水稻抗虫相关基因OsICS和OsHPL3的功能分析
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摘要
当植物受到病原菌及植食性昆虫危害后,会产生一系列的诱导防御反应。在一些双子叶植物如拟南芥和烟草等中的研究表明,水杨酸和绿叶性气味分别可作为信号分子及防御化合物在植物的诱导防御反应过程中发挥着重要作用。然而,作为单子叶模式植物的水稻,在这方面的研究还很少。为此,我们想通过实验进一步明确在单子叶模式植物水稻中是否也存在着一条异分支酸途径来合成水杨酸,以及该途径在抗虫防御反应中究竟发挥着什么作用?同时,与作为防御反应相关的重要物质茉莉酸竞争相同底物的绿叶性气味在水稻对白背飞虱的抗性防御反应也未见报道。因此,本文选择受褐飞虱取食诱导的水稻异分支酸合成酶基因OsICS和合成水稻绿叶性气味的脂氢过氧化物裂解酶基因OsHPL3为研究对象,结合利用反向遗传学、化学分析、分子生物学以及生物测定等研究方法,对OsICS的抗虫相关功能及OsHPL3调控的绿叶性气味在水稻对白背飞虱抗性防御反应中的作用进行了研究,获得的主要结果如下：
克隆了异分支酸合成酶基因OsICS,该基因的开放阅读框为2445bp,编码一个由814个氨基酸组成的蛋白质,预测分子量为88.024KD,pI值为5.92。OsICS编码的蛋白定位于叶绿体。组织表达谱分析发现OsICS的转录水平在水稻叶内要明显高于水稻中的其它部位；诱导表达谱分析发现褐飞虱(Nilaparvata lugens, rice brown planthopper, BPH)、白背飞虱(white back planthopper, WBPH)和二化螟(Chilo suppressalis, rice striped stem borer, SSB)取食、机械损伤处理均能诱导OsICS基因的表达,而外源激素茉莉酸(Jasmonic acid, JA)和水杨酸(Salicylic acid, SA)不诱导其表达,甚至后期有一定的抑制作用。我们利用农杆菌转化法,获得了水稻OsICS反义抑制突变体植株,经检测,该反义纯合品系中OsICS的转录水平比野生型水稻中分别降低了46.23%-50.86%。各品系表型研究结果表明,OsICS反义突变体植株在水稻萌发期及温室内种植时均显著矮于野生型水稻。同时,温室内种植的水稻反义品系本底SA含量明显低于野生型,且室内各品系受褐飞虱和二化螟取食诱导后,OsICS反义品系中SA含量也显著低于野生型水稻。我们接下来利用该突变体中SA含量的降低,对不同害虫进行了生测实验。结果表明,反义品系能明显增加水稻对BPH和WBPH的选择性,但产卵量并无明显差异,当外源SA处理后则可恢复BPH和WBPH对突变体的取食选择性；OsICS反义品系并不影响虫害诱导的水稻JA、胰蛋白酶抑制剂(trypsin protease inhibitors, TrypPIs)和乙烯(ethylene, ET)的合成,同时对BPH、WBPH的若虫发育历期和二化螟的生长均无明显影响。
水稻OsHPL3表达谱分析表明,机械损伤、SSB、BPH和WBPH为害均能够诱导OsHPL3的表达,而外源激素JA和SA处理对其无明显影响。我们利用相同方法获得了反义抑制OsHPL3单拷贝纯合品系。实验发现,OsHPL3表达水平的下调会引起突变体受机械损伤和白背飞虱取食诱导的GLV (Z)-3-hexenal、(E)-2-hexenal和1(Z)-3-hexen-1-ol的含量显著下降,该反义突变体明显提高了水稻对刺吸式口器害虫WBPH的抗性,可导致WBPH成虫对转基因水稻的选择性、产卵量及取食量均明显下降。同时,取食反义品系水稻的WBPH雌雄若虫发育历期均显著延长,存活率比野生型明显下降,以反义品系为食的若虫孵化后继续在该品系上产卵,其产卵量也显著低于以野生型植株为食的对照植株。当分别回补不同浓度的GLV后,发现(Z)-3-hexenal处理能恢复OsHPL3反义品系对WBPH的抗性影响。挥发物测定结果表明,反义品系能明显增加WBPH为害诱导的(+)-limonene,2-heptanol, sesquisabinene和β-sesquiphellandrene的释放量,其释放总量也有一定程度增加,但未达到显著差异水平,其他挥发性物质的释放量并无明显差异。对WBPH天敌稻虱缨小蜂的寄主选择性实验表明,小蜂更多地选择能合成释放较多GLV的野生型水稻植株,当外源回补3种不同浓度的GLV(Z)-3-hexenal、(E)-2-hexenal和(Z)-3-hexen-1-ol处理可以恢复反义品系对稻虱缨小蜂选择性的影响。综合以上结果表明,OsHPL3通过影响GLV的合成来调控水稻对WBPH的直接和间接防御反应。
Herbivore induced plant defense response, a complex physiological and biochemical processes, can produce when exposed to pathogen and phytophagous insects. Salicylic acid and green leaf volatiles, as signal molecules, play an important role in the process of herbivore induced plant defense responses in dicotyledonous plants such as Arabidopsis and tobacco. However, little to nothing is known in this aspect in rice, a model monocot species. So further research was made on whether isochorismate synthase was also involved in the synthesis of salicylic acid, and whether this pathway played an important role in rice defense responses. Also, little is known about functional analysis of OsHPL3, whose product competing the same substrate with jasmonic acid, against WBPH resistance. Therefore, we isolated two genes as the research object, one is isochorismate synthase gene OsICS, induced by brown planthopper feeding, and the other is hydroperoxide lyase gene OsHPL3, whose production green leaves volatile, combining with the methods of reverse genetics, molecular biology, chemical analysis and bioassay, in order to clarify OsICS function in rice herbivore induced resistance and OsHPL3against WBPH resistance defense responses. The main results are as follows:
A2445bp open reading frame of an herbivore specifically induced isochorismate synthase gene OsICS, which encodes a protein of814amino acids with a calculated molecular weight of88.024KDa and pI5.92, was cloned. OsICS coding protein is located in the chloroplasts. Tissue specific expression analysis revealed OsICS transcript levels in rice leaves was higher than that of other parts. Induced expression analysis showed that the feeding of Nilaparvata lugens (brown planthopper, BPH), Sogatella furrifera (white back planthopper, WBPH), Chilo suppressalis (rice striped stem borer, SSB) and mechanical wounding treatment can induce OsICS gene expression, and exogenous jasmonic acid (JA) and salicylic acid (SA) can not induce the expression, but have an opposite effect. Through agrobacterium-mediated transgenic method, we obtained the OsICS mutant plants. The OsICS expression levels in transgenic lines were decreased by46.23%-50.86%than the wild type. Phenotypic results of different lines revealed OsICS transgenic mutants are significantly shorter than the wild type during germination and grown in the field. Besides, the level of SA in transgenic mutants are significantly lower than the wild type both grown in the field and green house induced by the feeding of brown planthopper and rice stem borer. The bioassay of different pests results show that, the OsICS transgenic lines could obviously increase the feeding choice of BPH and WBPH, and the treatment of exogenous SA can restore BPH and WBPH feeding choice. However, the OsICS transgenic lines can not affect herbivore induced rice JA, trypsin inhibitor (TrypPIs) and ethylene (ET). And the transgenic lines has no significant effect on the nymph developmental duration and population growth of BPH and WBPH.
OsHPL3is a hydroperoxide lyase gene. Expression levels of OsHPL3could be significantly enhanced by mechanical wounding, SSB, BPH and WBPH infestation, whereas exogenous hormones JA and SA treatment had no significant effect. We use the same method to aquire the OsHPL3transgenic lines. The emission of GLV levels (Z)-3-hexenal,(Z)-3-Hexen-l-ol and (E)-2-hexenal were significantly lower in transgenic lines than in wild type rice plants after WBPH infestation and mechanical wounding treatment. At the same time, the transgenic lines can increase the resistance to WBPH. Bioassay results show that adult WBPH preferred to feed and oviposite on the transgenic lines. The nymph developmental duration in the transgenic lines were significantly longer, the survival rate was significantly decreased, and the number of eggs per day were significantly lower than in the wild type. While the treatment of exogenous (Z)-3-hexenal can restore the0sHPL3transgenic lines to the resistance of WBPH. The result of Chromatograms of volatiles obtained from various lines show that transgenic strains can increase the release of limonene after WBPH damage, while the decrease expression of OsHPL3in transgenic lines had no effect on other WBPH-induced plant volatiles. Compared with transgenic plants, Anagrus nilaparvatae, which is an egg parasitoid of WBPH, preferred the odors from wild type plants, releasing more GLV after WBPH infestation. However, exogenous different concentrations of GLV,(Z)-3-hexenal,(Z)-3-Hexen-l-ol and (E)-2-hexenal treatment can restore the Anagrus nilaparvatae selectively behavior between different lines.So GLV, the product of OsHPL3, play an important role in the rice resistance to WBPH.
克隆了异分支酸合成酶基因OsICS,该基因的开放阅读框为2445bp,编码一个由814个氨基酸组成的蛋白质,预测分子量为88.024KD,pI值为5.92。OsICS编码的蛋白定位于叶绿体。组织表达谱分析发现OsICS的转录水平在水稻叶内要明显高于水稻中的其它部位；诱导表达谱分析发现褐飞虱(Nilaparvata lugens, rice brown planthopper, BPH)、白背飞虱(white back planthopper, WBPH)和二化螟(Chilo suppressalis, rice striped stem borer, SSB)取食、机械损伤处理均能诱导OsICS基因的表达,而外源激素茉莉酸(Jasmonic acid, JA)和水杨酸(Salicylic acid, SA)不诱导其表达,甚至后期有一定的抑制作用。我们利用农杆菌转化法,获得了水稻OsICS反义抑制突变体植株,经检测,该反义纯合品系中OsICS的转录水平比野生型水稻中分别降低了46.23%-50.86%。各品系表型研究结果表明,OsICS反义突变体植株在水稻萌发期及温室内种植时均显著矮于野生型水稻。同时,温室内种植的水稻反义品系本底SA含量明显低于野生型,且室内各品系受褐飞虱和二化螟取食诱导后,OsICS反义品系中SA含量也显著低于野生型水稻。我们接下来利用该突变体中SA含量的降低,对不同害虫进行了生测实验。结果表明,反义品系能明显增加水稻对BPH和WBPH的选择性,但产卵量并无明显差异,当外源SA处理后则可恢复BPH和WBPH对突变体的取食选择性；OsICS反义品系并不影响虫害诱导的水稻JA、胰蛋白酶抑制剂(trypsin protease inhibitors, TrypPIs)和乙烯(ethylene, ET)的合成,同时对BPH、WBPH的若虫发育历期和二化螟的生长均无明显影响。
水稻OsHPL3表达谱分析表明,机械损伤、SSB、BPH和WBPH为害均能够诱导OsHPL3的表达,而外源激素JA和SA处理对其无明显影响。我们利用相同方法获得了反义抑制OsHPL3单拷贝纯合品系。实验发现,OsHPL3表达水平的下调会引起突变体受机械损伤和白背飞虱取食诱导的GLV (Z)-3-hexenal、(E)-2-hexenal和1(Z)-3-hexen-1-ol的含量显著下降,该反义突变体明显提高了水稻对刺吸式口器害虫WBPH的抗性,可导致WBPH成虫对转基因水稻的选择性、产卵量及取食量均明显下降。同时,取食反义品系水稻的WBPH雌雄若虫发育历期均显著延长,存活率比野生型明显下降,以反义品系为食的若虫孵化后继续在该品系上产卵,其产卵量也显著低于以野生型植株为食的对照植株。当分别回补不同浓度的GLV后,发现(Z)-3-hexenal处理能恢复OsHPL3反义品系对WBPH的抗性影响。挥发物测定结果表明,反义品系能明显增加WBPH为害诱导的(+)-limonene,2-heptanol, sesquisabinene和β-sesquiphellandrene的释放量,其释放总量也有一定程度增加,但未达到显著差异水平,其他挥发性物质的释放量并无明显差异。对WBPH天敌稻虱缨小蜂的寄主选择性实验表明,小蜂更多地选择能合成释放较多GLV的野生型水稻植株,当外源回补3种不同浓度的GLV(Z)-3-hexenal、(E)-2-hexenal和(Z)-3-hexen-1-ol处理可以恢复反义品系对稻虱缨小蜂选择性的影响。综合以上结果表明,OsHPL3通过影响GLV的合成来调控水稻对WBPH的直接和间接防御反应。
Herbivore induced plant defense response, a complex physiological and biochemical processes, can produce when exposed to pathogen and phytophagous insects. Salicylic acid and green leaf volatiles, as signal molecules, play an important role in the process of herbivore induced plant defense responses in dicotyledonous plants such as Arabidopsis and tobacco. However, little to nothing is known in this aspect in rice, a model monocot species. So further research was made on whether isochorismate synthase was also involved in the synthesis of salicylic acid, and whether this pathway played an important role in rice defense responses. Also, little is known about functional analysis of OsHPL3, whose product competing the same substrate with jasmonic acid, against WBPH resistance. Therefore, we isolated two genes as the research object, one is isochorismate synthase gene OsICS, induced by brown planthopper feeding, and the other is hydroperoxide lyase gene OsHPL3, whose production green leaves volatile, combining with the methods of reverse genetics, molecular biology, chemical analysis and bioassay, in order to clarify OsICS function in rice herbivore induced resistance and OsHPL3against WBPH resistance defense responses. The main results are as follows:
A2445bp open reading frame of an herbivore specifically induced isochorismate synthase gene OsICS, which encodes a protein of814amino acids with a calculated molecular weight of88.024KDa and pI5.92, was cloned. OsICS coding protein is located in the chloroplasts. Tissue specific expression analysis revealed OsICS transcript levels in rice leaves was higher than that of other parts. Induced expression analysis showed that the feeding of Nilaparvata lugens (brown planthopper, BPH), Sogatella furrifera (white back planthopper, WBPH), Chilo suppressalis (rice striped stem borer, SSB) and mechanical wounding treatment can induce OsICS gene expression, and exogenous jasmonic acid (JA) and salicylic acid (SA) can not induce the expression, but have an opposite effect. Through agrobacterium-mediated transgenic method, we obtained the OsICS mutant plants. The OsICS expression levels in transgenic lines were decreased by46.23%-50.86%than the wild type. Phenotypic results of different lines revealed OsICS transgenic mutants are significantly shorter than the wild type during germination and grown in the field. Besides, the level of SA in transgenic mutants are significantly lower than the wild type both grown in the field and green house induced by the feeding of brown planthopper and rice stem borer. The bioassay of different pests results show that, the OsICS transgenic lines could obviously increase the feeding choice of BPH and WBPH, and the treatment of exogenous SA can restore BPH and WBPH feeding choice. However, the OsICS transgenic lines can not affect herbivore induced rice JA, trypsin inhibitor (TrypPIs) and ethylene (ET). And the transgenic lines has no significant effect on the nymph developmental duration and population growth of BPH and WBPH.
OsHPL3is a hydroperoxide lyase gene. Expression levels of OsHPL3could be significantly enhanced by mechanical wounding, SSB, BPH and WBPH infestation, whereas exogenous hormones JA and SA treatment had no significant effect. We use the same method to aquire the OsHPL3transgenic lines. The emission of GLV levels (Z)-3-hexenal,(Z)-3-Hexen-l-ol and (E)-2-hexenal were significantly lower in transgenic lines than in wild type rice plants after WBPH infestation and mechanical wounding treatment. At the same time, the transgenic lines can increase the resistance to WBPH. Bioassay results show that adult WBPH preferred to feed and oviposite on the transgenic lines. The nymph developmental duration in the transgenic lines were significantly longer, the survival rate was significantly decreased, and the number of eggs per day were significantly lower than in the wild type. While the treatment of exogenous (Z)-3-hexenal can restore the0sHPL3transgenic lines to the resistance of WBPH. The result of Chromatograms of volatiles obtained from various lines show that transgenic strains can increase the release of limonene after WBPH damage, while the decrease expression of OsHPL3in transgenic lines had no effect on other WBPH-induced plant volatiles. Compared with transgenic plants, Anagrus nilaparvatae, which is an egg parasitoid of WBPH, preferred the odors from wild type plants, releasing more GLV after WBPH infestation. However, exogenous different concentrations of GLV,(Z)-3-hexenal,(Z)-3-Hexen-l-ol and (E)-2-hexenal treatment can restore the Anagrus nilaparvatae selectively behavior between different lines.So GLV, the product of OsHPL3, play an important role in the rice resistance to WBPH.
引文
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