泡桐丛枝植原体致病相关基因分子特征及其编码蛋白功能研究
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摘要
泡桐(Paulownia sp.)是我国重要的阔叶树种之一，泡桐丛枝病(Paulownia witches’broom disease)在我国发生普遍，是由泡桐丛枝植原体（Paulownia witches’ broomphytoplasma）引起的侵染性病害，表现为腋芽和不定芽大量萌发至簇生成团，有些病树上会出现花变叶症状，常常不能正常开花结实。为了深入了解植原体致病机理，本文通过植原体致病相关基因的克隆、原核表达、抗血清制备、蛋白活性测定、瞬时表达与病症分析等方法，对植原体致病基因的功能进行研究和分析，发现tRNA-ipt基因可能影响了植物细胞的分裂和增长，这为植原体致病分子机制提供了新的思路和线索。
本研究用PCR方法在我国四种16SrI组植原体中克隆到tRNA异戊烯基焦磷酸转移酶基因(tRNA-ipt)，并对其全长序列进行生物信息学分析，发现泡桐丛枝（Paulowniawitches’ broom，PaWB）、桑萎缩（Mulberry dwarf，MD）、长春花绿变（periwinkle virescence，PeV）及苦楝丛枝（Chinaberry witches’ broom，CWB）植原体的tRNA-ipt基因的开放读码框长度均为876bp，G+C含量分别为30%、29.5%、29.5%和29.3%；它们均编码291个氨基酸的蛋白，预测分子量均为33kDa。四种植原体的IPT蛋白N端皆含有ATP/GTP结合相关序列（GPTASGKT）。预测该蛋白无跨膜区，分布在细胞质中。四种植原体的tRNA-IPT之间的氨基酸序列相似性为99.1%-99.5%，与同组植原体蛋白相似性为95.4%-99.3%，与其他组植原体低于70%；系统进化分析显示，所有已知9个植原体tRNA-IPT共聚为一大支，且与无胆甾原体遗传关系更近。
对泡桐丛枝植原体tRNA-ipt基因进行原核表达并制备抗体。利用Western blot和FITC间接免疫荧光显微镜检测其在植原体中的表达。使用分光光度计分析该基因对大肠杆菌生长的影响，用ELISA测定转化菌株细胞分裂素含量。SDS-PAGE结果显示tRNA-IPT蛋白在大肠杆菌中得到表达。首次获得泡桐丛枝植原体tRNA-IPT抗体并检测到该蛋白在泡桐发病组织中的特异表达。经过对转化菌株生长曲线及玉米素含量的测定，发现该基因能促进大肠杆菌后期生长和玉米素核苷的积累。结果表明泡桐丛枝植原体tRNA-IPT蛋白能够在植原体中表达，根据该基因对异源菌株生长速率和激素合成的影响推断该蛋白可能参与植原体的细胞分裂素合成，并在致病过程中起到重要作用。
构建病毒表达载体pCAPE-ipt，转化农杆菌，注射浸润健康泡桐组培苗叶片，使植物细胞表达tRNA-IPT蛋白。结果显示感染pCAPE-ipt的泡桐离体叶片可以保持较长的生长和存活时间；感染pCAPE-ipt的泡桐植株，两周后开始长出腋芽，与泡桐丛枝病早期症状极为相似。同时Western blot证实tRNA-IPT已在植物中表达。根据此结果推测，tRNA-IPT蛋白可能影响寄主植物激素平衡，从而导致丛枝病，这为植原体致病分子机制提供了新的线索。
根据已发表植原体质粒序列设计引物，经两次PCR扩增，获得大小不同的两段DNA片段，测序后拼接得到了4种完整的环状质粒，分别是桑树萎缩植原体濮阳株系质粒（pMDPy）、长春花绿变植原体海南株系质粒（pPeVHn）、泡桐丛枝植原体山东株系质粒（pPaWBG33D）及苦楝丛枝植原体福清株系质粒（pCWBFq-2），其中pMDPy为首次测定，其质粒全长分别为3833bp、3943bp、3843bp和3913bp，4种质粒皆编码5个蛋白，ORF1和ORF5分别编码复制相关蛋白（RepA）和单链DNA结合蛋白（SSB），ORF2-4编码未知功能蛋白。4种质粒序列的非编码区分析表明，ORF1和ORF2之间的非编码区存在启动子和核糖体结合位点。将4种植原体质粒全序列与GenBank中登录的植原体质粒进行序列相似性比对和系统进化分析，结果表明这4种质粒与其它16SrI组的质粒聚为一个大的分支，与16S rDNA系统发育树基本一致。此研究为进一步深入了解不同植原体质粒的结构与功能、寄主专化性和质粒基因变异及进化奠定了基础，也为基于多基因分类鉴定提供新的理论依据。
本研究首次在泡桐丛枝植原体中获得胸苷酸激酶基因（tmk），并在4个地区不同品种感病泡桐中检测到不同tmk基因。以PaWB-G33D、PaWB-JAN、PaWB-PS和PaWB-BJ株系侵染的泡桐总DNA为模板，扩增并克隆了tmk基因。分别由591、600、639和588个核苷酸组成。基因序列比对结果显示，泡桐丛枝植原体与16SrI组的其它几种植原体tmk基因相似性均大于90%。不同地区和不同品种泡桐tmk基因各有不同，这体现出在泡桐丛枝植原体中存在着tmk基因丰富的遗传变异和多样性种群。
Paulownia sp. is a kind of important broad leaf species in China. Paulownia witches’broom disease, is infectious disease and caused by Paulownia witches’ broom phytoplasma, itstypical symptoms includes abnormal adventitious bud and axillary bud germination to formcluster, even changed from flowers to leaves and can not form normal flowers and fruits. Inorder to understand the pathogenic mechanism of phytoplasma, the related function researchand analysis were explored. Based on the related gene cloning and prokaryotic expression,antiserum preparation, protein activity determination, instantaneous expression and diseasesand symptoms analysis etc, it was found that the tRNA-ipt gene may influence plant celldivision and growth, which provides a new clue for the study of phytoplasma pathogenicmolecular mechanism.
Four kinds of full-length sequences of tRNA-ipts from aster yellows group(16SrI)phytoplasmas were amplified, sequenced and analyzed in-silico. It was found that the length ofopen reading frame of tRNA-ipt genes from paulownia witches’-broom，mulberry dwarf，periwinkle virescence and Chinaberry witches’-broom phytoplasmas is876bp, their G+Ccontents are30%,29.5%,29.5%and29.3%,respectively. All these genes encode291aminoacids, predicting the molecular weight is33kda. They contain an identical motif (GPTASGKT）at N-terminal region related to the ATP or GTP binding sites. It was predicted that the proteinsare without transmembrane region, distributed in the cytoplasm. Amino acid sequencehomology of four phytoplasma tRNA-IPTs was99.1-99.5%, sequence homology with thesame group phytoplasmas was95.4-99.3%, whereas with16SrX apple proliferation and16SrXII Australia grapevine yellows phytoplasmas was less than70%.Phylogenetic analysisrevealed that all nine known tRNA-IPTs of phytoplasmas were clustered into a common clade，and they had very close relationship with that of Acholeplasma sp.
The paulownia witches’-broom phytoplasma (PaWB) tRNA-ipt gene was expressed in E.coli, specific antibody was prepared and used for the gene expression analyses by Western blotand FITC indirect immunofluroscence microscopy. Then the growth curve and cytokinincontents of E. coli with PaWB tRNA-ipt were measured by photodensitometry and ELISA,respectively. The pET expression vector containing the PaWB tRNA-ipt gene was constructedand the induced fusion protein was expressed and detected by SDS-PAGE. By Western blottingand immunofluorescence detection, the expression and localization of the phytoplasmastRNA-IPT protein in the phloem of infected paulownia were confirmed. According to thedetermination of growth curve and zeatin content, we found that the gene can promote thegrowth of E. coli in the late stage and zeatin nucleotide accumulation. The determination ofgrowth curve suggested that the growth rate increase of E. coli was effected by thetransformation of exogenous tRNA-ipt gene, which might be associated with the cytokininaccumulation. This protein was assumed to involve in the cytokinin synthesis in phytoplasmasas well as other bacteria, which may play an important role in pathogenic processes ofphytoplasmas and symptom development.
The virus expression vector of pCAPE-ipt were constructed to transform into healthypaulownia plantlets leaf cells by agroinfiltrayion. Results showed that the detached paulownialeaves which infiltrated with GV3101-pCAPE-ipt could maintain a longer growing andsurvived period; When the PaWB tRNA-IPT was expressed in paulownia plants two weekslater, these treated plants showed symptoms of witches’ broom and dwarfism, which weretypical ones of phytoplasma infection. At the same time, Western blotting confirmed that theprotein tRNA-IPT was expressed in treated plants but not in control plants. According to thisresult, we speculateed that the phytoplasma tRNA-IPT protein may influence the host planthormone balance, and then lead to the symptom display of witches broom disease, whichprovides a new clue for the study of the phytoplasma pathogenic molecular mechanism.
On the basis of the known phytoplasma plasmid sequences, two pairs of specific primerswere designed and used for PCR. Two section of DNA amplified fragments with different sizes covering whole plasmid were obtained through two round of PCR amplification. Aftersequenced and spliced,four phytoplasmal full-length circular plasmid DNA sequences wereobtained, including the plasmids of mulberry dwarf(pMDPy) in Puyang of Henan province,periwinkle（Catharanthus roseus）virescence (pPEVHn) in Hainan province, paulowniawitches’-broom (pPaWBG33D) in Yanzhou of Shandong province and chinaberrywitches’-broom phytoplasma strains (pCWBFqHu) in Fuqing of Fujian province,with thesizes of3833bp,3943bp,3843bp and3913bp, respectively. They are predicted to encode fiveproteins,open reading frame ORF1encoding replication associated protein（RepA）,ORF5encoding binding protein（SSB）and ORF2-4encoding unknown function proteinss. pMDPy isthe first plasmid determined from mulberry dwarf phytoplasma.Sequence analysis ofnon-coding regions (NCR) of the four kinds of plasmids revealed that the NCR sequencesbetween ORF1and ORF2exist promoters and ribosome binding sites.The four phytoplasmaplasmid sequences were compared with the phytoplasma plasmid sequences found in GenBankthrough multiple alignment and phylogenetic analysis, the results showed that these fourplasmids and other16SrI group plasmid grouped into a large branch,which was consistent withthe phylogenetic tree created from16S rDNA comparisons. This research may not only providefurther insight into the structure and function, host specificity and plasmid variation andevolution within different phytoplasma species or strains,but also offers new theoretical basisfor multiple genes and index based system for phytoplasmal classification.
Four different thymidylate kinase(tmk) genes were also identified in different paulowniacultivars of various regions. It was found that the length of tmk genes from PaWB-G33D、PaWB-JAN、PaWB-PS and PaWB-BJ phytoplasmas is591、600、639and588bp respected.Four tmks share the90-98%nucleotide sequence homology with the same groupphytoplasmas.The tmks of PaWB from diferent regions were different, which suggested thatthere might exist abundant genetic variation and population diversity in paulowniawitches’-broom phytoplasmas.
本研究用PCR方法在我国四种16SrI组植原体中克隆到tRNA异戊烯基焦磷酸转移酶基因(tRNA-ipt)，并对其全长序列进行生物信息学分析，发现泡桐丛枝（Paulowniawitches’ broom，PaWB）、桑萎缩（Mulberry dwarf，MD）、长春花绿变（periwinkle virescence，PeV）及苦楝丛枝（Chinaberry witches’ broom，CWB）植原体的tRNA-ipt基因的开放读码框长度均为876bp，G+C含量分别为30%、29.5%、29.5%和29.3%；它们均编码291个氨基酸的蛋白，预测分子量均为33kDa。四种植原体的IPT蛋白N端皆含有ATP/GTP结合相关序列（GPTASGKT）。预测该蛋白无跨膜区，分布在细胞质中。四种植原体的tRNA-IPT之间的氨基酸序列相似性为99.1%-99.5%，与同组植原体蛋白相似性为95.4%-99.3%，与其他组植原体低于70%；系统进化分析显示，所有已知9个植原体tRNA-IPT共聚为一大支，且与无胆甾原体遗传关系更近。
对泡桐丛枝植原体tRNA-ipt基因进行原核表达并制备抗体。利用Western blot和FITC间接免疫荧光显微镜检测其在植原体中的表达。使用分光光度计分析该基因对大肠杆菌生长的影响，用ELISA测定转化菌株细胞分裂素含量。SDS-PAGE结果显示tRNA-IPT蛋白在大肠杆菌中得到表达。首次获得泡桐丛枝植原体tRNA-IPT抗体并检测到该蛋白在泡桐发病组织中的特异表达。经过对转化菌株生长曲线及玉米素含量的测定，发现该基因能促进大肠杆菌后期生长和玉米素核苷的积累。结果表明泡桐丛枝植原体tRNA-IPT蛋白能够在植原体中表达，根据该基因对异源菌株生长速率和激素合成的影响推断该蛋白可能参与植原体的细胞分裂素合成，并在致病过程中起到重要作用。
构建病毒表达载体pCAPE-ipt，转化农杆菌，注射浸润健康泡桐组培苗叶片，使植物细胞表达tRNA-IPT蛋白。结果显示感染pCAPE-ipt的泡桐离体叶片可以保持较长的生长和存活时间；感染pCAPE-ipt的泡桐植株，两周后开始长出腋芽，与泡桐丛枝病早期症状极为相似。同时Western blot证实tRNA-IPT已在植物中表达。根据此结果推测，tRNA-IPT蛋白可能影响寄主植物激素平衡，从而导致丛枝病，这为植原体致病分子机制提供了新的线索。
根据已发表植原体质粒序列设计引物，经两次PCR扩增，获得大小不同的两段DNA片段，测序后拼接得到了4种完整的环状质粒，分别是桑树萎缩植原体濮阳株系质粒（pMDPy）、长春花绿变植原体海南株系质粒（pPeVHn）、泡桐丛枝植原体山东株系质粒（pPaWBG33D）及苦楝丛枝植原体福清株系质粒（pCWBFq-2），其中pMDPy为首次测定，其质粒全长分别为3833bp、3943bp、3843bp和3913bp，4种质粒皆编码5个蛋白，ORF1和ORF5分别编码复制相关蛋白（RepA）和单链DNA结合蛋白（SSB），ORF2-4编码未知功能蛋白。4种质粒序列的非编码区分析表明，ORF1和ORF2之间的非编码区存在启动子和核糖体结合位点。将4种植原体质粒全序列与GenBank中登录的植原体质粒进行序列相似性比对和系统进化分析，结果表明这4种质粒与其它16SrI组的质粒聚为一个大的分支，与16S rDNA系统发育树基本一致。此研究为进一步深入了解不同植原体质粒的结构与功能、寄主专化性和质粒基因变异及进化奠定了基础，也为基于多基因分类鉴定提供新的理论依据。
本研究首次在泡桐丛枝植原体中获得胸苷酸激酶基因（tmk），并在4个地区不同品种感病泡桐中检测到不同tmk基因。以PaWB-G33D、PaWB-JAN、PaWB-PS和PaWB-BJ株系侵染的泡桐总DNA为模板，扩增并克隆了tmk基因。分别由591、600、639和588个核苷酸组成。基因序列比对结果显示，泡桐丛枝植原体与16SrI组的其它几种植原体tmk基因相似性均大于90%。不同地区和不同品种泡桐tmk基因各有不同，这体现出在泡桐丛枝植原体中存在着tmk基因丰富的遗传变异和多样性种群。
Paulownia sp. is a kind of important broad leaf species in China. Paulownia witches’broom disease, is infectious disease and caused by Paulownia witches’ broom phytoplasma, itstypical symptoms includes abnormal adventitious bud and axillary bud germination to formcluster, even changed from flowers to leaves and can not form normal flowers and fruits. Inorder to understand the pathogenic mechanism of phytoplasma, the related function researchand analysis were explored. Based on the related gene cloning and prokaryotic expression,antiserum preparation, protein activity determination, instantaneous expression and diseasesand symptoms analysis etc, it was found that the tRNA-ipt gene may influence plant celldivision and growth, which provides a new clue for the study of phytoplasma pathogenicmolecular mechanism.
Four kinds of full-length sequences of tRNA-ipts from aster yellows group(16SrI)phytoplasmas were amplified, sequenced and analyzed in-silico. It was found that the length ofopen reading frame of tRNA-ipt genes from paulownia witches’-broom，mulberry dwarf，periwinkle virescence and Chinaberry witches’-broom phytoplasmas is876bp, their G+Ccontents are30%,29.5%,29.5%and29.3%,respectively. All these genes encode291aminoacids, predicting the molecular weight is33kda. They contain an identical motif (GPTASGKT）at N-terminal region related to the ATP or GTP binding sites. It was predicted that the proteinsare without transmembrane region, distributed in the cytoplasm. Amino acid sequencehomology of four phytoplasma tRNA-IPTs was99.1-99.5%, sequence homology with thesame group phytoplasmas was95.4-99.3%, whereas with16SrX apple proliferation and16SrXII Australia grapevine yellows phytoplasmas was less than70%.Phylogenetic analysisrevealed that all nine known tRNA-IPTs of phytoplasmas were clustered into a common clade，and they had very close relationship with that of Acholeplasma sp.
The paulownia witches’-broom phytoplasma (PaWB) tRNA-ipt gene was expressed in E.coli, specific antibody was prepared and used for the gene expression analyses by Western blotand FITC indirect immunofluroscence microscopy. Then the growth curve and cytokinincontents of E. coli with PaWB tRNA-ipt were measured by photodensitometry and ELISA,respectively. The pET expression vector containing the PaWB tRNA-ipt gene was constructedand the induced fusion protein was expressed and detected by SDS-PAGE. By Western blottingand immunofluorescence detection, the expression and localization of the phytoplasmastRNA-IPT protein in the phloem of infected paulownia were confirmed. According to thedetermination of growth curve and zeatin content, we found that the gene can promote thegrowth of E. coli in the late stage and zeatin nucleotide accumulation. The determination ofgrowth curve suggested that the growth rate increase of E. coli was effected by thetransformation of exogenous tRNA-ipt gene, which might be associated with the cytokininaccumulation. This protein was assumed to involve in the cytokinin synthesis in phytoplasmasas well as other bacteria, which may play an important role in pathogenic processes ofphytoplasmas and symptom development.
The virus expression vector of pCAPE-ipt were constructed to transform into healthypaulownia plantlets leaf cells by agroinfiltrayion. Results showed that the detached paulownialeaves which infiltrated with GV3101-pCAPE-ipt could maintain a longer growing andsurvived period; When the PaWB tRNA-IPT was expressed in paulownia plants two weekslater, these treated plants showed symptoms of witches’ broom and dwarfism, which weretypical ones of phytoplasma infection. At the same time, Western blotting confirmed that theprotein tRNA-IPT was expressed in treated plants but not in control plants. According to thisresult, we speculateed that the phytoplasma tRNA-IPT protein may influence the host planthormone balance, and then lead to the symptom display of witches broom disease, whichprovides a new clue for the study of the phytoplasma pathogenic molecular mechanism.
On the basis of the known phytoplasma plasmid sequences, two pairs of specific primerswere designed and used for PCR. Two section of DNA amplified fragments with different sizes covering whole plasmid were obtained through two round of PCR amplification. Aftersequenced and spliced,four phytoplasmal full-length circular plasmid DNA sequences wereobtained, including the plasmids of mulberry dwarf(pMDPy) in Puyang of Henan province,periwinkle（Catharanthus roseus）virescence (pPEVHn) in Hainan province, paulowniawitches’-broom (pPaWBG33D) in Yanzhou of Shandong province and chinaberrywitches’-broom phytoplasma strains (pCWBFqHu) in Fuqing of Fujian province,with thesizes of3833bp,3943bp,3843bp and3913bp, respectively. They are predicted to encode fiveproteins,open reading frame ORF1encoding replication associated protein（RepA）,ORF5encoding binding protein（SSB）and ORF2-4encoding unknown function proteinss. pMDPy isthe first plasmid determined from mulberry dwarf phytoplasma.Sequence analysis ofnon-coding regions (NCR) of the four kinds of plasmids revealed that the NCR sequencesbetween ORF1and ORF2exist promoters and ribosome binding sites.The four phytoplasmaplasmid sequences were compared with the phytoplasma plasmid sequences found in GenBankthrough multiple alignment and phylogenetic analysis, the results showed that these fourplasmids and other16SrI group plasmid grouped into a large branch,which was consistent withthe phylogenetic tree created from16S rDNA comparisons. This research may not only providefurther insight into the structure and function, host specificity and plasmid variation andevolution within different phytoplasma species or strains,but also offers new theoretical basisfor multiple genes and index based system for phytoplasmal classification.
Four different thymidylate kinase(tmk) genes were also identified in different paulowniacultivars of various regions. It was found that the length of tmk genes from PaWB-G33D、PaWB-JAN、PaWB-PS and PaWB-BJ phytoplasmas is591、600、639and588bp respected.Four tmks share the90-98%nucleotide sequence homology with the same groupphytoplasmas.The tmks of PaWB from diferent regions were different, which suggested thatthere might exist abundant genetic variation and population diversity in paulowniawitches’-broom phytoplasmas.
引文
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