xa13基因在水稻—白叶枯病菌互作过程中的功能研究
摘要
水稻白叶枯病是由黄单胞杆菌水稻变种（Xanthomonas oryzae pv.oryzae, Xoo）引起，是水稻生产中的重要病害，每年造成巨大经济损失。多年生产实践表明，利用抗病基因资源，培育具有高效广谱抗性的优良品种是防治白叶枯病最经济有效的措施。对白叶枯病抗性基因机制和功能的深入研究，对利用主效抗病基因培育优良品种有着重要的指导意义。
本研究从病原和寄主两方面展开工作，对病原菌研究发现在所有的白叶枯病原菌中，菲律宾菌系生理小种6（PXO99）对铜离子最敏感。为了研究病原菌PXO99对铜敏感机制，构建了抗铜病原菌PXO61的全基因组文库，并导入到PXO99菌中，筛选抗铜突变体，对突变体测序比对发现该区段含有两个抗铜相关基因copA和copB。氨基酸比对发现，PXO61与PXO99的copB基因编码的氨基酸相同，而与PXO61的copA编码区相比，PXO99的copA基因编码区有六个氨基酸“DGTTQG”的缺失。将PXO99和PXO61的copA基因编码区进行互换，证实PXO99对铜离子的敏感是由于copA基因编码区的六个氨基酸“DGTTQG”缺失造成的。
为了研究copA和copB基因在病原菌中致病力的作用，将来自PXO61的copA和copB基因连接到pHMI载体上，并电转化PXO99菌株中，得到16株耐铜的突变菌株，接种水稻感病品种IR24发现其致病力增强，而接种抗病品种IRBB13，不能打败xa13介导的抗性。同时利用同源重组的原理，借助自杀性载体pK18mobsacB，将PXO99小种的copA和copB基因同时突变，得到对铜离子更加敏感的copAB的突变体7株，分别接种IR24和IRBB13，与野生型PXO99相比，致病力有所下降，说明copA和copB具有致病因子的功能。而抗铜突变体，不能使抗病品种完全恢复感病，说明铜离子只是部分参与xa13介导的抗病，暗示了xa13还存在未知功能。
通过对水稻XA13蛋白家族同源性分析发现，OsXa13与来自禾本科作物玉米的同源基因（ZmXa13）同源性最高，其在N-端（预测的跨膜区）的同源性高达90.5%，C-端的同源性为40.6%。通过酵母铜转运互补实验，证实OsXA13（1-220）可以和OsCOPT1和OsCOPT5形成三位一体的复合物，发挥其将铜从细胞外泵入细胞内的功能。同时酵母双杂交实验表明ZmXa13(56-100aa)第94位缬氨酸突变为丙氨酸影响其与OsCOPT5的互作。为进一步探索OsXA13C端的作用，获得PXa13: OsXA131-220、PXa13: ZmXa13、PXa13:NOs-CZM、PXa13: NZM-COs四种转基因植株，接种发现只有PXa13:NOs-CZM转基因植株能够恢复感病，说明OsXa13的C端是感病所必需的并且功能保守。
为了解析Xa13C端功能和作用机制，利用酵母双杂交的方法，筛选到两个与XA13相互作用的蛋白CacyBP和HMG，通过酵母双杂交和双分子荧光互补实验，分别在体外和体内证实XA13蛋白和CacyBP及HMG蛋白相互作用的真实性。同时，对CacyBP和HMG进行亚细胞定位分析，发现CacyBP定位于细胞膜区域，HMG定位于细胞膜和细胞核内。利用农杆菌介导的遗传转化，获得CacyBP和HMG超量和抑制表达的转基因水稻株系，接种试验表明，过量表达CacyBP或HMG的阳性株系的抗病性减弱而抑制表达CacyBP或HMG的阳性株系抗病性增强，说明CacyBP及HMG基因作为负调控因子参与水稻xa13介导的的抗病反应。
The rice bacterial blight (BB), caused by Xanthmonas oryzae pv. Oryzae,(Xoo),was oneof the most devastating diseases of rice world wide and causes tremendous agricultural yieldloss each year. Utilization of host resistance genes during agricultural production is the mosteffective and economic way. Understanding the function and mechanism on disease resistancegenes may have important significance in science and application for improving rice.
In the experiments of Xoo for copper stress response, high concentrations of copperwill inhibit the growth and reproduction of the bacterial blights. The strain of the Philippines6(PXO99) is the most sensitive to copper among all the Xoos tested. In order to study thecopper sensitive mechanism,firstly a genomic library constructed from PXO61which isresistant to copper and were transformed into PXO99.Secondly, anti-copper mutants whichgrow well in the medium containing1mM copper were screened. the cosmids of mutant wereextracted and sequenced. We found that the fragments contained two anti-copper gene copAand copB. Further studies showed that, pathogens of PXO99and PXO61have the same thecoding region of the copB. Compared with encoding region of copA genes in the PXO61,thecoding region of copA in the PXO99is lack of six amino acids which is “DGTTQG”.Byreplacing of the coding region of copA from pathogen of PXO99and PXO61,we found thesensitivity to copper of PXO99is caused by missing six amino acids of the coding region"DGTTQG"of copA gene.
Secondly, in order to study the role of the copA and copB genes in the pathogenicity ofXoo, we obtained the fragmants of copA and copB from PXO61.Then it was ligated into thevector pHM1.Transformation was conducted by electroporation to PXO99. We choosed16mutant strains which were more resistant to copper, and inoculated susceptible varietiesIR24.The result shows that the copper-resistant clones have stronger virulence, but can notrestore the susceptibility to IRBB13. At the same time, copA and copB were knocked by usinga suicide vector pK18mobsacB. We obtained7copA and copB genes deletion mutants, andfound that the mutants were more susensitive to copper. Mutans and wild type wereinoculated IR24and IRBB13.These results showed that the pathogenicity of the mutants weredeclined in IR24and IRBB13comparing with wild typre after14days. The results indicatedthat the copA and copB were as pathogenic factors. In addition, anti-copper mutants can notfully restore susceptible, indicating that copper was only partially involved in xa13-mediatedresitance to bacteaial blight disease, and suggesting xa13has still other unknown functions.
By homology analysis of rice XA13protein, we found that the gene frommaize(ZmXa13) was the highest homologous to OsXa13,which the homology of the N-terminal was up to90.5%and the homology of the C-terminal XA13was up to40.6%. Usingthe copper ions complementation experiments in S. cerevisiae ctr1Δctr3Δmutant MPY17indicated that combined with OsCOPT1and OsCOPT5, OsXA13(1-220)protein could beformed complex to play the role of transporting copper into cells from the extracellularenvironment. Yeast two-hybrid (Y2H)experiments showed that ZmXa13fragment (56-100aa)which is94valine to alanine impact the interaction with OsCOPT5protein.In order to furtherverify the role of C-terminal of OsXA13played in pathogenicity to rice, four trangenic plantsnamed PXa13: OsXA131-220、PXa13:ZmXa13、PXa13:NOs-CZM、PXa13:NZM-COswere inoculatedwith PXO99.We found that only the PXa13:NOs-CZMtrangenic plants restored susceptibility toPXO99. Thus, the C-terminus of the OsXa13is necessary and its function is conserved.
In order to resolve the function and mechanism of C-terminal XA13, We found twoproteins CacyBP and HMG could interact with XA13by using the Y2H. The interactionsbetween XA13and CacyBP or HMG were further confirmed by bimolecular fluorescencecomplementation(BiFC) experiment. To determine the subcellular location of CacyBP andHMG in plant cells, a construct containing a GFP fusion was transient expressed intobacco,we found that CacyBP located in the cell membrane area, and HMG located inmembrane and nucleus. To identify to the biological function of CacyBP and HMG, we over-express or suppress CacyBP and HMG in rice varieties Zhonghua11. Transgenic plants atbooting stage were measured the resistance to PXO99. The results showed that, comparedwith the wild-type varieties, CacyBP over-expression lines were more sensitive to PXO99,while CacyBP suppression lines exhibited high resistance to PXO99, and HMG transgeniclines had similar results to CacyBP. These results demonstrated that CacyBP and HMG genesare negative factors involved in regulating xa13-mediated resitantce to bacteaial blight disease.
本研究从病原和寄主两方面展开工作，对病原菌研究发现在所有的白叶枯病原菌中，菲律宾菌系生理小种6（PXO99）对铜离子最敏感。为了研究病原菌PXO99对铜敏感机制，构建了抗铜病原菌PXO61的全基因组文库，并导入到PXO99菌中，筛选抗铜突变体，对突变体测序比对发现该区段含有两个抗铜相关基因copA和copB。氨基酸比对发现，PXO61与PXO99的copB基因编码的氨基酸相同，而与PXO61的copA编码区相比，PXO99的copA基因编码区有六个氨基酸“DGTTQG”的缺失。将PXO99和PXO61的copA基因编码区进行互换，证实PXO99对铜离子的敏感是由于copA基因编码区的六个氨基酸“DGTTQG”缺失造成的。
为了研究copA和copB基因在病原菌中致病力的作用，将来自PXO61的copA和copB基因连接到pHMI载体上，并电转化PXO99菌株中，得到16株耐铜的突变菌株，接种水稻感病品种IR24发现其致病力增强，而接种抗病品种IRBB13，不能打败xa13介导的抗性。同时利用同源重组的原理，借助自杀性载体pK18mobsacB，将PXO99小种的copA和copB基因同时突变，得到对铜离子更加敏感的copAB的突变体7株，分别接种IR24和IRBB13，与野生型PXO99相比，致病力有所下降，说明copA和copB具有致病因子的功能。而抗铜突变体，不能使抗病品种完全恢复感病，说明铜离子只是部分参与xa13介导的抗病，暗示了xa13还存在未知功能。
通过对水稻XA13蛋白家族同源性分析发现，OsXa13与来自禾本科作物玉米的同源基因（ZmXa13）同源性最高，其在N-端（预测的跨膜区）的同源性高达90.5%，C-端的同源性为40.6%。通过酵母铜转运互补实验，证实OsXA13（1-220）可以和OsCOPT1和OsCOPT5形成三位一体的复合物，发挥其将铜从细胞外泵入细胞内的功能。同时酵母双杂交实验表明ZmXa13(56-100aa)第94位缬氨酸突变为丙氨酸影响其与OsCOPT5的互作。为进一步探索OsXA13C端的作用，获得PXa13: OsXA131-220、PXa13: ZmXa13、PXa13:NOs-CZM、PXa13: NZM-COs四种转基因植株，接种发现只有PXa13:NOs-CZM转基因植株能够恢复感病，说明OsXa13的C端是感病所必需的并且功能保守。
为了解析Xa13C端功能和作用机制，利用酵母双杂交的方法，筛选到两个与XA13相互作用的蛋白CacyBP和HMG，通过酵母双杂交和双分子荧光互补实验，分别在体外和体内证实XA13蛋白和CacyBP及HMG蛋白相互作用的真实性。同时，对CacyBP和HMG进行亚细胞定位分析，发现CacyBP定位于细胞膜区域，HMG定位于细胞膜和细胞核内。利用农杆菌介导的遗传转化，获得CacyBP和HMG超量和抑制表达的转基因水稻株系，接种试验表明，过量表达CacyBP或HMG的阳性株系的抗病性减弱而抑制表达CacyBP或HMG的阳性株系抗病性增强，说明CacyBP及HMG基因作为负调控因子参与水稻xa13介导的的抗病反应。
The rice bacterial blight (BB), caused by Xanthmonas oryzae pv. Oryzae,(Xoo),was oneof the most devastating diseases of rice world wide and causes tremendous agricultural yieldloss each year. Utilization of host resistance genes during agricultural production is the mosteffective and economic way. Understanding the function and mechanism on disease resistancegenes may have important significance in science and application for improving rice.
In the experiments of Xoo for copper stress response, high concentrations of copperwill inhibit the growth and reproduction of the bacterial blights. The strain of the Philippines6(PXO99) is the most sensitive to copper among all the Xoos tested. In order to study thecopper sensitive mechanism,firstly a genomic library constructed from PXO61which isresistant to copper and were transformed into PXO99.Secondly, anti-copper mutants whichgrow well in the medium containing1mM copper were screened. the cosmids of mutant wereextracted and sequenced. We found that the fragments contained two anti-copper gene copAand copB. Further studies showed that, pathogens of PXO99and PXO61have the same thecoding region of the copB. Compared with encoding region of copA genes in the PXO61,thecoding region of copA in the PXO99is lack of six amino acids which is “DGTTQG”.Byreplacing of the coding region of copA from pathogen of PXO99and PXO61,we found thesensitivity to copper of PXO99is caused by missing six amino acids of the coding region"DGTTQG"of copA gene.
Secondly, in order to study the role of the copA and copB genes in the pathogenicity ofXoo, we obtained the fragmants of copA and copB from PXO61.Then it was ligated into thevector pHM1.Transformation was conducted by electroporation to PXO99. We choosed16mutant strains which were more resistant to copper, and inoculated susceptible varietiesIR24.The result shows that the copper-resistant clones have stronger virulence, but can notrestore the susceptibility to IRBB13. At the same time, copA and copB were knocked by usinga suicide vector pK18mobsacB. We obtained7copA and copB genes deletion mutants, andfound that the mutants were more susensitive to copper. Mutans and wild type wereinoculated IR24and IRBB13.These results showed that the pathogenicity of the mutants weredeclined in IR24and IRBB13comparing with wild typre after14days. The results indicatedthat the copA and copB were as pathogenic factors. In addition, anti-copper mutants can notfully restore susceptible, indicating that copper was only partially involved in xa13-mediatedresitance to bacteaial blight disease, and suggesting xa13has still other unknown functions.
By homology analysis of rice XA13protein, we found that the gene frommaize(ZmXa13) was the highest homologous to OsXa13,which the homology of the N-terminal was up to90.5%and the homology of the C-terminal XA13was up to40.6%. Usingthe copper ions complementation experiments in S. cerevisiae ctr1Δctr3Δmutant MPY17indicated that combined with OsCOPT1and OsCOPT5, OsXA13(1-220)protein could beformed complex to play the role of transporting copper into cells from the extracellularenvironment. Yeast two-hybrid (Y2H)experiments showed that ZmXa13fragment (56-100aa)which is94valine to alanine impact the interaction with OsCOPT5protein.In order to furtherverify the role of C-terminal of OsXA13played in pathogenicity to rice, four trangenic plantsnamed PXa13: OsXA131-220、PXa13:ZmXa13、PXa13:NOs-CZM、PXa13:NZM-COswere inoculatedwith PXO99.We found that only the PXa13:NOs-CZMtrangenic plants restored susceptibility toPXO99. Thus, the C-terminus of the OsXa13is necessary and its function is conserved.
In order to resolve the function and mechanism of C-terminal XA13, We found twoproteins CacyBP and HMG could interact with XA13by using the Y2H. The interactionsbetween XA13and CacyBP or HMG were further confirmed by bimolecular fluorescencecomplementation(BiFC) experiment. To determine the subcellular location of CacyBP andHMG in plant cells, a construct containing a GFP fusion was transient expressed intobacco,we found that CacyBP located in the cell membrane area, and HMG located inmembrane and nucleus. To identify to the biological function of CacyBP and HMG, we over-express or suppress CacyBP and HMG in rice varieties Zhonghua11. Transgenic plants atbooting stage were measured the resistance to PXO99. The results showed that, comparedwith the wild-type varieties, CacyBP over-expression lines were more sensitive to PXO99,while CacyBP suppression lines exhibited high resistance to PXO99, and HMG transgeniclines had similar results to CacyBP. These results demonstrated that CacyBP and HMG genesare negative factors involved in regulating xa13-mediated resitantce to bacteaial blight disease.
引文
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