Chemical cues mediating aphid location by natural enemies

There is increasing evidence that chemical cues play a pivotal role in host selection by the natural enemies of aphids. We use Vinson's (1976) division of the host selection process into habitat location, host location and host acceptance for both parasitoids and predators and review what is known about the role of semiochemicals in aphid selection by natural enemies. For habitat location (i.e. detection of the host plant), volatiles emitted by plants after aphid attack have been described for a number of plant-aphid interactions. These synomones indicate not only the presence of an aphid host plant to the predator or parasitoid, but also the presence of aphids. Volatiles emitted from undamaged host plants are often attractive to aphid parasitoids, but less so for predators. Host location by the natural enemy on the food plant is guided by semiochemicals that mostly originate from the aphids, in particular aphid alarm pheromone, honeydew, or the smell of the aphid itself. Host acceptance is guided by contact chemicals for both predators and parasitoids. In parasitoids, host recognition may be based on visual cues or on contact chemicals on the aphid's cuticle, whereas host acceptance is ultimately based on as yet unknown substances within the aphid's hemolymph. While it appears that many predators and parasitoids are attracted to the same semiochemicals, synergistic and antagonistic interactions among chemical substances have only rarely been investigated. More research into model systems is needed, not only to identify important semiochemicals, but also to determine their range of attraction. Recent progress in the development of analytical techniques has created new opportunities to improve our understanding of the chemical ecology of aphid-natural enemy interactions in the coming years.