油菜素内酯促进番茄体内百菌清降解中谷胱甘肽相关解毒途径的机理研究
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摘要
农药的过量及不合理使用不仅对作物生长和品质造成了严重的影响,也造成了蔬菜和环境中农药残留的超标,对食品安全和生态环境已经构成了严重的威胁。随着经济的发展和生活水平的提高,人们对食品安全和生存环境问题越来越重视,因此,对农产品中的农药残留进行控制,使其降低至安全标准以下,已成为迫切需要解决的问题。已有研究表明,油菜素内酯BRs能够有效地降低植物体内的农药残留,但关于BRs促进植物体内农药降解代谢的机理还有待于进一步研究,对其机理的清晰了解是建立一条农作物安全生产调控新途径的必要前提,对于保证农产品安全和人类健康具有重大的意义。本文以番茄(Solanum lycopersicum Mill.)为材料,以生产中普遍使用的百菌清为目标化合物,利用基因沉默技术,针对谷胱甘肽相关解毒途径在植物体内农药降解代谢中的作用机理,以及在BRs促进植物体内农药降解过程中所起到的作用进行研究,并且对BRs通过诱导H2O2信号,启动植物对农药的解毒响应,调控植物体内解毒代谢的作用机理进行了探讨,以进一步阐述BRs促进植物体内农药降解代谢的相关机制。
所得主要结果如下：
1.建立并优化了基于TRV病毒诱导的基因沉默体系,获得了谷胱甘肽相关基因沉默的番茄植株材料。本研究利用VIGS技术,构建了目标基因的TRV病毒载体,获得了SIGSH1、SlGSH2、SlGR以及7个SlGST基因有效沉默的植株材料,并利用RT-PCR验证其沉默效率,结果表明,基因沉默对植株目标基因的抑制效率均高于一半以上,说明基于TRV的VIGS技术能有效诱导目标基因在番茄植株中发生系统性沉默。
2.研究了谷胱甘肽在植物体内百菌清解毒代谢中的作用机理。利用VIGS技术,对谷胱甘肽合成酶以及谷胱甘肽还原酶的编码基因GSH1, GSH2, GR分别进行沉默,研究发现谷胱甘肽合成与再生相关的基因沉默有效地降低了植株谷胱甘肽含量及GSH/GSSG比值,并且显著地抑制了番茄体内百菌清的降解代谢,尤其是GSH1和GR基因沉默的效果更为显著,降低了解毒反应底物谷胱甘肽的含量,或是改变了氧化还原状态均能显著地抑制了百菌清的降解代谢,同时,也抑制了植物体内农药解毒代谢相关的基因表达以及GST等解毒酶活力。说明谷胱甘肽不仅作为底物直接参与百菌清代谢反应,还能够通过Redox信号作用等诱导解毒基因表达,激活GST等解毒酶活性,促进植物体对农药的谷胱甘肽等解毒代谢反应的进行,从而有效地降低了植物体内农药残留量,在植物活体内证实了谷胱甘肽相关途径对体内农药降解代谢是至关重要的。
3.研究了谷胱甘肽在油菜素内酯促进番茄体内百菌清降解代谢中的作用。研究发现BRs能够促进植物体内谷胱甘肽等相关物质的合成,启动以谷胱甘肽为底物的轭合反应,BRs可以通过调控谷胱甘肽相关物质代谢而促进植物体内的农药代谢。另外,BRs也能够诱导植物体三相解毒代谢相关基因的表达,以及GST等解毒酶活力的提高,从而加速植物体谷胱甘肽对百菌清进行的轭合反应等解毒作用,促进百菌清在植物体内的降解代谢。
4.研究了H202在油菜素内酯促进番茄体内农药降解中的信号作用。采用外源化学试剂抑制内源H202的爆发,以及基因沉默方法调控内源BRIl及RBOH等基因的表达,对BRs促进植物体内农药代谢的作用机理以及信号调控进行研究,从不同角度证明了H202信号能够响应外源的EBR处理,并参与BRs介导的番茄体内农药的谷胱甘肽解毒代谢途径。BRs通过诱导RBOH基因的表达,调控NADPH氧化酶产生H202信号,以介导解毒反应底物谷胱甘肽的合成,启动相关的生理生化反应参与番茄对体内百菌清的轭合代谢,从而促进番茄体内百菌清的降解代谢。
5.研究了谷胱甘肽S-转移酶在BRs促进番茄体内百菌清降解中的作用。我们利用VIGS技术,针对已报道的番茄体内编码GST酶的7个基因GST1、GST2、 GST3、GST4、GST5、GST6、GST7分别进行沉默,研究发现各目标基因的沉默均显著地抑制了植株中其基因的转录水平,但部分基因的沉默对GST酶活性的抑制效果不明显。其中,GSTl与GST7基因沉默有效地抑制了GST酶活性,同时,对番茄体内百菌清代谢也具有显著的抑制作用,证明其编码的GST酶参与了番茄体内百菌清的解毒代谢,但GSTl基因沉默植株仍对EBR处理具有显著的响应。因此,BR能够通过诱导GST7等部分GST酶编码基因的有效表达,提高番茄GST酶活力,加速植物体对百菌清解毒反应,促进植物体内百菌清的降解代谢。
Pesticides are widely used in agriculture for its high efficacy in pathogens control, however, the excessive use of pesticide results in pollution to environment and agricultural products, which directly endangers human health. Therefore, it has become a serious problem to be solved in order to reduce pesticide residue following safety standards. Until now, extensive researches have focused on bioremediation of organic pollutants using enzymes produced from the microorganisms or plants. By contrast, there are only few studies on the metabolism or biotransformation of pesticides in plants, especially on the degradation of fungicides. Recent study shows that brassinosteroids (BRs) can reduce pesticides residue and promote pesticides detoxification in plant. However, the role and mechanism of BR promoting pesticide degradation is still not clear. Therefore, the present study was conducted to investigate the mechanism of BR promoting biotransformation and degradation of CHT in plant.
In this study, we investigated the mechanism of chlorothalonil (CHT) degradation in tomato (Solanum lycopersicum Mill.) plants involved in glutathione biosynthesis, regeneration and metabolism by VIGS. We also investigated the role and mechanism of brassinosteroids (BRs) alleviating pesticides phytotoxicity and promoting pesticide metabolism in glutathione-dependent gene silencing plants to understand the role of glutathione-dependent detoxification in BRs promoting pesticides degradation in plant. Meanwhile, we have investigated the role of H2O2in BRs promoting pesticides degradation to further clarify the mechanism of BRs promoting pesticides metabolism. The results are as follows:
1. In this study, tobacco rattle virus (TRV) and tomato plants(Solanum lycopersicum Mill. cv. Zheza205) were used to construct the gene silencing vector with the fragment of SlGSH1、SlGSH2、SlGR and seven SIGST genes. VIGS transform system of SIPDS reference gene and other genes were established. Quantitative RT-PCR analysis revealed that transcript levels of genes in silenced plants were decreased significantly, compared with TRV plants, which suggested an effective inhibition of target genes by VIGS.
2. To determine the role of glutathione biosynthesis and regeneration in the CHT metabolism, GSH1, GSH2and GR genes were silenced with VIGS. Silencing of GSH1, GSH2and GR1all resulted in decreased glutathione synthesis and ratio of reduced glutathione (GSH) to oxidized glutathione (GSSG), but increased CHT residues in plant tissues, while the GSH1-silenced plants and GR-silenced plants showed the lowest GSH levels and lowest ratio of GSH to GSSG, respectively. CHT exposure study indicated that glutathione-dependent pathway was involved in CHT detoxification and glutathione not only reacted as a substrate to conjugate with CHT, but also induced activities of detoxification enzymes, as well as expression of detoxification genes directly or indirectly. These observations provided evidences for the involvement of the glutathione-dependent pathway in the in vivo metabolism of pesticide in tomato plants.
3. Studies were conducted to understand the role of glutathione-dependent detoxification pathway in BRs promoting pesticides degradation in plant. The results demonstrated that BRs could increase glutathione synthesis and NPT, TT levels, and induced the detoxification metabolism reacts, in which GSH conjugated with CHT as a substrate. Meanwhile, BRs induced activities of detoxification enzymes, as well as the expression of detoxification genes directly or indirectly. Consequently, BRs could promote the CHT metabolism processes and finally reduces CHT residue in plant via glutathione-dependent detoxification metabolism.
4. The role of H2O2in BRs promoting pesticides degradation was also investigated to further clarify the mechanism of BRs promoting pesticides metabolism. We inhibited ROS burst in plant with the application of chemical inhibitors, and also regulated gene expression of BRI1and RBOH genes by VIGS approach to investigate the signal role of H2O2in BRs promoting pesticides metabolism. The studies confirmed that H2O2signal could be in response to EBR treatment, and induced glutathione-dependent detoxification pathway in BRs promoting pesticides degradation in plant. BRs induced the gene expression of RBOH, resulted in the production of H2O2signal by NADPH oxidase to regulate glutathione-dependent detoxification pathway to promote pesticide metabolism in plant. H2O2played a critical role in BR-induced glutathione-dependent detoxification to pesticide in plant.
5. The effects of GST in BRs promoting pesticides metabolism in plant were investigated. GST1、GST2、GST3、GST4、GST5、GST6、GST7genes that encode glutathione S-transferase in tomato, were silenced respectively in tomato plants by a virus induced gene silencing (VIGS) approach. The studies revealed that gene transcripts of seven GST genes were all significantly inhibited by VIGS, however, the inhibition in GST activities were different. GST1and GST7genes silencing inhibited both GST activity and CHT degradation metabolism in plants significantly, which demonstrated the participation of GST encoded by GST1and GST7genes in the detoxification of CHT in plant. However, GST1gene silenced plants could even be induced by EBR treatment significantly, while GST7gene silenced plants showed no response to EBR treatment, which demonstrated that BRs could induce the expression of some GST encoding genes, such as GST7, to increase GST activities to promote pesticides degradation metabolism.
所得主要结果如下：
1.建立并优化了基于TRV病毒诱导的基因沉默体系,获得了谷胱甘肽相关基因沉默的番茄植株材料。本研究利用VIGS技术,构建了目标基因的TRV病毒载体,获得了SIGSH1、SlGSH2、SlGR以及7个SlGST基因有效沉默的植株材料,并利用RT-PCR验证其沉默效率,结果表明,基因沉默对植株目标基因的抑制效率均高于一半以上,说明基于TRV的VIGS技术能有效诱导目标基因在番茄植株中发生系统性沉默。
2.研究了谷胱甘肽在植物体内百菌清解毒代谢中的作用机理。利用VIGS技术,对谷胱甘肽合成酶以及谷胱甘肽还原酶的编码基因GSH1, GSH2, GR分别进行沉默,研究发现谷胱甘肽合成与再生相关的基因沉默有效地降低了植株谷胱甘肽含量及GSH/GSSG比值,并且显著地抑制了番茄体内百菌清的降解代谢,尤其是GSH1和GR基因沉默的效果更为显著,降低了解毒反应底物谷胱甘肽的含量,或是改变了氧化还原状态均能显著地抑制了百菌清的降解代谢,同时,也抑制了植物体内农药解毒代谢相关的基因表达以及GST等解毒酶活力。说明谷胱甘肽不仅作为底物直接参与百菌清代谢反应,还能够通过Redox信号作用等诱导解毒基因表达,激活GST等解毒酶活性,促进植物体对农药的谷胱甘肽等解毒代谢反应的进行,从而有效地降低了植物体内农药残留量,在植物活体内证实了谷胱甘肽相关途径对体内农药降解代谢是至关重要的。
3.研究了谷胱甘肽在油菜素内酯促进番茄体内百菌清降解代谢中的作用。研究发现BRs能够促进植物体内谷胱甘肽等相关物质的合成,启动以谷胱甘肽为底物的轭合反应,BRs可以通过调控谷胱甘肽相关物质代谢而促进植物体内的农药代谢。另外,BRs也能够诱导植物体三相解毒代谢相关基因的表达,以及GST等解毒酶活力的提高,从而加速植物体谷胱甘肽对百菌清进行的轭合反应等解毒作用,促进百菌清在植物体内的降解代谢。
4.研究了H202在油菜素内酯促进番茄体内农药降解中的信号作用。采用外源化学试剂抑制内源H202的爆发,以及基因沉默方法调控内源BRIl及RBOH等基因的表达,对BRs促进植物体内农药代谢的作用机理以及信号调控进行研究,从不同角度证明了H202信号能够响应外源的EBR处理,并参与BRs介导的番茄体内农药的谷胱甘肽解毒代谢途径。BRs通过诱导RBOH基因的表达,调控NADPH氧化酶产生H202信号,以介导解毒反应底物谷胱甘肽的合成,启动相关的生理生化反应参与番茄对体内百菌清的轭合代谢,从而促进番茄体内百菌清的降解代谢。
5.研究了谷胱甘肽S-转移酶在BRs促进番茄体内百菌清降解中的作用。我们利用VIGS技术,针对已报道的番茄体内编码GST酶的7个基因GST1、GST2、 GST3、GST4、GST5、GST6、GST7分别进行沉默,研究发现各目标基因的沉默均显著地抑制了植株中其基因的转录水平,但部分基因的沉默对GST酶活性的抑制效果不明显。其中,GSTl与GST7基因沉默有效地抑制了GST酶活性,同时,对番茄体内百菌清代谢也具有显著的抑制作用,证明其编码的GST酶参与了番茄体内百菌清的解毒代谢,但GSTl基因沉默植株仍对EBR处理具有显著的响应。因此,BR能够通过诱导GST7等部分GST酶编码基因的有效表达,提高番茄GST酶活力,加速植物体对百菌清解毒反应,促进植物体内百菌清的降解代谢。
Pesticides are widely used in agriculture for its high efficacy in pathogens control, however, the excessive use of pesticide results in pollution to environment and agricultural products, which directly endangers human health. Therefore, it has become a serious problem to be solved in order to reduce pesticide residue following safety standards. Until now, extensive researches have focused on bioremediation of organic pollutants using enzymes produced from the microorganisms or plants. By contrast, there are only few studies on the metabolism or biotransformation of pesticides in plants, especially on the degradation of fungicides. Recent study shows that brassinosteroids (BRs) can reduce pesticides residue and promote pesticides detoxification in plant. However, the role and mechanism of BR promoting pesticide degradation is still not clear. Therefore, the present study was conducted to investigate the mechanism of BR promoting biotransformation and degradation of CHT in plant.
In this study, we investigated the mechanism of chlorothalonil (CHT) degradation in tomato (Solanum lycopersicum Mill.) plants involved in glutathione biosynthesis, regeneration and metabolism by VIGS. We also investigated the role and mechanism of brassinosteroids (BRs) alleviating pesticides phytotoxicity and promoting pesticide metabolism in glutathione-dependent gene silencing plants to understand the role of glutathione-dependent detoxification in BRs promoting pesticides degradation in plant. Meanwhile, we have investigated the role of H2O2in BRs promoting pesticides degradation to further clarify the mechanism of BRs promoting pesticides metabolism. The results are as follows:
1. In this study, tobacco rattle virus (TRV) and tomato plants(Solanum lycopersicum Mill. cv. Zheza205) were used to construct the gene silencing vector with the fragment of SlGSH1、SlGSH2、SlGR and seven SIGST genes. VIGS transform system of SIPDS reference gene and other genes were established. Quantitative RT-PCR analysis revealed that transcript levels of genes in silenced plants were decreased significantly, compared with TRV plants, which suggested an effective inhibition of target genes by VIGS.
2. To determine the role of glutathione biosynthesis and regeneration in the CHT metabolism, GSH1, GSH2and GR genes were silenced with VIGS. Silencing of GSH1, GSH2and GR1all resulted in decreased glutathione synthesis and ratio of reduced glutathione (GSH) to oxidized glutathione (GSSG), but increased CHT residues in plant tissues, while the GSH1-silenced plants and GR-silenced plants showed the lowest GSH levels and lowest ratio of GSH to GSSG, respectively. CHT exposure study indicated that glutathione-dependent pathway was involved in CHT detoxification and glutathione not only reacted as a substrate to conjugate with CHT, but also induced activities of detoxification enzymes, as well as expression of detoxification genes directly or indirectly. These observations provided evidences for the involvement of the glutathione-dependent pathway in the in vivo metabolism of pesticide in tomato plants.
3. Studies were conducted to understand the role of glutathione-dependent detoxification pathway in BRs promoting pesticides degradation in plant. The results demonstrated that BRs could increase glutathione synthesis and NPT, TT levels, and induced the detoxification metabolism reacts, in which GSH conjugated with CHT as a substrate. Meanwhile, BRs induced activities of detoxification enzymes, as well as the expression of detoxification genes directly or indirectly. Consequently, BRs could promote the CHT metabolism processes and finally reduces CHT residue in plant via glutathione-dependent detoxification metabolism.
4. The role of H2O2in BRs promoting pesticides degradation was also investigated to further clarify the mechanism of BRs promoting pesticides metabolism. We inhibited ROS burst in plant with the application of chemical inhibitors, and also regulated gene expression of BRI1and RBOH genes by VIGS approach to investigate the signal role of H2O2in BRs promoting pesticides metabolism. The studies confirmed that H2O2signal could be in response to EBR treatment, and induced glutathione-dependent detoxification pathway in BRs promoting pesticides degradation in plant. BRs induced the gene expression of RBOH, resulted in the production of H2O2signal by NADPH oxidase to regulate glutathione-dependent detoxification pathway to promote pesticide metabolism in plant. H2O2played a critical role in BR-induced glutathione-dependent detoxification to pesticide in plant.
5. The effects of GST in BRs promoting pesticides metabolism in plant were investigated. GST1、GST2、GST3、GST4、GST5、GST6、GST7genes that encode glutathione S-transferase in tomato, were silenced respectively in tomato plants by a virus induced gene silencing (VIGS) approach. The studies revealed that gene transcripts of seven GST genes were all significantly inhibited by VIGS, however, the inhibition in GST activities were different. GST1and GST7genes silencing inhibited both GST activity and CHT degradation metabolism in plants significantly, which demonstrated the participation of GST encoded by GST1and GST7genes in the detoxification of CHT in plant. However, GST1gene silenced plants could even be induced by EBR treatment significantly, while GST7gene silenced plants showed no response to EBR treatment, which demonstrated that BRs could induce the expression of some GST encoding genes, such as GST7, to increase GST activities to promote pesticides degradation metabolism.
引文
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