茉莉酸类化合物:从植物的诱导抗虫防御反应到生长-防御权衡

茉莉酸类化合物(jasmonates, JAs)可外源诱导植物产生抗虫防御反应.因此,JAs在植物生理及植物保护学领域具有十分重要的研究价值.本文归纳了近20年来在约40种植物上外源施用JAs对鳞翅目、半翅目等植食者及其天敌所产生的生态影响,并从JAs诱导植物产生的直接与间接抗虫反应、诱导系统抗性、诱导方法、田间诱导抗虫表现、应用现状等方面对JAs诱导植物抗虫的研究现状进行了系统的总结.同时,本文结合最新研究结果,从植物体内信号交流、JA信号途径节点调控等方面,对植物体内与JA途径相关的生长-防御“权衡”进行了综述.最后,本文对今后外源JAs诱导植物抗虫研究的发展方向与重点方面进行了分析和展望,以期促进相关研究及JAs田间应用的创新发展.     

茉莉酸类化合物在植物防卫反应中的作用

茉莉酸类化合物(jasmonate,JA)是诱导植物防卫反应的重要信号分子。JA不仅是系统素信号转导途径的重要组分,而且在植物长距离伤信号转导中发挥重要作用。JA还能单独或与其他激素相互作用调节与植物防卫反应相关的次生代谢物质芥子油苷的生物合成,从而影响植物的防卫反应。现对JA在植物防卫反应中的作用进行综述,并对今后这一领域的研究进行了展望。     

昆虫对植物蛋白酶抑制素的诱导及适应机制

植物蛋白酶抑制素是植物重要的防御物质之一,一般是分子量较小的多肽或蛋白质,能够与昆虫消化道内的蛋白酶形成复合物,阻断或削弱蛋白酶对食物中蛋白的水解,使昆虫厌食或消化不良而致死。植物蛋白酶抑制素在植物体内一般是诱导表达的,昆虫取食危害后,导致某些植物在伤口产生一种寡聚糖信息素-蛋白酶抑制素诱导因子,蛋白酶抑制素诱导因子诱导叶片局部产生植物蛋白酶抑制素,并刺激产生信号物质系统肽,通过十八烷酸途径在一系列酶的作用下产生茉莉酸,茉莉酸与受体结合,活化植物蛋白酶抑制素基因。昆虫在长期取食植物蛋白酶抑制素后会在生理及行为上产生适应性而导致不敏感,适应方式主要包括：(1)改变肠道蛋白酶对蛋白酶抑制素的敏感性;(2) 水解蛋白酶抑制素;(3)过量取食及干扰产生蛋白酶抑制素的信号通道。由于昆虫能够对植物蛋白酶抑制素产生适应,因此合理利用植物蛋白酶抑制素的抗虫作用显得十分重要。     

系统素、茉莉酸在番茄系统伤反应中的作用

当植物受到机械损伤或昆虫伤害时,植物体会在受伤部位产生伤信号分子启动防御基因的系统表达,蛋白酶抑制剂基因是防御基因的一典型代表.番茄是研究植物系统伤信号很好的模式植物,目前,三种类型的番茄系统伤信号突变体被鉴定出来,通过对番茄系统伤信号突变体进行功能分析并在它们之间进行相互嫁接实验,研究结果表明系统素和茉莉酸通过同一信号通路来激活防御基因的系统表达.系统素(或它的前体原系统素)在受伤部位激活茉莉酸的合成,使之达到系统反应的水平,应对外来伤害;茉莉酸或其衍生物是重要的系统伤信号分子,它诱导伤防御基因的系统表达.植物的系统伤反应可比做动物的炎症反应,它们之间有许多相似之处.     

韧皮部取食昆虫诱导的植物防御反应

刺吸式昆虫与寄主植物之间具有特殊的生物互作关系。本文对刺吸式昆虫取食韧皮部诱导的植物防御反应类型、 防御物质变化、 信号途径以及植物反应转录组学研究等方面进行综述。韧皮部取食昆虫取食诱导的植物防御反应机制主要包括： (1)改变自身的营养状况; (2)产生有毒的次生化合物; (3)产生防御蛋白。防御反应与植物水杨酸、 茉莉酸、 乙烯等信号分子密切相关。研究表明, 刺吸式昆虫取食诱导的植物防御反应主要引发以水杨酸为主的信号途径, 但相关分子互作机制还有待明确。日益丰富的基因组资源和不断发展的分子生物学技术为揭示植物防御反应中信号分子的作用机制、 找出植物内生抗性的特异因子以及阐明诱导防御机制奠定了基础。了解刺吸式昆虫取食诱导的植物防御反应, 为深入理解植物-昆虫间协同进化关系提供了依据, 为害虫治理和抗虫植物的培育提供了新的思路。     

外源茉莉酸和茉莉酸甲酯诱导植物抗虫作用及其机理

综述了茉莉酸(jasmonic acid, JA)和茉莉酸甲酯(methyl jasmo nate, MJA)的分子结构和应用其诱导的植物抗虫作用及其机制。植物受外源茉莉酸或茉莉酸甲酯刺激后,一条反应途径是由硬脂酸途径激活防御基因,另一条途径是直接激活防御基因。防御基因激活后导致代谢途径重新配置,并可能诱导植物产生下列4种效应：（1）直接防御,即植物产生对害虫有毒的物质、抗营养和抗消化的酶类,或具驱避性和妨碍行为作用的化合物;（2）间接防御,即产生吸引天敌的挥发物;（3）不防御,即无防御反应;（4）负防御,即产生吸引害虫的挥发物。     

茉莉酸对棉花单宁含量和抗虫相关酶活性的诱导效应

以植物生长调节物茉莉酸(Jasmonic acid,JA)为诱导子,以常规棉为研究对象,探讨了外源茉莉酸对棉花幼苗单宁和蛋白酶抑制素以及其它抗虫相关酶活性诱导的浓度依赖性和持久性,讨论了棉花抗虫相关物质的抗虫效果.结果表明,0.01、0.1和1.0 mmol/L茉莉酸都能在2周内诱导棉花单宁和胰蛋白酶抑制素(Proteinase inhibitors,PIs)含量增加,诱导多酚氧化酶(Polyphenol oxidase,PPO)、苯丙氨酸解氨酶(Phenylalanine ammonia-lyase,PAL)、过氧化物酶(Peroxidase,POD)和过氧化氢酶(Catalase,CAT)活性升高.对3种浓度茉莉酸的诱导效应进行分析表明,0.1 mmol/L茉莉酸对于诱导PIs、PPO、POD和CAT最有效,0.1和1.0 mmol/L茉莉酸对于诱导棉花单宁和苯丙氨酸解氨酶等效,二者的诱导效应均高于0.01 mmol/L.对茉莉酸诱导抗性的持久性进行分析表明,最佳诱导效应发生的时间各不相同:POD活性在JA处理后第1天最高,随后呈下降趋势,PIs和单宁含量分别在JA处理后第7天和第14天达最大值;JA处理后第1天和第7天的PPO活性无明显差异,但明显高于第14天;JA处理后第7天和第14天的PAL活性无明显差异,但明显高于第1天;JA处理后第1、7和14天棉花叶片的CAT活性均无明显差异.以上结果表明,茉莉酸可通过增加棉叶单宁和PIs含量、提高棉叶PAL、PPO、POD和CAT活性等增强棉花幼苗的抗虫性.     

外源茉莉酸诱导植物反应对菜蛾绒茧蜂寄生选择行为的影响

茉莉酸是植物体内重要的伤信号分子,向植物施用外源茉莉酸后, 可诱导植物产生各种防卫反应, 如挥发物组成发生改变等, 进而影响植食性昆虫及其天敌。该文报道用不同浓度外源茉莉酸处理白菜和甘蓝后,诱导植物反应所产生的挥发物对菜蛾绒茧蜂搜索及寄生选择行为的影响。外源茉莉酸处理白菜和甘蓝后,处理植株的挥发物对菜蛾绒茧蜂的引诱力增强;与在对照植株上相比,该蜂对经茉莉酸处理后白菜植株上的小菜蛾幼虫的寄生数显著要高。表明茉莉酸处理白菜及甘蓝后,植物诱导反应导致其挥发物的作用发生变化,进而可提高该蜂的搜索和寄生效率。     

拟南芥转录因子bHLH17负调控茉莉素介导的抗性反应

植物激素茉莉素作为抗性信号调控植物对腐生性病原菌和昆虫的抗性, 作为发育信号调控植物根的生长、雄蕊发育、表皮毛形成和叶片衰老。茉莉素受体COI1识别茉莉素分子, 进而与JAZ蛋白互作并诱导其降解, 继而调控多种茉莉素反应。拟南芥(Arabidopsis thaliana) IIId亚组bHLH转录因子(bHLH3、bHLH13、bHLH14和bHLH17)是JAZ的一类靶蛋白。与野生型相比, IIId亚组bHLH转录因子的单突变体对灰霉菌和甜菜夜蛾的抗性无明显差异, 而四突变体对灰霉菌和甜菜夜蛾的抗性增强。该文通过高表达bHLH17并研究其对灰霉菌和甜菜夜蛾的抗性反应, 结果显示, 被灰霉菌侵染的bHLH17高表达植株较野生型表现出更严重的病症。取食bHLH17高表达植株叶片的甜菜夜蛾幼虫体重大于取食野生型叶片的幼虫体重。bHLH17高表达抑制了茉莉素诱导的抗性相关基因(Thi2.1)和伤害响应基因(VSP2、AOS、JAZ1、JAZ9和JAZ10)的表达。原生质体转化实验显示bHLH17通过其N端行使转录抑制功能。研究结果表明, IIId亚组bHLH转录抑制因子bHLH17高表达会负调控茉莉素介导的对灰霉菌和甜菜夜蛾的抗性。     

一氧化氮与激发子诱导的植物抗病防卫反应

来源于真菌或植物细胞壁的激发子可以诱导植物的抗性反应。一系列的信号分子,如一氧化氮、活性氧、茉莉酸、水杨酸、乙烯等都参与了激发子诱导的植物抗性反应。它们在介导激发子刺激诱发胞内抗性反应的过程中起着重要的作用。本文介绍了激发子的种类,并简述了激发了受体以及植物细胞对激发子刺激的感受与传递;重点介绍了一氧化氮在激发子诱导植物抗性反应过程中的作用,以及它与其他信号分子之间相互关系的研究进展。     

Role and activity of jasmonates in plants under in vitro conditions

Jasmonates (JAs), such as jasmonic acid and its methyl ester, are lipid-derived compounds with signal functions in plant growth and development, as well as in responses to stress. JAs are widely distributed in plants as natural plant growth regulators. JAs do not work independently but work as a part of a complex signaling network with other phytohormones. They are deployed to induce response during wounding and are often used for elicitation and stimulation of secondary metabolites production in different in vitro culture systems. Application of JAs seems to be promising during different steps of the micropropagation system for different species. JAs stimulate proliferation rate of shoots, roots, callus and induce microtubers and bulblets formation. However, negative effects of JAs on the condition of plant tissues are also reported, e.g. leaf senescence, reduced growth and inhibited somatic embryogenesis. This review summarizes the current knowledge of the application and properties of jasmonates under in vitro conditions in terms of cell division, explant growth, proliferation ability, storage organ formation and stress response.

     

Regulation of stress hormones jasmonates and ethylene by MAPK pathways in plants

Plant stress hormones, such as jasmonates (JAs) and ethylene (ET) are essential in plant defence against stress conditions. JAs are used in cosmetics and food flavouring, and the recently demonstrated anti-cancer activity of JAs highlights their potential in health protection. It reinforces the need for a better understanding of biosynthetic regulation of JAs. Which mechanisms are involved in the regulation of the biosynthesis of JAs and ET? Production of stress hormones is induced in plants after wounding or herbivore attack. ET is a gaseous compound and plant JAs are oxylipins structurally similar to prostaglandins that are induced upon inflammation or injury in mammals. Wounding activates protein phosphorylation cascades involving mitogen-activated protein kinases (MAPKs). MAPKs regulate ET production. The induction of JA biosynthesis was suggested to require MAPK activation; however the defined roles of MAPKs in JA production remain unclear. Here we will highlight the most recent findings suggesting the regulation of JA biosynthesis and ethylene production by stress activated MAPKs and phosphatases that inactivate these MAPKs.     

Exploring the impact of wounding and jasmonates on ascorbate metabolism

Vitamin C (ascorbate, AsA) is the most abundant water-soluble antioxidant in plants. Ascorbate provides the first line of defense against damaging reactive oxygen species (ROS), and helps protect plant cells from many factors that induce oxidative stress, including wounding, ozone, high salinity, and pathogen attack. Plant defenses against these stresses are also dependent upon jasmonates (JAs), a class of plant hormones that promote ROS accumulation. Here, we review evidence showing that wounding and JAs influence AsA accumulation in various plant species, and we report new data from Arabidopsis and tomato testing the influence of JAs on AsA levels in wounded and unwounded plants. In both species, certain mutations that impair JA metabolism and signaling influence foliar AsA levels, suggesting that endogenous JAs may regulate steady-state AsA. However, the impact of wounding on AsA accumulation was similar in JA mutants and wild type controls, indicating that this wound response does not require JAs. Our findings also indicate that the effects of wounding and JAs on AsA accumulation differ between species; these factors both enhanced AsA accumulation in Arabidopsis, but depressed AsA levels in tomato. These results underscore the importance of obtaining data from more than one model species, and demonstrate the complexity of AsA regulation.     

Action of jasmonates in plant stress responses and development — Applied aspects

Jasmonates (JAs) are lipid-derived compounds acting as key signaling compounds in plant stress responses and development. The JA co-receptor complex and several enzymes of JA biosynthesis have been crystallized, and various JA signal transduction pathways including cross-talk to most of the plant hormones have been intensively studied. Defense to herbivores and necrotrophic pathogens are mediated by JA. Other environmental cues mediated by JA are light, seasonal and circadian rhythms, cold stress, desiccation stress, salt stress and UV stress. During development growth inhibition of roots, shoots and leaves occur by JA, whereas seed germination and flower development are partially affected by its precursor 12-oxo-phytodienoic acid (OPDA). Based on these numerous JA mediated signal transduction pathways active in plant stress responses and development, there is an increasing interest in horticultural and biotechnological applications. Intercropping, the mixed growth of two or more crops, mycorrhization of plants, establishment of induced resistance, priming of plants for enhanced insect resistance as well as pre- and post-harvest application of JA are few examples. Additional sources for horticultural improvement, where JAs might be involved, are defense against nematodes, biocontrol by plant growth promoting rhizobacteria, altered composition of rhizosphere bacterial community, sustained balance between growth and defense, and improved plant immunity in intercropping systems. Finally, biotechnological application for JA-induced production of pharmaceuticals and application of JAs as anti-cancer agents were intensively studied.     

Jasmonate Signal Pathway in Arabidopsis

Jasmonates （JAs）, which include jasmonic acid and its cyclopentane derivatives are synthesized from the octadecanoid pathway and widely distributed throughout the plant kingdom. JAs modulate the expression of numerous genes and mediate responses to stress, wounding, insect attack, pathogen infection, and UV damage. They also affect a variety of processes in many plant developmental processes. The JA signal pathway involves two important events： the biosynthesis of JA and the transduction of JA signal. Several important Arabidopsis mutants in jasmonate signal pathway were described in this review.