Silvia Dorn

Herbivore‐induced emissions of maize volatiles repel the corn leaf aphid, Rhopalosiphum maidis

When maize plants, Zea mays L., are mechanically damaged and the damaged sites are treated with caterpillar regurgitant, the plants will release a specific blend of volatiles. It is known that these volatiles can be attractive to natural enemies of herbivores. We hypothesise that the plant volatiles constitute part of the induced plant defence and that herbivores will be affected by the odours as well. In laboratory and semi‐field studies this hypothesis was tested for the aphid Rhopalosiphum maidis (Fitch) (Rhynchota, Sternorrhyncha, Aphididae).

SPECIALIZED BEES FAIL TO DEVELOP ON NON- HOST POLLEN: DO PLANTS CHEMICALLY PROTECT THEIR POLLEN?

Bees require large amounts of pollen for their own reproduction. While several morphological flower traits are known to have evolved to protect plants against excessive pollen harvesting by bees, little is known on how selection to minimize pollen loss acts on the chemical composition of pollen. In this study, we traced the larval development of four solitary bee species, each specialized on a different pollen source, when reared on non-host pollen by transferring unhatched eggs of one species onto the pollen provisions of another species. Pollen diets of Asteraceae and Ranunculus (Ranunculaceae) proved to be inadequate for all bee species tested except those specialized on these plants. Further, pollen of Sinapis (Brassicaceae) and Echium (Boraginaceae) failed to support larval development in one bee species specialized on Campanula (Campanulaceae). Our results strongly suggest that pollen of these four taxonomic groups possess protective properties that hamper digestion and thus challenge the general view of pollen as an easy-to-use protein source for flower visitors.

Long flights in Cydia pomonella L. (Lepidoptera: Tortricidae) measured by a flight mill: influence of sex, mated status and age

Abstract. The flight capacity of Cydia pomonella L. (Lepidoptera: Tortricidae) was measured in the laboratory by using computer‐linked flight mills. Codling moths showed a large variation in flight capacity between individuals. We defined arbitrarily a longest single flight (LSF) of more than 5 km as an index for long‐flyers. About 16.7% of virgin and 10.0% of mated males and 20.0% of virgin and 7.4% of mated females undertook such flights. Based on the LSF and the total distance flown (TDF^we concluded that males and females have little or no difference in flight capacity and that both the within‐and between‐habitat flights are similar in number and magnitude for both sexes. In the field, females are therefore potentially able to undertake flights of up to 11 km, as reported for males by other authors. This ability was highest at ages of 2–7 days after emergence, i.e. the first third of their lifetime, for virgin and mated male moths and for virgin female moths. Mated females showed peak flight capacity between 1 and 3 days after eclosion, which corresponded with the major egg‐laying period. Few long flights were undertaken before oviposition. These findings do not agree with the oogenesis flight syndrome described by other authors, and this theory is believed not to apply to C.pomonella. Our laboratory results are discussed in relation to field experiments in general and hypotheses are developed about the significance of long‐flyers for this species.

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.

Preimaginal learning determines adult response to chemical stimuli in a parasitic wasp

The behavioural responses of parasitic wasps to chemical cues from their hosts and host plants are known to be affected by genetic and environmental components. In a previous study of the codling moth ectoparasitoid Hyssopus pallidus, we found that the response of adult parasitoids to the frass of their host caterpillars depended on a learning process involving plant cues. In the present study, we investigated how and when learning takes place. A series of experiments was conducted involving exposure of parasitoids to fruit cues at different developmental stages. While parasitoids were not able to learn the fruit cues in the adult stage, exposure to fruit odour at early preimaginal stages significantly increased the adult response to frass from fruit–fed caterpillars. The olfactory memory persisted through metamorphosis, with a retention time of 14 days. Preimaginal learning was not confined to fruit cues but was also demonstrated for a host– and fruit–independent cue, menthol. Parasitoids exposed to menthol odour at the egg and larval stages no longer showed negative responses as adults. Sensitization to fruit cues and habituation to menthol are considered to be the mechanisms involved. This study provides evidence of true preimaginal learning of olfactory cues in a parasitic wasp.

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.

Sexual dimorphism in the olfactory orientation of adult Cydia pomonella in response to α-farnesene

The role of host plant‐derived volatile substances on the behaviour of adult codling moth, Cydia pomonella L. (Lepidoptera: Tortricidae) is poorly understood. We tested the response of females and males to a range of α‐farnesene dosages. Natural α‐farnesene from apples contains the E,E and Z,E isomers in varying proportions. No difference in the response of C. pomonella to preparations containing two different proportions of the E,E and Z,E isomers was noted (77:20.7 or 1.2:84.7% E,E and Z,E isomers respectively), indicating a similar bioactivity of E,E and Z,E α‐farnesene on codling moth. A marked sexual dimorphism was found to increasing dosages of α‐farnesene. Females were attracted to low dosages (starting from 63.4 ng) and repelled by high dosages (ending at 12 688 ng). The dose response over this concentration range was linear with a negative slope. Both mated and virgin females responded similarly in kind but differently in degree, both attraction and repellency being more pronounced in mated females. Males were neither attracted nor repelled over a large dose range (63.4 to 12 688 ng) except the highest rate which was attractive. This indicates a stronger dependency of females on plant‐derived volatiles.

Effect of plant volatile oils in protecting stored cowpea Vigna unguiculata (L.) Walpers against Callosobruchus maculatus (F.) (Coleoptera: Bruchidae) infestation.

Adult Callosobruchus maculatus (F.) were introduced into cowpea seeds which were stored in containers with volatile oils derived from Mentha arvensis, M. piperata, M. spicata and Cymbopogon nardus. The numbers of eggs laid, adult mortality, adult emergence and subsequent seed damage were studied for four months. All oils significantly influenced all parameters (P<0.05) and results with different parameters were generally parallel. Significant differences for at least some time/parameters combinations indicated an order of potency of M. spicata>M. piperata>M. arvensis>C. nardus.

Long foraging distances impose high costs on offspring production in solitary bees

1. Solitary bees are central place foragers returning to their nests several times a day with pollen and nectar to provision their brood cells. They are especially susceptible to landscape changes that lead to an increased spatial separation of suitable nesting sites and flower rich host plant stands. While knowledge of bee foraging ranges is currently growing, quantitative data on the costs of foraging flights are very scarce, although such data are crucial to understand bee population dynamics. 2. In this study, the impact of increased foraging distance on the duration of foraging bouts and on the number of brood cells provisioned per time unit was experimentally quantified in the two pollen specialist solitary bee species Hoplitis adunca and Chelostoma rapunculi. Females nesting at different sites foraged under the same environmental conditions on a single large and movable flowering host plant patch in an otherwise host plant free landscape. 3. The number of brood cells provisioned per time unit by H. adunca was found to decrease by 23%, 31% and 26% with an increase in the foraging distance by 150, 200 and 300 m, respectively. The number of brood cells provisioned by C. rapunculi decreased by 46% and 36% with an increase in the foraging distance by 500 and 600 m, respectively. 4. Contrary to expectation, a widely scattered arrangement of host plants did not result in longer mean duration of a foraging bout in H. adunca compared to a highly aggregated arrangement, which might be due to a reduced flight directionality combined with a high rate of revisitation of already depleted flowers in the aggregated plant arrangement or by a stronger competition and disturbance by other flower visitors. 5. The results of this study clearly indicate that a close neighbourhood of suitable nesting and foraging habitats is crucial for population persistence and thus conservation of endangered solitary bee species.

Synergism between aromatic compounds and green leaf volatiles derived from the host plant underlies female attraction in the oriental fruit moth

Blends of volatile compounds emitted by host plants are known to mediate the attraction of gravid female herbivores to oviposition sites, but the role of individual odor components is still little understood. We characterized the olfactory response of mated female Cydia (Grapholita) molesta (Busck) (Lepidoptera: Tortricidae) to synthetic mixtures of compounds emitted by peach shoot, a key host plant of this herbivore, and investigated the role of important constituents of bioactive mixtures in moth attraction. Relative ratios of constituents of the mixtures corresponded to the natural ratio of volatile compounds collected in the plants headspace. A significant attractant effect was found for a comparatively complex 10‐compound mixture that included four green leaf volatiles [(Z)‐3‐hexen‐1‐ol, 1‐hexanol, (E)‐2‐hexenal, and (Z)‐3‐hexen‐1‐yl acetate], five aromatics (benzaldehyde, methyl salicylate, methyl benzoate, benzonitrile, and phenylacetonitrile), and a carboxylic acid (valeric acid). Using a subtraction approach, the number of compounds was progressively decreased, resulting in a bioactive 5‐compound mixture composed of two constituents, green leaf volatiles and aromatic compounds. Further evaluations revealed that benzaldehyde and benzonitrile must be present in association with three distinct green leaf volatiles to produce an attractant effect on the female moths. This 5‐compound mixture was as attractive as natural peach shoot volatiles, which are known to comprise over 20 compounds. Results are discussed in light of the documented synergistic effect between the three general green leaf volatiles and the two specific aromatic compounds.