MAPK途径在盾壳霉产孢及重寄生作用中的功能研究
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摘要
核盘菌(Sclerotinia sclerotiorum (Lib.) de Bary)是一种-世界性分布的病原真菌,能导致多种重要作物的病害。盾壳霉(Coniothyrium minitans)是防治菌核病的一种重要生防菌。盾壳霉分生孢子的形成对其完成自身的生活史及重寄生作用非常重要。关于盾壳霉的产孢机理已有一些报道,但盾壳霉产孢是一个非常复杂的过程,分生孢子形成的调控网络目前尚不清晰。本论文主要以白盾壳霉T-DNA插入体库中筛选到的一株产孢缺陷型突变体ZS-1T1000为研究对象,通过TAIL-PCR等技术获得了T-DNA的侧翼序列,并以此展开研究了盾壳霉中与细胞壁完整性相关的MAPK途径(CWI-MAPK)在盾壳霉重寄生、产孢及其维系自身细胞壁稳定过程中的作用。同时也研究了与酵母中的另一MAPK途径,FUS3/KSS1MAPK途径中STE7的同源物CmSTE7在盾壳霉寄生核盘菌菌核中的作用,主要研究结果如下：
1)突变体ZS-1T1000丧失产孢能力系由T-DNA单拷贝插入破坏所致,T-DNA插入破坏了一个编码蛋白激酶激酶激酶的CmBCK1基因。CmBCK1基因的3’临近末端处含有一个内含子,基因编码区序列(CDS, coding sequence)为5361bp,编码大小为1786aa的蛋白质。T-DNA插入在该基因的5100bp位点,破坏了该蛋白质的蛋白激酶催化保守结构域.CmBCK1基因在盾壳霉基因组中为单拷贝基因,T-DNA的插入导致CmBCK1功能缺失,致使ZS-1T1000不能形成分生孢子器及分生孢子,寄生菌核能力显著下降,对细胞壁合成抑制剂及细胞壁降解酶敏感,菌落中央气生菌丝发生自溶,产色素能力显著下降。
2) CmBCK1基因敲除转化子丧失产孢能力,中央的气生菌丝出现自溶现象,产色素能力显著下降,对细胞壁合成抑制剂及细胞壁降解酶敏感,在寄生菌核能力方面存在缺陷,寄生能力显著下降,其表型与突变体ZS-1T1000类似。
3)根据序列同源性,自盾壳霉中克隆了酵母CWI-MAPK途径中BCK1下游Slt2的同源基因CmSlt2。CmSlt2基因含有5个内含子,CDS为1254bp,编码大小为417aa的蛋白质。与CmBCK1突变体类似,CmSlt2基因的敲除转化了也丧失了产孢能力,寄生核盘菌菌核能力显著下降,增加对细胞壁合成抑制剂及细胞壁降解酶的敏感性,影响色素的形成,菌落中央菌丝自溶。CmSlt2突变体的缺陷表型可被自身的CmSlt2基因互补所恢复。
4)结合CmBCK1及CmSlt2在盾壳霉中功能,表明CWI-MAPK途径参与调控盾壳霉的产孢、重寄生作用、色素形成及维持细胞壁稳定。
5)从盾壳霉中克隆了与酵母FUS3/KSS1MAPK途径中STE7同源的CmSTE7。 CmSTE7基因含有3个内含子,CDS为1362bp,编码大小为453aa的蛋白质。CmSTE7突变体菌落白色,不产生色素(后期在培养基基质中产生微量的色素),不能形成分生孢子器,气生菌丝增多,菌丝顶端分支增多,对核盘菌菌核的重寄生能力显著下降。结果表明CmSTE7参与调控盾壳霉的分生孢子器形成、重寄生作用及色素的产生。
综上所述,MAPK信号途径中的CWI-MAPK途径及FUS3/KSS1MAPK途径在盾壳霉产孢及重寄生过程中起着至关重要的作用,研究结果为阐明盾壳霉产孢及重寄生的作用机制提供了理论基础,丰富了丝状真菌的发育和致病相关的分子生物学内容,同时也为高效地规模化生产盾壳霉分生孢子提供了理论依据。
Sclerotinia sclerotiorum (Lib.) de Bary is a worldwide fungal pathogen causing serious diseases on many economically important crops. Coniothyrium minitans is an important biocontrol agent against Sclerotinia diseases. The conidiation of C. minitans is very important to its life cycle and mycoparasitism. Previous work had reported about the conidiation of C. minitans, however, the complex network that regulates the conidiation is still not clear. In this thesis, the conidiation deficient mutant ZS-1T1000, screened from T-DNA insertional mutagenesis library of ZS-1, was investigated. The flanking sequence of the T-DNA insertional site was amplified using TAIL-PCR, and based on the conidiation-related gene that blocked in the mutant ZS-1T1000, we investigated the functions of cell wall integrity MAPK pathway (CWI-MAPK) which participates in the mycoparasitism, conidiation and maintaining its cell wall integrity in C. minitans. Furthermore, we also analyzed the functions of CmSTE7, the homolog of STE7in the FUS3/KSS1MAPK pathway of Saccharomyces cerevisiae, in sclerotial mycoparasitism of C. minitans. Results achieved so far were summarized as following:
a) The conidiation deficient mutant ZS-1T1000was blocked by single copy of T-DNA insertion, and T-DNA inserted in the gene CmBCK1which encode a protein kinase kinase kinase. The CmBCK1contains one intron at the3'terminal end, and the coding sequence (CDS) of CmBCK1is5361bp which encodes a1786aa protein. The T-DNA insertional site exists at5100bp, and destroys the conservative catalytic domain. Only one copy of CmBCK1exists in the genome of C. minitans, and the lost function of CmBCKl leading to the lack of pycnidium and conidiation, significantly reduced the ability of sclerotial parasitization, hypersensitive to the cell wall inhibitor and cell wall lyases, the autolytic aerial mycelia and reduced pigmentation noticeably in the mutant ZS-1T1000.
b) Targeted disruption of the CmBCK1showed that the ΔCmBCK1mutant was uncapable of conidiation; autolytic mycelia in center of colony, decreased melanin production, hypersensitive to the cell wall inhibitor and cell wall lyases, and defective in the mycoparasitization, these deficiencies were similar to the mutant ZS-1T1000.
c) From C. minitans, the Slt2homolog CmSlt2which downstream of BCK1in the CWI-MAPK pathway was obtained. The CmSlt2contains5introns, and the CDS of CmSlt2is1254bp which encodes a417aa protein. Similar to the CmBCK1mutant, Cmslt2disruption mutants lost the ability to produce the conidia, significantly decreased in the sclerotial parasitization, increased in the susceptibility to the cell wall inhibitor and cell wall lyases and defective in the pigmentation. These defective phenotypes were restored by the complementation of CmSlt2gene.
d) Taken together, the results from CmBCK1and Cmslt2suggest that the CWI-MAPK pathway is essential for the conidiation, sclerotial mycoparasitism, pigmentation, and maintenance the stability of cell wall in C. minitans.
e) CmSTE7, the homolog of STE7from the FUS3/KSS1MAPK pathway of S. cerevisiae, was obtained from C. minitans. The CmSTE7contains3introns, and the CDS of CmSTE7is1362bp which encodes a453aa protein. CmSTE7disruption mutants showed a white colony, failed to produce melanin (slight melanin produced in the PDA medium after long time incubation), failed to form pycnidium, increased the number of aerial hyphae and multibranches, and extremely decreased the ability of sclerotial parasitization. These results suggest that CmSTE7involves in sclerotial mycoparasitism, the formation of pycnidium and melanin production in C. minitans.
The MAPK pathways involved in the regulation of the various development procedures in mycoparasite fungus C. minitans had been implicated in this thesis, the results suggested that CWI-MAPK and FUS3/KSS1MAPK pathway played critical roles in the conidiation and sclerotial mycoparasitism in C. minitans. These progeny could be a theoretical basis to elucidate the complex progress of the conidiation and sclerotial mycoparasitism in C. minitans, advance our knowledge of filamentous fungal biology and the molecular biology of pathogenicity, and also have important theoretical significance to the commercial usage of C. minitans as the biocontrol agent.
1)突变体ZS-1T1000丧失产孢能力系由T-DNA单拷贝插入破坏所致,T-DNA插入破坏了一个编码蛋白激酶激酶激酶的CmBCK1基因。CmBCK1基因的3’临近末端处含有一个内含子,基因编码区序列(CDS, coding sequence)为5361bp,编码大小为1786aa的蛋白质。T-DNA插入在该基因的5100bp位点,破坏了该蛋白质的蛋白激酶催化保守结构域.CmBCK1基因在盾壳霉基因组中为单拷贝基因,T-DNA的插入导致CmBCK1功能缺失,致使ZS-1T1000不能形成分生孢子器及分生孢子,寄生菌核能力显著下降,对细胞壁合成抑制剂及细胞壁降解酶敏感,菌落中央气生菌丝发生自溶,产色素能力显著下降。
2) CmBCK1基因敲除转化子丧失产孢能力,中央的气生菌丝出现自溶现象,产色素能力显著下降,对细胞壁合成抑制剂及细胞壁降解酶敏感,在寄生菌核能力方面存在缺陷,寄生能力显著下降,其表型与突变体ZS-1T1000类似。
3)根据序列同源性,自盾壳霉中克隆了酵母CWI-MAPK途径中BCK1下游Slt2的同源基因CmSlt2。CmSlt2基因含有5个内含子,CDS为1254bp,编码大小为417aa的蛋白质。与CmBCK1突变体类似,CmSlt2基因的敲除转化了也丧失了产孢能力,寄生核盘菌菌核能力显著下降,增加对细胞壁合成抑制剂及细胞壁降解酶的敏感性,影响色素的形成,菌落中央菌丝自溶。CmSlt2突变体的缺陷表型可被自身的CmSlt2基因互补所恢复。
4)结合CmBCK1及CmSlt2在盾壳霉中功能,表明CWI-MAPK途径参与调控盾壳霉的产孢、重寄生作用、色素形成及维持细胞壁稳定。
5)从盾壳霉中克隆了与酵母FUS3/KSS1MAPK途径中STE7同源的CmSTE7。 CmSTE7基因含有3个内含子,CDS为1362bp,编码大小为453aa的蛋白质。CmSTE7突变体菌落白色,不产生色素(后期在培养基基质中产生微量的色素),不能形成分生孢子器,气生菌丝增多,菌丝顶端分支增多,对核盘菌菌核的重寄生能力显著下降。结果表明CmSTE7参与调控盾壳霉的分生孢子器形成、重寄生作用及色素的产生。
综上所述,MAPK信号途径中的CWI-MAPK途径及FUS3/KSS1MAPK途径在盾壳霉产孢及重寄生过程中起着至关重要的作用,研究结果为阐明盾壳霉产孢及重寄生的作用机制提供了理论基础,丰富了丝状真菌的发育和致病相关的分子生物学内容,同时也为高效地规模化生产盾壳霉分生孢子提供了理论依据。
Sclerotinia sclerotiorum (Lib.) de Bary is a worldwide fungal pathogen causing serious diseases on many economically important crops. Coniothyrium minitans is an important biocontrol agent against Sclerotinia diseases. The conidiation of C. minitans is very important to its life cycle and mycoparasitism. Previous work had reported about the conidiation of C. minitans, however, the complex network that regulates the conidiation is still not clear. In this thesis, the conidiation deficient mutant ZS-1T1000, screened from T-DNA insertional mutagenesis library of ZS-1, was investigated. The flanking sequence of the T-DNA insertional site was amplified using TAIL-PCR, and based on the conidiation-related gene that blocked in the mutant ZS-1T1000, we investigated the functions of cell wall integrity MAPK pathway (CWI-MAPK) which participates in the mycoparasitism, conidiation and maintaining its cell wall integrity in C. minitans. Furthermore, we also analyzed the functions of CmSTE7, the homolog of STE7in the FUS3/KSS1MAPK pathway of Saccharomyces cerevisiae, in sclerotial mycoparasitism of C. minitans. Results achieved so far were summarized as following:
a) The conidiation deficient mutant ZS-1T1000was blocked by single copy of T-DNA insertion, and T-DNA inserted in the gene CmBCK1which encode a protein kinase kinase kinase. The CmBCK1contains one intron at the3'terminal end, and the coding sequence (CDS) of CmBCK1is5361bp which encodes a1786aa protein. The T-DNA insertional site exists at5100bp, and destroys the conservative catalytic domain. Only one copy of CmBCK1exists in the genome of C. minitans, and the lost function of CmBCKl leading to the lack of pycnidium and conidiation, significantly reduced the ability of sclerotial parasitization, hypersensitive to the cell wall inhibitor and cell wall lyases, the autolytic aerial mycelia and reduced pigmentation noticeably in the mutant ZS-1T1000.
b) Targeted disruption of the CmBCK1showed that the ΔCmBCK1mutant was uncapable of conidiation; autolytic mycelia in center of colony, decreased melanin production, hypersensitive to the cell wall inhibitor and cell wall lyases, and defective in the mycoparasitization, these deficiencies were similar to the mutant ZS-1T1000.
c) From C. minitans, the Slt2homolog CmSlt2which downstream of BCK1in the CWI-MAPK pathway was obtained. The CmSlt2contains5introns, and the CDS of CmSlt2is1254bp which encodes a417aa protein. Similar to the CmBCK1mutant, Cmslt2disruption mutants lost the ability to produce the conidia, significantly decreased in the sclerotial parasitization, increased in the susceptibility to the cell wall inhibitor and cell wall lyases and defective in the pigmentation. These defective phenotypes were restored by the complementation of CmSlt2gene.
d) Taken together, the results from CmBCK1and Cmslt2suggest that the CWI-MAPK pathway is essential for the conidiation, sclerotial mycoparasitism, pigmentation, and maintenance the stability of cell wall in C. minitans.
e) CmSTE7, the homolog of STE7from the FUS3/KSS1MAPK pathway of S. cerevisiae, was obtained from C. minitans. The CmSTE7contains3introns, and the CDS of CmSTE7is1362bp which encodes a453aa protein. CmSTE7disruption mutants showed a white colony, failed to produce melanin (slight melanin produced in the PDA medium after long time incubation), failed to form pycnidium, increased the number of aerial hyphae and multibranches, and extremely decreased the ability of sclerotial parasitization. These results suggest that CmSTE7involves in sclerotial mycoparasitism, the formation of pycnidium and melanin production in C. minitans.
The MAPK pathways involved in the regulation of the various development procedures in mycoparasite fungus C. minitans had been implicated in this thesis, the results suggested that CWI-MAPK and FUS3/KSS1MAPK pathway played critical roles in the conidiation and sclerotial mycoparasitism in C. minitans. These progeny could be a theoretical basis to elucidate the complex progress of the conidiation and sclerotial mycoparasitism in C. minitans, advance our knowledge of filamentous fungal biology and the molecular biology of pathogenicity, and also have important theoretical significance to the commercial usage of C. minitans as the biocontrol agent.
引文
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