细菌双组分调控系统基因的报道。
在完成了Xcc 8004菌株全基因组序列测定的基础上,为了弄清该菌基因组中是否存在colRS基因的同源基因,我们用已报道的假单胞菌colRS基因编码的蛋白质序列对Xcc 8004全基因组数据库进行Blast搜索。结果表明,在Xcc 8004菌株基因组中,搜索到与已报道的colR、colS相似性很高的三对基因,XC1049和XC1050、XC3125和XC3126、XC3451和XC3452。为了了解这些基因是否与定殖相关,通过定点整合突变的方式,构建了以上六个基因的突变体NK1049、NK1050、NK3125和NK3126、NK3451和NK3452。植株实验检测结果表明,突变体NK1049定殖能力显著下降,将带有XC1049基因的pLAFR3导入NK1049突变体,能成功互补NK1049。表明XC1049基因与Xcc在寄主叶表定殖有关,故将XC1049命名为colR_(Xcc)。植株实验还显示,NK1049致病性基本丧失,在寄主植物满身红萝卜中生长迟缓,在辣椒品种ECW-10R诱导过敏反应延迟、强度显著下降。其它基因的突变体表型与野生型Xcc 8004菌株表型基本一致。
生理生化检测发现,在重金属、有机溶剂、渗透压、酸碱胁迫下,突变体NK1049可以忍受的胁迫水平明显低于野生型菌株Xcc 8004,colR_(Xcc)对Xcc 8004抗逆性具有显著影响;与Xcc 8004、互补株CNK1049相比,突变体NK1049在NYGB培养基、MMX培养基中生长较慢。其它检测如营养缺陷型检测、泳动观察、生物膜形成检测、胞外蛋白酶,胞外淀粉酶、胞外纤维素酶和胞外多糖等Xcc的致病因子检测没有发现突变体与Xcc8004有明显差异。抗生素抗性分析结果表明,突变体NK1049比Xcc 8004对抗生素敏感,细菌细胞外膜渗透性可能改变;多粘菌素与细胞外膜脂多糖结合影响膜渗透性,NK1049对多粘菌素(polymyxin)敏感,colR_(Xcc)可能改变脂多糖功能而影响膜的渗透性。RT-PCR分析发现,colR_(Xcc)调节hrp,基因表达,colR_(Xcc)突变后hrpC、hrpE转录单元表达量显著降低。
总之,本研究鉴定了十字花科黑腐病菌的一个与叶表定殖有关的基因colR_(Xcc)。实验表明,colR_(Xcc)与定殖、致病性、抗逆性及细胞生长相关,且调控该菌的hrp致病系统基因的表达。
Xanthomonas campestris pathovar campestris(Xcc)is the causal agent of black-rot disease,one of the most destructive diseases of cruciferous plants worldwide.To prevent the disease intensive studies have been carried out to investigate the mechanism of this model phytopathogen.Colonization is the initial phase of infection that Plant pathogens need to success colonize on host surface before entering into plant.Two-component regulatory system is widely used by bacteria for regulation of various cellular processes such as colonization,adaptation,growth,virulence,etc.It had been reported that colRS two-component regulatory system plays an important role in regulation of colonization.In addition,the research on colRS showed colR has function in adaptation in Pseudomonas putida CD2 and Pseudomonas putida PaW85 and colRS participates regulation of Tn4652 transposition in Pseudomonas putida PaW85,respectively.
There are 3 pairs of predicted colRS homologous genes in X.campestris pv.campestris 8004.To investigate whether colRS homologous genes contribute to colonization of Xcc,the integration mutants of colR,colS genes NK1049,NK1050,NK3125,NK3126,NK3451 and NK3452 were constructed and applied to colonization test.It was found that the mutant NK1049 has reduced colonizing-ability on plant and the phenotypcs of NK1049 could be restored by XC1049 in trans.XC1049 was then designated as colR_(Xcc).Plant-infection test also showed that NK1049 was impaired in virulence and could only cause weak and delayed HR on non-host plant.
General stress assay revealed that colR_(Xcc)mutant was defective in adaptation to osmotic,heavy-metal,phenol,pH stress.Monitoring growth rate showed the mutant NK1049 persisting slow growth in medium and plant leaves.Antibiotic sensitivity determination indicates that membrane function of colR_(Xcc)is likely altered.RT-PCR showed that mutation in colR_(Xcc)reduced the transcriptional level of hrpC and hrpE operons.
This work identifies the colR_(Xcc)as a key gene which is involved in colonization on host leaves,stress adaptation,growth,infection of plant, and regulation of hrp genes in X.campestris pv.campestris.
引文
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