Below ground and above ground plant-insect-microorganism interactions are complex and regulate most of the developmental responses of important crop plants such as tomato. We investigated the influence of root colonization by a nonmycorrhizal plant-growth-promoting fungus on direct and indirect defenses of tomato plant against aphids. The multitrophic system included the plant Solanum lycopersicum ('San Marzano nano'), the root-associated biocontrol fungus Trichoderma longibrachiatum strain MK1, the aphid Macrosiphum euphorbiae (a tomato pest), the aphid parasitoid Aphidius ervi, and the aphid predator Macrolophus pygmaeus. Laboratory bioassays were performed to assess the effect of T. longibrachiatum MK1, interacting with the tomato plant, on quantity and quality of volatile organic compounds (VOC) released by tomato plant, aphid development and reproduction, parasitoid behavior, and predator behavior and development. When compared with the uncolonized controls, plants whose roots were colonized by T. longibrachiatum MK1 showed quantitative differences in the release of specific VOC, better aphid population growth indices, a higher attractiveness toward the aphid parasitoid and the aphid predator, and a quicker development of aphid predator. These findings support the development of novel strategies of integrated control of aphid pests. The species-specific or strain-specific characteristics of these below ground-above ground interactions remain to be assessed.
The prevailing reaction of plants to pest attack is the activation of various defense mechanisms. In tomato, several studies indicate that an 18 amino acid (aa) peptide, called systemin, is a primary signal for the systemic induction of direct resistance against plant-chewing pests, and that the transgenic expression of the prosystemin gene (encoding the 200 aa systemin precursor) activates genes involved in the plant response to herbivores. By using a combination of behavioral, chemical, and gene expression analyses, we report that systemin enhances the production of bioactive volatile compounds, increases plant attractivity towards parasitiod wasps, and activates genes involved in volatile production. Our data imply that systemin is involved in the systemic activation of indirect defense in tomato, and we conclude that a single gene controls the systemic activation of coordinated and associated responses against pests.
The leafminer Coptodisca sp. (Lepidoptera: Heliozelidae), recently recorded for the first time in Europe on Italian black and common walnut trees, is shown to be the North-American Coptodisca lucifluella (Clemens) based on morphological (forewing pattern) and molecular (cytochrome oxidase c subunit I sequence) evidence. The phylogenetic relatedness of three species feeding on Juglandaceae suggests that C. lucifluella has likely shifted, within the same host plant family, from its original North-American hosts Carya spp. to Juglans spp. Over the few years since its detection, it has established in many regions in Italy and has become a widespread and dominant invasive species. The leafminer completes three to four generations per year, with the first adults emerging in April-May and mature larvae of the last generation starting hibernation in September-October. Although a high larval mortality was recorded in field observations (up to 74%), the impact of the pest was substantial with all leaves infested at the end of the last generation in all 3 years tested. The distribution of the leafminer in the canopy was homogeneous. The species is redescribed and illustrated, a lectotype is designated and a new synonymy is established.
Flight responses of the aphid parasitoid Aphidius ervi to tomato volatiles have recently demonstrated that different plant stresses can lead to increases in attractiveness for this parasitoid. For example, infestation of tomato plants by the aphid Macrosiphum euphorbiae results in the overexpression of defensive genes, as well as the release of volatile compounds that attract aphid parasitoids. Here, we determine which of the induced compounds elicit a significant electrophysiological response from parasitoid antennae. Compounds shown to be detected at the antennal level were then tested at a range of doses in a wind tunnel assay. A significant electroantennogram response was demonstrated for three compounds, (8S,9R)-(E)-caryophyllene, methyl salicylate, and (Z)-3-hexen-1-ol, over four concentrations. These compounds proved to be significantly attractive in the wind tunnel at a rate not always proportionally dependent upon the dose. The practical implications of these findings are discussed in the framework of sustainable control for pest aphids in agriculture.
The blend of volatile compounds emitted by tomato plants (Solanum lycopersicum ) infested with the potato aphid (Macrosiphum euphorbiae ) has been studied comparatively with undamaged plants and aphids themselves. Aphid-infested plants were significantly more attractive towards Aphidius ervi than undamaged plants and aphids themselves. Oriented response towards host-damaged plant, from which aphids were removed just before running the bioassay, did not differ from that recorded for infested plants. Collection of the volatiles and analysis by gas chromatography revealed only quantitative differences between uninfested and aphid-infested plants. Nine compounds, a-pinene, (Z )-3-hexen-1-ol, a-phellandrene, limonene, (E )-b-ocimene, p-cymene, methyl salicylate, (E )-b-caryophyllene and an unknown compound, were emitted at higher levels from aphid-infested plants than from undamaged control plants, whilst no differences were noted for hexanal, 6-methyl-5-hepten-2-one, and humulene (0a-caryophyllene). Synthetic standards of these compounds were tested in wind tunnel bioassays and all elicited a significant increase in oriented flight and landings on the target by the aphid parasitoid Aphidius ervi . (E )-b-caryophyllene resulted the most attractive towards female wasps. These results corroborate the hypothesis that the volatiles produced by the plant in response to aphid attack derive from both jasmonic and salicylic acid pathways, and are exploited by A. ervi as olfactory cues to locate its hosts.
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