Identification of Volatiles That Are Used in Discrimination Between Plants Infested with Prey or Nonprey Herbivores by a Predatory Mite

Carnivorous arthropods can use herbivore-induced plant volatiles to locate their herbivorous prey. In the field, carnivores are confronted with information from plants infested with herbivores that may differ in their suitability as prey. Discrimination by the predatory mite Phytoseiulus persimilis between volatiles from lima bean plants infested with the prey herbivore Tetranychus urticae, or plants infested with the nonprey caterpillar Spodoptera exigua, depends on spider mite density. In this article, we analyzed the chemical composition of the volatile blends from T. urticae-infested lima bean plants at different densities of spider mites, and from S. exigua-infested plants. Based on the behavioral preferences of P. persimilis and the volatile profiles, we selected compounds that potentially enable the mite to discriminate between T. urticae-induced and S. exigua-induced volatiles. Subsequently, we demonstrated in Y-tube olfactometer assays that the relatively large amounts of methyl salicylate and (3E, 7E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene emitted by T. urticae-infested bean plants compared to S. exigua-infested plants enable the predators to discriminate. Our data show that specific compounds from complex herbivore-induced volatile blends can play an important role in the selective foraging behavior of natural enemies of herbivorous arthropods.     

Induction of Direct and Indirect Plant Responses by Jasmonic Acid, Low Spider Mite Densities, or a Combination of Jasmonic Acid Treatment and Spider Mite Infestation

Jasmonic acid (JA) and the octadecanoid pathway are involved in both induced direct and induced indirect plant responses. In this study, the herbivorous mite, Tetranychus urticae, and its predator, Phytoseiulus persimilis, were given a choice between Lima bean plants induced by JA or spider mites and uninduced control plants. Infestation densities resulting in the induction of predator attractants were much lower than thus far assumed, i.e., predatory mites were significantly attracted to plants that were infested for 2 days with only one or four spider mites per plant. Phytoseiulus persimilis showed a density-dependent response to volatiles from plants that were infested with different numbers of spider mites. Similarly, treating plants with increasing concentrations of JA also led to increased attraction of P. persimilis. Moreover, the duration of spider mite infestation was positively correlated with the proportion of predators that were attracted to mite-infested plants. A pretreatment of the plants with JA followed by a spider mite infestation enhanced the attraction of P. persimilis to plant volatiles compared to attraction to volatiles from plants that were only infested with spider mites and did not receive a pretreatment with JA. The herbivore, T. urticae preferred leaf tissue that previously had been infested with conspecifics to uninfested leaf tissue. In the case of choice tests with JA-induced and control leaf tissue, spider mites slightly preferred control leaf tissue. When spider mites were given a choice between leaf discs induced by JA and leaf discs damaged by spider mite feeding, they preferred the latter. The presence of herbivore induced chemicals and/or spider mite products enhanced settlement of the mites, whereas treatment with JA seemed to impede settlement.     

The Role of Methyl Salicylate in Prey Searching Behavior of the Predatory Mite Phytoseiulus persimilis

Many carnivorous arthropods use herbivore-induced plant volatiles to locate their prey. These plant volatiles are blends of up to hundreds of compounds. It is often unknown which compounds in such a complex volatile blend represent the signal to the foraging carnivore. We studied the role of methyl salicylate (MeSA) as part of the volatile blend in the foraging behavior of the predatory mite Phytoseiulus persimilis by using a Y-tube olfactometer. MeSA is one of the compounds released by lima bean, infested with Tetranychus urticae--a prey species of the predatory mite. MeSA attracted satiated predatory mites in a dose-dependent way with optimum attraction at a dose of 0.2 microg. Predatory mites did not discriminate between a prey-induced lima bean volatile blend (that contains MeSA) and a prey-induced volatile blend to which an extra amount of synthetic MeSA had been added. However, they preferred a MeSA-containing volatile blend (induced by T. urticae) to an otherwise similar but MeSA-free blend (induced by jasmonic acid). Adding synthetic MeSA to the MeSA-free blend significantly increased the mites' choice for this odor, suggesting an important role for MeSA. This study is a new step toward unraveling the role of herbivore-induced plant volatiles in the foraging behavior of predatory arthropods.     

Change in Behavioral Response to Herbivore-induced Plant Volatiles in a Predatory Mite Population

Damage by herbivorous spider mites induces plants to produce volatiles that attract predatory mites that consume the spider mites. A clear attraction to volatiles from Lima bean plants infested with the spider mite Tetranychus urticae has been consistently reported during more than 15 years for the predatory mite Phytoseiulus persimilis. We have monitored the response to volatiles from spider-mite infested Lima bean plants for a laboratory population of the predatory mite from 1991 to 1995 on a regular basis. A reduction in the level of attraction in the laboratory population of P. persimilis was recorded in mid-1992. The attraction of the laboratory population was weaker than that of a commercial population in the latter part of 1992, but the responses of these two populations were similarly weak in 1994 and 1995. Therefore, a behavioral change has also occurred in this commercial population. Experiments were carried out to address the potential causes of this change in attraction. The attraction of predators from a commercial population with a strong response decreased after being reared in our laboratory. Within a predator population with a low degree of attraction, strongly responding predators were present and they could be isolated on the basis of their behavior: predators that stayed on spider-mite infested plants in the rearing set-up had a strong attraction, while predators that had dispersed from the rearing set-up were not attracted to prey-infested bean plants. From our laboratory population with a low degree of attraction, isofemale lines were initiated and maintained for more than 20 generations. All isofemale lines exhibited a consistently strong attraction to spider mite-induced plant volatiles, similar to the attraction recorded for several populations in the past 15 years. Neither in a population with a strong attraction nor in two with a weak attraction was the response of the predators affected by a starvation period of 1–3 hr. Based on these results, possible causes for the observed reduction in predator attraction to spider mite-induced bean volatiles are discussed. The predatory mite P. persimilis is a cornerstone of biological control in many crops worldwide. Therefore, the change in foraging behavior recorded in this predator may have serious consequences for biological control of spider mites.     

Prey and Non-prey Arthropods Sharing a Host Plant: Effects on Induced Volatile Emission and Predator Attraction

It is well established that plants infested with a single herbivore species can attract specific natural enemies through the emission of herbivore-induced volatiles. However, it is less clear what happens when plants are simultaneously attacked by more than one species. We analyzed volatile emissions of lima bean and cucumber plants upon multi-species herbivory by spider mites (Tetranychus urticae) and caterpillars (Spodoptera exigua) in comparison to single-species herbivory. Upon herbivory by single or multiple species, lima bean and cucumber plants emitted volatile blends that comprised mostly the same compounds. To detect additive, synergistic, or antagonistic effects, we compared the multi-species herbivory volatile blend with the sum of the volatile blends induced by each of the herbivore species feeding alone. In lima bean, the majority of compounds were more strongly induced by multi-species herbivory than expected based on the sum of volatile emissions by each of the herbivores separately, potentially caused by synergistic effects. In contrast, in cucumber, two compounds were suppressed by multi-species herbivory, suggesting the potential for antagonistic effects. We also studied the behavioral responses of the predatory mite Phytoseiulus persimilis, a specialized natural enemy of spider mites. Olfactometer experiments showed that P. persimilis preferred volatiles induced by multi-species herbivory to volatiles induced by S. exigua alone or by prey mites alone. We conclude that both lima bean and cucumber plants effectively attract predatory mites upon multi-species herbivory, but the underlying mechanisms appear different between these species.     

Variation in Herbivory-induced Volatiles Among Cucumber (<Emphasis Type="Italic">Cucumis sativus</Emphasis> L.) Varieties has Consequences for the Attraction of Carnivorous Natural Enemies

In response to herbivory by arthropods, plants emit herbivory-induced volatiles that attract carnivorous enemies of the inducing herbivores. Here, we compared the attractiveness of eight cucumber varieties (Cucumis sativus L.) to Phytoseiulus persimilis predatory mites after infestation of the plants with herbivorous spider mites (Tetranychus urticae) under greenhouse conditions. Attractiveness differed considerably, with the most attractive variety attracting twice as many predators as the least attractive variety. Chemical analysis of the volatiles released by the infested plants revealed significant differences among varieties, both in quantity and quality of the emitted blends. Comparison of the attractiveness of the varieties with the amounts of volatiles emitted indicated that the quality (composition) of the blend is more important for attraction than the amount of volatiles emitted. The amount of (E)-β-ocimene, (E,E)-TMTT, and two other, yet unidentified compounds correlated positively with the attraction of predatory mites. Quantities of four compounds negatively correlated with carnivore attraction, among them methyl salicylate, which is known to attract the predatory mite P. persimilis. The emission of methyl salicylate correlated with an unknown compound that had a negative correlation with carnivore attraction and hence could be masking the attractiveness of methyl salicylate. The results imply that the foraging success of natural enemies of pests can be enhanced by breeding for crop varieties that release specific volatiles.     

Jasmonic Acid and Herbivory Differentially Induce Carnivore-Attracting Plant Volatiles in Lima Bean Plants

Lima bean plants respond to feeding damage of two-spotted spider mites (Tetranychus urticae) with the emission of a complex blend of volatiles that are products of several different biosynthetic pathways. These volatiles attract the carnivorous mite Phytoseiulus persimilis, a specialist predator of the spider mites that exterminates entire prey populations, and thus the volatiles contribute indirectly to plant defense. The volatile blend constitutes information to the carnivores, and blend composition is an important factor in this. Jasmonic acid (JA) is involved in the signal transduction of this induced defense. Application of JA through the petiole of Lima bean plants induces a volatile blend that is similar, but not identical, to that emitted by spider mite-infested plants. The induced volatiles originate from the lipoxygenase pathway, the shikimic acid pathway, and the isoprenoid pathway. Among the induced bean plant volatiles are nitriles and oximes. Of a total of 61 components, 10 are emitted at significantly different rates. Among these are the terpene (E)-4,8-dimethyl-1,3,7-nonatriene and the phenolic methyl salicylate, two compounds that are known to attract P. persimilis. A crucial test for comparing the effect of spider mite damage and JA application on volatile induction is the response of P. persimilis. The carnivore is attracted by volatiles from JA-treated plants. Moreover, even treatment of Lima bean plants with methyl jasmonate vapor made the plants attractive to the carnivorous mites. However, the predators prefer the volatiles from spider-mite-infested Lima bean plants over those from JA-treated plants. Thus, chemical as well as behavioral analyses demonstrate that spider mite damage and JA treatment have similar, although not identical, effects on volatile induction in Lima bean plants.     

Response of Predatory Insect Scolothrips takahashii Toward Herbivore-Induced Plant Volatiles Under Laboratory and Field Conditions

We studied the response of a predatory thrips, Scolothrips takahashii, towards herbivore-induced plant volatiles emitted by Lima bean plants infested by two-spotted spider mites Tetranychus urticae (green form). Tests were conducted with a Y-tube olfactometer in the laboratory and with traps under field conditions. The odor of artificially damaged and uninfested Lima bean leaves was not more attractive than clean air in the Y-tube olfactometer. The predatory insects showed a greater preference for Lima bean leaves infested by the two-spotted spider mites than for either clean air or uninfested bean leaves. They showed the same preference towards infested leaves from which all spider mites and their visible products had been removed. Neither the spider mites themselves nor their products attracted the predators. In a satsuma mandarin grove, two traps with infested Lima bean plants as an odor source attracted 42 adult S. takahashii in 55 days, whereas no S. takahashii were trapped in two control traps with uninfested Lima bean plants during the same period. No S. takahashii were found during this period in the vicinity of either the sample traps or the control traps (5-m radius of each trap). These data showed that S. takahashii use herbivore-induced plant volatiles in their foraging behavior in natural ecosystems.     

Genetic Variation in Jasmonic Acid- and Spider Mite-Induced Plant Volatile Emission of Cucumber Accessions and Attraction of the Predator <Emphasis Type="Italic">Phytoseiulus persimilis</Emphasis>

Cucumber plants (Cucumis sativus L.) respond to spider–mite (Tetranychus urticae) damage with the release of specific volatiles that are exploited by predatory mites, the natural enemies of the spider mites, to locate their prey. The production of volatiles also can be induced by exposing plants to the plant hormone jasmonic acid. We analyzed volatile emissions from 15 cucumber accessions upon herbivory by spider mites and upon exposure to jasmonic acid using gas chromatography—mass spectrometry. Upon induction, cucumber plants emitted over 24 different compounds, and the blend of induced volatiles consisted predominantly of terpenoids. The total amount of volatiles was higher in plants treated with jasmonic acid than in those infested with spider mites, with (E)-4,8-dimethyl-1,3,7-nonatriene, (E,E)-α-farnesene, and (E)-β-ocimene as the most abundant compounds in all accessions in both treatments. Significant variation among the accessions was found for the 24 major volatile compounds. The accessions differed strongly in total amount of volatiles emitted, and displayed very different odor profiles. Principal component analysis performed on the relative quantities of particular compounds within the blend revealed clusters of highly correlated volatiles, which is suggestive of common metabolic pathways. A number of cucumber accessions also were tested for their attractiveness to Phytoseiulus persimilis, a specialist predator of spider mites. Differences in the attraction of predatory mites by the various accessions correlated to differences in the individual chemical profiles of these accessions. The presence of genetic variation in induced plant volatile emission in cucumber shows that it is possible to breed for cucumber varieties that are more attractive to predatory mites and other biological control agents.     

Volatile Stimuli Related to Feeding Activity of Nonprey Caterpillars, Spodoptera exigua, Affect Olfactory Response of the Predatory Mite Phytoseiulus persimilis

The effect of volatiles related to feeding activity of nonprey caterpillars, Spodoptera exigua, on the olfactory response of the predatory mites Phytoseiulus persimilis was examined in a Y-tube olfactometer. At a low caterpillar density (20 caterpillars on 10 Lima bean leaves), the predators were significantly more attracted to volatiles from infested leaves on which the caterpillars and their products were present or from infested leaves from which the caterpillars and their products had been removed when compared to volatiles from uninfested leaves. The predators, however, significantly avoided odors from 20 caterpillars and their products (mainly feces) removed from bean leaves. In contrast, at a higher caterpillar density (100 caterpillars on 10 Lima bean leaves), the predators avoided volatiles from caterpillar-infested bean leaves. Volatiles from infested leaves from which the caterpillars and their products had been removed were not preferred over volatiles from uninfested leaves. Volatiles from feces collected from 100 caterpillars were strongly avoided by the predators, while the behavior of the predatory mites was not affected by volatiles from 100 caterpillars removed from a plant. The data show that carnivorous arthropods may avoid nonprofitable herbivores. This avoidance seems to result from an interference of volatiles from herbivore products with the attraction to herbivore-induced plant volatiles.     

Exposure of Lima Bean Leaves to Volatiles from Herbivore-Induced Conspecific Plants Results in Emission of Carnivore Attractants: Active or Passive Process?

There is increasing evidence that volatiles emitted by herbivore-damaged plants can cause responses in downwind undamaged neighboring plants, such as the attraction of carnivorous enemies of herbivores. One of the open questions is whether this involves an active (production of volatiles) or passive (adsorption of volatiles) response of the uninfested downwind plant. This issue is addressed in the present study. Uninfested lima bean leaves that were exposed to volatiles from conspecific leaves infested with the spider mite Tetranychus urticae, emitted very similar blends of volatiles to those emitted from infested leaves themselves. Treating leaves with a protein-synthesis inhibitor prior to infesting them with spider mites completely suppressed the production of herbivore-induced volatiles in the infested leaves. Conversely, inhibitor treatment to uninfested leaves prior to exposure to volatiles from infested leaves did not affect the emission of volatiles from the exposed, uninfested leaves. This evidence supports the hypothesis that response of the exposed downwind plant is passive. T. urticae-infested leaves that had been previously exposed to volatiles from infested leaves emitted more herbivore-induced volatiles than T. urticae-infested leaves previously exposed to volatiles from uninfested leaves. The former leaves were also more attractive to the predatory mite, Phytoseiulus persimilis, than the latter. This shows that previous exposure of plants to volatiles from herbivore-infested neighbors results in a stronger response of plants in terms of predator attraction when herbivores damage the plant. This supports the hypothesis that the downwind uninfested plant is actively involved. Both adsorption and production of volatiles can mediate the attraction of carnivorous mites to plants that have been exposed to volatiles from infested neighbors.     

Essential Compounds in Herbivore-Induced Plant Volatiles that Attract the Predatory Mite <Emphasis Type="Italic">Neoseiulus womersleyi</Emphasis>

Carnivorous arthropods use volatile infochemicals emitted from prey-infested plants in their foraging behavior. Although several volatile components are common among plant species, the compositions differ among prey–plant complexes. Studies showed that the predatory mite Neoseiulus womersleyi is attracted only to previously experienced plant volatiles. In this study, we identified the attractant components in prey-induced plant volatiles of two prey–plant complexes. N. womersleyi reared on Tetranychus kanzawai-infested tea leaves showed significant preference for a mixture of three synthetic compounds [mimics of the T. kanzawai-induced tea leaves volatiles: (E)-β-ocimene, (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT), and (E,E)-α-farnesene] at a level comparable to that for T. kanzawai-induced tea plant volatiles. However, mixtures lacking any of these compounds did not attract the predatory mites. Likewise, N. womersleyi reared on T. urticae-infested kidney bean plants showed a significant preference for a mixture of four synthetic compounds [mimics of the T. urticae-induced kidney bean volatiles: DMNT, methyl salicylate (MeSA), β-caryophyllene, and (E,E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene] at a level comparable to that for T. urticae-induced kidney bean volatiles. The absence of any of the four compounds resulted in no attraction. These results indicate that N. womersleyi can use at least four volatile components to identify prey-infested plants.     

The Predatory Mite Phytoseiulus persimilis Does Not Perceive Odor Mixtures As Strictly Elemental Objects

Phytoseiulus persimilis is a predatory mite that in absence of vision relies on the detection of herbivore-induced plant odors to locate its prey, the two-spotted spider-mite Tetranychus urticae. This herbivorous prey is feeding on leaves of a wide variety of plant species in different families. The predatory mites respond to numerous structurally different compounds. However, typical spider-mite induced plant compounds do not attract more predatory mites than plant compounds not associated with prey. Because the mites are sensitive to many compounds, components of odor mixtures may affect each other’s perception. Although the response to pure compounds has been well documented, little is known how interactions among compounds affect the response to odor mixtures. We assessed the relation between the mites’ responses elicited by simple mixtures of two compounds and by the single components of these mixtures. The preference for the mixture was compared to predictions under three conceptual models, each based on one of the following assumptions: (1) the responses elicited by each of the individual components can be added to each other; (2) they can be averaged; or (3) one response overshadows the other. The observed response differed significantly from the response predicted under the additive response, average response, and overshadowing response model in 52, 36, and 32% of the experimental tests, respectively. Moreover, the behavioral responses elicited by individual compounds and their binary mixtures were determined as a function of the odor concentration. The relative contribution of each component to the behavioral response elicited by the mixture varied with the odor concentration, even though the ratio of both compounds in the mixture was kept constant. Our experiments revealed that compounds that elicited no response had an effect on the response elicited by binary mixtures that they were part of. The results are not consistent with the hypothesis that P. persimilis perceives odor mixtures as a collection of strictly elemental objects. They suggest that odor mixtures rather are perceived as one synthetic whole.     

The Involvement of Volatile Infochemicals from Spider Mites and from Food-Plants in Prey Location of the Generalist Predatory Mite <Emphasis Type="Italic">Neoseiulus californicus</Emphasis>

We investigated volatile infochemicals possibly involved in location of the generalist predatory mite Neoseiulus californicus to plants infested with spider mites in a Y-tube olfactometer. The predators significantly preferred volatiles from lima bean leaves infested with Tetranychus urticae to uninfested lima bean leaves. Likewise, they were attracted to volatiles from artificially damaged lima bean leaves and those from T. urticae plus their visible products. Significantly more predators chose infested lima bean leaves from which T. urticae plus their visible products had been removed than artificially damaged leaves, T. urticae, and their visible products. These results suggest that N. californicus is capable of exploiting a variety of volatile infochemicals originating from their prey, from the prey-foodplants themselves, and from the complex of the prey and the host plants (e.g., herbivore-induced volatiles). We also investigated predator response to some of the synthetic samples identified as volatile components emitted from T. urticae-infested lima bean leaves and/or artificially damaged lima bean leaves. The predators were attracted to each of the five synthetic volatile components: linalool, methyl salicylate, (Z)-3-hexen-1-ol, (E)-2-hexenal, and (Z)-3-hexenyl acetate. The role of each volatile compound in prey-searching behavior is discussed.     

Response of Predatory Mites to a Herbivore-Induced Plant Volatile: Genetic Variation for Context-Dependent Behaviour

Plants infested with herbivores release specific volatile compounds that are known to recruit natural enemies. The response of natural enemies to these volatiles may be either learned or genetically determined. We asked whether there is genetic variation in the response of the predatory mite Phytoseiulus persimilis to methyl salicylate (MeSa). MeSa is a volatile compound consistently produced by plants being attacked by the two-spotted spider mite, the prey of P. persimilis. We predicted that predators express genetically determined responses during long-distance migration where previously learned associations may have less value. Additionally, we asked whether these responses depend on odors from uninfested plants as a background to MeSa. To infer a genetic basis, we analyzed the variation in response to MeSa among iso-female lines of P. persimilis by using choice-tests that involved either (1) MeSa presented as a single compound or (2) MeSa with background-odor from uninfested lima bean plants. These tests were conducted for starved and satiated predators, i.e., two physiological states, one that approximates migration and another that mimics local patch exploration. We found variation among iso-female lines in the responses to MeSa, thus showing genetic variation for this behavior. The variation was more pronounced in the starved predators, thus indicating that P. persimilis relies on innate preferences when migrating. Background volatiles of uninfested plants changed the predators’ responses to MeSa in a manner that depended on physiological state and iso-female line. Thus, it is possible to select for context-dependent behavioral responses of natural enemies to plant volatiles.     

Testing for Phytochemical Synergism: Arthropod Community Responses to Induced Plant Volatile Blends Across Crops

Using herbivore-induced plant volatiles (HIPVs) to attract specific natural enemies in the field has proven challenging, partly because of a poor understanding of: (i) which compound(s) to manipulate to attract specific taxa, and (ii) the ecological conditions over which HIPVs are effective. To address these issues, we quantified the response of a complex arthropod community to three common HIPVs (methyl salicylate, cis-3-hexen-1-ol, and phenylethyl alcohol) as individual compounds and equal part blends in corn and soybean fields. Of 119 arthropod taxa surveyed, we found significant responses by four species in corn fields (2 parasitoids, 1 herbivore, and 1 detritivore) and 16 in soybean fields (8 parasitoids, 3 predators, 4 herbivores, and 1 detritivore), with both attractive and repellent effects of the HIPVs observed. For example, tachinid flies were highly attracted to cis-3-hexen-1-ol (ca. 3-fold increase), but repelled by methyl salicylate (ca. 60?% decrease). Surprisingly, we found very few cases in which HIPVs acted synergistically; only two arthropod groups (ichneumonid wasps and phorid flies) were more attracted by a blend of the HIPVs than by the individual compounds composing the blend. Crop type, however, had a strong impact on the strength of arthropod responses to HIPVs. A few arthropod species were broadly affected across both crops (i.e., the herbivore Halticus bractatus was repelled by most of our treatments, regardless of crop background), but overall more arthropod groups responded to HIPVs released in soybean fields compared with corn. This was true despite the fact that taxa responding to HIPVs were present and abundant in both systems, suggesting that crop-based outcomes were likely driven by the plant matrix rather than mere differences in taxonomic composition of the arthropod community in corn vs. soybean fields. As a whole, these results suggest that: (i) repellent effects of HIPVs on natural enemies of herbivorous insects can be observed as frequently as attractive effects; (ii) odor blends may be no more effective than single-compound lures for some taxa; and (iii) crop background alters the magnitude of attraction to HIPVs, depending on the species being targeted.     

Volatiles from Psylla-Infested Pear Trees and Their Possible Involvement in Attraction of Anthocorid Predators

Previous work showed that anthocorid predators aggregate around gauze cages containing Psylla-infested trees in a pear orchard. Because anthocorids responded to odor from Psylla-infested leaves in a laboratory test, it was hypothesized that these aggregative responses in the field were triggered by olfaction of compounds associated with Psylla injury. We present chemical analyses of volatiles from damaged and undamaged plants and studies on behavioral responses of anthocorid predators to compounds released by damaged plants. Leaf headspace volatiles from clean and Psylla-infested pear trees were collected on Tenax and identified by GC-MS after thermodesorption. Twelve volatiles were found exclusively in headspace samples from Psylla-infested leaves. Six were present in significantly higher quantities in samples from infested leaves: the monoterpene, (E,E)-α-farnesene, the phenolic, methyl salicylate, and the green leaf compounds, (Z)-3-hexen-1-yl acetate, (Z)-3-hexen-1-ol, 1-hexyl-acetate, and 1-penten-3-ol. These compounds are known to be produced by plants, and damage by pear psyllids seems to trigger their emission. Blend composition varied and was partly correlated with tree or leaf age and degree of Psylla infestation. To study whether compounds associated with leaf injury elicit olfactory responses in anthocorid predators, apple-extracted (E,E)-α-farnesene, synthetic methyl salicylate, and (Z)-3-hexen-1-yl acetate were offered in a Y-tube olfactometer to field-collected adult Anthocoris spp. Significant positive responses were found to both the monoterpene and the phenolic, but not to the green leaf volatile. The results lend support to the hypothesis that predator attraction to herbivore-infested pear trees is mediated by herbivory-induced plant volatiles.     

The Response of Phytoseiulus persimilis to Spider Mite-Induced Volatiles from Gerbera: Influence of Starvation and Experience

When leaves of the ornamental crop Gerbera jamesonii are damaged by the spider mite Tetranychus urticae, they produce many volatile compounds in large quantities. Undamaged gerbera leaves produce only a few volatiles in very small quantities. In the headspace of spider mite-damaged gerbera leaves many terpenoids are present, comprising 65% of the volatile blend. In addition, a number of nitrogen containing compounds, such as oximes and nitriles, are produced.We studied the attraction of P. persimilis to the volatiles from spider mite-damaged gerbera leaves and how attraction is affected by starvation and previous experience. Phytoseiulus persimilis that were reared on spider mites (T. urticae) on Lima bean were not attracted to spider mite-induced volatiles from gerbera. Starvation did not influence the predator's response to these volatiles. In contrast, predators that were reared on spider mites on gerbera leaves were strongly attracted to volatiles from spider mite-infested gerbera. This was found also for predators that originated from a culture on spider mite-infested bean and were offered six days of experience with spider mites on gerbera leaves.     

Differences in Volatile Profiles of Turnip Plants Subjected to Single and Dual Herbivory Above- and Belowground

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.     

Herbivory induces systemic production of plant volatiles that attract predators of the herbivore: Extraction of endogenous elicitor

It was previously shown that in response to infestation by spider mites (Tetranychus urticae), lima bean plants produce a volatile herbivoreinduced synomone that attracts phytoseiid mites (Phytoseiulus persimilis) that are predators of the spider mites. The production of predator-attracting infochemicals was established to occur systemically throughout the spider mitein-fested plant. Here we describe the extraction of a water-soluble endogenous elicitor from spider mite-infested lima bean leaves. This elicitor was shown to be transported out of infested leaves and was collected in water in which the petiole of the infested leaf was placed. When the petioles of uninfested lima bean leaves were placed in water in which infested leaves had been present for the previous seven days, these uninfested lima bean leaves became highly attractive to predatory mites in an olfactometer when an appropriate control of uninfested lima bean leaves was offered as alternative. The strength of this effect was dependent on the number of spider mites infesting the elicitor-producing leaves. Higher numbers of spider mites resulted in an elicitor solution with a stronger effect. In addition, spider mite density was important. The elicitor obtained from one leaf with 50 spider mites had a stronger effect on the attractiveness of uninfested leaves than the elicitor obtained from three leaves with 17 spider mites each. This suggests that the stress intensity imposed on a plant is an important determinant of the elicitor quantity. While the elicitor has a strong effect on the attractiveness of uninfested leaves, spider mite-infested leaves are still much more attractive to predatory mites than elicitor-exposed leaves. The data are discussed in the context of systemic effects in plant defense and the biosynthesis of herbivore-induced terpenoids in plants.