HOG信号途径和VeA调控系统对小麦赤霉病菌DON毒素合成的调控机制研究
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摘要
赤霉病不仅影响小麦的安全生产,其病原禾谷镰刀菌(Fusarium graminearum)产生的真菌毒素严重威胁农产品质量安全。目前一些杀菌剂尽管能有效控制赤霉病,但反而会刺激镰刀菌产生更多的毒素。因此,在赤霉病持续防控上,开发既能防病又能抑制毒素的新型药剂显得尤为重要。前期研究初步表明,双组份组氨酸激酶信号途径(HOG信号途径)和VeA调控系统可能参与毒素的合成调控,为了更好的了解DON毒素合成的调控机制,本文系统的研究了小麦赤霉病菌HOG信号途径以及VeA调控系统中35个元件的生物学功能,研究发现：
1)赤霉病菌HOG信号途径的反应调控元件FgRrg1不仅参与调控病菌毒素和色素的合成,而且控制病菌的致病性以及对逆境的适应性。FgRRG-1基因敲除突变体对氯化钠、氯化钾、山梨醇等渗透胁迫以及金属离子胁迫都高度敏感,对二甲酰亚胺类药剂扑海因和吡咯类杀菌剂适乐时表现高抗,同时突变体在麦穗上的致病性显著降低,DON毒素合成量减少。
2)赤霉病菌的2C型丝氨酸/苏氨酸磷酸酶FgPtc3是HOG信号途径的负调控元件,FgPtc3不仅调控病菌毒素以及色素的合成,同时参与菌体生长发育过程以及多个代谢合成途径。PTC3基因缺失突变体生长缓慢,气生菌丝减少,产孢增加,细胞内甘油合成显著增加,脂肪体增多,对渗透以及细胞壁胁迫抗性增加,更重要的是,缺失突变体完全失去致病性,DON毒素合成显著降低,SAGE表达谱测定发现FgPtc3参与菌体多个代谢合成途径。FgPTC3基因能够恢复酵母PTc1突变体生长缺陷性,降低酵母体内Hog1磷酸化水平。并且研究发现FgPtc3具有PP2C磷酸酶活性,可以通过体外表达蛋白筛选该蛋白相应的活性抑制剂。
3)赤霉病菌的酪氨酸磷酸酶FgPtp2与HOG信号途径没有明显的关联,但是该蛋白参与调控病菌的毒素合成以及多个生长代谢途径。FgPTP2基因敲除突变体生长非常缓慢,气生菌丝减少,产孢减少,但是对各种胁迫的敏感性没有发生改变,并且突变体致病性降低,DON毒素合成也降低,SAGE测定突变体基因组表达谱发现,FgPtp2参与调控多个代谢途径,能够调控菌体核糖体上相关蛋白的合成,从而影响菌体正常生长。
4)赤霉病菌VeA调控系统中的FgVeA和FgVe1B在功能上有很多相似性,参与调控菌体的毒素合成以及多个生长代谢过程。FgVEA和FgVELB都与菌体生长发育密切相关,敲除突变体生长缓慢,气生菌丝减少,菌体表面疏水性降低。突变体产孢增加,对氯化钠,氯化钾等渗透胁迫,以及细胞壁胁迫抗性增加,对扑海因以及适乐时等杀菌剂敏感,并且菌体中甘油积累量也普遍高于野生型。FgVe1B与FgVeA都参与菌体的脂肪体,毒素,色素合成等多个次生代谢途径,并且控制病菌的致病性。赤霉病菌VeA调控系统中的LaeA同源基因不是菌体生长所必须的,基因的敲除除了对色素合成有微弱影响之外,对菌体的生长以及毒素的合成并没有影响。
本文研究结果表明小麦赤霉病菌HOG信号途径的FgRrg1和FgPtc3以及VeA调控系统中的FgVeA和FgVe1B元件控制病菌毒素合成和病菌的致病性,可能成为潜在的杀菌剂新药靶。本研究结果为开发既能防病又能抑制毒素的新型药剂创制提供新线索。
Fusarium graminearum is the major causal agent of Fusarium head blight (FHB) on wheat, which causes yield loss and affects the quality of grains. The mycotoxins, such as deoxynivalenol (DON) and its derivatives produced by the fungus in infected grains, pose a serious threat to human and animal health. Despite the FHB can be effectively controlled by some fungicides, fungicides application may stimulate toxin production by the fungus. Therefore, the development of novel anti-mycotoxin and antifungal agents is urgently needed for effective management of FHB and mycotoxin contamination in cereals. Previous research has revealed the involvement of two-component histidine kinase signaling pathway (HOG signaling pathway) and VeA regulatory system in the regulation of DON biosynthesis, to get a better insight into the regulation mechanism of DON synthesis in F. graminearum, the biological functions of35genes involved in the HOG signaling pathway and VeA regulatory system were studied, results of our research showed that:
1) The response regulator FgRrgl of F. graminearum in the HOG signaling pathway is involved in the regulation of DON and aurofusarin biosynthesis, playing important roles in pathogenicity and adaptation of fungi to a variety of environmental stresses. The FgRRG-1deletion mutant showed increased sensitivity to osmotic stress mediated by NaCl, KCl, sorbitol or glucose, and to metal cations Li+, Ca2+and Mg2+The mutant, however, was more resistant than the parent strain to dicarboximide and phenylpyrrole fungicides. Inoculation tests showed that the mutant exhibited decreased virulence on wheat heads, and produced a significantly low level of DON.
2) The type2C protein phosphatases FgPtc3is the negative regulator of the HOG signaling pathway in F. graminearum. FgPtc3is involved in the regulation of DON and aurofusarin biosynthesis, and also plays important roles in regulating many development processes and secondary metabolisms. The FgPTC3deletion mutant exhibited reduced aerial hyphae formation and deoxynivalenol (DON) production, but increased production of conidia. The mutant showed increased resistance to osmotic stress and cell wall-damaging agents, and accumulated a higher basal level of glycerol than the wild-type progenitor. Ultrastructural and histochemical analyses showed that conidia of the mutant contained an unusually high number of large lipid droplets. Pathogencity assays showed that the mutant is unable to infect flowering wheat head. Serial analysis of gene expression in the deletion mutant revealed that FgPtc3is associated with various metabolic pathways. Additionally, the FgPTC3partially rescued the growth defect of a yeast PTC1deletion mutant under various stress conditions, and also reduced the level of phosphorylated Hogl. The FgPtc3has the PP2C phosphatase activity, which may help in exploitation of drug targets based on screening of the protein activity inhibitors.
3) The tyrosine phosphatase FgPtp2is not associated with the HOG signaling pathway, but plays important roles in the regulation of DON biosynthesis and secondary metabolisms. The FgPTP2deletion mutant exhibited a significant decrease in mycelial growth and conidiation, but do not change the sensitivity to variety stresses. In addition, the deletion mutant was impaired dramatically in virulence and produced a low level of DON. Serial analysis of gene expression in the mutant revealed that FgPtp2is associated with various metabolic pathways, especially with the synthesis of ribosomal protein.
4) The FgVeA and FgVelB in the VeA regulatory system have some common functions in regulation of DON biosynthesis, and various cellular processes. The FgVEA and FgVELB deletion mutant exhibited a reduction in aerial hyphae formation, hydrophobicity, and deoxynivalenol (DON) biosynthesis, but increased the conidial production. Sensitivity tests to various stresses showed that the mutants exhibited increased resistance to osmotic stress and cell wall-damaging agents, but increased sensitivity to iprodione and fludioxonil fungicides, and the mutants contained an unusually high number of large lipid droplets, which is in agreement with the observation that the mutant accumulated a higher basal level of glycerol than the wild-type progenitor. Both FgVeA and FgVe1B play a critical role in pathogenicity,in a variety of biological functions and secondary metabolisms. The LaeA homologous genes are not essential for mycelial growth in F. graminearum, the deletion of the genes caused minor influence in pigment synthesis, and have no effect on DON production.
Results of the research indicated that the FgRrgl and FgPtc3in HOG signaling pathway, as well as FgVeA and FgVe1B in VeA regulatory system are involved in pathogenicity and DON biosynthesis, which may represent attractive antimicrobial targets for disease control purposes. The results aslo may help in exploitation of drug targets for the design of new anti-mycotoxin and antifungal agents.
1)赤霉病菌HOG信号途径的反应调控元件FgRrg1不仅参与调控病菌毒素和色素的合成,而且控制病菌的致病性以及对逆境的适应性。FgRRG-1基因敲除突变体对氯化钠、氯化钾、山梨醇等渗透胁迫以及金属离子胁迫都高度敏感,对二甲酰亚胺类药剂扑海因和吡咯类杀菌剂适乐时表现高抗,同时突变体在麦穗上的致病性显著降低,DON毒素合成量减少。
2)赤霉病菌的2C型丝氨酸/苏氨酸磷酸酶FgPtc3是HOG信号途径的负调控元件,FgPtc3不仅调控病菌毒素以及色素的合成,同时参与菌体生长发育过程以及多个代谢合成途径。PTC3基因缺失突变体生长缓慢,气生菌丝减少,产孢增加,细胞内甘油合成显著增加,脂肪体增多,对渗透以及细胞壁胁迫抗性增加,更重要的是,缺失突变体完全失去致病性,DON毒素合成显著降低,SAGE表达谱测定发现FgPtc3参与菌体多个代谢合成途径。FgPTC3基因能够恢复酵母PTc1突变体生长缺陷性,降低酵母体内Hog1磷酸化水平。并且研究发现FgPtc3具有PP2C磷酸酶活性,可以通过体外表达蛋白筛选该蛋白相应的活性抑制剂。
3)赤霉病菌的酪氨酸磷酸酶FgPtp2与HOG信号途径没有明显的关联,但是该蛋白参与调控病菌的毒素合成以及多个生长代谢途径。FgPTP2基因敲除突变体生长非常缓慢,气生菌丝减少,产孢减少,但是对各种胁迫的敏感性没有发生改变,并且突变体致病性降低,DON毒素合成也降低,SAGE测定突变体基因组表达谱发现,FgPtp2参与调控多个代谢途径,能够调控菌体核糖体上相关蛋白的合成,从而影响菌体正常生长。
4)赤霉病菌VeA调控系统中的FgVeA和FgVe1B在功能上有很多相似性,参与调控菌体的毒素合成以及多个生长代谢过程。FgVEA和FgVELB都与菌体生长发育密切相关,敲除突变体生长缓慢,气生菌丝减少,菌体表面疏水性降低。突变体产孢增加,对氯化钠,氯化钾等渗透胁迫,以及细胞壁胁迫抗性增加,对扑海因以及适乐时等杀菌剂敏感,并且菌体中甘油积累量也普遍高于野生型。FgVe1B与FgVeA都参与菌体的脂肪体,毒素,色素合成等多个次生代谢途径,并且控制病菌的致病性。赤霉病菌VeA调控系统中的LaeA同源基因不是菌体生长所必须的,基因的敲除除了对色素合成有微弱影响之外,对菌体的生长以及毒素的合成并没有影响。
本文研究结果表明小麦赤霉病菌HOG信号途径的FgRrg1和FgPtc3以及VeA调控系统中的FgVeA和FgVe1B元件控制病菌毒素合成和病菌的致病性,可能成为潜在的杀菌剂新药靶。本研究结果为开发既能防病又能抑制毒素的新型药剂创制提供新线索。
Fusarium graminearum is the major causal agent of Fusarium head blight (FHB) on wheat, which causes yield loss and affects the quality of grains. The mycotoxins, such as deoxynivalenol (DON) and its derivatives produced by the fungus in infected grains, pose a serious threat to human and animal health. Despite the FHB can be effectively controlled by some fungicides, fungicides application may stimulate toxin production by the fungus. Therefore, the development of novel anti-mycotoxin and antifungal agents is urgently needed for effective management of FHB and mycotoxin contamination in cereals. Previous research has revealed the involvement of two-component histidine kinase signaling pathway (HOG signaling pathway) and VeA regulatory system in the regulation of DON biosynthesis, to get a better insight into the regulation mechanism of DON synthesis in F. graminearum, the biological functions of35genes involved in the HOG signaling pathway and VeA regulatory system were studied, results of our research showed that:
1) The response regulator FgRrgl of F. graminearum in the HOG signaling pathway is involved in the regulation of DON and aurofusarin biosynthesis, playing important roles in pathogenicity and adaptation of fungi to a variety of environmental stresses. The FgRRG-1deletion mutant showed increased sensitivity to osmotic stress mediated by NaCl, KCl, sorbitol or glucose, and to metal cations Li+, Ca2+and Mg2+The mutant, however, was more resistant than the parent strain to dicarboximide and phenylpyrrole fungicides. Inoculation tests showed that the mutant exhibited decreased virulence on wheat heads, and produced a significantly low level of DON.
2) The type2C protein phosphatases FgPtc3is the negative regulator of the HOG signaling pathway in F. graminearum. FgPtc3is involved in the regulation of DON and aurofusarin biosynthesis, and also plays important roles in regulating many development processes and secondary metabolisms. The FgPTC3deletion mutant exhibited reduced aerial hyphae formation and deoxynivalenol (DON) production, but increased production of conidia. The mutant showed increased resistance to osmotic stress and cell wall-damaging agents, and accumulated a higher basal level of glycerol than the wild-type progenitor. Ultrastructural and histochemical analyses showed that conidia of the mutant contained an unusually high number of large lipid droplets. Pathogencity assays showed that the mutant is unable to infect flowering wheat head. Serial analysis of gene expression in the deletion mutant revealed that FgPtc3is associated with various metabolic pathways. Additionally, the FgPTC3partially rescued the growth defect of a yeast PTC1deletion mutant under various stress conditions, and also reduced the level of phosphorylated Hogl. The FgPtc3has the PP2C phosphatase activity, which may help in exploitation of drug targets based on screening of the protein activity inhibitors.
3) The tyrosine phosphatase FgPtp2is not associated with the HOG signaling pathway, but plays important roles in the regulation of DON biosynthesis and secondary metabolisms. The FgPTP2deletion mutant exhibited a significant decrease in mycelial growth and conidiation, but do not change the sensitivity to variety stresses. In addition, the deletion mutant was impaired dramatically in virulence and produced a low level of DON. Serial analysis of gene expression in the mutant revealed that FgPtp2is associated with various metabolic pathways, especially with the synthesis of ribosomal protein.
4) The FgVeA and FgVelB in the VeA regulatory system have some common functions in regulation of DON biosynthesis, and various cellular processes. The FgVEA and FgVELB deletion mutant exhibited a reduction in aerial hyphae formation, hydrophobicity, and deoxynivalenol (DON) biosynthesis, but increased the conidial production. Sensitivity tests to various stresses showed that the mutants exhibited increased resistance to osmotic stress and cell wall-damaging agents, but increased sensitivity to iprodione and fludioxonil fungicides, and the mutants contained an unusually high number of large lipid droplets, which is in agreement with the observation that the mutant accumulated a higher basal level of glycerol than the wild-type progenitor. Both FgVeA and FgVe1B play a critical role in pathogenicity,in a variety of biological functions and secondary metabolisms. The LaeA homologous genes are not essential for mycelial growth in F. graminearum, the deletion of the genes caused minor influence in pigment synthesis, and have no effect on DON production.
Results of the research indicated that the FgRrgl and FgPtc3in HOG signaling pathway, as well as FgVeA and FgVe1B in VeA regulatory system are involved in pathogenicity and DON biosynthesis, which may represent attractive antimicrobial targets for disease control purposes. The results aslo may help in exploitation of drug targets for the design of new anti-mycotoxin and antifungal agents.
引文
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