BackgroundAssociational herbivore resistance is potentiated by neighbouring heterogenic plant species that impact a focal plant’s attraction to herbivores or the damage that they cause. One mechanism to confer associational resistance is believed to be exposure to neighbour-emitted volatiles, the receivers of which range from intra- and interspecific neighbour plants to higher-trophic-level insects. In previous studies the passive adsorption of neighbour-emitted semivolatiles has been reported, but little is known regarding the mechanisms and ecological consequences on the receiver plant and its associated biota. To utilize volatile-based associational resistance for agricultural applications, it is imperative to know its effectiveness under varying diurnal temperatures and whether herbivore natural enemies, providing biological control, are impacted. Mimicking varying diurnal temperatures in a laboratory set-up, we assessed how the tritrophic model system Brassica oleracea var. italica (broccoli)–Plutella xylostella (crucifer specialist herbivore)–Cotesia vestalis (endoparasitoid of P. xylostella) is influenced by exposure to the natural semivolatile emitter plant Rhododendron tomentosum Harmaja.ResultsRhododendron tomentosum-exposed B. oleracea was less susceptible to P. xylostella oviposition at both night-time (12°C) and day-time (22°C) temperatures and less favoured and damaged by P. xylostella larvae at 12°C. Exposure did not interfere with indirect defence, i.e. attraction of the natural enemy C. vestalis on host-damaged, R. tomentosum-exposed B. oleracea under 22°C, while there was a reduction in attraction (marginal preference towards host-damaged B. oleracea) under 12°C.ConclusionsThe ability of R. tomentosum exposure to render associational resistance against an agriculturally important Brassica herbivore P. xylostella without severely compromising the specialist parasitoid C. vestalis host location encourages further studies on the potential of using this naturally abundant plant for biocontrol. The generality of our finding on temperature as a potential regulating mechanism for the efficacy of semivolatile emitter-based associational resistance towards specialist pest larval damage should be further studied in natural and agricultural associations. Our study emphasizes the need to develop techniques to compare volatiles at the leaf versus air interface and associate their appearance and ecological role with times of activity and level of specialisation of herbivores and their natural enemies.

Plants emit an extremely diverse bouquet of volatile organic compounds (VOCs) from their above-ground and below-ground parts. Emissions are constitutive or induced, e.g. by herbivores. VOCs can be classified as highly volatile, volatile and semivolatile compounds. Sesquiterpenes (SQTs) are typical semi-volatile organic compounds (sVOCs) released by plants. Similarly, herbivore-induced homoterpenes and methyl salicylate also have relatively low volatility. SVOCs have a high boiling point (> 240 °C) and a vapour pressure below 0.005 kPa at 25 °C. Glandular trichomes on plant surfaces can store SQTs in mixtures with more volatile VOCs, which are released into the air by diffusion or after gland rupture. The sVOCs stored in glandular trichomes often have repellent effects on herbivores, while herbivore-induced sVOCs are known for their attractiveness to natural enemies of herbivores, i.e. they act in indirect chemical defence of plants. Due to their low volatility, sVOCs produced by plants may easily adhere to the surfaces of emitter and neighbouring plants during the colder temperatures that plants face, e.g. at night. On the foliage of neighbouring receiver plants, sVOCs may act in direct and indirect defence of that plant species. When the temperature rises again, sVOCs are released into the atmosphere. The semi-volatile reaction products of highly volatile plant monoterpenes and photochemical pollutants such as ozone could constitute further sVOCs on plant leaf surfaces. Here, we review recent literature of the plant surface-environment interaction of biogenic sVOCs and particularly evaluate potential crop protection strategies such as intercropping and companion planting using sVOC-emitting species. Foliage typically forms the widest surfaces on crop plants, and foliar herbivory is a major type of pest damage during the vegetative stage of crop plants. Foliage is also a major source of herbivore-induced VOC emissions. Consequently, we focus on foliage-mediated sVOCs and their potential in pest management.

Environmentally acquired chemical camouflage is a phenomenon, where a plant growing close to a strong volatile organic compound (VOC) emitter will adsorb and re-emit the VOCs produced by the neighbouring plant. The re-emitted volatile bouquet may resemble more the VOC composition of the neighbour than plant’s own typical odour, and thus act as chemical camouflage against insect detection, potentially simultaneously providing associational resistance towards herbivory. We exposed a pest-sensitive horticultural crop, Brassica oleracea var. italica (broccoli) cv. Lucky, to the volatiles emitted by Rhododendron tomentosum [RT] twigs and assessed the host selection by ovipositing females and larval instars of the major caterpillar pest Pieris brassicae between RT-exposed and control plants. Potential impact of RT exposure on herbivore natural enemies was studied using behavioural tests with a parasitoid wasp Cotesia glomerata. P. brassicae females laid significantly less eggs and egg clusters were fewer on RT-exposed plants at both night-time (6 °C) and daytime (22 °C) temperatures. Larvae preferred leaves from control plants over RT-exposed plants at both temperatures. Preceding RT-exposure did not disturb orientation of parasitoid wasp Cotesia glomerata females towards B. oleracea plants damaged by its host P. brassicae. However, host-damaged control plants were favoured by the parasitoid over RT-exposed, host-damaged plants. Our results suggest that companion plant based chemical camouflage as a mechanism of pest suppression could be developed as an additional tool for the integrated pest management toolbox in agriculture.