禾谷镰孢菌（Fusarium graminearum）对多菌灵抗药性分子机制的研究
摘要
本文克隆了禾谷镰孢菌野生型和多菌灵抗药性菌株的p2-微管蛋白基因(β2-tub)全序列,明确了β2-tub突变类型与该菌对多菌灵抗药性水平间的关系,并通过基因功能互补验证确定β2-tub为禾谷镰孢菌对多菌灵的抗药性基因；进一步通过人工点突变技术研究了β2-tub双点突变中各个突变与抗药性的关系,探讨了p1-微管蛋白基因的关键氨基酸的突变与抗药性的关系及其生物学功能。
以禾谷镰孢菌(Fusarium graminearum)标准菌株PH-1的β2-微管蛋白基因(β2-tub)部分序列(genbank登录号：FG06611.1)为参考,扩增了野生型菌株2021的β2-微管蛋白基因全长cDNA序列和基因全长序列。测序结果表明,野生型菌株2021的β2-微管蛋白基因编码区共1713bp,含有6个内含子,与标准菌株PH-1的β2-微管蛋白基因编码区相比有13个核苷酸的差异；该基因可编码477个氨基酸,其编码的第7位氨基酸(Ile)与标准菌株PH-1(Val)有差异；该β2-微管蛋白与其它8种真菌的苯并咪唑抗药性相关p-微管蛋白的同源性为74-77%。通过比较禾谷镰孢菌野生型菌株和多菌灵抗药性菌株的β2-tub序列发现：多菌灵中抗(MBCMR)菌株2052和R9等,β2-Tub第167位氨基酸由苯丙氨酸突变为酪氨酸；MBCMB菌株NT-7等,β2-Tub第200位氨基酸由苯丙氨酸突变为酪氨酸；多菌灵高抗(MBCHR)菌株JT04的β2-Tub第73位和198位氨基酸分别由谷氨酰胺和谷氨酸突变为精氨酸和亮氨酸；多菌灵高抗诱导(MBCIHR)菌株ZF43-6、ZF43-17和MBCHR菌株J2的β2-Tub第198位氨基酸由谷氨酸突变为赖氨酸；MBCIHR菌株52-7的β2-Tub第17位和167位氨基酸分别由甘氨酸和苯丙氨酸突变为丝氨酸和突变为酪氨酸。半定量RT-PCR分析表明：禾谷镰孢菌β2-tub在芽管和菌丝中表达量最高,在分生孢子和子囊孢子中的表达量次之；β1-tub在以上各阶段的表达水平无明显差异。
利用PEG介导的原生质体转化系统,将禾谷镰孢菌β2-微管蛋白基因(β2-tub)敲除DNA片段“1-up:PtrpC-pht:1.2-dow”分别导入禾谷镰孢菌野生型菌株2021、MBCMR菌株R9和MBCHR菌株JT04,利用潮霉素筛选获得转化子,使用PCR方法快速筛选获得了β2-tub敲除突变体。所有β2-tub敲除突变体对MBC均表现为超敏感,其菌丝生长速率、产分生孢子能力、产子囊壳能力及致病力均显著下降。分别将源自禾谷镰孢菌的野生型菌株2021、MBCMR菌株R9和MBCHR菌株JT04的β2-tub回复插入至禾谷镰孢菌β2-tub敲除体后,β2-tub回复插入突变体表现出与导入β2-tub相应的抗药性水平,其菌丝生长速率、产分生孢子能力、产子囊壳能力及致病力也基本恢复。结果表明,禾谷镰孢菌β2-微管蛋白(β2-Tub)关键氨基酸的突变可导致禾谷镰孢菌对MBC的抗药性；β2-微管蛋白在禾谷镰孢菌菌丝生长和繁殖过程中具有重要作用。
利用体外人工点突变和两步同源置换法成功获得了禾谷镰孢菌β2-微管蛋白(β2-Tub)第17位氨基酸(Gly→Ser)点突变体、73位氨基酸(Gln→Arg)点突变体、167位氨基酸(Phe→Tyr),点、198位氨基酸(Glu→Leu)点突变体、第17位(Gly→Ser)和167位Phe→Tyr)氨基酸双位点突变体、73位(Gln→Arg)和198位(Glu→Leu)氨基酸双位点突变体和240位氨基酸(Phe→Leu)突变体。结果显示,禾谷镰孢菌β2-Tub第17位氨基酸突变(Gly→Ser)可导致该菌对多菌灵的低抗,第73位氨基酸突变(Gln→Arg)不影响该菌对多菌灵的敏感性,第167位氨基酸突变(Phe→Tyr)可导致该菌对多菌灵中抗,第198位氨基酸突变(Glu→Leu)可导致该菌对多菌灵高抗,在167位氨基酸突变(Phe→Tyr)的基础上,第17位氨基酸突变(Gly→Ser)可导致该菌对多菌灵高抗,240位氨基酸突变(Phe→Leu)可导致该菌对多菌灵超敏感；以上所有类型的突变均对禾谷镰孢菌的生活力无显著影响。
禾谷镰孢菌p1-微管蛋白基因(β1-tub)并非禾谷镰孢菌的多菌灵抗性基因,但与β2-微管蛋白基因(β2-tub)相比,β1-tub与其它真菌中的苯并咪唑抗药性p-微管蛋白基因的同源性更高。为更好了解禾谷镰孢菌对苯并咪唑类杀菌剂的抗性机制和禾谷镰孢菌81-微管蛋白(β1-Tub)生物学功能。本文使用同源重组二步置换法将p1-微管蛋白第167位、第198位及第200位氨基酸进行人工点突变,突变方式分别为Phe167Tyr、 Glu198Lys和Phe200Tyr,并用二步同源置换法导入野生型β1-tub作为对照,用以评估基因操作对菌株的影响。结果显示,与对照菌株相比,禾谷镰孢菌β1-Tub第167位氨基酸突变体(Phe→Tyr).第198位氨基酸突变体(Glu→Lys)以及第200位氨基酸突变体(Phe→Tyr)对多菌灵、苯菌灵、噻菌灵和甲基托布津的敏感性均有所下降,但生物学特性无明显变化。禾谷镰孢菌β1-tub敲除突变体对多菌灵的敏感性略有下降,但其菌丝生长速度、产分生孢子能力、产子囊孢子能力和致病力均显著下降。
The β2-tubulin gene ((32-tub) of Fusaium graminearum (teleomorph:Gibberella zeae) was cloned by reverse transcriptase and polymerase chain reaction. The coding region of the P2-tub is1713bp length, including6introns, encoding477amino acids. Difference at thirteen nucleotides in coding region of P2-tubulin gene was detected by comparison wild-type isolates of Fusaium graminearum with standard isolate PH-1.But this causes only one difference in β2-tubulin at residue7[Val(PH-1)→Ile(2021)]. The homology of P2-tubulin in F. graminearum with benzimidalzole resistance β-tubulin in some other fungi is between74%and77%. The site mutation of β2-tubulin at codon167(Phe→Tyr) and codon200(Phe→Tyr) in F. graminearum may confer carbendazim (MBC) medium resistance; Site-mutation of β2-tubulin at its codon198(Glu→Lys\Leu) or at both codon17(Gly→Ser) and codonl65(Phe→Tyr) may confer MBC high resistance. Semi-quantative RT-PCR of β1-, P2-tubulin indicated that expression level of β2-tubulin in germ tubes and mycelia was higher than that in conidia and ascospores; expression of β1-tubulin in ascosores, conidia, germ tubes and mycelia was not significantly diferent.
The P2-tub was confirmed as MBC-resistance gene in F. graminearum by gene deletion and complementation. The P2-tub was deleted by homologous recombination of "l-up:PtrpC-pht:1.2-down" DNA fragment with the β2-tubulin locus, and then complemented by inserting plasmid vector pAMDS-β2tub into the genome of P2-tub deletion mutant strains. The β2-tubulin deletion mutants of wild-type strain2021, MBCMR strain R9and MBCHR strain JT04were super-sensitive to MBC, and the fitness of these β2-tub deletion mutants were significantly decreased as well. The complementation mutants possessing β2-tub origin from wild-type and MBC-resistant strains exhibited a MBC-sensitivity corresponding to their parental strains, and had a comparative fitness to their original strains as well.
Site-directed mutagenesis of β2-tub in F. graminearum wild-type strain2021at codon17,73,167,198and240were performed to clarify the relation between MBC-resistance level and mutation types. Mutation at codon17(Gly→Ser) in β2-tub conferred MBC low resistance; mutation at codon167(Phe→Tyr) in β2-tub conferred MBC medium resistance, double mutations at codon17(Gly→Ser) and codon167(Phe→Tyr)conferred MBC high resistance; mutation at codon73(Gln→Arg) in β2-tub did not confer MBC resistance, but muation at codon198conferred high resistance; mutation at codon240(Phe→Leu) in β2-tub made the mutant strains super-sensitive to MBC. All the mutants had a comparative fitness to the progenitor strain2021.
Two β-tubulin genes, which were known as β1-tubulin gene (β1-tub) and β2-tub, had been found in Fusarium graminearum. Benzimidazole resistance in Fusarium graminearum in the field is conferred by mutation in p2-tub but than β1-tub.β1-tubulin in F. graminearum, however, was highly identical to β-tubulin genes conferring benzimidazole resistance in some other fungi. To determine whether β1-tub of F. graminearum has the potential to confer carbendazim resistance, site-directed mutagenesis followed by gene replacement was used to change β1tub of F. graminearum wild-type strain2021at codon167(Phe to Tyr), codon198(Glu to Lys), and codon200(Phe to Tyr). Compared to the wild-type β1-tub revertant strains, all the β1-tub mutant strains (Phe167Tyr, Glu198Lys, and Phe200Tyr) and β1-tub deletion mutant strains had an increased resistance to carbendazim, benomyl, thiabendazole, and thiophanate-methyl; the resistance ratio of the mutant to the benzimidazole fungicides, however, was low. There were no significant changes among wild-type strain2021and β1-tub mutant strains in other biological properties. The fitness of β1-tub deletion mutant strains was significantly decresed comparing to the progenitor strain2021. In general, β1tub in F. graminearum has a potential to confer resistance to benzimidazole fungicides; the β1tub site-mutant strains had a comparative fitness to F. graminearum wild-type strain2021; the β1tub deletion-mutant strains had an increased sensitivity to MBC and a decreased fitness, than parental strain2021.
以禾谷镰孢菌(Fusarium graminearum)标准菌株PH-1的β2-微管蛋白基因(β2-tub)部分序列(genbank登录号：FG06611.1)为参考,扩增了野生型菌株2021的β2-微管蛋白基因全长cDNA序列和基因全长序列。测序结果表明,野生型菌株2021的β2-微管蛋白基因编码区共1713bp,含有6个内含子,与标准菌株PH-1的β2-微管蛋白基因编码区相比有13个核苷酸的差异；该基因可编码477个氨基酸,其编码的第7位氨基酸(Ile)与标准菌株PH-1(Val)有差异；该β2-微管蛋白与其它8种真菌的苯并咪唑抗药性相关p-微管蛋白的同源性为74-77%。通过比较禾谷镰孢菌野生型菌株和多菌灵抗药性菌株的β2-tub序列发现：多菌灵中抗(MBCMR)菌株2052和R9等,β2-Tub第167位氨基酸由苯丙氨酸突变为酪氨酸；MBCMB菌株NT-7等,β2-Tub第200位氨基酸由苯丙氨酸突变为酪氨酸；多菌灵高抗(MBCHR)菌株JT04的β2-Tub第73位和198位氨基酸分别由谷氨酰胺和谷氨酸突变为精氨酸和亮氨酸；多菌灵高抗诱导(MBCIHR)菌株ZF43-6、ZF43-17和MBCHR菌株J2的β2-Tub第198位氨基酸由谷氨酸突变为赖氨酸；MBCIHR菌株52-7的β2-Tub第17位和167位氨基酸分别由甘氨酸和苯丙氨酸突变为丝氨酸和突变为酪氨酸。半定量RT-PCR分析表明：禾谷镰孢菌β2-tub在芽管和菌丝中表达量最高,在分生孢子和子囊孢子中的表达量次之；β1-tub在以上各阶段的表达水平无明显差异。
利用PEG介导的原生质体转化系统,将禾谷镰孢菌β2-微管蛋白基因(β2-tub)敲除DNA片段“1-up:PtrpC-pht:1.2-dow”分别导入禾谷镰孢菌野生型菌株2021、MBCMR菌株R9和MBCHR菌株JT04,利用潮霉素筛选获得转化子,使用PCR方法快速筛选获得了β2-tub敲除突变体。所有β2-tub敲除突变体对MBC均表现为超敏感,其菌丝生长速率、产分生孢子能力、产子囊壳能力及致病力均显著下降。分别将源自禾谷镰孢菌的野生型菌株2021、MBCMR菌株R9和MBCHR菌株JT04的β2-tub回复插入至禾谷镰孢菌β2-tub敲除体后,β2-tub回复插入突变体表现出与导入β2-tub相应的抗药性水平,其菌丝生长速率、产分生孢子能力、产子囊壳能力及致病力也基本恢复。结果表明,禾谷镰孢菌β2-微管蛋白(β2-Tub)关键氨基酸的突变可导致禾谷镰孢菌对MBC的抗药性；β2-微管蛋白在禾谷镰孢菌菌丝生长和繁殖过程中具有重要作用。
利用体外人工点突变和两步同源置换法成功获得了禾谷镰孢菌β2-微管蛋白(β2-Tub)第17位氨基酸(Gly→Ser)点突变体、73位氨基酸(Gln→Arg)点突变体、167位氨基酸(Phe→Tyr),点、198位氨基酸(Glu→Leu)点突变体、第17位(Gly→Ser)和167位Phe→Tyr)氨基酸双位点突变体、73位(Gln→Arg)和198位(Glu→Leu)氨基酸双位点突变体和240位氨基酸(Phe→Leu)突变体。结果显示,禾谷镰孢菌β2-Tub第17位氨基酸突变(Gly→Ser)可导致该菌对多菌灵的低抗,第73位氨基酸突变(Gln→Arg)不影响该菌对多菌灵的敏感性,第167位氨基酸突变(Phe→Tyr)可导致该菌对多菌灵中抗,第198位氨基酸突变(Glu→Leu)可导致该菌对多菌灵高抗,在167位氨基酸突变(Phe→Tyr)的基础上,第17位氨基酸突变(Gly→Ser)可导致该菌对多菌灵高抗,240位氨基酸突变(Phe→Leu)可导致该菌对多菌灵超敏感；以上所有类型的突变均对禾谷镰孢菌的生活力无显著影响。
禾谷镰孢菌p1-微管蛋白基因(β1-tub)并非禾谷镰孢菌的多菌灵抗性基因,但与β2-微管蛋白基因(β2-tub)相比,β1-tub与其它真菌中的苯并咪唑抗药性p-微管蛋白基因的同源性更高。为更好了解禾谷镰孢菌对苯并咪唑类杀菌剂的抗性机制和禾谷镰孢菌81-微管蛋白(β1-Tub)生物学功能。本文使用同源重组二步置换法将p1-微管蛋白第167位、第198位及第200位氨基酸进行人工点突变,突变方式分别为Phe167Tyr、 Glu198Lys和Phe200Tyr,并用二步同源置换法导入野生型β1-tub作为对照,用以评估基因操作对菌株的影响。结果显示,与对照菌株相比,禾谷镰孢菌β1-Tub第167位氨基酸突变体(Phe→Tyr).第198位氨基酸突变体(Glu→Lys)以及第200位氨基酸突变体(Phe→Tyr)对多菌灵、苯菌灵、噻菌灵和甲基托布津的敏感性均有所下降,但生物学特性无明显变化。禾谷镰孢菌β1-tub敲除突变体对多菌灵的敏感性略有下降,但其菌丝生长速度、产分生孢子能力、产子囊孢子能力和致病力均显著下降。
The β2-tubulin gene ((32-tub) of Fusaium graminearum (teleomorph:Gibberella zeae) was cloned by reverse transcriptase and polymerase chain reaction. The coding region of the P2-tub is1713bp length, including6introns, encoding477amino acids. Difference at thirteen nucleotides in coding region of P2-tubulin gene was detected by comparison wild-type isolates of Fusaium graminearum with standard isolate PH-1.But this causes only one difference in β2-tubulin at residue7[Val(PH-1)→Ile(2021)]. The homology of P2-tubulin in F. graminearum with benzimidalzole resistance β-tubulin in some other fungi is between74%and77%. The site mutation of β2-tubulin at codon167(Phe→Tyr) and codon200(Phe→Tyr) in F. graminearum may confer carbendazim (MBC) medium resistance; Site-mutation of β2-tubulin at its codon198(Glu→Lys\Leu) or at both codon17(Gly→Ser) and codonl65(Phe→Tyr) may confer MBC high resistance. Semi-quantative RT-PCR of β1-, P2-tubulin indicated that expression level of β2-tubulin in germ tubes and mycelia was higher than that in conidia and ascospores; expression of β1-tubulin in ascosores, conidia, germ tubes and mycelia was not significantly diferent.
The P2-tub was confirmed as MBC-resistance gene in F. graminearum by gene deletion and complementation. The P2-tub was deleted by homologous recombination of "l-up:PtrpC-pht:1.2-down" DNA fragment with the β2-tubulin locus, and then complemented by inserting plasmid vector pAMDS-β2tub into the genome of P2-tub deletion mutant strains. The β2-tubulin deletion mutants of wild-type strain2021, MBCMR strain R9and MBCHR strain JT04were super-sensitive to MBC, and the fitness of these β2-tub deletion mutants were significantly decreased as well. The complementation mutants possessing β2-tub origin from wild-type and MBC-resistant strains exhibited a MBC-sensitivity corresponding to their parental strains, and had a comparative fitness to their original strains as well.
Site-directed mutagenesis of β2-tub in F. graminearum wild-type strain2021at codon17,73,167,198and240were performed to clarify the relation between MBC-resistance level and mutation types. Mutation at codon17(Gly→Ser) in β2-tub conferred MBC low resistance; mutation at codon167(Phe→Tyr) in β2-tub conferred MBC medium resistance, double mutations at codon17(Gly→Ser) and codon167(Phe→Tyr)conferred MBC high resistance; mutation at codon73(Gln→Arg) in β2-tub did not confer MBC resistance, but muation at codon198conferred high resistance; mutation at codon240(Phe→Leu) in β2-tub made the mutant strains super-sensitive to MBC. All the mutants had a comparative fitness to the progenitor strain2021.
Two β-tubulin genes, which were known as β1-tubulin gene (β1-tub) and β2-tub, had been found in Fusarium graminearum. Benzimidazole resistance in Fusarium graminearum in the field is conferred by mutation in p2-tub but than β1-tub.β1-tubulin in F. graminearum, however, was highly identical to β-tubulin genes conferring benzimidazole resistance in some other fungi. To determine whether β1-tub of F. graminearum has the potential to confer carbendazim resistance, site-directed mutagenesis followed by gene replacement was used to change β1tub of F. graminearum wild-type strain2021at codon167(Phe to Tyr), codon198(Glu to Lys), and codon200(Phe to Tyr). Compared to the wild-type β1-tub revertant strains, all the β1-tub mutant strains (Phe167Tyr, Glu198Lys, and Phe200Tyr) and β1-tub deletion mutant strains had an increased resistance to carbendazim, benomyl, thiabendazole, and thiophanate-methyl; the resistance ratio of the mutant to the benzimidazole fungicides, however, was low. There were no significant changes among wild-type strain2021and β1-tub mutant strains in other biological properties. The fitness of β1-tub deletion mutant strains was significantly decresed comparing to the progenitor strain2021. In general, β1tub in F. graminearum has a potential to confer resistance to benzimidazole fungicides; the β1tub site-mutant strains had a comparative fitness to F. graminearum wild-type strain2021; the β1tub deletion-mutant strains had an increased sensitivity to MBC and a decreased fitness, than parental strain2021.
引文
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1. Davidse L C. Benzimidazole fungicides:mechanism of action and biological impact [J]. Annual Review of Phytopathology,1986,24:43-65
2. Fujimura M, Kanakura T, Yamaguchi I. Action mechanism of diethofencarb to a benzimidazole-resistant mutant in Neurospora crassa [J]. Journal of Pesticide Science,1992, 17:237-242
3. Yarden O, Katan T. Mutations leading to substitutions at amino acides 198 and 200 on beta-tubulin that correlates with benomyl resistance phenotypes of fied strains of Botrytis cinerea [J]. Phytopathology,1993,83:850-854
4. Yan K, Dickman M. Isolation of a β-tubulin gene from Fusarium moniliforme that confers cold-sensitive benomyl resistance [J]. Applied and Enviromental Microbiology,1996,62 (8): 3053-3056.
5. Fujimura M, Kamakura T, Inoue H, et al. Sensitivity of Neurospora crassa to benzimidazoles and N-phenylcarbamates:effect of amino acid substitutions at position 198 in β-tubulin. Pesticide Biochemistry Physiology,1992,44 (3):165-173
6. Koeneaadt H, Jones A L. Resistance to benomyl conferred by mutations in codon 198 or 200 in the beta-tubulin gene of Neurospora crassa and sensitivity to diethofencarb conferred by codon 198. Photopathology,1993,83:850-854
7. Yarden O, Katan T. Mutations leading to substitutions at amino acids 198 and 200 of beta-tubulin that correlate with benomyl-resistance phenotypes of field strains of Botrytis cinerea. Phytopathology,1993,83:1478-1483
8. Baraldi E, Mari M, Chierici E, Pondrelli M, Bertolini P, Pratella G C. Studies on thiabendazole resistance of Penicillium expansum of pears, pathogenic fitness and genetic characterization. Plant Pathology,2003,52:362-370
9. Gafur A, Tanaka C, Shimizu K, Ouchi S, Tsuda M. Molecular analysis and characterization of the Cochliobolus heterostrophus beta-tubulin gene and its possible role in conferring resistance to benomyl [J]. The Journal of General and Applied Microbiology,1998,44:217-223.
10. Wu H., Liu X. and Jaenisch R. Double replacement:Strategy for efficient introduction of subtle mutations into the murine Coila-i gene by homologous recombination in embryonic stem cells [J]. Proceedings of the National Academy of Sciences,1994,91:2819-2823
11. Yu J H, Hamari Z, Han K H, Seo J A, Reyes-Dominguez Y, Scazzocchio C. Double-joint PCR:a PCR-based molecular tool for gene manipulations in filamentous fungi [J]. Fungal Genetics and Biology,2004,41:973-981
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13. Ma Z, Yoshimura M, Michailides T J. Identification and characterization of benzimidazole resistance in Monilinia fructicola from stone fruit orchards in California [J]. Applied and Environmental Microbiology,2003,69:7145-7152
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