植物与植食性昆虫相互作用的分子机制

在长期的协同进化中,植物建立起应对昆虫取食为害的精密而又复杂的防御机制,植物转录组调控中防御应答基因的表达及防御物质的合成因不同的昆虫取食方式而异。一般来说,咀嚼式口器昆虫取食时造成大面积组织伤害,可诱导植物产生伤害反应;而刺吸式口器昆虫因其特殊的口针取食,诱导植物激活病原体相关的防御途径。不同的防御途径激活不同的识别机制和信号途径。本文从信号识别和转导上综述了不同食性的昆虫取食植物时所引发的防御反应,分析了昆虫-植物相互作用关系的分子机制。     

虫害诱导植物防御的分子机理研究进展

从虫害诱导的系统损伤信号、昆虫特异性激发子、间接防御、直接防御和负防御等方面,综述了虫害诱导植物防御的最新研究进展.在植物与昆虫的相互进化过程中,植物利用诱导防御物质对付昆虫的危害,昆虫则利用其特有的激发子降低植物的防御反应.文中比较了间接防御涉及的4种代谢途径,以及诱导挥发物释放的机制;阐明了虫害诱导植物直接防御的概念、防御物质及其作用机理;分析了虫害诱导植物负防御的机制.同时,也强调了虫害诱导林木防御反应的分子机理.     

虫害诱导植物间接防御反应的激发与信号转导途径

植物通过产生和释放挥发性物质增加植食性昆虫的天敌对其寄主或猎物的定位，减少植食性昆虫对植物的取食，从而达到间接防御的目的。植物对植食性昆虫所做出间接防御反应激发因子和信号转导途径的研究，对应用虫害诱导植物挥发物引诱害虫天敌，并进一步从植物、植食性昆虫及其天敌间三级营养关系，研究动植物协同进化机理和病虫害防治具有深远意义。本文根据国内外最新研究进展，对虫害诱导植物间接防御反应的激发因子，昆虫取食信号的转导途径及对植物间接防御相关基因的激活等方面进行了系统地综述。     

昆虫取食诱导的植物防御反应

植物被昆虫取食后可产生直接防御或间接防御。直接防御通过增加有毒的次生代谢产物或防御蛋白对昆虫生理代谢产生不利的影响,但对植物的消耗较大。间接防御通过释放挥发性化合物吸引天敌昆虫,并以此控制植食性昆虫。特异性的昆虫激发子(insect specific elicitors)能够诱导挥发性化合物的释放。多种信号途径参与昆虫取食诱导的植物防御反应,它们之间的相互作用协同或拮抗。了解昆虫取食诱导的植物防御反应,对于害虫综合治理策略的完善具有重要的意义。     

植食性昆虫对植物的反防御机制

本文综述了植食性昆虫对植物的反防御机制.一方面,植食性昆虫可通过其快速进化的寄主选择适应性,改变取食策略,调节生长发育的节律,以及规避自然天敌等抑制、逃避或改变植物的防御,即行为防御机制;另一方面,植食性昆虫可适应植物蛋白酶抑制剂、逃避植物防御伤信号、解毒植物次生物质,以及抑制植物阻塞反应来对植物防御进行反防御,即生理和生化防御机制.其中,昆虫抑制植物伤信号,防止植物阻塞反应是反防御机制的研究热点.昆虫反防御的研究有助于提高对昆虫-植物间协同进化关系的认识,并为害虫治理和抗虫植物的培育提供新的思路.     

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。     

植物诱导性直接防御

众所周知,植物对植食性昆虫危害的反应表现在3个方面：直接防御,间接防御,和耐害性。直接防御是指植物自身所具有的能影响寄主植物感虫性的所有特性。植物对昆虫危害的直接防御包括：限制食物供给,降低营养价值,减少偏嗜程度,破坏组织结构和抑制害虫代谢途径。目前已知的防御化合物主要包括植物次生代谢物质、昆虫消化酶（蛋白）抑制剂、蛋白酶、凝集素、氨基酸脱氨酶和氧化酶。植物在防御某种昆虫为害时多个因素往往具有累加效应或协同作用,并且对一种昆虫起主导作用的因素在防御另一种昆虫时可能仅仅起次要作用甚至根本不起作用。因此,对寄主植物基因表达、蛋白水平和活性以及代谢物含量在不同时空条件下进行广泛的定量和定性的高通量分析,不仅可以促进对植物直接防御机制的全面理解,而且有助于在农业生产中加快对作物抗性的特定靶标的鉴定。     

植食性昆虫适应植物防御反应的研究进展

在植物与植食性昆虫协同进化过程中,植物在不断完善其防御反应,同时植食性昆虫也在选择压下不断适应植物防御反应。植食性昆虫适应植物防御反应存在多样性。昆虫能够利用其唾液中的效应因子抑制或弱化植物防御反应,激活其肠道中的某些特异性蛋白阻断植物防御性次生代谢物的产生或者将其直接降解,以及通过其携带微生物间接抑制植物防御反应。此外,昆虫还能够通过产卵、虫害诱导植物挥发物、识别植物防御物质等方式适应植物的防御反应。本文综述了植食性昆虫如何利用各种效应因子适应寄主植物防御反应的研究进展。     

绿叶挥发物产生特征及其生态生理作用研究进展

该文系统综述了植物绿叶挥发物(green leaf volatiles,GLVs)的形成特征、对植物的生态生理作用及调控机制等方面的最新研究进展.GLVs是指植物经十八碳烷酸途径过氧化物酶分支途径产生的含6个碳原子的醛、醇及其酯.除叶片外,植物的根、茎、果实、种子等部位均可以合成该类化合物,合成调控存在转录和转录后水平的调控.在特定植物中,GLVs合成及其组分还受到植物生长发育阶段和生长季节等外源环境的影响.昆虫啃食、微生物感染以及有益真菌的定植等生物逆境与缺氮等物理逆境均具有诱导GLVs合成的作用.除了参与植物特有气味形成外,GLVs在植物直接和间接防御应答中发挥着重要的作用.GLVs不仅具有抑制微生物和多种昆虫繁殖的作用,还具有诱导多种防御化合物合成、预置(prime)有关信号途径的作用.基于GLVs在植物界的广泛存在性和在防御应答中的多层次作用,该文作者提出了GLVs在道地药材品质形成中的潜在作用及开展相关研究的必要性和紧迫性.     

茉莉素与植物生物胁迫反应

植物固定生长不能移动,时刻面临着害虫咬噬、病原微生物侵染等多种外界环境胁迫。为了应对这些胁迫,植物进化出了复杂且被精细调控的防御系统,包括利用植物激素调控抗性基因的表达以及抗性相关次生代谢产物的积累等。有趣的是,许多昆虫和病原微生物被发现能够采用各种策略来逃避、克服甚至操控植物防御系统,以促进其对宿主植物的利用。茉莉素是一种重要的脂质植物激素,调控生长发育的诸多方面,同时也在植物应对多种生物胁迫和非生物胁迫的防御反应中发挥重要作用。近年来,茉莉素的生物合成、信号转导和生理功能等均得到了广泛研究,并取得了一系列重要进展。概述了茉莉素的生物合成调控与信号转导途径,同时也探讨了茉莉素对植物生物胁迫反应的调控机制,并介绍了昆虫和病原微生物对植物茉莉素途径的操控策略,以期为深入理解茉莉素介导的植物与病原生物之间的相互作用提供参考。     

Low nutritive quality as defense against herbivores

Contrary to a widespread assumption in the literature on plant-herbivore interactions, individual plants do not necessarily benefit by possessing traits which lower herbivore fitness. In particular, genes conferring lowered nutritive quality could even increase herbivore damage under certain circumstances. Three special sets of conditions are outlined in which low nutritive quality would lower herbivore-induced damage to an individual plant. These sets are far from exhaustive. It is concluded that the adaptiveness of lowered nutritive quality in herbivore defense is widely possible but in no case demonstrated.     

The Myriad Plant Responses to Herbivores

Abstract Plant responses to herbivores are complex. Genes activated on herbivore attack are strongly correlated with the mode of herbivore feeding and the degree of tissue damage at the feeding site. Phloem-feeding whiteflies and aphids that produce little injury to plant foliage are perceived as pathogens and activate the salicylic acid (SA)-dependent and jasmonic acid (JA)/ethylene-dependent signaling pathways. Differential expression of plant genes in response to closely related insect species suggest that some elicitors generated by phloem-feeding insects are species-specific and are dependent on the herbivore's developmental stage. Other elicitors for defense-gene activation are likely to be more ubiquitous. Analogies to the pathogen-incompatible reactions are found. Chewing insects such as caterpillars and beetles and cell-content feeders such as mites and thrips cause more extensive tissue damage and activate wound-signaling pathways. Herbivore feeding is not equivalent to mechanical wounding. Wound responses are a part of the induced responses that accompany herbivore feeding. Herbivores induce direct defenses that interfere with herbivore feeding, growth and development, fecundity, and fertility. In addition, herbivores induce an array of volatiles that creates an indirect mechanism of defense. Volatile blends provide specific cues to attract herbivore parasites and predators to infested plants. The nature of the elicitors for volatile production is discussed.     

Plant allocation of carbon to defense as a function of herbivory,light and nutrient availability

We use modeling to determine the optimal relative plant carbon allocations between foliage, fine roots, anti-herbivore defense, and reproduction to maximize reproductive output. The model treats these plant components and the herbivore compartment as variables. Herbivory is assumed to be purely folivory. Key external factors include nutrient availability, degree of shading, and intensity of herbivory. Three alternative functional responses are used for herbivory, two of which are variations on donor-dependent herbivore (models 1a and 1b) and one of which is a Lotka–Volterra type of interaction (model 2). All three were modified to include the negative effect of chemical defenses on the herbivore. Analysis showed that, for all three models, two stable equilibria could occur, which differs from most common functional responses when no plant defense component is included. Optimal strategies of carbon allocation were defined as the maximum biomass of reproductive propagules produced per unit time, and found to vary with changes in external factors. Increased intensity of herbivory always led to an increase in the fractional allocation of carbon to defense. Decreases in available limiting nutrient generally led to increasing importance of defense. Decreases in available light had little effect on defense but led to increased allocation to foliage. Decreases in limiting nutrient and available light led to decreases in allocation to reproduction in models 1a and 1b but not model 2. Increases in allocation to plant defense were usually accompanied by shifts in carbon allocation away from fine roots, possibly because higher plant defense reduced the loss of nutrients to herbivory.     

Herbivore-specific plant volatiles prime neighboring plants for nonspecific defense responses

Plants produce species-specific herbivore-induced plant volatiles (HIPVs) after damage. We tested the hypothesis that herbivore-specific HIPVs prime neighboring plants to induce defenses specific to the priming herbivore. Since Manduca sexta (specialist) and Heliothis virescens (generalist) herbivory induced unique HIPV profiles in Nicotiana benthamiana, we used these HIPVs to prime receiver plants for defense responses to simulated herbivory (mechanical wounding and herbivore regurgitant application). Jasmonic acid (JA) accumulations and emitted volatile profiles were monitored as representative defense responses since JA is the major plant hormone involved in wound and defense signaling and HIPVs have been implicated as signals in tritrophic interactions. Herbivore species-specific HIPVs primed neighboring plants, which produced 2 to 4 times more volatiles and JA after simulated herbivory when compared to similarly treated constitutive volatile-exposed plants. However, HIPV-exposed plants accumulated similar amounts of volatiles and JA independent of the combination of priming or challenging herbivore. Furthermore, volatile profiles emitted by primed plants depended only on the challenging herbivore species but not on the species-specific HIPV profile of damaged emitter plants. This suggests that feeding by either herbivore species primed neighboring plants for increased HIPV emissions specific to the subsequently attacking herbivore and is probably controlled by JA.     

Indirect plant defense against insect herbivores: a review

Plants respond to herbivore attack by launching 2 types of defenses: direct defense and indirect defense. Direct defense includes all plant traits that increase the resistance of host plants to insect herbivores by affecting the physiology and/or behavior of the attackers. Indirect defense includes all traits that by themselves do not have significant direct impact on the attacking herbivores, but can attract natural enemies of the herbivores and thus reduce plant loss. When plants recognize herbivore‐associated elicitors, they produce and release a blend of volatiles that can attract predators, parasites, and other natural enemies. Known herbivore‐associated elicitors include fatty acid–amino acid conjugates, sulfur‐containing fatty acids, fragments of cell walls, peptides, esters, and enzymes. Identified plant volatiles include terpenes, nitrogenous compounds, and indoles. In addition, constitive traits including extrafloral nectars, food bodies, and domatia can be further induced to higher levels and attract natural enemies as well as provide food and shelter to carnivores. A better understanding of indirect plant defense at global and componential levels via advanced high throughput technologies may lead to utilization of indirect defense in suppression of herbivore damage to plants.     

激发子α-吡啶羧酸诱发拟南芥和水稻悬浮细胞中依赖于水杨酸、茉莉酸/乙烯和钙离子途径的防卫反应(英文)

α-吡啶羧酸(PA)是动物细胞程序化死亡的诱导物。我们前期的研究表明,PA可以激发单子叶模式植物水稻的过敏反应(HR)。进一步用双子叶模式植物拟南芥(Arabidopsis thaliana)进行的研究表明,PA是一个广谱的植物HR反应的激发子,包括诱导氧进发和细胞死亡。我们探究了PA诱导的拟南芥防卫反应途径,利用不同信号途径标志基因PR-1,PR-2和PDF1.2受诱导剂量和时间激活的结果,表明PA可以同时激活水杨酸和茉莉酸/乙烯依赖的防卫途径。我们也发现PA诱导水稻悬浮细胞产生活性氧是钙离子依赖性的。综合所有结果,我们认为PA可以作为一个非专化性的植物防卫反应激发子,可望用于系统获得性抗性激发的细胞模型的建立。