核盘菌菌核形成相关蛋白Ss-Sl2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植物病原真菌,能够侵染许多重要的农作物,引起多种作物如油菜、向日葵和大豆等的菌核病,给农业生产造成严重的损失。菌核是核盘菌越冬、越夏和繁殖的结构,也是菌核病的重要初侵染来源,抑制菌核的形成将切断菌核病的病害循环,因此研究核盘菌菌核形成的分子机理将为菌核病的防治提供新的思路和线索。
缺乏简便稳定的转化体系是核盘菌基因功能研究的重大障碍。本研究建立了以核盘菌菌丝为受体的农杆菌介导的遗传转化体系,即直接将含有载体pTFSS的农杆菌与新鲜的核盘菌菌丝块在乙酰丁香酮存在的条件下进行共培养,2d后将菌丝块丢弃并将共培养物置于含30μg/ml潮霉素的PDA培养基中进行选择性培养,3d后获得转化子。该方法操作简便,且转化效率较稳定,平均每个培养皿能获得1-3个转化子,为深入研究核盘菌菌核形成相关基因的功能建立了良好的平台。本研究还发现潮霉素磷酸转移酶基因在核盘菌转化子的无性阶段能稳定遗传,但在有性后代发生丢失。
为深入探索核盘菌菌核形成分子机理,本实验室前期利用Illumina-Solexa技术建立了核盘菌菌丝生长和菌核形成阶段的基因表达谱。分析该表达谱发现,Ss-Sl2(SS1G_05917, XM_001592945)的表达量在菌核形成阶段升高了400倍,推测其可能与核盘菌菌核形成密切相关。本研究表明Ss-S12蛋白全长352aa,具有一个信号肽结构和两个PAN模块,其同源蛋白仅仅出现在部分子囊菌中且均为功能未知蛋白。在核盘菌菌丝生长阶段,少量Ss-S12蛋白分布于细胞壁和隔膜上；在菌核形成阶段,Ss-Sl2表达量迅速升高,Ss-S12蛋白大量分布于细胞外加厚的细胞壁上。Ss-S12的表达水平受cAMP水平的抑制,而与环境pH无关。Ss-S12基因沉默转化子维持细胞完整性的能力下降,但是菌丝生长和致病力与野生型菌株没有差异。在菌核形成初期,Ss-Sl2沉默转化子可以形成白色绒毛状菌丝团,但是菌丝团不能固化和黑化并发育成熟的菌核,在菌丝团内部菌丝排列疏松且细胞壁加厚不明显,表明Ss-S12在核盘菌菌核形成过程中起着重要作用。
对Ss-S12互作蛋白进行鉴定和功能研究将有助于理解其控制菌核形成和维持细胞完整性的作用机理。本研究表明3-磷酸甘油醛脱氢酶(GAPDH, SS1G_07798)和沃鲁宁体主要组成蛋白(Hex1, SS1G_03527)可以与Ss-S12发生较强的直接相互作用,而转录延伸因子(EF-1α)则与Ss-S12存在着较弱的直接相互作用。GAPDH基因沉默转化子只能形成白色绒毛状菌丝团,而不能形成黑色的菌核,与Ss-Sl2沉默转化子菌核形成表型相似；Hex1基因沉默转化子保持细胞完整性能力下降,与Ss-Sl2沉默转化子菌丝表型相似,表明Ss-S12可能通过与GAPDH和Hexl的互作来控制核盘菌菌核的形成和维持细胞的完整性。
本研究深入解析了核盘菌Ss-S12蛋白的作用及其机理,为揭示菌核形成的分子机制提供了理论依据,也为菌核病的防治提供了重要线索；此外,本研究也为探索其它真菌中Ss-S12同源蛋白的功能提供了重要参考。
Sclerotinia sclerotiorum (Lib.) de Bary is a notorious fungal pathogen with worldwide distribution. This fungus may infect many important crops, such as oilseed rape, sunflower, soybean, and cause Sclerotinia diseases. Sclerotia are not only the important tissue for breeding and over summer and winter to S. sclerotiorum, but also the important source of primarily infection for Sclerotinia diseases. Repress the formation of sclerotia may interrupt the disease cycle induced by S. sclerotiorum and research on the molecular mechanism involved sclerotial development may provide some clues to control the Sclerotinia diseases.
Lack of efficient and easy transformation system is a great obstacle for studying the gene function in S. sclerotiorum. In this study, an Agrobacterium-medated transformation system was established for S. sclerotiorum by using the hyphae as receptor. A. tumefaciens cells containing pTFSS and fresh S. sclerotiorum mycelial plugs were co-cultured with acetovanillone for2days. Then the mycelial plugs were removed and the A. tumefaciens cells and S. sclerotiorum hypha were transferred into PDA medium containing30μg/ml of hygromycin B. The transformants emerged on the selective plates after3days. This method was simple and had a stable efficiency of transform. On average,1-3transformants can be obtained in each plate. This system provided an effective platform for gene functional study of involved in S. sclerotiorum. This research also showed that the HPH gene in transformants inherited steadily in the asexual stage of S. sclerotiorum, but it lost in the sexual progenies.
To study the molecular mechanism involved in sclerotial development, a gene expression pattern during the stage of hyphal growth and sclerotial development was contrasted with Illumina-Solexa in our laboratory. According to the gene expression pattern, a gene named Ss-Sl2(SSIG_05917, XM_001592945) had higher expression level during sclerotial development, with the relative expression during sclerotial development400-fold greater than that at the hyphal growth stage. Thus, Ss-Sl2is supposedly to have a close relation with sclerotial development. Our results showed that Ss-Sl2protein has352aa residues and contains a signal peptide and two PAN modules. Its homologies appeared only in the some ascomycotina fungi and all of those are of unknown function. At the stage of vegetative hyphal growth, only a few gold particles could be observed dispersed in the cell walls and septa of hyphae. During the stage of sclerotial development, the transcript level of Ss-Sl2rised rapidly and a large number of Ss-S12protein were secreted and located in the thicken cell wall of sclerotia. The expression level of Ss-Sl2is down-regulated by cAMP level but has no relation with the medium pH. The ability to maintain the cellular integrity of RNAi-mediated Ss-Sl2silenced strains was reduced, but the hyphal growth and virulence of Ss-Sl2silenced strains were not significantly different from the wild type strain. Ss-Sl2silenced strains could form interwoven hyphal masses at the initial stage of sclerotial development, but the interwoven hyphae could not consolidate and melanized to become mature sclerotia. Hyphae in these interwoven bodies were thin-walled, arranged loosely. These evidence showed that Ss-S12play an important role in sclerotial development of S. sclerotiorum.
Identification and functional study on proteins which interacted with Ss-S12may provide clues to understand its mechanism involved in controlling sclerotial development and cell integrity. Our results showed that glyceraldehyde-3-phosphate dehydrogenase (GAPDH), Woronin body major protein (Hexl) and elongation factor1alpha (EF-la) could interact with the Ss-S12directly, and EF-1a was found to have only a weak interaction with Ss-S12. GAPDH-knockdown strains did not produce typical sclerotia, but only forming interwoven hyphal masses. The phenotype was similar to that of Ss-Sl2silenced transformants. Hexl-knockdown strains showed similar impairment in maintenance of hyphal integrity as Ss-Sl2silenced strains. These evidences indicated that the Ss-S12may play the role in sclerotial development and the mantainence of cell integrity through its interaction with GAPDH and Hexl.
The research profoundly analysed the function and mechanism of Ss-S12protein in S. sclerotiorum, which not only provides the theory basis for revealing the molecular mechanism of sclerotial development, but also offers important clues to control Sclerotnia disease. Besides, this research also provides important references for studying the function of the homologies of Ss-S12in some other fungi.
缺乏简便稳定的转化体系是核盘菌基因功能研究的重大障碍。本研究建立了以核盘菌菌丝为受体的农杆菌介导的遗传转化体系,即直接将含有载体pTFSS的农杆菌与新鲜的核盘菌菌丝块在乙酰丁香酮存在的条件下进行共培养,2d后将菌丝块丢弃并将共培养物置于含30μg/ml潮霉素的PDA培养基中进行选择性培养,3d后获得转化子。该方法操作简便,且转化效率较稳定,平均每个培养皿能获得1-3个转化子,为深入研究核盘菌菌核形成相关基因的功能建立了良好的平台。本研究还发现潮霉素磷酸转移酶基因在核盘菌转化子的无性阶段能稳定遗传,但在有性后代发生丢失。
为深入探索核盘菌菌核形成分子机理,本实验室前期利用Illumina-Solexa技术建立了核盘菌菌丝生长和菌核形成阶段的基因表达谱。分析该表达谱发现,Ss-Sl2(SS1G_05917, XM_001592945)的表达量在菌核形成阶段升高了400倍,推测其可能与核盘菌菌核形成密切相关。本研究表明Ss-S12蛋白全长352aa,具有一个信号肽结构和两个PAN模块,其同源蛋白仅仅出现在部分子囊菌中且均为功能未知蛋白。在核盘菌菌丝生长阶段,少量Ss-S12蛋白分布于细胞壁和隔膜上；在菌核形成阶段,Ss-Sl2表达量迅速升高,Ss-S12蛋白大量分布于细胞外加厚的细胞壁上。Ss-S12的表达水平受cAMP水平的抑制,而与环境pH无关。Ss-S12基因沉默转化子维持细胞完整性的能力下降,但是菌丝生长和致病力与野生型菌株没有差异。在菌核形成初期,Ss-Sl2沉默转化子可以形成白色绒毛状菌丝团,但是菌丝团不能固化和黑化并发育成熟的菌核,在菌丝团内部菌丝排列疏松且细胞壁加厚不明显,表明Ss-S12在核盘菌菌核形成过程中起着重要作用。
对Ss-S12互作蛋白进行鉴定和功能研究将有助于理解其控制菌核形成和维持细胞完整性的作用机理。本研究表明3-磷酸甘油醛脱氢酶(GAPDH, SS1G_07798)和沃鲁宁体主要组成蛋白(Hex1, SS1G_03527)可以与Ss-S12发生较强的直接相互作用,而转录延伸因子(EF-1α)则与Ss-S12存在着较弱的直接相互作用。GAPDH基因沉默转化子只能形成白色绒毛状菌丝团,而不能形成黑色的菌核,与Ss-Sl2沉默转化子菌核形成表型相似；Hex1基因沉默转化子保持细胞完整性能力下降,与Ss-Sl2沉默转化子菌丝表型相似,表明Ss-S12可能通过与GAPDH和Hexl的互作来控制核盘菌菌核的形成和维持细胞的完整性。
本研究深入解析了核盘菌Ss-S12蛋白的作用及其机理,为揭示菌核形成的分子机制提供了理论依据,也为菌核病的防治提供了重要线索；此外,本研究也为探索其它真菌中Ss-S12同源蛋白的功能提供了重要参考。
Sclerotinia sclerotiorum (Lib.) de Bary is a notorious fungal pathogen with worldwide distribution. This fungus may infect many important crops, such as oilseed rape, sunflower, soybean, and cause Sclerotinia diseases. Sclerotia are not only the important tissue for breeding and over summer and winter to S. sclerotiorum, but also the important source of primarily infection for Sclerotinia diseases. Repress the formation of sclerotia may interrupt the disease cycle induced by S. sclerotiorum and research on the molecular mechanism involved sclerotial development may provide some clues to control the Sclerotinia diseases.
Lack of efficient and easy transformation system is a great obstacle for studying the gene function in S. sclerotiorum. In this study, an Agrobacterium-medated transformation system was established for S. sclerotiorum by using the hyphae as receptor. A. tumefaciens cells containing pTFSS and fresh S. sclerotiorum mycelial plugs were co-cultured with acetovanillone for2days. Then the mycelial plugs were removed and the A. tumefaciens cells and S. sclerotiorum hypha were transferred into PDA medium containing30μg/ml of hygromycin B. The transformants emerged on the selective plates after3days. This method was simple and had a stable efficiency of transform. On average,1-3transformants can be obtained in each plate. This system provided an effective platform for gene functional study of involved in S. sclerotiorum. This research also showed that the HPH gene in transformants inherited steadily in the asexual stage of S. sclerotiorum, but it lost in the sexual progenies.
To study the molecular mechanism involved in sclerotial development, a gene expression pattern during the stage of hyphal growth and sclerotial development was contrasted with Illumina-Solexa in our laboratory. According to the gene expression pattern, a gene named Ss-Sl2(SSIG_05917, XM_001592945) had higher expression level during sclerotial development, with the relative expression during sclerotial development400-fold greater than that at the hyphal growth stage. Thus, Ss-Sl2is supposedly to have a close relation with sclerotial development. Our results showed that Ss-Sl2protein has352aa residues and contains a signal peptide and two PAN modules. Its homologies appeared only in the some ascomycotina fungi and all of those are of unknown function. At the stage of vegetative hyphal growth, only a few gold particles could be observed dispersed in the cell walls and septa of hyphae. During the stage of sclerotial development, the transcript level of Ss-Sl2rised rapidly and a large number of Ss-S12protein were secreted and located in the thicken cell wall of sclerotia. The expression level of Ss-Sl2is down-regulated by cAMP level but has no relation with the medium pH. The ability to maintain the cellular integrity of RNAi-mediated Ss-Sl2silenced strains was reduced, but the hyphal growth and virulence of Ss-Sl2silenced strains were not significantly different from the wild type strain. Ss-Sl2silenced strains could form interwoven hyphal masses at the initial stage of sclerotial development, but the interwoven hyphae could not consolidate and melanized to become mature sclerotia. Hyphae in these interwoven bodies were thin-walled, arranged loosely. These evidence showed that Ss-S12play an important role in sclerotial development of S. sclerotiorum.
Identification and functional study on proteins which interacted with Ss-S12may provide clues to understand its mechanism involved in controlling sclerotial development and cell integrity. Our results showed that glyceraldehyde-3-phosphate dehydrogenase (GAPDH), Woronin body major protein (Hexl) and elongation factor1alpha (EF-la) could interact with the Ss-S12directly, and EF-1a was found to have only a weak interaction with Ss-S12. GAPDH-knockdown strains did not produce typical sclerotia, but only forming interwoven hyphal masses. The phenotype was similar to that of Ss-Sl2silenced transformants. Hexl-knockdown strains showed similar impairment in maintenance of hyphal integrity as Ss-Sl2silenced strains. These evidences indicated that the Ss-S12may play the role in sclerotial development and the mantainence of cell integrity through its interaction with GAPDH and Hexl.
The research profoundly analysed the function and mechanism of Ss-S12protein in S. sclerotiorum, which not only provides the theory basis for revealing the molecular mechanism of sclerotial development, but also offers important clues to control Sclerotnia disease. Besides, this research also provides important references for studying the function of the homologies of Ss-S12in some other fungi.
引文
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