Plant-mediated effects in the Brassicaceae on the performance and behaviour of parasitoids

Direct and indirect plant defences are well studied, particularly in the Brassicaceae. Glucosinolates (GS) are secondary plant compounds characteristic in this plant family. They play an important role in defence against herbivores and pathogens. Insect herbivores that are specialists on brassicaceous plant species have evolved adaptations to excrete or detoxify GS. Other insect herbivores may even sequester GS and employ them as defence against their own antagonists, such as predators. Moreover, high levels of GS in the food plants of non-sequestering herbivores can negatively affect the growth and survival of their parasitoids. In addition to allelochemicals, plants produce volatile chemicals when damaged by herbivores. These herbivore induced plant volatiles (HIPV) have been demonstrated to play an important role in foraging behaviour of insect parasitoids. In addition, biosynthetic pathways involved in the production of HIPV are being unraveled using the model plant Arabidopsis thialiana. However, the majority of studies investigating the attractiveness of HIPV to parasitoids are based on experiments mainly using crop plant species in which defence traits may have changed through artificial selection. Field studies with both cultivated and wild crucifers, the latter in which defence traits are intact, are necessary to reveal the relative importance of direct and indirect plant defence strategies on parasitoid and plant fitness. Future research should also consider the potential conflict between direct and indirect plant defences when studying the evolution of plant defences against insect herbivory.     

A meta‐analysis of insect pest behavioral manipulation with plant volatiles

Many insect pests utilize plant volatiles for host location and untangling the mechanisms of this process can provide tools for pest management. Numerous experimental results have been published on the effect of plant volatiles on insect pests. We used a meta‐analysis to summarize this knowledge and to look for patterns. Our goal was to identify herbivore and plant traits that might explain the herbivores’ behavioral response to plant volatiles in field applications. We scored a total of 374 unique plant volatile‐insect herbivore interactions obtained from 34 published studies investigating 50 herbivore pest species. Attractants had a significant effect on insect herbivore abundance but repellents did not; this latter result could be a result of the comparatively small number of field studies that tested plant volatiles as repellents (3%). Females were significantly more attracted to plant volatile baits than males. The diet breadth of herbivores was independent of a behavioral response to plant volatiles, but more case studies show effects of volatiles on chewers, followed by wood‐borers and sap‐feeders. There are more demonstrations of attraction to plant volatiles in Lepidoptera than in Thysanoptera. The method of plant volatile application had a significant effect on herbivore abundance and increasing the number of chemicals in individual baits attracted more herbivores. The magnitude of the response of herbivores to plant volatiles in forest and agricultural habitats was similar. We explore consistent patterns and highlight areas needing research in using plant volatiles to manage insect pests.     

Host plant scents attract rolled-leaf beetles to Neotropical gingers in a Central American tropical rain forest

Leaf volatile chemicals are known to reduce herbivory rates by repelling or intoxicating insect herbivores and by attracting the predators and parasitoids of herbivores. However, leaf volatiles may also be used by insect herbivores as cues to locate their host plants. Leaf volatiles are suggested to be important host search cues for herbivores in structurally complex and diverse habitats, such as tropical rain forests. A group of insect herbivores, the rolled-leaf beetles (Coleoptera: Chrysomelidae: Hispinae), have maintained a highly specialized interaction with Neotropical gingers (Zingiberales) for ca. 60 million years. In this study, we explored chemical attraction to host plants under controlled laboratory conditions, using four sympatric rolled-leaf beetle species, Cephaloleia dorsalis Baly, Cephaloleia erichsonii Baly, Cephaloleia fenestrata Weise, and Cephaloleia placida Baly. For each beetle species, we investigated (i) whether it was repelled or attracted by leaf scents produced by four host and four non-host plant species, including Neotropical gingers in the families Marantaceae, Costaceae, and Zingiberaceae; and (ii) its ability to use scents to detect its host plant. We found that rolled-leaf beetles can detect and are attracted by leaf volatiles from both host and non-host gingers. Additionally, when beetles were simultaneously exposed to leaf volatiles from host and non-host plants, three rolled-leaf beetle species were significantly more attracted by volatiles from their host plants than from non-hosts. Only one of the beetle species was not able to discriminate between host and non-host scents.     

Multitrophic effects of herbivore-induced plant volatiles in an evolutionary context

Herbivorous and carnivorous arthropods use plant volatiles when foraging for food. In response to herbivory, plants emit a blend that may be quantitatively and qualitatively different from the blend emitted when intact. This induced volatile blend alters the interactions of the plant with its environment. We review recent developments regarding the induction mechanism as well as the ecological consequences in a multitrophic and evolutionary context. It has been well established that carnivores (predators and parasitoids) are attracted by the volatiles induced by their herbivorous victims. This concerns an active plant response. In the case of attraction of predators, this is likely to result in a fitness benefit to the plant, because through consumption a predator removes the herbivores from the plant. However, the benefit to the plant is less clear when parasitoids are attracted, because parasitisation does usually not result in an instantaneous or in a complete termination of consumption by the herbivore. Recently, empirical evidence has been obtained that shows that the plant's response can increase plant fitness, in terms of seed production, due to a reduced consumption rate of parasitized herbivores. However, apart from a benefit from attracting carnivores, the induced volatiles can have a serious cost because there is an increasing number of studies that show that herbivores can be attracted. However, this does not necessarily result in settlement of the herbivores on the emitting plant. The presence of cues from herbivores and/or carnivores that indicate that the plant is a competitor- and/or enemy-dense space, may lead to an avoidance response. Thus, the benefit of emission of induced volatiles is likely to depend on the prevailing faunal composition. Whether plants can adjust their response and influence the emission of the induced volatiles, taking the prevalent environmental conditions into account, is an interesting question that needs to be addressed. The induced volatiles may also affect interactions of the emitting plant with its neighbours, e.g., through altered competitive ability or by the neighbour exploiting the emitted information.Major questions to be addressed in this research field comprise mechanistic aspects, such as the identification of the minimally effective blend of volatiles that explains the attraction of carnivores to herbivore-infested plants, and evolutionary aspects such as the fitness consequences of induced volatiles. The elucidation of mechanistic aspects is important for addressing ecological and evolutionary questions. For instance, an important tool to address ecological and evolutionary aspects would be to have plant pairs that differ in only a single trait. Such plants are likely to become available in the near future as a result of mechanistic studies on signal-transduction pathways and an increased interest in molecular genetics.     

The major volatile organic compound emitted from Arabidopsis thaliana flowers, the sesquiterpene (E)-β-caryophyllene, is a defense against a bacterial pathogen

Flowers have a high risk of pathogen attack because of their rich nutrient and moisture content, and high frequency of insect visitors. We investigated the role of (E)-β-caryophyllene in floral defense against a microbial pathogen. This sesquiterpene is a common volatile compound emitted from flowers, and is a major volatile released from the stigma of Arabidopsis thaliana flowers. Arabidopsis thaliana lines lacking a functional (E)-β-caryophyllene synthase or constitutively overexpressing this gene were challenged with Pseudomonas syringae pv. tomato DC3000, which is a bacterial pathogen of brassicaceous plants. Flowers of plant lines lacking (E)-β-caryophyllene emission showed greater bacterial growth on their stigmas than did wild-type flowers, and their seeds were lighter and misshapen. By contrast, plant lines with ectopic (E)-β-caryophyllene emission from vegetative parts were more resistant than wild-type plants to pathogen infection of leaves, and showed reduced cell damage and higher seed production. Based on in vitro experiments, (E)-β-caryophyllene seems to act by direct inhibition of bacterial growth, rather than by triggering defense signaling pathways. (E)-β-Caryophyllene thus appears to serve as a defense against pathogens that invade floral tissues and, like other floral volatiles, may play multiple roles in defense and pollinator attraction.     

Host plants alter their volatiles to help a solitary egg parasitoid distinguish habitats with parasitized hosts from those without

When attacked by herbivores, plants emit volatiles to attract parasitoids and predators of herbivores. However, our understanding of the effect of plant volatiles on the subsequent behaviour of conspecific parasitoids when herbivores on plants are parasitized is limited. In this study, rice plants were infested with gravid females of the brown planthopper (BPH) Nilaparvata lugens for 24 hr followed by another 24 hr in which the BPH eggs on plants were permitted to be parasitized by their egg parasitoid, Anagrus nilaparvatae; volatiles from rice plants that underwent such treatment were less attractive to subsequent conspecific parasitoids compared to the volatiles from plants infested with gravid BPH females alone. Chemical analysis revealed that levels of JA and JA-Ile as well as of four volatile compounds—linalool, MeSA, α-zingiberene and an unknown compound—from plants infested with BPH and parasitized by wasps were significantly higher than levels of these compounds from BPH-infested plants. Laboratory and field bioassays revealed that one of the four increased chemicals—α-zingiberene—reduced the plant's attractiveness to the parasitoid. These results suggest that host plants can fine-tune their volatiles to help egg parasitoids distinguish host habitats with parasitized hosts from those without.     

Damage to sagebrush attracts predators but this does not reduce herbivory

Emissions of volatiles increase following herbivory from many plant species and volatiles may serve multiple functions. Herbivore‐induced volatiles attract predators and parasitoids of herbivores and are often assumed to benefit plants by facilitating top‐down control of herbivores; this benefit of induced emissions has been tested only a few times. Volatile compounds released by experimentally clipped sagebrush shoots have been shown to reduce levels of chewing damage experienced by other shoots on the same plant and on neighboring sagebrush plants. In this study, I asked whether experimental clipping attracted predators of herbivorous insects to sagebrush shoots. I also evaluated aphid populations and chewing damage on clipped and unclipped shoots and whether predators were likely to have caused differences in aphids and chewing damage. Shoots that had been clipped recruited more generalist predators, particularly coccinellids and Geocoris spp. in visual surveys conducted during two seasons. Clipping also caused increased numbers of parasitized aphids in one season. Ants were common tending aphids but were not significantly affected by clipping. Despite the increase in generalist predators, clipped plants were more likely to support populations of aphids that increased during both seasons compared to aphids on unclipped control plants. Clipped shoots suffered less damage by chewing herbivores in the 1‐year in which this was measured. Chewing damage was not correlated with numbers of predators. These results suggest that predators and parasitoids were attracted to experimentally clipped sagebrush plants but that these predators were not effective at reducing net damage to the plant. This conclusion is not surprising as much of the herbivory is inflicted by grasshoppers and deer, herbivores that are not vulnerable to the predators attracted to sagebrush volatiles. More generally, it should not be assumed that predators that are attracted by herbivore‐induced volatiles necessarily benefit the plant without testing this hypothesis under field conditions.     

Oviposition by a moth suppresses constitutive and herbivore-induced plant volatiles in maize

Plant volatiles function as important signals for herbivores, parasitoids, predators, and neighboring plants. Herbivore attack can dramatically increase plant volatile emissions in many species. However, plants do not only react to herbivore-inflicted damage, but also already start adjusting their metabolism upon egg deposition by insects. Several studies have found evidence that egg deposition itself can induce the release of volatiles, but little is known about the effects of oviposition on the volatiles released in response to subsequent herbivory. To study this we measured the effect of oviposition by Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae) moths on constitutive and herbivore-induced volatiles in maize (Zea mays L.). Results demonstrate that egg deposition reduces the constitutive emission of volatiles and suppresses the typical burst of inducible volatiles following mechanical damage and application of caterpillar regurgitant, a treatment that mimics herbivory. We discuss the possible mechanisms responsible for reducing the plant’s signaling capacity triggered by S. frugiperda oviposition and how suppression of volatile organic compounds can influence the interaction between the plant, the herbivore, and other organisms in its environment. Future studies should consider oviposition as a potential modulator of plant responses to insect herbivores.     

Herbivore-induced plant volatiles mediate host selection by a root herbivore

In response to herbivore attack, plants mobilize chemical defenses and release distinct bouquets of volatiles. Aboveground herbivores are known to use changes in leaf volatile patterns to make foraging decisions, but it remains unclear whether belowground herbivores also use volatiles to select suitable host plants. We therefore investigated how above- and belowground infestation affects the performance of the root feeder Diabrotica virgifera virgifera, and whether the larvae of this specialized beetle are able to use volatile cues to assess from a distance whether a potential host plant is already under herbivore attack. Diabrotica virgifera larvae showed stronger growth on roots previously attacked by conspecific larvae, but performed more poorly on roots of plants whose leaves had been attacked by larvae of the moth Spodoptera littoralis. Fittingly, D. virgifera larvae were attracted to plants that were infested with conspecifics, whereas they avoided plants that were attacked by S. littoralis. We identified (E)-β-caryophyllene, which is induced by D. virgifera, and ethylene, which is suppressed by S. littoralis, as two signals used by D. virgifera larvae to locate plants that are most suitable for their development. Our study demonstrates that soil-dwelling insects can use herbivore-induced changes in root volatile emissions to identify suitable host plants.     

Subtile Verteidigungsstrategien in Pflanzen

Scents of survival: Subtle defense strategies in plants Plants are master chemists who synthesize an arsenal of compounds which efficiently defend against herbivore attack. In addition to chemicals which directly affect herbivores, attacked plants release characteristic bouquets of low molecular weight volatile compounds, mostly terpenes and fatty acid derivatives, into their environment. These volatiles serve as signals which can attract predators and parasitoids to attack herbivores, thus indirectly defending the plant. Volatiles may also be perceived by remote parts of the same plant, which can then prepare to defend themselves against imminent attack, and thus react more quickly when attacked. These natural phenomena suggest alternative strategies for agricultural pest management.     

Hyperparasitoids Use Herbivore-Induced Plant Volatiles to Locate Their Parasitoid Host

Plants respond to herbivory with the emission of induced plant volatiles. These volatiles may attract parasitic wasps (parasitoids) that attack the herbivores. Although in this sense the emission of volatiles has been hypothesized to be beneficial to the plant, it is still debated whether this is also the case under natural conditions because other organisms such as herbivores also respond to the emitted volatiles. One important group of organisms, the enemies of parasitoids, hyperparasitoids, has not been included in this debate because little is known about their foraging behaviour. Here, we address whether hyperparasitoids use herbivore-induced plant volatiles to locate their host. We show that hyperparasitoids find their victims through herbivore-induced plant volatiles emitted in response to attack by caterpillars that in turn had been parasitized by primary parasitoids. Moreover, only one of two species of parasitoids affected herbivore-induced plant volatiles resulting in the attraction of more hyperparasitoids than volatiles from plants damaged by healthy caterpillars. This resulted in higher levels of hyperparasitism of the parasitoid that indirectly gave away its presence through its effect on plant odours induced by its caterpillar host. Here, we provide evidence for a role of compounds in the oral secretion of parasitized caterpillars that induce these changes in plant volatile emission. Our results demonstrate that the effects of herbivore-induced plant volatiles should be placed in a community-wide perspective that includes species in the fourth trophic level to improve our understanding of the ecological functions of volatile release by plants. Furthermore, these findings suggest that the impact of species in the fourth trophic level should also be considered when developing Integrated Pest Management strategies aimed at optimizing the control of insect pests using parasitoids.     

Minor effects of two elicitors of insect and pathogen resistance on volatile emissions and parasitism of Spodoptera frugiperda in Mexican maize fields

Synthetic elicitors can be used to induce resistance in plants against pathogens and arthropod herbivores. Such compounds may also change the emission of herbivore-induced plant volatiles, which serve as important cues for parasitic wasps to locate their hosts. Therefore, the use of elicitors in the field may affect biological control of insect pests. To test this, we treated maize seedlings growing in a subtropical field in Mexico with methyl jasmonate (MeJA), an elicitor of defense responses against many insects, and benzo-(1,2,3)-thiadiazole-7-carbothioic acid S-methyl ester (BTH), an elicitor of resistance against certain pathogens. Volatile emission, herbivore infestation, pathogen infection, and plant performance (growth and grain yield) of treated and untreated maize plants were measured. Application of BTH slightly reduced volatile emission in maize, while MeJA increased the emission compared to control treatments. Despite the apparent changes in volatile emissions, the elicitor application did not consistently affect infestation by Spodoptera frugiperda larvae, the main insect pest found on the maize seedlings, and had only marginal effects on parasitism rates. Similarly, there were no treatment effects on infestation by other herbivores and pathogens. Results for the six replications that stretched over one summer and one winter season were highly variable, with parasitism rates and the species composition of the parasitoids differing significantly between seasons. This variability, as well as the severe biotic and abiotic stresses on young seedlings might explain why we measured only slight effects of elicitor application on pest incidence and biological control in this specific field study. Indeed, an additional field experiment under milder and more standardized conditions revealed that BTH induced significant resistance against Bipolaris maydis, a major pathogen in the experimental maize fields. Similar affects can be expected for herbivory and parasitism rates.     

二化螟盘绒茧蜂及稻虱缨小蜂对挥发物的嗅觉反应

【目的】揭示二化螟盘绒茧蜂Apanteles chilonis Munakata与稻虱缨小蜂Anagrus nilaparvatae对单一水稻挥发物组分的选择行为,了解水稻挥发物在其寄主搜寻过程中的作用,为水稻害虫寄生蜂引诱剂的研发提供技术基础。【方法】利用Y型嗅觉仪测定了二化螟盘绒茧蜂与稻虱缨小蜂对11种水稻挥发物组分(反-2-己烯醛、α-蒎烯、乙酸叶醇酯、芳樟醇、β-石竹烯、反-3-己烯醛、水杨酸甲酯、罗勒烯、苯甲醛、2-壬酮、柠檬烯)的嗅觉行为反应。【结果】β-石竹烯(10、50μg/kg)、罗勒烯(0.1、1、10μg/kg)、2-壬酮(10μg/kg)及反-2-己烯醛(50μg/kg)对二化螟盘绒茧蜂行为具有显著影响;稻虱缨小蜂对反-2-己烯醛(50μg/kg)、反-3-己烯醛(1、10、50μg/kg)、芳樟醇(0.1、1、10μg/kg)、β-石竹烯(0.1、50μg/kg)、罗勒烯(1μg/kg)、α-蒎烯(1μg/kg)和水杨酸甲酯(0.1、1、10、50μg/kg)有显著的嗅觉反应。【结论】寄生蜂对水稻挥发性气味的识别与挥发物的种类和浓度有很大关系,研发水稻害虫寄生蜂的引诱剂时需考虑挥发物的种类与浓度。     

Olfactory cues explain differential attraction of the striped rice stem borer to different varieties of rice plant

In Iranian rice fields, different varieties of rice are cultivated which are differentially impacted by females of the rice stem borer, Chilo suppressalis. To elucidate the role odours may play in their host plant finding behaviour, female Ch. suppressalis were exposed to four varieties of rice plants and their volatiles in a four‐arm olfactometer. In whole plant tests, Ch. suppressalis were significantly attracted to the variety previously characterized as most susceptible, least attracted to one characterized as semisusceptible, and showed no attraction to those varieties characterized as semi‐ and highly resistant. Tests using headspace volatile extracts yielded similar results in the case of the most susceptible variety, but showed no attraction to the semisusceptible and highly resistant varieties, and low attraction to the semiresistant variety. Subsequent analysis of the volatile composition identified a panel of 27 components, some of which were either unique to, or abundantly present in, particular varieties, and may explain the observed variation in their attractiveness. Our findings show that rice plant volatiles can play a role in the host selection behaviour of this pest species, and we suggest compounds which may be important to this process and the future application of volatiles in rice pest management programs.     

Caterpillars of Euphydryas aurinia (Lepidoptera: Nymphalidae) feeding on Succisa pratensis leaves induce large foliar emissions of methanol

A major new discovery made in the last decade is that plants commonly emit large amounts and varieties of volatiles after damage inflicted by herbivores, and not merely from the site of injury. However, analytical methods for measuring herbivore-induced volatiles do not usually monitor the whole range of these compounds and are complicated by the transient nature of their formation and by their chemical instability. Here we present the results of using a fast and highly sensitive proton transfer reaction-mass spectrometry (PTR-MS) technique that allows simultaneous on-line monitoring of leaf volatiles in the pptv (pmol mol(-1)) range. The resulting on-line mass scans revealed that Euphydryas aurinia caterpillars feeding on Succisa pratensis leaves induced emissions of huge amounts of methanol--a biogeochemically active compound and a significant component of the volatile organic carbon found in the atmosphere--and other immediate, late and systemic volatile blends (including monoterpenes, sesquiterpenes and lipoxygenase-derived volatile compounds). In addition to influencing neighboring plants, as well as herbivores and their predators and parasitoids, these large emissions might affect atmospheric chemistry and physics if they are found to be generalized in other plant species.     

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.     

Shared signals -'alarm calls' from plants increase apparency to herbivores and their enemies in nature

The attraction of natural enemies of herbivores by volatile organic compounds as an induced indirect defence has been studied in several plant systems. The evidence for their defensive function originates mainly from laboratory studies with trained parasitoids and predators; the defensive function of these emissions for plants in natural settings has been rarely demonstrated. In native populations and laboratory Y-tube choice experiments with transgenic Nicotiana attenuata plants unable to release particular volatiles, we demonstrate that predatory bugs use terpenoids and green leaf volatiles (GLVs) to locate their prey on herbivore-attacked plants. By attracting predators with volatile signals, this native plant reduces its herbivore load – demonstrating the defensive function of herbivore-induced volatile emissions. However, plants producing GLVs are also damaged more by flea beetles. The implications of these conflicting ecological effects for the evolution of induced volatile emissions and for the development of sustainable agricultural practices are discussed.     

Volatiles from host plant brinjal attract the brinjal Fruit and Shoot Borer -Leucinodes orbonalis Guenee

Brinjal Fruit and Shoot Borer- Leucinodes orbonalis Guenee is a major insect pest on brinjal- Solanum melongena worldwide. An effective strategy used in developing pest controlling agents is the synergism between insect pheromones and host plant volatiles, which can increase the attraction of insect pest. The present study was aimed at investigating the chemical constituents and attractant effects of the volatiles extracted from different parts of the host plant brinjal on the behavior of adult L. orbonalis. Bioassay using Y-shaped olfactometer revealed that the one-day old virgin female, gravid female and male insects respond positively to the host plant volatiles extracted from fruits, leaves and shoots but not to that of flowers. It was shown that the gravid females were significantly attracted to all three volatiles (p < 0.05). Bioassay using X-shaped olfactometer identified that all three types of insects highly preferred the volatiles from fruits (p < 0.05). Gas chromatography-mass spectrometry analysis of volatiles indicated that brinjal plant produces volatile secondary metabolites, which include 2,2′-(Ethane-1,2-diylbis(oxy))bis(ethane-2,1-diyl) dibenzoate (12.11%), 3,7-dimethylocta-1,6-dien-3-ol (22.38%), Benzyl alcohol (22.9%) and Benzyl alcohol (27.06%) as major constituents from fruits, shoots, leaves and flowers respectively. Responses of insects to the volatiles from host plant in the absence of visual cues direct us to focus on the importance of host plant volatiles to locate the plant. Results of this study emphasize the major role that host plant volatiles play in the attraction of insect pests towards the plant.     

Timing of induced volatile emissions in maize seedlings

Maize (Zea mays L.) releases specific volatiles in response to herbivory by caterpillars. These volatiles are known to serve as cues for parasitic wasps to locate the herbivores. In the present study the exact time of volatile emission after simulated herbivory (mechanical damage and treatment with caterpillar regurgitant) was measured for seedlings of the cultivars “Ioana Sweet Corn” and “LG11”. Odours were collected every 0.5 h for a total of 12 h. Typical “green leaf odours”, (Z)-3-hexenal, (E )-2-hexenal, (Z)-hexen-1-o1, and (Z)-3-hexen-1-yl acetate, were emitted immediately upon damage and their amounts dropped rapidly after the first collections. Several of the induced compounds were released within 2 h after treatment, while others (mainly sesquiterpenoids) started to be released after 4 h. The LG11 seedlings emitted several compounds (e.g. β-myrcene, (Z)-β-ocimene, benzyl acetate, β-caryophyllene, (E,E )-α-farnesene) that were not detected for Ioana. (E,E )-α-farnesene was continuously emitted by LG11 seedlings, even by undamaged plants. Timing of the release of volatile compounds that the two varieties had in common did not differ significantly, with the exception of indole for which the peak production was considerably earlier for LG11. These findings are discussed in the context of biosynthetic pathways and mechanisms involved in induced emissions of plant volatiles and the exploitation of the resulting odour by parasitoids and predators of herbivores. Received: 23 October 1997 / Accepted: 9 June 1998