大豆疫霉G蛋白和钙信号途径的功能分析
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摘要
大豆疫霉(Phytophthora sojae)侵染大豆引起的疫霉根腐病是大豆生产上的主要病害之一。大豆疫霉在田间的传播主要通过形成孢子囊,释放游动孢子,在水膜存在的条件下,通过识别植株分泌的异黄酮等化学信号分子,定向游动,寻找寄主植物,最终成功侵染感病植物。所以了解大豆疫霉菌无性发育过程中孢子囊和游动孢子的形成和游动孢子的趋化性机制,可以为寻找新的杀菌剂作用靶标,防止病害的发生和扩展提供理论依据。
大豆疫霉G蛋白效应因子和钙离子结合蛋白预测与分析。大豆疫霉孢子囊的形成和游动孢子的行为受到各种外界条件影响和胞内信号传递的控制。卵菌中陆续报道了G蛋白和钙信号途径的基因在孢子囊形成和游动孢子行为方面的作用,为大豆疫霉孢子囊的形成和游动孢子趋化性机制的研究提供了有利的借鉴作用。本文对大豆疫霉的基因组数据库进行了搜索,寻找到包括腺苷酸环化酶、G蛋白信号调解因子和磷脂酶D在内的G蛋白效应因子以及包括钙调素和钙离子调控的蛋白激酶在内的钙离子结合蛋白,并分析了其结构特点和基因组分布。发现了以腺苷酸环化酶为代表的G蛋白效应分子的特征。候选基因的EST分析结果表明,G蛋白和钙信号途径在大豆疫霉的侵染过程中可能起着更加重要的作用。
大豆疫霉G蛋白和钙信号途径的转录分析。为了研究G蛋白和钙信号途径在大豆疫霉发育和致病过程中的作用,并对前一章所进行的推测进行验证,本文挑选了20个可能与G蛋白和钙信号途径相关的基因,利用半定量RT-PCR对候选基因进行了营养胁迫、氧化压力、孢子囊产生和游动孢子等不同生长阶段以及侵染早期的转录分析,以揭示G蛋白和钙信号途径的表达规律。结果表明,包括G蛋白a和B亚基在内的10个基因在大豆疫霉孢子囊形成阶段的转录水平较营养生长阶段明显升高；13个基因在游动孢子中的转录水平较营养菌丝和孢子囊中有明显降低,其中10个基因编码的蛋白参与钙信号途径；在大豆疫霉侵染大豆早期,编码G蛋白a和B亚基的基因PsGPA1和PsGPBl表现出明显的上调表达。G蛋白a亚基是所有候选基因中唯一既在游动孢子中诱导表达,又在侵染阶段诱导表达的基因。深入分析PsGPA1和PsGPB1在大豆疫霉无性发育阶段的表达情况发现,在大豆疫霉的游动孢子游动阶段,G蛋白a亚基有可能抑制了胞内钙信号的激活。
G蛋白a亚基对于大豆疫霉游动孢子趋化性的作用研究。大豆疫霉的游动孢子对异黄酮类物质的趋化性是其识别和侵染寄主的前提。然而,这种趋化性的分子机制目前尚不清楚。G蛋白a亚基既能够在大豆疫霉的游动孢子中诱导表达,又能够在侵染早期诱导表达。为了研究其是否与游动孢子对异黄酮物质的趋化性机制相关,本研究利用基因沉默技术,获得了G蛋白a亚基的基因沉默突变体。突变体的表型分析结果表明,Ga的沉默不会影响茵丝的营养生长和孢子囊的产生,但却严重影响了游动孢子对不同浓度异黄酮类物质的趋化性。游动孢子趋化性的丧失和休止速度的加快降低了突变体在大豆根部的附着能力；休止胞萌发能力也随之改变,导致突变体游动孢子即使与大豆根部接触也很难成功侵染。此外,Ga沉默突变体游动孢子中钙调素基因表达量明显升高,说明Ga亚基有可能是钙信号途径的负调控因子。
Ga结合蛋白的筛选与受Ga调控的信号途径分析。大豆疫霉的G蛋白a亚基不仅影响着孢子囊的割裂和游动孢子的释放,而且控制着游动孢子的游动能力和休止胞的萌发能力。Ga亚基是通过调节哪些基因的表达、影响哪些信号通路、怎样传递信号来控制这些性状,尚未研究清楚。本文构建了大豆疫霉的cDNA文库,并以大豆疫霉Ga亚基为诱饵蛋白,利用酵母双杂交的方法,筛选到包括磷酸核酮糖差向异构酶和逆转录酶等7个可能与Ga相互作用的蛋白,说明G蛋白信号途径参与了大豆疫霉的糖代谢。同时,本文通过cDNA Macroarray的方法,比较分析了大豆疫霉野生型和Ga突变体游动孢子中的基因表达变化,得到了9个在Ga突变体游动孢子中转录水平发生变化的基因。这9个基因中有3个G蛋白信号调控因子(RGS)、一个Ca2+ATP酶和一个依赖于Ca2+/钙调素的蛋白激酶的转录水平发生下降；钙调素、两个Ca2+/钙调素的蛋白激酶和一个转录因子类似物的转录水平明显升高。这说明RGS和Ga在信号传递过程中可能是两个相互依赖、相互影响的因子。而Ga对钙信号途径并非单纯地起抑制作用。
Soybean root rot disease caused by oomycete pathogen Phytophthora sojae is one of main diseases which caused annual loss in agricultural production. In the field, sporangia formed on the tips of flooded mycelia, and cleavage to release mobile zoospores, which swim toward new host, through chemotaxis to the isoflavones secreted from soybean root. Consequently, to study the mechanism of sporangia formation and zoospore chemotaxis is crucial to disease control.
Bioinformatics analysis of G protein and calcium signal transduction pathways in Phytophthora sojae. G protein and calcium are the most ubiquitous cellular components in eukaryotic cells that execute many important roles in signaling. Genes encoding G protein binding effectors and calcium binding proteins required for signal transduction and metabolism have been characterized in many organisms, but only a few have been described for oomycetes. In this study, the genome sequences of Phytophthora sojae were explored to construct a genome wide inventory of genes involved in G protein and calcium signaling pathways, including those encoding adenylyl cyclase (AC), regulator of G protein signal (RGS), phospholipase D (PLD), calmodulin (CAM) and calcium dependent protein kinase (CMK). The most obvious missing link is a gene encoding G protein y subunit. In all eukaryotic genomes sequenced to date, Gy genes are annotated based on certain conserved features; however, these genes seem to be absent in Phytophthora spp. Analysis of the structural and regulatory domains and domain organization of the predicted isoforms of G protein, AC, RGS, PLD, CAM and CMK revealed many novel features compared with characterized representatives in other eukaryotes. Examples are transmembrane AC proteins with a N-terminal signal peptide. Compared with other sequenced eukaryotes, the genus Phytophthora clearly has several exceptional features in its ATP catalyze enzyme.
Transcriptional analysis of G protein and calcium signal pathways in Phytophthora sojae. Characterization of genes involved in development, stress response and pathogenesis is an important step in identifying methods to study Phytophthora sojae. In this study, the relative expression levels of 20 genes involved in G protein and calcium signals were studied using semi-quantitive RT-PCR. From the results of transcriptional analysis,10 genes including G protein a and B subunits were expressed upregulated in sporulating hyphae than that in nutrient hyphae.13 genes including 10 involved in calcium signal were expressed downregulated in zoospore than that in nutrient hyphae or sporulating hyphae. A Ga, a Gβ, a phospholipase, a calcium binding protein, a calcium binding protein kinase and a putative transcriptional factor were expressed upregulated in early infection stage of P. sojae. A Gβ,3 calcium binding protein kinase and 2 phospholipase D were differentially upregulated in early infection stage of P. sojae. Our results indicates that G protein and calcium signals are not only involved in P. sojae sporulating, but also important for early infection. G protein may temporarily suppress calcium signal activation in P. sojae zoospore.
Characterization of G protein alpha subunit. For the soybean pathogen Phytophthora sojae chemotaxis of zoospores to isoflavones is believed to be critical for recognition of the host and for initiating infection. However, the molecular mechanisms underlying this chemotaxis are unknown. To investigate the molecular mechanism of chemotaxis to isoflavone, expression of PsGPAl was silenced by introducing antisense constructs into P. sojae. PsGPAl silencing did not disturb hyphal growth or sporulation but severely affected zoospore behavior including chemotaxis to the soybean isoflavone daidzein. Zoospore encystment and cyst germinating were also alternated, causing that the PsGPAl silenced mutants were disabled in infecting soybean. Upregulation of a calmodulin gene in zoospores of PsGPAl-silenced mutants pointed to suppressor activity of Ga targeted at steps in the calcium signaling pathway, indicating that Ga subunit may be a negative regulator of calcium signal pathways.
Screening for Ga binding proteins and analysis of Ga regulated signal transduction pathways. G protein a subunit of Phytophthora sojae influences not only sporangia cleavage and zoospore release, but also zoospore motility and cyst germination. The mechanisms that how Ga binds with effectors and regulates downstream signal pathways are unknown. In this study, we constructed a P. sojae cDNA library and used yeast two hybrid method to screen for Ga binding proteins, and got 7 putative Ga binding proteins including ribulose-phosphate 3-epimerase and putative reverse transcriptase. Meanwhile, we compared gene expression profiles of P. sojae zoospores between wild type P6497 and Ga mutants A2 and A27 by cDNA Macroarray. From the results of comparison,6 genes including 3 regulator of G protein (RGS),1 Ca2+/calmodulin protein kinase and 1 Ca2+ ATPase were expressed down-regulated by Ga silencing, however,1 calmodulin,2 Ca2+/calmodulin protein kinase and 1 homolog of transcriptional factor were expressed up-regulated by Ga silencing. These results indicate that RGS activity may be dependent on the activity of Ga in P. sojae. And the relationship between G protein and calcium signal is very complex.
大豆疫霉G蛋白效应因子和钙离子结合蛋白预测与分析。大豆疫霉孢子囊的形成和游动孢子的行为受到各种外界条件影响和胞内信号传递的控制。卵菌中陆续报道了G蛋白和钙信号途径的基因在孢子囊形成和游动孢子行为方面的作用,为大豆疫霉孢子囊的形成和游动孢子趋化性机制的研究提供了有利的借鉴作用。本文对大豆疫霉的基因组数据库进行了搜索,寻找到包括腺苷酸环化酶、G蛋白信号调解因子和磷脂酶D在内的G蛋白效应因子以及包括钙调素和钙离子调控的蛋白激酶在内的钙离子结合蛋白,并分析了其结构特点和基因组分布。发现了以腺苷酸环化酶为代表的G蛋白效应分子的特征。候选基因的EST分析结果表明,G蛋白和钙信号途径在大豆疫霉的侵染过程中可能起着更加重要的作用。
大豆疫霉G蛋白和钙信号途径的转录分析。为了研究G蛋白和钙信号途径在大豆疫霉发育和致病过程中的作用,并对前一章所进行的推测进行验证,本文挑选了20个可能与G蛋白和钙信号途径相关的基因,利用半定量RT-PCR对候选基因进行了营养胁迫、氧化压力、孢子囊产生和游动孢子等不同生长阶段以及侵染早期的转录分析,以揭示G蛋白和钙信号途径的表达规律。结果表明,包括G蛋白a和B亚基在内的10个基因在大豆疫霉孢子囊形成阶段的转录水平较营养生长阶段明显升高；13个基因在游动孢子中的转录水平较营养菌丝和孢子囊中有明显降低,其中10个基因编码的蛋白参与钙信号途径；在大豆疫霉侵染大豆早期,编码G蛋白a和B亚基的基因PsGPA1和PsGPBl表现出明显的上调表达。G蛋白a亚基是所有候选基因中唯一既在游动孢子中诱导表达,又在侵染阶段诱导表达的基因。深入分析PsGPA1和PsGPB1在大豆疫霉无性发育阶段的表达情况发现,在大豆疫霉的游动孢子游动阶段,G蛋白a亚基有可能抑制了胞内钙信号的激活。
G蛋白a亚基对于大豆疫霉游动孢子趋化性的作用研究。大豆疫霉的游动孢子对异黄酮类物质的趋化性是其识别和侵染寄主的前提。然而,这种趋化性的分子机制目前尚不清楚。G蛋白a亚基既能够在大豆疫霉的游动孢子中诱导表达,又能够在侵染早期诱导表达。为了研究其是否与游动孢子对异黄酮物质的趋化性机制相关,本研究利用基因沉默技术,获得了G蛋白a亚基的基因沉默突变体。突变体的表型分析结果表明,Ga的沉默不会影响茵丝的营养生长和孢子囊的产生,但却严重影响了游动孢子对不同浓度异黄酮类物质的趋化性。游动孢子趋化性的丧失和休止速度的加快降低了突变体在大豆根部的附着能力；休止胞萌发能力也随之改变,导致突变体游动孢子即使与大豆根部接触也很难成功侵染。此外,Ga沉默突变体游动孢子中钙调素基因表达量明显升高,说明Ga亚基有可能是钙信号途径的负调控因子。
Ga结合蛋白的筛选与受Ga调控的信号途径分析。大豆疫霉的G蛋白a亚基不仅影响着孢子囊的割裂和游动孢子的释放,而且控制着游动孢子的游动能力和休止胞的萌发能力。Ga亚基是通过调节哪些基因的表达、影响哪些信号通路、怎样传递信号来控制这些性状,尚未研究清楚。本文构建了大豆疫霉的cDNA文库,并以大豆疫霉Ga亚基为诱饵蛋白,利用酵母双杂交的方法,筛选到包括磷酸核酮糖差向异构酶和逆转录酶等7个可能与Ga相互作用的蛋白,说明G蛋白信号途径参与了大豆疫霉的糖代谢。同时,本文通过cDNA Macroarray的方法,比较分析了大豆疫霉野生型和Ga突变体游动孢子中的基因表达变化,得到了9个在Ga突变体游动孢子中转录水平发生变化的基因。这9个基因中有3个G蛋白信号调控因子(RGS)、一个Ca2+ATP酶和一个依赖于Ca2+/钙调素的蛋白激酶的转录水平发生下降；钙调素、两个Ca2+/钙调素的蛋白激酶和一个转录因子类似物的转录水平明显升高。这说明RGS和Ga在信号传递过程中可能是两个相互依赖、相互影响的因子。而Ga对钙信号途径并非单纯地起抑制作用。
Soybean root rot disease caused by oomycete pathogen Phytophthora sojae is one of main diseases which caused annual loss in agricultural production. In the field, sporangia formed on the tips of flooded mycelia, and cleavage to release mobile zoospores, which swim toward new host, through chemotaxis to the isoflavones secreted from soybean root. Consequently, to study the mechanism of sporangia formation and zoospore chemotaxis is crucial to disease control.
Bioinformatics analysis of G protein and calcium signal transduction pathways in Phytophthora sojae. G protein and calcium are the most ubiquitous cellular components in eukaryotic cells that execute many important roles in signaling. Genes encoding G protein binding effectors and calcium binding proteins required for signal transduction and metabolism have been characterized in many organisms, but only a few have been described for oomycetes. In this study, the genome sequences of Phytophthora sojae were explored to construct a genome wide inventory of genes involved in G protein and calcium signaling pathways, including those encoding adenylyl cyclase (AC), regulator of G protein signal (RGS), phospholipase D (PLD), calmodulin (CAM) and calcium dependent protein kinase (CMK). The most obvious missing link is a gene encoding G protein y subunit. In all eukaryotic genomes sequenced to date, Gy genes are annotated based on certain conserved features; however, these genes seem to be absent in Phytophthora spp. Analysis of the structural and regulatory domains and domain organization of the predicted isoforms of G protein, AC, RGS, PLD, CAM and CMK revealed many novel features compared with characterized representatives in other eukaryotes. Examples are transmembrane AC proteins with a N-terminal signal peptide. Compared with other sequenced eukaryotes, the genus Phytophthora clearly has several exceptional features in its ATP catalyze enzyme.
Transcriptional analysis of G protein and calcium signal pathways in Phytophthora sojae. Characterization of genes involved in development, stress response and pathogenesis is an important step in identifying methods to study Phytophthora sojae. In this study, the relative expression levels of 20 genes involved in G protein and calcium signals were studied using semi-quantitive RT-PCR. From the results of transcriptional analysis,10 genes including G protein a and B subunits were expressed upregulated in sporulating hyphae than that in nutrient hyphae.13 genes including 10 involved in calcium signal were expressed downregulated in zoospore than that in nutrient hyphae or sporulating hyphae. A Ga, a Gβ, a phospholipase, a calcium binding protein, a calcium binding protein kinase and a putative transcriptional factor were expressed upregulated in early infection stage of P. sojae. A Gβ,3 calcium binding protein kinase and 2 phospholipase D were differentially upregulated in early infection stage of P. sojae. Our results indicates that G protein and calcium signals are not only involved in P. sojae sporulating, but also important for early infection. G protein may temporarily suppress calcium signal activation in P. sojae zoospore.
Characterization of G protein alpha subunit. For the soybean pathogen Phytophthora sojae chemotaxis of zoospores to isoflavones is believed to be critical for recognition of the host and for initiating infection. However, the molecular mechanisms underlying this chemotaxis are unknown. To investigate the molecular mechanism of chemotaxis to isoflavone, expression of PsGPAl was silenced by introducing antisense constructs into P. sojae. PsGPAl silencing did not disturb hyphal growth or sporulation but severely affected zoospore behavior including chemotaxis to the soybean isoflavone daidzein. Zoospore encystment and cyst germinating were also alternated, causing that the PsGPAl silenced mutants were disabled in infecting soybean. Upregulation of a calmodulin gene in zoospores of PsGPAl-silenced mutants pointed to suppressor activity of Ga targeted at steps in the calcium signaling pathway, indicating that Ga subunit may be a negative regulator of calcium signal pathways.
Screening for Ga binding proteins and analysis of Ga regulated signal transduction pathways. G protein a subunit of Phytophthora sojae influences not only sporangia cleavage and zoospore release, but also zoospore motility and cyst germination. The mechanisms that how Ga binds with effectors and regulates downstream signal pathways are unknown. In this study, we constructed a P. sojae cDNA library and used yeast two hybrid method to screen for Ga binding proteins, and got 7 putative Ga binding proteins including ribulose-phosphate 3-epimerase and putative reverse transcriptase. Meanwhile, we compared gene expression profiles of P. sojae zoospores between wild type P6497 and Ga mutants A2 and A27 by cDNA Macroarray. From the results of comparison,6 genes including 3 regulator of G protein (RGS),1 Ca2+/calmodulin protein kinase and 1 Ca2+ ATPase were expressed down-regulated by Ga silencing, however,1 calmodulin,2 Ca2+/calmodulin protein kinase and 1 homolog of transcriptional factor were expressed up-regulated by Ga silencing. These results indicate that RGS activity may be dependent on the activity of Ga in P. sojae. And the relationship between G protein and calcium signal is very complex.
引文
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