虫害诱导的植物挥发物代谢调控机制研究进展

长期受自然界的非生物/生物侵害,植物逐步形成了复杂的防御机制,为防御植食性昆虫的为害,植物释放虫害诱导产生的挥发性化合物(herbivore-induced plant volatiles,HIPVs)。HIPVs是植物-植食性昆虫-天敌三级营养关系之间协同进化的结果。HIPVs的化学组分因植物、植食性昆虫种类的不同而有差异。生态系统中,HIPVs可在植物与节肢动物、植物与微生物、虫害植物与邻近的健康植物、或同一植株的受害和未受害部位间起作用,介导防御性反应。HIPVs作为寄主定位信号,在吸引捕食性、寄生性天敌过程中起着重要作用。HIPVs还可以作为植物间信息交流的工具,启动植株的防御反应而增强抗虫性。不论从生态学还是经济学角度来看,HIPVs对于农林生态系中害虫综合治理策略的完善具有重要意义。前期的研究在虫害诱导植物防御的化学生态学方面奠定了良好基础,目前更多的研究转向阐述虫害诱导植物抗性的分子机制。为了深入了解HIPVs的代谢调控机制,主要从以下几个方面进行了综述。因为植食性昆虫取食造成的植物损伤是与昆虫口腔分泌物共同作用的结果,所以首先阐述口腔分泌物在防御反应中的作用。挥发物诱导素volicitin和β-葡萄糖苷酶作为口腔分泌物的组分,是产生HIPVs的激发子,通过调节伤信号诱发HIPVs的释放。接着阐述了信号转导途径对HIPVs释放的调节作用,并讨论了不同信号途径之间的交互作用。就HIPVs的代谢过程而言,其过程受信号转导途径(包括茉莉酸、水杨酸、乙烯、过氧化氢信号途径)的调控,其中茉莉酸信号途径是诱发HIPVs释放的重要途径。基于前人的研究,综述了HIPVs的主要代谢过程及其过程中关键酶类的调控作用。文中的HIPVs主要包括萜烯类化合物、绿叶挥发物和莽草酸途径产生的芳香族化合物,如水杨酸甲酯和吲哚等。作为化学信号分子,这些化合物中的一部分还能激活邻近植物防御基因的表达。萜烯合酶是各种萜烯类化合物合成的关键酶类,脂氧合酶、过氧化氢裂解酶也是绿叶挥发物代谢途径中的研究热点,而苯丙氨酸裂解酶和水杨酸羧基甲基转移酶分别是合成水杨酸及其衍生物水杨酸甲酯的关键酶类。这些酶类的基因在转录水平上调控着HIPVs代谢途径。最后展望了HIPVs的研究前景。     

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。     

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.     

茉莉酸在植物诱导防御中的作用

茉莉酸(JA)和茉莉酸甲酯(MeJA)作为与损伤相关的植物激素和信号分子,广泛地存在于植物体中,外源应用能够激发防御植物基因的表达,诱导植物的化学防御,产生与机械损伤和昆虫取食相似的效果。大量研究表明,用茉莉酸类化合物处理植物可系统诱导蛋白酶抑制剂(PI)和多酚氧化酶(PPO),从而影响植食动物对营养物质的吸收,还能增加过氧化物酶、壳聚糖酶和脂氧合酶等防御蛋白的活性水平,导致生物碱和酚酸类次生物质的积累,增加并改变挥发性信号化合物的释放,甚至形成防御结构,如毛状体和树脂导管。经茉莉酸处理的植物提高了植食动物的死亡率,变得更加吸引捕食性和寄生性天敌。挥发性化合物——茉莉酸甲酯可以从植物的气孔进入植物体内,在细胞质中被酯酶水解为茉莉酸,实现长距离的信号传导和植物间的交流,诱导邻近植物产生诱导防御反应。茉莉酸和茉莉酸甲酯分别具有4种立体异构,其中具有活性的是顺式结构,但顺式结构不稳定,会差向异构化为反式结构。茉莉酸的代谢物(Z)-茉莉酮(cis-Jasmone)具电生理活性,在植物诱导防御中起作用,并且在防御信号的作用上不同于茉莉酸和茉莉酸甲酯。     

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。     

植物的防御性萜类挥发信号分子

植物生长发育过程中会合成大量萜类化合物,其中多数具有挥发性。这些挥发性萜类成分除参与保护光合器官免受光合作用产生的热量灼伤之外,还作为信号分子参与了大量的化学生态过程,例如对昆虫和其他动物的吸引以利于花粉和种子的传播。另一方面,自上世纪80年代以来,逐渐有研究工作表明,挥发性萜类化合物也可以作为信号介导植物对植食性昆虫的防御反应,以及在植物．害虫．天敌的三级营养关系中发挥作用。同时,研究工作还显示萜类化合物可以作为信号在同株植物的不同部位之间及邻近植物之间进行有效传递以促进对虫害的防御,并证明了这些萜类分子在细胞和分子水平上的调控作用。这些工作为利用化学生态学手段对虫害进行有效控制提供了新的策略。文章仅就近年来对植物萜类挥发信号分子在合成、识别、传输及其作用等方面的研究进展做简要回顾。     

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。     

植物与植食性昆虫的分子互作:基础与应用

植物与植食性昆虫之间存在着复杂的分子互作.首先,植食性昆虫会利用自身的嗅觉和味觉化学感觉系统,通过对植物挥发性和非挥发性信息化合物的编码与解析,结合对植物颜色、形状等物理信息的感觉与编码,定位及确定寄主植物.其次,植物可以通过位于细胞膜的受体识别植食性昆虫相关模式分子和损伤相关模式分子,启动由早期信号事件和植物激素信号途径介导的防御反应,并由此而影响植食性昆虫的种群适合度.最后,为抵御寄主植物的防御反应,植食性昆虫会通过复杂多样的反防御策略适应或抑制寄主植物的防御反应.本文对如上所述的植物与植食性昆虫分子互作研究进展及由此而开发的一些害虫防控新技术进行了综述.     

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。     

虫害诱导挥发物的生态调控功能

虫害诱导挥发物（herbivore-induced plant volatiles, HIPVs）是植物受害虫胁迫后释放的挥发性物质,是植物与周围环境进行信息交流的媒介。环境中的天敌、害虫和植物通过感知HIPVs所携带的信息,对各自的行为或生理生化反应做出相应的调整。介绍了挥发物的种类及主要的生物合成途径,概括了影响天敌依据HIPVs搜寻寄主和猎物的主要因素。综述了这类挥发性物质对植食性昆虫寄主选择或产卵行为的影响,介绍了植物地上部分和地下部分受害后对彼此间接防御的影响,讨论了多种害虫加害同种植物后对天敌搜寻猎物或寄主行为的影响。另外,作为损伤信号,HIPVs还能诱导同株植物未受害部位和邻近植株的防御反应。最后,对HIPVs在害虫防治中的应用现状及前景作了介绍和讨论。     

植物种间和种内的化学作用

植物间相互作用是生态学基础科学问题之一,植物能感受和识别共存同种或异种植物,进而调整生长、繁殖和防御策略。植物种间和种内的感受和识别大多是由植物产生释放的次生物质所介导,这类化学识别通讯可以启动相应的植物化感作用机制。近年发现,植物亲属间也存在着化学识别、地下根系通讯调控地上开花繁殖等植物种间和种内的化学作用关系。目前植物通过地上挥发物介导的植物化学作用已基本澄清,但根分泌物介导的植物地下化学作用机制及其信号物质还所知甚少。地下化学作用不仅决定根系侵入(接近)和躲避(排斥)行为,也能调控地上开花时间和花期。这样,植物间的化学作用还涉及植物地下和地上的协调互作。本文以植物化感作用和植物化学识别通讯及相应的化感物质和信号物质为基点,从植物亲属识别、根系化学识别和行为模式、地下化学作用调控地上开花繁殖3个方面综述植物种间和种内化学作用的研究进展,为全面理解植物间相互作用提供新视野。     

Acquired immunity to herbivory and allelopathy caused by airborne plant emissions

Numerous plant species respond to volatile cues to adjust their defenses against herbivores. Some volatile chemicals, such as terpenoids and green leaf volatiles, that are responsible for communication between plants and arthropods are also required for intraspecific communication between plants and for coordination among branches within a single plant. We are now aware that some ‘receiver’ plants are able to eavesdrop on their neighbors and tailor their defenses to their current and expected risks caused by herbivores. By contrast, a suite of volatiles also serve as natural herbicides (allelochemicals) that are detrimental for receiver plants. Since various molecular and ecological mechanisms underlying these phenomena have been clarified, it is time to ask whether more plants eavesdrop on infochemical cues, and if these cues that allow them to adjust their defenses to suit their risk also increase their fitness as a result.     

Herbivore induced plant volatiles: Their role in plant defense for pest management

Plants respond to herbivory through different defensive mechanisms. The induction of volatile emission is one of the important and immediate response of plants to herbivory. Herbivore-induced plant volatiles (HIPVs) are involved in plant communication with natural enemies of the insect herbivores, neighboring plants, and different parts of the damaged plant. Release of a wide variety of HIPVs in response to herbivore damage and their role in plant-plant, plant-carnivore and intraplant communications represents a new facet of the complex interactions among different trophic levels. HIPVs are released from leaves, flowers, and fruits into the atmosphere or into the soil from roots in response to herbivore attack. Moreover, HIPVs act as feeding and/or oviposition deterrents to insect pests. HIPVs also mediate the interactions between the plants and the microorganisms. This review presents an overview of HIPVs emitted by plants, their role in plant defense against herbivores and their implications for pest management.     

Airborne signals of communication in sagebrush: a pharmacological approach

When plants receive volatiles from a damaged plant, the receivers become more resistant to herbivory. This phenomenon has been reported in many plant species and called plant-plant communication. Lab experiments have suggested that several compounds may be functioning as airborne signals. The objective of this study is to identify potential airborne signals used in communication between sagebrush (Artemisia tridentata) individuals in the field. We collected volatiles of one branch from each of 99 sagebrush individual plants. Eighteen different volatiles were detected by GC-MS analysis. Among these, 4 compounds; 1.8-cineol, β-caryophyllene, α-pinene and borneol, were investigated as signals of communication under natural conditions. The branches which received either 1,8-cineol or β-caryophyllene tended to get less damage than controls. These results suggested that 1,8-cineol and β-caryophyllene should be considered further as possible candidates for generalized airborne signals in sagebrush.     

Roles of (Z)-3-hexenol in plant-insect interactions

Green leaf C6-volatiles are among the most important herbivore-induced plant volatiles (HIPVs). They play important roles in mediating the behavior of herbivores and their natural enemies, and in triggering the plant-plant communication to prevent further attacks. Recently, wound-induced ubiquitous (Z)-3-hexenol, a C6-alcohol synthesized in the lipoxygenase/HPL pathway, was proved to be the most important info chemical for the herbivore repellence/attraction and natural enemy attraction in tritrophic interactions, as well as for the induction of gene expression in neighboring unattacked plants. In spite of the conflict functions of (Z)-3-hexenol in direct and indirect plant defenses, its positive roles in the indirect defense and the priming effect are consistent. Therefore, this compound can be used to develop novel insect pest control strategies.Key words: green leaf volatiles, (Z)-3-hexenol, direct defense, indirect defense, primingTo date, nearly 2000 volatile compounds have been identified in plant species from over 90 families.¹ These compounds are released from plant organs above or below the ground, and some are induced by biotic activities. Herbivore feeding stimulates the plants to release green leaf volatiles (GLVs), terpenoids, nitrogen-containing nitriles and oximes, methyl salicylate, etc. Production of these volatiles by plants involves at least three biosynthetic pathways: the fatty acid/lipoxygenase pathway for green leaf volatiles, the isoprenoid pathway for terpenoids, and the shikimic acid pathway for methyl salicylate.² Herbivore-damaged plants emit some of the most common GLVs and terpenoids that play important roles in mediating the behaviors of herbivores and their natural enemies, as well as in triggering the plant-plant communication.¹ Recently, functional studies on green leaf C6-volatiles have received wide attention and made exciting progresses. Especially, accumulating evidences on the C6-volatile (Z)-3-hexenol support its role in mediating indirect defense responses of plant.     

Tiadinil, a plant activator of systemic acquired resistance, boosts the production of herbivore-induced plant volatiles that attract the predatory mite Neoseiulus womersleyi in the tea plant Camellia sinensis

Plants respond with various defense mechanisms to pathogenic or herbivorous attack. Some chemicals called plant activators that induce the plant defense response against pathogens have been commercially used to protect plants. Here we studied the effects of tiadinil (TDL) on defense mechanisms against herbivores. TDL suppresses pathogenic fungi on tea leaves by inducing defense mechanisms. We used one of the major trophic systems in tea consisting of the herbivorous mite, Tetranychus kanzawai, and the predatory mite, Neoseiulus womersleyi. TDL enhanced the production of herbivore-induced plant volatiles that attract predatory mites. The predatory mites preferred the T. kanzawai-induced volatiles from TDL-treated plants to those produced by untreated plants. These results suggest that TDL activates the plant defense response via an indirect process mediated by plant volatiles that attract natural enemies of the herbivores. In contrast, the oviposition rate, adult maturation rate, and sex ratio of T. kanzawai were not affected by TDL treatment. These results suggest that TDL did not activate any direct defense against the herbivorous mite.     

Induction of priming by cold stress via inducible volatile cues in neighboring tea plants

Plants have evolved sophisticated defense mechanisms to overcome their sessile nature. However, if and how volatiles from cold‐stressed plants can trigger interplant communication is still unknown. Here, we provide the first evidence for interplant communication via inducible volatiles in cold stress. The volatiles, including nerolidol, geraniol, linalool, and methyl salicylate, emitted from cold‐stressed tea plants play key role(s) in priming cold tolerance of their neighbors via a C‐repeat‐binding factors‐dependent pathway. The knowledge will help us to understand how plants respond to volatile cues in cold stress and agricultural ecosystems.     

Plant–plant communication and community of herbivores on tall goldenrod

1. The volatiles from damaged plants induce defense in neighboring plants. The phenomenon is called plant–plant communication, plant talk, or plant eavesdropping. Plant–plant communication has been reported to be stronger between kin plants than genetically far plants in sagebrush.
2. Why do plants distinguish volatiles from kin or genetically far plants? We hypothesize that plants respond only to important conditions; the induced defense is not free of cost for the plant. To clarify the hypothesis, we conducted experiments and investigations using goldenrod of four different genotypes.
3. The arthropod community on tall goldenrods were different among four genotypes. The response to volatiles was stronger from genetically close plants to the emitter than from genetically distant plants from the emitter. The volatiles from each genotype of goldenrods were different; and they were categorized accordingly. Moreover, the arthropod community on each genotype of goldenrods were different.
4. Synthesis: Our results support the hypothesis: Goldenrods respond to volatiles from genetically close plants because they would have similar arthropod species. These results are important clues elucidating adaptive significance of plant–plant communication.

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Biochemical and physiological mechanisms mediated by allelochemicals

Allelochemistry, the production and release of toxic chemicals produced by one species that affect a receiving susceptible species, has been the subject of diverse degrees of scientific enquiry. Recent advances in plant biology have permitted the revamp of allelochemistry as a biologically and ecologically sound explanation for plant invasion and plant-plant communication in the rhizosphere. Recent progress has been made in understanding the biochemical and molecular changes that are induced by allelochemicals in susceptible plant species, and the complex mechanisms that are used by allelochemical-resistant plants to defend against this toxic insult.