Variation in natural plant products and the attraction of bodyguards involved in indirect plant defense

Plants can respond to feeding or egg deposition by herbivorous arthropods by changing the volatile blend that they emit. These herbivore-induced plant volatiles (HIPVs) can attract carnivorous natural enemies of the herbivores, such as parasitoids and predators, a phenomenon that is called indirect plant defense. The volatile blends of infested plants can be very complex, sometimes consisting of hundreds of compounds. Most HIPVs can be classified as terpenoids (e.g., (E)-beta-ocimene, (E,E)-alpha-farnesene, (E)-4,8-dimethyl-1,3,7-nonatriene), green leaf volatiles (e.g., hexanal, (Z)-3-hexen-1-ol, (Z)-3-hexenyl acetate), phenylpropanoids (e.g., methyl salicylate, indole), and sulphur- or nitrogen-containing compounds (e.g., isothiocyanates or nitrites, respectively). One highly intriguing question has been which volatiles out of the complex blend are the most important ones for the carnivorous natural enemies to locate ?suitable host plants. Here, we review the methods and techniques that have been used to elucidate the carnivore-attracting compounds. Electrophysiological methods such as electroantennography have been used with parasitoids to elucidate which compounds can be perceived by the antennae. Different types of elicitors and inhibitors have widely been applied to manipulate plant volatile blends. Furthermore, transgenic plants that were genetically modified in specific steps in one of the signal transduction pathways or biosynthetic routes have been used to find steps in HIPV emission crucial for indirect plant defense. Furthermore, we provide an overview on biotic and abiotic factors that influence the emission of HIPVs and how this can affect the interactions between members of different trophic levels. Consequently, we review the progress that has been made in this exciting research field during the past 30 years since the first studies on HIPVs emerged and we highlight important issues to be addressed in the future.