番茄白粉病抗性反应关键基因的鉴定及功能验证
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摘要
番茄白粉菌(Oidium neolycopersici)是一种产生单生分生孢子的真菌,主要分布在欧洲、非洲、南北美洲和亚洲的一种世界性植物病害病原菌。番茄白粉病严重影响了番茄的产量和质量,而杀菌剂的大量喷施造成了环境污染,防治该病害最关键有效的方法是培育抗病品种,而抗病品种选育的成效取决于抗病基因资源的挖掘和利用。本文利用建立的病毒诱导基因沉默(virus induced gene silencing, VIGS)的功能基因组学研究平台,对番茄白粉病抗性反应关键基因进行了鉴定及功能验证,在细胞和基因水平上研究了番茄对白粉病的抗性机理,为番茄抗病育种提供理论依据。主要研究结果如下:
1 .对番茄白粉菌商丘菌株进行了鉴定,在国内首次报道了番茄白粉菌(O.neolycopersici)。对河南商丘分离出的番茄白粉病病原菌进行了致病性检测、形态学及分子生物学鉴定。菌落形态、显微结构及核糖体DNA(rDNA)内转录间隔区(internal transcribed spacer, ITS)序列分析表明,该病原菌属于番茄白粉菌(O. neolycopersici)。番茄抗性材料特异性分析和品种抗病性评价表明,人工接种该病原菌导致番茄栽培品种Solanum lycopersicum致病,而野生品种S. habrochaites G1.1560(携带抗性基因Ol-1)和S. arcanum LA2172(携带抗性基因Ol-4)表现为高抗。
2.对番茄与白粉菌互作中活性氧的积累及生理生化反应进行分析。番茄与白粉菌亲和互作中感病番茄叶片细胞活性氧积累较慢、较少,无法完全抑制白粉菌的生长;而在非亲和互作中,抗病番茄叶片细胞活性氧积累较快、较多,能够在短时间内抑制白粉菌生长,使其无法完成生活周期,Ol-1和Ol-4介导的番茄白粉病抗性反应与快速过敏反应(hypersensitive reaction,HR)相关。通过防御反应相关酶活性分析,研究了核黄素诱导的番茄系统抗病性,发现细胞内活性氧积累速度和程度、膜脂过氧化、PAL和POD在核黄素诱导番茄对白粉菌的抗性反应中发挥重要作用。
3.建立番茄的VIGS体系,对前期研究获得的白粉菌诱导的上调番茄抗性候选基因序列进行功能鉴定,确定抗性反应的关键基因。对高抗白粉病番茄植株S. habrochaites G1.1560(携带抗性基因Ol-1)进行候选基因的沉默,观察到不同程度抗性丧失的表型。进而对沉默植株进行显微形态学观察,发现白粉菌可在沉默植株完成其整个无性生活周期,沉默植株对白粉菌表现为慢性过敏反应,而对照植株表现为快速过敏反应。根据沉默后表型变化鉴定出4个番茄白粉病抗性反应关键基因:ShGSTU1、ShME1、ShMADSTF、ShORR-1。BLAST分析预测关键基因的编码产物,其中ShGSTU1编码谷胱甘肽转移酶(glutathione transferases, GST),ShME1编码NADP苹果酸酶(NADP-malic enzyme, NADP-ME),ShMADSTF编码MADS盒转录因子(transcription factor, TF),而ShORR-1为功能未知的新基因。依据番茄全基因组序列对ShGSTU1、ShME1和ShORR-1进行基因步移,设计引物,PCR扩增,测序获得基因的全长(ShGSTU1 GengBank登录号:JF957860;ShME1 GengBank登录号:JF957861)。
Tomato powdery mildew caused by Oidium neolycopersici is a kind of fungus with a single-celled conidium produced on conidiosphore, which is a worldwide plant fungal disease distributed in Europe, African, South and North America and Asian. Tomato powdery mildew is responsible for a remarkable reduction in quality and yield of tomato, while insecticide spraying to stop the disease results in environmental contamination. The most effective way to prevent this disease is resistant breeding, which depends on the resistant materials and resistance genes. In the paper, virus induced gene silencing(VIGS)as a functional genomic research platform established was used to identify and functional analyze key genes for tomato resistance to powdery mildew, and resistance mechanism of tomato to powdery mildew was studied in the cellular and genetic levels, which provided theoretical basis for tomato resistant breeding. The main results were as follows:
1. The O. neolycopersici Shangqiu isolate was identified as the causual agent of tomato powdery mildew spreading in Shangqiu, which was the first report of O. neolycopersici in China. The pathogen of tomato powdery mildew was studied based on colony morphology, histological structure and internal transcribed spacer (ITS) sequence of rDNA. It indicated that the pathogen was O. neolycopersici. The wild types and cultivated varieties of tomato were inoculated with the fungi for disease test. Tomato cultivated varieties (Solanum lycopersicum) were susceptible to the fungi, while the tomato wild types S. habrochaites G1.1560 (carrying Ol-1 gene)and S. arcanum LA2172 (carrying Ol-4 gene)were highly resistant.
2. Active oxygen accumulation, physiological and biochemical changes in different tomato species to powdery mildew fungi were examined in the cellular level. It indicated that the slow accumulation speed and low quantity of active oxygen resulted in incompletely inhibiting the growth of powdery mildew in susceptive tomato, while the fast accumulation and high quantity of active oxygen can inhibit the growth of powdery mildew in resistant tomato successfully, in which the pathogen can’t finish it life cycle. The tomato resistance responses to powdery mildew mediated by Ol-1 or Ol-4 gene were involved with fast hypersensitive reaction. Systemic acquired resistance of tomato induced by unspecialized elicitor– riboflavin was studied by analyzing enzyme activities related to resistant pathway. The results suggested that riboflavin induced physiological changes including H2O2 accumulation, lipid peroxidation, POD and PAL could play a role in the increase of tomato resistance to O. neolycopersici.
3. VIGS system was used to analyze the function of the up-regulated candidate gene sequences associated with tomato resistance to powdery mildew. It was found silencing of candidate gene sequences resulted in resistance-breaking phenotype of different degrees in S. habrochaites G1.1560(carrying Ol-1 gene). Microscopic observation showed that O. neolycopersici can finish its whole life cycle and a slow hypersensitive reaction was displayed in epidermal cells of silenced plant leaves compared to fast hypersensitive reaction in control plant. Based on the results of VIGS analysis and microscopic observation, four genes ShGSTU1, ShME1, ShMADSTF and ShORR-1 were identified as key genes for tomato resistance response to powdery mildew. By BLAST, the putative encoding proteins of these genes were predicted, ShGSTU1 encoding glutathione transferases, ShME1 encoding NADP-malic enzyme, ShMADSTF encoding MADS box transcription factor, and ShORR-1 encoding an unknown function protein. Primers for obtaining complete sequences of ShGSTU1、ShME1 and ShORR-1 were designed based on gene walking in tomato genome sequence. PCR products of ShGSTU1、ShME1 and ShORR-1 amplified from S. habrochaites G1.1560 genemic DNA were sequenced to get the complete sequence. The GeneBank code of ShGSTU1 was JF957860, ShME1 was JF957861.
1 .对番茄白粉菌商丘菌株进行了鉴定,在国内首次报道了番茄白粉菌(O.neolycopersici)。对河南商丘分离出的番茄白粉病病原菌进行了致病性检测、形态学及分子生物学鉴定。菌落形态、显微结构及核糖体DNA(rDNA)内转录间隔区(internal transcribed spacer, ITS)序列分析表明,该病原菌属于番茄白粉菌(O. neolycopersici)。番茄抗性材料特异性分析和品种抗病性评价表明,人工接种该病原菌导致番茄栽培品种Solanum lycopersicum致病,而野生品种S. habrochaites G1.1560(携带抗性基因Ol-1)和S. arcanum LA2172(携带抗性基因Ol-4)表现为高抗。
2.对番茄与白粉菌互作中活性氧的积累及生理生化反应进行分析。番茄与白粉菌亲和互作中感病番茄叶片细胞活性氧积累较慢、较少,无法完全抑制白粉菌的生长;而在非亲和互作中,抗病番茄叶片细胞活性氧积累较快、较多,能够在短时间内抑制白粉菌生长,使其无法完成生活周期,Ol-1和Ol-4介导的番茄白粉病抗性反应与快速过敏反应(hypersensitive reaction,HR)相关。通过防御反应相关酶活性分析,研究了核黄素诱导的番茄系统抗病性,发现细胞内活性氧积累速度和程度、膜脂过氧化、PAL和POD在核黄素诱导番茄对白粉菌的抗性反应中发挥重要作用。
3.建立番茄的VIGS体系,对前期研究获得的白粉菌诱导的上调番茄抗性候选基因序列进行功能鉴定,确定抗性反应的关键基因。对高抗白粉病番茄植株S. habrochaites G1.1560(携带抗性基因Ol-1)进行候选基因的沉默,观察到不同程度抗性丧失的表型。进而对沉默植株进行显微形态学观察,发现白粉菌可在沉默植株完成其整个无性生活周期,沉默植株对白粉菌表现为慢性过敏反应,而对照植株表现为快速过敏反应。根据沉默后表型变化鉴定出4个番茄白粉病抗性反应关键基因:ShGSTU1、ShME1、ShMADSTF、ShORR-1。BLAST分析预测关键基因的编码产物,其中ShGSTU1编码谷胱甘肽转移酶(glutathione transferases, GST),ShME1编码NADP苹果酸酶(NADP-malic enzyme, NADP-ME),ShMADSTF编码MADS盒转录因子(transcription factor, TF),而ShORR-1为功能未知的新基因。依据番茄全基因组序列对ShGSTU1、ShME1和ShORR-1进行基因步移,设计引物,PCR扩增,测序获得基因的全长(ShGSTU1 GengBank登录号:JF957860;ShME1 GengBank登录号:JF957861)。
Tomato powdery mildew caused by Oidium neolycopersici is a kind of fungus with a single-celled conidium produced on conidiosphore, which is a worldwide plant fungal disease distributed in Europe, African, South and North America and Asian. Tomato powdery mildew is responsible for a remarkable reduction in quality and yield of tomato, while insecticide spraying to stop the disease results in environmental contamination. The most effective way to prevent this disease is resistant breeding, which depends on the resistant materials and resistance genes. In the paper, virus induced gene silencing(VIGS)as a functional genomic research platform established was used to identify and functional analyze key genes for tomato resistance to powdery mildew, and resistance mechanism of tomato to powdery mildew was studied in the cellular and genetic levels, which provided theoretical basis for tomato resistant breeding. The main results were as follows:
1. The O. neolycopersici Shangqiu isolate was identified as the causual agent of tomato powdery mildew spreading in Shangqiu, which was the first report of O. neolycopersici in China. The pathogen of tomato powdery mildew was studied based on colony morphology, histological structure and internal transcribed spacer (ITS) sequence of rDNA. It indicated that the pathogen was O. neolycopersici. The wild types and cultivated varieties of tomato were inoculated with the fungi for disease test. Tomato cultivated varieties (Solanum lycopersicum) were susceptible to the fungi, while the tomato wild types S. habrochaites G1.1560 (carrying Ol-1 gene)and S. arcanum LA2172 (carrying Ol-4 gene)were highly resistant.
2. Active oxygen accumulation, physiological and biochemical changes in different tomato species to powdery mildew fungi were examined in the cellular level. It indicated that the slow accumulation speed and low quantity of active oxygen resulted in incompletely inhibiting the growth of powdery mildew in susceptive tomato, while the fast accumulation and high quantity of active oxygen can inhibit the growth of powdery mildew in resistant tomato successfully, in which the pathogen can’t finish it life cycle. The tomato resistance responses to powdery mildew mediated by Ol-1 or Ol-4 gene were involved with fast hypersensitive reaction. Systemic acquired resistance of tomato induced by unspecialized elicitor– riboflavin was studied by analyzing enzyme activities related to resistant pathway. The results suggested that riboflavin induced physiological changes including H2O2 accumulation, lipid peroxidation, POD and PAL could play a role in the increase of tomato resistance to O. neolycopersici.
3. VIGS system was used to analyze the function of the up-regulated candidate gene sequences associated with tomato resistance to powdery mildew. It was found silencing of candidate gene sequences resulted in resistance-breaking phenotype of different degrees in S. habrochaites G1.1560(carrying Ol-1 gene). Microscopic observation showed that O. neolycopersici can finish its whole life cycle and a slow hypersensitive reaction was displayed in epidermal cells of silenced plant leaves compared to fast hypersensitive reaction in control plant. Based on the results of VIGS analysis and microscopic observation, four genes ShGSTU1, ShME1, ShMADSTF and ShORR-1 were identified as key genes for tomato resistance response to powdery mildew. By BLAST, the putative encoding proteins of these genes were predicted, ShGSTU1 encoding glutathione transferases, ShME1 encoding NADP-malic enzyme, ShMADSTF encoding MADS box transcription factor, and ShORR-1 encoding an unknown function protein. Primers for obtaining complete sequences of ShGSTU1、ShME1 and ShORR-1 were designed based on gene walking in tomato genome sequence. PCR products of ShGSTU1、ShME1 and ShORR-1 amplified from S. habrochaites G1.1560 genemic DNA were sequenced to get the complete sequence. The GeneBank code of ShGSTU1 was JF957860, ShME1 was JF957861.
引文
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