水稻对褐飞虱取食应答的蛋白质组学研究
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摘要
褐飞虱(Nilaparvata lugens Stal)是一种典型的植物韧皮部取食昆虫,是水稻(Oryza sativa L.)最严重的害虫之一。在感性水稻上,褐飞虱会造成植株的叶面积、光合速率、氮含量、叶绿素含量乃至干重的减少,同时,褐飞虱产卵量、卵和若虫存活率较高。田间褐飞虱的大量爆发通常导致“火虱烧”现象,即水稻叶和分蘖的枯萎变黄,造成水稻产量的急剧降低。褐飞虱还传播草状丛矮病和齿叶矮缩病等水稻病毒病。而在抗性水稻上,褐飞虱若虫的发育被扰乱,存活率明显比感性水稻上的低,成虫的产卵也被严重抑制。抗性水稻能够抑制褐飞虱种群的增长,降低植株受到的伤害并最终减少褐飞虱取食造成的产量损失。在水稻生产中,推广种植抗性品种是最有效和最环保的控制褐飞虱的方法。
蛋白质组学已成为在蛋白质水平上分析生物体生理变化强有力的工具,但目前应用此技术对水稻褐飞虱取食应答的研究还很少。与高灵敏度质谱技术联用的相对定量蛋白质组学分析,由于其高通量、重复性和灵敏度特点,越来越多地应用于关键生物问题的研究。为了探讨防御相关蛋白在抗感性水稻株系应答褐飞虱过程中的作用,我们利用同位素标记相对和绝对定量(isobaric tag for relative and absolute quantitation, iTRAQ)蛋白质组分析,筛选水稻抗褐飞虱相关的蛋白质。我们提取了褐飞虱取食96小时后的水稻叶鞘蛋白质,结果显示多种蛋白质表达量在褐飞虱取食后发生了显著性变化。经iTRAQ鉴定,这些蛋白包括3个JA合成蛋白质(alpah-DOX, AOS和AOC),7个氧胁迫应答蛋白(CATA, APX2和5个POXs),3个β-葡聚糖酶(Gns1, Gns4和Gns5),3个蛋白激酶(CRK5,CRK6和atypical RLK),1个蛋白网格蛋白,1个甘氨酸分解系统H蛋白,5个光合作用相关蛋白和4个水通道蛋白。其中AOC, CATA,3个POX, Gns1,Gns5,3个蛋白激酶和网格蛋白更多地在感性水稻株系中被诱导。随后,我们用荧光定量PCR分析了8个重要蛋白质基因在抗、感性水稻中受褐飞虱取食后的表达情况。这8个基因的表达趋势与蛋白质相似。蛋白质组和基因表达的结果显示,抗性水稻株系(含抗性基因BPH15)和感性水稻株系中褐飞虱诱导的与伤害胁迫,氧胁迫和病原物胁迫相关的应答反应非常相似。而与胼胝质分解相关的蛋白质和基因在抗性水稻株系中表达水平不受褐飞虱影响,但甘氨酸分解系统H蛋白在抗性水稻株系受褐飞虱取食后表达水平上升了2倍,暗示抗性水稻中存在着高效特异的防御机制。我们认为抗感性水稻针对褐飞虱采取了不同的防御策略,抗性基因BPH15在其中起着决定性的作用。
韧皮部是植物运输、分配光合作用合成的有机代谢产物的重要通道,它也是刺吸式昆虫取食植物的目标组织。除了糖和氨基酸这样的小分子外,高等陆生植物韧皮部汁液中还含有蛋白质。韧皮部汁液中的许多蛋白质参与了植物伤害和防御应答,并可能影响植物与昆虫间的相互作用。我们利用同位素标记的iTRAQ技术分析了感性水稻9311和抗性水稻B5叶鞘韧皮部汁液中的蛋白质。结果表明,韧皮部汁液蛋白质表达量受到了褐飞虱取食的显著影响。一共鉴定了238种韧皮部汁液蛋白质,涉及的功能包括糖代谢,活性氧／氧化还原反应,蛋白质修饰,信号传递等。其中,47种蛋白质在9311和B5之间表达量有着显著不同。在9311中54种蛋白质的含量在褐飞虱取食96小时后发生了显著变化；在B5中40种蛋白质发生了显著变化；11个蛋白质在9311和B5中含量变化的方向相反。我们的研究结果表明,在韧皮部汁液蛋白质水平上9311和B5对褐飞虱的应答是不同的。我们从中选取了5个重要的蛋白质,进行了原位杂交分析,试图从mRNA的水平研究这5个蛋白质在水稻韧皮部的表达模式。其中3个蛋白质在韧皮部检测到了阳性信号。
为了进一步筛选韧皮部特异表达基因,我们采用激光显微切割技术收集了水稻韧皮部细胞和薄壁细胞,提取微量RNA,结合RNA体外线性扩增,获得了用于RT-PCR分析的纯净组织细胞RNA,用于进一步研究RGAP、Serpin、WDS和TCTP这4个基因在水稻韧皮部的表达模式,结果显示它们在韧皮部特异表达。我们对Serpin和Serpin的启动子进行了不同长度5’端缺失分析,以寻找控制维管束特异表达的区段和元件。
The brown planthopper (Nilaparvata lugens Stal, BPH) is a typical vascular feeder and considered one of the most serious pests of rice(Oryza sativa L.). BPH causes reductions in leaf area, photosynthetic rate, leaf and stem nitrogen concentrations, chlorophyll contents and organic dry weight; the infestation ratios, egg numbers, and survival ratios of eggs and nymphs are high on susceptible rice plants. In the field, feeding by large numbers of BPHs usually causes "hopperburn", i.e. drying of the leaves and wilting of the tillers, resulting dramatic reductions in the yield of susceptible rice varieties. The BPH also transmits "grassy stunt" and "ragged" diseases as a virus vector. Nymph development is generally disrupted, oviposition is severely inhibited and the survival rate of nymphs is significantly lower on resistant rice plants than on susceptible plants. Thus population growth is suppressed, and a little or no physiological stress and yield loss are caused to resistant rice variety. For rice crops, growing resistant varieties is the most effective and environment-friendly way to control the BPH.
Proteomics has emerged as a powerful tool for gaining insight into physiological changes at the protein level, but few attempts have been made to apply this technique to study the response of rice to BPH attack. Relative quantification proteomic analysis, using highly sensitivity mass spectrometry techniques, is becoming increasingly popular, due to the high-throughputs, reproducibility and sensitivity it offers. To investigate the role of defense-responsive proteins in susceptible and resistant rice lines against BPH, we used an iTRAQ (isobaric tag for relative and absolute quantitation) proteomic analysis to screen for the pathological levels of proteins in rice. Proteins were extracted from leaf sheath 96 hours after BPH feeding. Several proteins involved in multiple pathways showed significant changes in expression in response to BPH feeding. The iTRAQ results identified them as three JA synthesis proteins (alpha-DOX, AOS, AOC), seven oxidative stress response proteins (CATA, APX2 and five POXs), three P-glucanases (Gns1, Gns4 and Gns5), three protein kinases (CRK5, CRK6 and atypical RLK), clathrin heavy chain protein, glycine cleavage system H protein, five photosynthesis proteins and four aquaporins. Of these proteins, AOC, CATA, three POXs, Gns1, Gns5, three protein kinases and clathrin heavy chain protein were induced more in compatible than in incompatible line. The corresponding genes of eight important proteins were further analyzed by quantitative RT-PCR. Proteomic and transcript responses that were related to wounding, oxidative and pathogen stress overlapped considerably between BPH-resistant (carrying the resistance gene BPH15) and susceptible rice lines. In contrast, proteins and genes related to callose metabolism remained unchanged and glycine cleavage system protein was up-regulated in the BPH resistant lines, indicating that they have an efficient and specific defense mechanism. We proposed that compatible and incompatible rice lines use different defense stratigies to defense BPH and the BPH resistance gene BPH15 is a decisive factor.
The phloem is the route for the translocation and distribution of organic metabolites assimilated during photosynthesis of plant. It is also the target of sucking and piercing insects. In addition to small molecules like sugars and amino acids, phloem sap of higher land plants contains proteins. Many phloem sap proteins have potential roles in wound and defense responses and may influence plant-insect interactions. The mass tagging strategy iTRAQ was applied to compare the phloem sap proteins of leaf sheath of the BPH compatible rice 9311 and the BPH incompatible rice B5. We found that the phloem sap proteins were significantly affected by feeding of BPH. In total,238 phloem sap proteins are identified; 47 showed significant differences in quantity between 9311 and B5 belong to function groups in sugar metabolism, reactive oxygen/redox, protein modification and signal transduct. The amount of 54 proteins changed significantly in 9311 after 96 hrs feeding by BPH and 40 in B5. Eleven of them changed in an opposite direction in the compatible and incompatible varieties. Our results show that 9311 and B5 respond to BPH differently at the protein levels. We further carried out in situ hybridization analysis for five important proteins to verify the expression of their corresponding proteins in rice phloem at the mRNA level. Three of them showed positive signals in the rice sieve elements.
To further screening phloem specific expressed genes, we applied laser capture microdissection to collect the rice phloem cells and parenchyma cells. Their RNA were extracted and though one round of in vitro RNA linear amplification we got sufficient pure phloem/parenchyma RNA for RT-PCR analysis. Four phloem specific expressed genes were identified:RGAP, Serpin, WDS and TCTP. The result showed that they were specific expressed in the phloem. The promoters of RGAP and Serpin were analyzed by constructing different length of 5'end deletion vectors, aiming at identifying domains and motifs that regulate genes of interest specifically expressed in rice phloem.
蛋白质组学已成为在蛋白质水平上分析生物体生理变化强有力的工具,但目前应用此技术对水稻褐飞虱取食应答的研究还很少。与高灵敏度质谱技术联用的相对定量蛋白质组学分析,由于其高通量、重复性和灵敏度特点,越来越多地应用于关键生物问题的研究。为了探讨防御相关蛋白在抗感性水稻株系应答褐飞虱过程中的作用,我们利用同位素标记相对和绝对定量(isobaric tag for relative and absolute quantitation, iTRAQ)蛋白质组分析,筛选水稻抗褐飞虱相关的蛋白质。我们提取了褐飞虱取食96小时后的水稻叶鞘蛋白质,结果显示多种蛋白质表达量在褐飞虱取食后发生了显著性变化。经iTRAQ鉴定,这些蛋白包括3个JA合成蛋白质(alpah-DOX, AOS和AOC),7个氧胁迫应答蛋白(CATA, APX2和5个POXs),3个β-葡聚糖酶(Gns1, Gns4和Gns5),3个蛋白激酶(CRK5,CRK6和atypical RLK),1个蛋白网格蛋白,1个甘氨酸分解系统H蛋白,5个光合作用相关蛋白和4个水通道蛋白。其中AOC, CATA,3个POX, Gns1,Gns5,3个蛋白激酶和网格蛋白更多地在感性水稻株系中被诱导。随后,我们用荧光定量PCR分析了8个重要蛋白质基因在抗、感性水稻中受褐飞虱取食后的表达情况。这8个基因的表达趋势与蛋白质相似。蛋白质组和基因表达的结果显示,抗性水稻株系(含抗性基因BPH15)和感性水稻株系中褐飞虱诱导的与伤害胁迫,氧胁迫和病原物胁迫相关的应答反应非常相似。而与胼胝质分解相关的蛋白质和基因在抗性水稻株系中表达水平不受褐飞虱影响,但甘氨酸分解系统H蛋白在抗性水稻株系受褐飞虱取食后表达水平上升了2倍,暗示抗性水稻中存在着高效特异的防御机制。我们认为抗感性水稻针对褐飞虱采取了不同的防御策略,抗性基因BPH15在其中起着决定性的作用。
韧皮部是植物运输、分配光合作用合成的有机代谢产物的重要通道,它也是刺吸式昆虫取食植物的目标组织。除了糖和氨基酸这样的小分子外,高等陆生植物韧皮部汁液中还含有蛋白质。韧皮部汁液中的许多蛋白质参与了植物伤害和防御应答,并可能影响植物与昆虫间的相互作用。我们利用同位素标记的iTRAQ技术分析了感性水稻9311和抗性水稻B5叶鞘韧皮部汁液中的蛋白质。结果表明,韧皮部汁液蛋白质表达量受到了褐飞虱取食的显著影响。一共鉴定了238种韧皮部汁液蛋白质,涉及的功能包括糖代谢,活性氧／氧化还原反应,蛋白质修饰,信号传递等。其中,47种蛋白质在9311和B5之间表达量有着显著不同。在9311中54种蛋白质的含量在褐飞虱取食96小时后发生了显著变化；在B5中40种蛋白质发生了显著变化；11个蛋白质在9311和B5中含量变化的方向相反。我们的研究结果表明,在韧皮部汁液蛋白质水平上9311和B5对褐飞虱的应答是不同的。我们从中选取了5个重要的蛋白质,进行了原位杂交分析,试图从mRNA的水平研究这5个蛋白质在水稻韧皮部的表达模式。其中3个蛋白质在韧皮部检测到了阳性信号。
为了进一步筛选韧皮部特异表达基因,我们采用激光显微切割技术收集了水稻韧皮部细胞和薄壁细胞,提取微量RNA,结合RNA体外线性扩增,获得了用于RT-PCR分析的纯净组织细胞RNA,用于进一步研究RGAP、Serpin、WDS和TCTP这4个基因在水稻韧皮部的表达模式,结果显示它们在韧皮部特异表达。我们对Serpin和Serpin的启动子进行了不同长度5’端缺失分析,以寻找控制维管束特异表达的区段和元件。
The brown planthopper (Nilaparvata lugens Stal, BPH) is a typical vascular feeder and considered one of the most serious pests of rice(Oryza sativa L.). BPH causes reductions in leaf area, photosynthetic rate, leaf and stem nitrogen concentrations, chlorophyll contents and organic dry weight; the infestation ratios, egg numbers, and survival ratios of eggs and nymphs are high on susceptible rice plants. In the field, feeding by large numbers of BPHs usually causes "hopperburn", i.e. drying of the leaves and wilting of the tillers, resulting dramatic reductions in the yield of susceptible rice varieties. The BPH also transmits "grassy stunt" and "ragged" diseases as a virus vector. Nymph development is generally disrupted, oviposition is severely inhibited and the survival rate of nymphs is significantly lower on resistant rice plants than on susceptible plants. Thus population growth is suppressed, and a little or no physiological stress and yield loss are caused to resistant rice variety. For rice crops, growing resistant varieties is the most effective and environment-friendly way to control the BPH.
Proteomics has emerged as a powerful tool for gaining insight into physiological changes at the protein level, but few attempts have been made to apply this technique to study the response of rice to BPH attack. Relative quantification proteomic analysis, using highly sensitivity mass spectrometry techniques, is becoming increasingly popular, due to the high-throughputs, reproducibility and sensitivity it offers. To investigate the role of defense-responsive proteins in susceptible and resistant rice lines against BPH, we used an iTRAQ (isobaric tag for relative and absolute quantitation) proteomic analysis to screen for the pathological levels of proteins in rice. Proteins were extracted from leaf sheath 96 hours after BPH feeding. Several proteins involved in multiple pathways showed significant changes in expression in response to BPH feeding. The iTRAQ results identified them as three JA synthesis proteins (alpha-DOX, AOS, AOC), seven oxidative stress response proteins (CATA, APX2 and five POXs), three P-glucanases (Gns1, Gns4 and Gns5), three protein kinases (CRK5, CRK6 and atypical RLK), clathrin heavy chain protein, glycine cleavage system H protein, five photosynthesis proteins and four aquaporins. Of these proteins, AOC, CATA, three POXs, Gns1, Gns5, three protein kinases and clathrin heavy chain protein were induced more in compatible than in incompatible line. The corresponding genes of eight important proteins were further analyzed by quantitative RT-PCR. Proteomic and transcript responses that were related to wounding, oxidative and pathogen stress overlapped considerably between BPH-resistant (carrying the resistance gene BPH15) and susceptible rice lines. In contrast, proteins and genes related to callose metabolism remained unchanged and glycine cleavage system protein was up-regulated in the BPH resistant lines, indicating that they have an efficient and specific defense mechanism. We proposed that compatible and incompatible rice lines use different defense stratigies to defense BPH and the BPH resistance gene BPH15 is a decisive factor.
The phloem is the route for the translocation and distribution of organic metabolites assimilated during photosynthesis of plant. It is also the target of sucking and piercing insects. In addition to small molecules like sugars and amino acids, phloem sap of higher land plants contains proteins. Many phloem sap proteins have potential roles in wound and defense responses and may influence plant-insect interactions. The mass tagging strategy iTRAQ was applied to compare the phloem sap proteins of leaf sheath of the BPH compatible rice 9311 and the BPH incompatible rice B5. We found that the phloem sap proteins were significantly affected by feeding of BPH. In total,238 phloem sap proteins are identified; 47 showed significant differences in quantity between 9311 and B5 belong to function groups in sugar metabolism, reactive oxygen/redox, protein modification and signal transduct. The amount of 54 proteins changed significantly in 9311 after 96 hrs feeding by BPH and 40 in B5. Eleven of them changed in an opposite direction in the compatible and incompatible varieties. Our results show that 9311 and B5 respond to BPH differently at the protein levels. We further carried out in situ hybridization analysis for five important proteins to verify the expression of their corresponding proteins in rice phloem at the mRNA level. Three of them showed positive signals in the rice sieve elements.
To further screening phloem specific expressed genes, we applied laser capture microdissection to collect the rice phloem cells and parenchyma cells. Their RNA were extracted and though one round of in vitro RNA linear amplification we got sufficient pure phloem/parenchyma RNA for RT-PCR analysis. Four phloem specific expressed genes were identified:RGAP, Serpin, WDS and TCTP. The result showed that they were specific expressed in the phloem. The promoters of RGAP and Serpin were analyzed by constructing different length of 5'end deletion vectors, aiming at identifying domains and motifs that regulate genes of interest specifically expressed in rice phloem.
引文
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