水稻白叶枯病菌和条斑病菌对噻枯唑和链霉素的抗药性机理研究
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摘要
水稻白叶枯病和水稻条斑病是水稻上的两种重要细菌性病害,噻枯唑和链霉素则是我国防治这两种病害的主要药剂。噻枯唑和链霉素在我国使用分别已有30年和50年的历史,在靶标病原群体中也均已出现抗药性。因此开展噻枯唑和链霉素抗性机理研究,对于抗药性治理和抗性诊断技术发展具有十分重要的意义。本论文围绕抗性机理进行了以下研究：1.水稻白叶枯病菌抗噻枯唑菌株粘粒基因组文库构建；2.抗噻枯唑水稻白叶枯菌株基因组文库的活体筛选和基因分析；3.水稻条斑病菌和白叶枯病菌抗链霉素分子机理研究；4.水稻白叶枯病菌、条斑病菌、大白菜软腐病菌以及柑橘溃疡疾菌抗药基因rpsL的分子检测。
1.水稻白叶枯病菌噻枯唑抗性菌株粘粒基因组文库构建
为了探索水稻白叶枯病菌Xanthomonas oryzae pv. oryzae (Xoo)对噻枯唑的抗性机制,本研究建立了白叶枯病菌噻枯唑抗性菌株2-1-1的粘粒基因组文库。通过对2-1-1基因组提取并不完全酶切,回收>23.13kb基因片段,用来自溶源菌BHB2688和BHB2690、效价达1×108pfu/μgDNA的包装蛋白包装,感染宿主大肠杆菌S17-1,在含LB+Km+Sp的平板上进行筛选,随机挑选1200个转化子进行扩增保存,库容达到2.7倍；随机挑选15个转化子进行酶切验证,均有外源片段插入粘粒载体中,插入片段大小23-40kb,成功构建了抗噻枯唑白叶枯病菌2-1-1的粘粒基因组文库。
2.水稻白叶枯菌噻枯唑抗性菌株基因组文库的活体筛选和基因分析
将构建好的白叶枯病菌噻枯唑抗性菌株2-1-1粘粒基因组文库,1200个转化子通过两亲交配转染白叶枯病菌野生敏感菌株PXO99,获得的结合子再剪叶接种经300μg/ml噻枯唑处理的稻苗,筛选发现646、518等2个结合子表现抗药性；同时通过在清水对照处理的水稻上筛选,发现5、33、115、152、119、102、172、519、537、611、643、501、694、518、612、654、308、919、670等多个转化子在致病性上强于2-1-1和PXO99。通过对646和518片段进行亚克隆,进一步缩小功能片段,发现抗性菌株有两个基因发生突变。两基因分别为raxR2基因和conserved hypothetical protein (Chp)基因发生突变。其中raxR2基因有7个碱基发生突变,但未造成氨基酸序列变化；Chp基因发生一个碱基突变,引起了氨基酸变化(Val变成Gly)。
3.水稻白叶枯病菌和条斑病菌链霉素抗性机理研究
通过室内紫外诱变和药剂驯服获得了水稻条斑病菌和白叶枯病菌链霉素抗性突变体。这些突变体可以在含100μg/mL链霉素平板上生长,而敏感菌株在5μg/mL浓度下则不能生长。通过田间致病力测定,发现抗性菌株在水稻叶片上的致病力与敏感出发菌株无明显差异。从水稻条斑病菌和白叶枯病菌野生敏感菌株及室内诱导抗性菌株中分别扩增到编码核糖体蛋白S12编码基因(rpsL)全序列,序列分析表明rpsL基因中第43位或第88位氨基酸由赖氨酸(AAG)突变为精氨酸(AGG)。rpsL基因全长375bp,编码125个氨基酸。序列分析结果表明水稻条斑病菌与水稻白叶枯病菌rpsL基因同源性达97%,编码氨基酸同源性达100%；与柑橘溃疡病菌氨基酸同源性达98%,与大白菜软腐病菌氨基酸同源性达99%。设计合成1对含有酶切位点(EcoRI)的特异性引物,从抗药性菌株UV-R-1(含rpsL基因43位点突变)和UV-R-3(含rpsL基因88位点突变)中扩增出含启动区的rpsL基因,成功构建了重组质粒pUFRRS知pUFRRX。功能验证证明,质粒pUFRRS和pUFRRX能够互补敏感菌株的抗药性功能,结合子RS1、RS2、RS3、PS1、PS2对链霉素敏感性测定结果证实rpsL基因序列发生点突变是水稻白叶枯病菌和条斑病菌对链霉素产生抗药性的主要原因。同时转化子的致病力与敏感出发菌株和抗药突变体的致病力无明显差异。
4.水稻白叶枯病菌、条斑病菌、大白菜软腐病菌以及柑橘溃疡病菌抗药基因rpsL的分子检测
通过室内药剂驯化和紫外诱导,获得大白菜软腐病菌链霉素抗性突变体12个,柑橘溃疡病菌突变体11个,番茄细菌性斑点病菌突变体12个。分别比较番茄细菌性斑点病菌、柑橘溃疡病菌、大白菜软腐病菌链霉素抗性突变体与他们的敏感亲本菌株的序列,发现所有抗链霉素菌株的rpsL基因中第43位或第88位氨基酸均由赖氨酸(AAG)突变为精氨酸(AGG)。由于psL基因43位点是MboⅡ的酶切位点,因此通过PCR-RFLP方法可以检测出上述细菌rpsL基因43位点突变情况。通过设计一对特异性引物,扩增rpsL基因部分片段304bp,若43位点发生突变,则酶切位点消失,304bp则被酶切成201bp和103 bp片段；若43位点不发生突变,304bp则被酶切成201bp、103bp和55 bp片段。研究结果表明PCR-RELP方法可以快速、准确的检测出rpsL基因43位点突变情况,检测结果与直接测序结果完全一致,且操作简便。
Bacterial blight of rice (BB) and bacterial leaf streak of rice (BLS) were caused by Xanthomonas oryzae pv. oryzae(Xoo) and X. oryzae pv. oryzicola (Xooc), respectively. BB is one of the most serious diseases of rice, and BLS is emerging in importance. Application of bactericides is an indispensable complementary tool, especially in regions where susceptible cultivars are frequently planted. Bismerthiazol and streptomycin are mainly bactericides which have been used for the control of bacterial plant disease. But bismerthiazol and streptomycin have been used for more than thirty years. Thus, to study the resistance mechanism becomes very important.In this paper, there are five points about resisitance mechanism that were studied:
Construction of cosmid gene library of bismerthiazol-resistant Xoo.
In vivo selection of the cosmid library of bismerthiazol-resistant Xoo and gene analysis.
Study on the molecule mechanism of the streptomycin resistance in Xoo and Xooc.
Detection of the resistance to streptomycin in Xoo、Xooc、Erwinia carotovora (Ec) and Xanthomonas axonopodis pv.citri (Xac) by PCR-RFLP within the mutation at codon 43 of the rpsL gene.
1. Construction of the cosmid library of bismerthiazol-resistant Xoo
In order to explore the resistance mechanism of bismerthiazol, the cosmid genome library of the bismerthiazol-resistant Xoo mutant 2-1-1 was constructed. Through incomplete digestion of the genome of 2-1-1, the>23.13 kb gene fragments were gathered. Then, the packaging protein was extracted with the lysogen BHB2688、BHB2690, and the potency came up to 1×108pfu/μgDNA. The gathered gene fragments were connected, packed with packaging protein, and infected with the host S17-1. Then the combined mutants were selected on the LB+Km+Sp plates. A total of 1200 transformants were randomly selected at last and stored. The reservoir storage came up to 2.7 times. Fifteen transformants were randomly selected and digested with enzymes, the results indicated all the cosmids contained heterogenous insertional fragments and the cosmid genome library was successfully constructed, with average 23-40 kb gene fragments inserted.
2. In vivo selection of the cosmid library of bismerthiazol-resistant Xoo and gene analysis
The 1200 transformants described above were transformed by conjugation into PXO99 (a bismerthiazol-sensitive strain). Then, the in-vivo selection was done on the treated rice plants (sprayed with 300μg/ml bismerthiazol or water) inoculated with the Xoo which had been transformed with exogenous fragment.
The results suggested that transformants numbered 646 and 518 confirmed their resistance to bismerthiazol, and the transformants numbered 5、33、115、152、119、102、172、519、537、611、643、501、694、518、612、654、308、919、670 were obviously stronger than that of the mutants 2-1-land PXO99. The transformants numbered 646 and 518 were subcloned, and the results revealed that two genes mutated. The two genes were raxR2 gene and conserved hypothetical protein(chp) gene. Seven bases mutated in the gene raxR2, but the amino acid sequence remained unchanged. Only one base mutated in the chp gene, resulting in the change of amino acid.
3. Study on the molecule mechanism of the streptomycin resistance in Xoo and Xooc
Xoo and Xooc were found to be sensitive to streptomycin, which is an inhibitor of protein synthesis resulting from interference with translational proofreading. Nineteen Xooc and eleven Xoo streptomycin resistant mutants were obtained by UV induction or streptomycin selection. These mutants could grow at 100μg mL-1 of streptomycin while the wild-type strain (RS105 and PXO99) cannot grow at 5μg mL"1 of streptomycin. The mutations at codon 43 and 88 in the rpsL gene might not be related to the reduced pathogenicity of Xoo and Xooc. Sequencing indicated that the rpsL gene has 375 bp encoding 125 amino acid residues. Blasting of the NCBI database indicated that the ORF had 97% similarity in nucleotide sequence and 100% similarity in deduced amino acid sequence with Xoo. In all resistant strains, a mutation in which AAG was substituted for AGG (Lys→Arg) occurred either at codon 43 or 88. Two plasmids, pUFRRS (containing a mutation at codon 43 of rpsL) and pUFRRX (containing a mutation at codon 88 of rpsL), were constructed by ligating the rpsL gene into the cosmid pUFR034. The plasmids pUFRRS and pUFRRX containing the Lys→Arg mutation of the rpsL gene conferred streptomycin resistance to the sensitive wild-type strain by electroporation. The transformants, RS1、RS2、PS1 and PS2, could grow in the medium containing 50μg mL-1 of streptomycin. A mutation at codon 43 or 88 in rpsL could result in resistance of Xoo to streptomycin. Meanwhile, The transformants with point mutation at codon 43 or 88 in the rpsL gene might not be related to the pathogenicity of Xoo and Xooc.
4. Detection of mutation at codon 43 of the rpsL gene in Xoo、Xooc、Ec and Xac by
PCR-RFLP
Twelve Ec、eleven Xac and twelve Ps streptomycin resistant mutants were obtained by UV induction or streptomycin selection. Sequencing indicated that the rpsL gene had a mutation at codon 43 or 88, which AAG was substituted for AGG (Lys→Arg).To develop a new method for detecting the point mutation of ribosomal S12 protein (rpsL) gene in streptomycin-resistant strains of Xoo, Xooc, Ec and Xac, the PCR-RELP method was developed to detect the point mutation of codon 43 of the rpsL gene in Xoo、Xooc、Ec and Xac. The 304-bp PCR product from the rpsL gene was digested by MboⅡinto two fragments (201 and 103 bp) if there was a mutation at codon 43, or into three fragments (146,103, and 55 bp) if there was no mutation. Compared with the results of nucleotide sequencing, the PCR-RFLP method was feasible in detecting the mutation in codon 43. The PCR-RFLP method could be used in the detection of the point mutation at codon 43 of the rpsL gene in streptomycin-resistant strains of Xoo, Xooc, Ec and Xac. PCR-RFLP is a simple, rapid, and reliable method for detection of the point mutation of codon 43 in the rpsL gene.
1.水稻白叶枯病菌噻枯唑抗性菌株粘粒基因组文库构建
为了探索水稻白叶枯病菌Xanthomonas oryzae pv. oryzae (Xoo)对噻枯唑的抗性机制,本研究建立了白叶枯病菌噻枯唑抗性菌株2-1-1的粘粒基因组文库。通过对2-1-1基因组提取并不完全酶切,回收>23.13kb基因片段,用来自溶源菌BHB2688和BHB2690、效价达1×108pfu/μgDNA的包装蛋白包装,感染宿主大肠杆菌S17-1,在含LB+Km+Sp的平板上进行筛选,随机挑选1200个转化子进行扩增保存,库容达到2.7倍；随机挑选15个转化子进行酶切验证,均有外源片段插入粘粒载体中,插入片段大小23-40kb,成功构建了抗噻枯唑白叶枯病菌2-1-1的粘粒基因组文库。
2.水稻白叶枯菌噻枯唑抗性菌株基因组文库的活体筛选和基因分析
将构建好的白叶枯病菌噻枯唑抗性菌株2-1-1粘粒基因组文库,1200个转化子通过两亲交配转染白叶枯病菌野生敏感菌株PXO99,获得的结合子再剪叶接种经300μg/ml噻枯唑处理的稻苗,筛选发现646、518等2个结合子表现抗药性；同时通过在清水对照处理的水稻上筛选,发现5、33、115、152、119、102、172、519、537、611、643、501、694、518、612、654、308、919、670等多个转化子在致病性上强于2-1-1和PXO99。通过对646和518片段进行亚克隆,进一步缩小功能片段,发现抗性菌株有两个基因发生突变。两基因分别为raxR2基因和conserved hypothetical protein (Chp)基因发生突变。其中raxR2基因有7个碱基发生突变,但未造成氨基酸序列变化；Chp基因发生一个碱基突变,引起了氨基酸变化(Val变成Gly)。
3.水稻白叶枯病菌和条斑病菌链霉素抗性机理研究
通过室内紫外诱变和药剂驯服获得了水稻条斑病菌和白叶枯病菌链霉素抗性突变体。这些突变体可以在含100μg/mL链霉素平板上生长,而敏感菌株在5μg/mL浓度下则不能生长。通过田间致病力测定,发现抗性菌株在水稻叶片上的致病力与敏感出发菌株无明显差异。从水稻条斑病菌和白叶枯病菌野生敏感菌株及室内诱导抗性菌株中分别扩增到编码核糖体蛋白S12编码基因(rpsL)全序列,序列分析表明rpsL基因中第43位或第88位氨基酸由赖氨酸(AAG)突变为精氨酸(AGG)。rpsL基因全长375bp,编码125个氨基酸。序列分析结果表明水稻条斑病菌与水稻白叶枯病菌rpsL基因同源性达97%,编码氨基酸同源性达100%；与柑橘溃疡病菌氨基酸同源性达98%,与大白菜软腐病菌氨基酸同源性达99%。设计合成1对含有酶切位点(EcoRI)的特异性引物,从抗药性菌株UV-R-1(含rpsL基因43位点突变)和UV-R-3(含rpsL基因88位点突变)中扩增出含启动区的rpsL基因,成功构建了重组质粒pUFRRS知pUFRRX。功能验证证明,质粒pUFRRS和pUFRRX能够互补敏感菌株的抗药性功能,结合子RS1、RS2、RS3、PS1、PS2对链霉素敏感性测定结果证实rpsL基因序列发生点突变是水稻白叶枯病菌和条斑病菌对链霉素产生抗药性的主要原因。同时转化子的致病力与敏感出发菌株和抗药突变体的致病力无明显差异。
4.水稻白叶枯病菌、条斑病菌、大白菜软腐病菌以及柑橘溃疡病菌抗药基因rpsL的分子检测
通过室内药剂驯化和紫外诱导,获得大白菜软腐病菌链霉素抗性突变体12个,柑橘溃疡病菌突变体11个,番茄细菌性斑点病菌突变体12个。分别比较番茄细菌性斑点病菌、柑橘溃疡病菌、大白菜软腐病菌链霉素抗性突变体与他们的敏感亲本菌株的序列,发现所有抗链霉素菌株的rpsL基因中第43位或第88位氨基酸均由赖氨酸(AAG)突变为精氨酸(AGG)。由于psL基因43位点是MboⅡ的酶切位点,因此通过PCR-RFLP方法可以检测出上述细菌rpsL基因43位点突变情况。通过设计一对特异性引物,扩增rpsL基因部分片段304bp,若43位点发生突变,则酶切位点消失,304bp则被酶切成201bp和103 bp片段；若43位点不发生突变,304bp则被酶切成201bp、103bp和55 bp片段。研究结果表明PCR-RELP方法可以快速、准确的检测出rpsL基因43位点突变情况,检测结果与直接测序结果完全一致,且操作简便。
Bacterial blight of rice (BB) and bacterial leaf streak of rice (BLS) were caused by Xanthomonas oryzae pv. oryzae(Xoo) and X. oryzae pv. oryzicola (Xooc), respectively. BB is one of the most serious diseases of rice, and BLS is emerging in importance. Application of bactericides is an indispensable complementary tool, especially in regions where susceptible cultivars are frequently planted. Bismerthiazol and streptomycin are mainly bactericides which have been used for the control of bacterial plant disease. But bismerthiazol and streptomycin have been used for more than thirty years. Thus, to study the resistance mechanism becomes very important.In this paper, there are five points about resisitance mechanism that were studied:
Construction of cosmid gene library of bismerthiazol-resistant Xoo.
In vivo selection of the cosmid library of bismerthiazol-resistant Xoo and gene analysis.
Study on the molecule mechanism of the streptomycin resistance in Xoo and Xooc.
Detection of the resistance to streptomycin in Xoo、Xooc、Erwinia carotovora (Ec) and Xanthomonas axonopodis pv.citri (Xac) by PCR-RFLP within the mutation at codon 43 of the rpsL gene.
1. Construction of the cosmid library of bismerthiazol-resistant Xoo
In order to explore the resistance mechanism of bismerthiazol, the cosmid genome library of the bismerthiazol-resistant Xoo mutant 2-1-1 was constructed. Through incomplete digestion of the genome of 2-1-1, the>23.13 kb gene fragments were gathered. Then, the packaging protein was extracted with the lysogen BHB2688、BHB2690, and the potency came up to 1×108pfu/μgDNA. The gathered gene fragments were connected, packed with packaging protein, and infected with the host S17-1. Then the combined mutants were selected on the LB+Km+Sp plates. A total of 1200 transformants were randomly selected at last and stored. The reservoir storage came up to 2.7 times. Fifteen transformants were randomly selected and digested with enzymes, the results indicated all the cosmids contained heterogenous insertional fragments and the cosmid genome library was successfully constructed, with average 23-40 kb gene fragments inserted.
2. In vivo selection of the cosmid library of bismerthiazol-resistant Xoo and gene analysis
The 1200 transformants described above were transformed by conjugation into PXO99 (a bismerthiazol-sensitive strain). Then, the in-vivo selection was done on the treated rice plants (sprayed with 300μg/ml bismerthiazol or water) inoculated with the Xoo which had been transformed with exogenous fragment.
The results suggested that transformants numbered 646 and 518 confirmed their resistance to bismerthiazol, and the transformants numbered 5、33、115、152、119、102、172、519、537、611、643、501、694、518、612、654、308、919、670 were obviously stronger than that of the mutants 2-1-land PXO99. The transformants numbered 646 and 518 were subcloned, and the results revealed that two genes mutated. The two genes were raxR2 gene and conserved hypothetical protein(chp) gene. Seven bases mutated in the gene raxR2, but the amino acid sequence remained unchanged. Only one base mutated in the chp gene, resulting in the change of amino acid.
3. Study on the molecule mechanism of the streptomycin resistance in Xoo and Xooc
Xoo and Xooc were found to be sensitive to streptomycin, which is an inhibitor of protein synthesis resulting from interference with translational proofreading. Nineteen Xooc and eleven Xoo streptomycin resistant mutants were obtained by UV induction or streptomycin selection. These mutants could grow at 100μg mL-1 of streptomycin while the wild-type strain (RS105 and PXO99) cannot grow at 5μg mL"1 of streptomycin. The mutations at codon 43 and 88 in the rpsL gene might not be related to the reduced pathogenicity of Xoo and Xooc. Sequencing indicated that the rpsL gene has 375 bp encoding 125 amino acid residues. Blasting of the NCBI database indicated that the ORF had 97% similarity in nucleotide sequence and 100% similarity in deduced amino acid sequence with Xoo. In all resistant strains, a mutation in which AAG was substituted for AGG (Lys→Arg) occurred either at codon 43 or 88. Two plasmids, pUFRRS (containing a mutation at codon 43 of rpsL) and pUFRRX (containing a mutation at codon 88 of rpsL), were constructed by ligating the rpsL gene into the cosmid pUFR034. The plasmids pUFRRS and pUFRRX containing the Lys→Arg mutation of the rpsL gene conferred streptomycin resistance to the sensitive wild-type strain by electroporation. The transformants, RS1、RS2、PS1 and PS2, could grow in the medium containing 50μg mL-1 of streptomycin. A mutation at codon 43 or 88 in rpsL could result in resistance of Xoo to streptomycin. Meanwhile, The transformants with point mutation at codon 43 or 88 in the rpsL gene might not be related to the pathogenicity of Xoo and Xooc.
4. Detection of mutation at codon 43 of the rpsL gene in Xoo、Xooc、Ec and Xac by
PCR-RFLP
Twelve Ec、eleven Xac and twelve Ps streptomycin resistant mutants were obtained by UV induction or streptomycin selection. Sequencing indicated that the rpsL gene had a mutation at codon 43 or 88, which AAG was substituted for AGG (Lys→Arg).To develop a new method for detecting the point mutation of ribosomal S12 protein (rpsL) gene in streptomycin-resistant strains of Xoo, Xooc, Ec and Xac, the PCR-RELP method was developed to detect the point mutation of codon 43 of the rpsL gene in Xoo、Xooc、Ec and Xac. The 304-bp PCR product from the rpsL gene was digested by MboⅡinto two fragments (201 and 103 bp) if there was a mutation at codon 43, or into three fragments (146,103, and 55 bp) if there was no mutation. Compared with the results of nucleotide sequencing, the PCR-RFLP method was feasible in detecting the mutation in codon 43. The PCR-RFLP method could be used in the detection of the point mutation at codon 43 of the rpsL gene in streptomycin-resistant strains of Xoo, Xooc, Ec and Xac. PCR-RFLP is a simple, rapid, and reliable method for detection of the point mutation of codon 43 in the rpsL gene.
引文
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