A survey of identified kairomones and synomones used by insect parasitoids to locate and accept their hosts

Summary The host utilization process of insect parasitoids can be described by three stages of (1)habitatlocation, (2)host-location, and (3)host-acceptance andoviposition. There are 19 systems in which chemicals used inhabitat-location have been identified, 12 systems in which chemical cues leading tohost-location have been identified, and 16 systems in which chemicals elicitinghost-acceptance andoviposition have been identified. Both the chemical class and the source of the infochemical change with the stage of the host utilization process. Semiochemicals identified in thehabitatlocation stage were predominantly aldehydes, alcohols, sulfur-containing compounds, esters and terpenes, and were equally likely to be from the host-plant of the host, or from the host itself. Semiochemicals identified in thehost-location stage were sugars, alkanes, terpenes and heterocyclic aromatic compounds and 3/4 of them were host-produced cues. In thehost-acceptance andoviposition stage the identified semiochemicals were all produced by the host and were proteins, amino acids, triglycerides and salts. The importance of recognizing specific cues involved in host utilization by parasitoids is discussed, and suggestions for future research are made.     

Why do plants ‘talk’?

Summary Plant defence can be induced by herbivory. This is true for both direct and indirect plant defence. Induced direct defence has been the most studied of the two. However, in most cases induced direct defence does not appear to be a water-tight defence option. In contrast, induced indirect defence through the production of herbivore-induced carnivore attractants can be a decisive factor in the extermination of herbivore populations. In this paper the main characteristics of induced attraction of carnivores by plants are reviewed. This includes the similarities and dissimilarities among tritrophic systems. There are two main patterns of induced carnivore attraction. (1) Through the emission of the same bouquet as that emitted by mechanically damaged plants, but in larger quantities and for a longer period of time after damage. (2) Through emission of large amounts of new volatiles that are synthesizedde novo in response to herbivore feeding andnot in response to mechanical wounding.Herbivore populations may be decimated by carnivores. Therefore it should be realized that herbivoreinduced carnivore attractants are essential in an important step in carnivore foraging,i.e. long-distance herbivore location. Once herbivores have started feeding on a plant and direct defence is not effective, induced indirect defence may be decisive for plant survival. Therefore, it is concluded that indirect defence is an essential aspect of induced plant defence directed at herbivorous arthropods.     

Dynamics of pheromone production and communication in the mountain pine beetle, Dendroctonus ponderosae Hopkins, and the pine engraver, Ips pini (Say) (Coleoptera: Scolytidae)

Summary. The mountain pine beetle, Dendroctonus ponderosae Hopkins, and the pine engraver, Ips pini (Say), often co-exist in lodgepole pine, Pinus contorta var. latifolia Engelmann. Intra- and interspecific semiochemical communication occurs in both species and their complete semiochemical repertoire and precise dynamics of pheromone production have not been elucidated. Porapak-Q extracts of captured volatiles from beetles of each species aerated at different attack phases (freshly emerged, pioneer sex alone in the log and both sexes paired in new galleries), followed by gas chromatographic-electroantennographic detection (GC-EAD) and GC-mass spectroscopic analyses identified 17 compounds (seven compounds common to both species, six present in D. ponderosae and four present in I. pini) that excited the antennae of either or both species. Seven compounds for D. ponderosae and nine for I. pini had not been assessed for behavioural activity. In field trapping experiments, 2-phenylethanol produced by both species inhibited the response of D. ponderosae to its aggregation pheromones. exo- and endo-Brevicomin produced by D. ponderosae significantly decreased the response of I. pini to its aggregation pheromone ipsdienol. Nonanal, a ubiquitous compound found in the volatiles of lodgepole pine, various nonhosts and in both beetle species deterred the response of I. pini to ipsdienol. The occurrence of cis-verbenol, trans-verbenol and verbenone in emergent I. pini, and verbenone and 2-phenylethanol in emergent D. ponderosae suggests that these compounds may inhibit aggregation and induce dispersal following emergence. Termination of aggregation in D. ponderosae appears to depend on the production of frontalin in combination with changes in the relative ratios of verbenone, exo-brevicomin, trans-verbenol and 2-phenylethanol. In I. pini, the cessation of ipsdienol production by males is probably the main factor in terminating aggregation. Received 16 November 1999; accepted 7 August 2000     

Evolution of plant volatile production in insect-plant relationships

Summary The production of volatile secondary plant substances during the evolution of terrestrial plants is reviewed in regard to the defensive systems of plants to microorganisms and herbivores. Plant volatiles can be produced by both anabolic and catabolic processes. Although attraction of pollinators is a well-studied phenomenon, functions of volatiles range from excretion of waste products to the production of compounds attracting natural enemies of herbivores. During the evolution of the angiosperms a diversity of volatiles were selected to defend generative parts against microorganisms. Many of these allomones were related to or even identical with sex pheromones of insects. As a result flowers of angiosperms became utilized as a mating site. Consequently insects visiting flowers became involved in pollination, facilitating the steps from anemophily to entomophily. The efficiency of entomophily was increased because of nutritional rewards.An evolutionary scenario for the impact of plant volatiles on insects is presented and the role of volatile allomones in the establishment of plant-insect relationships is emphasized by (1) their strong antimicrobial properties, (2) strategies to protect symbiotic microorganisms, (3) their function as repellents and deterrents, (4) the use of volatile allomones as kairomones. These facts speak for an adaptation of insects to plant physiology and a limited importance of phytophagous insects in selection pressure upon plants. Herbivorous insects have realized specific adaptations to be able to discriminate between complex odour blends, but the utilization of chemical groups among insect taxa is different.The main theories on plant chemical defence do not discuss the impact of volatiles on host plant selection and may be apt to revision when pheromones, allomones, kairomones and synomones are not taken into account.     

New insights in analysing parasitoid attracting synomones: early volatile emission and use of stir bar sorptive extraction

Summary. It is well known that feeding by Pieris brassicae caterpillars on cabbage leaves triggers the release of volatiles that attract natural antagonists such as the parasitoid Cotesia glomerata. The temporal dynamics in the emissions of parasitoid attracting volatiles has never been elucidated in this system. In a time course experiment, caterpillar infested leaves attracted the parasitoid within one hour after infestation. At such an early stage of infestation, as much as fifty percent of the parasitoids flew towards the infested plant in a wind tunnel bioassay, while only five percent flew towards the non-infested control plant. Three hours after infestation and later, the response to the volatiles from the infested plant reached its maximum and then continued at a constantly high level for the remaining 14 hours of the experiment. Chemical analyses of volatiles collected from infested leaves at short time intervals during the first 24 hours identified a total of ten compounds, comprising green leaf volatiles, terpenoids, and a nitrile. Significant increase of emission within the first 5 hours following initial herbivory was detected for (Z)-3-hexen-1-ol, (Z)-3-hexen-1-yl acetate, cineole and benzylcyanide. Subsequently, a coupled bioassay-chemical analysis procedure was developed allowing for testing and analyzing the same sample for future identification of the bioactive compounds. This was achieved by using stir bar sorptive extraction for the analysis of solvent extracts of caterpillar-damaged leaves.