植物气味多样性与昆虫关系的研究进展

植物气味是多种挥发物组成的混合物,具有丰富的多样性。由于植物气味多样性在植食性昆虫及其天敌与植物的关系中起着决定性作用,近年来将昆虫化学生态学、昆虫行为学与景观生态学研究相结合,探讨田间和不同景观尺度上植物气味多样性与昆虫嗅觉行为的关系受到关注。本文从植物气味多样性相关概念的分析、植物气味多样性与植食性昆虫的关系、植物气味多样性与天敌昆虫的关系、植物气味多样性研究方法等方面,介绍了国内外最新的研究进展,期望为推进我国该领域研究提供参考。     

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.     

虫害诱导的植物挥发物:植物与植食性昆虫及其天敌相互作用的进化产物

综述了植物、植食性昆虫及其天敌相互作用的进化过程。虫害诱导的植物挥发物的特征和功能是植物-植食性昆虫-天敌之间长期进化的结果。在植物、植食性昆虫与天敌相互作用的进化过程中,3个不同营养级,包括植物、植食性昆虫和天敌有着各自的调节和利用虫害诱导的植物挥发物的策略。但有一些问题,如通过实验研究得出的诱导防御在田间是否真正能起到保护作用等需进一步研究、阐明。     

冬虫夏草蝠蛾幼虫对几种植物的嗅觉反应初步研究

植食性昆虫主要利用嗅觉、味觉、视觉和触觉等对寄主植物进行选择,而植物挥发物是引导植食性昆虫寄主定向行为的主要因素。利用"Y"型嗅觉仪测试康定虫草蝠蛾Hepialus armoricanus二龄幼虫对适生地的几种植物材料的嗅觉反应;并用欧式距离和Q型聚类法对供试植物进行相似性分析和聚类分析。结果表明:蝠蛾幼虫对几种植物材料表现出强弱不同的趋性反应;根据植物的相似性可将供试植物分为4类,且同一类植物引导蝠蛾幼虫产生相似的嗅觉反应。     

植物与植食性昆虫防御与反防御的三个层次

在植物与植食性昆虫长期的进化过程中,双方形成了一系列的防御与反防御策略。本文将这些策略归为3个层次:第一层次起始于植物对植食性昆虫相关分子模式的识别,并由此激活植食性昆虫分子模式相关的免疫反应。这种免疫反应对于不能产生效应子的植食性昆虫种群是有效的;第二层次是一些植食性昆虫种群可以通过释放特异性效应子抑制植物产生的植食性昆虫分子模式相关的免疫反应,从而在植物上正常生长与繁衍;第三层次是一些植物基因型可以通过特异抗性基因识别植食性昆虫的效应子,进而激活效应子诱导的免疫反应,表现出特异的抗虫性。深入揭示植物与植食性昆虫间的这种分子互作机制,不仅在理论上有助于理解昆虫与植物的协同进化机制,而且在实践上可为作物抗性品种的培育提供重要的技术指导。     

寄生蜂基于嗅觉信号识别的宿主定位及其机制的研究进展

寄生蜂对宿主的成功定位是其繁衍后代的关键步骤之一，主要受挥发性化合物的调节。首先，雌性寄生蜂可根据寄主植物挥发物尤其是植食性昆虫诱导的植物挥发物(herbivore-induced plant volatiles, HIPVs)对宿主的栖息环境进行远距离定位，它们可根据HIPVs提供的信号缩小宿主范围，而HIPVs的混合物极为复杂，其成分和含量受多种因素的调节。尽管如此，萜类是HIPVs中常见的化合物，且被多数行为研究证实在寄生蜂宿主定位中发挥作用。随后，当寄生蜂发现并降落到与宿主相关的植物后，它们可利用宿主衍生的信号近距离定位宿主。宿主的虫体、茧和粪便等释放的挥发性化合物对寄生蜂具有吸引作用，有些宿主粪便的气味可以作为寄生蜂执行宿主定位行为的主信号，而且宿主粪便挥发物的成分因昆虫取食的植物种类的不同而发生变化。此外，来自寄生蜂自身的信息化学物质也有助于同种寄生蜂的其他个体对宿主的定位。寄生蜂对气味分子的感知依赖其主要位于触角上的嗅觉感器，而关于其嗅觉识别的分子机制的研究仍较为薄弱。由于气味结合蛋白(odorant-binding proteins, OBPs)是嗅觉系统中介导气味识别...     

虫害诱导的植物挥发物:基本特性、生态学功能及释放机制

植物在遭受植食性昆虫攻击时,能通过释放挥发物调节植物、植食性昆虫及其天敌三者间的相互关系,并由此而防御植食性昆虫。主要就虫害诱导的植物挥发物的基本特性、生态学功能及其释放机制进行了系统性综述,并提出了今后的研究方向。     

新旧创伤诱导的不同挥发物对寄生蜂选择性的影响

植物在植食性昆虫危害的压力下会释放出化合物,而寄生蜂却可以利用这些诱导出的植物挥发物定位寄主。植物叶表面受到创伤都会释放出最常见的绿叶挥发物（green leaf volatiles）,而植食性昆虫取食引诱的挥发物会因害虫和受害植物的不同而异。植物在受到害虫造成的创伤后会马上挥发出绿叶性化合物,而一些萜稀类化合物（terpenoids）只有在旧的创伤部位才会大量挥放。那么不同的寄生蜂可以识别这些化合物吗？     

对食性分类的商榷

昆虫的食性很复杂,但目前常用的分类,是按昆虫所取食物的性质分为:植食性、肉食性和杂食性。取食活植物的为植食性,取食动物的为肉食性,既能取食动物又能取食植物的为杂食性。植食性昆虫,又以选择食物种类多寡而分为单食性,寡食性和多食性。昆虫仅取食一种植物的为单食性。寡食性是指以某一类群的植物为食的昆虫。多食性是以多种不同类群的植物为食的昆虫。     

烟夜蛾和棉铃虫对高浓度烟草挥发物的电生理和行为反应

寡食性的烟夜蛾Helicoverpa assulta (Guenée)和广食性的棉铃虫H. armigera (Hübner)是夜蛾属Heliothis的两近缘种昆虫。主要的农作物中,棉花和番茄上极少发现烟夜蛾,而辣椒上几乎没有棉铃虫,只有烟草均为二者所嗜食,也唯独在烟草上它们可以稳定共存。为明确植物挥发物对昆虫寄主定向和选择行为的影响,本文运用触角电位（EAG）和风洞技术,测定了两种夜蛾的处女雌蛾、交配雌蛾及雄蛾对20种高浓度（0.1 mol/L）烟草挥发物的电生理和行为反应。结果表明：烟夜蛾和棉铃虫对测试的烟草挥发物均可产生EAG反应,而且都表现出绿叶气味＞脂类和芳香化合物＞单萜、倍半萜和杂环化合物的总体反应趋势;两种夜蛾对多数化合物的EAG反应既无显著的种内性别差异,也无显著的种间差异,表明二者嗅觉神经系统对寄主气味图谱的识别和感受能力具有一定的相似性,可对寄主植物的化学信息产生相同的“理解”;二者的行为反应结果与EAG反应结果基本一致,即两种夜蛾受到的嗅觉刺激与产生的行为反应具有统一性,结合两种夜蛾寄主范围的差异,推测烟草挥发物主要影响两种夜蛾的寄主定向行为,而寄主选择行为可能更依赖于二者与植物接触后对植物理化性质的评价过程。     

视觉信号在昆虫检测植物寄主中的作用

植物为数十万种昆虫提供各种资源,如食物、交配、产卵和躲避天敌的场所。目前对昆虫检测植物寄主的研究主要关注昆虫嗅觉系统和植物寄主挥发物之间的相互作用,对昆虫视觉系统发挥的作用关注较少。近年来,对昆虫视觉器官、光行为反应及分子生物学的研究表明,昆虫具有优异的视觉能力,能够辨别植物寄主的颜色、大小和轮廓,应该将视觉纳入昆虫检测植物寄主的研究中。昆虫能够利用视觉信号准确检测寄主,远距离时,主要依靠植物寄主轮廓检测寄主,近距离时,寄主的大小、颜色和形状发挥重要作用。利用昆虫视觉识别寄主的专一性研制诱捕装置,可为害虫的监测和防治提供一定的理论基础。     

Perception of plant volatile blends by herbivorous insects--finding the right mix

Volatile plant secondary metabolites are detected by the highly sensitive olfactory system employed by insects to locate suitable plants as hosts and to avoid unsuitable hosts. Perception of these compounds depends on olfactory receptor neurones (ORNs) in sensillae, mostly on the insect antennae, which can recognise individual molecular structures. Perception of blends of plant volatiles plays a pivotal role in host recognition, non-host avoidance and ensuing behavioural responses as different responses can occur to a whole blend compared to individual components. There are emergent properties of blend perception because components of the host blend may not be recognised as host when perceived outside the context of that blend. Often there is redundancy in the composition of blends recognised as host because certain compounds can be substituted by others. Fine spatio-temporal resolution of the synchronous firing of ORNs tuned to specific compounds enables insects to pick out relevant host odour cues against high background noise and with ephemeral exposure to the volatiles at varying concentrations. This task is challenging as they usually rely on ubiquitous plant volatiles and not those taxonomically characteristic of host plants. However, such an odour coding system has the advantage of providing flexibility; it allows for adaptation to changing environments by alterations in signal processing while maintaining the same peripheral olfactory receptors.     

Habitats as Complex Odour Environments: How Does Plant Diversity Affect Herbivore and Parasitoid Orientation?

Plant diversity is known to affect success of host location by pest insects, but its effect on olfactory orientation of non-pest insect species has hardly been addressed. First, we tested in laboratory experiments the hypothesis that non-host plants, which increase odour complexity in habitats, affect the host location ability of herbivores and parasitoids. Furthermore, we recorded field data of plant diversity in addition to herbivore and parasitoid abundance at 77 grassland sites in three different regions in Germany in order to elucidate whether our laboratory results reflect the field situation. As a model system we used the herb Plantago lanceolata, the herbivorous weevil Mecinus pascuorum, and its larval parasitoid Mesopolobus incultus. The laboratory bioassays revealed that both the herbivorous weevil and its larval parasitoid can locate their host plant and host via olfactory cues even in the presence of non-host odour. In a newly established two-circle olfactometer, the weeviĺs capability to detect host plant odour was not affected by odours from non-host plants. However, addition of non-host plant odours to host plant odour enhanced the weeviĺs foraging activity. The parasitoid was attracted by a combination of host plant and host volatiles in both the absence and presence of non-host plant volatiles in a Y-tube olfactometer. In dual choice tests the parasitoid preferred the blend of host plant and host volatiles over its combination with non-host plant volatiles. In the field, no indication was found that high plant diversity disturbs host (plant) location by the weevil and its parasitoid. In contrast, plant diversity was positively correlated with weevil abundance, whereas parasitoid abundance was independent of plant diversity. Therefore, we conclude that weevils and parasitoids showed the sensory capacity to successfully cope with complex vegetation odours when searching for hosts.     

Root herbivores influence the behaviour of an aboveground parasitoid through changes in plant-volatile signals

It is widely reported that plants emit volatile compounds when they are attacked by herbivorous insects, which may be used by parasitoids and predators to locate their host or prey. The study of herbivore-induced plant volatiles and their role in mediating interactions between plants, herbivores and their natural enemies have been primarily based on aboveground systems, generally ignoring the potential interactions between above and belowground infochemical- and food webs. This study examines whether herbivory by Delia radicum feeding on roots of Brassica nigra (black mustard) affects the behaviour of Cotesia glomerata , a parasitoid of the leaf herbivore Pieris brassicae , mediated by changes in plant volatiles. In a semi-field experiment with root-damaged and root-undamaged plants C. glomerata prefers to oviposit in hosts feeding on root-undamaged plants. In addition, in a flight-cage experiment the parasitoid also prefers to search for hosts on plants without root herbivores. Plants exposed to root herbivory were shown to emit a volatile blend characterized by high levels of specific sulphur volatile compounds, which are reported to be highly toxic for insects, combined with low levels of several compounds, i.e. beta-farnesene, reported to act as attractants for herbivorous and carnivorous insects. Our results provide evidence that the foraging behaviour of a parasitoid of an aboveground herbivore can be influenced by belowground herbivores through changes in the plant volatile blend. Such indirect interactions may have profound consequences for the evolution of host selection behaviour in parasitoids, and may play an important role in the structuring and functioning of communities.     

植物挥发性次生物质在植食性昆虫、寄生植物和昆虫天敌关系中的作用机理

植物挥发性次生物质在植食性昆虫、寄生植物和昆虫天敌关系中的作用机理杜永均,严福顺（中国科学院动物研究所,北京１０００８０）植物挥发性次生物质是一些分子量在１００—２００的有机化学物质,包括烃类、醇类、醛类、酮类、有机酸、内酯、含氮化合物以及有机硫等化...     

植食性昆虫寄主植物嗅觉搜寻述介

植食性昆虫与寄主植物关系的本质是化学。植食性昆虫搜寻寄主的嗅觉媒介是植物气味即化学信息物质。在介绍植物气味构成及其扩散模型基础上，阐述了植物气味在地上植食性昆虫成虫、幼虫和地下植食性昆虫搜寻寄主过程中的嗅觉导向作用，并指出了今后相关研究需要注意的问题。从植物与环境因子的关系来看，植物气味包括构成性气味和诱发性气味两类，这两类气味的概念既相联系而又不同。构成性气味组分及构成因植物分类地位等而不同。诱发性气味组分因植食性昆虫取食、植物病原微生物、机械致伤等因子的胁迫而变化，这种变化性状随植物属和/或种、植株生长发育阶段、胁迫因子性质及其作用方式而不同。无论是哪种植物气味，其释放均具有节律性。气味扩散过程比较复杂，扩散状态可用数学模型表征。对于地上植食性昆虫成虫，植物气味对其寄主搜寻行为具有导向特异性，重点分析了这种特异性形成的两个假说；鳞翅目昆虫幼虫，能够利用植物化学信息物质趋向寄主植物或回避非寄主植物；地下植食性昆虫搜寻寄主，既与寄主植物地下组织释放或分泌的次级代谢物有关，又与一些初级代谢物有关。初级代谢物中的CO₂,起着“搜寻触发器”作用。有助于增强人们对昆虫与植...     

Cockchafer Larvae Smell Host Root Scents in Soil

In many insect species olfaction is a key sensory modality. However, examination of the chemical ecology of insects has focussed up to now on insects living above ground. Evidence for behavioral responses to chemical cues in the soil other than CO₂ is scarce and the role played by olfaction in the process of finding host roots below ground is not yet understood. The question of whether soil-dwelling beetle larvae can smell their host plant roots has been under debate, but proof is as yet lacking that olfactory perception of volatile compounds released by damaged host plants, as is known for insects living above ground, occurs. Here we show that soil-dwelling larvae of Melolontha hippocastani are well equipped for olfactory perception and respond electrophysiologically and behaviorally to volatiles released by damaged host-plant roots. An olfactory apparatus consisting of pore plates at the antennae and about 70 glomeruli as primary olfactory processing units indicates a highly developed olfactory system. Damage induced host plant volatiles released by oak roots such as eucalyptol and anisol are detected by larval antennae down to 5 ppbv in soil air and elicit directed movement of the larvae in natural soil towards the odor source. Our results demonstrate that plant-root volatiles are likely to be perceived by the larval olfactory system and to guide soil-dwelling white grubs through the dark below ground to their host plants. Thus, to find below-ground host plants cockchafer larvae employ mechanisms that are similar to those employed by the adult beetles flying above ground, despite strikingly different physicochemical conditions in the soil.     

Insect host location: a volatile situation

Locating a host plant is crucial for a phytophagous (herbivorous) insect to fulfill its nutritional requirements and to find suitable oviposition sites. Insects can locate their hosts even though the host plants are often hidden among an array of other plants. Plant volatiles play an important role in this host-location process. The recognition of a host plant by these olfactory signals could occur by using either species-specific compounds or specific ratios of ubiquitous compounds. Currently, most studies favor the second scenario, with strong evidence that plant discrimination is due to central processing of olfactory signals by the insect, rather than their initial detection. Furthermore, paired or clustered olfactory receptor neurons might enable fine-scale spatio-temporal resolution of the complex signals encountered when ubiquitous compounds are used.     

捕食螨化学生态研究进展

捕食螨是重要的生物防治因子。早在20世纪70年代就发现了捕食螨的性信息素,许多研究证明植物挥发物在捕食螨向猎物定位过程中发挥着至关重要的作用,影响捕食螨寻找猎物的植物挥发物来源于未受害植物、机械损伤植物、猎物危害植物、非猎物危害植物。人工合成的植物挥发物组分对捕食螨具有引诱作用,但引诱活性低于虫害诱导植物释放的挥发性混合物。捕食螨的饲养条件、饥饿程度、学习与经验行为等会影响捕食螨对植物挥发物的反应。介绍了信息素与植物挥发物对捕食螨的作用,并讨论了目前存在的问题和研究前景。