棘孢木霉（Trichoderma asperellum）MAPK家族基因的克隆及生物防治功能研究
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摘要
棘孢木霉（T.asperellum）是重要的生物防治菌，通过多种机制防治植物土传病菌的危害。这些机制已经被广泛研究，但是其生物防治的分子机制仍不清楚。为了掌握棘孢木霉（T.asperellum）生物防治的分子调控机制，本论文以棘孢木霉（T.asperellum）菌丝体时期的cDNA文库为基础，克隆了与生物防治相关的调控基因：丝裂原活化蛋白激酶基因task1和裂原活化蛋白激酶激酶激酶基因taskkk，并对其生物防治功能进行研究。
首先克隆了task1基因和taskkk基因及两基因的侧翼序列，并对两基因进行序列分析。然后构建双元重组质粒，以潮霉素抗性基因作为筛选标记，采用根癌农杆菌介导法转化棘孢木霉（T.asperellum，）获得task1基因敲除突变体Δtask1和taskkk基因敲除突变体Δtaskkk。
将突变体Δtask1和Δtaskkk分别与棘孢木霉野生菌体进行对比，研究task1和taskkk基因对棘孢木霉菌体形态和生长的影响。结果表明：task1基因影响棘孢木霉菌丝生长、分裂、孢子的形成和菌体的生长状态，对生长速度和生长量没有影响；taskkk基因影响孢子的形成，而与菌丝分裂、菌体生长状态、生长速度和生长量无关。
然后，对task1和taskkk基因对棘孢木霉重寄生作用的影响进行了研究。结果表明：棘孢木霉突变体Δtask1能识别病原菌，但不能寄生病原菌，只是沿菌丝表面生长，产生细胞壁水解酶，水解病原菌，说明task1基因与棘孢木霉对病原菌的寄生有关；突变体Δtaskkk并不识别致病菌，与病原菌各自生长，说明taskkk基因与棘孢木霉重寄生过程中对致病菌的识别相关。对棘孢木霉、突变体Δtask1和Δtaskkk重寄生过程中的水解酶表达和代谢产生的水解酶活性分析表明：task1基因对纤维素酶基因，β-1，3葡聚糖酶基因，β-1，6葡聚糖酶基因，N-乙酰-氨基葡萄糖苷酶1，β-1，4葡聚糖酶基因和聚酮合酶基因的表达起负调控作用；taskkk基因对几丁质酶基因，N-乙酰-氨基葡萄糖苷酶2，β-1，3葡聚糖酶基因，β-1，6葡聚糖酶基因和聚酮合酶基因起正调控作用。对棘孢木霉、突变体Δtask1和Δtaskkk在不同碳源诱导条件下的水解酶活性分析表明：task1基因与N-乙酰氨基葡萄糖苷酶、β-1，3葡聚糖酶、纤维素酶和β-葡萄糖苷酶的产生有关，taskkk基因与几丁质酶、N-乙酰氨基葡萄糖苷酶和β-1，3葡聚糖酶的产生有关。
最后，对task1和taskkk基因对棘孢木霉抗生作用的影响进行了研究。结果表明：task1基因负调控棘孢木霉聚酮合酶基因的表达，而taskkk基因对聚酮合酶基因起正调控作用，所以task1和taskkk基因在聚酮合酶基因的表达上不在同一条信号传导途径上；敲除task1基因后突变体抗真菌能力提高，而敲除taskkk基因后突变体抗真菌能力降低，说明两基因与抗真菌代谢物的合成都有关，但是两基因作用相反；然后，研究了task1和taskkk基因对棘孢木霉代谢产生的抗生素6-戊基-α-吡喃酮的影响，棘孢木霉野生型代谢产生的6-戊基-α-吡喃酮的量为1.32mg/g菌丝干重，突变体Δtask1为2.80mg/g菌丝干重，是野生型的2.12倍，突变体Δtaskkk代谢产生的6-戊基-α-吡喃酮的量为0.11mg/g菌丝干重，是野生型的0.08倍，说明task1基因对6-戊基-α-吡喃酮合成酶基因起负调控作用，taskkk基因是棘孢木霉代谢产物6-戊基-α-吡喃酮合成酶基因的正调控基因，task1和taskkk基因在6-戊基-α-吡喃酮的合成上不在同一条信号传导途径上。
Trichoderma asperellum is important biocontrol fungus and control plant soil-borne pathogens through a variety of mechanisms. Though those mechanisms have been researched extensively, its biocontrol molecular mechanisms remain unclear. In order to master the biocontrol molecular mechanism of T. asperellum, the regulating genes related to biological control:taskl and taskkk were cloned based on cDNA library from T. asperellum mycelium and their biocontrol related functions were studied.
Firstly, gene taskl, taskkk and their flanking sequences were cloned; the sequences of two genes were analysed. Recombinant binary vectors were constructed using hygromycin resistance gene as a selection marker. Transformation of T. asperellum utilized the method of Agrobacterium tumefaciens-mediated homologous recombination, the gene knock out mutants Ataskl and Ataskkk were obtained
Through comparing mutant Δtask1and Δtaskkk with T. asperellum wild type, the influences of gene taskl and taskkk on T. asperellum growth were studied. The result showed that gene taskl had an effect on mycelia growth, mitosis, spore formation and colony growth state, did not affect colony growth speed and quality; gene taskkk was influential to spore formation, did not influence mycelia growth, mitosis, colony growth state, colony growth speed and quality.
Then, the influences of gene taskl and taskkk on T. asperellum mycoparasitic effects were studied. The experiment results indicated that mutant Ataskl could recognize and adhere to R. solani, produce cell wall hydrolytic enzyme and hydrolyze R. solani, but could not mycoparasite it, that means gene taskl has anything to do with mycoparasitic effects of T. asperellum; mutant Ataskkk could not recognize and mycoparasite R. solani and growth separately, which means gene taskkk has anything to do with recognition and mycoparasitic effects of T. asperellum. The hydrolases gene expression and activity analysis of T. asperellum, mutant Ataskl and Ataskkk showed that gene taskl down regulated expression of cellulase, β-1,3glucanase, β-1,6glucanase, N-acetyl glucosaminidasel and β-1,4glucanase gene; taskkk gene up regulated expression of gene chitinase, N-acetyl glucosaminidase2, β-1,3glucanase and β-1,6glucanase gene. The hydrolases activity analysis related to biocontrol of T. asperellum, mutant Ataskl and Ataskkk indicated that gene taskl related to production of N-acetyl-glucosaminidase, β-1,3glucanase, cellulose and β-glucosidase, gene taskkk related to production of chitinase, N-acetyl-glucosaminidase and β-1,3glucanase.
Finally, the influences of gene taskl and taskkk on T. asperellum antibiosis were studied. The experiment results indicated that gene taskl down regulated expression of polyketide synthase, but gene taskkk up regulated expression of it, which means that gene taskl and taskkk is not on the same signaling pathways. Antifungal ability of mutant Ataskl was higher than T. asperellum and mutant Ataskkk was lower than T. asperellum, which indicated the two genes related to the production of antifungal metabolites, only with the opposite effect. The influences of gene taskl and taskkk on6-amyl-a-pyrone from T. asperellum were studied. The yield of6-amyl-a-pyrone from T. asperellum wild type was1.32mg/g mycelial dry weight, mutant Ataskl was2.80mg6-amyl-α-pyrone/g mycelial dry weight,2.12times of T. asperellum wild type, and mutant Ataskkk was0.11mg6-amyl-a-pyrone/g mycelial dry weight,0.08times of T. asperellum wild type. Those showed that gene tasklwas negative regulation gene of6-amyl-a-pyrone, gene taskkk was up regulation gene of it, gene taskl and taskkk was not on the same signaling pathways.
首先克隆了task1基因和taskkk基因及两基因的侧翼序列，并对两基因进行序列分析。然后构建双元重组质粒，以潮霉素抗性基因作为筛选标记，采用根癌农杆菌介导法转化棘孢木霉（T.asperellum，）获得task1基因敲除突变体Δtask1和taskkk基因敲除突变体Δtaskkk。
将突变体Δtask1和Δtaskkk分别与棘孢木霉野生菌体进行对比，研究task1和taskkk基因对棘孢木霉菌体形态和生长的影响。结果表明：task1基因影响棘孢木霉菌丝生长、分裂、孢子的形成和菌体的生长状态，对生长速度和生长量没有影响；taskkk基因影响孢子的形成，而与菌丝分裂、菌体生长状态、生长速度和生长量无关。
然后，对task1和taskkk基因对棘孢木霉重寄生作用的影响进行了研究。结果表明：棘孢木霉突变体Δtask1能识别病原菌，但不能寄生病原菌，只是沿菌丝表面生长，产生细胞壁水解酶，水解病原菌，说明task1基因与棘孢木霉对病原菌的寄生有关；突变体Δtaskkk并不识别致病菌，与病原菌各自生长，说明taskkk基因与棘孢木霉重寄生过程中对致病菌的识别相关。对棘孢木霉、突变体Δtask1和Δtaskkk重寄生过程中的水解酶表达和代谢产生的水解酶活性分析表明：task1基因对纤维素酶基因，β-1，3葡聚糖酶基因，β-1，6葡聚糖酶基因，N-乙酰-氨基葡萄糖苷酶1，β-1，4葡聚糖酶基因和聚酮合酶基因的表达起负调控作用；taskkk基因对几丁质酶基因，N-乙酰-氨基葡萄糖苷酶2，β-1，3葡聚糖酶基因，β-1，6葡聚糖酶基因和聚酮合酶基因起正调控作用。对棘孢木霉、突变体Δtask1和Δtaskkk在不同碳源诱导条件下的水解酶活性分析表明：task1基因与N-乙酰氨基葡萄糖苷酶、β-1，3葡聚糖酶、纤维素酶和β-葡萄糖苷酶的产生有关，taskkk基因与几丁质酶、N-乙酰氨基葡萄糖苷酶和β-1，3葡聚糖酶的产生有关。
最后，对task1和taskkk基因对棘孢木霉抗生作用的影响进行了研究。结果表明：task1基因负调控棘孢木霉聚酮合酶基因的表达，而taskkk基因对聚酮合酶基因起正调控作用，所以task1和taskkk基因在聚酮合酶基因的表达上不在同一条信号传导途径上；敲除task1基因后突变体抗真菌能力提高，而敲除taskkk基因后突变体抗真菌能力降低，说明两基因与抗真菌代谢物的合成都有关，但是两基因作用相反；然后，研究了task1和taskkk基因对棘孢木霉代谢产生的抗生素6-戊基-α-吡喃酮的影响，棘孢木霉野生型代谢产生的6-戊基-α-吡喃酮的量为1.32mg/g菌丝干重，突变体Δtask1为2.80mg/g菌丝干重，是野生型的2.12倍，突变体Δtaskkk代谢产生的6-戊基-α-吡喃酮的量为0.11mg/g菌丝干重，是野生型的0.08倍，说明task1基因对6-戊基-α-吡喃酮合成酶基因起负调控作用，taskkk基因是棘孢木霉代谢产物6-戊基-α-吡喃酮合成酶基因的正调控基因，task1和taskkk基因在6-戊基-α-吡喃酮的合成上不在同一条信号传导途径上。
Trichoderma asperellum is important biocontrol fungus and control plant soil-borne pathogens through a variety of mechanisms. Though those mechanisms have been researched extensively, its biocontrol molecular mechanisms remain unclear. In order to master the biocontrol molecular mechanism of T. asperellum, the regulating genes related to biological control:taskl and taskkk were cloned based on cDNA library from T. asperellum mycelium and their biocontrol related functions were studied.
Firstly, gene taskl, taskkk and their flanking sequences were cloned; the sequences of two genes were analysed. Recombinant binary vectors were constructed using hygromycin resistance gene as a selection marker. Transformation of T. asperellum utilized the method of Agrobacterium tumefaciens-mediated homologous recombination, the gene knock out mutants Ataskl and Ataskkk were obtained
Through comparing mutant Δtask1and Δtaskkk with T. asperellum wild type, the influences of gene taskl and taskkk on T. asperellum growth were studied. The result showed that gene taskl had an effect on mycelia growth, mitosis, spore formation and colony growth state, did not affect colony growth speed and quality; gene taskkk was influential to spore formation, did not influence mycelia growth, mitosis, colony growth state, colony growth speed and quality.
Then, the influences of gene taskl and taskkk on T. asperellum mycoparasitic effects were studied. The experiment results indicated that mutant Ataskl could recognize and adhere to R. solani, produce cell wall hydrolytic enzyme and hydrolyze R. solani, but could not mycoparasite it, that means gene taskl has anything to do with mycoparasitic effects of T. asperellum; mutant Ataskkk could not recognize and mycoparasite R. solani and growth separately, which means gene taskkk has anything to do with recognition and mycoparasitic effects of T. asperellum. The hydrolases gene expression and activity analysis of T. asperellum, mutant Ataskl and Ataskkk showed that gene taskl down regulated expression of cellulase, β-1,3glucanase, β-1,6glucanase, N-acetyl glucosaminidasel and β-1,4glucanase gene; taskkk gene up regulated expression of gene chitinase, N-acetyl glucosaminidase2, β-1,3glucanase and β-1,6glucanase gene. The hydrolases activity analysis related to biocontrol of T. asperellum, mutant Ataskl and Ataskkk indicated that gene taskl related to production of N-acetyl-glucosaminidase, β-1,3glucanase, cellulose and β-glucosidase, gene taskkk related to production of chitinase, N-acetyl-glucosaminidase and β-1,3glucanase.
Finally, the influences of gene taskl and taskkk on T. asperellum antibiosis were studied. The experiment results indicated that gene taskl down regulated expression of polyketide synthase, but gene taskkk up regulated expression of it, which means that gene taskl and taskkk is not on the same signaling pathways. Antifungal ability of mutant Ataskl was higher than T. asperellum and mutant Ataskkk was lower than T. asperellum, which indicated the two genes related to the production of antifungal metabolites, only with the opposite effect. The influences of gene taskl and taskkk on6-amyl-a-pyrone from T. asperellum were studied. The yield of6-amyl-a-pyrone from T. asperellum wild type was1.32mg/g mycelial dry weight, mutant Ataskl was2.80mg6-amyl-α-pyrone/g mycelial dry weight,2.12times of T. asperellum wild type, and mutant Ataskkk was0.11mg6-amyl-a-pyrone/g mycelial dry weight,0.08times of T. asperellum wild type. Those showed that gene tasklwas negative regulation gene of6-amyl-a-pyrone, gene taskkk was up regulation gene of it, gene taskl and taskkk was not on the same signaling pathways.
引文
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