植物次生代谢在抗氧化胁迫和有机污染物毒害中的作用及调控机制
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摘要
设施栽培业已成为我国现代农业中的重要组成部分,但是由于农作物生长发育对环境条件的较高要求以及农业基础设施、生产管理水平的相对落后,导致农作物的产量和品质常常受到低温、高温、干旱、盐害等非生物逆境和病虫害等生物逆境的影响。次生代谢是植物在长期进化中与环境相互作用的结果,次生代谢产物在植物提高自身保护和生存竞争能力、协调与环境关系上充当着重要的角色。因此,研究次生代谢的合成途径以及其与环境胁迫的关系,探索一条植物的抗性、产量及品质安全调控的新途径对于建立可持续农业具有非常重要的理论和实践意义。本文先以拟南芥(Arabidopsis thaliana)野生型和pall pal2突变体为材料,研究了拟南芥PAL1和PAL2双基因缺失突变对次生代谢和氧化胁迫的影响;又以黄瓜(Cucumis sativus L.)为材料,研究了叶面喷施24-表油菜素内酯(EBR)对黄瓜次生代谢的影响以及叶面喷施EBR缓解黄瓜有机污染物毒害的生理与分子机理研究。所取得的主要研究结果如下:
1.拟南芥PAL1和PAL2双基因缺失突变能够抑制苯丙烷类代谢途径的进程从而降低对氧化胁迫的抗性。结果表明,缺失PAL1和PAL2基因能有效改变拟南芥植株表观形态和生物量,且pall pal2突变体的苯丙氨酸解氨酶(PAL)活性显著下降。同时,PAL1和PAL2双基因缺失突变阻碍了总酚、类黄酮和木质素的合成,显著降低DPPH自由基清除活性和CAD、G-POD、CA-POD、CGA-POD、PPO活力。当pall pal2突变体受到氧化胁迫后叶绿素荧光参数显著降低,说明由于PAL1和PAL2基因的存在能够缓解百草枯胁迫、紫外辐射胁迫对植株造成的伤害。因此,PAL通过调控次生代谢对植物抗逆反应具有积极作用。
2.叶面喷施EBR能够促进黄瓜叶片和根系次生代谢的进程。结果表明,EBR处理后短时间内即诱导黄瓜总酚、类黄酮、木质素含量以及DPPH自由基清除活性的上升,直至2 d后次生代谢物质含量显著下降,然后随着时间的延长恢复到正常水平。同时,EBR处理使黄瓜体内的次生代谢相关酶活性呈现出类似的周期性变化。然后,选择次生代谢变化幅度最明显的时间点进行测定,肉桂酸、对香豆酸、咖啡酸、阿魏酸的变化规律与总酚、类黄酮、木质素一致,而次生代谢相关基因的表达量也与次生代谢相关酶活性一致。值得注意的是,叶面喷施EBR导致黄瓜叶片中的次生代谢变化均早于根系。因此,BR可能是通过可移动运输的物质作为信号从而促进植株整体的次生代谢。
3.EBR能够加速有机污染物在黄瓜体内的代谢从而缓解毒害。首先,确定了三种具有代表性的有机污染物的实验使用浓度,三氯苯酚(属于氯代酚类化合物)为5μM,毒死蜱(属于农药)为10μM,土霉素(属于抗生素)为50μM。结果表明,有机污染物能够明显抑制黄瓜的地上部、地下部生长量,而不同浓度的EBR预处理都显著缓解有机污染物对生长量的受抑现象,以0.1μM缓解效果最佳。然后,对抗氧化能力、谷胱甘肽S-转移酶和次生代谢进行测定,EBR预处理显著降低膜质过氧化(MDA)程度,明显提高有机污染物处理后SOD、CAT、APX的活性和基因表达量;EBR显著提高GSH+GSSG、GSH的含量以及GST、GR的活性和基因表达量,调节GSH/GSSG氧化还原状态;EBR促进有机污染物处理后次生代谢物质的积累,显著提高PAL、PPO、G-POD的活性和基因表达量。因此,BR作为环境友好的天然化合物在解决有机环境污染的问题上有广阔的应用前景。
Greenhouse horticulture is an improtant integrative part of modern agriculture in China.The growth and development of crops are dependent on the suitable environment conditions.However,the productivity and quality of crops are always negatively affected by abiotic stresses(such as chill,high temperature,drought,and salinity) and biotic stresses(pathogen infection,insect attack) due to the relatively poor basic facility and management.Secondary metabolism is a consequence of the interaction between plants and environments in the long process of plant evolution. Secondary metabolites play important roles for plants to improve defensive and competitive abilities as well as correspond to the environments.Therefore,it is of great importance to investigate the synthetic way of secondary metabolism and its response to environment stresses and explore the effective strategy to enhance the tolerance of plants and improve the productivity,quality and safety of agricultural produces.This study plays theoretical and practical roles in the development of sustainable agriculture.To begin with,wild type and pal1 pal2 mutants of Arabidopsis thaliana were used as the material to investigate the effects of PAL1 and PAL2 on secondary metabolism and oxidative stresses.And then cucumber(Cucumis sativus L.) was used as the material to study the effect of 24-epibrassinolide(EBR) on secondary metabolism and analyze physiological and molecular mechanism of EBR on tolerance to toxicities of organic pollutants.The main results are as follows:
1.Tolerance to oxidative stresses in pal1 pal2 mutants decreased due to the inhibition of the phenylpropanoid pathway.First the lack of PAL1 and PAL2 resulted in effective changes in plant morphology and biomass,and these effects were associated with the declines of phenylalanine ammonia-lyase(PAL) activity,phenols, flavonoids,lignin and DPPH activity.Moreover,the activities of CAD,G-POD, CA-POD,CGA-POD and PPO were inhibited in pal1 pal2 mutant leaves.Since oxidative stresses induced reductions of chlorophyll fluorescence parameters in pal1 pal2 mutants,it was found that PAL1 and PAL2 played important roles in protecting Arabidopsis from paraquat stress and UV-B stress.The results demonstrated that PAL may be directly involved in plant stress responses through regulating secondary metabolism.
2.EBR had a positive effect on the induction of secondary metabolism in cucumber leaves and roots.Levels of phenols,flavonoids,lignin,DPPH activity and enzyme activities related to secondary metabolism were elevated in a short time and reduced at 2 d and returned to basal levels gradually after EBR treatment.We selected the timing of the most remarkable changes in cucumber secondary metabolism, meanwhile,it was found that EBR played the similar roles in regulating the contents of cinnamic acid,ρ-coumaric acid,caffeic acid and ferulic acid.The change of gene expression was closely associated with enzyme activities related to secondary metabolism.In addition,EBR-induced changes were earlier in treated leaves than in roots significantly.These results provided evidence that BRs may be involved in promoting the whole plant secondary metabolism by a novel long-distance signal transmitted from shoots to roots.
3.EBR protected cucumber from toxicities of organic pollutants through accelerating the whole metabolism.First of all,cucumber seedlings were supplied with three kinds of typical organic pollutants with different concentrations.It was shown that plant shoot and root growth rate decreased accompanied by increasing concentrations,moreover,we selected experimental concentrations of 2,4,6-trichlorophenol(TCP,5μM),chlorpyrifos(CPF,10μM) and oxytetracycline (OTC,50μM) respectively.EBR at different concentrations had a positive effect on tolerance to toxicities of organic pollutants and 0.1μM EBR was most effective.In addition to its roles in promoting glutathione metabolism and secondary metabolism, EBR application alleviated the increase of MDA and regulated the redox ratio of GSH/GSSG after treatment with TCP,CPF or OTC.EBR induced enzyme activities (such as SOD,CAT,APX,GST,GR,PAL,PPO,and G-POD) and gene expression. The results suggested that BRs may be promising,environmentally friendly,natural substances suitable for wide application to reduce the risks of human and environment exposure to organic pollutants.
1.拟南芥PAL1和PAL2双基因缺失突变能够抑制苯丙烷类代谢途径的进程从而降低对氧化胁迫的抗性。结果表明,缺失PAL1和PAL2基因能有效改变拟南芥植株表观形态和生物量,且pall pal2突变体的苯丙氨酸解氨酶(PAL)活性显著下降。同时,PAL1和PAL2双基因缺失突变阻碍了总酚、类黄酮和木质素的合成,显著降低DPPH自由基清除活性和CAD、G-POD、CA-POD、CGA-POD、PPO活力。当pall pal2突变体受到氧化胁迫后叶绿素荧光参数显著降低,说明由于PAL1和PAL2基因的存在能够缓解百草枯胁迫、紫外辐射胁迫对植株造成的伤害。因此,PAL通过调控次生代谢对植物抗逆反应具有积极作用。
2.叶面喷施EBR能够促进黄瓜叶片和根系次生代谢的进程。结果表明,EBR处理后短时间内即诱导黄瓜总酚、类黄酮、木质素含量以及DPPH自由基清除活性的上升,直至2 d后次生代谢物质含量显著下降,然后随着时间的延长恢复到正常水平。同时,EBR处理使黄瓜体内的次生代谢相关酶活性呈现出类似的周期性变化。然后,选择次生代谢变化幅度最明显的时间点进行测定,肉桂酸、对香豆酸、咖啡酸、阿魏酸的变化规律与总酚、类黄酮、木质素一致,而次生代谢相关基因的表达量也与次生代谢相关酶活性一致。值得注意的是,叶面喷施EBR导致黄瓜叶片中的次生代谢变化均早于根系。因此,BR可能是通过可移动运输的物质作为信号从而促进植株整体的次生代谢。
3.EBR能够加速有机污染物在黄瓜体内的代谢从而缓解毒害。首先,确定了三种具有代表性的有机污染物的实验使用浓度,三氯苯酚(属于氯代酚类化合物)为5μM,毒死蜱(属于农药)为10μM,土霉素(属于抗生素)为50μM。结果表明,有机污染物能够明显抑制黄瓜的地上部、地下部生长量,而不同浓度的EBR预处理都显著缓解有机污染物对生长量的受抑现象,以0.1μM缓解效果最佳。然后,对抗氧化能力、谷胱甘肽S-转移酶和次生代谢进行测定,EBR预处理显著降低膜质过氧化(MDA)程度,明显提高有机污染物处理后SOD、CAT、APX的活性和基因表达量;EBR显著提高GSH+GSSG、GSH的含量以及GST、GR的活性和基因表达量,调节GSH/GSSG氧化还原状态;EBR促进有机污染物处理后次生代谢物质的积累,显著提高PAL、PPO、G-POD的活性和基因表达量。因此,BR作为环境友好的天然化合物在解决有机环境污染的问题上有广阔的应用前景。
Greenhouse horticulture is an improtant integrative part of modern agriculture in China.The growth and development of crops are dependent on the suitable environment conditions.However,the productivity and quality of crops are always negatively affected by abiotic stresses(such as chill,high temperature,drought,and salinity) and biotic stresses(pathogen infection,insect attack) due to the relatively poor basic facility and management.Secondary metabolism is a consequence of the interaction between plants and environments in the long process of plant evolution. Secondary metabolites play important roles for plants to improve defensive and competitive abilities as well as correspond to the environments.Therefore,it is of great importance to investigate the synthetic way of secondary metabolism and its response to environment stresses and explore the effective strategy to enhance the tolerance of plants and improve the productivity,quality and safety of agricultural produces.This study plays theoretical and practical roles in the development of sustainable agriculture.To begin with,wild type and pal1 pal2 mutants of Arabidopsis thaliana were used as the material to investigate the effects of PAL1 and PAL2 on secondary metabolism and oxidative stresses.And then cucumber(Cucumis sativus L.) was used as the material to study the effect of 24-epibrassinolide(EBR) on secondary metabolism and analyze physiological and molecular mechanism of EBR on tolerance to toxicities of organic pollutants.The main results are as follows:
1.Tolerance to oxidative stresses in pal1 pal2 mutants decreased due to the inhibition of the phenylpropanoid pathway.First the lack of PAL1 and PAL2 resulted in effective changes in plant morphology and biomass,and these effects were associated with the declines of phenylalanine ammonia-lyase(PAL) activity,phenols, flavonoids,lignin and DPPH activity.Moreover,the activities of CAD,G-POD, CA-POD,CGA-POD and PPO were inhibited in pal1 pal2 mutant leaves.Since oxidative stresses induced reductions of chlorophyll fluorescence parameters in pal1 pal2 mutants,it was found that PAL1 and PAL2 played important roles in protecting Arabidopsis from paraquat stress and UV-B stress.The results demonstrated that PAL may be directly involved in plant stress responses through regulating secondary metabolism.
2.EBR had a positive effect on the induction of secondary metabolism in cucumber leaves and roots.Levels of phenols,flavonoids,lignin,DPPH activity and enzyme activities related to secondary metabolism were elevated in a short time and reduced at 2 d and returned to basal levels gradually after EBR treatment.We selected the timing of the most remarkable changes in cucumber secondary metabolism, meanwhile,it was found that EBR played the similar roles in regulating the contents of cinnamic acid,ρ-coumaric acid,caffeic acid and ferulic acid.The change of gene expression was closely associated with enzyme activities related to secondary metabolism.In addition,EBR-induced changes were earlier in treated leaves than in roots significantly.These results provided evidence that BRs may be involved in promoting the whole plant secondary metabolism by a novel long-distance signal transmitted from shoots to roots.
3.EBR protected cucumber from toxicities of organic pollutants through accelerating the whole metabolism.First of all,cucumber seedlings were supplied with three kinds of typical organic pollutants with different concentrations.It was shown that plant shoot and root growth rate decreased accompanied by increasing concentrations,moreover,we selected experimental concentrations of 2,4,6-trichlorophenol(TCP,5μM),chlorpyrifos(CPF,10μM) and oxytetracycline (OTC,50μM) respectively.EBR at different concentrations had a positive effect on tolerance to toxicities of organic pollutants and 0.1μM EBR was most effective.In addition to its roles in promoting glutathione metabolism and secondary metabolism, EBR application alleviated the increase of MDA and regulated the redox ratio of GSH/GSSG after treatment with TCP,CPF or OTC.EBR induced enzyme activities (such as SOD,CAT,APX,GST,GR,PAL,PPO,and G-POD) and gene expression. The results suggested that BRs may be promising,environmentally friendly,natural substances suitable for wide application to reduce the risks of human and environment exposure to organic pollutants.
引文
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