小麦应答禾谷丝核菌的3个激酶基因的分离及功能分析
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摘要
小麦纹枯病是小麦的重要土传真菌病害，主要是由禾谷丝核菌引起的。小麦纹枯病抗性遗传基础研究薄弱，常规抗病育种进展缓慢，目前生产中的抗纹枯病小麦品种匮乏，迫切需要研究小麦抗纹枯病反应机制，挖掘抗纹枯病的相关基因。蛋白激酶是植物免疫反应中的一类磷酸基团转移酶类，在植物免疫反应中发挥重要的作用。目前还没有蛋白激酶参与小麦抗纹枯病反应的报道。本研究通过基因芯片、RNA-Seq技术与半定量RT-PCR相结合的方法筛选出3个与小麦纹枯病抗性程度相关联的蛋白激酶基因，分别为TaCRK1基因、TaWAK5基因及TaPK-R1基因。
TaCRK1基因编码由680个氨基酸构成的富含半胱氨酸的受体蛋白激酶，受禾谷丝核菌和植物激素脱落酸诱导表达。TaWAK5基因编码由703个氨基酸构成的细胞壁关联的受体蛋白激酶，受禾谷丝核菌和植物激素水杨酸、脱落酸和茉莉酸甲酯诱导表达。TaCRK1蛋白与TaWAK5蛋白定位于细胞膜。利用病毒诱导的基因沉默(VIGS)技术分别沉默在抗纹枯病小麦CI12633中TaCRK1基因和TaWAK5基因，结果表明TaCRK1基因或TaWAK5基因表达量的下调并未减弱沉默植株对纹枯病的抗性，说明TaCRK1基因和TaWAK5基因并非小麦防御禾谷丝核菌反应所必需的。
TaPK-R1基因由13个外显子和12个内含子组成，编码由557个氨基酸构成的小麦AGC蛋白激酶。TaPK-R1基因受禾谷丝核菌、植物激素乙烯及过氧化氢的诱导表达；该基因在禾谷丝核菌接种21天的抗纹枯病小麦和感纹枯病小麦中的表达模式与小麦的纹枯病抗性程度相关联。洋葱表皮细胞与小麦原生质体的亚细胞定位分析表明TaPK-R1蛋白定位于细胞质与细胞核；通过体外自磷酸化分析明确了TaPK-R1蛋白具有自磷酸化活性。TaPK-R1表达量下降的抗纹枯病小麦CI12633植株对纹枯病的抗性减弱；TaPK-R1过表达的转TaPK-R1基因的小麦植株对纹枯病抗性显著提高，说明TaPK-R1基因是小麦防御纹枯病反应中所必需的。本研究首次证明了AGC家族蛋白激酶参与小麦对腐生营养型真菌的抗性反应，并通过遗传转化获得抗纹枯病的小麦新种质，为小麦抗病育种提供了新型的抗性材料和潜在的基因资源。
Wheat sharp eyespot, mainly caused by Rhizoctonia cerealis, is one of the most devasting diseases ofwheat in the world. Because the genetics of wheat resistance to R. cerealis is not fully understood andthe applied wheat cultivars are susceptible to sharp eyespot, it is urgent to unravel the mechanism ofwheat defense response to R. cerealis and to identify the genes involved in wheat defense response to R.cerealis. Protein kinase played important roles in perceiving pathogen infection, subsequent signalingpathways and ultimately inducing plant defense response. However, no kinase gene involved in wheatresistance to R. cerealis had yet been identified. In this study, using microarray, RNA-Sequencing(RNA-Seq), and semi-RT-PCR analysis, three protein kinase genes, including TaCRK1, TaWAK5, andTaPK-R1, were identified to be associated with the resistance degrees of wheat lines to R. cerealisinfection.
The TaCRK1gene encoded a cystein-rich receptor protein kinase with680amino acid residues.Transcription of TaCRK1was induced in CI12633after infection with R. cerealis and exogenousabscisic acid (ABA) treatment. The TaWAK5gene encoded a wall-associated kinase of703amino acidresidues. The expression of TaWAK5was induced by infection with R. cerealis and treatments withexogenous salicylic acid (SA), abscisic acid, and methyl jasmonate (MeJA). Subcellular localizationanalysis in onion epidermal cells or wheat protoplasts indicated that the TaCRK1protein and TaWAK5protein were localized to the plasma membranes. Characterization of TaCRK1silencing and TaWAK5induced by virus-mediated method in CI12633, respcetively showed that the down-regulation ofTaCRK1or TaWAK5transcript did not obviously impair resistance to R. cerealis, suggesting that thesetwo genes were not the major genes involved in the wheat defense response to R. cerealis.
The TaPK-R1gene, including13exons and12introns, encoded a wheat protein A/proteinG/protein C with557amino acid residues. The expression of the TaPK-R1transcript was induced by R.cerealis and by exogenous ethylene and hydrogen peroxide. Moreover, transcript abundance ofTaPK-R1were higher in the R. cerealis-resistant lines (CI12633, Shanhongmai, Navit14, Shannong0431, and Xifeng) than that in the R. cerealis-susceptible lines (Yangmai158, Wenma6, and Zhoumai18). Transient expression of TaPK-R1in onion epidermal cells or wheat protoplast indicated that theTaPK-R1protein was localized to the cytoplasm and nucleus. The protein kinase domain ofrecombinant GST-TaPK-R1showed autophosphorylated activity. Characterization of TaPK-R1silencinginduced by virus-mediated method in CI12633showed that down-regulation of TaPK-R1gene inCI12633significantly compromise wheat resisitance to R. cerealis. Over-expression of TaPK-R1gene intransgenic wheat plants confers increased resistance to wheat sharp eyespot accompanied withdecreased R. cerealis abundance. These results indicated that the TaPK-R1gene played an importantrole in wheat defense response to R. cerealis. This study indicated that the AGC protein kinase wasinvolved in plant defense response to necrotrophic pathogen in wheat for the first time. The TaPK-R1transgenic plants improved wheat resistance to R. cerealis and lay a foundation for unraveling the mechanis mof wheat defense response to sharp eyespot.
TaCRK1基因编码由680个氨基酸构成的富含半胱氨酸的受体蛋白激酶，受禾谷丝核菌和植物激素脱落酸诱导表达。TaWAK5基因编码由703个氨基酸构成的细胞壁关联的受体蛋白激酶，受禾谷丝核菌和植物激素水杨酸、脱落酸和茉莉酸甲酯诱导表达。TaCRK1蛋白与TaWAK5蛋白定位于细胞膜。利用病毒诱导的基因沉默(VIGS)技术分别沉默在抗纹枯病小麦CI12633中TaCRK1基因和TaWAK5基因，结果表明TaCRK1基因或TaWAK5基因表达量的下调并未减弱沉默植株对纹枯病的抗性，说明TaCRK1基因和TaWAK5基因并非小麦防御禾谷丝核菌反应所必需的。
TaPK-R1基因由13个外显子和12个内含子组成，编码由557个氨基酸构成的小麦AGC蛋白激酶。TaPK-R1基因受禾谷丝核菌、植物激素乙烯及过氧化氢的诱导表达；该基因在禾谷丝核菌接种21天的抗纹枯病小麦和感纹枯病小麦中的表达模式与小麦的纹枯病抗性程度相关联。洋葱表皮细胞与小麦原生质体的亚细胞定位分析表明TaPK-R1蛋白定位于细胞质与细胞核；通过体外自磷酸化分析明确了TaPK-R1蛋白具有自磷酸化活性。TaPK-R1表达量下降的抗纹枯病小麦CI12633植株对纹枯病的抗性减弱；TaPK-R1过表达的转TaPK-R1基因的小麦植株对纹枯病抗性显著提高，说明TaPK-R1基因是小麦防御纹枯病反应中所必需的。本研究首次证明了AGC家族蛋白激酶参与小麦对腐生营养型真菌的抗性反应，并通过遗传转化获得抗纹枯病的小麦新种质，为小麦抗病育种提供了新型的抗性材料和潜在的基因资源。
Wheat sharp eyespot, mainly caused by Rhizoctonia cerealis, is one of the most devasting diseases ofwheat in the world. Because the genetics of wheat resistance to R. cerealis is not fully understood andthe applied wheat cultivars are susceptible to sharp eyespot, it is urgent to unravel the mechanism ofwheat defense response to R. cerealis and to identify the genes involved in wheat defense response to R.cerealis. Protein kinase played important roles in perceiving pathogen infection, subsequent signalingpathways and ultimately inducing plant defense response. However, no kinase gene involved in wheatresistance to R. cerealis had yet been identified. In this study, using microarray, RNA-Sequencing(RNA-Seq), and semi-RT-PCR analysis, three protein kinase genes, including TaCRK1, TaWAK5, andTaPK-R1, were identified to be associated with the resistance degrees of wheat lines to R. cerealisinfection.
The TaCRK1gene encoded a cystein-rich receptor protein kinase with680amino acid residues.Transcription of TaCRK1was induced in CI12633after infection with R. cerealis and exogenousabscisic acid (ABA) treatment. The TaWAK5gene encoded a wall-associated kinase of703amino acidresidues. The expression of TaWAK5was induced by infection with R. cerealis and treatments withexogenous salicylic acid (SA), abscisic acid, and methyl jasmonate (MeJA). Subcellular localizationanalysis in onion epidermal cells or wheat protoplasts indicated that the TaCRK1protein and TaWAK5protein were localized to the plasma membranes. Characterization of TaCRK1silencing and TaWAK5induced by virus-mediated method in CI12633, respcetively showed that the down-regulation ofTaCRK1or TaWAK5transcript did not obviously impair resistance to R. cerealis, suggesting that thesetwo genes were not the major genes involved in the wheat defense response to R. cerealis.
The TaPK-R1gene, including13exons and12introns, encoded a wheat protein A/proteinG/protein C with557amino acid residues. The expression of the TaPK-R1transcript was induced by R.cerealis and by exogenous ethylene and hydrogen peroxide. Moreover, transcript abundance ofTaPK-R1were higher in the R. cerealis-resistant lines (CI12633, Shanhongmai, Navit14, Shannong0431, and Xifeng) than that in the R. cerealis-susceptible lines (Yangmai158, Wenma6, and Zhoumai18). Transient expression of TaPK-R1in onion epidermal cells or wheat protoplast indicated that theTaPK-R1protein was localized to the cytoplasm and nucleus. The protein kinase domain ofrecombinant GST-TaPK-R1showed autophosphorylated activity. Characterization of TaPK-R1silencinginduced by virus-mediated method in CI12633showed that down-regulation of TaPK-R1gene inCI12633significantly compromise wheat resisitance to R. cerealis. Over-expression of TaPK-R1gene intransgenic wheat plants confers increased resistance to wheat sharp eyespot accompanied withdecreased R. cerealis abundance. These results indicated that the TaPK-R1gene played an importantrole in wheat defense response to R. cerealis. This study indicated that the AGC protein kinase wasinvolved in plant defense response to necrotrophic pathogen in wheat for the first time. The TaPK-R1transgenic plants improved wheat resistance to R. cerealis and lay a foundation for unraveling the mechanis mof wheat defense response to sharp eyespot.
引文
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