Role of host feeding niches and host refuges in habitat-related behaviour of Hyssopus pallidus (Hymenoptera: Eulophidae), a larval parasitoid of the codling moth.

Parasitoid fitness depends largely on the capability to locate a host in an ecosystem. A parasitoid of a polyphagous host might not be able to find or to access the host in all its feeding niches. This study evaluated the niche selection of Hyssopus pallidus (Askew), a larval parasitoid of Cydia pomonella (Linnaeus), at the plant level with the goal of assessing its potential for biological control on different fruit crops throughout the plant cycle. Parasitoid behaviour during host location and reproduction rate were investigated on host caterpillars actively feeding on apple, pear, apricot or plum, and on caterpillars diapausing under the bark. Under laboratory conditions, the host searching behaviour of H. pallidus varied depending on the fruit species offered and the infestation of the fruits. Parasitoid females searched longer on apples than on other fruit species, and they searched longer on infested than on uninfested apples. Female wasps were able to locate and parasitize host caterpillars under the tree bark, and their behaviour did not vary with host accessibility. The numbers of caterpillars attacked by H. pallidus depended on the fruit species. The highest numbers of caterpillars were parasitized in apples and apricots. Their accessibility (i.e. position) within the fruit or on the branch did not influence parasitism success. Although hosts were parasitized throughout the season, the best results were achieved with early and late releases. Therefore, the host niche selection behaviour of H. pallidus most likely co-evolved with the host C. pomonella on apples, which renders H. pallidus a valuable biocontrol agent for successful release at different times of the season into apple orchards.

Response of female Cydia molesta (Lepidoptera: Tortricidae) to plant derived volatiles.

Peach shoot volatiles were attractive to mated female oriental fruit moth, Cydia molesta (Busck), in a dual choice arena. No preference was observed between leaf odours from the principle host plant, peach, and the secondary host plant, apple. Twenty-two compounds were identified in headspace volatiles of peach shoots using gas chromatography-mass spectrometry. Green leaf volatiles accounted for more than 50% of the total emitted volatiles. A bioassay-assisted fractionation using different sorbent polymers indicated an attractant effect of compounds with a chain length of 6-8 carbon atoms. The major compounds of this fraction were tested either singly or in combinations for behavioural response of females. Significant bioactivity was found for a three-component mixture of (Z)-3-hexen-1-yl acetate, (Z)-3-hexen-1-ol and benzaldehyde in a 4:1:1 ratio. This synthetic mixture elicited a similar attractant effect as the full natural blend from peach shoots as well as the bioactive fraction.

Systemically induced plant volatiles emitted at the time of "danger".

Feeding by Pieris brassicae caterpillars on the lower leaves of Brussels sprouts (Brassica oleracea var. gemmifera) plants triggers the release of volatiles from upper leaves. The volatiles are attractive for a natural antagonist of the herbivore, the parasitoid Cotesia glomerata. Parasitoids are attracted only if additional damage is inflicted on the systemically induced upper leaves and only after at least three days of herbivore feeding on the lower leaves. Upon termination of caterpillar feeding, the systemic signal is emitted for a maximum of one more day. Systemic induction did not occur at low levels of herbivore infestation. Systemically induced leaves emitted green leaf volatiles, cyclic monoterpenoids, and sesquiterpenes. GC-MS profiles of systemically induced and herbivore-infested leaves did not differ for most compounds, although herbivore infested plants did emit higher amounts of green leaf volatiles. Emission of systemically induced volatiles in Brussels sprouts might function as an induced defense that is activated only when needed, i.e., at the time of caterpillar attack. This way, plants may adopt a flexible management of inducible defensive resources to minimize costs of defense and to maximize fitness in response to unpredictable herbivore attack.

beta-Glucosidase: an elicitor of herbivore-induced plant odor that attracts host-searching parasitic wasps.

Cabbage plants respond to caterpillar (Pieris brassicae) herbivory by releasing a mixture of volatiles that makes them highly attractive to parasitic wasps (Cotesia glomerata) that attack the herbivores. Cabbage leaves that are artificially damaged and subsequently treated with gut regurgitant of P. brassicae caterpillars release a volatile blend similar to that of herbivore-damaged plants. We demonstrate the presence of beta-glucosidase in P. brassicae regurgitant. Leaves treated with commercial beta-glucosidase (from almonds) release a volatile blend similar to that of leaves treated with P. brassicae regurgitant. In a flight bioassay, leaves treated with almond beta-glucosidase are highly attractive to the parasitic wasp C. glomerata. Furthermore, the wasps do not discriminate between cabbage leaves treated with almond beta-glucosidase and leaves treated with larval regurgitant. beta-Glucosidase was also recorded in cabbage leaf extract, but this is not as effective as caterpillar beta-glucosidase in eliciting the volatile production. Caterpillars that feed on a beta-glucosidase-free diet secrete the enzyme, and their regurgitant is an effective elicitor of the plant response. These findings show that beta-glucosidase is a P. brassicae-secreted elicitor of the defense response of cabbage plants to herbivore injury, inducing the emission of volatiles that are used by parasitoids of the herbivore to locate their victims.

Induction of parasitoid attracting synomone in brussels sprouts plants by feeding ofPieris brassicae larvae: Role of mechanical damage and herbivore elicitor.

Induction of plant defense in response to herbivory includes the emission of synomones that attract the natural enemies of herbivores. We investigated whether mechanical damage to Brussels sprouts leaves (Brassica oleracea var.gemmifera) is sufficient to obtain attraction of the parasitoidCotesia glomerata or whether feeding byPieris brassicae caterpillars elicits the release of synomones not produced by mechanically damaged leaves. The response of the parasitoidCotesia glomerata to different types of simulated herbivory was observed. Flight-chamber dual-choice tests showed that mechanically damaged cabbage leaves were less attractive than herbivore-damaged leaves and mechanically damaged leaves treated with larval regurgitant. Chemical analysis of the headspace of undamaged, artificially damaged, caterpillar-infested, and caterpillar regurgitant-treated leaves showed that the plant responds to damage with an increased release of volatiles. Greenleaf volatiles and several terpenoids are the major components of cabbage leaf headspace. Terpenoids are emitted in analogous amounts in all treatments, including undamaged leaves. On the other hand, if the plant is infested by caterpillars or if caterpillar regurgitant is applied to damaged leaves, the emission of green-leaf volatiles is highly enhanced. Our data are in contrast with the induction of more specific synomones in other plant species, such as Lima bean and corn.

A long-range attractant kairomone for egg parasitoidTrissolcus basalis, isolated from defensive secretion of its host,Nezara viridula.

A short-chain α,ß-unsaturated aldehyde, (E)-2-decenal, present in the defensive metathoracic gland ofNezara viridula (L.) (Heteroptera: Pentatomidae), stimulates a behavioral response in the egg parasitoidTrissolcus basalis (Woll.) (Hymenoptera: Scelionidae). Preliminary studies showed thatT. basalis are attracted to an area containing adultN. viridula, but we also found that femaleT. basalis would examine and probe glass beads coated with an acetone extract of the metathoracic gland from males or females. Using this bioassay, the kairomone was isolated by bioassay directed by preparative gas chromatography and identified by NMR and mass spectrometry as (E)-2-decenal. The biological activity of the identified aldehyde was compared with analogs to determine specificity. An unstable Z isomer was found to be more active but not present in detectable or behaviorly relevant levels in the host, based on the bell-shaped dose-response curve of the two isomers. An investigation was also designed to determine if theE isomer was also responsible for the egg recognition kairomone activity previously reported. However, no 2-decenal isomers were detected in host egg extracts and the chemical characteristics of the 2-decenal isomers differ from the unidentified egg recognition kairomone. The role of the (E)-2-decenal in attracting femaleT. basalts toN. viridula was demonstrated in a Y-tube olfactometer; this alk-2-enal appears to act as a long-range kairomone orientingT. basalis toNezara populations.