Differences in volatile profiles of turnip plants subjected to single and dual herbivory above- and belowground

Abstract

Plants attacked by herbivorous insects emit volatile organic compounds that are used by natural enemies to locate their host or prey. The composition of the blend is often complex and specific. It may vary qualitatively and quantitatively according to plant and herbivore species, thus providing specific information for carnivorous arthropods. Most studies have focused on simple interactions that involve one species per trophic level, and typically have investigated the aboveground parts of plants. These investigations need to be extended to more complex networks that involve multiple herbivory above- and belowground. A previous study examined whether the presence of the leaf herbivore Pieris brassicae on turnip plants (Brassica rapa subsp. rapa) influences the response of Trybliographa rapae, a specialist parasitoid of the root feeder Delia radicum. It showed that the parasitoid was not attracted by volatiles emitted by plants under simultaneous attack. Here, we analyzed differences in the herbivore induced plant volatile (HIPV) mixtures that emanate from such infested plants by using Orthogonal Partial Least Squares-Discriminant Analysis (OPLS-DA). This multivariate model focuses on the differences between odor blends, and highlights the relative importance of each compound in an HIPV blend. Dual infestation resulted in several HIPVs that were present in both isolated infestation types. However, HIPVs collected from simultaneously infested plants were not the simple combination of volatiles from isolated forms of above- and belowground herbivory. Only a few specific compounds characterized the odor blend of each type of damaged plant. Indeed, some compounds were specifically induced by root herbivory (4-methyltridecane and salicylaldehyde) or shoot herbivory (methylsalicylate), whereas hexylacetate, a green leaf volatile, was specifically induced after dual herbivory. It remains to be determined whether or not these minor quantitative variations, within the background of more commonly induced odors, are involved in the reduced attraction of the root feeder's parasitoid. The mechanisms involved in the specific modification of the odor blends emitted by dual infested turnip plants are discussed in the light of interferences between biosynthetic pathways linked to plant responses to shoot or root herbivory.

Fig. 1

Two dimensional bubble plot representation of volatiles emitted after different forms of herbivore infestations in Brassica rapa subsp. rapa plants: discriminate functions of root herbivory by Delia radicum larvae (D) and leaf herbivory by Pieris brassicae caterpillars (P) plotted against each other. Numbers correspond to compounds listed in Table 1. The size of each bubble representing a volatile corresponds to the weight vector value of that volatile for the dual herbivory treatment, i.e., herbivory by Delia radicum larvae and Pieris brassicae caterpillars (DP). The color shows if the volatile increases (filled) or decreases (open) in the DP treatment. The three arrows indicate examples of compounds given in the text. The table below presents the number of latent variables (LVs) prescribed by cross-validation and the P-values for the significance (P ≤ 0.050) of the system-wide effect brought about by each form of herbivory