水稻抗虫相关基因OsWRKY24、OsWRKY70和OsNPR1的功能解析
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摘要
在自然条件下,水稻经常会受到各种害虫的取食,比如二化螟(SSB,Chilo suppressalis)、褐飞虱(BPH,Niaparvata lugens)、稻纵卷叶螟(LF,Cnaphalocrocis medinalis)等。植食性昆虫取食造成了水稻转录组发生了很大的变化,虫害诱导的防御反应信号途径被激活,抗性基因得以表达,其中转录调控因子起了非常重要的调控作用。然而,目前对在诱导抗虫反应中起作用的转录调控因子还了解得很少。本文中我们选取了两个水稻中植食性昆虫诱导的转录因子以及OsNPR1基因,对它们在植物抗虫途径中的作用机理进行了分析：
1) OsWRKY70基因可以被机械损伤、二化螟取食强烈诱导,而褐飞虱取食微弱诱导该基因的表达。OsWRKY70定位在细胞核中,可以和W-box结合,具有转录激活活性。体外GST pull-down实验证明OsWRKY70可以和OsMPK3和OsMPK6互作,并且位于MAPK的下游。OsWRKY70正调控茉莉酸(jasmonic acid, JA)和胰蛋白酶抑制剂(TrypPIs)合成及水稻对SSB的抗性,而负调控赤霉素(gibberellin acid, GA)、水杨酸(salicylic acid,SA)、过氧化氢(H2O2)合成及水稻对BPH的抗性。其中我们首次发现了GA信号途径参与了植物应答植食性昆虫的抗性。OsWRKY70是一个各种虫害诱导防御网络中的关键节点。
2) OsWRKY24基因可以被机械损伤、二化螟取食诱导上调,而褐飞虱取食微弱的诱导该基因的表达。该基因也是位于MAPK下游、微弱的正调控了JA和TrypPIs合成及水稻对SSB的抗性,而对BPH的抗性影响不大。通过构建ir-wrky24/70双突变体,我们发现OsWRKY24基因在二化螟诱导的抗性途径中贡献不大。
3) OsNPR1基因可以被机械损伤、二化螟取食、稻纵卷叶螟取食和信号分子SA、JA诱导上调,而机械损伤和SA诱导后该基因的表达要明显早于虫害诱导。反义抑制该基因后提高了二化螟诱导的JA、ET含量以及二化螟诱导的JA合成酶OsHI-LOX以及ET合成酶OsACS2基因的表达,并且提高了虫害诱导的TrypPIs的含量、挥发物的含量以及对二化螟的抗性。我们的结果表明OsNPR1是一个虫害诱导早期的响应基因,并且可以通过调节激素信号途径来有效的、特异的调控植食性昆虫诱导的抗性。
我们的研究结果表明三个基因通过调控激素信号途径参与了水稻植食性昆虫诱导的防御反应。
In natural, rice plants are always suffered by different herbivores, for example, rice striped stem borer (SSB) Chilo suppressalis, rice brown planthopper (BPH) Niaparvata lugens, rice leaf folder (LF) Cnaphalocrocis medinalis, and so on. Rice large-scale transcriptional changes include activating herbivore induced defense signal pathways and some defense-related genes after infesting by herbivores. Among this, the transcription regulators play a central role. Here, we selected two herbivore induced transcription factors and OsNPRl gene and study the function in herbivore induced resisitance.
1) OsWRKY70is strongly induced by mechnical wounding and SSB infested, but weakly induced by BPH infested. OsWRKY70localized in the nucleus, binding W-box motif and is a transcription activator. In vitro pull-down assay demonstrated that OsWRKY70interacted with OsMPK3and OsMPK6, and act downstream of MAPKs. OsWRKY70positively regulated JA, TrypPIs levels and plant resistance to SSB, whereas negatively regulated GA, SA and H2O2levels and rice resistance to BPH. It's the first time we demastrated that GA signal pathway involved in herbivore induced defense. OsWRKY70is a key node in different herbivore induced signal network.
2) OsWRKY24is strongly induced by mechnical wounding and SSB infested, but weakly induced by BPH infested. OsWRKY24also act downstream of MAPKs, slight positively affect JA and TrypPIs levels and rice resistance to SSB, however it has no affection to BPH. We found OsWRKY24contribute little to rice SSB resistance by detected OsWRKY24and OsWRKY70doubt mutant ir-wrky24/70.
3) OsNPR1expression levels were up-regulated in response to infestation by the rice striped stem borer (SSB) Chilo suppressalis and rice leaf folder (LF) Cnaphalocrocis medinalis, and to mechanical wounding and treatment with jasmonic acid (JA) and salicylic acid (SA). Moreover, mechanical wounding induced the expression of OsNPR1more quickly than did herbivore infestation. The antisense expression of OsNPR1(as-nprl), which reduced the expression of the gene by50%, increased elicited levels of JA and ethylene (ET) as well as of expression of a lipoxygenase gene OsHI-LOX and an ACC synthase gene OsACS2. The enhanced JA and ethylene signaling in as-nprl plants increased the levels of herbivore-induced trypsin protease inhibitors (TrypPIs) and volatiles, and reduced the performance of SSB. Our results suggest that OsNPR1is an early responding gene in herbivore-induced defense and that plants can use it to activate an effective and specific defense against invaders by modulating signaling pathways. Our result indicated that all three genes involved in rice herbivore induced denfense by regulate plant hormone signal pathways.
1) OsWRKY70基因可以被机械损伤、二化螟取食强烈诱导,而褐飞虱取食微弱诱导该基因的表达。OsWRKY70定位在细胞核中,可以和W-box结合,具有转录激活活性。体外GST pull-down实验证明OsWRKY70可以和OsMPK3和OsMPK6互作,并且位于MAPK的下游。OsWRKY70正调控茉莉酸(jasmonic acid, JA)和胰蛋白酶抑制剂(TrypPIs)合成及水稻对SSB的抗性,而负调控赤霉素(gibberellin acid, GA)、水杨酸(salicylic acid,SA)、过氧化氢(H2O2)合成及水稻对BPH的抗性。其中我们首次发现了GA信号途径参与了植物应答植食性昆虫的抗性。OsWRKY70是一个各种虫害诱导防御网络中的关键节点。
2) OsWRKY24基因可以被机械损伤、二化螟取食诱导上调,而褐飞虱取食微弱的诱导该基因的表达。该基因也是位于MAPK下游、微弱的正调控了JA和TrypPIs合成及水稻对SSB的抗性,而对BPH的抗性影响不大。通过构建ir-wrky24/70双突变体,我们发现OsWRKY24基因在二化螟诱导的抗性途径中贡献不大。
3) OsNPR1基因可以被机械损伤、二化螟取食、稻纵卷叶螟取食和信号分子SA、JA诱导上调,而机械损伤和SA诱导后该基因的表达要明显早于虫害诱导。反义抑制该基因后提高了二化螟诱导的JA、ET含量以及二化螟诱导的JA合成酶OsHI-LOX以及ET合成酶OsACS2基因的表达,并且提高了虫害诱导的TrypPIs的含量、挥发物的含量以及对二化螟的抗性。我们的结果表明OsNPR1是一个虫害诱导早期的响应基因,并且可以通过调节激素信号途径来有效的、特异的调控植食性昆虫诱导的抗性。
我们的研究结果表明三个基因通过调控激素信号途径参与了水稻植食性昆虫诱导的防御反应。
In natural, rice plants are always suffered by different herbivores, for example, rice striped stem borer (SSB) Chilo suppressalis, rice brown planthopper (BPH) Niaparvata lugens, rice leaf folder (LF) Cnaphalocrocis medinalis, and so on. Rice large-scale transcriptional changes include activating herbivore induced defense signal pathways and some defense-related genes after infesting by herbivores. Among this, the transcription regulators play a central role. Here, we selected two herbivore induced transcription factors and OsNPRl gene and study the function in herbivore induced resisitance.
1) OsWRKY70is strongly induced by mechnical wounding and SSB infested, but weakly induced by BPH infested. OsWRKY70localized in the nucleus, binding W-box motif and is a transcription activator. In vitro pull-down assay demonstrated that OsWRKY70interacted with OsMPK3and OsMPK6, and act downstream of MAPKs. OsWRKY70positively regulated JA, TrypPIs levels and plant resistance to SSB, whereas negatively regulated GA, SA and H2O2levels and rice resistance to BPH. It's the first time we demastrated that GA signal pathway involved in herbivore induced defense. OsWRKY70is a key node in different herbivore induced signal network.
2) OsWRKY24is strongly induced by mechnical wounding and SSB infested, but weakly induced by BPH infested. OsWRKY24also act downstream of MAPKs, slight positively affect JA and TrypPIs levels and rice resistance to SSB, however it has no affection to BPH. We found OsWRKY24contribute little to rice SSB resistance by detected OsWRKY24and OsWRKY70doubt mutant ir-wrky24/70.
3) OsNPR1expression levels were up-regulated in response to infestation by the rice striped stem borer (SSB) Chilo suppressalis and rice leaf folder (LF) Cnaphalocrocis medinalis, and to mechanical wounding and treatment with jasmonic acid (JA) and salicylic acid (SA). Moreover, mechanical wounding induced the expression of OsNPR1more quickly than did herbivore infestation. The antisense expression of OsNPR1(as-nprl), which reduced the expression of the gene by50%, increased elicited levels of JA and ethylene (ET) as well as of expression of a lipoxygenase gene OsHI-LOX and an ACC synthase gene OsACS2. The enhanced JA and ethylene signaling in as-nprl plants increased the levels of herbivore-induced trypsin protease inhibitors (TrypPIs) and volatiles, and reduced the performance of SSB. Our results suggest that OsNPR1is an early responding gene in herbivore-induced defense and that plants can use it to activate an effective and specific defense against invaders by modulating signaling pathways. Our result indicated that all three genes involved in rice herbivore induced denfense by regulate plant hormone signal pathways.
引文
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