Plant and insect cuticular lipids serve as behavioral cues for insects

The roles of plant and insect cuticular lipids in insect and plant interactions are reviewed. Emphasis is given to the influence that the host plant and the surface lipids of the host plant have upon insect herbivores and the predators and parasitoids of these herbivores. Variations in cuticular lipids of herbivorous insects are dependent upon the host plant, and these variations may affect the behavior of predators and parasitoids. The cuticular lipids of species which interact on multiple trophic levels are compared. Similarities were found between the hydrocarbons of herbivorous insects, their host plants, and their predators or parasitoids.     

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...     

Phytophagous insect fauna tracks host plant responses to exotic grass invasion

The high dependence of herbivorous insects on their host plants implies that plant invaders can affect these insects directly, by not providing a suitable habitat, or indirectly, by altering host plant availability. In this study, we sampled Asteraceae flower heads in cerrado remnants with varying levels of exotic grass invasion to evaluate whether invasive grasses have a direct effect on herbivore richness independent of the current disturbance level and host plant richness. By classifying herbivores according to the degree of host plant specialization, we also investigated whether invasive grasses reduce the uniqueness of the herbivorous assemblages. Herbivorous insect richness showed a unimodal relationship with invasive grass cover that was significantly explained only by way of the variation in host plant richness. The same result was found for polyphagous and oligophagous insects, but monophages showed a significant negative response to the intensity of the grass invasion that was independent of host plant richness. Our findings lend support to the hypothesis that the aggregate effect of invasive plants on herbivores tends to mirror the effects of invasive plants on host plants. In addition, exotic plants affect specialist insects differently from generalist insects; thus exotic plants affect not only the size but also the structural profile of herbivorous insect assemblages.     

Phytochemical defences and performance of specialist and generalist herbivores: a meta-analysis

1. Phytochemical coevolution theory, a long-standing paradigm in plant–insect interactions, predicts that specialist herbivores are less negatively affected by the allelochemicals of their host plants than are generalist herbivores. Although this theory is prevalent in plant–insect science, it is not always supported by empirical studies measuring the performance of specialist and generalist insects in response to allelochemicals. 2. The present study aimed to investigate: (i) whether there a difference between specialist and generalist performance in response to allelochemicals and (ii) whether the effect of allelochemicals on specialists and generalists depend upon allelochemical class or insect order. 3. A meta-analysis was conducted incorporating 76 effect sizes drawn from studies that directly compared the performance of specialist and generalist insects in response to treatment and control diets. Most of the effect sizes were related to the performance metric growth, the insect order Lepidoptera, and the allelochemical class nitrogen-containing compounds. 4. As predicted by phytochemical coevolution theory, specialist insects responded less negatively to allelochemicals of their hosts than generalist insects in terms of growth. There were no significant differences in terms of fecundity or survival, or among allelochemical classes or insect orders. 5. These results support the prediction of phytochemical coevolution theory that specialist insects respond less negatively to allelochemicals of their hosts than generalists, although only in terms of growth.     

植食性昆虫的嗅觉选食过程及其机制研究进展

植食性昆虫的嗅觉在其选择食物的过程中发挥了重要的作用,它能通过对植物挥发物的感受来定向和定位食物源并产生趋近行为,进而根据特殊的化合物或者多种化合物的特异浓度组合来区分寄主和非寄主植物.在这个过程中,昆虫嗅觉器官上相关的嗅觉感受蛋白被植物挥发物激活,形成特异的嗅觉感受通路,在行为上调控昆虫嗅觉选食的能力.本文主要从植食...     

Transcriptional profile and differential fitness in a specialist milkweed insect across host plants varying in toxicity

Interactions between plants and herbivorous insects have been models for theories of specialization and co‐evolution for over a century. Phytochemicals govern many aspects of these interactions and have fostered the evolution of adaptations by insects to tolerate or even specialize on plant defensive chemistry. While genomic approaches are providing new insights into the genes and mechanisms insect specialists employ to tolerate plant secondary metabolites, open questions remain about the evolution and conservation of insect counterdefences, how insects respond to the diversity defences mounted by their host plants, and the costs and benefits of resistance and tolerance to plant defences in natural ecological communities. Using a milkweed‐specialist aphid (Aphis nerii) model, we test the effects of host plant species with increased toxicity, likely driven primarily by increased secondary metabolites, on aphid life history traits and whole‐body gene expression. We show that more toxic plant species have a negative effect on aphid development and lifetime fecundity. When feeding on more toxic host plants with higher levels of secondary metabolites, aphids regulate a narrow, targeted set of genes, including those involved in canonical detoxification processes (e.g., cytochrome P450s, hydrolases, UDP‐glucuronosyltransferases and ABC transporters). These results indicate that A. nerii marshal a variety of metabolic detoxification mechanisms to circumvent milkweed toxicity and facilitate host plant specialization, yet, despite these detoxification mechanisms, aphids experience reduced fitness when feeding on more toxic host plants. Disentangling how specialist insects respond to challenging host plants is a pivotal step in understanding the evolution of specialized diet breadths.     

Evolutionary dynamics of specialisation in herbivorous stick insects

Understanding the evolutionary dynamics underlying herbivorous insect mega‐diversity requires investigating the ability of insects to shift and adapt to different host plants. Feeding experiments with nine related stick insect species revealed that insects retain the ability to use ancestral host plants after shifting to novel hosts, with host plant shifts generating fundamental feeding niche expansions. These expansions were, however, not accompanied by expansions of the realised feeding niches, as species on novel hosts are generally ecologically specialised. For shifts from angiosperm to chemically challenging conifer hosts, generalist fundamental feeding niches even evolved jointly with strong host plant specialisation, indicating that host plant specialisation is not driven by constraints imposed by plant chemistry. By coupling analyses of plant chemical compounds, fundamental and ecological feeding niches in multiple insect species, we provide novel insights into the evolutionary dynamics of host range expansion and contraction in herbivorous insects.     

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.     

Does host‐plant diversity explain species richness in insects? A test using Coccidae (Hemiptera)

1. The megadiverse herbivores and their host plants are a major component of biodiversity, and their interactions have been hypothesised to drive the diversification of both. 2. If plant diversity influences the diversity of insects, there is an expectation that insect species richness will be strongly correlated with host‐plant species richness. This should be observable at two levels (i) more diverse host‐plant groups should harbour more species of insects, and (ii) the species richness of a group of insects should correlate with the richness of the host groups it uses. However, such a correlation is also consistent with a hypothesis of random host use, in which insects encounter and use hosts in proportion to the diversity of host plants. Neither of these expectations has been widely tested. 3. These expectations were tested using data from a species‐rich group of insects – the Coccidae (Hemiptera). 4. Significant positive correlations were found between the species richness of coccid clades (genera) and the species richness of the host‐plant family or families upon which the clades occur. On a global scale, more closely related plant families have more similar communities of coccid genera but the correlation is weak. 5. Random host use could not be rejected for many coccids but randomisation tests and similarity of coccid communities on closely related plant families show that there is non‐random host use in some taxa. Overall, our results support the idea that plant diversity is a driver of species richness of herbivorous insects, probably via escape‐and‐radiate or oscillation‐type processes.     

Does ant predation favour leaf beetle specialization on toxic host plants?

Generalist predators are frequently seen as evolutionary forces that narrow the host range in herbivorous insects. Predators may favour specialization of herbivores on host plants containing toxic chemicals (which can be used by herbivores for their own defence) if host plant‐derived defences provide better protection from enemies than do autogenously produced defences. We compared the effectiveness of these two defensive strategies in the larvae of six species of leaf beetle (Chrysomelidae) against wood ants (Formica rufa group) in field experiments. Ants were more strongly repelled by larvae with host plant‐derived, salicylaldehyde‐containing secretions than by larvae with various autogenous secretions, but collectively foraging ants ultimately overcame any type of chemical defence by social interactions, chemical signalling, and olfactory learning. As a result, ants killed all larvae of Chrysomela lapponica defended by salicylaldehyde‐containing secretions within 2 days of their introduction to willows within 15 m of ant nests. We conclude that in the field neither type of chemical defence provides complete protection against wood ants in the vicinity of their nests, and that evolutionary shifts from autogenous production of secretion to sequestration of plant allelochemicals in leaf beetles may be favoured mostly at low ant densities on the periphery of ant foraging areas.     

用现代分子生物学方法揭示植物与昆虫的相互关系

昆虫与植物之间相互关系的研究由来已久。1 964年Ｅｈｒｌｉｃｈ和Ｒａｖｅｎ提出了协同进化 (ｃｏｅｖｏ ｌｕｔｉｏｎ) [1 ] 的概念后 ,更大大促进了植物与昆虫相互关系的研究。马世骏高度重视这一领域的研究 ,指出协同进化的观点是研究生物进化的方法论的基础之一[2 ] 。钦俊德对昆虫与植物的关系作了系统的论述和总结[3] 。以往研究昆虫与植物之间的相互关系 ,主要依靠分析昆虫的习性与相关植物的外部形态和内含次生物质之间的关系 ,辅以数学手段及计算机工具来进行 ,虽然取得了大量卓有成效的成果 ,但都还未能直接从绝大多数生物的遗传…     

The influence of host and non‐host companion plants on the behaviour of pest insects in field crops

Companion plants grown as ‘trap crops’ or ‘intercrops’ can be used to reduce insect infestations in field crops. The ways in which such reductions are achieved are being described currently using either a chemical approach, based on the ‘push‐pull strategy’, or a biological approach, based on the ‘appropriate/inappropriate landing theory’. The chemical approach suggests that insect numbers are reduced by chemicals from the intercrop ‘repelling’ insects from the main crop, and by chemicals from the trap‐crop ‘attracting’ insects away from the main crop. This approach is based on the assumptions that (1) plants release detectable amounts of volatile chemicals, and (2) insects ‘respond’ while still some distance away from the emitting plant. We discuss whether the above assumptions can be justified using the ‘appropriate/inappropriate landing theory’. Our tenet is that specialist insects respond only to the volatile chemicals released by their host plants and that these are released in such small quantities that, even with a heightened response to such chemicals, specialist insects can only detect them when a few metres from the emitting plant. We can find no robust evidence in the literature that plant chemicals ‘attract’ insects from more than 5 m and believe that ‘trap crops’ function simply as ‘interception barriers’. We can also find no evidence that insects are ‘repelled’ from landing on non‐host plants. Instead, we believe that ‘intercrops’ disrupt host‐plant finding by providing insects with a choice of host (appropriate) and non‐host (inappropriate) plant leaves on which to land, as our research has shown that, for intercropping to be effective, insects must land on the non‐host plants. Work is needed to determine whether non‐host plants are repellent (chemical approach) or ‘non‐stimulating’ (biological approach) to insects.     

Glutathione S-transferases in the adaptation to plant secondary metabolites in the Myzus persicae aphid

Glutathione S-transferases (GST) in insects play an important role in the detoxification of many substances including allelochemicals from plants. Induction of GST activity in Myzus persicae in response to secondary metabolites from Brassica plants was determined using different host plant species and confirmed using artificial diet with pure allelochemicals added. The 2,4-dinitro-1-iodobenzene (DNIB) was found to be a useful substrate for identifying particular GSTs in insects. GSTs from M. persicae were purified using different affinity chromatography columns and related kinetic parameters were calculated. GST isoenzymes were characterised using electrophoretic methods. Although SDS-PAGE results indicated similarity among the purified enzymes from each affinity column, biochemical studies indicated significant differences in kinetic parameters. Finally, the GST pattern of M. persicae was discussed in terms of insect adaptation to the presence of plant secondary substances such as the glucosinolates and the isothiocyanates, from Brassicaceae host plants.     

Glucosinolates in oilseed rape: secondary metabolites that influence interactions with herbivores and their natural enemies

Glucosinolates are sulphur‐containing secondary metabolites characteristic of Brassicaceous plants. Glucosinolate breakdown products, which include isothiocyanates, are released following tissue damage when hydrolytic enzymes act on them. The isothiocyanates have toxic effects on generalist herbivores when they attempt to feed on oilseed rape, Brassica napus, and also function as repellents. However, specialist herbivores such as Brevicoryne brassicae aphids, flea beetles, Psylliodes chrysocephala and the Lepidopteran pest, Pieris rapae, are adapted to the presence of glucosinolates and thrive on plants containing them. They may do this by avoiding tissue damage to prevent the formation of isothiocyanates or by metabolising or tolerating glucosinolates. For many specialist herbivores, the isothiocyanates function as attractants and glucosinolates can even be sequestered for defence against predatory insects. Thus, these herbivores have evolved resistance to host‐plant secondary metabolites and this type of evolutionary history may have given some insects an enhanced ability to adapt to xenobiotics. In an agricultural context, this may make pests better able to evolve resistance to artificially applied pesticides. The effect of increased glucosinolate content in making oilseed rape cultivars more susceptible to specialist pests was highlighted in a seminal article in the Annals of Applied Biology in 1995. This review of the literature considers developments in this area since then.     

媒介昆虫-病毒-植物互作对生物入侵的影响

媒介昆虫-病毒-植物互作关系复杂多样。虽然相关的研究较多, 然而有关三者互作对于生物入侵的影响还知之甚少。已有证据表明, 寄主植物对病毒的敏感性和对媒介昆虫的适合性、媒介昆虫对寄主的适应能力等因素影响三者互作关系。当寄主植物易感病并且对媒介昆虫的适合性低, 而媒介昆虫对寄主植物的适应能力强时, 媒介昆虫与植物病毒之间很可能建立间接互惠关系, 这种互惠可促进媒介昆虫入侵和病毒病流行。此外, 媒介昆虫与植物病毒之间中性或偏害的互作关系对于外来生物入侵的促进作用也不容忽视。鉴于三者互作对于生物入侵的重要性, 今后需要对不同物种所组成的多种组合进行比较研究, 并采用多种方法揭示互作的生理和分子机制。     

Conserved oviposition preferences in alpine leaf beetle populations despite host shifts and isolation

Abstract.  1. Choosing the plant on which to lay their eggs is the last act of care that most female herbivorous insects bestow upon their offspring. These decisions play a pivotal role in insect–plant interactions, placing host preference under strong selection and contributing to the diversity of phytophagous insects as one of the first traits to adapt to new hosts.
2. This study presents a test of whether extreme isolation and exposure to different host plants can produce intra-specific divergence in oviposition preference in alpine insects. Geographic variation should impose selection to fine-tune host plant ranking and specificity to the plants normally encountered, to avoid wasting time during the very limited reproductive season experienced at high altitudes.
3. Beetles from five populations of Oreina elongata differing in host availability were offered three natural hosts: Cirsium spinosissimum , Adenostyles alliariae , and Adenostyles glabra . A novel application of a continuation ratio model (logistic regression) was made to sequential no-choice experiments, combined with quasi-likelihood analysis of multiple-choice experiments.
4. The results show little geographic variation in host plant choice: all populations strongly preferred Cirsium in multiple-choice trials, and in no-choice experiments laid around 47% of their remaining eggs during each stage, almost regardless of the host present.
5. Enemy-free space seems to explain the preference for Cirsium , but isolation and exposure to different plants has clearly not caused local adaptation in host plant ranking or specificity. Reasons for this conservatism despite divergence in other characteristics are discussed.     

Dietary proteinase inhibitors alter complement of midgut proteases

Plant serine proteinase inhibitors (Pls) have the potential to restrict the growth and/or development of herbivorous insects. However, there are limitations to the efficacy of these Pls. An insect's susceptibility to a Pl is determined, at least in part, by the relative proportion of proteolytic enzyme activity in the midgut that is suppressed by that inhibitor. Insects adapt to dietary trypsin inhibitor in their host plant by secreting “inhibitor-resistant” trypsin(s). These “inhibitor-resistant” enzyme(s) may be the standard proteinase(s) secreted into the midgut (e.g., Pieris rapae), or may be enhanced following ingestion of proteinase inhibitor (e.g., Helicoverpa zea). In addition, insects may be pre-adapted to specific Pl(s), following adaptation to a Pl from the same family. For example, Pieris rapae is a crucifer specialist that is resistant to cabbage Pl, but is also resistant to Kunitz soybean trypsin inhibitor, a Pl in the same family as cabbage Pl, but from a non-host plant. The ultimate value of this pre-adaptation to herbivory by a species of insect will be determined by the number of different families of Pl in host plant(s) to which the species has adapted, and the distribution of those families among other species of plants. Thus, it is possible that the presence of a plant Pl limits herbivory by insect(s). However, multiple inhibitors, matched to the complement of enzymes in the insect's midgut, may be required to enhance this resistance of plants to herbivorous insects. © 1996 Wiley-Liss, Inc.     

Phylogenetic test of speciation by host shift in leaf cone moths (Caloptilia) feeding on maples (Acer)

The traditional explanation for the exceptional diversity of herbivorous insects emphasizes host shift as the major driver of speciation. However, phylogenetic studies have often demonstrated widespread host plant conservatism by insect herbivores, calling into question the prevalence of speciation by host shift to distantly related plants. A limitation of previous phylogenetic studies is that host plants were defined at the family or genus level; thus, it was unclear whether host shifts predominate at a finer taxonomic scale. The lack of a statistical approach to test the hypothesis of host‐shift‐driven speciation also hindered studies at the species level. Here, we analyze the radiation of leaf cone moths (Caloptilia) associated with maples (Acer) using a newly developed, phylogeny‐based method that tests the role of host shift in speciation. This method has the advantage of not requiring complete taxon sampling from an entire radiation. Based on 254 host plant records for 14 Caloptilia species collected at 73 sites in Japan, we show that major dietary changes are more concentrated toward the root of the phylogeny, with host shift playing a minor role in recent speciation. We suggest that there may be other roles for host shift in promoting herbivorous insect diversification rather than facilitating speciation per se.     

不同寄主植物叶片上蚜虫的形态适应

昆虫对寄主植物之间的形态适应性是两者相互关系的重要组成部分, 本文以植食性蚜虫为研究对象, 研究其对不同种类寄主植物叶片的形态适应。选取寄生于禾本科、 杨柳科、 壳斗科和松科4科7属植物叶片上的26种蚜虫, 基于有翅孤雌蚜和无翅孤雌蚜, 在光学显微镜下观察、 测量并统计比较了37个形态特征及其种内变异。以克隆平均值为基础数据,选取与蚜虫取食和附着于植物表面相关的形态特征（喙末端、跗节Ⅰ、Ⅱ和爪）为变量,分别得出无翅、 有翅孤雌蚜和所有蚜型3个聚类分支图,并将各种蚜虫的寄主植物科、 属分别对应到分支图上。结果表明：体型、腹管和触角的形态在不同蚜虫的科间有一定差别, 喙末端、 跗节Ⅰ、跗节Ⅱ及爪的形态则在不同的寄主植物间存在差异; 在蚜虫种内各形态特征存在一定变异, 变异系数的范围为1.89%~26.08%。3个聚类分析的结果显示,不同种类蚜虫形成的分支基本对应不同科、属的寄主植物;而杨一条角蚜Doraphis populi、三堡瘿绵蚜Epipemphigus imaicus和杨柄叶瘿绵蚜Pemphigus matsumurai则处在较为特殊的分支位置, 那是因为前者为次生寄主型, 后两者均能产生虫瘿。研究结果表明寄生于不同植物相同部位上的蚜虫存在形态特征的明显分异, 对应不同的寄主植物, 蚜虫与寄主植物之间存在着形态适应关系。     

寄主植物影响害虫药剂敏感性的研究进展

害虫取食不同寄主植物后,对杀虫剂的反应可归为3类：敏感性降低、增高和无明显变化。害虫对药剂的敏感性变化与不同植物中次生物质诱导激活/抑制昆虫体内相关解毒酶活性有关。这种诱导作用可受到植物营养、次生物质种类及其含量分布、害虫种类与发育阶段、以及环境温度等多种因素影响。经诱导的昆虫解毒酶对不同类型杀虫剂的代谢能力并不相同,进而导致对不同药剂的敏感性变化有明显差异。解毒酶系的诱导激活在害虫抗药性形成早期被认为有利于提高隐性抗性基因频率,从而可促进害虫抗药性的发展。最后,就寄主植物影响害虫对药剂敏感性在害虫治理中的应用作了探讨。